Turfgrass Physiology and Ecology

Advanced Management Principles

This book is dedicated to my wife, Diana, who has supported me in all things
for over 35 years and to my mother and father who taught me the value of
hard work and determination that made this effort possible.
Turfgrass Physiology and Ecology
Advanced Management Principles

Gregory E. Bell
Oklahoma State University
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A catalogue record for this book is available from the British Library, London, UK.

Library of Congress Cataloging-in-Publication Data

Bell, Gregory E.
Turfgrass physiology and ecology : advanced management principles / Gregory E. Bell.
p. cm. -- (Modular text)
ISBN 978-1-84593-648-8 (alk. paper)
1. Turfgrasses--Physiology. 2. Turfgrasses--Ecology. 3. Turf management. I. Title. II. Series: Modular texts.

SB433.B29 2011
635.9'642--dc22
2010028031

ISBN-13: 978 1 84593 648 8

Commissioning editor: Meredith Carroll/Sarah Mellor

Production editor: Fiona Chippendale

Typeset by SPi, Pondicherry, India

Printed and bound in the UK by Cambridge University Press
Contents

About the Author ix

1 Diagnosing Plant Need 1
1.1 An Introduction to Using Scientific Concepts 1
1.2 A Look at Terminology 2
1.3 Using the Science 3
1.4 Observing Turf Performance 6
1.5 Photosynthesis 8
1.6 Respiration 8
1.7 Transpiration 9
1.8 Diagnosing Plant Need 10
1.9 Chapter Summary 11
2 Understanding Photosynthesis 12
2.1 Encourage Photosynthesis to Encourage Turfgrass Growth 12
2.2 The Importance of Light 13
2.3 The Importance of Carbon Dioxide 17
2.4 The Importance of Water 21
2.5 Chapter Summary 22
3 Why C3 and C4 Grasses Require Different Management 24
3.1 Use Species Adaptation to Your Advantage 24
3.2 Photorespiration 25
3.3 Kranz Anatomy 30
3.4 Characteristics of C3 and C4 Plants 33
3.5 Chapter Summary 35
4 Respiration and Transpiration 36
4.1 Encouraging Efficient Use of Carbohydrates and Water 37
4.2 Plant Respiration 37
4.3 Transpiration 51
4.4 Chapter Summary 57
5 Why Our Management Practices Affect Our Turf 58
5.1 Understanding What We Know From a Turfgrass Perspective 58
5.2 Mowing Causes Chronic Damage to Turf 60
5.3 Thatch and Thatch Management 66
5.4 Issues Pertaining to Soil Compaction 72
5.5 Managing Wear Caused by Traffic 72
5.6 Chapter Summary 73
6 The Importance of Light and Managing Shade 75
6.1 The Practice of Making Adjustments to Improve Photosynthesis in Shade 75
6.2 The Influence of Irradiance 76
6.3 Managing Turf in Shade 81
6.4 Chapter Summary 86

v
 7 Understanding and Prescribing Nutrition 88
7.1 Manage Your Fertilization to Match Your Turfgrass Needs 88
and Local Conditions
7.2 Basic Soil Attributes 89
7.3 Plant Nutrition 93
7.4 Chapter Summary 105
8 Irrigation and Water Management 107
8.1 Manage Your Irrigation and Drainage for Site-specific Objectives 107
8.2 Irrigation Management 108
8.3 Species Adaptation to Low Water Use 123
8.4 Managing Water Overload 124
8.5 Chapter Summary 127
9 Adjusting for Seasonal Conditions and Temperature Stress 129
9.1 Optimal Turfgrass Health is a Result of Flexible 130
Management Under Differing Temperatures
9.2 Seasonal Turfgrass Management 130
9.3 High-temperature Stress in Grasses 131
9.4 Cold-temperature Stress in Grasses 139
9.5 Chapter Summary 144
10 Growing Grass on Soil, Sand and Salt 146
10.1 Different Media Require Different Management 146
10.2 Problems With Soil 147
10.3 Problems With Sand 150
10.4 Sand System Design and Construction 152
10.5 Managing Turfgrass on Soil and Sand 154
10.6 Managing Turf on Soils High in Salts 156
10.7 Chapter Summary 161
11 The Ecology of Turfgrass Management 163
11.1 How to Make Nature Work For You 164
11.2 Introduction to Ecological Theory 164
11.3 Turfgrass Ecology 170
11.4 Chapter Summary 174
12 Managing Competition Among Plant Species 175
12.1 Manage Your Sites to Favor Turfgrass and Discourage Other Plant Species 175
12.2 Competition Among Turfgrass Species 176
12.3 Competition Between Turfgrass and Other Plants 177
12.4 Biological Herbicides 183
12.5 Chapter Summary 185
13 Managing Competition Between Turf and its Pests 187
13.1 How Can We Affect Relationships Between Predator and Prey? 187
13.2 Predator/Prey Relationships 187
13.3 Managing Turfgrass Predators 188
13.4 Affecting Relationships Between Predators and Turf 193
13.5 Making a Management Plan 196
13.6 Chapter Summary 199

vi Contents
14 Making the Right Decisions 201
14.1 Learn to Use Your Knowledge to Your Advantage 201
14.2 Planning the Project 202
14.3 Turfgrass Establishment 205
14.4 Making a Management Plan 207
14.4 Chapter Summary 210

References 211
Index 227

Contents vii
This page intentionally left blank
About the Author

Dr. Greg Bell is the Wayne and Jean Huffine Endowed Professor of Turfgrass Science at Oklahoma State
University, Stillwater, Oklahoma. After many years as an industrial supervisor and business owner, Dr. Bell
returned to school at the Ohio State University to finish a B.S. in Turfgrass Management, followed by a
Masters and Ph.D. in Agronomy with specialty in turfgrass science. Greg has published a number of scientific
papers, book chapters and trade journal articles in turfgrass science, and was part of a research team that
was awarded the United States Department of Agriculture Secretary’s Honor Award in 2002. He was named
the Outstanding Undergraduate Student Advisor in the College of Agricultural Sciences and Natural
Resources at Oklahoma State University in 2001 and was awarded the Faculty Phoenix Award dedicated to
the outstanding graduate student advisor of the year at Oklahoma State University in 2003. Dr. Bell teaches
classes in Introductory Turfgrass Management, Turfgrass Physiology and Ecology, and Personnel and
Financial Management in Horticulture.

ix
This page intentionally left blank
 Diagnosing Plant Need
1

Key Terms
Best management practices (BMP) for turfgrass are the most effective techniques known at any given time for
managing high-quality turf with little or no off-site effects.
Capillary action is the result of adhesive and cohesive forces that cause water to rise upward against gravity in
a very small tube.
Osmosis is the movement of water through a permeable membrane from high water potential to low water poten-
tial such as when roots take up soil water.
Photosynthesis is the process plants use to convert light energy into chemical energy.
Respiration is the process plants use to release the energy trapped by photosynthesis.
Transpiration is the transport and evaporation of water that is used to dissipate the heat that is produced in the
plant as a result of its metabolic processes, such as photosynthesis and respiration.
Turfgrass biology is the set of growth and developmental patterns that can be expected of turfgrass plants in
response to all environments conducive to life.
Turfgrass ecology is the interaction of turfgrasses with other living organisms and other non-living entities, both
natural and artificial, within the turfgrass environment.
A turfgrass environment is the sum of all natural and artificial conditions under which a particular stand of turf is
expected to live and grow.
Turfgrass physiology is the awareness of the inner physical mechanisms and chemical processes that perpetuate
life in a turfgrass plant.

1.1 An Introduction to Using Scientific

that fertilizer and water and how to keep it healthy
Concepts
under a variety of both natural and artificial
We all learn from experience. In fact, we may be conditions.
able to learn everything we ever need to know from Start with the power of observation, add some
experience. However, if we combine our knowledge experience, factor in some knowledge, and assign
learned from classrooms, from books, from friends some responsibility to yourself and you have the
and family and from other sources with our beginning of a formula for diagnosing plant need.
experiences, we can become truly wise. First, you have to know what the plant should look
There is no secret to managing turfgrass reason- like at any given time. This you learn mostly from
ably well most of the time. Just add fertilizer and experience and a little knowledge. Second, you
water. However, if you intend to be a turfgrass need to know how to make the plant look better
manager, your job is to manage turf exceptionally and grow better if it is not meeting your customer’s
well all of the time and do it economically with the expectations. This you do mostly with knowledge
least amount of input and with little or no impact and a little from experience. Finally, you need to
on people, animals and off-site locations. know how to identify the problem if the plant is
Consequently, you need to know when to add fer- not responding to your input. That is why not just
tilizer and water and how much to add. You also anybody can do this job. That is why your cus-
need to know how the turfgrass should react to tomer needs you.

©CAB International 2011. Turfgrass Physiology and Ecology (G. Bell) 1

 1.2 A Look at Terminology protection practices. Dr. McCarty and others, for
instance, authored the outstanding book that
As an instructor, I am constantly reminded that
I referred to earlier using the term BMP to include
undergraduate students, sometimes graduate stu-
a wide range of environmental management tech-
dents and, often, practitioners, do not like to hear
niques. Those techniques may or may not include
terms like “plant biology”, “plant physiology” and
chemical applications and water-protection prac-
“plant ecology”. Perhaps that is because they do
tices. So a contemporary definition of BMP for
not fully understand what those terms mean. There
agronomic purposes could be: the best-known
are lots of popular terms that are confusing and
procedure for encouraging or maintaining plant
don’t seem to make sense. Let’s take a look at three
health without detrimentally affecting off-site
terms that are often used to describe conditions or
conditions. In simple terms, contemporary turf
situations, but are not clearly defined.
BMPs are protocols that we use to grow good
grass without hurting anything else in our
Best management practices environment.
One of those terms that begs for a clear definition
Environment
is “best management practice” (BMP). Literally,
BMP identifies a supervisory or technical proce- “Environment” is another of those constantly
dure that cannot be improved upon because it is evolving terms. In this case, however, the term has
already the best. However, that is not what the also been highly misused and abused. Environment
term really means. The true connotation of the has a very simple definition: the aggregate of all
term is not evident in its literary meaning. surrounding conditions that influence an organism
According to Balogh and Walker in Golf Course (Beard, 1973). Environment can be used to describe
Management and Construction: Environmental the conditions that exist on a dish, either a petri
Issues (Balogh and Walker, 1992) and McCarty dish or your dinner dish, or it can be used to
et al. in Best Golf Course Management Practices describe conditions that exist across an entire coun-
(McCarty, 2001), a BMP used in golf course man- try or around the world. The petri dish or dinner
agement, and presumably in other turf endeavors, dish are better applications of the term than the
should achieve five basic goals: (i) decrease off- more commonly used version, which refers to all
site transport of pesticides and nutrients; (ii) con- natural entities that exist over a nearly unlimited
trol the application of pesticides and nutrients; location such as a country, a continent or the
(iii) decrease total chemical loads; (iv) use both world. In fact, an environment does not have to
biological and mechanical soil conservation plans; consist of natural conditions. If you place the petri
and (iv) educate both managers and the public dish in an oven, it now exists in an artificial envi-
about the relationship between environmental ronment, not a natural one. If microbes happen to
issues and golf course management. Although we be living on that dish, they are natural organisms
may still be a little confused about BMPs, we at living in an artificial environment. The same could
least know that the term does not necessarily refer be said for certain turfgrass conditions. For instance,
to a “best” practice unless it happens to be the a golf course putting green built to United States
best practice known at that particular time for Golf Association recommendations could be said
that particular location. We also now know that a to include natural organisms, one of which is the
BMP specifically refers to chemical application grass, living in an artificial environment (USGA
management – or does it? Actually, the term was Green Section Staff, 1993). So the question becomes:
originally used to refer to techniques used to man- if the environment is artificial, who or what is con-
age waste water and storm water runoff. Its first trolling it?
use may have been in the 1987 amendments to the On a USGA golf course putting green, the envi-
U.S. Clean Water Act (U.S. Congress, 1987). The ronment is primarily controlled by the grounds
term later evolved to include practices that reduce manager or golf course superintendent. There are
chemical contamination of surface and ground some conditions, usually temperature for instance,
water following application. However, it appears that the manager cannot control or has minimal
that BMP has evolved further into its more literal influence over. There are other conditions, such
translation to include more than just water- as shade and traffic, that the manager could

2 Chapter 1
control and would like to control but may be 1.3 Using the Science
forced to accept. Because the manager can con-
This discussion began with an introduction of
trol or influence nearly all of the environmental
three terms, “plant biology”, “plant physiology”
conditions on the golf course putting green,
and “ecology”. After the three scientific terms
the turfgrass growing there is at the mercy of the
were introduced, three popular terms were
manager. If the manager is knowledgeable in the
introduced. It should be clear that “BMP” has a
needs of the turf, the turf will thrive, if not, it will
lofty connotation and represents highly desirable
probably die. The same could be said for a home
principles, but it tends to take on the definition of
lawn, a park or any other location where turf-
the person using the term at the time. The same is
grass is cultivated for human use. The putting
true of “environment”.
green is a good example because even the grow-
“Environment” was never meant to mean all
ing medium is highly artificial and because the
natural entities in a given area. According to
conditions required to satisfy the golfer’s expecta-
Merriam-Webster (2009), environment is the
tions are severe. In fact, the closer the turf is to
circumstances, objects or conditions by which one
death, the better the golfers seem to like it.
is surrounded. For our purposes, a more scientific
Especially competent turfgrass managers like to
definition is appropriate and can also be found in
work with these artificial environments because
Merriam-Webster: “the complex of physical, chem-
they control, or nearly control, most, or all, of
ical, and biotic factors (as climate, soil, and living
the environmental conditions. The more environ-
things) that act upon an organism or an ecological
mental influence the knowledgeable manager has,
community and ultimately determine its form and
the better the manager likes it. Less competent
survival”. Apparently, if a BMP is going to have a
managers, homeowners for instance, would rather
positive effect on turfgrass growth or survival, it
manage plants under more natural conditions. It
must also affect the environment in which the turf
is easier and it requires less knowledge. The best
lives. Consequently, a BMP has to affect the
looking home lawns are managed by knowledge-
turfgrass environment but should not affect the
able homeowners or by professional lawn-care
surrounding environment detrimentally.
managers.
Our third popular term, “best looking”, is
extremely important to turfgrass management but
Best looking
it is the hardest to define. “Best looking” is a mat-
“Best looking” is a relative term. It is a term rela- ter of opinion or personal preference and can differ
tive to an individual and, therefore, to location. substantially by individual. However, turfgrass and
The best looking home lawn in Europe is not the other horticultural plants are used not only for
same as the best looking home lawn in Australia, is their functional characteristics but also for their
not the same as the best looking home lawn in the aesthetic value. Therefore, “best looking” is not
USA. The lawns differ because of differing cus- just a result of good management, it is a condition
tomer expectations, different rules and regulations, that turfgrass managers strive to affect. “Best
and different environments. Notice that the term looking” often determines the BMPs that the
“environments” is not used here to describe the turfgrass manager incorporates into the turfgrass
environment in Europe or the environment in management plan. “Best looking” determines when
Australia, it is used to describe the environment of to irrigate, when to fertilize, and when and how to
the best looking lawn in Europe or the best looking apply other management practices. Yet, “best
lawn in Australia. Consequently, if we can identify looking” is a matter of personal preference and is
the best looking lawn, we can characterize the best seldom the same for different locations or cultures.
environment for growing home-lawn turf on that European turfgrass managers, for instance, do not
particular medium in that particular vicinity. We share the same preference for dark green turf that
can also identify the best management practices for their counterparts in the USA desire.
growing home-lawn turf on that particular medium If these last few paragraphs were somewhat
in that particular vicinity and we could possibly confusing, don’t be alarmed. They were meant to
improve on those BMPs with a general knowledge be. I introduced a term that was coined not for its
of plant biology, plant physiology and plant literary value, but for its “feel good” effects, BMP.
ecology. I also discussed a term that has been twisted by

Diagnosing Plant Need 3

society to mean things that it was never supposed science”. One of the crop production scientists in
to mean, environment, and finally, a term that has the front row jokingly made the comment “isn’t
a substantially different meaning depending on turfgrass science something of an oxymoron?”.
who is using it at the time, best looking. Certainly, I immediately flashed up a slide of researchers
scientific terms such as plant biology, plant physiol- applying treatments to a golf course and before
ogy and plant ecology are not so confusing as these I could say anything another turfgrass scientist said
three popular terms and others like them that we “that sure beats working in a hay field doesn’t it?”.
are comfortable using daily. Yes, we do practice science and most of those tech-
In order to be the best turfgrass managers that niques for managing turfgrass that you have
you can be, you have to make decisions based on learned on and off the job came from us. As scien-
plant biology, plant physiology and plant ecology. tists go, turfgrass scientists tend to do a lot of work
We already know the basic principles that we need for little money and we tend to do a lot more
to know to make decisions based on these scientific applied work, research that can be used in the field,
factors. In the following 200 pages or so, I will help than we do basic work, research that leads to
you relearn these basic principles, not in any great greater understanding but is not immediately
detail, but in enough detail that you can use them applicable in the field. Both types of research are
successfully. We all learn best from experience, but important. As you will see, we are going to use
we can learn so much more from our experiences if basic science throughout this book to help us make
we use our knowledge as well. It is good to know decisions that we can apply in the field.
how to accomplish a task successfully. It is better to
know why the procedures that we used to complete
Turfgrass research
the task worked so well. For the time being, let’s
forget about how you do something and focus on We use research for a variety of reasons, including
why you do it instead. testing new products, testing new grasses, testing
new maintenance procedures alone or in combina-
tion with old procedures, and gaining knowledge in
Plant biology
plant growth or metabolism that may lead to
Plant biology is the same as botany and can be improved varieties, improved cultural practices,
defined as a science involving the study of plants improved weed control and other improvements.
(Stern, 1991). I suppose that means that if you stare Experiments are effective because they measure
at a plant for a few hours then you are performing treatment differences under comparable condi-
botany. Actually, no; if you stare at a plant for a few tions, because they answer questions posed by
hours, you are performing basically nothing, which specific objectives and because they answer ques-
is not a bad thing as long as you don’t make a habit tions deemed important. An experiment is a planned
of it. As botany is a science, it involves more than inquiry to obtain new facts or to test previous
staring and as it is a science, there is actually more results that aid in decision-making (Waddington
research involved than there is learning. People like et al., 1992). Consequently, scientific experiments
university instructors do the research so that people are governed by a fixed set of research protocols.
like university students can learn from it. Obviously, Although a great deal of useful information has
the instructors, researchers and scientists who did been accumulated by simple observation, an equal
this research did it so that they and newer instruc- amount of misinformation has probably been accu-
tors could learn from it and pass it on to their stu- mulated in the same manner. Research protocols
dents. Consequently, we need to know a little about are specifically designed to minimize misinforma-
research to understand botany. As you might have tion and are consequently more likely to overlook
guessed, we are not interested in research done on a positive result than to verify a result that is not
all of the plants in the universe; we are only inter- true. Research can be as simple as applying a her-
ested in the research done on turfgrass. bicide to see how many weeds are killed or it can
be as complex as mapping the human genome.
Although the complexity of research projects varies
Turfgrass science
widely, the same scientific protocol applies in all
I was giving a presentation to a group of scientists cases. Based on scientific protocol, if a turfgrass
several years ago when I used the term “turfgrass scientist reports that a particular herbicide is

4 Chapter 1
effective for controlling a particular weed, the sci- not the biological principles that are difficult, it is
entist should be right 95% of the time. That will the flexibility to make decisions and the confidence
always be true, regardless of the experiment, as to know you are right that is difficult.
long as the scientist works within the generally
accepted 95% confidence interval.
Plant physiology
Most of what we know about turfgrass manage-
ment has been learned either by simple observation Plant physiology is the science that studies plant
or by experimentation. What we know about turf- function (Salisbury and Ross, 1992). That is a sim-
grass biology, however, has all been learned through ple, yet extremely broad, definition similar to the
experimentation. In other words, we aren’t guess- definition that we used for plant biology.
ing. We are basing our conclusions on experiences Fortunately, we do not need to know a great deal
that occur repeatedly under the same circum- about the physiological processes of turfgrass in
stances. Each biological principle has had to with- order to make sound management decisions.
stand rigorous tests. If we are aware of these Photosynthesis, respiration and transpiration are
principles, we can expect a plant to respond to a physiological plant processes. They are activities
given stimulus in the same way each time the that take place inside the plant but result in exter-
stimulus occurs, provided that the same environ- nal activity or aesthetics that we can often see or
mental conditions are present each time. Using measure. It is not difficult for an experienced man-
scientific information, we know when the plant ager to determine when transpiration has slowed
will grow rapidly, when it is likely to be dormant and the turf needs water. However, it is difficult for
or quiescent, when it is susceptible to damage and, a manager to determine when photosynthesis is
most importantly, when it needs fertilizer and occurring too slowly or respiration has been com-
water. We know how low we can mow it under a promised. Poor photosynthesis or poor respiration
given set of conditions, we know when we can manifest themselves in basically the same symp-
perform aggressive cultural management, and we toms that occur when turfgrass is under nearly any
know when we should leave it alone. That’s right; type of environmental stress. Like disease symp-
doing nothing can sometimes be an excellent man- toms, there are nuances that can help us identify
agement practice; one that is often overlooked. the source of the stress, but the symptoms are gen-
Biologically speaking, we can expect a turfgrass erally similar because they all affect photosynthe-
plant to respond to a given set of environmental sis, respiration and/or transpiration.
conditions in the same way each time that those When a turfgrass is exposed to saturated soil
conditions occur. Therefore, managing turfgrass is conditions for an extended period of time, it turns
easy. All we have to do is perform the same man- yellow, then brown, then dies in a matter of days
agement practices at the same time in the same depending on species. How does this stress from
amount each season and our turfgrass will always saturated soil differ from the stress experienced
grow well and look good; or will it? Biologically when turfgrass is exposed to very dry soil for an
speaking, that is true provided that the same set of extended period of time? Actually, there is very lit-
environmental conditions occurs each year at the tle difference in the effects of very wet and very dry
same time and in the same amount. That doesn’t soil. In fact, in both cases, the turf dies of drought.
happen. Yet we are still looking for that magic set When the soil is dry, there is no water for the plant
of management practices that makes our turf per- to take up. When the soil is wet, the roots are sur-
form to our expectations year after year regardless rounded by water but they have no air. No air
of environmental differences. That magic recipe means no oxygen, no oxygen means no respiration,
doesn’t exist, but if we consider the environment no respiration means no energy to take up water
independently of what we did last year or last and the plant dies of drought. How do you keep
week, we can make decisions based on simple bio- this turf alive in saturated soil until the soil has
logical principles that will help our grasses perform time to drain?
as best as can be expected nearly all of the time. In All that you need to know to solve this problem
this text we will pursue that goal. The biological is a miniscule amount of plant physiology. You
principles needed are often as simple as “turfgrass need to know that roots have to respire to take up
grows well when it has water but not well when water and that oxygen has to be present for the
there is too much water or too little water”. It is roots to respire. Once you have realized that these

Diagnosing Plant Need 5

conditions are not present, you know that you have that are best adapted will survive, and because
to get water to the plants or they will die. intraspecific competition also occurs, only the strong-
Consequently, your only option is to apply water in est individuals of that species and their offspring will
foliar sprays that the plants can take up through remain. Although I would not want to be the one to
the leaves. The long-term fix is to provide better try to prove this theory, the concept is a good one
drainage to the area, but the short-term fix is to and extremely useful for plant management.
frequently spray water on to the leaves of plants In order to use ecology to your advantage, you
whose roots are sitting in saturated soil. The only have to have a sound knowledge of your target plant,
way that you can make that conclusion is through your turf, and at least a basic knowledge of its com-
a simple knowledge of plant physiology. Neither petitors, as well as of the neutral, predatory and
experience nor common sense would lead you to beneficial organisms with which it shares their envi-
the conclusion that you should spray water on to ronment. That sounds complex and it is. Ecological
turf that is sitting in saturated soil. However, as a management is not easy but we all practice it. The
person who knows a little about plant physiology, more knowledge you can use and the more experi-
a simple assessment of the problem should lead you ences that you have, the easier it becomes.
to that conclusion even if you have never experi-
enced such a condition. That is the power of simple
1.4 Observing Turf Performance
scientific knowledge.
Turfgrass performance varies considerably accord-
ing to species, climate, weather, time of year and
Plant ecology
several other factors. Therefore, it is difficult to
Ecology is the study of the interactions of organ- know exactly what your turfgrass should look like
isms with one another and with their environment at any given time unless you have considerable expe-
(Starr, 2000). That does not sound like much, but rience with a particular turf in a specific location.
ecology is powerful. Understanding which environ- Site-specific factors such as low areas, shaded areas,
mental factors favor a weed and which favor turf south-facing slopes and many other conditions also
can save a turfgrass manager considerable time and affect turfgrass performance. As part of our research
money. The objective of the turfgrass manager is to (Xiong et al., 2007), we observed the performance
influence the environmental conditions so that they of a cool-season grass, creeping bentgrass (Agrostis
favor the turf at the expense of competing organ- stolonifera), and a warm-season grass, bermuda-
isms. Disease, for instance, occurs when the envi- grass (Cynodon dactylon), over two growing sea-
ronmental conditions favor the growth of a sons in the south-central USA using the normalized
pathogen more than they favor the growth of turf. difference vegetation index (NDVI), an objective
If the manager can minimize or postpone the occur- measure of reflected irradiance, to determine the
rence of certain environmental conditions that approximate color of those grasses over two grow-
favor the pathogen instead of the turf, the manager ing seasons (Fig. 1.1). An NDVI of 0.6 is considered
can reduce the need for pesticides and minimize the green but slightly yellow and an NDVI of 0.9 is very
amount of disease that occurs. dark green. Based on our observations, the NDVI
For purposes of this book, we will refer to plant and the color of these grasses changed considerably
ecology as turfgrass ecology, thereby limiting the during the growing season in spite of the fact that
interactions in a given environment to those most they were fertilized consistently and watered suffi-
important to turf. There could be a hundred or more ciently. Consequently, unless you are very familiar
species living in an ecological system containing turf, with a certain grass in your location, it is unlikely
but we are primarily concerned with those species that you could accurately predict what that grass
that have the greatest influence on our turf. This should look like during a particular time of year. In
ecological system, simply called an ecosystem, con- addition, if weather conditions are different this year
tains microbes, insects, plants and other living organ- from last year, and they almost always are, it is likely
isms constantly competing for nutrients and space that these patterns will be, at the very least, slightly
(Danneberger, 1993). The organisms that are best different between the two seasons. Making turfgrass
adapted to the environment of the ecosystem flourish management decisions based on visual assessment
and those that are not so well adapted decline. requires a highly experienced manager, yet that is
Theoretically, given enough time, only the species how most of us attempt to do it.

6 Chapter 1
 0.9

0.8

NDVI
0.7

0.6 Creeping bentgrass

Common bermudagrass

0.5
06 Feb 28 Mar 17 May 06 Jul 25 Aug 14 Oct 03 Dec
Day of year

Fig. 1.1. Approximate color of creeping bentgrass (Agrostis stolonifera) and bermudagrass (Cynodon dactylon)
measured using the normalized difference vegetation index (NDVI) averaged over two growing seasons in Stillwater,
Oklahoma. A score of 0.6 is slightly yellow-green turf and a score of 0.9 is very dark green turf.

Earlier in this chapter, I suggested the use of called plant physiology and it can help us identify
biology, physiology and ecology for making plant need and manage turf accordingly.
management decisions pertaining to turf. According There are three physiological processes that are
to biology, a grass should respond to a given set of of particular importance when making plant
environmental conditions in approximately the management decisions. Those are photosynthesis,
same way every time those conditions are present. respiration and transpiration. Photosynthesis is
However, plant biology is obviously a very com- the means by which the plant makes food, respira-
plex method for making turfgrass management tion is the means by which the plant turns that
decisions, especially if we consider the huge scope food into energy and transpiration is the means by
of a turfgrass environment and the number of which the plant cools itself sufficiently to perform
changes that could occur. We need a decision- photosynthesis and respiration continuously.
making system that is simpler than basic plant bio- There are an enormous number of physiological
logy and yet more attainable than relying on a processes that occur in a grass plant but, for now,
multitude of experience. In order to find that sim- those three processes will be sufficient for our
pler system, let us look a little deeper into the purposes.
cause-and-effect relationship between biology and Physiological processes are far from simple. They
environment. are extremely complex. Scientists can spend their
According to science, biological responses are entire lives trying to unravel the biochemical reac-
primarily predetermined by genetic code. The tions in a small portion of the photosynthetic path-
genetic code encrypted on the plant’s DNA regu- way. Those reactions are in constant flux and the
lates which proteins and enzymes, and the amount pathways that regulate those reactions can be more
of those proteins and enzymes, that are present in complex than the reactions themselves. It is
the plant under a given set of conditions. The understandable why many students get confused by
enzymes produced encourage chemical reactions these systems. However, the presentation of photo-
that enable or disable metabolic pathways. These synthesis, respiration and transpiration need not be
metabolic pathways determine the health of a complex. It is far beyond the scope of this text to
plant, its appearance, how well it competes with try to dissect these three processes. Instead, we only
other organisms and how quickly it grows and need to know the basics of these systems to make
spreads. The study of these metabolic pathways is them useful to us.

Diagnosing Plant Need 7

 1.5 Photosynthesis where it was produced. However, glucose is not the
only sugar produced by photosynthesis. The other
There are two types of photosynthesis, warm-
sugars produced are primarily used to synthesize
season and cool-season photosynthesis, that occur
other compounds that the plant needs. Yes, photo-
in grasses. Warm-season photosynthesis is called C4
synthesis produces the energy that powers all living
photosynthesis and cool-season photosynthesis is
things. However, it also produces the sugars that
called C3 photosynthesis. Like it or not, these dis-
are used to synthesize amino acids and fatty acids
tinctions are important to you and we will discuss
for the construction of proteins, lipids and the
them in a later chapter. In simple terms, photosyn-
other compounds necessary for plant survival.
thesis is the means the plant uses to turn radiant
Glucose is also combined to make cellulose and
energy into chemical energy. In fact, all of the bio-
other products used to form structural plant tissue.
logical energy on our planet comes from the sun
The glucose that is not used for energy or the syn-
and is channeled to us by the process of photosyn-
thesis of other products in the cell where it was
thesis. Plants trap energy, then we eat the plants or
produced is converted to sucrose and transported
we eat an animal that ate the plants and that is how
to areas of the plant where energy is needed. If the
we survive. Consequently, photosynthesis is not
plant has an excess of energy, the sucrose is trans-
just important to you because you want to be a
ported to cells, usually in the stems and roots,
turfgrass manager, it is also important because it
where it is converted back to glucose and stored as
keeps you alive.
starch (C4 plants) or is converted to fructose and
Photosynthesis requires light, water and carbon
glucose and stored as fructans (C3 plants). There
dioxide. If one of these important components is
the stored products remain until energy is needed
low or missing, photosynthesis will slow or stop. If
or it is used for conversion to other compounds.
the condition does not improve, the plant will
Because a plant can photosynthesize, it does not
become sick and eventually die. Other nutrients are
have to forage for food and it can make everything
required to perform photosynthesis as well but they
it needs as long as sufficient nutrients are present.
are rarely in low supply and are not nearly as
All the plant requires is a medium (soil) that con-
important as light, water and carbon dioxide.
tains the minimum nutrients necessary for survival,
Consequently, it is our job, above all else, to make
plus light, water and carbon dioxide.
sure that our turf has sufficient light, water and
Photosynthesis is so important that it should
carbon dioxide. You may believe that light and
always be our primary focus. Nothing else occurs
carbon dioxide are beyond your control, but that is
without it. We must always maintain sufficient
not correct. We can have a substantial influence on
light, water and carbon dioxide for our turf. If the
those components in most situations. If, for some
turf declines, we should first eliminate the possibil-
reason, it is impossible to deliver sufficient light,
ity that light, water or carbon dioxide are not suf-
water or carbon dioxide to an area where you want
ficient before we look for deficiencies elsewhere.
to grow turf, give up and mulch it.
Photosynthesis takes a portion of the energy the
plant requires in light, converts it to chemical 1.6 Respiration
energy, and stores it in carbon bonds. You are Respiration is the process by which the plant
probably familiar with the general principle repre- releases the energy stored in the chemical bonds
sented in the following chemical equation: produced by photosynthesis, and oxygen is required
for respiration to occur. Other nutrients, especially
6CO2 + 6H2O + light ® C6H12O6 + 6O2
phosphorus, must also be present but oxygen is the
which shows that carbon dioxide (CO2) plus water most important. Respiration consumes oxygen and
(H2O) in the presence of light yields glucose (C6H12O6) gives off carbon dioxide, as represented in the fol-
and oxygen (O2). Glucose, of course, is a sugar and lowing equation:
it has five carbon bonds that store energy. C6H12O6 + 6O2 + light ® 6CO2
The glucose equation represents the major pur-
+ 6H2O + energy
pose and events in photosynthesis, but it is an
oversimplification. In fact, much of the energy which shows that glucose (C6H12O6) plus oxygen
from light is converted into glucose, which is (O2) yields carbon dioxide (CO2) plus water
temporarily stored as starch in the chloroplast (H2O) plus energy. For that reason, respiration

8 Chapter 1
complements photosynthesis. Photosynthesis takes Water uptake for transpiration
in carbon dioxide and gives off oxygen, whereas
If you add salt, sugar, or any other soluble com-
respiration takes in oxygen and gives off carbon
pound to water, it lowers its water potential. So if
dioxide. It would seem that a turfgrass plant would
you were to place a permeable barrier between
be in complete balance with its environment. It
pure water and water containing solutes, the pure
takes in what it needs and gives it back in another
water would move toward the water containing
process. However, we must remember that respira-
solutes. The water contained in a plant’s soil
tion produces nothing except carbon dioxide, water
medium is high in solutes. In fact, many of the
and energy for plant functions. Photosynthesis,
solutes in soil water are plant nutrients.
however, produces sugars for energy and sugars
Consequently, these solutes are very important to
used to synthesize other compounds as well as all of
our plant. The soil water has a water potential
the structural components of the plant. Consequently,
lower than pure water. However, the water inside
photosynthesis must take in considerably more car-
the plant contains even more solutes than the water
bon dioxide than respiration produces.
in the soil and nearly always has a lower water
Respiration occurs in all plant parts. There is
potential than soil water. Consequently, the soil
more than sufficient oxygen for respiration to
water is attracted through the roots into the turf-
occur in the plant parts located above ground.
grass. Once inside the turfgrass, the plant can
After all, the air is 21% oxygen. The area where
remove whatever nutrients it needs and fill its cells
oxygen could be deficient is below the soil surface.
with water. Most of the water, however, is not used
Compacted or saturated soils are examples of situ-
for metabolism; it is drawn through the grass and
ations where sufficient oxygen is not present for
evaporated into the air. This evaporative process is
adequate root respiration. That is why turf declines
used to cool the plant. Otherwise it would die from
rapidly when those conditions occur and why
the heat produced by its own metabolic processes.
sports field managers aerify at every opportunity.
Energy transfer is not a perfectly efficient proc-
Aerification, of course, is a procedure where the
ess. Any time that energy is transferred or trans-
manager opens the soil to air infiltration thereby
formed, a certain amount of the initial energy is
helping to provide oxygen to the roots. Aerification
lost. A huge number of chemical reactions are
is also used to break up soil layers and to encour-
required to perform plant photosynthesis, respira-
age water infiltration, but those are subjects for
tion and other important functions. Each time one
another chapter. Suffice to say, that if we perceive
of these reactions occurs, a small amount of energy
no barriers to photosynthesis above ground then
is lost. It follows then that the amount of light
we should be making sure that there are no barriers
energy accumulated by photosynthesis is not the
to respiration below ground.
same amount that is released through respiration.
Only a portion of that energy is available for use by
1.7 Transpiration the plant, the rest is mostly heat. That heat has to
be dissipated otherwise our plant will die of some-
Transpiration is a simple plant system, but also a
thing like a very high fever. When we work hard,
very important one. In basic terms, it is a movement
our metabolism increases and we produce excessive
of water from the soil through a plant and into the
heat. We dissipate that heat through perspiration.
air. It occurs because of osmosis, cohesive forces,
A plant dissipates heat through transpiration. Some
adhesive forces and evaporation, and is fueled by
heat is also dissipated through conduction and
water potential. If you have had a soils class, you
convection, but transpiration is the most important
have studied water potential and you know that
system for plant cooling.
water moves from high water potential to low water
potential. So for transpiration to work, the water
potential in the plant has to be lower than the water
Water movement from roots to leaves
potential in the soil, and the water potential of the
air has to be lower than the water potential of the Once soil water is in the plant, it circulates among
plant. During transpiration, water moves from the and around the cells mostly by diffusion, a relatively
soil into the plant by diffusion, through the plant by slow process. It can also move upward into the leaves
cohesion, adhesion and water potential, and into the and other plant parts as it is drawn by the relatively
air by evaporation. low water potential of the air. You may be familiar

Diagnosing Plant Need 9

with the capillary tubes that are used for a variety light? Why is turf difficult to manage in the shade
of purposes and often to demonstrate adhesive and of trees? Why does turf require more water when it
cohesive forces in chemistry class. A capillary tube is warm and windy than it does when it is cool and
is a glass tube with a very small opening. You could calm?
call it a little glass pipe. If you put this capillary tube We have already answered those questions, not
in water, the water fills the tube even if you are literally, but logically, in the preceding sections of
holding the tube vertical and only the low end is in this chapter. Diagnosing the needs of your turf
the water. So the tube causes the water to move requires some thought. The preceding three ques-
upward against gravity. That is called capillary tions are relatively easy compared with those we
action. Plant scientists are quick to point out that often face in the field. However, they still require
capillary action is not the means by which plants thought.
move water for transpiration, but the same princi- Turf surrounded by buildings is difficult to man-
ples are used to help it along. The water molecules age because the air movement at ground level is
adhere or stick to the sides of the tube. They also restricted. Our planet’s air is only 0.035% carbon
stick to each other, a force called cohesion. The dioxide. Photosynthesis requires light, carbon diox-
adhesive and cohesive forces hold the water to the ide and water. In this situation, we have light, we
sides of the tube and to itself so strongly that the can add water, but we are limited in the amount of
water overcomes gravity and moves up the tube. If carbon dioxide available. As photosynthesis pro-
the tube was larger, the cohesive forces would not ceeds, our plants take in carbon dioxide and give
be strong enough to hold the water together across off oxygen. If air movement is restricted, fresh air,
the width of the tube. In that case, a little bit of hence fresh carbon dioxide, is not available.
water would stick to the sides of the tube but the Consequently, the air above the turf becomes high
rest would fall to the bottom. The xylem that par- in oxygen and low in carbon dioxide, and photo-
tially makes up the veins of a plant is like a capillary synthesis stalls.
tube, but it is not open to the air, which is another Turfgrasses don’t grow well in the shade of trees
requirement of capillary action. Consequently, cap- because sufficient light is not present. Again, we
illary action does not drive transpiration but the have a problem with photosynthesis. Our turf
two forces that explain capillary action must be needs more water when it is warm and windy
present for transpiration to occur. It is generally because transpiration is based on evaporation and
believed that water potential drives transpiration the warmer it is and the windier it is, the faster
and cohesive and adhesive forces help hold the evaporation occurs. There are other reasons as
water columns together in the plant and keep them well, that you can probably figure out, but we will
from falling. There is also a root pressure effect that save those for later chapters.
sometimes forces water upward, usually during the Biological problems are nearly always compli-
night, but this is not a major mechanism for the cated and more is rarely better. Plants are biological
movement of water in transpiration. systems just like us. Food is a good thing for us but
We are all familiar with the cooling effect of too much makes us unhealthy. If we have a head-
evaporation, but did you also realize that the air ache, taking one or two aspirin might relieve the
has a water potential like the water potential of the pain and make us feel better, but taking ten or 12
soil and the water potential of the plant? Typically, aspirin will almost certainly be harmful. Exercise is
the water potential of the air is very low and evapo- good for us but working too much and too hard is
ration occurs readily. However, when humidity is harmful. Our turfgrass has the same qualities that
high, the water potential increases and evaporation we have. Light is good but too much light can be
occurs more slowly. Hence, hot, humid conditions harmful, water is good but too much water is harm-
are difficult for plants because the humidity inhib- ful, and fertilizer is good but too much fertilizer is
its transpiration and the heat causes the internal harmful. We not only have to know what our grass
plant temperature to build more rapidly. needs, we have to know how much is too much.

1.8 Diagnosing Plant Need Single-tactic systems

Why is turf difficult to manage when it is sur- If you are familiar with integrated pest management
rounded by buildings even when it gets sufficient (IPM), you have probably heard of the dangers of

10 Chapter 1
relying on single-tactic systems. IPM refers to a 1.9 Chapter Summary
system of pest control measures that combine for
The objective of this text is to teach the reader
effective control of invading pests. For us, healthy
how to use scientific or logical thought to make
turf is always our number one pest control measure.
technical decisions. In other words, it is designed
If a particular pest problem is severe, we may com-
to demonstrate why management techniques work
bine management practices that improve the health
rather than how they work. The most important
of our turf with mechanical practices, cultural prac-
part of any decision is gathering information. The
tices, biological control techniques and pesticides
more information available, the more likely the
that are designed to specifically manage a particular
manager will be able to make a sound decision.
pest. A single-tactic system would refer to one of
However, we rarely have as much information as
these techniques, usually pesticides, as our only
we would like to have to make important decisions
control measure. Single-tactic systems, even pesti-
and we certainly cannot predict the future.
cides, are ineffective in comparison with IPM. So
Consequently, our knowledge and decision-making
are single-tactic diagnoses. If we are having prob-
procedures must be sound enough to help overcome
lems managing our turf to customer expectations,
our lack of information. To this end, turfgrass
there will nearly always be more than one defi-
management decisions and practices can be
ciency that needs to be addressed. We might blame
improved by the study of the basic scientific
the problem on the soil, on the weed competition,
concepts and plant characteristics introduced in
on a disease problem or on the lack of some nutri-
this chapter.
ent, but it is almost always a combination of prob-
lems that exists. Therefore it is difficult to address
the needs of a particular stand of turf based entirely Suggested Reading
on things that we have done at other times at other
Introductory turfgrass textbooks
locations. We also need to think through the situa-
tion for this particular time and this particular Beard, J.B. (1973) Turfgrass Science and Culture.
place. We need some independent assessment. Prentice-Hall, Englewood Cliffs, New Jersey.
Think photosynthesis, respiration and transpira- Christians, N.E. (2007) Fundamentals of Turfgrass
tion. Are sufficient light, water and carbon dioxide Management, 3rd edn. John Wiley and Sons,
Hoboken, New Jersey.
present? If so, is there sufficient oxygen in the soil?
Duble, R.L. (1989) Southern Turfgrasses: Their Manage-
If we have sufficient water, are the environmental
ment and Use. TexScape, College Station, Texas.
conditions conducive for evaporation to occur Emmons, R.D. (2008) Turfgrass Science and Manage-
rapidly enough to cool our plants? Once we have ment, 4th edn. Thompson Delmar Learning, Clifton
eliminated or minimized barriers to photosynthesis, Park, New York.
respiration and transpiration we can start to Turgeon, A.J. (2008) Turfgrass Management, 8th edn. Pear-
consider other deficiencies. son Prentice Hall, Upper Saddle River, New Jersey.

Diagnosing Plant Need 11

 Understanding Photosynthesis
2

Key Terms
nm is an abbreviation for nanometer, 1 billionth of a meter.
Light is the sun’s radiant energy that we can see and that plants can use for photosynthesis. Ultraviolet and
infrared radiance are not light.
PAR is an abbreviation for photosynthetically active radiation and is the energy in light or radiance.
PPF is an abbreviation for photosynthetic photon flux and refers to the particles or quantum in light or radiance.
Ultraviolet refers to the sun’s radiance below the 400 nm wavelength. These wavelengths have very high energy
and can be destructive to biological tissues.
Infrared refers to the sun’s radiance above the 700 nm wavelength. These wavelengths are relatively low in radiant
energy and very close to the energy level of heat.
Affinity is the force of attraction between objects or particles that causes them to enter into a physical or chemical
combination.
NADPH (the reduced form of nicotinamide adenine dinucleotide phosphate) is a cellular energy molecule in which
the light reaction of photosynthesis stores light energy. It is formed by the donation of electrons to NADP+
(nicotinamide adenine dinucleotide phosphate).
ATP (adenosine triphosphate) is a second cellular energy molecule in which light energy is stored by the light
reaction of photosynthesis. It is formed from ADP (adenosine diphosphate) during the process of photophos-
phorylation.
Thylakoid is an inner compartment of a chloroplast separated from the stroma by a membrane where the light
reaction takes place.
Stroma is the name given to the cytoplasm of the chloroplast.
A sink is a part of a plant, such as a root, that uses more energy than it produces.
A source is a part of a plant, such as a mature leaf, that produces more energy than it uses.
RuBP carboxylase, also called Rubisco is the enzyme that catalyzes the combination of ribulose bisphosphate
(RuBP) and carbon dioxide in the dark reaction of photosynthesis to produce sugars for energy storage and
for plant use in other processes.
A boundary layer refers to a layer of air that surrounds turfgrass leaves that, owing to the occurrence of
photosynthesis and transpiration, is low in carbon dioxide and high in oxygen and water vapor.

2.1 Encourage Photosynthesis

As mentioned earlier, photosynthesis is simply
to Encourage Turfgrass Growth
defined as the process plants use to convert light
Photosynthesis can be an extremely complex energy into chemical energy. This chemical energy
subject. Biologists have been studying photosyn- is then used to power metabolic processes. The
thesis for 100 years or more and have learned energy may be used in the cell where it was pro-
many of its finest details. For our purposes, there duced, it may be transported to another portion of
is no need to go into great detail about this the plant, or it may be transported to a storage
important plant process, but we will study the location. Regardless, if photosynthesis is proceed-
basic concepts so that you can use them to your ing normally at near maximum rate, the plant will
advantage. typically be healthy and capable of resisting stress.

12 ©CAB International 2011. Turfgrass Physiology and Ecology (G. Bell)

As you know, light, water and carbon dioxide are The light spectrum includes all of the colors we
the most important components required for see and ranges from the highest energy light, violet,
photosynthesis. to lowest energy light, red (Fig. 2.2). When all of the
light energy in the light spectrum is combined, we
call it white light, because we cannot differentiate
2.2 The Importance of Light
the colors. Consequently, a perfectly white object is
We know that the sun’s radiance exists in a range reflecting all colors simultaneously. A perfectly black
from at least 50 nm to over 3000 nm in wavelength object absorbs all colors and reflects none. The por-
(NASA/Cool Cosmos EPO Group, 2010). The tion of the light that we see and know as PAR (pho-
term “wavelength” refers to radiant energy’s exist- tosynthetically active radiation) not only differs by
ence in waves of a specific length (Giancoli, 1998) wavelength (400–700 nm), it also differs by color.
(Fig. 2.1). The abbreviation “nm” stands for You will notice in Fig. 2.2 that there is more
nanometer, or 1 billionth of a meter, a very short PAR, measured in watts per meter squared, in the
distance. Not all of the sun’s radiance is light. The blue–green range of the spectrum than any other.
term “light” refers to radiance that we can see. We That might lead you to believe that more blue–
cannot see individual light waves. In fact, we can- green light is used for photosynthesis than any
not see much of the sun’s radiance at all. For pur- other. However, that would not be correct. In fact,
poses of photosynthesis, light occurs in a spectral less green light is used for photosynthesis than
band from the 400 nm wavelength to the 700 nm light from any other part of the PAR (Zscheile and
wavelength and is also called PAR (photosyntheti- Comar, 1941). Peak light absorption by plants is
cally active radiation) or PPF (photosynthetic believed to occur in two wavelength bands, one in
photon flux). Radiance that we can see is slightly the blue range of the spectrum and one in the red
different from what plants use for photosynthesis. range. For that reason, and for simplicity, we nor-
Visible light occurs from about the 400 nm wave- mally divide the plant absorption spectrum into
length to about the 750 nm wavelength (Starr, only three colors: blue, green and red (Fig. 2.3).
2000). We cannot see wavelengths below 380 nm, The sun’s radiance not only exists in wave bands of
called ultraviolet, and although we can see light up energy, it also exists as particles (Giancoli, 1998). We
to around 750 nm, we do not see wavelengths call those particles photons and it is the photons that
above 700 nm very well (Kandel et al., 1991). plants absorb. When a plant absorbs a photon, it also
Radiance above 700 nm is called infrared. If we absorbs the energy in that photon. Consequently,
could see any part of the infrared spectrum as well when a plant absorbs a blue photon, it absorbs more
as we can see green light (500–600 nm), plants energy than it does when it absorbs a red photon. We
would look dark red, not green. That is because call the particle distribution of light the photon flux,
plants reflect substantially more infrared radiance and this is usually measured as micromoles per meter
than they do green light (Chappelle et al., 1992). squared per second. The distribution of photon flux
Because we have such difficulty distinguishing within the spectrum is different than the light energy
light below 400 nm and above 700 nm, many peo- distribution in the spectrum (Fig. 2.4). There is more
ple, including some instructors, refer to the visible green and red photon flux available than there is blue.
spectrum as the same as PAR. Technically, that is So, presumably, under normal conditions, more green
not correct, but it is easy to remember and in a and red photon flux would be absorbed by a plant
practical sense it works. than blue.

Red light
Wavelength Wavelength
650 nm 650 nm

Sun

Fig. 2.1. Radiant energy exists in waves of specific length. Shorter wavelengths contain higher energy than long
wavelengths.

Understanding Photosynthesis 13
 0.8
Visible light
0.7
0.6

Energy (W/m2)
0.5
Yellow
0.4
Orange
0.3 Green
Red
0.2 Blue

0.1 Infrared
Violet
0.0
350 400 450 500 550 600 650 700 750 800 850
Wavelength (nm)

Fig. 2.2. The approximate amount of energy in visible light that strikes the earth’s surface on a clear day in a temperate
climate. Although radiance in the violet band has the highest energy, only a small amount of violet radiance occurs in
sunlight.

2.0
Blue Green Red

1.6
Absorption

1.2

0.8

0.4

0
350 400 450 500 550 600 650 700 750
Wavelength (nm)

Fig. 2.3. Peak absorption of PAR (photosynthetically active radiation) wavelengths by creeping bentgrass (Agrostis stolonifera).
All of the PAR spectrum can be used for photosynthesis, but portions of the blue band and the red band are preferred.

3.5
Visible light
3.0
Photon flux (μ mol/m2/s)

2.5

2.0

1.5 Green Red

Red

1.0 Infrared
Blue

0.5 Violet

0.0
350 400 450 500 550 600 650 700 750 800 850
Wavelength (nm)

Fig. 2.4. Photon flux at solar noon on a sunny day in Columbus, Ohio. Notice that there are more green and red
photons available for photosynthesis than there are blue photons.

14 Chapter 2
 Green light is used for photosynthesis, but chlo- plants high in chlorophyll b. However, all plants that
rophyll, the plant pigment most closely associated contain chlorophyll reflect green light in preference
with energy absorption in plants, accepts blue or to red or blue and, consequently, plants look green
red photons in preference to green. A biochemist or blue–green. Why a plant is green has minor
would say that chlorophyll has a higher affinity for importance, but the concept of affinity is extremely
blue or red photons than it does for green photons. important and very useful (Fig. 2.5).
That does not mean that green photons are not In the demonstration in Fig. 2.5, the enzyme has
absorbed. It means that if blue, green and red pho- twice the affinity for the black molecule as it does
tons are equidistant from a chlorophyll molecule, for the white molecules, but the white molecules
the chlorophyll will bind the blue photons or the are half the distance from the enzyme receptor site.
red photons in preference to the green photons. If the black molecule was closer, the likelihood of
However, if the green photons are considerably the enzyme binding the black molecule would
closer, the chlorophyll will bind the green photons. increase because the black molecule would have
This same type of affinity occurs with enzymes. preference. However, because the distance from the
Many enzymes will bind more than one com- receptor cancels the affinity, all of the molecules
pound. However, the receptor site of an enzyme have an equal opportunity for binding. That means
usually has a strong affinity for only one com- that the probability of a white molecule binding is
pound or may be specific for only one compound. two out of three and the probability of a black
If the enzyme receptor is specific for only one com- molecule binding is one out of three. Remember
pound, then no other compound will bind. If the this concept, it will be useful.
enzyme receptor is not specific for a particular Based on the knowledge of chlorophyll absorp-
compound, then it will have a strong affinity for tion that was just discussed, you should be able to
one compound but will also bind others. If a com- answer this question: if you were managing an
pound with a lesser affinity is near, the enzyme athletic field and you wanted to paint a logo on the
receptor may bind that compound rather than the field that was red, blue or green, which color
one it has the most affinity for. This concept is very would be least likely to damage your turf? Take a
important and we will visit it over and over moment to consider the question.
throughout this text.
Remember that plant processes and all chemical
reactions are dynamic. Molecules are constantly in
motion, and the environment near a chlorophyll
molecule, an enzyme or even a soil particle changes
moment by moment. Consequently, given equal
distributions of red, blue and green photons, red or
blue photons will be bound by chlorophyll most
often. However, if there are substantially more
green photons present than red or blue, the like-
lihood of a chlorophyll molecule binding a green
photon increases.
In the case of chlorophyll and PPF (photosyn-
thetic photon flux), there are more red photons
available than blue and chlorophyll has a higher
affinity for red than green, so red photons are the
most likely to be bound. For that reason, we see
plants as green or blue–green, not red. Red photons
are highly available and highly absorbed. There are
two types of chlorophyll, chlorophyll a and chloro-
Enzyme receptor
phyll b. Some plants have more chlorophyll a than
others, and chlorophyll a has a relatively low affinity Fig. 2.5. This enzyme has twice the affinity for the
for blue photons compared with red (Zscheile and black compound as it does the white compounds, but
Comar, 1941). Consequently, plants high in chloro- the white compounds are half the distance from the
phyll a reflect a larger portion of the blue band than receptor site. Which compound will the enzyme bind?

Understanding Photosynthesis 15
 If the paint reflects green light before it can be smaller amounts of phycoerythrin, phycocyanin
absorbed by the plants, that would be the least and others. All of these pigments are in substan-
damaging to the turf because green is the least tially lower concentrations than chlorophyll. As
likely to be needed for photosynthesis. Red paint already explained, the reflection of green light from
would be most damaging because the plant depends chlorophyll determines the green color of green
on the presence of a substantial number of red plants. When chlorophyll degrades as a result of
photons. Blue would be intermediate. You could stress or seasonal change, turfgrass turns yellow.
answer that question and be reasonably certain The yellow color is reflected by xanthophylls, a
that you were correct simply by applying the specific form of carotenoids. Under normal circum-
knowledge you just learned. That has value. stances, chlorophyll is in a high enough concentra-
tion to mask the color of the xanthophylls. Other
plants, such as trees, reflect yellow, orange or red
The light reaction
light when chlorophyll degrades, depending on
The process of photosynthesis requires two dis- which normally masked pigments are present in
tinctly different pathways. They are called the light greatest concentration. We can encourage the pro-
reaction or z-scheme and the dark reaction or duction of chlorophyll by applying iron (Fe), a
Calvin cycle. The light reaction converts light precursor for the chlorophyll synthesis pathway,
energy into chemical energy and the dark reaction and nitrogen (N), a necessary component of the
stores the energy in sugar molecules for later use chlorophyll molecule and of components of the
according to plant need. You have probably seen a chlorophyll synthesis pathway, and sometimes by
diagram of the z-scheme before. I include it here to applying magnesium (Mg), another important
refresh your memory (Fig. 2.6). component of the chlorophyll molecule (Fig. 2.7).
The purpose of the light reaction is to capture A simple knowledge of biochemistry is sometimes
energy for use in the dark reaction. The initial useful for the turfgrass manager.
receptor of light energy is chlorophyll. Chlorophyll When a light photon combines with a chloro-
is one of many plant pigments. Green plants also phyll molecule, the molecule becomes excited as it
contain significant amounts of carotenoids, and absorbs the light energy of the photon. The amount
of energy absorbed depends on the wavelength of
the photon. As you know, the light reaction
4e– depends on electrons being passed among the com-
ponents of the z-scheme. However, chlorophyll is
PSI NADP+

NADPH Fe
CH2

ADP CH CH3
Cytochrome
O2 5-Amino levulinate
b6–f
4e – CH3 CH2 CH3
ATP
Porphobilinogen N N
M
Mgg
4H+ Uroporphyrinogen III N N

CH3 CH3
PSII Protoporphyrin IX
CH2
O
Mg-protoporphyrin IX CH2 CO O CH3
2H2O monomethyl ester
CO

Fig. 2.6. A basic diagram of the light reaction or Protochlorophyllide O

z-scheme. The purpose of the light reaction is to

capture energy for use in the dark reaction. ADP = Chlorophyllide Chlorophyll a

adenosine diphosphate; ATP = adenosine triphosphate;

NADP+ = nicotinamide adenine dinucleotide Fig. 2.7. Iron (Fe) is an important precursor for
phosphate+; NADPH = reduced form of nicotinamide the chlorophyll synthesis pathway shown here, and
adenine dinucleotide phosphate; PSI = photosystem I; nitrogen (N) and magnesium (Mg) are important
PSII = photosystem II. components.

16 Chapter 2
not the component that donates electrons, it is sim- fluorescence, both forms of radiance, but it is most
ply an energy carrier (Voet and Voet, 1990). The commonly lost as heat (Salisbury and Ross, 1992).
energy of the photon causes the chlorophyll to Hence, transpiration is required to cool the plant.
enter a highly excited state. In this state, the chlo- Photosynthesis can only proceed temporarily
rophyll becomes unstable and will pass the energy unless transpiration occurs simultaneously as the
on to another chlorophyll molecule or to other process of photosynthesis produces too much heat.
compounds that it touches. Hence, the light energy The plant has to be cooled. Otherwise, it will, in
is passed among chlorophyll molecules until it is simple terms, burn itself up. Turfgrasses, even
accepted by one of the reaction centers, PSI (photo- warm-season species, yellow during the summer as
system I) or PSII (photosystem II), in the z-scheme a matter of seasonal acclimation (Xiong et al.,
(Fig. 2.8). 2007). This yellowing is caused by high photosyn-
A photon of blue light may have energy equiva- thesis and high energy levels among the chlorophyll
lent to that contained in a 450 nm wavelength. molecules resulting in chlorophyll degradation
A photon of red light will have considerably less (Demmig-Adams, 1990). The long day lengths and
energy such as the energy equivalent to that of a intense light common to summer reduce the need
650 nm wavelength (Fig. 2.2). The reaction centers, for chlorophyll and increase the need for caroten-
PSI and PSII, cannot accept energy of that magni- oid pigments that help to quench excess energy
tude. PSI accepts energy equivalent to approxi- (Adams and Demmig-Adams, 1992). During sum-
mately 700 nm and PSII accepts energy equivalent mer, grasses need less chlorophyll to absorb light
to 680 nm. Consequently, the blue light and red energy because light is available at high intensity
light in this example are too powerful for the reac- and for long periods. However, the plants need
tion centers to accept. The light energy absorbed by more carotenoid pigments to quench excess light
the chlorophyll cannot be absorbed by one of the energy when the photosynthesis pathway becomes
reaction centers until the energy degrades to an overloaded. Consequently, your turf turns yellow–
acceptable magnitude. When a chlorophyll mole- green instead of the dark-to-medium green color
cule absorbs a photon the energy degradation proc- that occurs in the spring and usually in the fall. It
ess begins immediately (Salisbury and Ross, 1992). also becomes resistant to fertilization. During sum-
Every time the energy passes from one chlorophyll mer, less fertilizer is needed to encourage plant
molecule to another, a portion of the energy is lost. metabolism. Because of the high heat of summer
This energy can be lost as phosphorescence or that dramatically increases the rate of chemical
reactions, and because light and day length are
driving photosynthesis to optimum and beyond,
Chlorophyll
little fertilizer is needed (Xiong et al., 2007).
Chlorophyll

Ch Sun In summer, light is optimal, and except in shade,

ll
hy

lor
op
op

hy Ch light is present in greater amounts than turfgrass

or

ll lor
hl
ll

op
hy

hy requires. However, water for transpiration becomes

op

ll
or

Chlorophyll Chlorophyll increasingly important because of the high heat and

hl

Ch ll
C

lor hy
op
hy rop
Chlorophyll

Ch
ll Ch
lo the high rate of activity in the plant. High
lor Chlorophyll
op Chlorophyll transpiration rates are common and required for
Chlorophyll

hy
ll
plant cooling. Carbon dioxide is also needed in
ll

l
hyl
hy

op Reaction
larger quantities during the summer because of the
op

lor
or

Chlorophyll Ch Center
ll
hl

hy

high rate of activity, especially the increases in

C

op

PSI or PSII Ch
or
Chlorophyll

lor
Chlorophyll

hl

op photosynthesis that accompany increases in light

ll

hy
phy

ll
Ch
intensity and day length (Adams and Demmig-
loro

lor
op
Chlorophyll

hy
Ch

ll Chlorophyll Adams, 1992).

y ll
ph

Ch
lor
ro

op
o
hl

hy
C

ll
2.3 The Importance of Carbon Dioxide
Fig. 2.8. Light energy from the sun is absorbed by As mentioned earlier, the light reaction provides the
chlorophyll and passed among chlorophyll molecules energy that fuels the dark reaction or Calvin cycle.
until it is absorbed by a reaction center, either PSI As electrons pass through the z-scheme, energy rich
(photosystem I) or PSII (photosystem II). molecules called NADPH (the reduced form of

Understanding Photosynthesis 17
nicotinamide adenine dinucleotide phosphate) called the oxygen-evolving complex (Fig. 2.9). This
and ATP (adenosine triphosphate) are produced splitting of water is what causes oxygen to be
(Fig. 2.6). The NADPH is produced when NADP+ released during photosynthesis. When two water
(nicotinamide adenine dinucleotide phosphate) molecules are split, four electrons (e−) are released
is reduced by an electron from the z-scheme. along with four protons (H+) and one oxygen (O2)
A chemical reduction is the name given to a molecule. This activity helps to make the inside of
chemical reaction when an electron is accepted by the thylakoid positively charged compared with the
a molecule. A chemical oxidation occurs when an outside. Reactions that occur in the z-scheme also
electron is released from a molecule. Molecules of drive a proton-producing process by chemical
ATP are formed from ADP (adenosine diphos- reaction. Each time four electrons pass through the
phate) by a proton pump that is activated when z-scheme eight protons are transferred from
electrons flow through the z-scheme. The energy in the stroma to the thylakoid, thus further increasing
these molecules is used to convert carbon dioxide the difference in polarity between the inside and
to sugar in the dark reaction. outside. Because of this electrical gradient, protons
The actions of proton pumps and similar mecha- are attracted to the stroma and negative solutes are
nisms that work across cellular membranes, attracted to the thylakoid.
organelles and internal membranes are very impor- Because the inside of the thylakoid is relatively
tant concepts in biology and deserve consideration. positive and the outside is relatively negative, a
The light reaction of photosynthesis occurs in the hole in the membrane would be all that is needed
wall of a membrane inside the chloroplast. This to provide the passage of positive solutes out and
membrane is called the thylakoid membrane and it negative solutes in. However, as a hole in the mem-
separates the thylakoid, an inner compartment of brane would destroy the cell, special carrier pro-
the chloroplast, from the stroma. The stroma is the teins are employed to move protons from the inside
name given to the cytoplasm of the chloroplast. to the outside. This movement provides the energy
The electrons that pass through the z-scheme inside necessary to synthesize ATP from ADP. All proton
the thylakoid membrane are obtained from water. pumps work on this principle and are important
Water inside the thylakoid is split by a complex for various functions in plant metabolism. As you

Stroma
H+
+ NADP+ NADPH
H
ADP ATP
+
H
4e–
Cytochrome
b6–f H+
+
H

PSΙ H+ PSΙΙ H+
4e–
H+
8H+
4H+
H+ H+
H+
2H2O
O2 Membrane
Thylakoid

Fig. 2.9. The oxygen-evolving complex. The z-scheme in the light reaction of photosynthesis produces ATP
(adenosine triphosphate) by the action of a proton pump. The splitting of water and the passage of electrons along
the z-scheme cause an electrical difference across the thylakoid membrane. The electrical difference, which is
positive inside, negative outside, provides the energy that fuels the proton pump. ADP = adenosine diphosphate;
ATP = adenosine triphosphate; NADP+ = nicotinamide adenine dinucleotide phosphate; NADPH = reduced form of
nicotinamide adenine dinucleotide phosphate; PSI = photosystem I; PSII = photosystem II.

18 Chapter 2
will learn, turf maintenance practices or environ- sugars. Many of these sugars will be converted to
mental conditions that affect the passage of solutes sucrose for transport to plant parts that need
through membranes or through the proton pumps energy. As you probably know, we call those plant
in membranes can drastically affect plant health. If parts that require energy “sinks” and those plant
the proton pumps are damaged or holes occur in parts that produce energy “sources”. Other sucrose
the thylakoid membrane, photosynthesis no longer molecules will be used to form starch or fructans
works. When all is working properly, each time for long-term energy storage. This energy storage
three protons pass through the pump their passage occurs primarily in chloroplasts, in roots, or in
provides sufficient energy to convert one molecule plant stems. Turfgrasses have three types of stems.
of ADP to one of ATP. The stem that holds the reproductive parts – the
flowers and seeds, stems called rhizomes and stems
called stolons. We rarely see the seed stalks because
The dark reaction
most turfgrasses will not produce flowers or seeds
The dark reaction of photosynthesis is a complex at the height at which we mow. In addition, some
process that provides the plant with a number of grasses do not produce rhizomes or stolons, and
three-, five-, six- and seven-carbon sugars that are store their energy mostly in roots. All of these
used for various functions (Fig. 2.10). The dark stems are produced by the turfgrass crown, and
reaction provides the chemical bonding of carbon you should recall from introductory turfgrass
molecules that store or use the energy sequestered classes that the crown is the source of all turfgrass
in the light reaction. Carbon dioxide provides the growth. We will study these processes further in
carbon that the dark reaction needs to produce Chapter 4.

ADP
NADPH
ATP +
BPG NADP

3PGA 1,3-Bisphoglycerate
CO2
3-Phosphoglycerate Gylceraldehyde-3- GAP Storage sugars
phosphate
ADP Sugar
RuBP
ATP Ribulose-1, 5-
Dihydroxyacetone
bisphosphate
phosphate
DHAP
Ribulose-5-
bisphosphate

Ru5BP
Fructose-1,6-
biphosphate
FPB

Ribulose-5-
phosphate Fructose-6-
phosphate
Ru5P
F6P
Xyulose-5-
R5P phosphate E4P
Xu5P Erythrose-4-
Ribose-5-
phosphate
phosphate

SBP
S7P Sedoheptulose-1,7-
bisphosphate
Sedoheptulose-7-
phosphate

Fig. 2.10. The dark reaction of photosynthesis or Calvin cycle is a complex process that requires CO2 and both ATP
(adenosine triphosphate) and NADPH (the reduced form of nicotinamide adenine dinucleotide phosphate) from the light
reaction. The ATP and NADPH are converted back to ADP (adenosine diphosphate) and NADP+ (nicotinamide adenine
dinucleotide phosphate), and the energy so released is used to drive the Calvin cycle.

Understanding Photosynthesis 19
 In this chapter, we are primarily concerned with vapor and low in carbon dioxide is called a bound-
the processes that sequester energy for future plant ary layer (Turgeon, 2008). There is always a
use. The production of energy storage compounds boundary layer surrounding plant leaves, and
and sugars for use in other processes requires the because this layer is high in oxygen and low in
combination of carbon dioxide (CO2) with ribulose carbon dioxide, it deters photosynthesis. Because it
bisphosphate (RuBP), a 5-carbon sugar. This com- is also high in water vapor, it slows transpiration.
bination is catalyzed by an enzyme called RuBP Consequently, poor air movement reduces two of
carboxylase, commonly called Rubisco, and the three most important plant functions, photo-
believed to be the most pervasive enzyme in exist- synthesis and transpiration.
ence. Needless to say, Rubisco is important. A thin boundary layer is normal and not particu-
The first product of the dark reaction of photosyn- larly detrimental. A thick boundary layer, however,
thesis is an unstable 6-carbon sugar that immediately can slow or even halt photosynthesis entirely.
breaks down into two 3-carbon sugar molecules When this occurs, the lack of carbon dioxide has
called PGA (phosphoglycerate). Although PGA is become so limiting that photosynthesis will no
produced each time carbon dioxide combines with longer occur in spite of otherwise conducive envi-
RuBP, not all of the PGA produced is used to seques- ronmental conditions. In this case, you have failed
ter energy. As RuBP has to be preserved so that the as a turfgrass manager. It is imperative that you
cycle can continue and because the cycle also pro- provide enough air movement across your turf to
duces sugars that are needed to synthesize tissue and allow adequate photosynthesis to occur.
other compounds, it actually takes three cycles to Scientists have sophisticated instruments that
produce one PGA or six cycles to produce one can be used to estimate photosynthesis in the
6-carbon sugar such as glucose. The production of field. However, it is difficult for us to determine
two PGA molecules requires the energy from 18 ATP how much photosynthesis is enough under a
molecules and 12 NADPH molecules that came from given situation and changing environmental con-
the light reaction (Voet and Voet, 1990). The light ditions. Unless technological advances occur,
reaction may be producing these energy molecules at turfgrass managers will not have the equipment
a very high rate and all of the constituents of the to measure photosynthesis anytime soon. If you
pathway may be available, but the dark reaction suspect that air movement across your turfgrass is
does not occur unless carbon dioxide is present. not sufficient to perform adequate photosynthe-
sis, it probably isn’t. Your turfgrass will tell you.
If carbon dioxide is deficient the turf will yellow
Maintaining the presence of carbon dioxide
more quickly than normal when day lengths
The presence of carbon dioxide should be extend beyond optimum. The turf will be weak.
paramount in any high-quality turfgrass mainte- It will be easily damaged by traffic, disease,
nance plan. Light may be available, water may be insects or any other stress. Once damaged, it will
available, but if carbon dioxide is not available, be very slow to recover. Root mass and root
photosynthesis does not occur. Carbon dioxide is depth will be poor. Density will suffer as there is
often the limiting factor for photosynthesis. not enough plant energy available to produce
new tillers and/or daughter plants. These plant
responses are typical of turfgrass stress caused by
Boundary layers
a number of different conditions. However, in
Carbon dioxide is most often deficient in areas this case, the symptoms will be pervasive, exten-
where air movement across the turf is restricted. sive and contiguous. Growing conditions nor-
The air is only 0.035% carbon dioxide. Therefore, mally conducive to plant health will have little
during daylight when sufficient water is present, a effect. Other areas of turfgrass managed similarly
dense turf can use up the carbon dioxide in stag- will respond well to management, but the same
nant air rather quickly. As photosynthesis proceeds, management will have no effect on the air-
the carbon dioxide in this stagnant air is replaced restricted turfgrass. Planting a different species,
with oxygen. As the plant transpires to cool itself applying more or less water, more or less fertilizer
from the heat of photosynthesis, the stagnant air or more pesticides is not the answer. In this case,
also becomes high in water vapor. The air sur- as in most, you have to fix the problem, not just
rounding leaves that is high in oxygen and water treat the symptoms.

20 Chapter 2
 depends on the climate and the situation. However,
Improving air movement
a major improvement in air movement at the site
Restricted air movement is usually caused by sur- is likely to result in substantially healthier looking
rounding vegetation, buildings or problems with turf. If not, there may be additional negative fac-
the contour of the land in the immediate vicinity. tors to consider.
Surrounding vegetation is the easiest to deal with. Buildings can be a real problem to deal with.
All it takes is removal. You don’t necessarily have Courtyards and stadiums present unique problems
to remove trees, only low-growing vegetation. to air movement over turfgrass. In this case, man-
Removing low vegetation and debris does not usu- made structures surround the turf. Sometimes the
ally meet with a great deal of customer resistance. solution is as simple as opening or removing doors
In fact, in many cases it makes the site look better. on opposite ends of the courtyard or stadium.
All low-growing vegetation and debris should be Opening doors or windows to the outside may not
removed. Fences and hedges are particularly restric- solve an air restriction problem completely, but it
tive. It is most important to remove vegetation and will certainly help. In some instances open passage-
debris facing the predominant winds and on the ways to the outside can be constructed fairly easily,
leeward side of the site. It usually does not do any but in most cases a major reconstruction is required.
good to remove restrictions on only one side. In that case it depends on the value of the turf, and
Restrictions on both sides should be removed to sometimes on your salesmanship, to convince the
allow air to flow across the site. Clearing only one owners of the importance of new passages. Fans are
side of the site is similar to opening a door on one a last resort. Fans are expensive and require electri-
side of a house. Opening a single door has a minor cal installation. They are noisy and need constant
effect on ventilation but opening doors or windows upkeep and repair. However, if the turf is of signifi-
on opposite sides of the house allows air to circu- cant value, a fan or multiple fans will keep it alive
late freely into one side of the house and out of the and make it easier to manage and maintain at an
other. If possible, remove low-growing vegetation acceptable level.
and debris on all sides of the site. There are times when the contour of the land
Although trees do not necessarily need to be does not allow air movement across your turf.
removed to improve air circulation, low-hanging I have, in fact, seen golf courses with putting
branches are restrictive. Remove all branches greens designed for placement in such a situation.
within at least 10 feet (3.2 m) of the ground. It simply does not work to place a high-value stand
Higher removal is better but more difficult and not of turf that is required to be maintained at near-
often necessary. Coniferous trees are usually more perfect condition in the bottom of a hole. If it can
restrictive than deciduous trees and more likely to be maintained at an acceptable level, the man-
require additional trimming. A small, inexpensive power and materials required are not worth the
anemometer (wind gauge) is useful during and effort. Fortunately there are few situations where
after this operation. Use the anemometer to meas- land contour restricts air movement severely
ure wind speed just above the turf at the wind- enough to influence turfgrass growth. In those few
restricted site and compare that measurement with instances where improving air movement is
wind speed at an unrestricted site immediately required, the only good option is to regrade the
before or after measuring the restricted site. site. Fans can also be useful. In most situations, the
Complete this operation a few different times each turfgrass can be managed, but it will always be
day for several days. Record the measurements weak and require more than normal care. If you
each time they are made and compare the average learn your lessons well and take advantage of as
wind speed at the restricted site with the aver- much practical experience as possible, you will be
age wind speed at the unrestricted site. The differ- able to handle it.
ence may be substantial. A 50% decline is not
unlikely. Continue to make these measurements as
2.4 The Importance of Water
you clear the restricted site and after you think
that you have improved the site sufficiently. There Water is important for all plant functions.
is no hard, fast number short of 0.0% decline in Turfgrass leaves, where most of the photosynthe-
air movement that is sure to be effective. As in sis of the plant occurs, are mostly water. Dry clip-
most situations involving turfgrass management, it pings of turfgrass leaves, for instance, weigh

Understanding Photosynthesis 21
considerably less than fresh clippings. Not only is absorbs PPF (photosynthetic photon flux). When
water necessary to provide the medium for all chlorophyll absorbs a photon, it also absorbs its
chemical reactions in photosynthesis and to cool energy. Blue photons have high energy, red photons
the plant because of the heat produced in photo- have low energy and green photons have interme-
synthesis, it also has to provide the electrons for diate energy. Nearly all of the photons absorbed by
the light reaction of photosynthesis. Too little chlorophyll have energy that is too high for use in
water can also cause indirect consequences for the light reaction. Therefore, this energy has to
photosynthesis. As you will learn later in the text, degrade to a level equivalent to either 680 nm for
dry conditions cause plant stomates to close, thus use in the PSII (photosystem II) pathway or 700 nm
limiting the amount of carbon dioxide that enters for use in the PSI (photosystem I) pathway. As this
the leaf and severely reducing photosynthesis. The energy degrades it gives off heat. That is why tran-
importance of water will be a recurring theme spiration is so important for photosynthesis. The
throughout this text. plant must be protected from the heat produced
during this release of energy.
Once energy is absorbed by the reaction centers,
2.5 Chapter Summary
PSI and PSII, the z-scheme is activated and the light
Photosynthesis is necessary for plant survival and reaction begins. Water is split by the oxygen-
for the survival of all living things on earth. Like all evolving complex, producing four electrons, four
biological physiology, it is an extremely complex protons, and one oxygen molecule for every two
process of molecular activity. The concept, how- water molecules. The electrons enter the z-scheme
ever, is relatively simple. The process of photosyn- and carry the energy through the pathway. The
thesis is the physiological pathway that plants use protons enter the thylakoid where they and addi-
to transform light energy into chemical energy. tional protons produced by the passage of electrons
This energy may be used immediately, it may be through the z-scheme cause a difference in polarity
transported to other plant parts to be used as across the thylakoid membrane. Because of the dif-
energy or converted to important plant products, ference in polarity between the inside of the thyla-
or it may be transferred to a storage area and koid and the outside protons move across the
sequestered for later use. thylakoid membrane through a proton pump and
There are two major pathways that constitute provide the energy necessary to produce ATP
photosynthesis, the light reaction or z-scheme and (adenosine triphosphate) from ADP (adenosine
the dark reaction or Calvin cycle. Light and water diphosphate). The ATP and NADPH (the reduced
are the major components required to fuel the form of nicotinamide adenine nucleotide phos-
light reaction. Light provides the energy that is phate) are the two energy molecules formed during
sequestered by the plant during the process and the light reaction. NADPH is produced from the
water provides the electrons that carry the energy reduction of NADP+ (nicotinamide adenine nucle-
through the z-scheme. Water also provides a otide phosphate) by an electron from the
medium for chemical reactions to occur and is z-scheme.
consumed by the transpiration process that cools The ATP and NADPH are used to provide the
the plant of the excessive heat produced by the energy to fuel the dark reaction or Calvin cycle.
light reaction. Light is not required in the dark reaction but car-
Light called PAR (photosynthetically active radi- bon dioxide must be present. During the dark
ation) is the source of energy for photosynthesis. reaction, RuBP carboxylase, an enzyme more com-
PAR occurs in a spectral band from 400 nm to monly called Rubisco, catalyzes the combination
700 nm wavelengths and is divided into three of carbon dioxide with RuBP (ribulose bisphos-
major colors, blue, 400 to 500 nm, green 500 to phate), a 5-carbon sugar. Each time one carbon
600 nm, and red 600 to 700 nm. All three colors are dioxide is bound, one more carbon enters the
active for photosynthesis, but red and blue are cycle. This carbon accumulation results in the syn-
preferred. thesis of many different sugars for a variety of
PAR not only contains energy but also occurs in plant uses.
particles called photons. These photons are the The combination of RuBP and carbon dioxide
particles actually absorbed by chlorophyll. So results in 3-carbon sugar molecules called PGA
chlorophyll does not actually absorb PAR, it (phosphoglycerate). As the Calvin cycle proceeds,

22 Chapter 2
some PGA is used to synthesize glucose and As turfgrass managers, one of our most impor-
fructose, which can be combined to make tant functions is to make sure that the light, water
sucrose. The sucrose is transported throughout and carbon dioxide necessary for healthy photo-
the plant as an energy source or combined into synthesis are present in our turfgrass environment.
long chains of starch or fructans for energy stor- If this essential process cannot be performed at a
age and later use. high level on a near constant basis, our turf will
Carbon dioxide is imperative for the perform- always be weak or nonexistent.
ance of the Calvin cycle. It is up to the turfgrass
manager to make certain that a lack of carbon
dioxide does not interfere with photosynthesis. Suggested Reading
Carbon dioxide becomes limiting when a boundary Kelly, G.J. and Latzko, E. (2006) Thirty Years of Photo-
layer high in oxygen and water vapor and low in synthesis. Springer, New York.
carbon dioxide builds up around the turf. Managing Starr, C., Evers, C.A. and Starr, L. (2006) Biology: Con-
for good air movement across the turf to refresh cepts and Applications, 6th edn. Thomas, Brooks/
the air and bring in new carbon dioxide is a neces- Cole, Belmont, California.
sary practice in some situations. Low-growing tree Stern, K.R., Bidlack, J. and Jansky, S. (2008) Introduc-
tory Plant Biology, 11th edn. McGraw-Hill Higher
limbs and other vegetation and debris may need to
Education, New York.
be removed from a site to enhance air movement. Voet, D.V. and Voet, J.G. (2004) Biochemistry, 3rd edn.
Sometimes regrading of the site is necessary. In John Wiley and Sons, New York.
courtyards or stadiums air channels from the inside
to the outside may need to be constructed. There is
always a thin boundary layer around plant leaves Suggested Websites
as they perform photosynthesis, but when the NASA/Cool Cosmos EPO (Education and Public
boundary layer gets too thick, photosynthesis is Outreach) (2010) Infrared Astronomy: Near, Mid and
affected and steps must be taken to create fresh air Far Infrared. Available at: http://coolcosmos.ipac.
circulation. Otherwise, the turfgrass may become caltech.edu/cosmic_classroom/ir_tutorial/irregions.
so weak that it can not be managed. html (accessed 27 July 2010).

Understanding Photosynthesis 23
 Why C3 and C4 Grasses Require
3 Different Management

Key Terms
A warm-season grass is a grass that uses the C4 photosynthesis pathway.
A cool-season grass is a grass that uses the C3 photosynthesis pathway.
Photorespiration occurs when Rubisco, the enzyme that binds carbon dioxide in the Calvin cycle, binds oxygen
instead.
Mitochondria are organelles where oxidative respiration takes place in both plants and animals. Mitochondria also
function in photorespiration in C3 plants.
A peroxisome is a plant organelle that, among other things, is part of the photorespiration pathway.
The light saturation point is the amount of light present when photosynthesis is occurring as rapidly as
possible.
Vapor pressure is the pressure exerted by a vapor in a closed container that is in dynamic equilibrium with its
liquid form.
Dynamic equilibrium is a state of balance in which opposing processes occur at the same rate at the same time
so that it appears as if nothing is happening. In fact, a great deal is happening but there is no net effect.
Kranz anatomy is the name given to the special anatomy of a C4 plant that allows the light reaction of
photosynthesis and the dark reaction of photosynthesis to take place in two separate cells.
A bundle sheath cell is a small cell located near the vascular system of grasses where the Calvin cycle takes
place in C4 plants. The bundle sheath cell is the basic component of Kranz anatomy.

3.1 Use Species Adaptation to Your

is more efficient than C3 photosynthesis. In each
Advantage
case, I was living in a temperate climate at the time.
The C4 (warm-season) pathway of photosynthesis Consequently, I asked them both to tell me why
is rare in plant species in general but quite common there weren’t any perennial C4 grasses growing
in turfgrasses (Matsuoka et al., 2001). The grasses outside and they both explained that most C4
that we call warm-season, the C4 plants, and the grasses could not survive cold temperatures. Then
grasses that we call cool-season, the C3 plants, I asked why, if the C4 pathway is more efficient,
perform photosynthesis differently. They also differ don’t cold-adapted plants use it as well as warm-
in their adaptation to climatic conditions. Whether adapted plants? Neither scientist had an explana-
warm-season grasses adapted to the climate that C4 tion for that. Both of those scientists know a lot
photosynthesis was best suited for, or the grasses more about photosynthesis and other plant func-
adopted C4 photosynthesis because it was more tions than I do, but until they can answer that ques-
efficient for their climate is unknown. However, we tion for me, I will continue to believe that C3
know for certain that C3 and C4 grasses require photosynthesis is more efficient in cool tempera-
different management procedures because of tures and C4 photosynthesis is more efficient in
the differences in photosynthesis and climatic warm temperatures. If you are a homeowner in a
adaptation between them. temperate climate, you will probably want to know
I have been told by at least one biochemist and how to manage cool-season grasses. If you are a
at least one plant physiologist that C4 photosynthesis homeowner in a tropical climate you will want to

24 ©CAB International 2011. Turfgrass Physiology and Ecology (G. Bell)

Table 3.1. Some of the most common cool- and warm-season grasses used for turf throughout the world. Common
names are those used in the USA. This list does not include all grasses used for turfgrass.

Common name Scientific name

Cool-season (C3) grasses

Velvet bentgrass Agrostis canina L.
Colonial bentgrass Agrostis capillaris L.
Creeping bentgrass Agrostis stolonifera L.
Tall fescue Festuca arundinacea Schreb.; synonym, Schedonorus phoenix (Scop.)
Holub.
Fine fescues
Sheep fescue Festuca ovina L.
Creeping red fescue Festuca rubra L.
Chewing’s fescue Festuca rubra L. ssp. fallax (Thuill.) Nyman; synonym Festuca rubra L. ssp.
commutata Gaudin
Hard fescue Festuca trachyphylla (Hackel) Krajina; synonym, Festuca brevipila R. Tracey
Annual ryegrass Lolium multiflorum Lam.
Perennial ryegrass Lolium perenne L.
Alpine bluegrass Poa alpina L.
Annual bluegrass Poa annua L.
Texas bluegrass Poa arachnifera Torr.
Bulbous bluegrass Poa bulbosa L.
Canada bluegrass Poa compressa L.
Kentucky bluegrass Poa pratensis L.
Supina bluegrass Poa supina L.
Roughstalk bluegrass Poa trivialis L.
Warm-season (C4) grasses
Common carpetgrass Axonopus affinis Chase
Tropical carpetgrass Axonopus compressus (Swartz) Beauv.
Buffalograss Buchloe dactyloides (Nutt.) Engelm.; synonym, Bouteloua dactyloides (Nutt.)
Columbus
Bermudagrass Cynodon dactylon (L.) Pers.
African bermudagrass Cynodon transvaalensis Burtt-Davy
Centipedegrass Eremochloa ophiuroides (Munro) Hack.
Bahiagrass Paspalum notatum Flugge
Seashore paspalum Paspalum vaginatum Swartz
Kikuyagrass Pennisetum clandestinum Hochst ex Chiov.
St. Augustinegrass Stenotaphrum secundatum S. (Walt.) Kuntze
Zoysiagrasses
Japanese lawngrass Zoysia japonica Steud.
Manilagrass Zoysia matrella (L.) Merr.
Mascarenegrass Zoysia tenuifolia Willd. ex Thiele; synonym, Zoysia pacifica Willd.

know how to manage warm-season grasses. If you reactions of photosynthesis, also called the Calvin
are a turfgrass manager, you need to know how to Cycle. Rubisco is more correctly called ribulose
manage both types of grasses and how to manage bisphosphate carboxylase oxygenase because it
them both well (Table 3.1). not only binds carbon dioxide, it binds oxygen.
When Rubisco binds oxygen instead of carbon
dioxide, the resulting process is called photore-
3.2 Photorespiration
spiration – an inefficient, energy-consuming
In Chapter 2, you learned about Rubisco, techni- process. Considering that our air is 21% oxygen
cally called ribulose bisphosphate carboxylase, the but only 0.035% carbon dioxide, Rubisco would
enzyme that binds carbon dioxide in the dark be considerably more likely to bind oxygen than

C3 and C4 Grasses Management 25

carbon dioxide, except that it has a much higher bisphosphate), a combination resulting in an unstable
affinity for carbon dioxide than it does for oxygen 6-carbon molecule that immediately splits into two
(Fig. 3.1). Enzyme affinity is a concept that you 3-carbon molecules of PGA (phosphoglycerate).
also learned about in Chapter 2 (Fig. 2.5). Rubisco’s When Rubisco binds oxygen instead of carbon diox-
high affinity for carbon dioxide overcomes the ide, a different result occurs. The combination of
amount and proximity of oxygen and increases the RuBP and oxygen results in one molecule of PGA
likelihood that carbon dioxide will be bound in and one molecule of PGL (phosphoglycolate), a
preference to oxygen. However, sometimes, envi- 2-carbon molecule:
ronmental conditions – mainly intense light and
Carbon assimilation = RuBP +CO2
high temperature – increase the likelihood for oxy-
gen to be bound by Rubisco and for photorespira- ⎯⎯⎯⎯
Rubisco
→ 2PGA
tion to occur. Oxygen assimilation = RuBP +O2
Only C3 plants photorespire, and the likelihood
⎯⎯⎯⎯
Rubisco
→ PGA + PGL
for photorespiration to occur increases with increas-
ing temperature and usually with increasing light. The phosphoglycolate that results from the
Although C4 plants have the pathways and enzymes combination of RuBP and oxygen is not useless
to accommodate photorespiration, it does not occur but it must be processed through a complicated
in a measurable amount, and we assume that it is pathway, after which it is converted to PGA and
nonexistent in C4 plants. In order to understand why enters the Calvin cycle. This is actually a respira-
photorespiration occurs and how C4 plants avoid it, tory process, hence the name photorespiration,
you need to learn a little about the process. and except for the end product, PGA, it has little
Photorespiration is an inefficient form of plant to do with photosynthesis. The process requires
respiration that occurs when Rubisco binds oxygen energy and results in the loss of one carbon diox-
instead of carbon dioxide. As you know, carbon ide molecule. To salvage photosynthetic carbon
assimilation has to occur during photosynthesis or from glycolate, photorespiration combines two
there is no conversion of light energy to chemical glycolates to produce one PGA and one carbon
energy. When the Calvin cycle is working properly, dioxide. The PGA enters the Calvin cycle and the
Rubisco binds carbon dioxide and RuBP (ribulose carbon dioxide is lost to the atmosphere.
Consequently, two combinations of RuBP and
oxygen result in three PGAs and the loss of one
carbon. If Rubisco had bound carbon dioxide
O2 CO2 twice, the result would be four PGAs for the
O2
Calvin cycle, with no loss of carbon and no need
for additional energy from ATP (adenosine tri-
O2 O2
phosphate) to recover carbon otherwise lost as
O2
PGL. Therefore, you could call photorespiration a
destructive process.
High O2 O2
The photorespiratory process occurs in three
affinity different organelles, the chloroplast, the
mitochondrion and the peroxisome (Fig. 3.2).
Low Phosphoglycolate is transported from a chloroplast
O2
affinity to a peroxisome, then to a mitochondrion (via
glycine), back to a peroxisome (via serine) and,
finally, back to a chloroplast (via glycerate), where it
enters the Calvin cycle as PGA. During photorespira-
Rubisco tion then, two PGLs are converted to one PGA, with
an expenditure of energy, and one carbon dioxide is
Fig. 3.1. Under normal environmental conditions,
considerably more oxygen (O2) than carbon dioxide
lost. Because less carbon is accumulated and energy
(CO2) is in close proximity to the enzyme Rubisco is actually used when photorespiration occurs, the
(ribulose bisphosphate carboxylase), but Rubisco has plant may have to use stored energy reserves for
such a strong affinity for CO2 that it is more likely to metabolic processes, even though conditions are suit-
bind CO2 than to bind O2. able for photosynthesis.

26 Chapter 3
 Peroxisome

PGL

Glycerate Glycine

Serine
PGL
RuBP
O
O22
Rubisco PGA Glucose
Calvin Cycle
Calvin cycle
Glycerate

PGA

Glycine
Chloroplast
Serine

Mitochondrion

Fig. 3.2. Photorespiration is a complicated process that requires phosphoglycolate (PGL) to be transferred from
a chloroplast to a peroxisome, then to a mitochondrion (via glycine), back to a peroxisome (via serine) and finally,
back to a chloroplast (via glycerate), where it enters the Calvin cycle as phosphoglycerate (PGA). RuBP = ribulose
bisphosphate; Rubisco = ribulose bisphosphate carboxylase.

Light affects photorespiration These conditions do not occur during the winter or
during the spring or fall. Under natural conditions,
Photorespiration is most common under conditions
the formation of boundary layers due to rapid photo-
of high light and high heat. When light is intense, the
synthesis can only occur during the summer. Summer
light reactions of photosynthesis occur rapidly and
is when sunlight is most intense and day length is the
oxygen is released in high concentrations. Because
longest. Consequently, if you are managing cool-sea-
carbon dioxide is being assimilated rapidly, the air in
son grasses, you must prepare them by doing the right
and around our plants becomes high in oxygen and
things in the fall, winter and spring so that they can
low in carbon dioxide. These are perfect conditions
survive the next summer. Summer is a very stressful
for photorespiration. Warm-season grasses have a
period for cool-season grasses (Liu and Huang,
definite advantage over cool-season grasses under
2001). If your plants are not healthy in late spring,
such conditions because they do not photorespire
they will not survive the summer.
and because they can assimilate carbon dioxide even
when its concentration in the surrounding air is very
Light saturation
low (Krenzer and Moss, 1969).
In Chapter 2 we called this high oxygen low carbon You may be familiar with the “light saturation
dioxide condition a boundary layer and discussed it point”. The light saturation point of a particular
as a barrier to photosynthesis where air movement plant is the amount of light present when
was restricted. In this case, the boundary layer is not photosynthesis is occurring as rapidly as possible. An
a result of restricted air movement but of rapid increase in light beyond the light saturation point will
photosynthesis. It follows that when the air holds not result in an increase in photosynthesis. Light
higher than normal oxygen and lower than normal saturation points are affected by a number of factors,
carbon dioxide Rubisco is more likely than normal to including temperature and carbon dioxide availabil-
bind oxygen and less likely to bind carbon dioxide. ity. Consequently, a light saturation point is not
As photorespiration occurs when Rubisco binds constant for a particular species and may not even be
oxygen instead of carbon dioxide, boundary layers constant for two different leaves on the same plant
inhibit photosynthesis and enhance photorespiration. (McLendon and McMillen, 1982). Therefore, the

C3 and C4 Grasses Management 27

 1600

1400

1200
Light saturation
PPF (μmol/s/m2)

1000

800

600

400

200

0
7:30 8:30 9:30 10:30 11:30 12:30 13:30 14:30 15:30 16:30 17:30 18:30 19:30
Time

Fig. 3.3. Photosynthetic photon flux (PPF) during a summer day compared with the approximate light saturation point
of creeping bentgrass (Agrostis stolonifera). Notice that PPF exceeds the light saturation point for about 5 hours from
10:30 to 15:30. This situation encourages photorespiration in cool-season grasses. Source: Gaussoin et al. (2005).

light saturation point is a value that can be measured sealed container, the pressure of the gaseous form
on a given leaf, plant or group of plants at a single of that liquid trying to escape the container is
point in time that will change when internal or called vapor pressure. Liquids like alcohol and
external changes occur. Nonetheless, light saturation gasoline have higher vapor pressures than water.
is a useful concept. Increases in light beyond the light That means that they evaporate more easily and in
saturation point do not result in greater photosynthesis a sealed container there will be more pressure
(Fig. 3.3). In fact, increases in light beyond the light above alcohol or gasoline than there will be above
saturation point are detrimental. Just like all biologi- water at the same temperature (Fig. 3.4).
cal conditions, too much light can be as detrimental
as too little light. Photorespiration is one of those
Dynamic equilibrium
detrimental conditions encouraged by too much light.
You will learn of others in succeeding chapters. When a liquid in a closed container reaches a
steady vapor pressure it has achieved a dynamic
Temperature affects photorespiration Pressure Pressure

High temperature encourages photorespiration.

Remember back in chemistry class when you
learned that molecular activity increases with
increasing temperature. You learned a term called
“vapor pressure” and you learned that vapor pres-
sure increases with increasing temperature (Brown
et al., 1991). Consequently, you can heat a liquid Water
Water
Alcohol
Alcohol
such as water until it boils and turns to steam and
the water will quickly evaporate away. However,
Fig. 3.4. In a sealed container partially filled with
water also evaporates without being heated. Unless alcohol there will be more vapor pressure above the
it is in a sealed container under pressure, water alcohol than there will be above the same amount of
evaporates as long as it is not frozen. It evaporates water under the same environmental conditions. Heating
because there are always a certain number of mol- the water or the alcohol causes more molecules to
ecules escaping the liquid. If you put a liquid in a escape the liquid, and vapor pressure increases.

28 Chapter 3
equilibrium. Dynamic equilibrium is an important Because of this increase in activity, it is more likely
concept. Remember from Chapter 2 that during the for warm water molecules to escape from the liquid
discussion about affinity you learned that chemical into the air than it is for cold water molecules.
reactions and activities are dynamic, meaning that
they are constantly occurring and often changing
Oxygen and carbon dioxide in water
with changes in the environment. When a steady
vapor pressure is achieved, that does not mean that The recent discussion concerning temperature,
the molecules of liquid stop evaporating and that evaporation, vapor pressure and dynamic equilib-
the molecules of vapor stop condensing. It means rium was leading up to this point: oxygen and
that the number of liquid molecules that are carbon dioxide exist in water. They dissolve in
becoming vapor is the same as the number of vapor water and they also exist as whole molecules in
molecules that are becoming liquid in a particular water. When water warms, every molecule in the
period of time. Consequently, there is a lot of activ- water, including oxygen and carbon dioxide,
ity occurring but there is no net change in the becomes more active and more likely to leave the
amount of liquid or the amount of vapor present. liquid. The reason that water temperatures in your
Because there is no change in the amount of liquid plant affect photorespiration so greatly is because
or vapor, it appears that nothing is happening. carbon dioxide is not bound as tightly by water as
However, the vapor and water molecules are in is oxygen. As water gets warm the concentration of
constant motion, making the equilibrium dynamic, carbon dioxide in the water decreases more rapidly
not static. than the concentration of oxygen (Hull, 1992). The
Because of this vapor pressure property of liq- ratio of carbon dioxide to water becomes smaller
uids, water in a sealed container does not evapo- and the difference between the concentrations of
rate beyond equilibrium. However, water exposed carbon dioxide and oxygen in the water becomes
to the atmosphere evaporates readily. In a sealed greater. Therefore, there is greater likelihood for
container, the water vapor is in equilibrium with oxygen to be bound by Rubisco when the tempera-
the water and the air above the water is at 100% ture is high than there is when the temperature is
humidity. In the open, the air above the water is low. As temperature increases from 50 °F to 95 °F
continually refreshed. Therefore, water and water (10 °C to 35 °C), the rate of photorespiration in C3
vapor cannot reach dynamic equilibrium unless all plants more than doubles (Hall and Keys, 1983).
of the air in the surrounding area is at 100% Because all of the chemical reactions in the plant
humidity. So, except under rare circumstances, take place in water, the properties of water have a
water exposed to the atmosphere is evaporating direct influence on what occurs in the plant. As
continuously. Warm air can hold more water than water heats up, both oxygen and carbon dioxide
cold air so water evaporates faster when the air escape the liquid, but carbon dioxide escapes more
above it is warm. Warm water also evaporates rapidly and the remaining concentrations are rela-
faster than cool water. tively higher in oxygen and lower in carbon diox-
As we know, water molecules are in constant ide than they were when the water was cool.
motion. As water warms, movement increases. It is A scientist would say that the ratio of carbon diox-
this increase in movement that allows a water mol- ide to oxygen in water decreases as the temperature
ecule to attain a high enough velocity to literally fly of the water increases.
out of the water and into the air. Water molecules High light and high heat encourage photorespi-
in the air are also in constant motion, and if they ration in C3 plants. Therefore summer is a stressful
strike a body of water, they are likely to stick to it period for C3 grasses and is the season when they
and become part of it. Hence, there is a constant are most difficult to manage. Photosynthesis in C4
exchange of water molecules in both directions grasses is more efficient than in C3 grasses when
between the liquid water and the air. That exchange daytime temperature is high, but that does not
becomes faster as the temperature of the water mean that C4 grasses are unaffected by high tem-
increases. Heat is a form of energy, so when water perature. It means that they can perform photosyn-
absorbs heat and becomes warm it has more energy thesis efficiently when C3 grasses cannot. This also
than it does when it is cold. Consequently, water means that C4 grasses can metabolize normally at
molecules become more active and move greater higher temperatures than C3 grasses. It does not
distances with greater velocity as they get warmer. mean that C4 grasses are completely unaffected by

C3 and C4 Grasses Management 29

high temperature. It is widely believed that C3 Bundle sheath cells
grasses perform best at temperatures up to about
The cells that make the C4 pathway work so well
80 °F (27 °C) and that C4 grasses perform best at
are called bundle sheath cells (Rogers et al., 1976).
temperatures up to about 95 °F (35 °C) (Beard,
Cool-season plants may have bundle sheath cells,
1973). Beyond 95 °F, healthy growth of C4 grasses
but they do not perform photosynthesis in them.
begins to decline, but the decline is not a result of
Bundle sheath cells are small cells located deep in
photorespiration.
the leaf near the vascular system (Fig. 3.5). You
may recall that the xylem and phloem make up the
vascular system of a plant. The xylem primarily
3.3 Kranz Anatomy
transfers water through the plant for transpiration.
Warm-season C4 grasses not only perform photo- The phloem carries carbohydrates and other essen-
synthesis differently, they have different anatomy tial compounds through the plant from source to
from cool-season C3 grasses. Photosynthesis is com- sink. Mature leaves are sources because that is
partmentalized in C4 grasses. In C3 grasses, both the where most photosynthesis occurs. In grasses, pho-
light reaction of photosynthesis and the dark reac- tosynthesis may also occur in stolons, in above-
tion of photosynthesis occur in the same cell. In C4 ground stems and in sheaths, but production of
grasses the light reaction and the dark reaction carbohydrates at these sites is slight compared with
occur in different cells. C4 grasses also bind carbon that in mature leaves. Most stems, as well as roots,
differently. In C3 grasses, carbon is bound only in a flowers and seeds are sinks. They don’t produce
mesophyll cell, usually near the surface of the leaf, carbohydrates, they only use them. Because the
whereas in C4 grasses it is bound first in a meso- bundle sheath cells are located right next to the
phyll cell and is then transferred to a bundle sheath phloem, carbohydrates produced in the bundle
cell where it is released as carbon dioxide. Although sheath cells are easily transferred to the phloem for
this process requires extra steps to bind the carbon distribution.
and transfer it from one cell to another, it is more In C4 plants, the Calvin cycle takes place in the
efficient than C3 photosynthesis when plants are bundle sheath cells. It is the same Calvin cycle that
exposed to high light and high heat. occurs in C3 plants but it is located in these

Epidermis Spongy mesophyll

Palisade mesophyll

Xylem Phloem
Bundle sheath

Fig. 3.5. A representation of a cross section of a leaf from a C4 grass. The light reaction and carbon uptake occur
primarily in the palisade mesophyll cells near the upper surface of the leaf. The dark reaction takes place in the
bundle sheath cells near the vascular system, which is composed of the xylem and phloem.

30 Chapter 3
specialized cells. The only reason for the existence ATP and NADPH are used to fuel the Calvin cycle
of a bundle sheath cell appears to be to perform the and other plant processes. There is no Calvin cycle
dark reaction of photosynthesis in C4 plants. in the mesophyll cells of a C4 plant. The Calvin
Because bundle sheath cells are located deep in the cycle, the dark reaction, only occurs in the bundle
leaf they are relatively unaffected by light and by sheath cells.
atmospheric oxygen compared with mesophyll Carbon is bound in the mesophyll cells of C4
cells. In addition, the main source of oxygen, the plants just as it is in C3 plants but a different
light reaction of photosynthesis, does not occur in process is used to accomplish this binding. As you
the bundle sheath cells. Consequently, although the recall, in C3 plants the Calvin cycle exists in the
same Rubisco enzyme is used to bind carbon diox- mesophyll cells so carbon dioxide is bound by
ide for the Calvin cycle in the bundle sheath cells as Rubisco as it enters the cell. In C4 plants, the meso-
the Rubisco that is prone to photorespiration in C3 phyll cells do not bind carbon dioxide, they bind
plants, the Rubisco in the bundle sheath cell does bicarbonate. Bicarbonate (HCO3−) is carbon diox-
not, or at least only rarely, binds oxygen. The ide dissolved in water. Consequently, unless the pH
explanation for that is quite simple. There is very of the water is extremely low there is always a sub-
little oxygen present to be bound. stantial amount of bicarbonate in water exposed to
Bundle sheath cells are small. They do not rely air. A pH low enough to restrict the formation of
on atmospheric carbon dioxide for use in the bicarbonate would also be low enough to kill the
Calvin cycle. Instead, all of the carbon dioxide used plant. As bicarbonate enters the mesophyll cells, it
in the cycle is transferred to the bundle sheath cells is bound by an enzyme called PEP (phospho-
after being assimilated in the mesophyll cells. As enolpyruvate) carboxylase. The PEP carboxylase
bundle sheath cells are not directly exposed to enzyme catalyzes the combination of bicarbonate
atmospheric oxygen and because there is no oxygen and PEP, resulting in a 4-carbon molecule; hence,
production from the light reaction in the bundle the name C4 pathway. The C3 pathway is so named
sheath cells, the cytoplasm of a bundle sheath cell because the first stable molecule produced by fixing
is highly concentrated in carbon dioxide. High carbon in cool-season photosynthesis is a 3-carbon
carbon dioxide concentration and low oxygen is a compound, PGA. The C4 pathway is named C4
perfect situation for Rubisco. Its high affinity for because the first product of fixing carbon in warm-
carbon dioxide in an environment rich in carbon season photosynthesis is a 4-carbon compound,
dioxide makes it highly unlikely that Rubisco will which is called oxaloacetate. Oxaloacetate cannot
ever bind oxygen. Hence, C4 plants do not move through cell membranes so it is converted to
photorespire. malate, another 4-carbon compound, and shipped
off to a bundle sheath cell (Fig. 3.6). In the bundle
sheath cell, the malate is split into a molecule of
Carbon binding in C4 plants
pyruvate, which returns to a mesophyll cell, and a
Now that you know what a bundle sheath cell is molecule of carbon dioxide, which enters the
and why the Calvin cycle is so efficient in C4 plants, Calvin cycle. When the pyruvate returns to a meso-
we need to discuss how the carbon dioxide for the phyll cell, it is phosphorylated to resynthesize PEP
Calvin cycle gets to the bundle sheath cells. First and the process starts again.
let’s talk about the light reaction. The light reaction We have already discussed the advantages of
in C4 plants does not differ from the light reaction Kranz anatomy. There are also advantages to
in C3 plants. It takes place in the mesophyll cells binding carbon by combining bicarbonate and PEP.
near the surface of the leaves because that is where In water, carbon dioxide exists in three forms,
the light is most intense. Just as in C3 plants, chlo- carbonic acid, bicarbonate and carbon dioxide
rophyll in the mesophyll cells absorbs light energy (Box 3.1). Unless the pH of the water is extremely
and transfers that energy to reaction centers where high or low, bicarbonate is the predominate form.
it enters the z-scheme. Water is split producing Carbon dioxide and bicarbonate exist in
oxygen that vents to the air through stomates and equilibrium, but the equilibrium strongly favors
electrons that carry the light energy through the bicarbonate. Consequently, we believe that the
z-scheme. The z-scheme produces ATP and NADPH concentration of bicarbonate in the cytoplasm of
(the reduced form of nicotinamide adenine the mesophyll cells is always sufficient to perform
dinucleotide) just as it does in C3 plants, and the sustained photosynthesis regardless of temperature.

C3 and C4 Grasses Management 31

 CO2 HCO3–

Palisade mesophyll
PEP

Pi

Oxaloacetate

Malate

Malate

Pyruvate

CO2
Bundle sheath Calvin
cycle

Fig. 3.6. In C4 plants, carbon is fixed in a mesophyll cell by combining bicarbonate (HCO3−) and phosphoenolpyruvate
(PEP). This combination results in oxaloacetate, a 4-carbon compound that is converted to malate and shipped
to a bundle sheath cell. In the bundle sheath cell, the malate splits into pyruvate and carbon dioxide. The carbon
dioxide enters the Calvin cycle and the pyruvate returns to a mesophyll cell where it is again converted to PEP by
combination with phosphate (Pi), and the process starts again.

Box 3.1. Is the pH of pure water really 7.0?

Presumably the pH of pure water is 7.0. You know pure water in the presence of air becomes about
that because you have been told that many times by 5.7. If we were to add more acid to that water the
chemists, physiologists and botanists. However, you equilibrium would move in reverse and carbon
may not know that the difference between a pH of dioxide gas would evolve. If the concentration of
7.0 for pure water and a pH of 5.7 is simply the carbon dioxide in the air was to increase, more
presence of air. Carbon dioxide (CO2) dissolves in carbon dioxide would enter the water and form
water. When carbon dioxide dissolves in water, it more carbonic acid and more bicarbonate,
forms carbonic acid (H2O + CO2 ↔ H2CO3). The decreasing the pH further. In chemistry, assuming
carbonic acid and carbon dioxide reach equilibrium pure water has a pH of 7.0 works very well, and
with some of the carbon dioxide remaining in the that assumption is the right one to make. In
water as a gas, but most becomes carbonic acid. nature, water should have a measured pH of
Most of the carbonic acid dissociates to bicarbonate about 5.7, but because water in nature is not pure,
(H2CO3 ↔ H+ + HCO3−), and so the pH of otherwise it rarely does.

Therefore, carbon is rarely a limiting factor for C4 Warm-season grasses easily outperform cool-
photosynthesis. Although air movement over C4 season grasses when atmospheric carbon dioxide
grasses needs to be maintained, restricted air concentrations are low (Moss and Smith, 1972). In
movement over these grasses is not as great a fact, C4 photosynthesis is maintained at carbon
concern as restricted air movement over C3 grasses. dioxide concentrations as low as 2% of normal

32 Chapter 3
(Hesketh, 1963). Although high temperature can temperatures exceeding roughly 80 °F (27 °C).
cause a high enough loss of carbon dioxide from However, nature appears to have chosen the C3
water to restrict C3 photosynthesis, bicarbonate in pathway for plants in temperate climates.
the water will still be plentiful and C4 photosynthe- Buffalograss (Buchloe dactyloides), for instance, is
sis will not be restricted. the most cold-tolerant warm-season (C4) species
The enzyme PEP carboxylase does not bind oxy- that we use for turfgrass in the USA (Turgeon,
gen. What does that mean? It means that C4 grasses 2008). Although it survives extremely cold temper-
can perform photosynthesis in intense light for long atures, such as −20 °F (−29 °C) or lower, it has a
periods. It means that no matter how long the day very short growing season in temperate climates
or how intense the sun, C4 photosynthesis will not compared with C3 grasses. The same can be said of
be affected as long as carbon dioxide is present to Japanese lawngrass (Zoysia japonica), another rela-
dissolve and maintain sufficient bicarbonate levels tively cold-tolerant C4 grass. Its growing season is
in water. No other molecules compete for the bind- considerably shorter in temperate climates than the
ing site of the PEP carboxylase enzyme. Therefore, growing season of C3 grasses in the same location.
oxygen and other compounds do not affect the rate Perhaps C3 grasses perform better than C4 grasses in
of carbon binding in the mesophyll cells. Even cool weather because the C4 pathway includes extra
when plant photosynthesis is evolving oxygen rap- binding steps that require more energy (Black et al.,
idly, the air will not get rich enough in oxygen for 1973). A C3 plant requires 3 ATP molecules to bind
PEP carboxylase to bind it. Therefore, C4 photo- a carbon dioxide molecule. After first binding car-
synthesis eliminates the perceived deficiencies of bon in the mesophyll cell, and its then transfer,
Rubisco. The pathway sequesters Rubisco in small release and rebinding by Rubisco in the bundle
cells deep in the interior of the leaf and surrounds sheath cell, the C4 pathway requires 5 ATP mole-
the enzyme with highly concentrated carbon diox- cules. It is possible that a C3 plant must be photore-
ide. Rubisco is unlikely to bind oxygen in a bundle spiring at a particularly high rate before the extra
sheath cell. Instead of using Rubisco to bind car- energy required to perform C4 photosynthesis
bon initially, C4 plants use PEP carboxylase, which equals the energy lost to photorespiration in a C3
binds bicarbonate exclusively, so it is not affected plant. Although we know a great deal about photo-
by high oxygen concentrations. The use of bicarbo- synthesis, there are still questions to be answered.
nate as a carbon source ensures a sufficient supply Both C3 and C4 plants have unique characteris-
of carbon even when temperatures get exception- tics that influence management practices. In the
ally warm. The net result is that C4 photosynthesis following section, we will discuss the general differ-
is considerably more effective in the intense light ences between cool- and warm-season grasses.
and high heat of summer in temperate climates and Specific management techniques will be discussed
in tropical climates. later in individual sections of the text.

3.4 Characteristics of C3 and C4 Plants Temperature tolerance

After reading to this point, you probably have the It is clear that warm-season grasses are more heat
impression that C4 photosynthesis is a more efficient tolerant than cool-season grasses and that cool-
process than C3 photosynthesis. That would seem season grasses are more cold-tolerant than warm-
to be true and many experts believe it to be true. season grasses. Cool-season grasses differ in their
However, the fact remains that the number of C4 tolerance for cold and warm-season grasses differ
plants known to exist is very small compared with in their tolerance for heat (Turgeon, 2008). If you
the number of C3 plants in existence. All of these live near the equator or near the Arctic Circle that
plants have had considerable time to adapt to their would be very important to you as there are few
environments, and the C3 pathway is still the major grasses that can tolerate those extremes. However,
system of photosynthesis. Nature has determined most of us are more concerned about the heat
that C4 photosynthesis is best for plants in tropical tolerance of our cool-season grasses and the cold
regions. The performance of grasses during mid- tolerance of our warm-season grasses. Warm-
summer periods in temperate regions certainly season grasses differ in their tolerance to cold and
demonstrates that C4 is the most effective pathway cool season grasses differ in their tolerance to
for conditions of intense light, long day length and heat (Turgeon, 2008). However, how you manage

C3 and C4 Grasses Management 33

the C3 pathway when temperatures are high for tolerance will become increasingly important in the
your area, and how you manage the C4 pathway future, although it is important to note that they
when temperatures are low for your area differs are two different characteristics. The perfect turf-
little by species. grass for dry conditions would have good aesthetic
value, a low water use rate, a high drought toler-
ance and remain green during dry periods.
Adaptation to light
Of the grasses that we use most commonly for
Generally speaking, cool-season grasses are better turfgrass, the warm-season grasses tend to have
adapted to shade or cloudy conditions and tend to lower water use rates and higher drought toler-
grow better in those situations than do warm-season ances. Our warm-season grasses are genetically
grasses. You learned earlier that warm-season grasses adapted for producing aggressive root systems that
are much better adapted to intense light and long provide drought tolerance. It is not uncommon for
days than cool-season grasses. Also, generally speak- warm-season grasses to develop root systems that
ing, cool-season grasses grow better in low light and extend over 3 feet (91 cm) deep (Christians, 2007).
short days than warm-season grasses. There is no Consequently, these grasses can reach water when
such thing as a shade-loving grass. All grasses like the upper soil layers have dried out. Their low
full sun but some grasses like more sun than others. water use rates can be explained by the efficiency of
Most cool-season species are somewhat shade toler- the C4 pathway (Black et al., 1969). Warm-season
ant, and because of photorespiration prefer clouds grasses transpire less water (Kneebone et al., 1992).
rather than intense sunlight. Warm-season plants Because C4 grasses can bind bicarbonate, which is
tend to decline when cloudy conditions prevail for usually plentiful in water, stomates can be partially
more than a few days. St. Augustinegrass closed when needed to conserve water without
(Stenotaphrum secundatum) is an exception to these affecting the supply of carbon dioxide.
statements. This grass is a warm-season grass that,
in certain cultivars, exhibits shade tolerance that
Nitrogen use efficiency
rivals some of the cool-season grasses (Beard, 1973).
Select species and cultivars of zoysiagrass (Zoysia Warm season grasses use nitrogen more efficiently
spp.) also exhibit some shade tolerance, but they do than cool-season grasses (Brown, 1978). The
not appear to be as shade tolerant as most cool- leaves of C4 grasses contain less nitrogen per unit
season grasses (Duble, 1989). area or mass than C3 grasses, suggesting that less
nitrogen is required for each carbon assimilated
during photosynthesis (Hull, 1992). The reason
Drought tolerance and water use rates
for the C4 increase in nitrogen efficiency could be
If a plant is drought tolerant that does not neces- because a considerably smaller amount of Rubisco
sarily mean that it has a low water use rate. Some is needed in C4 leaves than in C3 leaves (Brown,
of the most drought-tolerant grasses escape drought 1978). A second possible explanation could be the
by entering a state of semi-dormancy. Warm-season allocation of nitrate reductase and assimilation
grasses like bermudagrass (Cynodon dactylon) and enzymes between mesophyll and bundle sheath
buffalograss and cool-season grasses like Kentucky cells (Moore and Black, 1979). Regardless, we
bluegrass (Poa pratensis) are good examples of don’t know the whole story.
plants that can remain dormant for long periods of Although research demonstrates that warm-
time during dry conditions. When these plants are season grasses are more efficient users of nitrogen
active, they do not necessarily have low water use compared with cool-season grasses, such is not the
rates but they are relatively good at surviving a case in practice. For instance, bermudagrass is a C4
drought. A cool-season plant like tall fescue grass that presumably is an efficient user of nitrogen
(Festuca arundinacea) has very good drought toler- but, in practice, it requires 0.8 to 1.5 pounds
ance for a cool-season grass, but a high water use N/1000 square feet each growing month (39–73 kg/
rate when water is plentiful (Qian et al., 1997). ha a month) to perform up to most customer expec-
Low water use rates and drought tolerance are tations (Beard, 1973; Duble 1989). A quick review
important characteristics of modern turfgrasses. of introductory turfgrass textbooks (see Suggested
Irrigation water is in high demand in most parts Reading, Chapter 1) reveals that the nitrogen
of the world, and water use rates and drought recommendations for C3 grasses do not exceed the

34 Chapter 3
nitrogen recommendations for C4 grasses. In this oxygen. Warm-season plants concentrate the car-
case, we are missing something. We have failed to bon dioxide by assimilating carbon from bicarbo-
consider other factors. Bermudagrass is extremely nate in the mesophyll cells and transferring it to the
aggressive. It has excellent recuperative ability, bundle sheath cells where they release it as carbon
spreads rapidly, forms a deep, extensive root sys- dioxide. This C4 process eliminates photorespira-
tem, and a vast network of stolons and rhizomes. tion and is considerably more efficient than C3
Perhaps it needs more nitrogen because of these photosynthesis when temperatures are high and
qualities, yet uses it efficiently. I honestly don’t light is intense. Consequently, C4 photosynthesis is
know. Again, we don’t know everything that we better adapted to tropical and subtropical climates
would like to know. than C3 photosynthesis. Because of the C4 pathway,
In this case, knowing the physiology, the fact warm-season grasses use water and nitrogen more
that C4 grasses are more efficient users of nitro- efficiently and they can perform photosynthesis
gen, does not help us in the field. The physiology when the carbon dioxide concentrations in the air
leads us to believe that C4 grasses require less are very low. However, they do not adapt well to
nitrogen fertilizer for optimum performance. Such low temperatures and low light, suggesting that the
is not the case. We need more information. C4 pathway is not as efficient under those condi-
Usually the physiology will lead you in the right tions as the C3 pathway.
direction. In this case it does not; there are always The warm-season grasses have either adopted
exceptions. the C4 pathway of photosynthesis because of their
environment or they have adapted to their environ-
ment because of the C4 pathway. In either case, the
3.5 Chapter Summary
warm-season grasses have characteristics not
Photorespiration occurs when Rubisco binds oxy- directly related to photosynthesis that differ from
gen. This process only occurs in C3 (cool-season) cool-season grasses. The C4 grasses tend to have
plants. Because of photorespiration, cool-season deeper root systems than C3 grasses and tend to be
grasses do not perform well when temperatures are more drought tolerant. In general, they do not per-
high and light is intense. Rapid photosynthesis in form well in low light and cannot survive extremely
intense light leads to high oxygen and low carbon low temperatures. Consequently, warm-season
dioxide concentrations in and around the plant, grasses require different management strategies
making the binding of oxygen by Rubisco more than cool-season grasses. The turfgrasses selected
likely. Also, high temperatures reduce the concen- for a particular site and the management practices
tration of gases, including oxygen and carbon required to meet customer expectations at that site
dioxide, in water. Carbon dioxide is less tightly are largely determined by the pathway that they
bound by water than oxygen. Consequently, as use for photosynthesis and the adaptation that
water gets warm the ratio of carbon dioxide to accompanies the pathway.
oxygen in the water decreases, further increasing
the likelihood that Rubisco (ribulose bisphosphate
Suggested Reading
carboxylase) will bind oxygen.
Warm-season (C4) grasses avoid photorespira- Hull, R.J. (1992) Energy relations and carbohydrate
tion by adding extra steps in carbon assimilation partitioning in turfgrass. In: Waddington, D.V.,
and by performing the light reaction and the dark Carrow, R.N. and Shearman, R.C. (eds) Turfgrass.
reaction of photosynthesis in separate cells. Warm- ASA-CSSA-ASSA (American Society of
Agronomy-Crop Science Society of America-Soil
season grasses have Kranz anatomy, meaning that
Science Society of America), Madison, Wisconsin,
they have bundle sheath cells located near their leaf
pp. 175–206.
vascular system where the Calvin cycle occurs. The Salisbury, F.B. and Ross, C.W. (1992) Plant Physiology,
light reaction takes place in the mesophyll cells 4th edn. Wadsworth Publishing, Belmont, California.
near the surface of the leaf and the dark reaction
takes place in the bundle sheath cells. Both reac-
tions are the same as the reactions performed in C3 Suggested Websites
plants, but they occur in different cells. The bundle NTEP (National Turfgrass Evaluation Program) (2009)
sheath cell where Rubisco resides is highly concen- Available at: http://www.ntep.org (accessed 17
trated in carbon dioxide so Rubisco does not bind December 2009).

C3 and C4 Grasses Management 35

 Respiration and Transpiration
4

Key Terms
A source is a part of a plant such as a mature leaf that produces more energy than it uses.
A sink is a part of a plant such as a root that uses more energy than it produces.
Carbohydrate partitioning refers to the allocation of carbohydrates from source leaves to other organs such as
roots and stems in a priority determined by season and environment.
Respiration is the breakdown of carbohydrates and other compounds into smaller units of cellular energy such as
ATP (adenosine triphosphate).
Stomates are small holes in grass leaves controlled by guard cells that open and close as necessary to permit or
restrict transpiration.
Transpiration is the transport and evaporation of water that is used to dissipate the heat that is produced in the
plant as a result of its metabolic processes, such as photosynthesis and respiration.
Photosynthates are the carbohydrate products of photosynthesis.
Starch is a long chain of glucose molecules and is the storage carbohydrate used by C4 grasses.
Sucrose is table sugar and results from combinations of a glucose molecule and a fructose molecule. Sucrose is
the predominate form of carbohydrate movement through a plant.
Amyloplasts are organelles where starch is stored.
Fructans are long chains of fructose molecules and are the storage carbohydrate used by C3 grasses. Fructans
are soluble, starch is not.
Vacuoles are organelles responsible for maintaining cell turgor pressure and are the site where most fructans are
stored.
Carbon dioxide compensation point is the amount of carbon dioxide present when the binding of carbon dioxide
for photosynthesis equals the evolution of carbon dioxide during respiration.
Phloem consists of sieve tube elements and companion cells and is the part of the vascular bundle, also called a
vein, through which plants move sucrose to where it is needed.
A pathogen is a disease-causing organism.
Passive transport means that plant energy is not required to move compounds from one location to
another.
Active transport means that plant energy is required to move compounds from one place to another.
The Apoplast is the area between cells also called intercellular space. It is, in fact, everything outside the cell
membranes including cell walls and xylem. It is the dead part of the plant.
The Symplast is the interior of a cell. It is the living part of the plant or the protoplasm inside the cell mem-
branes.
Aerobic means that air, in the case of respiration, specifically oxygen, is necessary.
Anaerobic means that air, in the case of respiration, specifically oxygen, is not necessary.
NADH (the reduced form of nicotinamide adenine dinucleotide) is an energy-rich electron donor molecule used in
respiration much like NADPH (the reduced form of nicotinamide adenine dinucleotide phosphate) is used in
photosynthesis. It is formed by the donation of electrons to its oxidized form, NAD+.
Capillary action is the result of adhesive and cohesive forces that cause water to rise upward against gravity in
a very small tube.
Xylem is the tubular system of tracheids and vessels that carries water from the roots to other parts of the
plant.
Symplastic movement is movement from cell to cell as occurs through phloem, or plasmodesmata.
Apoplastic movement is movement through intercellular spaces, through cell walls, or through xylem.

36 ©CAB International 2011. Turfgrass Physiology and Ecology (G. Bell)

 4.1 Encouraging Efficient Use plant locations where the energy is most useful.
of Carbohydrates and Water Consequently, respiration is of little use unless the
carbohydrates of photosynthesis are translocated
Plants have only five organs; leaves, stems, roots,
to where energy is needed.
flowers and seeds. Of the five plant organs only
leaves, except for a minor amount in stolons, per-
form photosynthesis. Leaves are a source; the other Components of plant energy
organs are sinks. The source provides chemical
Carbohydrates formed by active photosynthesis
energy in the form of carbohydrates. The sinks use
can either be respired by the photosynthesizing cell
it or store it. The translocation and fate of plant
for energy in that cell or be translocated immedi-
carbohydrates is called carbohydrate partitioning.
ately to other plant parts. However, most of the
The sink that is growing the fastest during periods
photosynthates produced are stored as starch in the
of growth or storing the most during periods of
chloroplast where they were formed, and then
storage has priority. Grasses, both C3 (cool season)
translocated during the night (Geiger and Servaites,
and C4 (warm season), for instance, slow shoot
1994). Both warm- and cool-season grasses use
growth and produce carbohydrates for storage in
starch as their temporary storage mechanism in
their roots and stems as winter approaches (McKell
chloroplasts, and translocate carbohydrates as
et al., 1969).
sucrose (table sugar). If carbohydrate supply
Release of chemical energy stored in carbohy-
exceeds demand, sugar is translocated mostly to
drates for use by the plant is called respiration.
stems and to roots for long-term storage and future
Respiration is not a particularly efficient process
use. In warm-season grasses, the sugar is combined
and much of the chemical energy in the respired
into units of starch (long chains of glucose) and
carbohydrates is lost as heat. When you work hard
stored in organelles called amyloplasts (Chatterton
you perspire because respiration in your cells is
et al., 1989). In cool-season grasses, the sugar is
occurring rapidly and heat is building up inside
combined into fructans (long chains of fructose)
your body. Perspiration is one of your body’s meth-
and stored in vacuoles (Pollock and Cairns, 1991).
ods for cooling itself. Photosynthesis and respira-
Vacuoles are organelles responsible for maintaining
tion in plants create heat so plants cool themselves
cell turgor and are unique to plants. Fructans are
by transpiration. Plants move water from their
soluble, but starch is not. Consequently, the storage
roots to their leaves where it exits the plant through
of fructans in vacuoles aids in maintaining cell tur-
stomates and evaporates. This translocation and
gor pressure in cool-season plants. Warm-season
evaporation for cooling purposes is called transpi-
plants use other solutes such as potassium to main-
ration and occurs consistently during daylight
tain turgor pressure in their vacuoles and cells.
hours and sometimes but rarely during warm
As mesophyll cells in C3 plants or bundle sheath
nights. If you have done everything that you can do
cells in C4 plants fix carbon dioxide, the resulting
to encourage healthy photosynthesis in your turf-
carbohydrates are stored in the chloroplasts as
grass, it is time to look at respiration and transpira-
starch. Plants respire continuously during both day
tion. Aside from photosynthesis, respiration and
and night, and a cell performing photosynthesis is
transpiration are the two physiological processes
also performing respiration at the same time
that can have the most influence on the health of
(Krömer, 1995). Consequently, some photosynt-
your turf.
hates are used or translocated immediately to
accommodate normal respiration. However, if pho-
tosynthesis is proceeding at a rate that exceeds the
4.2 Plant Respiration
carbon dioxide compensation point, which is the
In order for respiration to occur there must be car- point at which carbon dioxide fixation equals car-
bohydrate or another form of chemical energy bon dioxide evolution, excess photosynthates are
present. Plants can also respire lipids and proteins, produced. These photosynthates are stored as
but carbohydrates are preferred (Plaxton, 1996). If starch in the chloroplast where they were produced
a plant is respiring more than a small amount of and translocated later as they are needed.
(usually damaged) lipids and proteins, it is proba- Starch is a huge molecule and it cannot be
bly under stress. In order for plants to use respira- transported through cell walls and membranes.
tion effectively, the respiration must take place in So in order for the plant to translocate starch, it

Respiration and Transpiration 37

 Box 4.1. How do vacuoles help to maintain cell turgor pressure?
Pure water in the presence of air contains carbon concentration of potassium is separated by a cell
dioxide in the form of bicarbonate as a solute. No membrane permeable to water but not to potassium,
other solutes are present in pure water. What hap- and there is a solution on the other side of the cell
pens to osmotic water potential when we add a sol- membrane with a low concentration of potassium,
ute such as sugar or salt to otherwise pure water? water will move through the membrane toward the
Any general chemistry book will tell you that at least higher potassium concentration until both concen-
four things happen: (i) the freezing point of the solu- trations are equal. In other words, the water will
tion decreases; (ii) the boiling point increases; (iii) move until the ratio of solute to water is the same on
the vapor pressure decreases; and (iv) the osmotic both sides of the membrane. Consequently, there
pressure increases. Chemists call these colligative will be less water on the (originally) low potassium
properties. All of those colligative properties are concentration side of the membrane and more water
important to us in the study of biological systems on the (originally) high potassium concentration side
and in understanding how best to manage them. so that the potassium to water ratio is the same on
Everything that we have discussed so far in this text both sides. In the case of guard cells (cells that sur-
depends on osmosis. Osmosis may seem confusing, round the pore of a stomate), the movement of water
but it is actually quite simple. Colligative properties from outside a guard cell to inside that guard cell
occur when a solute is mixed with a solvent. In our causes the cell to swell which, in turn, causes the
case, the solvent is always water. Anything that dis- stomate to open. Remember: water always moves
solves in water is a solute. The solutes most impor- from a low concentration of a particular solute to a
tant to us in terms of osmosis are sugars, salts and high concentration of that solute. Solutes attract
nutrients. Potassium, for instance, is a solute. You water. In a tug of war for water, the high concentra-
will learn shortly that potassium is instrumental in tion of solute always wins. Vacuoles maintain high
the opening and closing of stomates by managing concentrations of solutes, mostly sugar and/or
osmosis. In biology, water is never pure. It always potassium. The sugar and potassium draw water
contains solutes, so it is always a solution. If a solu- through the vacuole membranes so that the vacuole
tion has a lot of potassium in it, we say it has a high can maintain turgor pressure. That is how vacuoles
concentration of potassium. If a solution with a high help to maintain cell turgor pressure in a plant.

must first be converted to molecules that can be used for cellular energy or combined into starch or
moved. The transport carbohydrate of choice in fructans for storage and later use (Plaxton, 1996)
most plants is sucrose, a sugar made of one mol- (Fig. 4.1).
ecule of glucose combined with one molecule of
fructose. Sucrose is what we know as table sugar.
Sources and sinks
It dissolves readily in water and moves easily
through cell walls and membranes. Some plants Mature grass leaves are sources, meaning that they
also transport raffinose, stachyose or verbascose, produce more photosynthates than they use
but turfgrasses transport mostly sucrose (Lalonde (Fig. 4.2). Very young leaves are sinks because they
et al., 2004). have yet to develop their full photosynthetic
Starch is broken down (hydrolyzed) one glucose potential. Very old leaves have outlived their use-
unit at a time in a chloroplast through chemical fulness and are also sinks for a short time until
processes catalyzed by enzymes called amylase and they senesce, a plant science term meaning that
starch phosphorylase. The resulting glucose is then they die and fall away. The sheaths of turfgrass
transported through the chloroplast membrane plants are normally green and can be a source of
into the cytoplasm. In the cytoplasm some of the photosynthesis, but they are not a major source of
glucose is converted to fructose and the two simple photosynthates. The same can be said for stolons.
sugars are combined to make sucrose. The sucrose Most stolons are green and perform photosynthe-
is loaded into the phloem and transported to a sink sis, but they are not major sources of photosynt-
where it is unloaded and either converted to hates for plant metabolism. Plant roots are sinks.
another carbohydrate, an amino acid or a lipid, They require energy for growth and for water

38 Chapter 4
 Chloroplast
NADP+ 2PGA Glucose
NADPH Calvin cycle
Starch synthesis
z-scheme

4e– Starch
O2
H2O Starch hydrolysis
4H+
Glucose

Glucose + Fructose

Cytoplasm Sucrose

To phloem

Fig. 4.1. Starch in a chloroplast is hydrolyzed to glucose and enters the cytosol (the fluid part of the cytoplasm)
where some of it is converted to fructose and combined with glucose to form sucrose. The sucrose is loaded into
phloem cells and transported to areas of need called sinks. NADP+ = nicotinamide adenine dinucleotide phosphate;
NADPH = reduced form of NADP+; PGA = phosphoglycerate.

This is the newest leaf;

new leaves are sinks

Mature leaves are sources

Flowers are sinks Seeds are sinks

Roots are sinks

Stems are sinks

Fig. 4.2. Mature leaves are the primary source of photosynthates for plant energy in grasses. Sheaths and stolons
may provide some photosynthates but are not major sources. All other plant parts are sinks.

Respiration and Transpiration 39

uptake. The root tip where meristematic growth annual and perennial biotypes, and is a good
occurs is an active sink. Energy is also necessary example of how much energy is required to pro-
for root water uptake so if photosynthates or oxy- duce flowers and seeds.
gen are not available for respiration, water uptake Technically, sinks are organs that use more
does not occur. energy than they produce. Sinks do not have to be
Reproductive parts are major sinks. Turfgrass organs that are rapidly using energy, they can be
growth slows, sometimes stops, as plants develop organs that are storing energy. Grasses primarily
flowers and seeds. Many crop plants are annual store carbohydrate energy in stems and secondarily
grasses. They produce considerable vegetation in in roots (Hull, 1992). During seasonal periods or
the weeks after establishment, but growth slows periods of low carbohydrate accumulation, such as
and stops as flowers and seeds are produced summer for C3 plants, these stems and roots can
(Barden et al., 1987). With few exceptions, turf- become sources rather than sinks (Pollock and
grasses do not flower and seed at the mowing Cairns, 1991). Spring, for instance, is a period of
heights that we commonly use. Those few that rapid shoot growth and root growth in grasses.
do waste energy. We would prefer that those More vegetative growth occurs in our turfgrasses
plants produce vegetation and spread rather during the spring than at any other time of year.
than produce flowers and seeds as flowers and Photosynthesis is occurring rapidly during periods
seeds are strong sinks for energy that could be of sunlight, but spring days are relatively short and
used for other purposes. A good example of a often cloudy in many locations. So where does all
plant that wastes energy is annual bluegrass this energy for spring growth come from? It comes
(Poa annua). from storage.
Annual bluegrass is ubiquitous throughout the Phloem transport is not always downward from
world and is often considered a weed. However, shoots to roots. In the spring, it is often upward from
under some conditions it makes an exceptionally roots to shoots. Every fall, turfgrasses slow in growth
nice turf (Huff, 1998). Annual bluegrass is best and move carbohydrates into storage to maintain
suited for golf course putting greens in areas winter metabolism (Hull, 1976). During winter,
where summers are cool, and it is competitive photosynthesis slows and often stops depending on
with perennial grasses such as creeping bentgrass the species and climate. Plants use stored carbohy-
(Agrostis stolonifera) and others. Under the right drates to maintain life over the winter and use what’s
situation, annual bluegrass can exist as an annual left in the spring to spread and grow. If we under-
or a perennial (Turgeon, 2008). There is also a stand this movement and when it occurs, we can use
perennial species of the plant that we call annual it to our advantage. For instance, spring is not the
bluegrass (Poa annua f. reptans). A perennial best time to apply a foliar-absorbed herbicide, but it
plant called “annual” bluegrass is a misnomer is the best time to apply a root-absorbed herbicide.
but, nonetheless, the name exists. One reason Most herbicides are meant to be applied to actively
that annual bluegrass persists as a weed and as a growing weeds. In the fall, turfgrass weeds move
turf in putting greens is because it can seed at photosynthates downward and foliar-absorbed
extremely low mowing heights. As a volunteer pesticides move with them. In the spring, phloem
plant, annual bluegrass germinates in the fall and movement is upward and foliar-applied pesticides do
continues rapid vegetative growth until winter not work quite as well. Although today’s herbicides
temperatures become too low for healthy metab- are effective in both spring and fall, you can increase
olism. During the fall and early spring annual their effectiveness by applying them at the right time
bluegrass appears green and healthy. As spring of year. Unfortunately, you don’t usually have a
progresses, however, the species begins to flower choice.
and seed prolifically. As the reproductive phase
continues, the plants turn yellow and look
Phloem transport
unhealthy because most of the available photo-
synthate energy goes toward reproduction rather Phloem transport occurs from source to sink.
than tissue growth and maintenance (Fig. 4.3). Phloem is the portion of the plant’s vascular system
This rapid vegetative growth stage followed by responsible for the translocation of sucrose and
prolific seeding is common in annual plants. In other sugars to areas of need. The phloem also
annual bluegrass, however, it is common in both transports larger molecules such as proteins, RNA

40 Chapter 4
 (a)

(b)

Fig. 4.3. Annual bluegrass (Poa annua) surrounded by a Kentucky bluegrass (Poa pratensis) lawn in early spring
(a) and late spring (b). Notice the difference in the healthy appearance compared with the lawn in early spring
versus a relatively unhealthy appearance in the late spring.

(ribonucleic acid) and even viruses (Oparka and However, this pressure also encourages rapid passive
Cruz, 2000). The phloem is under tremendous pres- movement of phloem sap from source to sink; the
sure because of the difference in solute concentra- word “passive” means that no plant energy is
tion (remember osmosis) at the source compared needed for transport. “Active” transport means that
with its concentration at the sink. This pressure cre- plant energy is required for transport. Phloem load-
ates problems for us when we mow. When we cut ing is an active process resulting in high pressure at
the phloem the nutritious contents leak out and the source but phloem transport is mostly passive.
become food for any number of microorganisms, The phloem consists of sieve cells and companion
including pathogens (disease-causing agents). cells (Fig. 4.4). Macromolecules such as proteins

Respiration and Transpiration 41

 phloem near those cells, sucrose will move passively
toward the phloem and enter the phloem. A second
and more common passage is through the apoplast,
Sieve plate
or intercellular space, and through cell walls and
membranes by active transport. In this case, the
sucrose is part of a co-transport system with pro-
tons in a process similar but backward to the ATP
(adenosine triphosphate) pumps in the thylakoid
membrane that you learned about in the z-scheme
of photosynthesis (Lalonde et al., 2004). In this
case, ATP is used rather than made. The energy in
ATP is used to fuel a proton pump. The ATP pump,
Sieve tube member
activated by an ATPase enzyme, is in the membrane
of the phloem companion cells. The ATPase enables
the pumping of protons from inside the cell to out-
side the cell. As a result, the apoplast becomes rich
in protons that diffuse back into the symplast (the
interior of the cell) as a result of the acid gradient
formed across the membrane. Sucrose enters the cell
with the protons in a co-transport relationship.
Consequently, the sucrose concentration in the
phloem can be substantially higher than it is in sur-
rounding cells. That means that the source cells do
not have to maintain the high concentration of
sucrose necessary to perpetuate a passive transport
gradient into the phloem. Owing to the systems of
Companion cells active and passive transport into the phloem, plants
are able to perpetuate high concentrations of
sucrose in the phloem for transport from sources to
sinks during both day and night.
As sucrose concentrations are high at the sources
and low at the sinks, sucrose can move passively
Fig. 4.4. The basic anatomy of phloem tissue. In through the phloem traveling with the concentra-
the plant’s vascular system, the phloem transports tion gradient. The faster the sink is using or storing
photosynthates, protein, RNA and, sometimes, by sucrose, the greater the gradient becomes and the
accident, microorganisms. more pressure that is placed on the phloem. In fact,
there is some evidence that sink demand can
and RNA that are transported in the phloem are actually influence the rate of photosynthesis in
synthesized in the companion cells. This is neces- leaves (Gifford and Evans, 1981).
sary because those molecules are too large to pass The basic idea behind phloem transport is fairly
through typical plant cell walls and membranes. simple if you remember your chemistry. Remember
The membrane between the sieve cell and compan- that water moves toward the lowest water potential
ion cell is not a normal cell membrane. It is believed and that water potential decreases with increasing
to consist of a thin line of organelles (Oparka and solute concentration. So if you add table sugar to a
Turgeon, 1999). The gaps between these organelles glass of water, its water potential becomes lower. If
permit the passage of large molecules. you have one glass of water straight from the tap
Phloem loading is both a passive and active and a second glass of tap water in which you added
process in grasses. Sucrose passes relatively easily two tablespoons of sugar, the water potential in the
through plasmodesmata, the intersecting cell-to-cell sugar water is lower than the water potential of the
passive transport system. As long as the sucrose pure tap water. What would happen if you were to
concentration in the cytoplasm of source cells is put tap water into a larger container, then take a
greater than the concentration of sucrose in the membrane permeable to water but not to dissolved

42 Chapter 4
sugar, partially submerge the membrane in the new companion cells is the result of the concentration
container, and pour the sugar water into it? The tap gradient from source cell to companion cell. More
water would move into the membrane until likely, however, the high concentration of sucrose
the water potential in the membrane was equal to in the companion cell is the result of active phloem
the water potential outside of the membrane. In loading by ATPase pumps, in which case phloem
this case, nearly all of the tap water would probably loading is considered active but transport is still
move into the membrane because the tap water is considered passive.
not likely to have enough solutes in it to equilibrate
with two tablespoons of sugar. So, except for some
Carbohydrate partitioning
water clinging to the sides of the container, nearly
all of it would move into the membrane to dilute Carbohydrate partitioning is the term used to
the sugar (Fig. 4.5). If the permeable membrane describe the priority distribution of carbohy-
was rigid and could not expand, the water moving drates throughout a plant. We actually know lit-
into it would create a great deal of pressure against tle about carbohydrate partitioning. Many
the side of the membrane and sugar water would scientific projects have attempted to use carbohy-
push out any available opening except back into drate content as a measure of turfgrass health.
the tap water. However, after multiple attempts over many
Because plant cells have walls and those walls years it appears that the amount of carbohydrate
are relatively rigid, osmotic pressure can build in a plant or plant part does not always correlate
within them as it did inside the membrane in the with plant health or visual turf quality (Xu and
tap water example above. The osmotic pressure Huang, 2003). In some cases, for the measure-
that builds near sources as sucrose fills companion ment of certain parameters under specific condi-
cells is the primary force behind phloem transport tions, there may be a relationship, but more
(Box 4.2). Although the pressure is obviously the often, there is not. We have known for many
result of an energy source, plant energy is not used years that the amount of photosynthate translo-
to move sap through the phloem and the transport cated to roots is negatively correlated with shoot
is considered passive. This pressure buildup growth (Mehall et al., 1984). Consequently, as
can also be passive if the only sucrose entering we reduce mowing height, we encourage shoot

Phloem

Apoplast

Source Sink

Fig. 4.5. As sucrose moves into the companion cells of the phloem either by active or passive transport the sucrose
solute lowers the water potential of the companion cell and attracts water from outside the cell. The water moving into
the cell causes pressure that forces water and solutes into the sieve tubes and instigates rapid flow toward the sinks.

Respiration and Transpiration 43

 Box 4.2. The pressure of the phloem.
How much pressure is actually present in phloem tis- weeping willow (Salix babylonica) was estimated to be
sue? Considerably more pressure is exerted on the about 145 pounds per square inch (1.0 MPa) (Wright
sides of plant phloem tissue than is exerted within the and Fisher, 1980). However, that pressure varies by
arteries and veins of the human circulatory system. It height and varies considerably by species according
is a good thing that plant cells have walls. Otherwise to the size of the sieve tubes and other variables.
this pressure could not be contained. The velocity of It is also interesting to note that the phloem
phloem transport is in the neighborhood of 20–40 pressure concept was first introduced by a German
inches/h (500–1000 mm/h) (Salisbury and Ross, scientist back in 1927 and has changed little since
1992). The pressure on the walls of the sieve tubes in then (Münch, 1927, 1930).

growth to replenish the portion lost to mowing tends to encourage shoot growth at the expense
and root growth suffers. The amount of carbohy- of root growth (Dudeck and Peacock, 1992). It is
drate relative to root mass has not changed. probably also reasonable to assume that the plant
However, the roots are fewer and shorter and the parts that are growing fastest are the plant parts
plant is less tolerant of dry conditions and less that are receiving the most carbohydrates.
capable of nutrient uptake. We have also known However, those plant parts will not have the
for many years that total nonstructural carbohy- highest TNC unless they are receiving carbohy-
drate (TNC) relative to tissue mass declines with drates faster than they are using them for
increasing shoot growth, suggesting that healthy growth.
plants can have low TNC (Youngner and Nudge,
1976).
Glycolysis
In 1996 and 1997, we attempted to use total
TNC content of entire grass plants as an objective Once an energy source such as sucrose reaches its
measure of shade stress (Bell and Danneberger, destination in a plant, it can be used for energy,
1999). There were four treatments, morning used for the synthesis of another compound or
shade, afternoon shade, all day shade and no stored for later use. When a compound is broken
shade. The no shade treatment was dense and down for its energy, we call that respiration.
healthy and the all day shade treatment had Respiration releases the energy stored in chemical
extremely poor cover and appeared unhealthy, bonds and converts it to cellular energy, usually
yet the TNC content did not differ between the ATP that is used to fuel cellular metabolism. There
shade and no shade treatments. We reasoned that are four kinds of plant respiration: the photorespi-
grass plants probably do not store carbohydrates ration that you already learned about, glycolysis,
during the growing season beyond a certain pre- pentose phosphate respiration and aerobic respira-
determined level. Instead, they use whatever tion. As you know, photorespiration is an effort to
excess carbohydrates that are available to form salvage as much carbon and energy as possible
new tillers and daughter plants. In fact, as stated when Rubisco (ribulose bisphosphate carboxylase)
earlier, research indicates that during periods of binds oxygen. The other three types of respiration
rapid growth TNC levels in grass plants are lower provide energy and the synthesis of plant com-
(Hull, 1992). Our conclusion may or may not be pounds for plant growth and maintenance.
true, but based on past research findings, it is a To extract all of the energy possible from a mol-
reasonable explanation and a good example of ecule of carbohydrate, protein or lipid, the com-
the thought process and information-gathering pound or its components must pass through the
effort required to make decisions when there is citric acid cycle and through an electron transport
no prior experience with a given situation. Had system in a process of aerobic respiration. In order
we measured TNC content in the shoots and for aerobic respiration to occur, air, specifically
TNC content in the roots, we might have found oxygen, must be present. Glycolysis is an anaerobic
more TNC in the shoots of the shade plants and process, meaning that oxygen is not required.
more TNC in the roots of the sun plants, as shade Glycolysis provides cellular energy in small doses

44 Chapter 4
very rapidly. Its strength is that it can provide so there are two GAPs passing through glycolysis
energy under the anaerobic conditions that can for every glucose or fructose that entered the path-
occur when respiration is moving so rapidly that way. Near the end of the pathway, 2-phosphoglyc-
oxygen is used faster than it can be refreshed. erate is converted to phosphoenolpyruvate (PEP),
Plant and animal respiration are not quite the the molecule that is so important for C4 photosyn-
same, but they are very similar. If you were to lift a thesis. A PEP molecule can enter late in the pathway
heavy weight repeatedly with the same arm in the and generate one molecule of ATP when it is con-
same way, your muscles would soon tire. You would verted to pyruvate. Pyruvate is the end product of
begin to feel pain as your muscles exhausted them- glycolysis. Two ATP molecules are required to con-
selves and, sooner or later, you would no longer be vert glucose or fructose to two GAP molecules. The
able to lift the weight. What you would have expe- two GAP molecules move through glycolysis in a
rienced is a balancing act between aerobic and series of three-carbon compounds, with ultimate
anaerobic respiration. Eventually, if you repeated conversion to two pyruvates. In this process, the
the exercise long enough, your arm would have tried two GAPs generate four ATPs, so the net product of
but failed to lift the weight and you would have had glycolysis is two ATPs.
to give up. If you had waited a few minutes, no mat-
ter how tired your arm became, you would have Glucose + 2ATP → 2 pyruvate + 4ATP
been able to lift the weight again. What happened? + 2NADH + 2H 2O
Your muscles lifted the weight easily the first time
you tried because they were well supplied with oxy- The NADH (reduced form of nicotinamide adenine
gen and aerobic respiration was not inhibited. Plenty dinucleotide, an electron donor like NADPH, nico-
of energy was available to lift the weight. As you tinamide adenine dinucleotide phosphate) pro-
lifted repeatedly, respiration went into overdrive in an duced in glycolysis can enter the electron transport
effort to supply the energy needed to complete the system of aerobic respiration where each will yield
weight-lifting act. Soon, your muscles were using two ATPs.
more oxygen than could be refreshed and aerobic Glycolysis is capable of producing pyruvate faster
respiration began to slow. Glycolysis was also provid- than the citric acid cycle can accept it. That is what
ing energy during the activity, and as respiration happened when your arm started to hurt. The
slowed, glycolysis was able to provide enough energy buildup of pyruvate reached a key level and because
to complete the activity. However, as the oxygen con- it could not enter the citric acid cycle, it entered the
tent became lower and lower in the muscles, aerobic fermentation process and was converted to lactic
respiration became completely ineffective and glycol- acid (lactate). In animals, fermentation usually
ysis was required to support the energy needed for the results in lactic acid, in plants it usually results in
entire task. As glycolysis can only provide a limited ethanol. Both products are toxic at high levels and
supply of energy, it was quickly over-tasked, the ATP should be metabolized quickly. The buildup of
in the muscle cells was exhausted, and you could no toxin is what caused your arm to hurt. Once the
longer lift the weight. The pain in your muscles was toxin had been partially metabolized and more oxy-
caused by fermentation. gen was supplied to your muscles, you could lift the
As far as metabolic pathways are concerned, gly- weight again. Glycolysis produces energy rapidly
colysis is quite simple (Fig. 4.6). The components of but it only produces two ATPs per six-carbon sugar.
sucrose, glucose and fructose, can enter glycolysis as The glycolysis process can provide energy when
glucose 6-phosphate or fructose 1,6-bisphosphate, oxygen is not present, such as in waterlogged soil,
both of which have to use a molecule of ATP in but it cannot sustain a plant for very long. If fer-
their preparation. Another molecule of ATP is mentation does not occur, the NADH produced
required to convert fructose 1,6-bisphosphate to during glycolysis is not consumed and will produce
glyceraldehyde 3-phosphate (GAP) or to dihy- two ATPs for each NADH in aerobic respiration. If
droxyacetone phosphate (DHAP). You may recog- aerobic respiration cannot occur, the NADH is con-
nize GAP and DHAP as important components of sumed by fermentation and glycolysis produces
the Calvin cycle. These compounds can enter glyco- only two ATPs. The process of aerobic respiration,
lysis directly from the Calvin cycle if needed. At this including the products of glycolysis, can convert a
point, the six-carbon sugars glucose and fructose six-carbon sugar to 36 ATPs, which is 18 times
have been split into two three-carbon compounds more energy than glycolysis alone.

Respiration and Transpiration 45

 Glucose
ATP
ADP

G6P FBP
Glucose-6-phosphate Fructose-1, 6-bisphosphate

ATP
ADP

GAP DHAP

Glyceraldehyde-3-phosphate Dihydroxyacetone phosphate

NAD+
NADH + H+

BPG
1,3-Bisphosphoglycerate

ATP
ADP

2PG 3PG
2-Phosphoglycerate 3-Phosphoglycerate
Acetyl-CoA
H2O

PEP Pyruvate
ADP
Phosphoenolpyruvate
ATP
NAD+ NADH + H+
NADH + H+ Citric acid cycle
CO2
Fermentation NAD+

Ethanol lactate

Fig. 4.6. Glycolysis is an anaerobic process that results in pyruvate, which can either be combined with carbon
dioxide or converted to acetyl-CoA and enter the citric acid cycle for aerobic respiration. However, if the citric acid
cycle is overloaded and cannot accept acetyl-CoA, the pyruvate degrades by fermentation to lactic acid or ethanol.
Both products are toxic at high concentrations and must be metabolized quickly. ADP = adenosine diphosphate;
ATP = adenosine triphosphate; NAD+ = nicotinamide adenine dinucleotide; NADH = reduced form of NAD+.

Many of the components of glycolysis are also pathway is in the business of degrading sugars.
precursors for other plant compounds (Table 4.1). A quick study of this pathway and you probably
realize that these compounds could be shared
The pentose phosphate pathway
with the Calvin cycle if the two processes occurred
A second form of respiration is the pentose in the same place. Apparently, the Calvin cycle
phosphate pathway (Fig. 4.7). As you examine provides these important compounds for the
Fig. 4.7, you probably recognize many of those chloroplast or bundle sheath cell. The pentose
compounds from earlier discussions. The pentose phosphate pathway provides them in the cytosol.
phosphate pathway is much like the Calvin cycle. The Calvin cycle uses these processes to synthesize
The Calvin cycle, however, is in the business of sugars and the pentose phosphate pathway uses
sugar synthesis, whereas the pentose phosphate the same reactions in reverse to break sugars

46 Chapter 4
Table 4.1. The products of glycolysis are used to synthesize other plant compounds.

Product Fate

Triose phosphates (phosphoglycerate (PGA), etc.) Glycerol (fats, oils, phospholipids), serine, cysteine
Phosphoenolpyruvate (PEP) Phenolic amino acids, auxin
Pyruvate Ethanol, lactic acid, alanine
Acetyl-CoA Fatty acids, carotenoids, gibberellins, terpenes

+
H NADP
+

6PGlu
H2O NADPH + CO2
6-Phosphoglutonate
(2) (3)

+
NADPH + H

+ 6PGuL Ribulose-5-
Ru5P
NADP phosphate
6-Phosphoglucono-
(1) lactone
(5) (4)
Xyulose-5- Ribose-5-
phosphate Xu5P R5P phosphate

G6P (6)
Glucose-6-
phosphate Fructose-6- Sedoheptulose-7-
phosphate F6P S7P phosphate

+
Glyceraldehyde-3- Glyceraldehyde-3-
phosphate GAP GAP phosphate

(7)
(8)

Erythrose-4-
F6P + E4P phosphate
Fructose-6-
phosphate

Fig. 4.7. The pentose phosphate pathway contains many of the same compounds that appear in the Calvin cycle.
NADP+ = nicotinamide adenine dinucleotide phosphate; NADPH = reduced form of NADP+.

Table 4.2. The products of the pentose phosphate

down. Many of these compounds are important
pathway are important components of other plant
for the synthesis of other products and for the functions and precursors for other important plant
functioning of other pathways (Table 4.2). products such as ATP (adenosine triphosphate),
The GAP produced by the pentose phosphate DNA and RNA.
pathway can enter glycolysis directly and
potentially yield 19 ATPs after full aerobic Product Fate
respiration. Fructose-6-phosphate (F6P) can also GAP (glyceraldehyde Glycolysis to ATP
enter glycolysis, potentially yielding 36 ATPs. 3-phosphate)
NADPH can be oxidized in the mitochondria to F6P (fructose- Glycolysis to ATP
produce ATP but it is also necessary as an 6-phosphate)
electron donor in pathways that synthesize fatty NADPH (reduced form Electron donor
acids and in other important processes. Erythrose- of nicotinamide adenine reactions
4-phosphate (E4P) is a necessary precursor for dinucleotide phosphate) Mitochondria to ATP
many phenolic compounds, and ribose- E4P (eryththrose- Phenolic compounds
4-phosphate)
5-phosphate is a required component for the
R5P (ribose-5-phosphate) DNA, RNA
synthesis of RNA and DNA.

Respiration and Transpiration 47

 Aerobic respiration Uses for components of aerobic respiration
Although plants have at least one other method Like the Calvin cycle, glycolysis and the pentose
for producing compounds that fuel aerobic respi- phosphate pathway, the citric acid cycle provides
ration, glycolysis is the dominant pathway for precursors for many important plant compounds
that purpose (Plaxton, 1996). Aerobic respira- (Fig. 4.9). Citric acid (citrate) for instance, is a pre-
tion is a two-part process much like photosyn- cursor for cholesterol and fatty acids. Sometimes
thesis in reverse. The first process of aerobic electron donor molecules like NADH from the
respiration is the citric acid cycle, also called the citric acid cycle or NADPH from the pentose phos-
Krebs cycle, and it is similar to the Calvin cycle phate pathway are required for the synthesis of
in reverse. The citric acid cycle yields the electron amino acids, fatty acids and other compounds
donors NADH and ubiquinol (UQH2). These necessary for plant life that originate from the citric
donors fuel an electron transport system like the acid cycle. Cellular energy, usually ATP from
z-scheme. As in the z-scheme the electrons aerobic respiration or glycolysis is almost always
traverse a series of reactions that moves hydro- required. Succinyl-CoA is a precursor for amino
gen protons from one side of a membrane to the acids such as isoleucine, methionine and valine. It is
other, setting up an electrical gradient. The same also a precursor for odd-chain fatty acids and for
type of ATP synthase spans the membrane and porphyrins, structural components of chlorophyll
with the passage of protons combines ADP and other pigments. Fumarate is a precursor for
(adenine diphosphate) and Pi (phosphate) to aspartic acid, phenylalanine and tyrosine, which
form ATP. Curiously, the final electron acceptor you probably recognize as amino acids. Malate is
is oxygen, and this is reduced to form water. a precursor for glucose. Oxaloacetate and
Recall that in photosynthesis water provides a-ketoglutarate are precursors for more amino
electrons and oxygen is evolved. In respiration, acids. A portion of the carbon fixed by photosyn-
oxygen is reduced to form water and carbon thesis provides the plant with energy through respi-
dioxide is evolved. ration. A large portion is also removed at various
stages of the energy pathways to provide all of the
structural and nonstructural components of the
The citric acid cycle plant (Plaxton, 1996). Therefore, your grasses must
take in considerably more carbon dioxide during
The citric acid cycle or Krebs cycle initiates aerobic
photosynthesis than they give off in respiration.
respiration. Although the citric acid cycle is consid-
ered part of aerobic respiration, oxygen is not
required for this portion of the aerobic respiration
The respiratory electron transport system
process. Remember that in photosynthesis the elec-
tron transport system, the z-scheme, was used first The final step in aerobic respiration is the electron
to capture light energy. The captured energy was transport system. The respiratory electron transport
then used to power the Calvin cycle. The Calvin system is located within an inner membrane in the
cycle stored whatever energy that it could recover mitochondria. The process is very much like the
in carbon bonds for use when it was needed to fuel z-scheme of photosynthesis (Fig. 4.10). Electrons are
metabolic processes. As the function of respiration donated to the pathway by NADH from the citric acid
is to release that energy into usable forms of cycle. UQH2 from the citric acid cycle and NADH
cellular energy, primarily ATP, it would stand to from glycolysis can also donate electrons. In the inner
reason that respiration would work backwards membrane of the mitochondria, the electrons pass
from photosynthesis. In respiration, the citric acid through a series of oxidation–reduction reactions that
cycle occurs before electron transport, forming transfer protons from the space between the inner and
electron-rich NADH and UQH2 for use in an elec- outer membranes (called the intermembrane space) to
tron transport system. When NADH and UQH2 the space inside the inner membrane (called the
are oxidized they donate electrons to the respira- matrix). An ATP synthase mechanism just like the one
tory electron pathway that occurs in the inner across the thylakoid membrane in the chloroplast
membrane of the mitochondria. The citric acid converts ADP to ATP as protons move from the
cycle also results in many components that are used matrix to the intermembrane space as a result of the
in the synthesis of other compounds (Fig. 4.8). electrical gradient across the inner membrane.

48 Chapter 4
 NADH + H+
Acetyl-CoA Citrate H2O
NAD+

Pyruvate
Oxaloacetate cis-Aconitate

NADH + H + H2O

NAD+

Malate Isocitrate
NAD+

NADH + H+
Fumarate Oxalosuccinate

UQH2

UQ CO2
Succinate α-Ketoglutarate

NAD+
ATP
Succinyl-CoA
ADP NADH + H+

Fig. 4.8. The citric acid cycle provides electron donor molecules for the respiratory electron pathway in the inner
membrane of the mitochondria. The cycle also produces components that are used in the synthesis of many other
plant compounds. ADP = adenosine diphosphate; ATP = adenosine triphosphate; NAD+ = nicotinamide adenine
dinucleotide; NADH = reduced form of NAD+; UQ = ubiquinone; UQH2 = ubiquinol.

Cholesterol
Pyruvate Acetyl-CoA Fatty acids
Citrate

Oxaloacetate Isocitrate

Amino acids Amino acids

Glucose Malate α-Ketoglutarate

Fatty acids

Fumarate Succinyl CoA Porphyrins

Amino acids
Amino acids
Succinate

Fig. 4.9. Some of the plant compounds that require precursors from the citric acid cycle.

Respiration and Transpiration 49

 Intermembrane Inner
space membrane Matrix
NADH + H+

H+ FMN
From citric acid cycle
H+ FMNH2
FP NAD+
NADH + H+
Fe2+ –S Fe3+–S
From glycolysis
H+ H2O
NAD+ FPPH2
–
UQ UQH2 OH
H+
H+
H+ ATP
ADP
Fe2+ Fe3+

H+ H+

H+ Fe3+ –S Fe2+–S
H+
H+ H2O
UQH2 –
UQ OH
H+
H+
H+ ADP ATP
Fe3+ Fe2+
+
H
H+ H+

Fe2+ Fe3+
H+ H2O
½O2
H+ Fe2+ Fe3+
H2O
Fe3+ H+
H+ 2+
Fe OH–

Fig. 4.10. The respiratory electron transport system in the mitochondria is much like the z-scheme of photosynthesis
and synthesizes ATP in a similar manner. The oxidation of water to oxygen begins the z-scheme and the reduction of
oxygen to water is the final step in respiration. ADP = adenosine diphosphate; ATP = adenosine triphosphate; FMN =
flavin mononucleotide; FMNH2 = reduced form of FMN (an electron donor); NAD+ = nicotinamide adenine dinucleotide;
NADH = reduced form of NAD+; UQ = ubiquinone; UQH2 = ubiquinol.

Oxygen must be present for the electron trans- oxygen was released. You learned that this oxygen
port system to occur. If there is no receptor at the evolution can cause problems when the air around
end of the electron transport pathway, electrons do your C3 grasses becomes high in oxygen and low in
not flow, and oxygen must be present to accept the carbon dioxide, resulting in photorespiration. It is
electrons as they exit the pathway. Oxygen will too bad that your plants cannot inject the oxygen
accept two electrons and two hydrogen protons to from photosynthesis into the soil where they really
form water. Remember that in photosynthesis, need it.
water was split to provide the electrons for the Boundary layers high in oxygen and low in car-
z-scheme. The split also resulted in two protons bon dioxide form over turfgrass during periods of
that increased the acidity of the space inside the high photosynthesis and poor air movement. Air
thylakoid, helping to establish an electrical gradient movement in soil is poor by definition, but it is
across the thylakoid membrane. During the process, adequate as long as the soil does not become

50 Chapter 4
compacted. We will talk more about compaction in cooling. Research demonstrates that grasses with
later chapters. In short, however, compaction is the high transpiration rates tolerate summer heat bet-
process of turning large soil pores into small soil ter than those with lower transpiration rates
pores through traffic or some other situation that (Perdomo et al., 1996; Abraham et al., 2008).
causes pressure on the soil. Large pores drain water
and are normally filled with air. Small pores hold
The components of transpiration
water by capillary action and are usually filled with
water. As the large pores are compacted to small To understand how transpiration works, it is neces-
pores, they stop draining and hold water. Where sary to revisit the concepts of the apoplast and
there is water there is no air. Consequently, there is symplast. Plant fluids exist in three different
no oxygen for root respiration. If a root cannot systems: the apoplast, the symplast and vacuoles
respire, it cannot provide energy for water uptake, (Canny, 1977). Vacuole fluids function to keep
which is an active process. So if the soil becomes individual cells turgid and to help prevent collapse.
highly compacted your turf has difficulty taking up They also have other important functions such as
water in spite of being surrounded by soil with regulating ion concentrations and storing carbohy-
plenty of water in it. If your turf has everything drates that we will discuss in later chapters.
that it needs to perform photosynthesis, the next Vacuoles are separated from the cell protoplasm by
thing that you need to evaluate is respiration. a special membrane called the tonoplast. Each
vacuole exists in only one cell and is not connected
to any other cell. The symplast, living protoplasm,
4.3 Transpiration
and apoplast, non-living fluid, are continuous sys-
It has been estimated that 60% of the energy seques- tems. The protoplasm of a root cell is connected to
tered during photosynthesis is lost as heat during the protoplasm of a leaf cell and to all other cells
aerobic respiration (Salisbury and Ross, 1992). We through the plasmodesmata (interconnecting cell
can probably assume that at least 60%, probably ports) of all the cells in between them (Fig. 4.11).
more, of the light energy absorbed by chlorophyll is The phloem is part of the symplast, but the xylem,
lost as heat before or during photosynthesis. the system through which a plant transpires, is part
Accordingly, plants get hot; plants get very hot. The of the apoplast. The symplast is separated from the
temperature of a turfgrass canopy on a warm day apoplast by interconnecting cell membranes.
with less than sufficient soil water uptake reaches Water passes through the cell membrane readily,
well over 104 °F (40 °C), the temperature at which but most solutes are restricted. As we discussed
enzymes and proteins begin to denature (Steinke earlier, some solutes can pass through cell mem-
et al., 2009). Extended exposure at this temperature branes passively, without requiring plant energy,
each day for several days will probably kill your but most have to pass through active pumping
grass. An extended temperature of 122 °F (50 °C) mechanisms. These active pumping mechanisms
will kill cool-season grasses in 2–3 hours and warm require cellular energy, usually ATP. These ATPase
season grasses in 10–12 hours (DiPaola and Beard, pumps remove nutrients from the apoplast for bio-
1992). A temperature of 140 °F (60 °C) will kill grass chemical activity in the symplast. Consequently, the
within a few minutes. If transpiration stalls because symplast tends to be highly concentrated in solutes
soil water is not present, a grass plant must enter a whereas the apoplast is relatively dilute. Water in
state of semi-dormancy, reach deep into the soil for the apoplast circulates throughout the plant cross-
more water, or die. ing cell membranes passively to rehydrate the sym-
Many botanists seem to feel that transpiration is plast when necessary. The water not used for
of limited importance. In fact, it has been stated metabolism is transpired.
that transpirational cooling is not necessary and is The apoplast is a lot more complicated than just
a waste of water (Salisbury and Ross, 1992). If intercellular spaces. It includes cell walls, spaces
transpiration did not occur, plants presumably between walls and spaces between membranes and
have other means of cooling themselves. If you walls. Cell walls are not impervious to water. In fact,
have been tasked to keep C3 turfgrasses green in the cell walls of a plant are always wet and their
temperatures greater than 90 °F (32 °C) or C4 pores are filled with water. Consequently, the walls
grasses green in temperatures over 105 °F (41 °C), themselves are part of the apoplast. This system of
you know how important it is to have transpirational cell walls and spaces is hydrated by the xylem.

Respiration and Transpiration 51

 Plasmodesmata
Symplast
Protoplasm

Intracellular space

Apoplast Vacuole
Intercellular space

Cell wall
Tonoplast

Fig. 4.11. Fluid in plants exists in three systems: a system of vacuoles, one per cell, that do not connect with each
other; the symplast, living material; and the apoplast, non-living material. The three systems are interconnected. The
symplast fluid inside the cell membranes interconnects from cell to cell through plasmodesmata. The fluid outside of
the cell membranes includes the cell walls, the space between the membranes and walls, and the space between the
walls, and is called the apoplast.

The xylem is the tubular structure, similar to the In grasses, as in deciduous trees, the xylem
phloem that carries water quickly from the roots comprises systems of cells called the tracheids and
to the shoots. The xylem and phloem exist in the vessels (Fig. 4.12). These cells have both primary
veins of the plant and are easily seen in turfgrass (normal) and secondary walls for added strength.
leaves. The veins extend throughout the plant, The secondary walls are thick and strong, but have
connecting all of the organs and tissues. Technically, small openings like valves to the primary wall so
a vein is called a vascular bundle and includes that passage of water and certain solutes can
xylem, phloem, and bundle sheath cells and/or fib- proceed normally to surrounding cells. Tracheids
ers (Fig. 4.12). In biology or plant biology it is are long and narrow; vessels are short and fat. Both
common to study the vascular systems of trees, cell types are hollow. Once they are formed, the
and considerable scientific study has also been protoplasm is translocated to other cells, and the
done with vascular systems of trees. If you are tracheid and vessel cells die, leaving only the walls
familiar with the vascular system of a tree, you will intact. Like irrigation pipes, the small tracheids are
notice from Fig. 4.12 that the vascular system of a more resistant to water flow than the larger vessels,
grass plant is not quite the same. A tree has one so vessels move water much faster. Water moves up
large vascular bundle that forms its trunk. The against gravity through the tracheids and vessels on
shoot of a grass plant has many small bundles principles similar to capillary action combined with
spaced in its sheath, similar to the spacing in the the attractive forces of water potential.
leaf. The sheath, in fact, is composed of the visible
leaves wrapped around each other. The newest leaf
The mechanisms of transpiration
is on the inside and the oldest leaf makes the out-
side layer. Consequently, the grass plant is not Trying to understand how water moves through
supported by a single stalk or stem but by multiple xylem can be confusing. Primarily because bota-
leaves wrapped around each other. Water does not nists do not fully agree on how water gets from the
pass up through the center of the shoot but all soil to the top of a 300-foot (96-m) tree. We really
around it. Except for late summer (C4) or fall (C3), don’t need to know how this occurs in order to
symplastic movement is downward and apoplastic manage turfgrass, but the same principles, osmosis,
movement is upward. adhesion, cohesion and water potential apply to

52 Chapter 4
 Bundle sheath cells

Sieve tube
elements

Companion
Phloem
cells

Xylem

Tracheids

Parenchyma

Vessel elements

Fig. 4.12. A representation of a vascular bundle in a warm-season (C4) grass plant. The sieve tube elements and
companion cells of the phloem share the vascular bundle, also called a vein, with the tracheids and vessels of the
xylem. The bundle is surrounded by bundle sheath cells where the Calvin cycle takes place in C4 plants.

the theory of xylem transport as apply to many the surface. Although that is interesting, it does not
other processes that we must understand to become concern us. There are principles that apply to
the best plant managers that we can be. grasses that cannot, according to physics, apply to
Consequently, the principles of water uptake and tall trees. One such principle is a very important
xylem transport deserve attention. one called root pressure.
The first thing that we need to do is realize that Root pressure is very similar to the osmotic
we are not dealing with a tall tree. We are manag- potential of the phloem. Before water can move
ing a grass no more than a few inches tall. Scientists through the apoplast, it must first be absorbed
have spent considerable time trying to work out from the soil. Root absorption is based on osmotic
how water can get from tree roots 20 (6 m) feet movement and water potential. If the water poten-
below the soil surface to the leaves 300 feet above tial of the root is the same as the water potential of

Respiration and Transpiration 53

the soil surrounding it, root water uptake does not These fluids contain sugar, proteins and other
occur. The root must have a lower water potential solutes that microorganisms love to feed on. Most
than the soil around it. So the root lowers its water of these microorganisms are either neutral or
potential by pumping solutes into its cell mem- beneficial, but some are pathogens. It is clear from
branes, thus encouraging water uptake by diffu- multiple research studies that this guttation fluid,
sion. Pumping these solutes across membranes is an if not removed from the grass, increases the likeli-
active process and the reason that roots cannot hood of disease occurrence (Couch, 1995; Ellram
take up water unless they can respire aerobically to et al., 2007).
produce ATP: no oxygen, no uptake. Once in the Root pressure is one principle that explains
symplast, the water moves through the cells and water movement in grasses, but it is not the most
passively diffuses into the apoplast, including the important. Capillary action is another possibility
xylem. When the grass is transpiring, the water that presumably cannot occur in trees. Based on
moves into the xylem because the xylem is rela- capillary theory and the size of tracheal tubes, cap-
tively dry. Hence, its water potential is much lower illary action can only account for water movement
than that of adjacent cells. At night, when the plant to approximately 1.5 feet (0.5 m) off the ground
is not transpiring because the cells are full, water (Salisbury and Ross, 1992). That would be suffi-
drawn into the roots has nowhere to go; not a very cient for grass, but capillary movement also requires
scientific statement but it is true, nonetheless. In that the upper end of the capillary tube be exposed
this case, water is moving into the apoplast because to the atmosphere. Based on plant architecture and
of pressure from the excess water in the cells and leaf hydraulics it does not seem likely that the apo-
we call this root pressure. plastic path is sufficiently clear to be considered
Root pressure can be substantial, but it is not open to the air in a capillary sense (Sack and
great enough to move water more than several Holbrook, 2006). However, the principles of adhe-
inches off the ground, it does not occur unless the sion and cohesion that are so important for capil-
soil is well watered and is rarely present during the lary action to occur also occur in the xylem. In fact,
day when stomates are open (Salisbury and Ross, as already mentioned, plant cell walls are porous
1992). Accordingly, root pressure is of no conse- and absorb water. So unless the plant is dehydrated
quence in most plants. It can have very dire conse- and about to die, the cell walls are filled with water,
quences for turfgrass, however. The dew that thus improving adhesion. The water in the xylem is
forms on grasses overnight does not all come from also dilute compared with the fluid in the cells, so
the atmosphere, some of it comes from the soil. We the osmotic potential of the symplast also works to
call it guttation fluid and it is a result of root pres- draw xylem water upward. Finally, the cohesion of
sure (Box 4.3) (Duell and Markus, 1977). The root water molecules is very strong and unless an air
pressure forces fluids from the leaves of grasses. bubble occurs in a xylem vessel or tracheid, the

Box 4.3. What is a hydathode?

Grasses don’t have to be mowed to exude guttation cells no longer open and close. Consequently, they
fluid. The guttation fluid is forced out through small appear to act as a pressure relief device. When
pores at the tips of leaf veins called hydathodes pressure builds up in a vein, the sap forces its way
(Stern, 1991). Although it may seem that stomates through the hydathodes and onto the leaf surface.
should occur at the tips of the veins, such is not the We continue to teach students that guttation fluid
case. Stomates evaporate water from adjacent cell from turfgrasses is exuded through hydathodes when,
walls and other apoplast components. The xylem is in fact, we mow most of them off. Each time we mow,
not exposed directly to the air through stomates, we open the xylem and phloem to the atmosphere.
which is why most botanists feel that xylem does not These wounds close quickly just as they would on our
have sufficient exposure to the atmosphere for capil- skin, but they are weak and root pressure can force
lary action to be a driving force of transpiration. them open to exude guttation fluid. The leaves may
Hydathodes are very much like stomates, and may regrow and form new hydathodes, but if we are
have evolved from stomates. However, their guard mowing regularly, we continually cut them off.

54 Chapter 4
xylem is always filled with a long column of water, practices that have a small effect. We will discuss
each molecule bound to the other about as tightly those practices more thoroughly in later chapters. Air
as intermolecular bonding can occur. movement and temperature have a definite influence
The most common hypothesis of xylem water on evaporation and anything that affects evaporation
movement is called the cohesion–tension theory also affects transpiration. The faster the atmosphere
(Steudle, 2001). Based on hydraulic and physical evaporates water, the lower its water potential. The
principles, the cohesion–tension theory demon- lower its water potential the faster the flow of water
strates an accumulation of forces sufficient to move through the xylem provided that the stomates are
water from the roots to the leaves of the tallest open and the roots are providing sufficient water.
trees. Although it is widely accepted by botanists, With that statement, it should become clear that
critics of the theory are not uncommon (Zimmerman many factors, plant, atmospheric and soil factors
et al., 2000). The cohesion–tension theory includes included, affect transpiration. The only means that a
the principles of adhesion and cohesion but is plant uses to control its rate of transpiration is by
based in water potential. In short, water is pulled opening and closing its stomates. Stomate control, of
by the molecular forces of adhesion and cohesion course, is not a conscious act, it is a response to
but the most important molecular force of the two external and internal plant conditions.
is the cohesive force of water. As long as water Stomates respond to light, specifically blue light
exists in a column of molecules bound to one (Zeiger and Hepler, 1977), to plant carbon dioxide
another, the difference in water potential between concentration and to increases in abscisic acid (ABA).
the air and the roots is nearly always great enough The opening and closing of stomates is controlled by
to cause water movement through the xylem from guard cells on either side of the stomate (Fig. 4.13).
the roots to the leaves. In the case of a grass, espe- When guard cells swell as a result of increased turgid-
cially a mown grass, that is not very far. You can ity, the stomates close. When guard cells lose water
think of the column of water in the xylem as a rope and shrink, their stomates open. That is easy to
held together by cohesive forces. The air applies remember. If the plant cannot get enough water and
tension to the rope pulling it upward through the its cells begin to shrink, the guard cells get smaller
xylem. If the rope does not break, the process con- and the stomates close so that more water is not lost
tinues as long as the air keeps pulling. to the atmosphere. So dry means less turgor and the
stomates close; wet means more turgor and the
stomates open. If you understand hydraulics,
The regulation of transpiration
Now that you know what the apoplast is, what the
When guard cells lose turgor, radial bands of
symplast is, what the components of the apoplast cellulose microfibrils relax and the stomate closes
and xylem are, and how water is transported through
the xylem, it is time for the important part. How is
water transport regulated? If we understand how
transpiration is regulated and how environmental
factors influence it, we can devise means to affect it.
The primary components of transpiration regula- Guard cell Guard cell
tion are miraculous little holes in grass leaves called
stomates. Stomates open to expose the apoplast to
the atmosphere. Unless the air around our grass is at
almost 100% relative humidity, the water potential Stomate
of the air will always be less than the water potential
of our plant’s apoplast. Obviously, relative humidity
is a very important environmental factor that affects
transpiration. Many grasses cannot tolerate high When the guard cells fill with water, the radial
microfibril bands pull the stomate open
humidity and most grasses are adversely affected by
it. We can influence humidity to some extent by Fig. 4.13. From above, the stomates in grass plants
encouraging air movement, adjusting irrigation prac- appear like sets of barbells suspended on two bars. When
tices, reducing shade periods and through other the guard cells expand, bands of microtubules attached to
minor management practices or management the bars pull the bars apart, opening the stomate.

Respiration and Transpiration 55

however, that procedure does not make sense. You shrink, the bars fall back together closing the sto-
would think that the guard cells would swell together mate. The opening and closing of stomates is medi-
closing the stomate, but that is not how it happens. ated by potassium ions (Zeiger and Hepler, 1977).
In most plants, guard cells are shaped like lips When conditions are right for stomates to open,
with the mouth serving as the stomate. The guard potassium pumps in the guard cell membranes
cells are wrapped with cellulose microfibrils that pump potassium into the cell. The increase in solute
restrict swelling. Think of a long balloon wrapped lowers the osmotic potential of the cell and water
with tape everywhere except on the ends. As you moves through the membranes causing the cells to
add air to the balloon, the tape restricts expansion expand and the stomates to open.
and expansion is channeled to the ends. If two of Light causes stomates to open (Fig. 4.14).
these balloons are placed together longitudinally Stomates are normally closed during the night. Of
inside a cardboard box where their ends cannot course, they are hardly ever closed completely and
expand sideways, the middle will separate as the there is almost always a small exchange between
ends expand pushing the centers of the balloons into the apoplast and the air. Stomates open when
half moon shapes in both directions leaving a hole in guard cells sense light and not only cool the plant
the middle. That is how the stomates of trees and during photosynthesis but admit the carbon dioxide
many other plants work, but it is not how the guard that is necessary for photosynthesis to occur.
cells of grasses work. In grasses, cytokinesis (cell Consequently, during daylight there is a constant
division) stops before completion so the guard cells exchange of carbon dioxide and water between the
never fully separate. The incomplete division results plant and the atmosphere. Carbon dioxide concen-
in a bridge between the cells that surround the sto- trations in the plant also affect stomatal opening.
mate (Fig. 4.13). The guard cells resemble a set of When light levels are low, carbon dioxide concen-
barbells with two bars instead of one. The stomate tration appears to dominate stomate response. In
is in between the two bars. In this case, the microfi- high light, however, the light level appears to domi-
brils are wrapped around the barbells with ends nate and carbon dioxide concentration can be high
attached to the bars. When the guard cells swell, the without affecting stomate status (Sharkey and
microfibrils pull the bars apart. When the guard cells Raschke, 1981). Stomates close when guard cell

Epidermis Spongy mesophyll

Palisade mesophyll

Guard
cells

Open Xylem Phloem ABA in

H2O in Bundle sheath cells K+ out

K+ in CO2 H2O out

Blue light H2O Closed

Fig. 4.14. When the guard cells of a stomate sense light, chemical reactions in the cell turn on the potassium pumps
in the membranes and the cell water potential declines. As water diffuses into the guard cells, they swell and cause
the stomates to open. A drought situation triggers a buildup of abscisic acid (ABA) in plant leaves that turns off the
potassium pumps, allowing the guard cells to lose turgor and the stomates close to conserve water.

56 Chapter 4
components sense an increase in ABA. This is a use, more than half is again lost, mostly as heat,
protective measure during drought stress, and it during respiration. If our grasses are to survive this
occurs regardless of light and carbon dioxide sta- heat when temperatures are high, they must tran-
tus. The ABA increases in response to drought spire. Not only must we provide water when neces-
stress in the leaf and triggers the closing of sto- sary if we want our plants to stay green, they must
mates (Wang et al., 2004). As you will learn later also have a strong root system to take up the water
in the text, heat stress is more severe when drought they need. If we do not intend to keep our plants
stress is also present (Jiang and Huang, 2000b). green during drought conditions, they still need a
good root system to maintain root and crown
hydration and remain alive until the rains occur.
4.4 Chapter Summary
Although photosynthesis is a very important
Photosynthesis fixes carbon into chemical bonds process, it is only part of the very complicated
providing a source of energy for use in building and chemical systems required to maintain plant life. In
maintaining a plant’s physiological systems. this text, we are also concerned with two other
However, photosynthesis, for the most part, only very important processes, respiration and transpi-
occurs in leaves. There has to be a system available ration. By learning how to encourage photosynthe-
in plants to transfer the photosynthetic energy to sis, respiration and transpiration, we also learn to
other plant parts in need. The energy is transported maintain the other complicated systems that sup-
in units of sucrose through veins containing phloem. port these three. In the remaining chapters you will
Phloem is a tube-like system comprising sieve tube be asked to use the knowledge gained from these
elements and companion cells that carries sucrose first four chapters to understand why management
to the plant parts needing it most. Plant parts that practices work and how to adjust them to your
need more energy than they produce are called advantage. By learning the underlying factors
sinks. Plant parts like mature leaves that produce rather than just the cosmetic symptoms that make
more energy than they use are called sources. So your grasses respond the way they do, management
the sources produce the energy and then the energy decisions become relatively easy.
is shipped to the sinks via the phloem.
The energy in the carbon bonds of sucrose and
Suggested Reading
other plant compounds is released for use through
a process called respiration. There are four types of DiPaola, J.M. and Beard, J.B. (1992) Physiological effects
respiration: (i) photorespiration; (ii) glycolysis; (iii) of temperature stress. In: Waddington, D.V., Carrow,
the pentose phosphate pathway; and (iv) aerobic R.N. and Shearman, R.C. (eds) Turfgrass. ASA-CS-
respiration. Respiration not only produces energy SA-ASSA (American Society of Agronomy-Crop Sci-
ence Society of America-Soil Science Society of
in the form of ATP (adenosine triphosphate), or
America), Madison, Wisconsin, pp. 231–268.
other energy molecules, it produces compounds Dudeck, A.E. and Peacock, C.H. (1992) Shade and
that are used to synthesize other important plant turfgrass culture. In: Waddington, D.V., Carrow, R.N.
components such as amino acids, lipids, RNA and and Shearman, R.C. (eds) Turfgrass. ASA-CSSA-
DNA. Carbohydrate energy is used for all plant ASSA (American Society of Agronomy-Crop Science
maintenance and growth. One of the most impor- Society of America-Soil Science Society of America),
tant processes requiring energy is root uptake of Madison, Wisconsin, pp. 269–284.
water and nutrients. Our grass leaves take in car- Hull, R.J. (1992) Energy relations and carbohydrate
bon dioxide and give off oxygen. Our grass roots partitioning in turfgrass. In: Waddington, D.V., Carrow,
take in oxygen and give off carbon dioxide as they R.N. and Shearman, R.C. (eds) Turfgrass. ASA-CS-
SA-ASSA (American Society of Agronomy-Crop Sci-
take up water. If we expect our leaves to perform
ence Society of America-Soil Science Society of
photosynthesis we must make carbon dioxide America), Madison, Wisconsin, pp. 175–206.
available. If we expect our roots to take up water,
we must make oxygen available.
Our third important process is transpiration. Suggested Websites
Grasses have to have a means of cooling them- University of Hamburg (2009) Botany Online – The Inter-
selves. Photosynthesis traps the energy of the sun. net Hypertextbook. Available at: http://www.biologie.
Much of this energy is lost as heat before it is fixed uni-hamburg.de/b-online/e00/default.htm (accessed
in carbon bonds. As this energy is released for plant 8 May 2009).

Respiration and Transpiration 57

 Why Our Management Practices
5 Affect Our Turf

Key Terms
Xeriscapes are landscapes designed to minimize or eliminate the need for irrigation.
Soil compaction is the compression of soil pores, usually caused by traffic, resulting in less soil air and greater
soil water-holding capacity.
Topdressing is the act of applying a thin layer of soil material, usually sand, directly over the turf and encouraging
it to penetrate the thatch (see below) by dragging a mat, heavy irrigation or some other process. Topdressing
can be used to fill soil pores following core aerification, or to smooth the surface, but it is mainly used to
encourage the microbial degradation of thatch.
Thatch is a layer of dead and living turfgrass roots, stems and crowns that exists between the grass shoots and
the soil.
Creeping grasses are those that spread not only by tillers but also by stems called stolons and/or rhizomes.
Shoot priority is a term used in this book to refer to the natural tendency of turfgrasses to favor shoot growth over
root growth and stem growth.
Humus is organic material that has been degraded over a long period to a semi-stable colloidal form.
Colloids are very fine particles of one substance somewhat evenly distributed throughout another. As used in this
book, clay particles and humus are colloids that help bind soil together to form structure.
Soil structure is the aggregation of soil particles to form larger units with small pores that hold water. These larger
units, called aggregates, have larger pores between each other that hold air.
Mat is thatch mixed with soil and is located between the thatch and the soil.
Aerification is a process used to help increase the air porosity of the soil. Core aerification, where circular probes
penetrate the soil to remove soil cores, is the most common aerification practice in turfgrass management.
However, there are several other protocols used as well. I prefer the term aerification rather than aeration so that
the cultivation of soil by aerification is not confused with the process of aeration of water and other substances.

5.1 Understanding What We Know From

detrimental practices and the stresses caused by
a Turfgrass Perspective
the natural and artificial environments in which
You have spent considerable time during the first turfgrass is expected to persist. Highly managed
four chapters of this book learning about cellular grasses are not natural. They are biological enti-
mechanics and important plant physiological ties growing in artificially managed environ-
processes. You may or may not have understood ments. We like our turf to be weed free, dense
the importance of these concepts but I hope that and uniform in color and height. There are no
you learned them well because now you are monocultures and very little uniformity in nature.
going to begin applying them. There are several When we manage our grasses for human aes-
common turf management practices that are thetic value and functional sports characteristics,
designed to either make turfgrass look better, we have to fight nature to do it. If we are not
make turfgrass more functional or keep turfgrass persistent and precise, we lose.
healthy. Many practices that make turf look Turgeon (2008) defines turfgrass as plants that
good are detrimental to turfgrass health. We use form a more or less contiguous cover that persists
the other practices to help relieve the stress of the under regular mowing and traffic. Notice that

58 ©CAB International 2011. Turfgrass Physiology and Ecology (G. Bell)

Dr. Turgeon used two detrimental human prac- irrigation, fertilization or cultural management
tices, mowing and traffic, to describe turfgrass. practices and will provide functional characteristics
Apparently turfgrass is a system of plants under that other plants lack. Functional characteristics are
continuous stress because of human intervention, further discussed in Box 5.1. The turf will not,
and that is exactly right. According to Beard however, look as good or play as good as we would
(1973) turfs have a number of functional charac- like. As turfgrass managers, it is our responsibility
teristics. They reduce wind and water erosion, to make turfgrasses look good and to provide ade-
mud and dust. Turfs reduce glare, noise, air pollu- quate playing surfaces. They do not need our help
tion, heat buildup and visual pollution problems to survive. They only need our help to survive the
(Beard, 1973). These functional characteristics, stress that we apply to them or to survive an envi-
however, are not unique to the grasses that we ronment for which they are not adapted (Fig. 5.1).
mow. These are natural characteristics and they According to our adopted definition, turfgrasses
occur with any mature grass sward. Dr. Beard also have to be mown and have to survive traffic.
describes turf as having recreational and ornamen- Therefore, mowing is a management practice that
tal value. The recreational and ornamental charac- has to be performed or we are not working with a
teristics of turfgrass make it desirable to us for turfgrass. Mowing by cattle, sheep, goats, rabbits
more than just functional characteristics and or other animals doesn’t count. Although animals
encourage its widespread use. Consequently, turf- were the original mowers, grasses mowed by
grasses are used all over the world for ornamental domestic animals are now called forages and
and recreational purposes, while also providing grasses mowed by wild animals are just called
functional advantages. In fact, the functional char- grasses. Were you aware that at one time, the Royal
acteristics of turf are often forgotten in favor of and Ancient Golf Club of St. Andrews in Scotland,
other types of supposedly environmentally sound believed to be the oldest surviving golf course, once
landscapes. Turfgrass managers have become so calculated the number of cunnigers or rabbits
incredibly good at what they do that people have (Oryctolagus cuniculus) required to keep the course
forgotten that grass does not really need our help at a particular mowing height (Beard and Beard,
to survive. In many locations, turfgrass is believed 2005)? It was estimated that 150 to 200 cunnigers
to require artificial irrigation and is, therefore, were required to graze 2.5 acres (1 ha) of bentgrass
undesirable. That is only true in particularly arid (Agrostis spp.)/fescue (Festuca spp.) turf to 0.4
regions for which only desert plants are adapted. inches (10 mm). Thanks to Dr. and Mrs. Beard for
In most situations, the plants that are recom- that interesting tidbit and an interesting paper. The
mended for landscapes because they require little study of turf does not always have to be boring.
irrigation require more water than turfgrass (Park Hopefully you can find the book that includes that
et al., 2005). paper at your library. See the list of references at
Grass needs our help to look and play as we the back of this book for more information.
want it to. It does not need our help to survive in By definition, we are also faced with traffic prob-
areas for which it is adapted. We use certain grasses lems on turfgrass. Whether it is a lawn, park, golf
as turfgrasses for a reason. They are highly adapted course or athletic field, turf is meant to be walked
to most of the regions in which they are grown and on, sometimes driven on. If you live in an agricul-
they are highly resistant to stress. If the proper tural area, you know that it is considerably easier
turfgrass is chosen for a site, based on its adapta- and safer to walk or drive on a grass field when it
tion, it will survive easily without our help, but is wet than it is to walk or drive on a dirt road
there are some regions where turfgrasses are not turned to mud. Turfgrass roots and sometimes rhi-
adapted and will not grow without help. zomes are extensive and extremely effective for
Turfgrasses will not survive desert conditions holding soil together and providing traffic areas
unless we provide water. In that situation we should resistant to separation and to compaction. However,
consider whether or not the functional characteris- repeated traffic can overcome the root/rhizome
tics of turf are sufficiently important to justify the system of turf, turning the grass field into the dirt
use of water for irrigation. In most regions of the road. The soil will separate, allowing a vehicle to
world, turfgrasses can exist in low maintenance, sink when the soil is wet enough or the vehicle is
mow-only situations. Turfgrasses adapted to the heavy enough. Turf areas can only stand a finite
region in which they are used will not need amount of repeated vehicle or foot traffic until the

Management Practices 59
 Box 5.1. Have people forgotten about the functional characteristics of turfgrasses and turf?
Although it may appear that I use turfgrasses and system of labor, water, fertilizer and perhaps even
turf interchangeably they are not actually the same pesticides. So I ask you: is this environmentally
thing. Turfgrasses are plants. A turf is an ecological sound management?
system that includes the plants, the dead material Let’s consider another potential system, a xeris-
accumulated because of the plants and the soil that cape: a system of drought-resistant, pest-resistant
surrounds the turfgrass roots. Consequently, func- plants that requires no fertilizer growing naturally on
tional characteristics as well as recreational and sand or native soil. The only maintenance performed
ornamental characteristics are qualities of the turf, on this site is trimming and mechanical weed
not the turfgrass. So why do people seem to have removal. Compare this with a low-maintenance turf-
forgotten the functional characteristics of turf? grass species that is mowed as needed to maintain
I have very definite opinions on this matter, but I a 3-inch (8-cm) mowing height and weeded on a
would rather you form your own opinions and be similar interval to the xeriscape. Which is most
able to support them with logical thought. Everyone aesthetically pleasing – the turf or the xeriscape
knows how nice a manicured golf course, home lawn (Fig. 5.1)? Which is most environmentally desirable?
or commercial area looks under intensive turfgrass Which system has the greatest functional value? Is
management. These areas are especially attractive one system always the best alternative compared
if they include other ornamental plants designed to with the other? Is there some reason other than
attract attention, with the manicured turf used as a aesthetic value that a drought-tolerant turfgrass has
background. Aesthetically it would be very difficult to to be irrigated? Which system would you recom-
improve on that system. People like it; you and I like mend to a neighbor who does not want to irrigate,
it. In this scenario we not only are managing a good- fertilize or apply pesticides? Can you support your
looking area, we are perpetuating the functional conclusion with a logical argument? As a plant
characteristics of turf. However, there is a price for manager, these are questions that you should be
aesthetic value. The beauty is maintained through a comfortable with and able to discuss intelligently.

soil becomes compacted and the grasses thin or die. we mow when the grass reaches a height of
Consequently, we use cultivation techniques to help 3 inches thereby removing one third its height.
relieve soil compaction and we employ aggressive The grass shoot is not all photosynthetic leaf
sod-forming grasses in areas where traffic is area. It is a combination of sheath and leaves.
expected to be severe. We use other techniques, The sheath is not a major source of photosynt-
such as specialized mowing and topdressing to hates. So when we remove 1⁄3 of the canopy
smooth playing surfaces and reduce thatch, a measured by height from the soil surface, we are
buildup of dead plant material unique to turf. All removing around ½ of the primary photosyn-
of our management practices affect turf either posi- thetic leaf area. As photosynthesis is one of the
tively or negatively. The negative practices make three important plant processes that we try to
the turf look better or make it available to us for encourage as managers, removing ½ of the
recreation. The positive practices help us counter- plant’s photosynthetic capacity is highly detri-
act the negative practices. mental and counterproductive to our goals.
However, it is not a turfgrass if it is not mowed.
Consequently, we have to mow and we have to
5.2 Mowing Causes Chronic
help our turf overcome the detrimental affects
Damage to Turf
of mowing. Let us consider just how detrimen-
Christians (2007) made a profound comment tal mowing really is.
when he wrote “turfgrasses do not thrive on
mowing – they tolerate it”. Mowing is an injury.
The negative aspects of mowing
We mow our grasses according to the 1⁄3 rule.
We never like to remove more than 1⁄3 of the When we mow, we not only remove a large portion
canopy of our turfgrasses at any one time. For of photosynthetic material, we also create an injury
instance if our target mowing height is 2 inches (Madison, 1962; Krans and Beard, 1985). This

60 Chapter 5
 (a)

(b)

Fig. 5.1. (a) A planned low-maintenance xeriscape, and (b) a low maintenance turf. If the turfgrass is naturally
adapted to the region, it will require little if any maintenance and provide functional value that the xeriscape cannot
provide. However, a beautiful xeriscape like this one also has value.

injury has to be healed. Luckily, plants, especially withstand poor environmental conditions better
grasses, are very good at growing new parts. If a than most ornamental plants. Imagine what would
grass is healthy it can grow new leaves or fix dam- happen to most ornamental plants if you removed
aged leaves. Even if the grass is scalped so low that half their leaf area weekly. This could be a good
all of the leaves are removed, it can grow new ones. student project; do you own any bushes that you
The effort required of a previously healthy plant to would like to challenge?
grow a new leaf system may weaken it so much Although grasses are quite capable of healing
that it dies as a result of other damaging factors. themselves, the frequency at which we mow
However, it will most likely survive if growing exposes them to continuous, or at least recurring,
conditions are conducive. Turfgrasses are survi- chronic injury. A golf course putting green, for
vors. They are not easy to kill and will normally instance, that is double cut for a tournament has

Management Practices 61
lower photosynthetic efficiency than a green cut
only once (Howieson and Christians, 2008).
A portion of a turfgrass plant’s energy is continu-
ously required to heal its injuries from mowing.
Mowing requires the removal of photosynthetic
area. Mowing seriously affects photosynthesis,
causing, among many other things, a serious
decline in root length (Youngner and Nudge,
1976). The shorter the mowing height, the shorter
the root system becomes. Imagine how much more
energetic these grasses could be if we were not
removing half their leaf area on a regular basis. In
addition to affecting water uptake by reducing
root length, the injury also creates further problems
with another of the three primary physiological
functions, transpiration.
Grasses control transpiration by adjusting their
stomates to allow more or less transpiration to
occur and more or less carbon dioxide to enter the
leaf. When the leaf cells begin to dry out severely,
a buildup of abscisic acid (ABA) occurs, closing
stomates and conserving water. When we mow, we
open the xylem and other apoplastic areas to
evaporation. Substantial soil water is lost for a
short time after mowing, and the closing of sto-
mates cannot prevent it. If the soil is dry to begin
with and the plant is near drought status, dehydra-
tion occurs rapidly and leaf tips dry beyond recov-
ery (Fig. 5.2). Under those circumstances you will
notice the leaf tips turn white then brown as their
cells die over a period of hours or days. Shredding
Fig. 5.2. Leaf tips lose water uncontrollably for a short
of the leaf tip can cause an identical condition.
time following mowing. If sufficient water is not available
Dull mower blades tear rather than cut, shredding
to replenish this loss quickly the cells near the injury
leaf blades and exposing more than twice the leaf dehydrate and die.
area to desiccation as would occur with a clean
sharp cut. These leaf blades desiccate and partially
die even when sufficient water is available. photosynthetic potential has been recently reduced
Regardless, water use rate increases for a short by half and whose leaf tips are desiccated and in
time immediately following mowing, wasting soil need of repair are increasingly susceptible to
water that could be used for normal cooling disease.
processes.
Mowing wounds allow microorganisms to enter
The positive aspects of mowing
leaf blades (Vargas, 1994). Nearly all of the
economically important diseases of turfgrass are Mowing disrupts two of our primary plant func-
caused by fungi. Most fungi though are not patho- tions, photosynthesis and transpiration, but the
gens. Most are either beneficial or neutral to turf, mowing process is not entirely negative. Mowing
but some fungi cause nasty diseases in our grasses has some beneficial aspects as well. These benefi-
(Fig. 5.3). Pathogenic fungi are capable of degrading cial aspects are ornamental and recreational, not
cellulose to enter grasses, but their entry is easier and agronomic, and only occur because mowing is not
faster through wounds. Consequently, turfgrasses as disruptive to photosynthesis as you might sus-
with wounds from mowing, traffic and other injuries pect. One of the principles of ecology that we will
are more susceptible to disease. Turfgrasses whose discuss later in the text has to do with maintaining

62 Chapter 5
 In this case, the price is root growth. The lower
turf is mowed, the shorter the root system becomes
(Liu and Huang, 2002b). By spreading, a grass can
increase its photosynthetic potential. However, the
photosynthetic potential is less than the potential
that existed at its former higher height of cut.
Although the photosynthetic potential increases as
more shoot area is added, the transpiration poten-
tial is reduced by shorter root growth. Consequently,
the plants are now more susceptible to drought
stress and heat stress, and more precise irrigation
management is required. Turfgrasses tend to
exhibit shoot priority; this means that in most situ-
ations, shoot growth is favored at the expense of
root growth. When excessive nitrogen is applied,
for instance, an increase in shoot growth normally
occurs, sometimes at the expense of root growth
(Dunn et al., 1995). When photosynthetic area is
reduced, shoot area is partially maintained later-
ally, but root growth suffers (Tucker et al., 2006).
Shoot priority is a concept that you need to
remember. It influences how grasses respond to
stress and how they respond to your management
practices.
Mowing has one more important advantage,
Fig. 5.3. This tall fescue (Festuca arundinacea) leaf is this time an agronomic one. Mowing helps to
infected with brown patch disease caused by Rhizoctonia control weed competition. If you are thinking
solani. Based on the spread of the symptoms, the ahead, you might want to argue that issue, as
pathogen probably entered through the leaf tip. some of the weeds that compete with turfgrasses
need to perceive light in order to germinate. The
consistent plant biomass. Mature turfgrass plants notorious summer annual, crabgrass (Digitaria
strive to maintain their biomass at levels within a spp.) and winter annual, annual bluegrass (Poa
certain range. It helps to think that their physiology annua) are two of those weeds (although you may
is designed to operate best when their shoot struc- remember from Chapter 4 that annual bluegrass
ture constitutes a particular mass. When we mow, is a suitable turfgrass under some conditions).
the plant not only tries to heal the wound and grow Opening the canopy allows more light to reach
new leaves, the mowing encourages new growth. It the soil surface and encourage weed seed germi-
is believed that the mowing of the canopy allows nation. Some of that germination can be discour-
more light to strike the lower leaves of grass plants, aged by proper irrigation practices, but nature
thus encouraging the formation of new tillers and, has a way of providing those species with the
in the case of creeping grasses, new daughter plants conditions that they need at the time that they
(Duble, 1989). It is well known and accepted that need them. Consequently, those weeds are likely
mowing turfgrass lower within its range of adapt- to germinate.
ability results in greater shoot density (Juska and Although weed pressure is severe and generally
Hanson, 1961). If the grass plant cannot increase constant, we should not underestimate the
its biomass vertically, the biomass increases hori- aggressive nature of most turfgrasses. An
zontally. So a very good way to increase the aes- immature weed is not much competition for a
thetic value of your turf is to mow it near the low mature turfgrass in the act of spreading. Naturally
end of its range of adaptability, thereby increasing some weed encroachment will occur when mow-
its density and ornamental characteristics. Increases ing height is lowered but, eventually, turfgrass
in aesthetic value or playability, however, always density will increase and the weed population
seem to exact a price. is likely to decline rather than increase. Unless

Management Practices 63
 Mowing management
herbicide application or time-consuming, back-
breaking mechanical removal are practiced, Mowing affects photosynthesis by removing por-
100% or even 80% weed control is unlikely. tions of leaves, the most important plant organ for
Nature does not like a monoculture. Nature likes photosynthesis. It also results in temporarily uncon-
diversity (Fig. 5.4). If we mow our grasses lower trolled transpiration. By minimizing the detrimental
than our grass species can tolerate, weeds are effects of mowing on these two important plant
likely to out-compete the turf (Voigt et al., functions you are doing most of what you can to
2001). We are fortunate that most weeds cannot help your turf survive and prosper under chronic
compete with turf at common mowing heights. mowing. If each time that you injure the leaves by
So mowing alone provides a substantial amount mowing uncontrolled transpiration occurs and pho-
of weed control. tosynthesis is reduced, how often should you mow?

(a)

(b)

Fig. 5.4. (a) A common bermudagrass (Cynodon dactylon) managed using irrigation, fertilization and herbicide weed
control as well as regular mowing. (b) A common bermudagrass where the only management practice used is regular
mowing. People like uniformity, nature likes diversity.

64 Chapter 5
 This is a good time to start thinking about such 1988). That minimizes injury and water loss by pro-
things and preparing yourself to start making deci- longing the period between losses of photosynthetic
sions that solve problems. Decisions are made by material. If you thought about that, you probably
considering alternatives. Management practices are came to a similar conclusion. If not, no problem,
often developed through trial and error, but not keep trying. What else can we do to reduce the nega-
without forethought. You should begin to practice tive effects of mowing? Think about it.
your skills in creating solutions to problems and in What else can we do to reduce the negative agro-
organizing plans now while you have an instructor nomic effects of mowing? We need to use the right
or author to help you. All too soon, you will have equipment and the equipment needs to be in good
to make those decisions and plans without help working order and operated correctly (Box 5.2).
and with considerably greater risk. Practice your Both operational training and safety training will
skills now while you can. Mistakes are part of be an important part of your job as a turfgrass
learning. We learn more from our mistakes than we manager. Most of you are going to have to stop
do from our successes. being the worker and start being the manager, and
As far as mowing frequency is concerned, you that is a difficult transition. It is far easier to do the
mow as infrequently as possible while maintaining work yourself than it is to successfully encourage
the turf to your customer’s expectations and abiding and train others to do it for you. You will not only
by the 1⁄3 rule (Johnson et al., 1987; Soper et al., be responsible for the work you do, you will be

Box 5.2. Know your mower.

We usually use one of three types of machines for the cutting action occurs at the tip of the blade as the
mowing turfgrass, a reel mower, a rotary mower or a mower moves forward. If the front of the deck is
flail. Each has strengths and weaknesses that are lower, the grass is cut normally but if the back of the
clearly explained in introductory turfgrass texts. deck is lower, the grass is cut twice, once in the front
Rotary mowers are the most common. They have at a higher height and once in the back at a lower
single blades mounted horizontally on a spindle height. Not only will the mower have to work harder
underneath the mower deck that cut as the blade to cut grass when the back of the deck is lower, the
rotates. All mowers have to be treated with respect grass will be injured twice with each pass.
to avoid accidents and potentially serious injuries, Many rotary mowers are mulching mowers with a
but the rotary mower is the most dangerous of the blade design that keeps the clippings under the deck
three. It is reasonably easy to slip or otherwise acci- for multiple cuts. Therefore you need to sharpen
dentally extend a foot under a rotary mower deck more than just the tip of the blade but the initial cut-
and it is common for debris to be thrown from under ting action still occurs at the tip.
the deck as the blade hits it. Large rotary mowers You might also notice that when you are using a
have multiple blades mounted under large decks to rotary mower with multiple blades under a wide
mow wide patterns and finish large areas quickly. deck and you are traveling too fast while you turn,
It is interesting to note that the blade under a rotary the outside blade cuts more raggedly than the
mower deck is not sharp across its entire length. In inside blade and the area looks uneven. For one
fact, the very tip of the blade is all that needs to be reason, the physical forces of the turn cause pres-
sharp to cut grass with a rotary mower at slow speed. sure on the outside of the deck pushing it down
High speed mowers may need slightly more blade lifting the inside of the deck up so the outside blade
sharpened because the rotational speed of the blade is cutting lower and the inside blade is cutting
may not be fast enough to cut with only the tip when higher. In addition, the outside blade has to travel
the mower is moving forward. Under normal circum- farther and cut more grass than the inside blade.
stances, the blade is rotating so fast and the forward Both blades have the same cutting (rotational)
speed of the mower is slow enough that only the tip speed. Consequently, the outside blade is expected
of the mower blade touches unmown grass as the to cut more grass at the same rotational speed and
mower moves forward. For the same reason, you can may not be able to keep up when the forward speed
adjust the mower deck one step lower in the front of the mower is excessive. Therefore, in order to get
than in the back to drop the mowing height approxi- an even finish, the appearance of the turns
mately half a normal step. You can do that because determines how fast you can drive.

Management Practices 65
responsible for the work of others. Take a quick insect invasion because it has to be mowed more
word of advice from a person who was an indus- often, creating chronic wounding and because it is
trial supervisor for many years before becoming a under more stress and is therefore more vulnerable.
professor. Your people are important. They are the Low mowing requires that irrigation be provided
only means you have of completing your assigned and that irrigation be more frequent because root
responsibilities. The saying “take care of your peo- systems are weaker. For that same reason, low mown
ple and your people will take care of you” is just as turf usually needs more nutrients and more frequent
true today as it was when it was first spoken by nutrient applications. Finally, low mown turf requires
someone many years ago. a good manager; a manager with agronomic knowl-
Your operators should be trained on the safe and edge and the ability to think through problems.
proper use of mowing equipment. That will elimi- Another factor to consider in mowing manage-
nate a lot of agronomic problems. They should be ment is the design of landscape features. This is a
trained to look for signs of mower wear and time and labor decision rather than an agronomic
improper operation. They should be alert to decision. This topic is discussed in Box. 5.3.
conditions in the field that relate to mower per-
formance and to overall turfgrass health. Each
5.3 Thatch and Thatch Management
piece of equipment should have a checklist assigned
to it (Fig. 5.5). The checklist should contain the Although our definition of a turfgrass does not
pre-use, mid-use and post-use responsibilities of the include thatch, thatch is unique to grasses and
operator, and should be quickly reviewed by the thatch control is unique to turfgrasses. Thatch is an
operator before each use and at the completion of important consideration for any turfgrass manage-
the assignment. ment plan. Many people seem to believe that thatch
Mowing equipment has to be kept in good repair has no benefit when, in fact, it is very beneficial to
and mower blades have to be kept sharp. Otherwise soil conditions in a long-term biological system.
agronomic problems will occur. A cut in your fin- The most fertile soils are those that have sustained
ger caused by a knife heals faster with less trauma grass for long periods of time, meaning hundreds
than a ragged cut made by a sharp stone. Turfgrass or thousands of years. Turfgrass managers tend to
cut with a sharp blade heals more rapidly than have a negative concept of thatch because many of
turfgrass cut with a dull blade and causes the least our turfgrass management practices lead to exces-
disruption to photosynthesis and the least desicca- sive thatch layers that have to be controlled.
tion. A sharp cut covers less surface area than a However, thatch is mostly a positive influence in
ragged one, so fewer cells are affected by the injury the natural environment and also provides some
(Fig. 5.6). Consequently, a sharp cut exposes less benefits to managed turf.
surface area to air, causing less desiccation and less Thatch is a combination of dead and living plant
browning of the leaf tip. Mowing when the soil is material deposited by grass between the grass
wet may cause scalping as the mower sinks into the shoots and the underlying soil. It consists of stems
turf. Mowing during extremely dry periods should and roots and, rarely, leaves. As long as mowing
be followed by irrigation if possible to reduce leaf frequency is practiced according to the 1⁄3 rule,
tip desiccation. Remain within recommended mow- clippings degrade quickly and do not add to the
ing heights for the species being mowed. Refer to thatch layer (Soper et al., 1988). Thatch is actually
the textbooks suggested at the end of Chapter 1 for dead material, but living material has to grow
recommended mowing height by species. See the through or across the thatch to reach the soil and
suggested reading at the end of this chapter for consequently becomes a part of it. For that reason,
websites of major equipment manufacturers, where any definition of thatch should include living plant
considerable equipment information can be found. material. The type and amount of plant material
Remember that mowing is a stress. The lower you growing in the thatch differs depending on how
mow the more stress that occurs. Low-mown turf- thick the thatch layer becomes. If the layer of dead
grass requires more management and care than plant material becomes thick enough it can house a
higher mown turf within the range of species adapta- substantial amount of roots, stolons and sometimes
tion (Toler et al., 2007). Low-mown turf is more even crowns and rhizomes, and begins to cause
susceptible to desiccation because it has a shorter problems with turfgrass health (Ledeboer and
root system. It is more susceptible to disease and Skogley, 1967). In natural situations, this rarely

66 Chapter 5
 Equipment Instruction Sheet
Equipment Name: ____________________________

Equipment Number: ____________________________

Safety equipment required for operation of this machine:

 Hardhat
 Safety glasses
 Ear protection
 Pesticide personal protective gear

Pre-use procedures inside:

 Inspect for obvious damage, report to mechanic if damage observed
 Visually inspect tires for sufficient air
 Check the fuel level
 Check the oil level
 Check hydraulic fluid level if applicable
 Quickly inspect cables for signs of wear
 Quickly inspect hoses for signs of wear
 Sign out on equipment log

Pre-use procedures outside:

 Check where machine was parked for oil leaks or other problems
 Evaluate engine operation
 Does machine move forward and backward without any problem
 Check operation of blades, booms,buckets, bed lifts, as applicable

During operation:
 Operate safely according to your training on this machine
 Be alert for broken hoses, cables or equipment malfunctions no matter how minor
 Note any equipment problems both major and minor, return to shop if necessary

Post-use procedures:
 Quickly inspect cables for signs of wear
 Quickly inspect hoses for signs of wear
 Report major or minor malfunctions to mechanic
 Clean equipment according to training
 Park the machine in its assigned location
 Sign in on equipment log

Equipment Log
Equipment Name: ____________________________

Equipment Number: ____________________________

Out In
Operator Odometer Odometer
Date name Time miles/hours Time miles/hours Equipment status

Fig. 5.5. A sample checklist of requirements and operations that an operator should review and perform before and
after using a piece of equipment.

Management Practices 67
 (a) (b)

Fig. 5.6. A shredded leaf tip exposes more surface area to air immediately following mowing than a cleanly cut leaf
tip. The shredded leaf tip usually sustains severe damage. (a) This leaf tip was cut cleanly with a sharp rotary mower
blade. (b) This leaf tip was cut using a rotary mower with a dull blade.

occurs, but managed turfgrass systems commonly Turfgrass species, even cultivars, differ in their
require forms of thatch management to keep thatch propensity to produce thatch based on anatomical
layers thin enough to be beneficial rather than differences that affect the speed of microbial break-
detrimental. down (Stiff and Powell, 1974). In addition, the
Thatch is naturally degraded by saprophytic more aggressive grasses are often the ones that
microorganisms, so an environment that encour- thatch up most quickly (Shearman et al., 1980).
ages microbial activity speeds the degradation of Consequently, species and cultivar selection for
thatch. Theoretically, it should be possible to accel- your region is a viable method for maintaining
erate thatch degradation by inoculating turfgrass thatch at desirable levels. Management procedures
with microorganisms that feed on thatch. Research that encourage aggressive turfgrass growth tend to
performed many years ago suggested that inoculation result in rapid thatch accumulation. If the environ-
is a possibility for reducing thatch in turfgrass ment favors rapid plant growth and slow microbial
(Sartain and Volk, 1984). However, to date, a suc- activity, thatch increases rapidly. This all means
cessful program or commercial product that can that the faster your grass is growing, the more
accelerate thatch degradation consistently has not likely you are to have a thatch problem. The thick-
been found (McCarty et al., 2007). est, greenest grass is probably going to require

68 Chapter 5
 Box 5.3. Design landscape features to facilitate mowing.
By designing landscape features that are easy to and trimming time by 10 minutes per event results
mow around, you can save yourself a considerable in about a 5-hour time saving each year if you
amount of time and labor. Even in a situation as mow 30 times per year. Consider the examples
small as a single home lawn, reducing your mowing below:

All that you have to do to save yourself considera- paint or a garden hose then paint and mow along the
ble trimming time is to design your bed-to-lawn inter- edge, maintaining easy turning radii that roughly
face using your mower. Once you have determined follow the pattern. The mower determines the finished
the shape that you would like to install in the land- edge and you will save substantial mowing and trim-
scape feature, lay out the edges of the feature using ming time with every mowing event.

cultural management for thatch control. So let us

review some of the lessons that you have probably
learned about thatch in earlier classes, previous
reading, or on-the-job-training.
A reasonably thin thatch layer of approximately
0.5 inches (13 mm) on most turfgrass areas, or
about 0.3 inches (8 mm) on a golf course putting
green, helps to cushion turfgrass crowns and roots
from traffic damage (Fig. 5.7). Thatch heats and
cools rapidly, but as long as plant roots and crowns
are growing in the soil instead of the thatch, the
thatch layer actually provides an extra layer of
insulation that helps prevent the rapid heating and
cooling that is stressful to biological systems.
Thatch is hydrophobic (water repellent) when it is
dry but it is also porous (Hurto et al., 1980). As a
result, a thin layer of thatch is not particularly
detrimental to water infiltration and is unlikely to
Fig. 5.7. The soil profile of a sand putting green. The
cause a significant increase in water runoff. Once
thatch on this green is very thin, but the mat layer, the
thatch is partially degraded to humus, it becomes a
mixture of thatch and soil below it, is very thick. During
somewhat stable part of the soil system and summer, the roots of this grass probably do not extend
provides exceptional benefits. below the mat. If you were responsible for managing
The richest, most fertile soils appear dark this green what changes would you make to the
brown, even black, as a result of high levels of previous management plan?

Management Practices 69
organic material that has been degraded to humus, more protection for turfgrass crowns and roots,
a semi-stable complex of organic compounds in and mat retains more water and nutrients than
colloidal form. Soils high in organic material that thatch. Because water is slower to evaporate from
was once thatch have high cation exchange capaci- mat than from thatch and because mat is less water
ties, meaning that they are capable of holding a lot repellent, mat is likely to retain plant-available
of nutrients (Carrow et al., 2001). These soils also water. Because water is slower to heat and cool,
hold a considerable amount of plant-available mat has better insulating properties than thatch.
water, yet are highly aggregated to provide larger Most importantly, degradation occurs more rapidly
air-filled pores (Brady, 1990). It is the thatch and in mat than in thatch.
other organic compounds in various levels of deg- As mentioned earlier, management methods that
radation that bind clay soil particles together to increase microbial activity reduce thatch by encour-
form aggregates. These aggregates are the basic aging degradation of mat to humus (Berndt, 2008).
units of soil structure. They contain small pores It stands to reason that management methods that
that hold plant-available water and are separated encourage aggressive turfgrass growth would lead
by larger pores that drain readily to provide soil to more plant biomass, hence more dead biomass,
air. The oxygen in this air is used by plant roots in hence more thatch. High applications of nitrogen
the respiration process. Respiration provides fertilizer have been linked to increases in thatch
energy for water uptake. This water provides the (Soper et al., 1988). However, multiple research
cooling medium for the transpiration mechanism projects have demonstrated that nitrogen rate and
and provides nutrients for plant growth and nitrogen source do not affect thatch (Dunn et al.,
development. 1981; Carrow et al., 1987; Johnson et al., 1987;
Although a relatively thin thatch layer has many Hollingsworth et al., 2005). From these conflicting
benefits, an unreasonably thick thatch layer, results, it appears that nitrogen can have an effect
approximately 1 inch (25 mm), is highly detrimental on thatch accumulation but, as usual, it is not the
(Fig. 5.7). That is why thatch generally is viewed as only factor that matters. Turfgrass management
a negative influence by turfgrass managers. In truth, and turfgrass growth and development rarely have
before its degradation, the thatch layer has not much simple relationships. There are nearly always mul-
to offer a turfgrass plant other than a little cushion tiple factors involved. In this case, the acidity of the
from traffic and some insulating value for crowns soil is also a factor that influences thatch develop-
and roots. The primary advantages of water and ment (Sartain, 1985). Soil that is too acidic or too
nutrient retention, along with soil structuring, do basic can slow microbial activity. Soil and thatch
not take place until the thatch has been or is well that is too wet or too dry affects microbial
into the process of degradation to humus. As degra- respiration in the same way that it affects root
dation occurs, a layer of partially degraded thatch respiration. Too little oxygen or too little water
mixed with soil called mat forms below the thatch slows degradation. Temperature affects microbial
layer (Williams and McCarty, 2005). The mat layer activity as well. As you learned earlier, the speed of
has much higher nutrient and water retention prop- chemical reactions increases geometrically with
erties than those that occur in thatch. Consequently increasing temperature. The increases in microbial
turfgrass managers tend to view mat more positively activity occur at about the same rate as increases in
than thatch. They might even call it beneficial. chemical reaction rates up to about 100 °F (38 °C).
In our business, mat layers are more likely to be Consequently, there are times when grasses are
man-made than natural. Natural forces such as growing exceptionally well but microbial activity is
earthworm activity, burrowing insects and water or slow, and times when microbial activity is rapid but
air deposition result in the mixing of dust, soil or turfgrass growth is slow. Cool-season grasses, for
water-borne silt deposits with thatch, resulting in instance, grow best at about 65 °F–80 °F (18 °C–27 °C).
mat. However, turfgrass managers also cause mat At those temperatures microbial activity is relatively
formation by adding topdressing materials to turf. slow, but at 100 °F (38 °C) microbial activity is
By spreading a thin layer of soil or sand that closely rapid and cool-season grass growth is slow or
matches the soil at the site into the thatch, the man- nonexistent.
ager can encourage the formation of mat from Turfgrass managers often feel that they are prac-
thatch. Mat has all of the advantages of thatch and ticing aerification and topdressing to prevent thatch
more. It is more resilient than thatch so it provides from occurring. That is certainly not the case.

70 Chapter 5
Aerification and topdressing are practiced to
increase thatch degradation, not decrease its occur-
rence. In situations where turf is managed for high
expectations, like golf greens, bowling greens or
professional athletic fields, thatch is a part of the
process. It can’t be helped. If you are managing the
turf for high aesthetic and functional value, includ-
ing good health and rapid recovery from injury,
thatching is going to happen. A certain amount of
thatch will occur and thatch management will have
to be a part of your management program. Turf
that is poorly maintained or not growing well does
not produce excessive thatch. Only turf that is
managed for good health and rapid growth pro- Fig. 5.8. Vertical mowers have rigid blades something
duces excessive thatch. If you are practicing quality like saw blades. This power rake has blades mounted
thatch management, but the turf is still producing on hinges so that it can cut deeper with less chance of
injury to the operator or the equipment if a blade hits a
excessive thatch, it is time that you consider your
rock or a similar hard object.
fertilization, irrigation and other management
practices to slow the production of thatch. However,
under a high-quality maintenance program, thatch destructive nature of vertical mowing for thatch
will be produced but controlled through practices removal is too destructive to be performed frequently
that encourage rapid microbial degradation. (Hollingsworth et al., 2005).
As you probably know, there are three practices Research suggests that core aerification may be
that we use to manage thatch: dethatching, which is more or equally effective for thatch management as
most often called vertical mowing, aerification and vertical mowing and it is usually less destructive
topdressing. Dethatching is a process of thatch (Murray and Juska, 1977). Core aerification
removal but also has some influence on microbial improves the environment for microbial growth
activity. Aerification is a process that encourages and reproduction in the same manner that it
microbial activity but also has some thatch removal improves the environment for root growth.
characteristics, and it is used to relieve soil compac- Aerification allows the carbon dioxide produced by
tion and may also improve water infiltration plants and microbes during respiration to escape
(Murphy et al., 1993a; McCarty et al., 2007). the soil and encourages the entry of fresh air con-
Topdressing is primarily used to promote microbial taining oxygen. Consequently, thatch and mat
activity, but it is also used to help relieve compaction degradation improve. When core aerification is
following aerification, to smooth and level the turf/ used on a golf course putting green or other highly
soil surface, or to modify soil characteristics. managed area the soil cores are usually removed
Dethatching is accomplished using either a vertical and the holes in the soil are usually filled with top-
mower or a power rake (Fig. 5.8). The only real dif- dressing material that is applied in an amount
ference between these is that vertical mowers have slightly greater than that required to fill the holes.
rigid blades and power rake blades work on hinges. This topdressing procedure effectively mixes top-
You might even see a dethatching device that uses dressing material with thatch, forming mat.
vertical chains or something similar to dig out Topdressing appears to be the best of the thatch
thatch. The purpose is the same regardless of the management techniques (Carrow et al., 1987;
equipment used, and that is to remove existing Johnson et al., 1988), and four applications per year
thatch. In doing so, the mowing device has to cut are better than one (White and Dickens, 1984). In
into the top of the soil and consequently may home lawns, parks and other medium-maintenance
positively affect aeration and water infiltration. turf, cores are usually left on-site and allowed to
Vertical mowing, as this process is most commonly reincorporate naturally. Leaving the cores to incor-
called, regardless of the equipment used, was once porate naturally is a slightly less effective form of
the most commonly recommended process for topdressing. Topdressing is believed to provide an
thatch control. Deep vertical mowing for thatch environment more conducive for microbial thatch
removal, however, is extremely destructive. The degradation than a typical thatch layer. Although

Management Practices 71
topdressing alone is a fairly effective form of thatch and smaller pores holding air, there is limited soil
control, the combination of light vertical mowing, air exchange with the atmosphere, resulting in the
aerification and topdressing has proved most effec- same effect on root respiration that a thick bound-
tive for managing thatch on golf course putting ary layer resulting from poor air movement causes
greens (McCarty et al., 2007). for leaf photosynthesis. As roots and soil microbes
respire, oxygen in the soil air is depleted and the
carbon dioxide concentration increases. High car-
5.4 Issues Pertaining to Soil
bon dioxide concentrations in soil air can cause
Compaction
reductions in root length and mass (Bunnell et al.,
Turfgrass and turfgrass soils usually have to sustain 2002). As normal air is about 21% oxygen, it is not
moderate-to-intense traffic. Turf most certainly has likely that oxygen concentration would become
to sustain a certain amount of mowing traffic, but low enough in the soil air to affect respiration.
it is usually also required to sustain considerable However, compaction results in more capillary
foot traffic and sometimes vehicle traffic. This traf- water and considerably less soil air. Although the
fic results in soil compaction, a compression of the concentration of oxygen in the air probably remains
soil that destroys its structure and compresses large sufficient, the availability of air in general could
air-containing pores, converting them to smaller become a limiting factor for root and microbial
water-containing pores. Compaction has a detri- respiration. That would not only affect root growth,
mental effect on root growth by reducing root it would affect also microbial activity, resulting in
respiration. As respiration declines, transpiration is less thatch degradation. Multiple research studies
affected. A compacted soil is also dense, making it have indicated that turfgrass water-use rates decline
difficult for roots to grow through it. The concept in compacted soil (O’Neil and Carrow, 1982,
of constructing playing fields of specialized sand 1983; Agnew and Carrow, 1985).
systems resulted from the poor turf performance
and poor drainage of compacted soils. Pure sand
5.5 Managing Wear Caused by Traffic
has no structure and, unless it is extremely fine, is
unlikely to compact to the point that turfgrass Traffic on turfgrass systems not only causes com-
growth is affected. paction, it also causes damage due to wear. Research
Aerification is a term that refers to the multiple on creeping bentgrass (Agrostis stolonifera) and
processes used to help relieve soil compaction. velvet bentgrass (Agrostis canina) suggested that
Aerification includes coring, spiking, slicing, water wear from traffic on those grasses caused more
injection and other procedures that are used to damage than soil compaction (Samaranayake et al.,
fracture the soil, causing large channels that con- 2008). However, this research was performed on
tain air and drain water. Most soil compaction on sandy loam soil. Had it been done on clay loam,
turfgrass systems is surface compaction that usu- the compaction from traffic may have been the
ally does not extend deeper than 2–3 inches major cause of decline. The amount of damage that
(6–8 cm). For that reason, the most common aerifi- occurs from wear and compaction is not only influ-
cation practices are designed to primarily affect enced by the amount of traffic at the site, it is also
shallow soil layers. Aerification procedures lower affected by environmental conditions that deter-
soil bulk density, improve subsoil drainage and mine plant and soil status at the site. Saturated soil,
promote air circulation, which admits oxygen and for instance, does not hold up well to traffic
vents carbon dioxide. Aerification is also effective because the soil is easily shifted, causing roots and
for disrupting soil layers that interfere with water stems to fracture or break. Plant succulence
percolation and air exchange. increases with high fertilization and in shade.
As soil is compressed and its pores become Succulent plants are easily damaged because water
smaller, the number of soil pores that contain water does not compress and is likely to push out through
increase and the number that contain air decrease. plant tissue when the tissue is compressed. Although
Small soil pores hold capillary water, meaning that wear tolerance is affected by the major plant proc-
they bind water against gravity. You learned about esses, photosynthesis, respiration and transpira-
capillary action in Chapter 1. Larger pores are tion, species selection and simple traffic management
needed to drain the soil and it is the large pores that are also important considerations for managing
hold the oxygen for root respiration. With fewer turf under traffic.

72 Chapter 5
 Research has suggested that Kentucky blue- spread out over 3 days per week with only a day
grasses (Poa pratensis) that have more vertical or two of recovery time between events (Minner
shoot orientation, are less succulent and have and Valverde, 2005b). Managing traffic patterns
stronger cell walls are more tolerant to wear to permit the longest period between damaging
(Brosnan et al., 2005). Although this work indi- events is probably the most important mainte-
cates that cultivars differ in wear tolerance, it is nance practice for improving turfgrass quality
equally useful for selecting management practices under traffic.
to help relieve traffic stress. Alternating mowing
patterns to encourage upright growth, and apply-
5.6 Chapter Summary
ing only enough nitrogen fertilizer and water to
encourage recovery is good management to reduce Over the years, turfgrass researchers and managers
traffic damage. It is important that the grass cho- have developed cultural practices that help
sen for the site is well adapted for the environ- turfgrasses to survive and flourish in spite of
ment. A poorly adapted grass will not survive chronic stress. Species selected for turfgrasses are
traffic (Dunn et al., 1994). Both wear tolerance those that tolerate close mowing and can survive
and recuperative ability are important considera- soil compaction and wear. Our expectations for
tions when choosing a species for a site where these grasses are high. The grasses are highly
intense traffic is likely. However, it would be unu- adapted for most of the environments in which we
sual to find a turfgrass species that is both highly use them, but they seldom meet our expectations
wear tolerant and rapidly recovers from damage. unless we provide management. The development
Cool-season grasses with reasonably good wear of cultural practices and other management tech-
tolerance are perennial ryegrass (Lolium perenne), niques have made it possible for us to grow grasses
tall fescue (Festuca arundinacea) and Kentucky that meet our aesthetic and functional expectations
bluegrass (Shearman and Beard 1975a,b; Minner even in situations for which they are poorly
and Valverde, 2005a). Of the three, Kentucky adapted. The turfgrass must be healthy to tolerate
bluegrass recovers most rapidly so generally holds regular mowing and it must be healthy to resist and
up best in high traffic situations. Among the recover from traffic. Proper turfgrass selection and
warm-season grasses, zoysiagrass (Zoysia spp.) cultural management practices make it possible for
has outstanding wear tolerance, but most zoysia- turf to survive our expectations. We aerify, topdress
grasses tend to be slow to recover from damage. and vertical mow to improve water infiltration,
Bermudagrass (Cynodon dactylon) is usually the which encourages transpiration, and we apply soil
most effective warm-season grass for high traffic aerification, which encourages root respiration. We
areas (Duble, 1989). adjust mowing height and frequency to allow for
Traffic on turfgrass may cause reductions in adequate photosynthesis. In the future, it is likely
leaf water content, turf density and leaf chloro- that we will have to further adjust our practices to
phyll content (Han et al., 2008). These responses conserve more water and to further reduce fertilizer
suggest declines in photosynthesis, transpiration and pesticide applications. Techniques we develop
and possibly root respiration. High shoot density, for meeting our aesthetic and functional
a product of good photosynthesis, transpiration expectations, while reducing water, fertilizer and
and respiration management has been linked to pesticide use, will only be effective if they encourage
wear tolerance (Trenholm et al., 1999). Obviously, photosynthesis, respiration and transpiration.
management practices that encourage photosyn-
thesis, transpiration and respiration will affect Suggested Reading
the ability of a turf to recover from traffic dam-
age, so managing to encourage these practices Beard, J.B. and the United States Golf Association
(2002) Turf Management for Golf Courses. Ann Arbor
should improve performance. Turfgrasses should
Press, Chelsea, Michigan.
also be allowed sufficient time to recover between
Carrow, R.N. and Petrovic, A.M. (1992) Effects of traffic
damaging events. A traffic simulation study indi- on turfgrass. In: Waddington, D.V., Carrow, R.N. and
cated that intense traffic applied to Kentucky Shearman, R.C. (eds) Turfgrass. ASA-CSSA-ASSA
bluegrass weekly with 7 days recovery time (American Society of Agronomy-Crop Science Soci-
between damaging events caused less turfgrass ety of America-Soil Science Society of America),
decline than the same amount of weekly traffic Madison, Wisconsin, pp. 285–330.

Management Practices 73
McCarty, L.B. (2001) Best Golf Course Management Prac- Suggested Websites
tices. Prentice-Hall, Upper Saddle River, New Jersey.
Some major mower manufacturers
McCarty, L.B. and Miller, G.L. (2002) Managing Ber-
mudagrass Turf: Selection, Construction, Cultural Gravely (2009) Mowers and some other turf manage-
Practices and Pest Management Strategies. Ann ment equipment. Available at: www.gravely.com/
Arbor Press, Chelsea, Michigan. (accessed 5 June 2009).
Puhalla, J., Krans, J. and Goatley, M. (1999) Sports Husqvarna (2009) Mowers and most other turf manage-
Fields: A Manual for Design, Construction and ment equipment. Available at: www.husqvarna.com/
Maintenance. Ann Arbor Press, Chelsea, Michigan. (accessed 5 June 2009).
Rieke, P.E. and Murphy, J.A. (1989) Advances in turf Jacobsen (2009) Mowers and most other turf manage-
cultivation. International Turfgrass Society Research ment equipment. Available at: www.jacobsen.com/
Journal 8, 49–54. (accessed 5 June 2009).
Waddington, D.V. (1992) Soils, soil mixtures, and soil John Deere (2009) Turfgrass, crop production, construc-
amendments. In: Waddington, D.V., Carrow, R.N. and tion, irrigation and other equipment (includes video
Shearman, R.C. (eds) Turfgrass. ASA-CSSA-ASSA instruction). Available at: www.deere.com/ (accessed
(American Society of Agronomy-Crop Science 5 June 2009).
Society of America-Soil Science Society of America), Toro (2009) Mowers, irrigation and most other turf man-
Madison, Wisconsin, pp. 331–384. agement equipment (includes video instruction).
White, C.B. (2000) Turf Managers’ Handbook for Golf Available at: www.toro.com/ (accessed 5 June
Course Construction, Renovation and Grow-In. Ann 2009).
Arbor Press, Chelsea, Michigan. Wikco.com (2009) Mowers, including flail mowers and
Witteveen, G. and Bavier, M. (1998) Practical Golf Course most other turf management equipment (includes
Maintenance: The Magic of Greenkeeping. Ann Arbor video instruction). Available at: www.wikco.com/
Press, Chelsea, Michigan. (accessed 5 June 2009)

74 Chapter 5
 The Importance of Light
6 and Managing Shade

Key Terms
Irradiance is the combination of direct, diffuse and reflected sunlight (solar radiance) striking an object at any
particular time.
Direct solar radiance is the portion of sunlight striking an object that arrived on a straight line from the sun. Direct
sunlight constitutes the largest portion of irradiance.
Diffuse solar radiance is the portion of sunlight that has been scattered or diffused by atmospheric particles. Blue
light suffers the most diffusion, hence the sky is blue.
Reflected solar radiance is the sun’s energy that bounces from one object to another without being absorbed.
When we look at an object, it is the light reflected from that object that we see.
An aerosol is a suspension of fine solid or liquid particles in a gas. Aerosols are the cause of atmospheric diffusion.
Light quality is not the same as light quantity. Light quality is the spectral distribution of light striking an object. It
could be a very small quantity or a very large quantity but both quantities, large and small, could have the
same spectral distribution. In this chapter we are concerned about the proportions of blue, green, red and far
red that make up the irradiance that penetrates shade.
Light quantity is a measure of the amount of light that strikes an object. We are mostly interested in the amount
of light that strikes an area of turf, especially in the shade.
Reflectance is the portion of irradiance striking an object that is reflected from the object. The remainder of the
irradiance striking the object is either absorbed or transmitted. In our case, reflectance is the amount of
irradiance reflected from a turfgrass canopy divided by the amount of irradiance that is striking the canopy.
Because reflectance, absorbance and transmittance are all fractions of the total amount of irradiance
available, they are often expressed as a percentage of irradiance or a percentage of full sun.
Absorbance is the portion of irradiance striking an object that is absorbed by the object.
Transmittance is the portion of irradiance striking an object that passes through the object.
Morphological plant characteristics are those characteristics such as size and color that we can see. Some of the
most important turfgrass morphological characteristics are color, texture, density or cover, and uniformity.
Density and cover are not characteristics of turfgrass plants but they are characteristics of turf.
Physiological characteristics are characteristics that we can not see. It is the physiological processes that
determine morphological characteristics. You already know the most important physiological processes.
Those would be photosynthesis, respiration and transpiration.

6.1 The Practice of Making Adjustments

shade than it does in full sun. Under nearly all
to Improve Photosynthesis in Shade
circumstances, turfgrasses prefer full sun to shade
Light provides biological energy. Without light there and often will not grow satisfactorily in shade. In
is no life as we know it. Plants must have sufficient Chapter 2 you learned the basics of light absorption
light to provide the energy that fuels metabolism. and conversion to chemical energy. In this chapter
Some plants grow proficiently in low light but most we will expand on that knowledge and help you
do not. Although some turfgrasses tolerate low light understand how to use what you have learned.
better than others, it is a rare environmental situa- For the purpose of plant study and management,
tion when a particular turfgrass grows better in the solar spectrum is divided into six categories

©CAB International 2011. Turfgrass Physiology and Ecology (G. Bell) 75

called bands, ultraviolet, blue, green, red, near at any one time. Those terms may be new to you
infrared (NIR) and infrared. The ultraviolet band is and require explanation. If you were to look
high-energy light that can be destructive to plants directly at the sun, not a good idea, your eyes
and to us. Ultraviolet radiation causes genetic muta- would be absorbing direct radiation. Direct radia-
tions and tissue destruction but there is little that we tion is very intense. However, it does not exist
can do to affect that, at least on our plants. Actually, outside a fairly straight line between an object, in
some of our top-performing grasses are the result of this case you, and the sun. If you were to stand
natural or artificial genetic mutation caused by directly behind a tall building when the sun was on
radiation. Otherwise the ultraviolet band is of little the other side of it, there would be no direct radia-
concern to us and we need not discuss it further. tion in that area. All of the direct radiation would
The infrared band has some potential for nonde- be blocked by the building. However, you would
structive assessment of tissue and/or soil water in turf still be able to see. Why can you still see?
but, as yet, has not been fully developed for that pur- The reason that you can see when you are on the
pose. As a source of energy it is important to us, but side of a building opposite the sun is because there
not in a positive manner. Infrared radiation degrades are two other kinds of irradiance at work. When
rapidly to heat. Much of the heat that builds in turf- you look at the sky on a clear day, it looks blue. It
grass leaves on bright summer days is a result of looks blue because there are aerosols – suspended
infrared radiation. Fortunately, most of the infrared fine particles or molecules – in the air that scatter
radiation that strikes a green plant is reflected and, blue light (Gates, 1966). A portion of the blue light
therefore, is inconsequential. That infrared radiation that is projected directly from the sun is detoured
which is absorbed quickly degrades to heat and is by diffusion throughout the atmosphere (Brine and
dissipated mostly through active transpiration, and Iqbal, 1983). Consequently, the atmosphere looks
partially by conduction and convection. blue. Diffuse blue light strikes the earth from all the
The light bands blue, green and red are impor- directions of the sky’s hemisphere (Smith, 1982). It
tant spectral bands for us to consider. The NIR, can strike an object in the open from any direction.
which, for the most part, is not considered light If you are standing on a lawn in deep shade behind
because we can’t see it, is also an important band. the building that we mentioned, the turfgrass that
The NIR is generally considered to extend from a you are standing on is absorbing blue light from
wavelength of 700 nm to a wavelength of 1350 nm any direction in which you can see sky (Fig. 6.1).
(see Chapter 2 to refresh your memory concerning Diffuse blue light is one of many reasons that some
wavelengths). However, there is a narrower portion turfgrasses can grow reasonably well in shade.
of the NIR from 700 nm to 800 nm that is more Reflected light is another, and this is the main rea-
useful for plant study. We call the band from son that you can see on the shaded side of a
700 nm to 800 nm the far red band and we can building.
refer to it as light. Depending on the authority you White or shiny objects reflect nearly all of the
would like to believe, we can see wavelengths of light that strikes them. We have trouble looking at
up to either 750 nm or 780 nm (Starr, 2000). those objects in bright sunlight because they hurt
Consequently, we can see half or more of the far our eyes. Direct sunlight reflected from a pure
red band, so we can loosely refer to it as light. The white object is almost as intense as direct sunlight
portion of the far red that we can see appears to us itself. When we look at such an object, all that we
as dark to very dark red. It is the blue, green, red see is blinding white light. Such an object is very
and far red bands that directly affect our turf. The rare. We usually see off-white objects and multiple
blue, green and red bands provide energy through colors of objects. Pure black objects, another rare
photosynthesis, and the red and far red bands are occurrence, reflect no light at all. We see them as a
absorbed by phytochrome, the pigment that helps hole in a field of colors. In other words, we don’t
to release or control many plant functions (Grant, really see pure black objects, we see where they are
1997; Nagy and Schäfer, 2002). based on the position of surrounding and
background objects that reflect light. We see most
objects because, other than pure black ones, they
6.2 The Influence of Irradiance
all reflect light. The types or wavelengths of light
Irradiance is the sum of direct, diffuse and reflected that objects reflect determine what color they are
solar radiance (sunlight) striking a particular object to us. Turfgrass is green. Turfgrass is green to us

76 Chapter 6
 Turfgrass color has always been and still is a valu-
able indicator of plant health. Experienced profes-
sionals know when their grass is too green and
when it is not green enough. In addition to its use
as a discriminatory tool, reflected green and other
colors of reflected light are sources of light for
photosynthesis. Reflected light is capable of pene-
trating shade because, like diffuse light, reflection
can occur from any direction. Shade occurs when
the direct rays of the sun are intercepted by an
object. However, direct irradiance is not the only
light available to turfgrass. In shade there may still
be sufficient diffuse and reflected light to maintain
a small amount of photosynthesis. For some plants
diffuse and reflected light is sufficient for growth.
Not so for turf: turfgrasses, even shade-tolerant
ones, require daily periods of full sunlight to
maintain growth. You will not find turfgrasses
growing directly under large trees or in the north
Fig. 6.1. Only a small portion of the direct rays of the inside section of U-shaped buildings. There is not
sun can pass through a tree canopy. Consequently, enough light.
there is shade on the side of this tree opposite the sun.
However, notice that there is an entire hemisphere of
sky surrounding this tree and consider that diffuse blue The amount of irradiance available in shade
light radiates from all points in the hemisphere. That
Three things can happen to solar radiation when it
diffuse blue light penetrates the shade and helps to
stimulate plant growth. strikes a plant leaf or any other object. The irradi-
ance may be reflected, it may be absorbed, or it
may be transmitted. The reflectance of an object (in
because it reflects more green light than any other this case a turfgrass canopy) is the portion of irra-
color that we can see and because our eyes are diance striking it that is reflected from it; the
more sensitive to green than to any other color. As remainder is either absorbed or transmitted. The
you learned in Chapter 2, red and blue light are absorbance is the portion of irradiance striking the
highly absorbed by chlorophyll. Some of the blue object that is absorbed by it, and the transmittance
and red light striking turfgrass is also reflected, but is the portion of irradiance striking the object that
usually only a small portion. Turfgrass reflects a passes through it. Very little light is transmitted by
much larger portion of the green light that strikes a plant canopy such as that of a tree (Fig. 6.2).
it than the red or blue light, hence the plants appear Although substantially more far red is transmitted
green. The high concentration of chlorophyll in by a vegetation canopy than are the other light
turfgrass leaves masks the color of other pigments. bands, the proportion of the far red transmitted
The xanthophylls make up the second largest compared with the portion that strikes the canopy
concentration of pigments in turfgrass leaves is very small (Bell et al., 2000). Very little of the
(Turgeon and Lester, 1976; McElroy et al., 2006). blue, green and red bands, and little of the far red
Consequently, when turf experiences stress and band, penetrate right through a plant canopy. No
chlorophyll begins to degrade, the plants turn light penetrates right through a building or other
yellow. Yellow is the color of the xanthophyll substantial structure. Consequently, shade from
pigments. When our turf begins to yellow, we take buildings, and for the most part shade from vegeta-
notice. We know that when we begin to see the tion, are substantially devoid of direct sunlight.
color of the xanthophylls, the plant’s chlorophyll is Turfgrass growing under shade conditions must
degrading and its photosynthesis is slowing. survive on diffuse and reflected light until a time
Something needs to be fixed. during the day when it can receive direct sunlight.
The greenness of turf has long been the factor If the period of direct sunlight is long enough, the
most commonly used to assess its health (Box 6.1). turf will survive. If the period of sunlight is longer

Importance of Light and Managing Shade 77

 Box 6.1. The role of green light in photosynthesis.
We generally refer to green light as light occurring in Using this calculation, as the amount of red
a band from a wavelength of 500 nm to a wavelength reflectance increases or the amount of near infrared
of 600 nm. That band is part of what we call photo- reflectance decreases, the NDVI gets smaller. A large
synthetically active radiation (PAR). However, NDVI indicates a green plant and a small NDVI
because plants are green and obviously reflect a indicates a yellow or brown plant. The same is true of
substantial amount of green light, there is a common GNDVI, an abbreviation for another vegetation index
misconception that plants do not use green light for called the green normalized difference vegetation
photosynthesis. That is simply not true. index (Shanahan et al., 2001). The GNDVI is exactly
Plants appear green because they reflect more the same calculation as the NDVI except that a green
green than any other color. Chlorophyll has peak light band is substituted for the red light band:
absorption points in the blue light range at approxi-
mately 410, 430, 455 and 460 nm, and in the red GNDVI = (RNIR − Rgreen)/(RNIR + Rgreen)
light range at approximately 640 and 660 nm (French,
1961). Therefore, chlorophyll absorbs more blue When the amount of green reflectance increases,
light and red light than it does green light. the GNDVI declines. A high GNDVI indicates a green
Consequently, the amount of reflected green light is plant and a low GNDVI indicates a yellow or brown
much greater than that of any other color that we plant. If chlorophyll reflected more green light than it
can see, so plants appear green. Green, however, is absorbed, the GNDVI would increase when NDVI
also quite active for photosynthesis in grasses. decreased, but it does not. A low red reflectance
Measures of plant reflectance can be used to means that substantial red light is being absorbed and
estimate plant health based on greenness (Bell and NDVI increases, a low green reflectance means that
Xiong, 2008). That concept was introduced in substantial green light is being absorbed and the
Chapter 1, as were vegetation indices. A vegetation GNDVI increases. Both a high NDVI and a high
index is a combination of two or more bands GNDVI indicate very green plants. If more green light
reflected from plants that when combined in a was being reflected from a green plant than was being
specific way provide an accurate measure of plant absorbed, the GNDVI would not work in the same way
greenness. The most common vegetation index as the NDVI. The GNDVI has been thoroughly tested
used is the normalized difference vegetation index on wheat (Moges et al., 2004), corn (Dellinger et al.,
(Rouse et al., 1973), which is abbreviated NDVI. The 2008), creeping bentgrass (Bell et al., 2004) and ber-
NDVI is calculated as the difference between near mudagrass (Xiong et al., 2007).
infrared reflectance (RNIR) and red reflectance (Rred) More green light is absorbed by chlorophyll than is
divided by their sum: reflected. Green light helps our plants assimilate
carbon as an active component of the photosynthesis
NDVI = (RNIR − Rred)/(RNIR + Rred) process.

than that minimally required for growth of the spe- the reflection of blue light. Blue light is highly
cies at the site, the turfgrass will prosper and reflected from objects that appear blue or white,
spread. Because little direct sunlight can penetrate and red is highly reflected from objects that appear
shade, trimming a tree canopy to allow more light red or white. For that reason, turfgrass may grow
to penetrate is not nearly as effective for turfgrass slightly better in the shade of a building that is
shade management as is strategically removing painted blue, red or white than a building painted
individual trees to extend the period of direct sun- a different color. I would not count on that as a
light (Bell and Danneberger, 1999). management practice though unless all other
We know that blue and red light is highly options were exhausted. Then again, turfgrass
absorbed by turfgrass leaves because they contain areas of athletic fields painted blue or red, espe-
high concentrations of chlorophyll. What you may cially red, may decline more rapidly than turfgrass
not know is that blue and red light is also reflected areas painted other colors. In that case, too much
from turfgrass. The bluish color of buffalograss of the most important photosynthetic light is being
(Buchloe dactyloides) and slight blue tint of reflected and is not available for absorption. If
Kentucky bluegrass (Poa pratensis) is caused by marking paints specially formulated for application

78 Chapter 6
 40
Shade Full sun

Photon flux (umol/m2/s)

30

20

10

0
330 380 430 480 530 580 630 680 730 780 830
Wavelength (nm)

Fig. 6.2. A tree canopy such as the one under which these data were collected substantially restricts the penetration
of direct sunlight.

to turfgrass are used, however, blue and red colors higher the reflection of far red that occurs (Gausman
should not be a major problem. Green light is et al., 1969; Maas and Dunlap, 1989). As our
highly reflected from turf, but a substantial amount vision is not very sensitive to most of the NIR
of it is also absorbed for photosynthesis. Because band, its reflection has little influence on the color
of that, reflected green light can be used as a meas- of turfgrass. However, its ability to penetrate veg-
ure of chlorophyll concentration and potential etation canopies at a greater rate than blue, green
photosynthesis. Contrary to popular opinion, the or red light is a point of major concern. We will
greenest turf absorbs the most green light. Green spend considerable time on that phenomenon dur-
reflectance actually declines as turfgrass gets more ing the discussion on light quality. For now,
green (Shanahan et al., 2001; Bell et al., 2004; remember that phytochrome absorbs irradiance in
Moges et al., 2004; Xiong et al., 2007; Dellinger a band centered near 730 nm in the far red and
et al., 2008). Reflectance is not an amount of another band centered near 660 nm in the red
reflection, it is the proportion of available (Grant, 1997).
irradiance that is reflected, so although the turf
turns greener owing to reflected light, it is also
The quality of irradiance available in shade
absorbing a larger portion of the total green light
available. The turfgrass turns more green because With a little instruction, light quality is reasonably
the grass is also absorbing a greater portion of the easy to understand. Turfgrass does not grow well
available blue and red light, allowing the green in shade because there is not enough light for it to
reflection to be more dominant. In simple terms, perform adequate photosynthesis. However, that is
the more green your turf, the more light that is not the only light-related factor that causes
being absorbed. turfgrass to decline in shade. Light quality is also
The far red band is transmitted through vegeta- a consideration.
tion at a higher rate than the other light bands and Light quality is nothing more than the amount of
it is also highly reflected (Bell et al., 2000). each spectral band of interest in a given amount of
Substantially more far red is reflected than blue, irradiance (Fig. 6.3). As turfgrass managers, the
green or red, and little far red is absorbed. The light bands of interest are four: blue, green, red and
reflection of far red light presumably has little or far red. Actually, these bands account for less than
nothing to do with pigments, and is believed to be half the total photon flux in sunlight, but they are
strongly influenced by cell structure (Knipling, the only bands known to have a major influence,
1970). It has been suggested that the smaller the except for genetic mutations and heat, on plant
cells in the leaf blades, the higher the proportion of growth (to refresh your memory on photon flux,
cell walls compared with cell contents and the see Chapter 2).

Importance of Light and Managing Shade 79

 Full sun
Blue
we cannot see often makes plant study easier to
Far red
27%
19% understand. The things that we can see, the mor-
phological effects, are important indicators of the
things that we cannot see, the physiological effects.
Turfgrass management has historically, and may
Red
Green always, be based on what our plants look like at
26%
28% any given time combined with our knowledge of
Shade
Blue
recent environmental conditions and our knowl-
Far red 24% edge and experience of typical plant responses to
32%
those conditions. In short, if our turfgrass looks
good morphologically, it is probably in good shape
physiologically.
Green
Red 24%
Many of the responses of turfgrass to shade result
20%
from phytochrome activity (Box 6.2). As you prob-
Fig. 6.3. The spectral distribution of irradiance
ably recall, phytochrome is a pigment that absorbs
determines its light quality. Irradiance in full sun, for red and far red light (Furuya, 2004). Another pig-
instance, contains more red light than it does in shade ment called cryptochrome absorbs blue light (Lin
from vegetation. The chart on the left depicts irradiance and Shalitin, 2003). Cryptochrome is a relatively
containing only blue, green, red and far red spectral recent discovery compared with phytochrome and its
bands, and their typical proportions in full sun. The activities are not well known. It is unlikely that cryp-
chart on the right displays the light quality typical of tochrome influences turfgrass growth in shade, but
shade from vegetation. we do know that the cryptochrome or some other
blue light-absorbing pigment must be present for
Wilkinson, Beard and Krans conducted land- stomates to open (Shimazaki et al., 2007).
mark shade research and identified many of the Cryptochrome has also been linked to phototropism,
morphological and physiological responses of turf- and both phytochrome and cryptochrome have been
grass to shade (Wilkinson and Beard, 1975; linked to photoperiodism and circadian rhythms in
Wilkinson et al., 1975). Some of these effects are plants (Christie, 2007; Bae and Choi, 2008).
caused by low light but many are caused by The ratio of red to far red light in sun and shade
changes in light quality. These and other turfgrass influences phytochrome status. With the absorption
shade responses are summarized in Table 6.1. In of red light, phytochrome enters an activated state
truth, physiological and morphological effects can called Pfr. The Pfr state is sensitive to far red light
not be separated as easily as this table implies. and enters the inactivated state called Pr when it
Morphological effects are the result of physiologi- absorbs far red light. At any given time within a
cal effects which are the result of chemical reac- single turfgrass plant, a certain portion of the
tions caused by plant response to environmental plant’s phytochrome is in an inactived state (Pr) and
stimuli predetermined by genetic code. However, the rest is in the activated state (Pfr). Under normal
separating physiological effects and morphological conditions, phytochrome reverts to the inactive (Pr)
effects into things that we can see and things that state at night when light is not present and is inac-
tivated during the day in proportions roughly equal
Table 6.1. Turfgrass morphological and physiological to the proportion of far red to red light in the irradi-
responses to the low light and poor light quality in ance striking the plant (Fig. 6.4). The amount of
shade from vegetation. light present during the day is not particularly
important for phytochrome-mediated reactions, as
Physiological effects Morphological effects
phytochrome-mediated reactions are primarily
Decreased photosynthesis Thinner, more delicate influenced by the balance between red and far red
leaves irradiance, a characteristic of light quality rather
Decreased carbohydrate Increased succulence than the amount of light present (light quantity).
reserves The diurnal qualities of phytochrome enable
Decrease transpiration Vertical shoot growth plant processes, such as flowering, that respond to
Changes in pigment Reduced root growth
photoperiod. Phytochrome enables the regulation
concentrations
of other processes, such as shade responses, based

80 Chapter 6
 Box 6.2. The difference between shade from a building and shade from vegetation.
If you have ever seen bermudagrass (Cynodon activity to grow upright quickly and expends a large
dactylon) growing on the north side of a building proportion of its energy to produce rapid shoot
where it only gets full sun for 1 to 2 hours in the growth at the expense of root growth. In the absence
morning and 1 to 2 hours in the evening you might of mowing that reaction would be a positive one. In
have asked how it could possibly grow in that much the presence of mowing, however, it is a negative
shade. reaction. Because of mowing, the turfgrass cannot
It is possible because the grass is able to allocate grow the taller shoots that would be more satisfac-
resources better in the shade of a building than it tory for performing additional photosynthesis. As
can in the shade of a tree. Some of the far red light soon as the shoots begin to reach a more satisfac-
striking the tree penetrates the canopy but none can tory height, we mow them off. Therefore, the plants
penetrate the building. For that reason more phyto- in the shade of a building have an advantage
chrome is inactivated in the shade of a tree than is because the light quality is different enough for less
inactivated in the shade of a building. The turf in the energy to be allocated to upright growth and so
shade of the tree is encouraged by phytochrome more is available to grow roots and stems.

on the amount of phytochrome inactivated during upright than normal as if they were trying to
daylight. In other words, phytochrome is an out-compete their neighbors and reach the sun.
indicator of light quality. You learned earlier in this Photosynthesis is severely limited in shade,
chapter that shade, especially shade from vegetation, consequently less than normal energy is available.
has a higher proportion of far red light and a lower The situation results in thinner leaf blades and
proportion of red light than full sunlight. thinner cuticles as the turf distributes most of its
Consequently, phytochrome tells our turf when energy to rapid upright growth. This shade response
shade is present. The turf responds by partitioning is common in plants. For instance, a tree, grown in
a higher portion of its total energy into shoot full sun will be shorter and have a thicker trunk
growth with less going to root and stem growth. than the same tree grown in shade. Under natural
Shoots are stimulated to grow rapidly and more conditions, this redistribution of energy is a benefi-
cial response. Ecologically speaking, the strongest,
in this case the most shade-resistant individuals,
Pfr
will garner the most sunlight and perpetuate the
species. Theoretically, if some individuals survive,
Pr the species will become more shade tolerant.
However, this naturally beneficial shade response is
highly detrimental to mown turf.
Day

6.3 Managing Turf in Shade

Consider how the redistribution of energy in
Night response to shade works against a grass that is
mown regularly (Fig. 6.5). As the plant is attempt-
ing to grow shoots rapidly, we are removing leaf
material by mowing. Not only does our grass have
Pr to regrow that shoot material removed by mowing,
it must also direct energy to heal the mowing
Fig. 6.4. Most of the phytochrome in a plant reverts
injury. The period immediately following mowing
to the inactive form (Pr) overnight. During the day, a is usually accompanied by a period of shock that
portion of the phytochrome is then activated to Pfr results in very low metabolism. The shock period is
according to the balance of far red and red irradiance followed by a period of slow growth as the plants
striking the plant. repair the mowing injury. Following this period of

Importance of Light and Managing Shade 81

 Sun Consequently, we must provide an artificial advantage
Stolon
to all of our plants if we expect them to survive and
growth Shoot grow reasonably well under shade and mowing. One
growth of the most effective means of increasing shade toler-
ance in turfgrasses is to increase their photosynthetic
leaf area by raising the mowing height.
Root When we increase mowing height, we increase
growth
the potential for sufficient photosynthesis to occur.
Shade Increasing the mowing height even a small amount
Shoot
growth can increase photosynthesis substantially (Fig. 6.6).
Raising the mowing height on a golf course putting
Stolon green by as little as 0.031 inches (0.8 mm), from
growth
Root 0.125 inches (3.2 mm) to 0.156 inches (4.0 mm)
growth increases photosynthetic capacity by 25%. Raising
the mowing height on a home lawn from 2.0 inches
Fig. 6.5. Phytochrome signals grasses and other plants
(50 mm) to 3.0 inches (75 mm), increases its photo-
to redistribute energy to encourage rapid vertical shoot
growth when the plants are exposed to shade. This synthetic capacity by 50%. If we mow shaded turf
naturally beneficial response is detrimental to mown turf at the same height that we mow the same species in
because each time meaningful shoot growth occurs, we full sun, we are not giving our shade turf much
mow it off and the plants have to start all over again. In chance of survival.
this situation, the small amount of photosynthetic energy The interesting effect of shade versus sunny con-
available in shade is perpetually partitioned to shoot ditions on the speed of a putting green is outlined
growth, causing a rapid decline in stems and roots. in Box 6.3.

slow growth, the plants can again direct most of

Managing air movement in shade
their energy into shoot growth. However, previous
growth progress has been removed, and the plants The detrimental effects of restricted air movement
are, in essence, beginning over again. have already been discussed in Chapter 2. However,
because restricted air movement and shade
commonly exist together, air movement deserves
Mowing in shade
another mention here. The effects of restricted air
In a natural environment, where mowing did not movement and its accompanying boundary layer of
occur, the individual grass plants best adapted to oxygen-rich, carbon dioxide-poor air can be as
shade would eventually out-compete their neighbors detrimental to turfgrass growth as shade (Koh et al.,
for available light. As the neighbors declined, more 2003). When restricted air movement and
space and nutrients would be available to the best
adapted plants. The tallest, best adapted plants
would further shade their neighbors and as their
neighbors died, more light would be available to the
+50% Ps
best adapted plants. In addition, the leaf area that
Photosynthesis (Ps)

+25% Ps
the adapted plants accumulated during rapid upright
growth would increase their ability to gather light
and perform photosynthesis. Consequently, they 3.0
3.0 in.
in.
2.5 in.
would eventually have more energy available to 2.0 in.
75 mm
75 mm
partition into root growth and stem growth. 51 mm
64 mm

Eventually, their energy distribution would return to

one more conducive to overall growth. That distri- Low Medium High
bution would be more similar to the distribution Mowing height
usually present in full sun. When we mow, we
remove that intraspecies competition. Under mown Fig. 6.6. A comparison of mowing heights. Notice that
conditions, the plants best suited for rapid vertical even a slight increase in mowing height results in a
shoot growth no longer have a natural advantage. substantial increase in potential photosynthesis.

82 Chapter 6
 Box 6.3. Which putting green is faster, one in full sun or one in shade?
Most people would probably say that a putting green as a golf ball rolls across the surface it is touching
in shade is slower than one in full sun because the more leaf tips and fewer leaf blades. The tips have
one in shade stays wetter than the one in full sun. That less surface area than the blades so they cause
is not correct. The green in shade is usually wetter, but less resistance to the motion of the ball. In addi-
it is not slower. In fact, the green in shade is faster and tion, the grass is nearly always less dense in
you can prove that to yourself with a stimpmeter, an shade, which causes even less resistance to the
instrument specifically designed to measure putting motion of the ball. Therefore, shaded greens are
green speed, and multiple trials on greens that do not usually faster than sunny greens, and a slight
differ except for their location in either shade or sun. increase in mowing height does not affect their
The leaf blades on a shaded green are growing playability in comparison with the other greens on
more upright than those in full sun. Consequently, the course.

shade exist together, the combination causes an root systems in shade are usually shorter and may
environment sufficiently detrimental to turfgrass need more frequent irrigation, but a lower amount
growth that it is very difficult to overcome, even of water is required compared with the same
with shade-resistant grasses. turfgrass in full sun.
The buildings and vegetation that cause shade Although a turfgrass requires less water in shade
often cause restricted air movement as well. Reducing than in full sun, it may have to be watered more
or eliminating either the shade component or the under trees or other vegetation. Turfgrass in shade
air-restriction component is sometimes sufficient to is often competing for water with the trees or other
promote turfgrass growth. As it is usually easier to vegetation creating the shade. Then again, if the
encourage air movement rather than to remove a shade is caused by a building or some other struc-
shade source, the recommendations for managing ture, root competition does not occur. Considering
air movement presented in Chapter 2 should always all of these possibilities, and how they interact with
be followed on shaded turfgrass sites. Start by trim- each other and with climate and weather, causes
ming tree branches to a height of at least 10 feet great difficulty when trying to determine how
(3 m) from the ground. Remove all low-growing much water is required by your turf on any particu-
vegetation, including ornamental plants. In wooded lar shaded site. There is no standard requirement
and other areas where debris tends to accumulate, and these choices are far from easy.
try to keep the surface of the ground free of this litter Let us consider the needs of the turf in a shaded
to allow good air flow. Pay particular attention to situation. Shade affects all three of our most impor-
both the leeward and windward sides of the site that tant functions. Photosynthesis is reduced in shade
correspond with the regions of predominant wind compared with full sun because less light is availa-
direction. Finally, remove or level hills and valleys ble. The shade environment is cooler than that of
that affect air flow. Good air movement encourages full sun, consequently, respiration is slower. Because
all three of our most important processes: photosyn- a shaded site is cooler and often includes air restric-
thesis, respiration and transpiration. tion, evaporation and transpiration decline. However,
because photosynthesis and respiration proceed
more slowly, creating less heat, and because less light
Managing irrigation in shade
is present to create heat, less transpiration is needed,
Irrigation management is as much an art form as it and because transpiration is by far the greatest use
is a science. There are so many variables to consider of turfgrass water, less water is needed. How much
with irrigation scheduling that it often becomes a less water is needed, however, is a very difficult
trial and error pursuit. Irrigation scheduling is question to answer. I suppose that if we were dealing
affected by many factors, including shade. Plants in with similar soils, similar climate, similar weather,
shade grow more slowly and need less water. In the same turfgrass species and the same type of
addition, shade sites are cooler and evaporation is shade at most turfgrass sites, we could study the
slower. Consequently, less water is needed. Turfgrass problem and find a reasonably good irrigation

Importance of Light and Managing Shade 83

determination based on the amount of time that the under stress. It requires more care than a plant in
turf is in full sun and in shade during each day. full sun and it is highly susceptible to drought stress
However, these conditions are rarely, if ever, the in its weakened condition. Consequently, long peri-
same and hardly ever similar. For that reason, there ods between irrigation events will harm the plant
are no valid recommendations for irrigating turf- rather than encourage it to grow a more extensive
grasses in shade. Turfgrass plants in shade require root system. Plants with weak root systems require
less water, but because of root competition from frequent irrigation. In shade, it is not advisable to
other plants they often require more irrigation than allow more than the top 1 inch (25 mm) of soil to
the same plants in full sun. In many cases, we have dry before the next irrigation event. Because shaded
to irrigate trees or other vegetation as well as our sites exhibit poor drying conditions, this may take
turf. Otherwise our turf will not get enough water. 3 days or more, whereas in full sun, it would nor-
Healthy turfgrasses have excellent root systems. mally occur in 1 day. Severe root competition,
Compared with overall plant size, most turfgrass however, will shorten this period considerably.
root systems are stronger than those of most trees. When attempting to grow turfgrasses on shaded
The plains in North America, South America and sites it is often necessary to irrigate lightly but often.
Africa, for instance, are covered with grasses but Tree root competition is an important factor in
do not have sufficient water for trees except in determining how often and how much irrigation is
lowland areas or near rivers and streams. Trees are required. Tree roots can usually be restricted from
huge plants, however, and a shaded turf has diffi- important turf sites using a technique called root
culty competing with the shear size of the root pruning. Root pruning is a simple process of cutting
system of such a large plant. Because turfgrasses tree roots between the tree and the turf you intend
are best adapted to full sun, they have a difficult to protect. It is normally accomplished by digging a
time competing with trees for water in shade. If we ditch, but in some cases can be accomplished using
hope to grow turf in the shade of a tree, we have to machines with thin knives or saws designed to cut
provide sufficient water to satisfy the tree before through tree roots in the upper 2 to 3 feet (60–90 cm)
we can provide sufficient water for the turf. Again, of the soil. As long as the pruning takes place no
there is no standard calculation to determine how closer that 10 feet (3 m) from the trunk of the tree,
much water is needed. It depends on the species of and on only one side of the tree, the tree will not
turf, the amount of shade, the type and size of the sustain lingering damage. In order to be effective,
tree and all of the other factors mentioned in the this process must be repeated once every 3 years or
previous paragraph. Regular observations and so as the tree roots tend to grow back fairly quickly.
probably some trial and error will be required to Because this process is time-consuming and expen-
determine the irrigation needs of such a site. sive, it is usually only used on areas of high-value
Irrigation frequency is also a consideration on turfgrass, such as bowling greens, tennis courts, or
shaded sites. Because shoot growth is favored in golf course greens and tees.
shade, turfgrass root systems are usually weak. The Growing turfgrass in shade takes knowledge and
root system of a tall fescue plant (Festuca arundi- care. Irrigating that turf requires redundant obser-
nacea) mowed at 3.0 inches (76 mm) may have a vation. Turfgrass quality and soil moisture should
root system that is 12 inches (30 cm) deep in full be checked daily during dry weather until the man-
sun, but that root system may be less than 6 inches ager is confident that the situation has been prop-
(15 cm) in shade (Wherley et al., 2005). The shorter erly assessed and that an acceptable program has
turfgrass root systems generally found in shade been established.
require more frequent irrigation than the deeper
root systems that are usually present in full sun. In
Managing fertilization in shade
most cases, it is advisable to allow the top 1 to 3
inches (25–76 mm) of soil to dry between irrigation Turfgrass grown in shade requires very little ferti-
events. If a grass plant is healthy and growing lizer (Baldwin et al., 2009). In fact, it is probably
conditions are conducive, a more extensive root better to not fertilize at all than it is to fertilize the
system will form if the plant experiences a slight same as you would in full sun. I once executed a
drought stress between irrigation events. A plant in simple study in which I covered a creeping bent-
shade, though, does not have the energy to chase grass (Agrostis stolonifera) putting green with
water deeper into the soil profile. This plant is shade cloth and allowed it to grow under that cloth

84 Chapter 6
for two full growing seasons. During that period, be quite detrimental to root growth. Nitrogen
I applied a normal amount of nitrogen, phosphorus fertilization in shade should be practiced in low
and potassium to some plots, and normal phospho- amounts and monitored closely for detrimental
rus, potassium and no nitrogen to others. At the effects. In some situations, nitrogen fertilizer may
end of two seasons, all of the turf was poor in qual- not be required to maintain acceptable turf.
ity because of the shade stress. However, the plots
that had not received nitrogen fertilization were
Factors that affect root growth in shade
significantly better than the plots that received a
normal rate of nitrogen. As you know, nitrogen is With the exception of St. Augustinegrass
our most important fertilizer nutrient for turfgrass (Stenotaphrum secundatum) and to a lesser extent,
management. However, in this case, nitrogen was some zoysiagrasses (Japanese lawngrass, Zoysia
detrimental. Let me explain why. japonica) warm-season (C4) turfgrasses have poor
As I stated earlier in this chapter, the light quality shade tolerance. However, warm-season grasses, in
present in shade causes responses that encourage general, form deeper root systems than cool-season
rapid vertical shoot growth. Nitrogen fertilization (C3) grasses and appear to have stronger root sys-
tends to do the same thing (Stanford et al., 2005). tems in shade. In addition, the warm summer
In a different study, in 1998, we fertilized plots of months are normally very conducive to warm-sea-
a creeping bentgrass putting green at an exception- son turfgrass growth, so it is easier for one of the
ally high rate (2 lb/1000 sf (sq. ft.); 97 kg ha−1) of rare shade-tolerant warm-season grasses to main-
urea nitrogen during the summer and irrigated tain an acceptable root system even though the
immediately (Bell et al., 2004). A week later the shade environment encourages rapid vertical shoot
plots had turned exceptionally green and looked growth instead. Cool-season grasses are detrimen-
beautiful to most people. To a turf manager, how- tally affected by high summer temperatures, high-
ever, they looked too green to be healthy. The plots intensity summer light and long photoperiods, and
remained that beautiful green color for about 10 root decline during the summer is a common occur-
days until they died. They died because their root rence in these grasses. Consequently, shade tends to
systems all but disappeared. The putting green was make an already poor situation worse.
irrigated daily but the root systems on our highly Cool-season grasses usually grow better in full
fertilized plots were so weak that they could not sun than in shade, but most will tolerate light shade
sustain water uptake through the day. Had the and some will tolerate moderate shade. Tall fescue
plants not been under summer stress they probably and the four fine fescue species, creeping red fescue
would have survived the excessive nitrogen appli- (Festuca rubra), sheep fescue (Festuca ovina), hard
cation and the reduction in root growth. However, fescue (Festuca trachyphylla) and Chewings fescue
the combination of stressful summer conditions, (Festuca rubra ssp. fallax) are probably the most
photorespiration and excessive fertilization was shade tolerant of the cool-season grasses (Gardner
enough to affect the root system so severely that and Taylor, 2002). However, even the fescue spe-
the plants died. cies suffer weaker than normal root systems in
Nitrogen fertilization in shade at normal full-sun shade (Wherley et al., 2005). Because these are
rates has a similar combination effect. The balance cool-season grasses, root growth can be encour-
between shoot growth, root growth and stem aged without enhancing shoot growth during cool
growth has already been upset by the shade condi- periods. In shade caused by deciduous trees, some
tions. Consequently, even a fertilizer program typi- root growth also occurs after leaf fall when sun-
cal of full sun can cause turfgrass to decline in light is less restricted.
shade. The plant’s photosynthesis, respiration and In cool-season grasses, shoot growth occurs best
transpiration rates have slowed considerably owing when the air temperature is between 60 °F and
to the shaded environment. Because of that, the 75 °F (16 °C and 24 °C), but root growth occurs
plants do not need much fertilizer. In addition, best when soil temperature is between 55 °F and
nitrogen fertilization tends to encourage rapid ver- 65 °F (13 °C and 18 °C) (Beard, 1973). For that
tical shoot growth, as does light quality and per- reason, there are periods during the year when
haps light quantity in shade (Tan and Qian, 2003). nitrogen fertilization encourages root growth with-
Consequently, fertilizer in an amount greater than out encouraging shoot growth (Fig. 6.7). Root
about half what is normally applied in full sun, can growth will continue in the fall after shoot growth

Importance of Light and Managing Shade 85

 60
Sun Shade
50

Root mass (mg)

40
30
20
10
0
Sept Oct Nov Apr May Jun Jul Aug Sep Oct
Month

Fig. 6.7. Root growth of creeping bentgrass (Agrostis stolonifera) managed as a putting green over the course of a
year from fall 1996 to fall 1997. Shade cloth on shaded sites was removed in the fall to coincide with deciduous leaf fall
and replaced in the spring as deciduous leaves developed. Notice the amount of winter root growth from November to
April in both shade and sun, and the difference in root mass between shade and full sun during the summer.

has slowed to almost nothing, and will continue to not enough light to perform adequatephotosynthesis,
grow slowly throughout the winter and into the there is often insufficient air movement to refresh
spring when the turf is exposed to light and the soil the boundary layer and encourage transpiration,
is not frozen. During late fall, winter and early and in many cases there is significant root competi-
spring, nitrogen fertilization encourages root tion with shade-producing vegetation. In addition,
growth rather than shoot growth. For that reason, plants, including turfgrasses, are programmed to
low applications of nitrogen fertilizer in soluble respond to shade by partitioning most of their
form or slow-release isobutylidene diurea (IBDU) available energy to rapid vertical shoot growth.
are recommended to encourage root growth in The result is poor root growth, poor stem growth
shade. Such a program is also effective in full sun. and thin, succulent vertical leaf blades with weak
cuticles. These leaf blades are easily damaged and
slow to heal because the low temperature and high
6.4 Chapter Summary
humidity normally present in shade interferes with
Turfgrasses do not particularly like shade but some respiration of the limited carbohydrates available.
will tolerate light shade and some will tolerate For instance, turfgrasses rarely tolerate traffic in
more. We can’t grow grass in a forest or under an shade because they are too easily damaged and too
opaque canopy, but we can often grow it beside a slow to heal. Consequently, shade areas should be
building or under a single tree or small group of protected from foot and vehicle traffic. If used, the
trees. Whether or not your turf will grow in shade shaded areas need to have sufficient time to heal
depends on many factors. However, the primary before they are used again. Damaged leaf blades
factors that determine shade tolerance are the are more attractive to feeding insects and more
adaptability of the species selected and the duration susceptible to disease. Because of this susceptibility,
of shade each day. Even the most shade-tolerant shaded turfgrass requires frequent observation and
turfgrasses will require at least a short period, usu- may need special pest protection.
ally 2 to 3 hours, of full sunlight each day to main- Trimming tree canopies is probably not worth
tain acceptable density. Non-tolerant species such the effort; it may not be sufficient to significantly
as bermudagrass (Cynodon dactylon) or buffalo- improve photosynthesis and if the trimming is
grass (Buchloe dactyloides) will require periods successful, it will have to be repeated regularly,
much longer than that. If you are responsible for perhaps as often as every year. Tree removal or,
growing grass in shade, and conditions and species better stated, specific tree removal may be required
are suitable for that purpose, there are manage- in order to grow turf on the site. The trees selected
ment practices that will make the turf more pre- for removal should be those that will allow the
sentable and improve its functional value. longest periods of full sun to irradiate the site.
There are three main factors that reduce the However, most people are resistant to tree removal
effectiveness of turfgrass growth in shade. There is and it may be a battle that you can’t win. In that

86 Chapter 6
case, you need to have other options available. height. That is often all that is required to grow
Shade gardens, ground covers, decorative stone good grass in shade. Managing fertilization, irriga-
patios, or even simple mulch are examples. In the tion and other factors then become practices that
case of playing fields or golf courses where alterna- improve the aesthetic and functional value of
tives to turfgrass are not an option, the decision- shaded turf rather than practices that improve its
makers will have to be educated about turfgrass chances for survival.
and shade. Your arguments may be easier to sup-
port if you invite the opinion of an outside Suggested Reading
consultant.
Air movement is often restricted on shaded sites Dudeck, A.E. and Peacock, C.H. (1992) Shade and turf-
grass culture. In: Waddington, D.V., Carrow, R.N. and
and should be encouraged by the removal of low-
Shearman, R.C. (eds) Turfgrass. ASA-CSSA-ASSA
growing tree limbs, vegetation and debris. The (American Society of Agronomy-Crop Science Soci-
simple act of removing a fence may be all that is ety of America-Soil Science Society of America),
needed to improve air flow and grow acceptable Madison, Wisconsin, pp. 269–284.
turf. Shaded turf requires little water unless it has Puhalla, J., Krans, J. and Goatley, M. (1999) Sports
to compete for water with other vegetation. Fields: a Manual for Design, Construction and Main-
Turfgrass grown on shaded sites has to be moni- tenance. Ann Arbor Press, Chelsea, Michigan.
tored closely and soil moisture sampled regularly Stier, J.C. and Gardner, D.S. (2008) Shade stress and
until you are confident that the irrigation program management. In: Pessarakli, M. (ed.) The Handbook
is acceptable. Fertilization with nutrients other of Turfgrass Management and Physiology. CRC
Press, Boca Raton, Florida, pp. 447–472.
than nitrogen should be based on growth. If the
shaded turf is growing about half as rapidly as the
same species in full sun then fertilization at half the Suggested Websites
rate is a good starting point. Nitrogen fertilization NASA Goddard Space Flight Center. Electromagnetic
that encourages root growth is desirable, nitrogen Spectrum. Available at: http://imagine.gsfc.nasa.gov/
fertilization that encourages shoot growth is not docs/science/know_l1/emspectrum.html (accessed 6
desirable. Most importantly, raise the mowing April 2010).

Importance of Light and Managing Shade 87

 Understanding and Prescribing
7 Nutrition

Key Terms
The basic plant nutrients are carbon, hydrogen and oxygen.
The macronutrients consist of two groups of three nutrients each, the primary macronutrients and the secondary
macronutrients.
The primary macronutrients are nitrogen, phosphorus and potassium.
The secondary macronutrients are calcium, magnesium and sulfur.
Turfgrasses need micronutrients, iron, copper, molybdenum, chlorine, zinc, boron and manganese in very small
quantities.
Soil texture is the proportion of three minerals, sand, silt and clay that exist in a specific soil.
Soil structure occurs when clay particles combine with organic material to form soil aggregates.
Cation exchange capacity is a measure of a soil’s ability to attract and hold nutrients.
Organic matter is dead plant material that exists in the soil in various stages of degradation. As you learned in
Chapter 5, mature turfgrass sites nearly always have organic soil layers of thatch and mat.
Soil compaction is the compression of soil pores, usually caused by traffic, resulting in less soil air and greater
soil water-holding capacity.
Humus is the fairly stable form of soil organic matter that remains after the major portions of the plant and animal
matter have been decomposed. Humus is usually dark in color.
Soil aggregates form when clay combines with organic matter. Aggregates are as large or larger than sand
particles enabling them to form large drainage pores between aggregates and small capillary pores within
aggregates.
Macropores are the large pores formed between sand particles or soil aggregates. These pores drain quickly
following soil saturation, providing air for root respiration.
Micropores are very small pores capable of holding water against gravity by capillary action. Although water
drains readily from soil macropores, it remains in micropores and provides the water for plant metabolism and
transpiration.
Compounds that volatilize (i.e. are volatile), like nitrogen fertilizer, have a propensity to change from a solid to
gaseous state under normal atmospheric conditions.

7.1 Manage Your Fertilization

nutrients. The ranking of turfgrass nutrients by
to Match Your Turfgrass Needs
importance depends on the amounts that are found
and Local Conditions
in turfgrass plants. Carbon, hydrogen and oxygen
On some of the old science fiction television shows are called basic nutrients because they are the most
and movies it was common for aliens to refer to common elements found in animals and plants,
humans as carbon-based life forms. That is an apt including turfgrasses. The second category of
description. In fact, all life on earth, including turfgrass nutrients is the macronutrients which con-
turfgrass, is carbon based. Biochemistry is a study sist of the primary macronutrients nitrogen, phos-
in carbon combinations. Consequently, it is curious phorus and potassium, and the secondary
that we tend to forget that carbon is our most basic macronutrients calcium, magnesium and sulfur.
nutrient. Oxygen and hydrogen are also basic Turfgrasses contain more primary macronutrients

88 ©CAB International 2011. Turfgrass Physiology and Ecology (G. Bell)

than secondary macronutrients and more secondary plastic, still use either natural soil or an imported
macronutrients than the third category of nutrients, sand mix. For that reason, basic soil properties are
called micronutrients. The micronutrients are made very important to us. We not only need to manage
up mostly of metals and our grasses only require our plants, in most cases, we need to manage our
trace amounts of them. Nonetheless, a missing soil to achieve our best plant growth.
micronutrient can make the difference between
acceptable turf quality and an unacceptable stand.
Cation exchange capacity
You can find standard fertilizer recommendations
for each turfgrass species in books, but most scien- Natural soils are made up primarily of three miner-
tists and managers are reluctant to make general als, sand, silt and clay. These mineral classifications
statements concerning fertilization. Different situa- are based on size. Sand is the largest of the mineral
tions, climates and overall environments require particles, followed by silt and then clay (Fig. 7.1).
custom fertilization practices to grow the best turf Sand particles are huge compared with clay parti-
possible. For that reason, standard recommenda- cles. Large particles have great volume but small
tions are a good place to start but not a good place particles have high surface area to volume ratios.
to end. As turfgrass managers, you are required to A beach ball, for instance, has a much greater vol-
develop the best fertilization practices for your sites ume than a golf ball. However, the golf ball has a
and to be flexible enough to alter those practices as greater surface to volume ratio than the beach ball.
the environment dictates. This chapter will help you If you were to fill a large dump truck with beach
to make those adjustments. balls it would probably only accept 20 to 30 balls
In order to determine our most effective fertilizer before it was full. However, if you were to fill it
plan, we need to consider a number of factors and with golf balls, it would accept hundreds of them
variables. The most important factor is probably before it was full. The golf balls have lower volume
soil. Everything about the soil affects nutrition. so more of them fit into the truck, but the sum of
A basic understanding of soil pH, soil texture, soil the surface area surrounding those golf balls is
structure and other soil attributes is not just impor- much greater than the total surface area surrounding
tant, it is absolutely necessary to designing and the beach balls that fit in the same truck (Fig. 7.2).
implementing a successful custom fertilization pro- This surface area property and the highly weath-
gram. Factors such as climate, rainfall, temperature ered condition of clay make it very much more
and seasonal growth rates are also necessary con- reactive than sand. In other words, clay will attract
siderations when designing a program. If these considerably more nutrients than sand.
variables are not considered, adjusting your pro- As you probably recall, when some chemical
gram to your situation is nothing more than trial compounds, primarily salts, are added to water,
and error management, which will probably fail, if they disassociate to form cations and anions. Table
not this year, next year or some future year. salt (sodium chloride, NaCl) disassociates to form
Managing a fertilization program for turfgrass is Na+, a cation, and Cl−, an anion. It is the molecular
not easy. It requires knowledge of the soil and bond between the positive cation and the negative
plant, consideration of the climate and weather, anion that hold the two together to form table salt.
and specific knowledge of the site. Experience also However, when they are added to water, the attrac-
helps a lot, so, in addition to reading this book, ask tion of the polar water molecules is strong enough
questions of experienced teachers and managers
until you have acquired additional knowledge of
various practices and procedures that work.

7.2 Basic Soil Attributes Clay

Plants have to have a medium in which their roots Silt

can find the nutrients and water that enable them Sand
to grow. In hydroponics, this medium is simply aer- Fig. 7.1. These three cylinders represent the size
ated water. Container plants are grown in various differences between sand, silt and clay. This drawing is
types of soilless medium, but turfgrass people, not to scale; there is a much greater difference in size
except for some experimental growth of sod on among these particles than is depicted here.

Understanding and Prescribing Nutrition 89

 Sand Clay really not necessary. You simply need to know the
difference between a high CEC and a low CEC.
Sandy soil has a very low CEC and clay loam has a
very high CEC. The CEC of soil organic matter that
has been mostly degraded is extremely high
(Table 7.1). Consequently, organic matter such as
peat, a highly degraded material, is usually added to
pure sand systems for athletic fields and golf courses
to provide some much needed nutrient and water
retention characteristics.
The highly degraded organic material and clay
particles in a soil are called soil colloids. Do you
remember soil colloids from Chapter 5? Colloids
are fine particles of one substance somewhat evenly
distributed throughout another. When a soil scien-
Fig. 7.2. It takes a much larger number of clay
tist or instructor refers to soil colloids, he/she is
particles to fill a container than it does sand particles. referring to the highly reactive portion of a soil, the
However, the clay particles that fill the container on portion that determines the CEC.
the right (not to scale) constitute a much greater total
surface area than the sand particles on the left.
The influence of soil texture and soil structure
In the preceding paragraphs I introduced the terms
to pull the cation and anion in opposite directions “loam” and “clay loam” assuming that you knew
and separate them. We say that salt dissolves in or what they were. These are terms that refer to soil
forms a solution in water. Once in solution, if a texture, and soil texture is the proportionate com-
dissolved cation encounters a stronger negative bination of sand, silt and clay that exists in a par-
force than the water molecules it will bind to the ticular soil. Soil texture is a description of the size
stronger force. Clay particles provide a stronger of the particles in the soil and has a major influence
force. We call clay particles reactive because they on soil properties. You are probably familiar with
are highly charged. Although the surface of clay the soil texture triangle of Fig. 7.3. Soil structure is
has some positive charges it is mostly covered with a property unique to combinations of clay and
negative charges that attract cations more strongly organic matter. Consequently, it is influenced in
than does water. The charges on the surface of clay part by soil texture and also by soil compaction.
particles are numerous and are called exchange
sites. The number of exchange sites in a given
Soil texture
amount of soil determines the soil’s cation exchange
capacity, commonly called CEC. Soil compaction is a very important problem
There are six nutrient anions: phosphate, sulfate, associated with growing turfgrass on common
nitrate, borate, molybdate and chloride. These six areas. It is rare for a turfgrass area to be dis-
negative nutrient-containing anions either bond played and not used. Most turfgrass stands have
weakly to the positive sites of soil particles or enter
complex reactions that bind them to the soil in other
ways. The rest of the plant nutrients are cations. Table 7.1. General approximations of cation exchange
These cations are attracted to the negative exchange capacities (CECs) common to certain types of soils.
sites on clay particles. The number of negative
Soil texture CEC (cm kg−1)
exchange sites in a soil, the CEC, is a measure of its
potential fertility. Cation exchange capacity is meas- Sand 2
ured in units called milliequivalents (meq) per 100 Sandy loam 7
grams of soil or centimoles (cmol) per kilogram. Loam 25
Clay loam 35
One meq per 100g is equal to one cmol per kg. I am
Clay 100
sure that you would understand these units if I
Organic matter 250
explained them to you but understanding them is

90 Chapter 7
 100

90 10

80 20

70 30
Clay

60 40

Pe
lay

rc
tc

en
50
en

ts
50
rc

ilt
Pe

Silty clay
Sandy
40 clay 60
Silty clay
Clay loam
30 loam 70
Sandy clay loam

20 80

Sandy loam Loam Silty loam

10 90
Loamy
Sand 0
Sand 10
10 90 80 70 60 50 40 30 20 10
0
Percent sand

Fig. 7.3. The soil triangle figuratively describes the soil texture classifications and is found in most introductory soils texts.

to sustain a certain amount of traffic which It now contains silt and clay and has a different soil
results in compaction. Compaction will be dis- texture classification.
cussed more thoroughly in Chapter 10. For now, Soil texture affects many important soil factors,
understand that soil compaction is the compres- such as bulk density, hydraulic conductivity and
sion of soil particles and results in reduced soil air others, which you can study in soil textbooks or
space and increased soil water-holding capacity. soil classes. For purposes of this text, air-holding
Because air space containing oxygen is an impor- capacity, water-holding capacity and nutrient reten-
tant requirement for root respiration, compaction tion of soils are the important factors that satisfy
can be a major problem on turfgrass sites. our purposes.
Compaction does not influence soil texture but In the preceding section, you learned that sand
soil texture determines how easy soil is to has very poor nutrient retention compared with
compact. clay. We refer to sand as inert and to clay as reac-
Sandy soil is not easily compacted and pure sand tive. Silt is also inert and organic material in the
cannot be compacted enough to cause a decline in form of humus (highly degraded organic material)
turfgrass growth. That said, however, there are is extremely reactive (Brady, 1990). Clay and
other factors such as natural accumulations of humus by themselves hold lots of water, very little
dust, sediment and organic material that can con- air and lots of nutrients. Silt also holds lots of
taminate pure sand enough to encourage compac- water and very little air, but few nutrients. Sand
tion sufficient to restrict root growth and water holds lots of air, little water and few nutrients. The
uptake (Murphy et al., 1993b). In that case, the properties of silt are mostly negative (Table 7.2). It
original pure sand is no longer the same texture. is easily compacted, holds very little air and few

Understanding and Prescribing Nutrition 91

Table 7.2. Some of the advantages and disadvantages of the mineral fractions that determine soil texture.

Mineral Advantages Disadvantages

Sand High compaction resistance Poor nutrient retention

Rapid drainage Poor water-holding capacity
High oxygen retention
Silt High water-holding capacity Poor nutrient retention
Poor oxygen retention
Poor compaction resistance
Poor drainage
Clay High nutrient retention Poor drainage if compacted
High water-holding capacity Poor oxygen retention
Good drainage if not compacted Poor compaction resistance

nutrients. Sand and clay, however, have advan- or more of the soil are compressed, destroying
tages depending on the expectations of the site. structure. This compaction also helps to seal the
Sand does not compact substantially and provides soil surface, thus interfering with water infiltration
superior aeration. Clay and humus hold high and encouraging water runoff. For that reason,
amounts of water and nutrients and are desirable turfgrass managers regularly aerify, the process
for that reason. We will discuss these properties introduced in Chapter 5 primarily for thatch con-
further but before we do we need to review soil trol. Aerification helps to relieve soil compaction
structure. and promote water infiltration (McCarty, 2001).
Although aerification does not necessarily promote
soil structure, it does promote root growth, which
Soil structure
leads to better structure.
Soil structure occurs when clay and organic matter, Refer to the representation of soil structure in
primarily humus, combine to form large particles Fig. 7.4. Notice that the aggregates are large com-
called aggregates (Fig. 7.4). These aggregates are plexes that consist of much smaller particles. These
the units that combine to form soil structure. There complexes are too large to fit tightly together, much
are at least seven different types and subtypes of
aggregates that form from various combinations of
materials and determine the various forms of soil
structure (Brady, 1990). There are normally at least
two types of structure present in a mineral soil.
However, that is not our basic concern. We need to
know how structure affects turfgrass and how we
Macropores
can sustain it and make it work for us.
Plants are facilitators of soil structure. Root
growth helps to break up large compacted units of
soil and helps to form smaller, more useful aggre-
gates. Plant roots, mostly root hairs, secrete organic Micropores
materials that help to cement soil aggregates
together (Brady, 1990). Theoretically then, a soil
with a high number of plants and plant roots, such
as a turfgrass stand, maintains soil structure better
than a soil containing few or larger plants with less
Fig. 7.4. Soil structure occurs when clay and organic
extensive rooting. However, turfgrass areas tend to material combine to form particles as large or larger
sustain high levels of pedestrian and sometimes than sand called aggregates. Aggregates are the
vehicle traffic that negatively affect structure. As building blocks of soil structure. Large pores between
pedestrian and/or vehicle traffic moves across the aggregates (macropores) hold air and small pores
turfgrass sites, the upper 1 to 2 inches (2.5–5.1 cm) within the aggregates (micropores) hold water.

92 Chapter 7
like the cylinders that represent the sand particles more thought than action. You not only have to
in Fig. 7.2. Large rounded objects cannot physi- select the right product, determine how much to
cally fit together no matter how tightly packed apply, calibrate your equipment and select the
without leaving relatively large gaps or pores proper time, you also have to take precautions that
between them. They can only fit tightly together the fertilizer is not going to volatilize, runoff or
along flat surfaces. Small rounded particles cannot leach before the plant has a chance to use it.
fit tightly together either, but there are much Incidentally, how do you apply carbon, hydrogen
smaller gaps between them (Fig. 7.2). The large and oxygen, and what do you do if your soil pH is
gaps that form between sand particles, between soil too high or too low?
aggregates and between sand particles and soil
aggregates are called soil macropores, and the
Managing the basic nutrients
small gaps that form between clay particles and
within soil aggregates are called soil micropores. Can we manage carbon, hydrogen and oxygen?
Soil aggregates are large enough to have the Carbon, yes; oxygen, yes; and hydrogen does not
attributes of sand as well as the attributes of clay. matter. Hydrogen is ubiquitous. It is the most avail-
I remember my introductory soils instructor asking able element in nature and it is everywhere. Our
the class a question many years ago. The question plants can get hydrogen from water, from air and
was “Which soil drains fastest, a sandy loam or a from nearly any compound that they assimilate. We
highly aggregated clay loam?”. Of course, we all don’t have to manage hydrogen but we do have to
answered sandy loam but, according to him, a manage carbon and oxygen if we expect our turf to
highly respected soil scientist, we were wrong. look and perform at its optimum. The three most
A highly structured clay loam forms aggregates as important plant physiological functions are photo-
large or larger than sand. Consequently, the soil synthesis, respiration and transpiration, and these
assumes the drainage properties of sand. The large require carbon dioxide, oxygen and water, respec-
particles cannot fit together tightly and macropores tively. If we cannot manage carbon and oxygen then
are left between them. Yet, the aggregates are com- all we can do is mow and hope for the best.
posed of clay particles and organic matter that
maintain very small pores. These micropores hold
Carbon
water against gravity because of capillary action, a
physical phenomenon that was described in How do grasses accumulate carbon? Grasses
Chapter 1 and again in Chapter 4. Hence, a highly accumulate carbon by assimilating carbon dioxide
aggregated clay loam has both the drainage during photosynthesis. Consequently, management
advantages of sand and the water and nutrient procedures that encourage photosynthesis also
retention capabilities of clay. Unfortunately, the encourage carbon nutrition. We may not be able
clay loam does not maintain the sand characteristic to control photosynthesis but we can certainly
of resistance to compaction and its structure is affect it.
easily destroyed by traffic. If the site has to In review, photosynthesis requires light, water
maintain regular traffic, the compression of the and carbon dioxide. Light is reasonably easy to
structure in the upper soil layers cannot be avoided manage in most situations. There is not much we
and aerification must be performed regularly to can do about buildings or other shade-producing
help relieve the inevitable soil compaction. structures, but we can affect trees and other vegeta-
tion. In Chapter 6, you learned some techniques of
shade management. There is a limit to the amount
7.3 Plant Nutrition
of shade that a turfgrass can stand. That limit has
In order for a plant to assimilate a nutrient, the a lot to do with you, or your customer’s expecta-
nutrient must be soluble. The air, for instance, is tions. The better you want the turf to look and the
about 78% nitrogen, but that nitrogen is not avail- lower that you want to mow it, the more shade you
able to plants because it is not soluble. The soil in will have to eliminate. You can often grow a sparse
my area of the USA is often red, indicating high turfgrass cover in shade but in most cases a sparse
iron content. That is mostly iron oxide, however, cover is not acceptable. If trees need to be removed
and it is neither soluble nor plant available. in order to grow good grass, it is up to you to edu-
Consequently, applying fertilizer to turf requires cate your customer on his/her alternatives. It is also

Understanding and Prescribing Nutrition 93

up to you to know which trees need to be removed oxygen. Consequently, oxygen is not limiting for
and which trees can remain. In the case of tree respiration in aboveground plant organs. However,
removal your role is advisor. Tree removal deci- it can become limiting in the soil. As you learned
sions have to be made by your customer. Otherwise, earlier, oxygen is necessary for root respiration and
you may be jeopardizing your job. However, the root respiration is necessary for water and nutrient
customer must realize that growing acceptable turf uptake. Roots do not function unless they respire.
in the shade currently present is simply not possi- Hence, saturated soil around turfgrass roots shuts
ble. The decision is to save the tree or save the down root uptake. Turfgrass managers must pro-
grass. In some cases, saving both is not possible. vide adequate drainage for their turf. There may be
As you learned in earlier chapters, we can also situations where adequate drainage is not possible
provide more carbon dioxide to our turf by pro- but those situations are extremely rare. Like trying
moting air movement. Unrestricted airflow refreshes to grow grass in near perpetual shade, you will not
the boundary layer, replacing air that is rich in be successful consistently growing turfgrass in
oxygen and water and low in carbon dioxide with areas that cannot be drained properly. During any
air containing normal gaseous proportions. Good rainy season, your turf will struggle or die.
airflow also encourages transpiration. You learned Soil compaction is another major factor contrib-
in Chapter 6 that you can help to mitigate low uting to poor soil oxygen levels. Compaction is
photosynthesis in shade by enhancing airflow. similar to poor drainage in that the soil holds too
Sometimes, enhancing air movement on shaded much water and too little air. In the case of com-
sites is sufficient to provide adequate carbon for paction, however, better drainage will not be ade-
acceptable turfgrass nutrition. quate to completely relieve the problem. Compaction
Mowing height has a very noticeable effect on results in smaller pores that hold water more
carbon accumulation in stressful situations. The tightly. In effect, compaction turns macropores into
lower you mow the turf, the more efficient it has to micropores. So following compaction, many of the
be to accumulate carbon. A long leaf provides sub- macropores that previously held air become micro-
stantially more photosynthetic potential than a pores that hold water. If these pores give up water
short one. Finally, grasses have to have water to to transpiration or evaporation and become filled
accumulate carbon. If yellow to brown grass is not with air, there is sufficient oxygen present for respi-
acceptable during extended dry periods, irrigation ration, but there is no longer water present for
must be provided. When turfgrasses do not have uptake. Aerification and traffic control are the two
sufficient water to transpire or to perform photo- primary practices that can help control compac-
synthesis, they enter a period of quiescence that we tion. We have already discussed those procedures.
call dormancy, or they die. Some species can sus- Soils high in salt also have a restrictive effect on
tain long periods of dormancy, others cannot. soil oxygen. Salts, especially sodium, destroy soil
Although we do not list water as a nutrient, it is structure and attract and hold excess water. Similar
obviously important for all plant functions. to compacted soils, sodic soils hold too much water
Yes, you can affect carbon nutrition and oxygen and too little air (Harivandi et al., 1992). Sodium
nutrition, but the processes required are not nearly as is a large element that is larger when water is avail-
simple as a fertilizer application. Most management able. When water is present, sodium is surrounded
practices are designed to improve carbon and oxygen by it and holds it tightly. Consequently, where large
assimilation. It is your responsibility to assess envi- complexes of sodium and water are bound to the
ronmental factors and choose the correct manage- soil CEC, the colloids cannot move close enough to
ment for your situation(s). Managing the basic each other to form aggregates. Hence, soil structure
nutrients requires evaluation, planning and effective is destroyed and few macropores are formed. With
timing of the practices required. Do not forget that no macropores to hold air, the effects of the salt are
your decisions not only enhance the assimilation of similar to those of compaction. However, even if
carbon and oxygen, they can also cause a decline. oxygen was present for respiration, much of the
water in the soil would not be plant available
because the turfgrass roots would not be able to
Oxygen
create osmotic potentials sufficient to break the
As photosynthesis proceeds, oxygen is produced. strong bonds between water and salt. We will
Normal air also holds a substantial amount of discuss these phenomena further in Chapter 10.

94 Chapter 7
For now, remember that the best management is best to apply lime in the spring and the fall, and
practice for salty soil is applications of gypsum – a avoid applications in the summer when the weather
common name for calcium sulfate. is hot. A soil test is required to determine how
much sulfur or how much lime will be required to
change soil pH. It may be an incredibly large
What to do with soil pH
amount. If sulfur/lime applications are not practi-
Soil pH can have a detrimental affect on nutrient cal, foliar feeding will be necessary. Foliar feeding
availability (Carrow et al., 2001). Most turfgrasses is simply the application of nutrients in light, fre-
grow adequately in a wide range of pH, but all quent fine sprays. The objective is to make applica-
appear to grow adequately at a soil pH of about 6.5 tions in amounts small enough and at times that
(Waddington, 1992). A pH that is too high affects are conducive for the plants to absorb the nutrients
the solubility of micronutrients such as iron, copper, through the foliage instead of through the roots.
zinc, boron and manganese. Nitrogen becomes less Foliar feeding is effective, but it is an expensive,
available at a pH of over about pH 8.0 or less than labor-intensive practice.
about pH 6.0. Phosphorus has a more narrow range
than nitrogen, with decline in availability beginning
Soil test reports
at about pH 7.5, and a sharp decline at about pH
6.5. Acidic soils of pH below 6.0 tend to limit the Fertilizer decisions should be based on soil test
availability of important nutrients such as potassium, reports. Over-application of nutrients such as
calcium, sulfur and magnesium, and the micronutri- nitrogen, and particularly phosphorus, can be an
ent molybdenum. Consequently, managing pH is an environmental hazard and should be avoided
important practice. (Soldat and Petrovic, 2008). Although P is bound
Agricultural sulfur (elemental sulfur, S2) is used tightly to the soil, fertilizer P applications are solu-
to lower pH. Microbial breakdown of S2 results in ble and run off easily into surface water, thereby
sulfuric acid, which eventually lowers the pH of the creating biological hazards (Bell and Moss, 2008).
soil. Sulfur applications can burn turfgrass so they Not all soil reports are created equal. There are
should be limited to less than 10 pounds per 1000 several different methods available for determining
square feet (488 kg ha−1) of turf per application and the soil content of major nutrients. Particular soils
only applied in spring and fall when the weather is are better tested with one method than they are
cool. Be careful not to overlap passes, resulting in with others. Consequently, it is best to get your soil
strips of double applications. Applications of iron tested at a laboratory in your general region.
sulfate or aluminum sulfate can also help to acidify A laboratory close by is most likely to use the best
the soil. Iron is an important micronutrient of turf- testing methods for your location and should be
grass and is preferable to aluminum, which rarely familiar with the specific soils in your area. A local
adds anything positive to the soil. laboratory will also be familiar with your turfgrass
Soils are highly buffered, meaning that they species and be able to make competent recommen-
strongly resist changes in pH. A downward, more dations for the amount of fertilizer to apply. Unless
acidic change in pH is more difficult to accomplish you are very familiar with soil science, it is best for
than an upward one. On some highly buffered you to at least start with laboratory recommenda-
soils, it could take a lifetime of sulfur applications tions rather than trying to make your own. With
to make even a minor change in pH. Under those experience and enough recommendations, you will
circumstances it is not worth the effort, although probably learn how to determine fertilizer require-
sometimes, the application of acidifying fertilizers ments for your site, or at least the requirements for
such as ammonium nitrate, ammonium sulfate or macronutrient fertilization.
potassium nitrate, particularly ammonium sulfate, Some laboratories report soil nitrate nitrogen
are enough to make sufficient nutrients available. (Fig. 7.5), and some report total soil nitrogen. For
Lime (CaCO3) is applied to raise soil pH. It is purposes of turfgrass management, however,
usually possible to raise the pH of acidic soil but it measures of soil nitrate or total soil nitrogen have
often requires a substantial amount of lime. Lime is no real value. Soil N changes daily and is affected
safer on turfgrass than sulfur and can be applied at by numerous factors. Consequently, we have to
rates of up to about 40 pounds per 1000 square apply the proper amount of N fertilizer based on
feet (1950 kg ha−1) in a single application. Again, it knowledge and practical experience. Soil reports

Understanding and Prescribing Nutrition 95

Fig. 7.5. A typical soil report. Notice that this laboratory reports soil nitrate and other nutrients in parts per million
(ppm) or pounds per acre (lbs./acre).

96 Chapter 7
are generally meaningless for determining applica- testing is superior to soil testing. Tissue testing,
tions of N fertilizer to turfgrass. Soil reports for P however, is exceedingly more complicated than soil
and K, however, are accurate and useful. Reports testing and requires a thorough knowledge and/or
of calcium, magnesium and sulfur are useful for history of what nutrient levels are necessary to
pointing out potential deficiencies, and micronutri- maintain your expectations. It gets complicated
ent tests may lead you to the answer for a difficult- because adequate tissue levels differ with season,
to-diagnose problem. just as response to fertilizer differs with season. If
While some laboratories report nutrient content you are willing to keep good records and thor-
per unit of soil in parts per million and/or pounds oughly assess responses to fertilizer input, tissue
per acre (Fig. 7.5), some may use kilograms per testing may be right for you. Regardless of the
hectare. Others report nutrient content as percent method you use to assess your nutrient require-
base saturation. Still others issue reports that ments, you will be money ahead and exceedingly
include both nutrient content per unit of soil and more effective if guesswork is mostly eliminated.
nutrient content in percent base saturation
(Fig. 7.6). There are general guidelines for nutrient
The primary macronutrients
content per unit soil for both cool- and warm-
season grasses but many specific circumstances On any bag of consumer fertilizer you will nor-
apply and it is my opinion that you are best served mally find a formulation. The formulation describes
to ask the lab for recommendations. If you don’t the amount of the three primary macronutrients,
completely understand the report, call the labora- nitrogen (N), phosphorus (P) and potassium (K), in
tory and ask questions. Your success may depend the bag. These amounts refer to percentages and
on knowing the answer. are presented in a form consistent with N-P-K. The
If your laboratory reports base saturation, they formulation 14-3-6 means that the bag contains
are telling you the proportion of soil exchange sites 14% pure N, 3% P2O5, and 6% K2O. The 3%
that are theoretically occupied by each nutrient. P2O5 in that bag is not the same as 3% P. In fact,
That is very useful information because it ignores P2O5 is only 44% P and K2O is 84% K, which
soil fertility and reports nutrient retention in com- results from a simple comparison of the molecular
parison with other nutrients. In most soils, calcium weights of P, O and K. So the bag really contains
occupies the majority of exchange sites and there is 14% N, 3% × 0.44 = 1.32% P and 6% × 0.84 =
a high percentage of magnesium. If that is not the 5.04% K. Turfgrass managers apply fertilizers
case in your soil, it indicates problems that need to based on pure N applied, so we need to know these
be addressed. Other deficiencies may also be appar- calculations so that we can choose formulations
ent. A quality soil laboratory knows what your that result in the proper application of P and K
base saturation should look like for growing turf- when we apply the expected amount of N. That
grass and will make recommendations to correct sounds complicated, but it really is not; it is just
existing problems. simple mathematics as we use every day.
Choose a laboratory before you sample your
soil. A laboratory is generally chosen based on
Nitrogen
reputation and the recommendations of other turf-
grass managers and horticultural or crop produc- Nitrogen is the most important nutrient for turf-
tion specialists. Nearby turfgrass managers are grass. If water is present and a minimal amount of
your best source of information. Call the labora- the other important nutrients are present, an
tory before you collect samples. Ask them for application of N will encourage turfgrass growth
advice on how to proceed with sampling. Ask for and improve green color. Adequate N fertilization
recommendations concerning the proper size of will improve turfgrass overall health, and increase
sampling areas and the number of samples to col- turfgrass cover and density. However, N fertiliza-
lect on each area. Also ask for directions on sam- tion at a rate above optimum is detrimental to
pling depth; this is important and differs by turfgrass health and can result in increased traffic
laboratory. In addition, ask for any other advice damage, pest susceptibility, decreased root growth
that the laboratory might feel is important. and even death.
Many soil laboratories also do tissue testing, and Turfgrass gets nitrogen from soil organisms that
many turfgrass managers are convinced that tissue remove it from the air. It can also get nitrogen from

Understanding and Prescribing Nutrition 97

Fig. 7.6. A typical soil report from a laboratory that reports base saturation as well as soil nutrient content in pounds
per acre. Notice the recommendations at the bottom of the report.

organic material as microorganisms degrade it. selection, are based on climate. Climate, however, is
Nitrogen is a part of rainfall, especially during substantially different from weather. Climate is the
thunderstorms, when lightning causes N in the air average of weather conditions over an extended
to combine with precipitation. Turfgrasses rarely period. In the USA, the period is 30 years. Weather
need N in addition to that supplied by nature to changes substantially on an annual basis and
survive. However, they normally do need addi- weather determines the conditions under which
tional N to look and function the way that people your turfgrass must thrive, not climate. When we
want them to. We can make a huge difference in talk about ecology in later chapters, you will learn
turfgrass by adding N fertilizer at the right time that disturbances, sometimes caused by weather,
and in the right amount. can change the species composition of an ecosys-
You can find turfgrass fertilization guidelines on tem. Consequently, weather can certainly influence
university websites in your region based on species the health of a turfgrass stand and the amount of
and climate. Introductory turfgrass textbooks also fertilizer required to sustain acceptable growth and
include general guidelines for turfgrass fertilization. development. You must be able to adjust to a
You can often find guidelines in trade magazines changing environment to grow your best turf.
and you can surely get advice from other turfgrass One of the most important weather factors that
managers in your area. However, you must remem- influences fertilization is temperature. Temperature
ber that these are just general guidelines, they are influences all three important physiological processes,
not always the best programs for your site and they photosynthesis, respiration and transpiration, there-
should be adjusted according to recent weather fore it affects the amount of nutrients required for
conditions. Fertilizer recommendations, like species plant maintenance. In Chapter 6, we discussed how

98 Chapter 7
applying fertilizer at the proper time of year could Warm-season grasses are better adapted to sum-
encourage root growth in shaded cool-season (C3) mer conditions and apparently have been naturally
grasses. This same procedure is useful for cool- selected for deep root systems (Christians, 2007).
season grasses in full sun (Wehner et al., 1988). Although, similar to cool-season grasses, the natu-
Fertilization, especially nitrogen fertilization, tends ral tendency of warm-season grasses is to favor
to encourage shoot growth, sometimes at the expense shoot growth over root growth, there appears to
of root growth (Hull, 1992; Bowman, 2003). If we be a greater energy partition to root growth in C4
hope to use N fertilizer to encourage root growth in grasses than there is in C3 grasses. For that reason,
a cool-season grass, the best time to affect that proc- and because photorespiration does not occur in
ess is when it is too cool for shoot growth but warm the C4 photosynthetic pathway, warm-season grass
enough for root growth (Moore et al., 1996; Grossi fertilization is more simple. Growth slows in
et al., 2005). As we discussed in Chapter 6, those warm-season grasses when daytime high tempera-
conditions occur in late fall, winter and early spring tures exceed 95 °F (35 °C) for extended periods.
(Wehner and Haley, 1993). However, warm-season grasses can be fertilized to
Summer is a poor time to fertilize cool-season match seasonal growth rates or can be fertilized on
grasses. During summer, high temperatures encour- a regular periodic basis throughout the growing
age rapid photosynthesis, respiration and transpi- season with few detrimental effects. Most manag-
ration, increasing the need for nutrients. In C3 ers caution against fall N fertilization of warm-
grasses, however, high temperatures also encourage season grasses. Nitrogen fertilization tends to
photorespiration. As photorespiration increases make plant leaves more succulent. In view of this,
during summer (see Chapter 3), plant energy and because warm-season grasses are very sensi-
declines. During this period, cool-season grasses tive to cold temperatures, it is believed that dam-
will use most of the available energy to maintain aging ice crystals might form in succulent
shoot growth, and partition less available energy warm-season plants at higher temperatures than in
into root growth (Hull, 1992). Consequently, drier plants. Although this reasoning is sound,
although some new roots are formed many roots scientific tests have not demonstrated such an
die and are not replaced. This process results in a occurrence (Reeves et al., 1970; Goatley et al.,
gradual loss of the root system of a cool-season 1994). In spite of that, even scientists are cautious
grass during summer (Huang and Liu, 2003). If the about recommending fall N fertilization of warm-
root system was not adequate when the summer season grasses.
began, the plant will probably die. If the root sys- Rainfall is also a factor for consideration during
tem was healthy when the summer began, the plant N fertilization. Nitrogen fertilizers may be lost to
will survive. When nitrogen is added, the balance volatilization or to leaching, but the most likely
of shoot growth to root growth shifts farther loss of N fertilizer is in surface runoff (Petrovic,
toward shoot growth, causing an increase in stress 1990). Runoff occurs when rainfall exceeds the
on plant roots (Bushoven and Hull, 2005). As the infiltration rate of the soil. Once the soil pores fill
root system declines the plant’s ability to transpire with water, surface runoff begins. If the soil is satu-
also declines. Eventually, the root system may rated or partially saturated before rainfall begins,
become so weak that it can no longer keep the runoff occurs more rapidly than it would if the soil
plant hydrated during hot, dry days and the plant was dry. Consequently, fertilizer on saturated or
dies. Fertilizer timing of cool-season grasses is partially saturated soil may be wasted if rainfall
important and should be based more on the times occurs within the next 24 to 48 hours. In addition,
when root growth can be encouraged as well as or periods of rainy weather are usually accompanied
in place of shoot growth. A nitrogen rate that is by long periods of cloud cover which reduce pho-
typically healthy in the fall can damage a cool- tosynthesis. The temperature is usually cooler than
season grass in the summer. The best time to ferti- normal as well, and as long as the turfgrass is wet
lize cool-season grasses to encourage the root it stays cool. Therefore, respiration slows and tran-
growth that will help them survive summer heat is spiration is not required. Less nutrition is required
after the weather cools to the point that the shoots too. Periods of wetter than normal weather require
stop growing. adjustments to your fertilizer program. Periods of
The timing of warm-season (C4) grass fertiliza- exceptionally dry weather also require alterations
tion differs from cool-season grass fertilization. to your fertilizer program.

Understanding and Prescribing Nutrition 99

 If irrigation is available, fertilization should con- form and provide a slow fairly constant N release
tinue normally during periods of dry weather. over a long period. Slow-release N provides a
However, roots do not take up fertilizer unless it is consistent turfgrass response over time, but with
dissolved in water, so fertilizing non-irrigated turf- one exception, IBDU (isobutylidine diurea), has
grass when the weather is dry is a waste of time and little effect when applied in cool weather (Volk and
money. During dry weather, a large amount of N Horn, 1975). Turfgrass managers tend to be loyal
fertilizer may volatilize before rainfall occurs and to the types of fertilizer they use. Not just to the
soil water becomes available. Other fertilizers (P, K, quick-release, slow-release type of fertilizer used,
etc.) will also degrade, but degradation will occur but often to the manufacturer and formulation.
more slowly. Humidity is another factor that These managers trust the products that they use
should be considered before fertilization. but, even more important, they trust their ability to
High humidity slows transpiration and also accurately determine how best and when to apply
encourages fungal diseases. When N fertilizer is them. With these particular products, they are con-
added to turfgrass the grass becomes greener. The fident that they can determine how their turfgrass
grass greens because N fertilizer encourages chloro- will respond under different environmental condi-
phyll synthesis. As chlorophyll levels increase, tions and to different rates of application. That
turfgrasses absorb more light energy and have to should give you some idea of how important they
dissipate more heat. If transpiration is slow because deem N fertilization to be in a management scheme
humidity is high, the turf becomes too hot and its for providing outstanding turf (Box 7.1).
overall health declines. Therefore, N fertilization There are several types of quick-release and slow-
should be adjusted during periods of higher than release N sources to choose from. Some of the most
normal humidity. If disease occurs, depending on common forms are listed in Table 7.3. These and
the pathogen present, N fertilization may make the other N sources are often mixed to provide specific
disease worse. formulations and combinations that include both
Nitrogen fertilizer and water are the two major slow-release and quick-release N in a bag of ferti-
inputs that make the most difference between man- lizer. Explanations of how these fertilizers are manu-
aged and unmanaged turfgrass. Nitrogen fertilizer factured and combined, and the special properties of
makes turfgrass green. It also encourages turfgrass each, can be found in many sources (Carrow et al.,
to grow and spread, and increases the overall 2001; Christians, 2007; Turgeon, 2008).
health of the plant. However, there are occasions One of the most difficult aspects of N fertiliza-
when N fertilization should be reduced in compari- tion is its propensity for N being lost to the atmos-
son with the normal amounts applied. Periods of phere or to runoff or soil water. Nitrogen fertilizer
high temperature, high rainfall and high humidity volatilizes easily, meaning it is lost to the atmos-
call for less than normal fertilizer. As in late-fall phere as a gas (Petrovic, 1990). The warmer the
fertilizer applications to cool-season turf, some- temperature, the more likely that volatilization will
times the timing of the application is as important occur. Losses of N to the atmosphere are more
as the amount applied. A turfgrass manager should likely to occur from quick-release N sources than
always know what weather is expected for the next from slow-release sources (Torello et al., 1983).
3 or 4 days following a fertilizer application. Because quick-release N is highly soluble, it is also
A severe storm within 24 to 48 hours of a fertilizer more likely to be lost to surface or soil water.
application could result in substantial losses to A major rainfall event that is sufficient to produce
runoff (Shuman, 2002). runoff will carry surface-applied soluble fertilizers
in solution off the target site and into surface water
NITROGEN FERTILIZERS Nitrogen fertilizers for turf- (Cole et al., 1997). However, irrigating with
grass are separated into two categories, quick approximately 0.25 inches (6 mm) of water imme-
release and slow release. Quick-release fertilizers diately following fertilizer application will prevent
are soluble in water, relatively inexpensive, can be most volatilization and runoff losses (Titko et al.,
applied in spray or granular form, can be applied in 1987; Shuman, 2002).
any season and provide a quick turfgrass response. Although quick-release N has advantages, it also
Slow-release N fertilizers are not readily soluble, has the disadvantage of being easily lost to volatil-
are moderately to highly expensive, with one or ity or water movement. Quick-release N is less
two exceptions can only be applied in granular expensive than slow-release N but must be applied

100 Chapter 7
 Box. 7.1. Nitrogen fertilizer sources: believe it or not.
Turfgrass managers have all kinds of opinions con- quick-release source. Although that opinion may
cerning fertilizer sources. I have heard experienced have some basis, it is not scientifically sound and is
golf course superintendents claim that quick-release just an opinion.
granular N fertilizer is more likely to burn creeping Although these opinions have no scientific basis
bentgrass (Agrostis stolonifera) putting greens than and may seem unreasonable, there is something to
spray applications of the same source. That does be said for having confidence in the product that you
not seem to make sense, but when more than one use and less confidence in those that you do not.
experienced superintendent makes that statement Considerable experience with a good fertilizer prod-
you have to believe that there might be something uct that you believe in is substantially better than
to it. I asked one experienced superintendent, one using a supposedly more reliable product in which
of my old students, if he believed that a granular you have little experience and little faith. Knowing
slow-release fertilizer would burn his greens. He your product intimately is a huge advantage when
said “probably not but why take the chance”. I have environmental conditions warrant a change in your
heard many managers claim that quick-release N normal fertilization plan. Unsupported opinions are
sources, when applied properly, result in a slightly usually destructive, but there are rare times when
greener, slightly denser canopy than an equally they can be advantageous. In this case, there are a
good slow-release source. There is no scientific number of good fertilizer products and companies
basis for that opinion. Nonetheless, I also have that seem to be making them better and better. Choose
opinion. But I also believe that a turfgrass fertilized the one that works best for you, learn its performance
regularly (monthly or so) with a slow-release ferti- characteristics under a variety of conditions and use
lizer is healthier than one fertilized properly with a it with confidence.

Table 7.3. Common quick-release and slow-release may be lost to leaching after watering in, but most
sources of nitrogen for turfgrass fertilization. will bind to the soil CEC strongly enough to resist
leaching. As soils high in sand have low CEC, leach-
Quick-release N
sources Slow-release N sources
ing is more probable in sand than in soil (Petrovic,
1990). However, if a moderate amount of soluble
Ammonium nitrate Natural organics (manure, etc.) fertilizer is watered in with about ¼ inch of irrigation
Ammonium sulfate Methylene urea or rainfall, as mentioned earlier, it will be assimilated
Potassium nitrate Sulfur-coated urea by the plants quickly. Consequently, after 2 to 3 days,
Calcium nitrate Polymer-coated urea
there will be little fertilizer left to leach.
Isobutylidene diurea (IBDU)
Quick- and slow-release N sources cause slightly
different color/growth responses from turf (Fig. 7.7).
at lower rates more frequently. Slow release N is Quick-release sources elicit strong color and growth
more expensive but is less labor intensive because it responses that reach a peak in about 10 to 14 days,
can be applied less frequently. Turfgrass managers depending on environmental conditions, and then
often choose the type of fertilizer they use because decline at about the same rate. A slow-release N
they like the turfgrass response that occurs because source elicits more gradual color and growth
of it. However, those on a tight budget must also responses that are sustained over a longer period
consider the economics of the fertilizer they choose, (Waddington and Duich, 1976). Consequently,
including the labor expense required to apply it. If monthly applications of slow-release fertilizer result
your labor is expensive, slow-release sources may in a consistent growth rate and a more consistent
be more economical than quick-release sources. If color response compared with monthly applications
labor is relatively inexpensive, then quick-release of quick-release sources (Zhang et al., 1998). For
sources are likely to be cheaper. Regardless, you that reason, quick-release sources need to be applied
can save expensive fertilizer movement off the site at lower rates more frequently than slow-release
by handling the product properly. Once fertilizer is sources to elicit consistent turfgrass responses.
watered into the soil it is not likely to volatilize and A steady growth rate is preferable and considered
once in the soil it will not run off. Some fertilizer healthier for turfgrass than a series of peaks and

Understanding and Prescribing Nutrition 101

 9 polymer-coated particle is affected. Another factor
Biweekly
that should not be forgotten is that if the coating
Monthly
on a coated urea particle is damaged, it becomes a
Visual color rating

8
quick-release fertilizer. Mowing, aerification and
other cultural practices can damage coated parti-
7
cles of urea, and the N application that you
expected to release over a long period may release
6 much quicker. Coated particles need to be watered
in before mowing and fertilization should not be
5 followed by other cultural practices that are likely
Apr May Jun Jul Aug Sep Oct
to cause damage to the particle. Both quick-release
Month
and slow-release N sources provide satisfactory
Fig. 7.7. Actual color response of bermudagrass results for the N fertilization of turfgrass. However,
(Cynodon dactylon) fertilized with urea nitrogen, a application timing and application methods vary
quick-release N source every 2 weeks (biweekly) at 0.3 and need to be seriously considered when develop-
pounds N/1000 square feet (15 kg ha−1) averaged over ing a fertilizer program.
two seasons in Stillwater, Oklahoma (black line). The Clippings are a primary source of slow-release N.
gray line represents what the response curve would When the temperature is warm, clippings are bro-
probably look like (estimated) if the fertilizer was applied ken down fairly rapidly by microbial activity and
at 0.6 pounds N/1000 square feet monthly. Notice the
nutrients are returned to the soil and plants (Starr
peaks and valleys as the monthly applications reach
maximum color then decline until the next application
and DeRoo, 1981). Research has indicated that the
cycle. A slow-release N source at the same rate would N recovered from clippings of Kentucky bluegrass
closely approximate the smooth response curve of the (Poa pratensis), perennial ryegrass (Lolium per-
biweekly applications. Color was rated visually on a enne) and tall fescue (Festuca arundinacea) was
scale of 1–9, where 1 = brown and 9 = dark green. often greater than the N fertilizer applied (Liu and
Hull, 2006). It has been estimated that N fertilizer
could be reduced by 50% on a mixed stand of
valleys that occur with each fertilizer application. Kentucky bluegrass, perennial ryegrass and creep-
However, it must be remembered that each slow- ing red fescue (Festuca rubra) if clippings were
release product has slightly different attributes, and returned instead of collected (Kopp and Guillard,
some research among suppliers and producers is 2002). Unless clippings must be collected to pro-
required to determine the product that best meets mote the playability of a surface, or the aesthetic
your needs (Carrow, 1997). value of the turf requires collection, clippings
It should also be remembered that most slow- should be left in place; they are a great source of
release products are affected by microbial activity fertilizer.
and therefore by temperature. Release of all slow-
release fertilizers except for IBDU is delayed by
Phosphorus
cold temperatures (Wilkinson, 1977). The N release
of IBDU is controlled by water and determined by Phosphorus (P) is the least important of the
particle size; the larger the particle, the slower the primary macronutrients for turfgrass fertilization.
release. Microbial activity need not be present. A primary use for plant P is in cellular energy mol-
However, N release from natural organics such as ecules such as ATP (adenosine triphosphate) that
manure, or from methylene urea, is controlled by are in high demand during periods of rapid growth
microbial activity and does not occur in cold or reproduction. Because turfgrasses are normally
weather (Wilkinson, 1977). Release of N from mowed below the heights at which they flower and
sulfur-coated or resin-coated urea is partially con- seed, P is not as important for a reproductive
trolled by imperfections in the particle coating and energy boost in turfgrass as it is in most flowering
partially by microbial breakdown. Therefore release plants. In addition, turfgrasses have very extensive
slows but does not stop in the winter. The same is root systems for their size and are capable of
true of polymer-coated urea particles that work by sequestering soil P better than most plants
diffusion. As water cools it becomes more vis- (Christians, 2007). Research has demonstrated
cous, and the diffusion of urea in and out of the that turfgrass roots actually seek out P and will

102 Chapter 7
grow toward the greatest concentration of it desired. Sometimes additional P at establishment
(Lyons et al., 2008). makes no difference at all (Frank et al., 2002). Soil
Soil tests determine soil P availability and whether tests for P are quite accurate, inexpensive fertilizers
or not P fertilization is required. A very general are reasonably easy to find and mature turfgrasses
estimate of the P subsistence level for adequate are very good at finding P when they need it.
turfgrass growth is around 12 ppm (parts per mil- Consequently, planning a P fertilization program is
lion). Phosphorus levels recommended by local relatively easy.
laboratories are usually accurate. However, subsist-
ence levels can be affected by many factors, and a
Potassium
little experimentation is sometimes required. If
your turfgrass is not responding to fertilization as Maintaining adequate potassium (K) levels on fine
you think it should, at least one nutrient may be textured soil is fairly easy, but it can be more dif-
deficient. Soil tests and laboratory recommenda- ficult on soils with very high sand contents. On
tions may often lead you to the missing nutrient. sand, potassium is not tightly bound and leaches
Apply fertilizer containing the suspected nutrient to easily. Potassium soil analysis and laboratory rec-
some small areas at the rate used previously with ommendations are accurate and it is widely
unacceptable results, and fertilizer containing a believed that excess K is not toxic to turfgrass, nor
high rate of the nutrient to other small areas and does it have detrimental environmental effects.
compare responses. That procedure will usually Consequently, a little too much doesn’t hurt, or so
demonstrate whether or not the suspected nutrient we believe.
is deficient in the fertilizer mix. If P is deficient it is Potassium fertilization has been linked to par-
often because of turfgrass competition with trees or tial relief from numerous turfgrass stresses
other flowering plants for soil P or because soil pH (Turner and Hummel, 1992). As yet, we do not
is below 5.5 or above 7.5. know exactly why potassium has such desirable
There was a time when we recommended that stress relief qualities, but we know it works.
turfgrass managers determine an adequate annual Potassium is second to nitrogen in its concentra-
P to apply based on turfgrass use and soil P present, tion in turfgrass plants (Turner and Hummel,
and make one or two applications per year to apply 1992). It is not a constituent of important bio-
sufficient nutrient. Because of the dangerous envi- logical molecules, but it is involved in many
ronmental effects of excess P in surface water men- metabolic reactions and in the maintenance of
tioned earlier, we no longer recommend that cell turgor, and is often responsible for maintain-
procedure. Now we recommend that P be applied ing electrical gradients across membranes.
in regular small applications and watered in imme- Potassium is sometimes applied at rates equal to
diately with about ¼ inch (6.4 mm) of irrigation those of N, especially on sandy soil. However,
whenever possible. We also recommended applica- that is typically more than is needed.
tions of P during establishment, but this too has Potassium is sometimes applied to warm-season
changed. It is true that seedling turfgrass has a dif- turfgrasses as they enter dormancy to help protect
ficult time assimilating P in its early stages of root them from cold and desiccation during the winter.
growth. However, the establishment period is also However, there have been mixed reports on whether
the best time for P runoff to occur because the soil or not such a program is actually effective. Research
is practically bare and the turfgrass root system is in Massachusetts determined that late-fall applica-
not strong enough to hold the soil together. After tions of K reduced winter damage on perennial
seedlings have begun to grow, it may be wise to ryegrass, a cool-season grass, but results in North
apply a light application of P for support, but the Carolina on hybrid bermudagrass (Cynodon dacty-
old recommendations of starter fertilizers high in P lon × Cynodon transvaalensis) suggest that late-
are no longer valid. We have not experienced a season K applications had no effect on winter
decline in growth or delay of turfgrass establish- hardiness (Peacock et al., 1997; Webster and
ment substantial enough to warrant a P application Ebdon, 2005).
at establishment. Soils low in P should be adjusted Fertilization of K at about half the rate of N
before establishment by tilling P into the topsoil. applied is usually adequate. However, K fertiliza-
As long as adequate P is present in the soil, germi- tion should be adjusted for soil tests and other site
nation and seedling growth should proceed as conditions based on observation and site history.

Understanding and Prescribing Nutrition 103

 The secondary macronutrients now include micronutrient fertilizers in their prod-
and the micronutrients uct lines. That is primarily because there has been
such a strong industry movement toward sand-
Soil tests from respectable laboratories are typi-
based sports fields and putting greens that are
cally very accurate. However, most laboratory
considerably more likely to be micronutrient defi-
analyses, including those involving soil nutrients,
cient than native soil. Native soils, especially those
become less accurate with decreasing concentra-
polluted by human activities, are as likely to have
tions. Reported amounts of the primary and sec-
toxic levels of micronutrients as they are to be
ondary macronutrients, except perhaps for sulfur,
deficient.
are highly accurate, but reports of micronutrients
Iron (Fe) is the most important micronutrient for
are harder to evaluate and may have marginal
turfgrass growth and maintenance. Iron directly
accuracy. That is not to say that micronutrient
promotes chlorophyll synthesis, so its addition to
reports are not useful. More often than not, a soil
turfgrass almost always results in a nearly instant
test report will identify micronutrient deficiencies.
color change. The turf color usually turns blackish
If a micronutrient deficiency is suspected, test
at first, but changes to green after about 24 hours.
areas (as described earlier) are a good method for
Some scientists believe that the black response is
determining whether or not the nutrient should be
actually a staining response rather than a response
included in the fertilizer program. A small amount
to increasing chlorophyll or heme synthesis.
of a safe-for-turf chemical containing the nutrient
Whether or not a staining effect is involved, Fe
of interest can be applied with normal fertilizer to
clearly encourages chlorophyll synthesis and will
some areas and not to others, and the turfgrass
nearly always add green color to turf (Lee et al.,
response evaluated.
1996). For that reason, light applications of
chelated or soluble Fe may be used to cause turf to
Calcium, magnesium and sulfur green without encouraging shoot growth. Deficiency
testing for Fe by spraying some plots and not oth-
Calcium (Ca) is a primary constituent of most soils ers is not productive because a change in chloro-
but not all Ca is plant available. At low soil pH, Ca phyll concentration and color nearly always occurs.
is often present in insoluble forms that turfgrass A high pH will usually result in an Fe deficiency,
cannot assimilate. However, as you should recall, and the missing Fe causes the plant to yellow
lime (CaCO3) is the product best used to increase noticeably as a result of chlorophyll deprivation.
soil pH and it also contains soluble Ca. If the soil Many turfgrasses, especially those that prefer acid
pH is not too low but Ca is still not available, gyp- soils, need supplemental Fe in very small amounts.
sum (CaSO4), the same product as used to remove If your turf is reasonably healthy but yellow for no
soil salts, is the product of choice for Ca applica- apparent reason, it may need Fe. Centipedegrass
tion. Gypsum is considered to be pH neutral. (Eremochloa ophiuroides) is notorious for needing
Magnesium (Mg) may also be deficient at low supplemental Fe (Carrow et al., 1988). However,
soil pH. Epsom salts (MgSO4), available at most excess Fe can cause slight growth declines in creep-
drug stores, is a good product to use for testing of ing bentgrass (Agrostis stolonifera), annual blue-
Mg deficiencies. However, it is probably not the grass (Poa annua) and possibly other grasses (Xu
best product to use for treating Mg deficiencies and and Mancino, 2001). Growth may also slow fol-
may not be legal for direct soil application in your lowing an Fe application when air temperatures are
area. A Mg-containing fertilizer is a better choice high (Schmidt and Snyder, 1984). Light Fe applica-
once a Mg deficiency is identified. Sulfur (S) can tions, however, are relatively safe, and if growth
also be unavailable at low soil pH and this should slows, it does not have a severe negative effect on
be tested using agricultural sulfur (S2). Agricultural the turf.
sulfur or an S-containing fertilizer can be used to Manganese (Mn) is very important for the split-
correct deficiencies. ting of water molecules during photosynthesis.
However, it is rarely deficient and is one of those
nutrients that is typically difficult to assess by soil
Iron and other micronutrients
testing. A Mn-containing fertilizer should be tested
Micronutrient deficiencies are rare in native soils before investing in a potentially expensive program
but can occur. Most commercial fertilizer producers of Mn fertilization. Deficiencies in plant available

104 Chapter 7
Mn may be related to high soil pH. In that case, amount of K in the soil. However, excessive K can
lowering soil pH to below 7.0 will probably solve cause deficiencies of other macronutrients (Miller,
the Mn problem and also positively affect the avail- 1999). Potassium competes with Ca, Mg and other
ability of other nutrients (Snyder et al., 1979). nutrient cations for exchange sites on the soil CEC.
Copper, zinc and boron are rarely deficient, but If K is excessive it can cause soil deficiencies of Ca
may be unavailable at high pH. Many micronutri- and Mg (Woods et al., 2005).
ent fertilizers contain trace amounts of these ele- Earlier, I recommended that a K fertilization
ments. Test for deficiencies before investing in a program should be approximately half that of the
micronutrient program. Soil tests for micronutrients N applied. Based on research and observation that
are inexpensive and have value, but are not always is a good starting point for your fertilizer program
accurate (Faust and Christians, 2000). If a nutrient (Christians et al., 1979; Snyder and Cisar, 2000).
deficiency or toxicity is observed through soil test- In fact, your entire fertilizer program should be
ing, or suspected, simple experiments should be based on the amount of N applied. Basically, noth-
designed for verification before applying micronu- ing happens in turf without nitrogen. If, for
trients on a large scale. If tissue tests are used to instance, your soil is deficient in S or Mg your turf
determine micronutrient content, it should be may not take up those nutrients if N is not also
remembered that micronutrients, like macronutri- present (Goss et al., 1979). If green turfgrass is an
ents exist in plant tissue in different concentrations important goal for your site, application of mix-
depending on season and environment (Belesky and tures of Fe and N will increase green color better
Jung, 1982). than applications of either nutrient alone (Yust
et al., 1984). Iron is the nutrient that has to be
present for the heme synthesis pathway, the path-
Nutrient interactions
way that also produces chlorophyll, to occur.
Nutrient uptake and concentrations within turf- Nitrogen is an important part of the chlorophyll
grass plants are affected by each other (Christians, molecule and also encourages chlorophyll synthe-
1993). Turfgrass, for instance, may not take up K sis when present. The nutrients in combination are
if N is deficient (Petrovic et al., 2005). In fact, more likely to encourage chlorophyll synthesis
interaction between K and N for plant maintenance together than either would alone. You may be able
is common. Potassium is needed for a number of to grow green grass with less N input if Fe is also
physiological processes to occur and also for other applied. However, N must always be present for Fe
purposes within a turfgrass plant. Those purposes to be most useful. That is the case for nearly all
include helping to maintain cell turgor pressure and turfgrass activity. Nitrogen must be present in
osmotic potential, and helping to preserve electrical adequate supply for maximum turfgrass perform-
gradients across cell membranes (Carroll et al., ance. Otherwise, the remaining nutrients will not
1994). These functions are extremely important, encourage turfgrass performance as efficiently.
and if K is not present to perform them plants will Begin your fertilization program based on the
use sodium (Na) as a substitute. Plant use of Na in amount of N needed to meet expectations, then
place of K can lead to a decline in turfgrass quality adjust additional nutrients to combine with the
(Snyder and Cisar, 2005). Therefore, K fertilization program most effectively.
is extremely important if you expect to manage
turfgrass at optimum levels.
7.4 Chapter Summary
When sufficient K is part of an N fertilizer pro-
gram, the turf will use N more efficiently (Fitzpatrick Carbon, hydrogen and oxygen make up most of
and Guillard, 2004). For that reason, less N may be the dry matter in a turfgrass plant. To supply those
needed in the program and it can be adjusted to nutrients we must encourage photosynthesis, respi-
match your demands. As far as we know, K is not ration and transpiration. Sufficient light must be
damaging to the natural environment as excessive available for photosynthesis to occur. Once carbon
N and P can be. However, it can reach levels detri- dioxide is removed from the air surrounding a turf-
mental to plants. It would not really be accurate to grass canopy, fresh air must be available to replace
say that K can reach toxic levels, because that it. As transpiration cools our plants, the humidity
would suggest that excessive K could kill plants; in the air around them increases and transpiration
that is highly unlikely and would require a huge slows unless fresh air is available. Oxygen must be

Understanding and Prescribing Nutrition 105

present for respiration to occur. Air is limited in to manage your turf at acceptable levels. Proper
some soils and fresh air infiltration may also be planning, execution and adjustment of that pro-
limited. As oxygen is used by turfgrass roots, fresh gram results in healthy turfgrass and satisfied
soil air must be available to replace the air that has customers.
become oxygen deficient. Soil drainage, aerifica-
tion and other soil cultivation and management
practices are required to encourage adequate oxy- Suggested Reading
gen for sufficient root respiration. Brady, N.C. and Weil, R.R. (2008) The Nature and Prop-
Nitrogen fertilizer is required to synthesize chlo- erties of Soils, 14th edn. Prentice Hall, Upper Saddle
rophyll molecules, and high N availability tends to River, New Jersey.
encourage chlorophyll synthesis. Hence fertilized Carrow, R.N., Waddington, D.V. and Rieke, P.E. (2001)
turfgrass becomes more green. Nitrogen is also a Turfgrass Soil Fertility and Chemical Problems:
Assessment and Management. Ann Arbor Press,
part of every enzyme that catalyzes metabolic reac-
Chelsea, Michigan.
tions and a part of every protein. It is the most Hull, R.J. (1992) Energy relations and carbohydrate par-
conspicuous nutrient in turfgrass plants (Jones, titioning in turfgrasses. In: Waddington, D.V., Carrow,
1980). Properly applied N fertilizer programs R.N. and Shearman, R.C. (eds) Turfgrass. ASA-CS-
encourage both shoot growth and root growth. SA-SSSA (American Society of Agronomy-Crop Sci-
They also encourage high turfgrass density and a ence Society of America-Soil Science Society of
nice green color. A good N fertilization program America), Madison, Wisconsin, pp. 175–206.
helps turfgrass out-compete weeds and resist path- Turner, T.R. and Hummel, N.W. Jr. (1992) Nutritional
ogen and insect invasion. Too much N, however, requirements and fertilization. In: Waddington, D.V.,
can encourage certain diseases and other problems. Carrow, R.N. and Shearman, R.C. (eds) Turfgrass.
ASA-CSSA-SSSA (American Society of Agronomy-
Too much N causes plants to retain more water,
Crop Science Society of America-Soil Science Soci-
becoming more succulent and more easily damaged ety of America), Madison, Wisconsin, pp. 385–440.
by traffic. Excessive N also encourages rapid shoot
growth, usually at the expense of root growth and
that can result in a very weak root system. A typi- Suggested Websites
cal N fertilization plan should be a part of the Duble, R.L. (2010) Turfgrass Fertilization. Texas A&M
maintenance schedule, but it should be flexible and University. Available at: http://aggie-horticulture.tamu.
adjusted for weather conditions such as tempera- edu/plantanswers/turf/publications/fer tiliz.html
ture, rainfall and humidity. (accessed 12 March 2010).
If sufficient N is present, turfgrass plants will Georgia Turf (2010) A Balanced Soil Fertility Program is
respond to P and K fertilization when needed. Essential in Turfgrass Management. Available at:
Phosphorus is important for the formation of cel- http://mulch.cropsoil.uga.edu/fertility/ (accessed 3
lular energy and for DNA and RNA synthesis. August 2010).
Potassium helps to maintain electrical gradients Mississippi State University Extension Service (2010)
Home Lawn and Turf in Mississippi. Available at:
across cell membranes and cell turgor. It also has
http://msucares.com/lawn/lawn/fertilization/index.
to be present for many metabolic reactions to html (accessed 12 March 2010).
occur. Sometimes Ca, Mg or S are required and Penn State Cooperative Extension, Berks County (2010)
occasionally micronutrients are missing from the Turfgrass – Fertilization. Available at: http://berks.
soil solution. A regular program of soil testing extension.psu.edu/mg/hgargd/turffert.html (accessed
helps to determine the fertilizer program required 12 March 2010).

106 Chapter 7
 Irrigation and Water Management
8

Key Terms
A water window is the length of time that you have available to irrigate your site each day. It is not the length of
time required, but the length of time available.
The term precipitation is normally used to describe natural rainfall. It may also be used to refer to other naturally
occurring climatic forms of water such as hail, snow, sleet and even dew. We also use it to refer to irrigation
and other forms of simulated rainfall.
Static pressure refers to the pressure in a closed irrigation system when no water is moving. Although static
pressure is constant throughout the system, it is really only useful for irrigation design when it is considered
at the water source. Once the water starts moving, friction becomes a factor and the pressure declines.
Dynamic pressure, also called working pressure, is the pressure that exists at a given point in an irrigation system
when it is operating. Dynamic pressure changes throughout the system, declining as a result of friction loss
as it travels from the source and increasing or decreasing with elevation. A system must be designed to have
adequate dynamic pressure at each sprinkler head.
A backflow preventer is a sophisticated check valve that prevents the siphoning of water from an irrigation system
backwards into a potable water supply. Without a backflow preventer, pesticides, fertilizers and any number
of other materials could flow into sprinkler heads and siphon back into a public water source.
The friction factor is a calculated number that irrigation designers use to help them choose the proper pipe size
required to supply the individual irrigation heads in a zone.
Evapotranspiration (or ET) is the term used to describe the total water lost through a combination of evaporation
and transpiration from a turfgrass or any other cropping system.
The ETo is the evapotranspiration reference standard determined by entering specific weather parameters into a
mathematical model developed to calculate ET from a reference crop. The reference crop chosen for this
calculation happens to be a grass.
The Kc is a crop coefficient determined by scientific study of a particular species in a particular location.
The ETc is the estimated evapotranspiration of a particular crop on site or at a location nearby.
Precision irrigation management is the mapping of a large turfgrass area for sectioning into management units
based on irrigation requirements, followed by adjustment of the irrigation system to best accommodate the
needs of each unit. Once the site is mapped and the units determined, similar units are combined and
irrigated independently based on need.
Water use rate is the amount of water a turfgrass species extracts from the soil over a given period during
adequate precipitation.
Leaf fire is a measure of leaf browning of a turfgrass caused by long periods without precipitation.
Drought tolerance is the ability of a turfgrass plant to survive long periods without precipitation.

8.1 Manage Your Irrigation and Drainage

fell in less than 30 minutes. It was not a tornado, a
for Site-specific Objectives
hurricane or a cyclone, it was just a rain storm. It
Over the course of my life, I have been fortunate to was incredible and, needless to say, drainage was a
live in areas of the USA that have either too much problem. Drainage was a particular problem
or too little rainfall. Not long ago, I experienced a because the event occurred in a relatively dry por-
summer storm when over 2 inches (5.1 cm) of rain tion of the USA where drainage is often ignored.

©CAB International 2011. Turfgrass Physiology and Ecology (G. Bell) 107
 It has been my experience that regions where make your job easier, but that knowledge is really
large amounts of rainfall occur tend to have poor not necessary. It is your job to know how irrigation
irrigation systems and regions where little rainfall systems work, not how to install or repair them.
occurs tend to have poor drainage systems. As a You need to know the design. You can learn the
turfgrass manager, you need to be intimately famil- installation and repair later and in far less time.
iar with both irrigation and drainage. If you are In earlier times, I was hired as a supervisor into
managing high-quality, high-value turf, both irriga- two different industries where I had absolutely no
tion and drainage are major concerns for you. In experience. However, in spite of the fact that I did
most areas of the world, periods of dry weather not know how to make the products, I did know
and periods of wet weather occur annually regard- how to manage people, organize a work force and
less of overall climate. If you are not prepared for make intelligent decisions. Therefore, in both cases,
both, you are going to lose grass. I was very successful and once I earned their
In this chapter, we will address both too much respect, my employees were happy to help me learn
water and too little water. In nearly all regions, dry how to do their jobs. Knowing how to do their jobs
periods occur, and turfgrass requires irrigation to made my job easier but it wasn’t absolutely
look and function best during those periods. necessary.
Irrigation is not only a science but an art. It requires I don’t wish to offend our agricultural colleagues,
complete flexibility for weather changes and site- but a central pivot irrigation system is the simplest
specific differences in soil, light and airflow envi- of irrigation systems. If you don’t understand how
ronments. Plant competition and ecological systems central pivot irrigation works, you can’t possibly
have to be considered, and the limitations of water hope to understand a simple home lawn system
use or irrigation system design have to be part of and most certainly not a system for a golf course or
irrigation planning. Drainage is more of a common- athletic field complex. Those systems are far too
sense technology than irrigation, but in some situa- complicated for you. Several of my students have
tions it requires some creative ideas and problem commented to me after they entered management
solving. Although most turfgrasses can survive sev- positions that they wished they had paid more
eral weeks of drought and recover, it only takes 1 or attention in irrigation class. Now they have to
2 weeks to die under submerged or waterlogged design a system, make an addition to a system, or
conditions. Remember, even if you are dealing with oversee a design and installation by an independent
an extremely well-drained soil, like pure sand in a contractor. If you don’t understand basic irrigation
dry climate, it is still going to rain occasionally. No system design, you are not qualified for those tasks.
matter how well drained your soil is, the water has In the opinion of many who might consider hiring
to have someplace to go or you are going to be deal- you, being proficient in irrigation design is part of
ing with saturation or submersion. You have to your job.
have a plan, even if that plan is as simple as drain- When it comes to irrigation systems, turfgrass
ing the area into a collection swale or cistern where people are the very best. We may lag behind the
excess water can be pumped off-site. expertise of other agricultural industries in some
plant management methods, but we are the very
best in irrigation. This text is not going to provide
8.2 Irrigation Management
you with an education in turfgrass irrigation sys-
Students often ask why most turfgrass college cur- tems. I hope that you will further your education in
riculums include an irrigation class where they that area with more informative books and classes.
have to learn about the calculations required of However, we will discuss the basics of irrigation
irrigation design and where they might have to design because you need to know those basics to
learn about methods involved in central pivot irri- make good management decisions.
gation or other non-turf systems. The answer is
this: you are not training or taking classes or read-
Basic irrigation design
ing this book so that you can be a technician. You
are training to make decisions for technicians. No Everything about an irrigation system is affected by
matter how deeply you respect a highly qualified its water source. The water source determines how
technician, and you should, that is not your job. much area you can irrigate, how large your pipe
Knowing how to be an irrigation technician would has to be, how many heads you can put on a zone,

108 Chapter 8
and how many zones you can run at one time. The irrigation systems, large or small. The water win-
whole idea of water source is based on a simple dow determines how large your water source must
concept called the “water window”. be to fulfill your irrigation needs. You may use a
The water window is the length of time available public water source, an irrigation pond, a well or
to irrigate everything that you need to irrigate each some other source. Assuming that there is enough
day. It is not the length of time required to irrigate water available at the source to satisfy all of your
your site, it is the length of time available to irrigate irrigation needs, the size of the pipe and the pres-
your site. In other words, if you want to be able to sure in the pipe determine how much water you
irrigate your entire athletic complex in one night have to work with. Your public system or your
and the complex closes at 20:00 and opens at 7:00 pump must be able to fill that pipe continuously at
during the longest day of the year, your water win- the pressure required. Before you start buying
dow is 11 hours. If you only require the capability pumps or pipe, however, you must first determine
of irrigating half your complex each night, your how much water is required to irrigate the target
water window is still 11 hours but you are allowing area. To keep this discussion simple, let us assume
two windows (22 hours) to complete the job. If it that you will be irrigating a home lawn with water
requires 16 hours to pump all of the water that is from a public water source.
required to irrigate your complex and you need the Just because you are using a public water source
capability to irrigate it all in one night, then your from a municipal system does not mean that you
window is still 11 hours but your system is too have an unlimited supply of water. Some systems
small. You need a new water source. Hence, your are barely adequate to supply drinking water. You
water window determines how large your water could be dealing with a system that has a small
source has to be to irrigate your site. It is always supply line on 30 pounds of pressure per square
best to have a larger water source than you need inch (psi) or 26 bar. That source will not supply
because you never know when you might lose your very much water and you will have to plan accord-
water or have to replace a pump or get into some ingly. Assuming that you have a 5⁄8-inch (16-mm)
other situation where your turf has not received water meter on 30 psi, your source will theoreti-
water for some time, and you need to get it all cally supply about 4 gallons per minute (gpm) (or
irrigated as fast as you can. If your water source is 0.908 m3/h) from a 1-inch (25-mm) supply pipe
a public water supply, you do not have much con- (Hunter Irrigation Innovators, 2009a). Now let us
trol over the rate of supply. However, if your water assume that the lawn you intend to irrigate is
source is a pumping station from a lake or other 10,000 square feet (sf) (929 m2) and that during the
water feature with a nearly unlimited supply of hottest, driest periods, you will have to supply irri-
water, the supply rate is mostly up to you. In that gation at about 2 inches (51 mm) per week. Under
case, the supply usually depends on how much you a worst-case scenario, the system would break
or your sponsors are willing to spend for the sys- down and the homeowner would be forced to
tem. When making those decisions, the reliability apply all 2 inches of irrigation in 1 day as soon as
of the pumping equipment and the rest of the sup- the system was repaired to keep the grass alive.
ply system are of profound importance. In addi- A few calculations bring you to the conclusion that
tion, a backup pumping system must be considered at the maximum rate of supply, the lawn will
and included in the purchase. require 12,467 gallons (47 m3) of water and 52
You can find instruction manuals and tables at hours to irrigate completely with 2 inches of pre-
most irrigation supplier websites (see the list at the cipitation. Consequently, the homeowner cannot
end of this chapter); these are very helpful for irrigate in 1 day, but if nothing unforeseen hap-
understanding irrigation principles and helping you pens, the homeowner could potentially supply the
learn to design and install an irrigation system. amount of irrigation needed during the driest sea-
There are also books available on turfgrass irriga- son by running the system all night for four nights
tion systems and drainage which are listed at the per week. However, if the homeowner happens to
end of this chapter. I will not attempt to instruct prefer the capability to irrigate everything in 1
you on how to design or install an irrigation sys- night (8 hours), a total irrigation time of 52 hours
tem, but you need a basic knowledge of irrigation is not workable. This particular system, because of
systems to use them most efficiently. Certain basic a poor water source, would require a booster pump
principles and construction apply to all turfgrass and probably a storage tank to supply enough

Irrigation and Water Management 109

water to irrigate the entire site within an changes. However, if you were to check the pres-
8-hour water window. If you are not in control of sure at any point when the system was at rest, and
the water supply, you have to adjust to the situa- then check the pressure at the same point when the
tion. Now, let us assume that you have the same system was operating, the pressure would be lower
system, but a new water source that can meet the during operation because water is moving through
8-hour water window, and you will typically run the system. When the system is operating, the pres-
all zones on the system for 4 hours, two nights per sure is not fixed or static, it is moving or dynamic.
week during the driest time of the year. During operation, you are not measuring static
Our new water source has a 1-inch (25 mm) pressure, you are measuring dynamic pressure.
water meter and 60 psi (52 bar) pressure on a Dynamic pressure, the actual working pressure,
1.25 inch (32-mm) supply line. This particular changes throughout the system based on elevation
source has a maximum output of about 26 gpm: and friction (Fig. 8.1). Pressure decreases as water
over six times greater than the old source (Hunter is pushed uphill but increases as water is pushed
Irrigation Innovators, 2009a). In that case, we can downhill. Any restrictions inside the pipe cause
supply 12,467 gallons (47 m3) of water to the lawn turbulence and friction, which result in pressure
in 8 hours, which will meet the requirements that loss. Our theoretical home lawn application, for
we determined for the water window. instance, will have to have a backflow preventer
Assume that, in this example, we used a measure on the main line before any water flows to the
of static pressure to estimate the water available to irrigation heads. The backflow preventer is a spe-
operate our system. However, static pressure is a cial valve that only allows irrigation water to flow
measure of potential pressure, not actual pressure. in one direction. Consequently, groundwater or
Dynamic pressure, often called working pressure, surface water cannot flow backwards from the
is the pressure that you actually have to work with irrigation heads to the potable water system and
at any particular point in your system. If you were contaminate it. Valves have a manufacturer’s rat-
to check the pressure on the system when it was at ing describing how much pressure is lost as water
rest with no irrigation running, that would be the moves through them. A backflow preventer on a
static pressure of the system. The static pressure small system might cause a 5 to 10 psi pressure
would be basically the same throughout the system, loss, so the pressure in the system could be 10 psi
except for the effects of gravity when elevation lower than that of the source after water passes

42 psi 40 psi
B C

Pressure
Pressure
increases
decreases
12 ft.
14 ft.
D
A 44 psi
50 psi

Pressure at point A = 50 psi (pounds/sq.in.)

Water passes through 2 fittings (–0.4 psi), 16 ft of pipe (–2.2 psi) and climbs 14 ft (–6.1 psi)
Pressure at point B = 42 psi
Water passes through 10 ft of pipe (–1.4 psi)
Pressure at point C = 40 psi
Water passes through 2 fittings (–0.4 psi), 12 ft of pipe (–1.7 psi) and falls 12 ft (+5.2 psi)
Pressure at point D = 44 psi

Fig. 8.1. Water flowing through pipe causes friction, resulting in pressure loss. Pressure is also lost when water is
forced uphill but pressure increases as water flows downhill.

110 Chapter 8
through the backflow preventer. Pipe fittings also All but the simplest of irrigation systems are
cause friction, so each time water flows through a divided into sections called zones. Each zone is
tee or an elbow, pressure is lost. Last, and possibly operated independently by a valve. The amount of
most important, the pipe itself causes friction. The water available from the source determines how
amount of pressure lost is affected by the roughness many irrigation heads can be controlled by a single
of the pipe and the distance that the water has to zone. Zones may also vary by the type of head used
travel through the pipe. The material used to make in the zone or by the type of plant material in the
a pipe, plastic, steel, etc., differs in its roughness zone. Planting beds, for instance, should be on dif-
and in the amount of friction caused as water ferent zones from turf because bedding plants and
flows through it (Table 8.1). The size of the pipe turf are usually irrigated on different schedules.
also affects the amount of friction that occurs. The friction factor helps to determine when pipe
A small pipe causes greater friction than a sizes can be reduced without seriously affecting the
large pipe. Consequently, less pressure is lost in a flow rate at the heads on a zone. Each head should
1.25 inch pipe than is lost in a 1.00 inch pipe. distribute approximately the same amount of water
Therefore, pipe size is extremely important to the uniformly on an area basis (e.g. gallons precipita-
amount of water that can be supplied to the irriga- tion per square foot). How this is accomplished is
tion heads that are farthest from the source. not vital to you learning how to irrigate effectively,
Irrigation designers use a number they call “the but it is interesting. Let us suppose that you have
friction factor” to help them determine what pipe calculated a friction factor of 1.50 (see Box 8.1)
size is required to adequately supply each head in and that you plan to use CL 200 PVC pipe. You
the system (Box 8.1). can use a friction factor short cut chart (Table 8.2)

Table 8.1. The amount of friction as water flows through a pipe is influenced by the velocity of the water flowing,
the size of the pipe and the type of pipe used. The velocity and the pipe size determine the flow rate. Irrigation
supply companies provide friction loss charts that make it relatively easy to calculate pressure from one point in a
pipe to another. A sample of a friction loss chart for comparison of polyvinylchloride (PVC) schedule 40 plastic pipe,
polyethylene (PE) SDR (standard dimension ratio)-pressure rated tubing, schedule 40 standard steel pipe (steel) and
type K copper tubing (copper) is given below. Notice that velocity does not change with the type of pipe used. That is
because velocity is a function of flow rate and pipe size, and is not affected by friction. fps = feet per second;
gpm = gallons per minute; psi = pressure per square inch.

Type of pipe PVC PE Steel Copper

Size of pipe 1 in. 1 in. 1 in. 1 in.

Velocity Pressure Velocity Pressure Velocity Pressure Velocity Pressure

Flow (gpm) (fps) loss (psi) (fps) loss (psi) (fps) loss (psi) (fps) loss (psi)

1 0.37 0.03 0.37 0.04 0.37 0.07 0.37 0.05

2 0.74 0.12 0.74 0.14 0.74 0.26 0.74 0.18
3 1.11 0.26 1.11 0.29 1.11 0.55 1.11 0.38
4 1.48 0.44 1.48 0.50 1.48 0.93 1.48 0.65
5 1.85 0.66 1.85 0.76 1.85 1.41 1.85 0.98
6 2.22 0.93 2.22 1.06 2.22 1.97 2.22 1.37
7 2.59 1.24 2.59 1.41 2.59 2.63 2.59 1.82
8 2.96 1.59 2.96 1.80 2.96 3.36 2.96 2.33
9 3.33 1.97 3.33 2.24 3.33 4.18 3.33 2.90
10 3.70 2.40 3.70 2.73 3.70 5.08 3.70 3.53
11 4.07 2.86 4.07 3.25 4.07 6.07 4.07 4.21
12 4.44 3.36 4.44 3.82 4.44 7.13 4.44 4.94
14 5.19 4.47 5.19 5.08 5.19 9.48 5.19 6.57
16 5.93 5.73 5.93 6.51 5.93 12.14 5.93 8.42
18 6.67 7.13 6.67 8.10 6.67 15.10 6.67 10.47
20 7.41 8.66 7.41 9.84 7.41 18.35 7.41 12.73

Irrigation and Water Management 111

 Box 8.1. A friction factor helps to determine pipe size.
A friction factor is used to determine the size of Pv = the pressure variation between the valve and the
lateral line pipes required to operate within a last sprinkler head on the zone, not to exceed
desired range of flow rates without exceeding a 10% or certainly no greater than 20%; and
detrimental amount of pressure loss (Hunter Lc = the length of pipe from the control valve to the
Irrigation Innovators, 2009b). To maintain consist- farthest head in hundreds of feet.
ency and adequate coverage, all of the sprinklers
on a zone should be operating within ±10% of and Let us assume that we have a zone of sprinklers that
certainly no worse than ±20% of their desired oper- operate best at 30 psi (pounds per square inch) (Po =
ating pressure. 30 psi) and we want to operate all of the sprinklers on
The friction factor is calculated by: the zone within 20% of the recommended pressure
(Pv = 0.20). The farthest sprinkler head is 400 feet
Ff = (Po × Pv)/Lc from the valve (Lc = 400 ft./100 ft. = 4.0). Then the
friction factor is:
Where:
Ff = the friction factor, which is the loss of pressure Ff = (Po × Pv)/Lc = (30 × 0.20)/4.0 = 1.5 psi/100 ft.
that you are willing to accept per 100 feet of
pipe; Once the friction factor has been determined, a friction
Po = the sprinkler operating pressure suggested by factor short cut chart (Table 8.2) is used to determine the
the manufacturer; proper pipe size for each section of the zone (Fig. 8.2).

Table 8.2. A portion of a typical friction factor short cut chart. Once a friction factor is calculated and the flow rate
that is needed for that particular section of a zone is known, the pipe size required can be estimated using the chart.
A pipe that is too large will always work, but it is also more expensive. gpm = gallons per minute.

Friction factor

Pipe 1.00 1.25 1.50 1.75 2.00

Size (in.) Type Max. gpm Max. gpm Max. gpm Max. gpm Max. gpm

0.50 CL 315 PVC 2.30 2.60 2.80 3.10 3.30

0.75 CL 200 PVC 4.50 5.10 5.70 6.10 6.60
1.00 CL 200 PVC 8.70 9.80 10.80 11.70 12.60
1.25 CL 200 PVC 16.00 18.10 19.90 21.70 23.30
1.50 CL 200 PVC 22.90 25.80 28.50 30.90 33.20
2.00 CL 200 PVC 41.00 46.30 51.10 55.50 59.70
2.50 CL 200 PVC 67.80 76.40 84.30 91.70 98.50

to determine the size of pipe required to supply an higher pressures than spray heads, usually at about
adequate amount of water to each section of a zone 50 psi (43 bar) for residential applications. Spray
(Fig. 8.2). On a system with small zones, such as heads usually work best at about 30 psi (26 bar).
our lawn example, using the same pipe size through- Rotor heads are tasked to cover much larger areas
out a zone is typical. However, on systems with than spray heads, and although they use water at a
large zones, such as that shown in Fig. 8.2, reduc- much higher rate, they also take much longer to
ing pipe size in the farthest sections of the zone can apply precipitation over their assigned area because
save a considerable amount of money on the area is so much larger. Therefore, if spray heads
installation. and rotor heads are mixed in the same zone, the
Rotor-type sprinkler heads (the heads that rotate area covered by the spray heads always receives
as they spray) should not be included on the same more water than the areas covered by the rotors
zone as common spray heads (those that do not and the turf is either too wet at the spray heads or
rotate as they spray). Rotor heads work best at too dry under the rotors.

112 Chapter 8
 Main line (30 psi)

Pipe diameter

1.50 in.
20 gpm
1.25 in. 1.25 in. 1.25 in. 1.25 in.
18 gpm 16 gpm 14 gpm 12 gpm

10 gpm
1.00 in.
2 gpm 4 gpm 6 gpm 8 gpm

0.50 in. 0.75 in. 1.00 in. 1.00 in.

Pipe Friction factor 40 ft.

1.50
Size (in.) Type Max. gpm
0.50 CL 315 PVC 2.80
Irrigation heads
0.75 CL 200 PVC 5.70
(each head requires 2 gpm)
100 CL 200 PVC 10.80
1.25 CL 200 PVC 19.90
1.50 CL 200 PVC 28.50

Fig. 8.2. A simple schematic of a typical zone in a large irrigation system. Irrigation heads are set 40 ft. apart making
the total length of pipe from the main line to the last head 400 ft. Each section of pipe is labeled with the flow that it
is required to sustain and the pipe size that is required to carry that flow based on a friction factor of 1.5 psi/100 ft.
A friction factor table is included for reference. gpm = gallons per minute; psi = pounds per square inch.

Getting back to our home lawn irrigation sys- assigned to supply a 180° radius because the 90°
tem, it is now time to choose the types of irrigation head only has to cover half the area that the 180°
heads that we need to accomplish the tasks in each head has to cover. Most irrigation companies sell
zone. We will need both spray heads and rotors to nozzles for spray heads that vary in output so as to
complete the system because we have small lawns supply the same precipitation rate under the same
on each side of the house and large lawns at the pressure and flow regardless of radius. You can
front and back (Fig. 8.3). If we ignore the planting usually purchase those nozzles in fixed radii of 45°,
beds, we need at least two zones, one for spray 90°, 180°, 270° and 360°. However, not all spray
heads and one for rotors. However, as we design nozzles are designed to deliver uniform flow at dif-
our system, we will probably find that our water ferent radii, so you have to be aware of what you
source is not sufficient to supply both the front are buying. The most versatile spray nozzles are
yard and the back yard at the same time, and we those that can be adjusted for radius from just a
will have to irrigate them individually on different few degrees to 360°. On those nozzles, you some-
zones. times have to adjust the flow rates when you adjust
We will use typical residential spray heads and the radii to match your needs. When using rotors,
rotors. Spray heads are adjustable by using nozzles you have to adjust nozzle size to accommodate
that differ by radius and area of coverage, and uniform precipitation rates. A rotor tasked to cover
adjustable for flow rate by adjusting a screw that a 180° arc, for instance, needs a nozzle with twice
moves to fill the supply tube and works much like the output of a rotor tasked to cover a 90° arc on
an adjustable valve. As each spray head on a zone the same zone.
can potentially supply an equal amount of water, a Rotors are highly adjustable for radius, spray
head in the corner of a zone that sprays a 90° angle, precipitation rate and distribution pattern.
radius will supply twice as much water to its The radius on most rotors is adjustable over almost
assigned area as a head on the side of a zone 360° and the spray angle that controls the height of

Irrigation and Water Management 113

 will be some deterioration in the system and the
water source over time.
Zone 5 Our next step should be to choose the heads that
we will use and lay out the system based on the
amount of water that we have available. We will
use spray heads to apply precipitation in arcs up to
17 feet (5 m) from the heads and use rotors for
larger arcs up to 40 feet (12 m). On a perfect sys-
tem, the arc of one spray head or rotor should
extend to the head(s) adjacent to it. We call that
head-to-head spacing (Fig. 8.4). In that case, every
Zone 4 part of the turfgrass area to be irrigated is covered
by at least two sprinklers. In a rectangular area of
turf that type of coverage is relatively easy to
accomplish, but in most areas some creative place-
ment will be necessary to achieve uniform cover-
age. It must also be remembered that even in the
very best equipment the edges of the spray pattern
supply slightly lighter precipitation than the inner
Zone 2 Zone 3 portions of the pattern. The portion of the pattern
Zone 1 closest to the head supplies the largest amount of
precipitation and the pattern diminishes slightly as
the distance from the head increases. For that rea-
son, we can sometimes adjust the arc on opposing
1 square = 1 ft. (0.3 m) Rotor head heads to provide uniform distribution when head-
Spray head to-head coverage is not entirely possible.
Consider the diagram in Fig. 8.3. To adequately
Fig. 8.3. A simple home lawn irrigation system irrigate our turf, we need 16 spray heads to cover
designed to supply precipitation to the various lawn the side lawns, zones 1 and 4. Each 180° head in
sections (white areas). Zones 1 and 4 contain spray
zone 1 must irrigate a radius of 12 feet (3.7 m) and
heads and zones 2, 3 and 5 contain rotor heads.
according to manufacturer’s specifications applies
Additional zones would need to be added to supply
precipitation to the planting beds (dotted areas). approximately 1.26 gpm (4.8 l/min) at 30 psi (26
bar). The heads in zone 4 must irrigate a 13 feet
(4 m) arc and require 1.86 gpm (7 l/min) at 30 psi
the spray pattern above the ground is adjustable by (26 bar). The 90° heads need half the water of the
using high- or low-angle nozzles. The precipitation 180° heads, so the total supply required for zone 1
rate can also be adjusted by the size of the nozzle is 7.56 gpm (29 l/min) and for zone 4 is 11.16 gpm
chosen and the distribution pattern can be slightly (42 l/min). As our source can supply 26 gpm, we
altered by adjusting a deflector screw in the head. could run all of these sprinklers on one zone, but
Some of the adjustments made to a spray head or our system will be more versatile if we split them
rotor alter the amount of water that it needs to into two zones. It is likely that these two sections of
complete its assignment and in nearly every case
this alteration can be estimated reasonably closely.
By referring to manufacturer’s specifications for
nozzle output and summing the water requirements
of each nozzle on the zone, you can estimate the
amount of water needed to supply the entire zone
and determine whether your water supply is suffi- Fig. 8.4. In a well-designed turfgrass irrigation system,
cient to meet the needs of the zone or whether the each spray head or rotor covers an area that ends
zone will need to be divided into smaller zones. at the head(s) adjacent to it. Arcs of 90° and 180°
Some designers also build a comfort factor into are normally used to accomplish this head-to-head
each zone to account for the likelihood that there coverage. Arcs of 45°, 270° and 360° are also common.

114 Chapter 8
lawn will have different irrigation requirements, identified easily by puddles in the direction the
and by placing them on different zones we will be rotor is facing when it stops turning. However, if it
able to accommodate our water needs more effi- turns intermittently the fault will not be noticed so
ciently. If we use small nozzles in the rotors on the easily and it will still create problems. Although
front lawn, we can irrigate the entire lawn on one portable sprinklers will probably always be more
zone, zone 2, but we can probably increase water common than automatic irrigation systems on small
use efficiency by splitting it into two zones, zones 2 sites, the days of manually operated permanent
and 3. That also makes it possible to operate the systems are long over. However, automatic systems
system with less water flow in case our water source still have to be monitored and inspected regularly or
diminishes over time. We can also try long-radius there will be problems that require immediate atten-
nozzles or short-radius nozzles on the outside edges tion every time the weather gets dry.
to get the best possible uniformity with the short-
radius nozzles that we will use in the center.
Determining your irrigation needs
Uniform precipitation rates are an important
goal for an irrigation system and are the reason Irrigation efficiency is becoming increasingly
that we have discussed basic system design. important as demands for potable water become
Irrigation systems employ multiple heads to irrigate greater. For most people, turfgrass irrigation is a
single areas to gain uniformity of precipitation. luxury. The importance of adequate water supplies
Consequently, when water is plentiful and natural for home and industry far exceed the importance of
rainfall is close to adequate, irrigation system turfgrass irrigation. Irrigation for the production of
deformities are rarely evident; however, they tend food crops is also more important than turfgrass
to stand out when environmental conditions are irrigation. However, there are alternatives to the
more severe. Extended dry periods usually define use of potable water for irrigating turf (Thomas
the inadequacies of a system rather clearly, but et al., 2006). Irrigating turf with used or partially
once they appear, it may be too late. At the very treated wastewater is becoming more common and
least, expensive hand watering will be required in techniques for the desalination of seawater have
the areas of turf where the system fails to meet improved and are becoming more cost-effective
expectations. If the system is not designed properly (Throssell et al., 2009). In many cases, effluent
to provide uniform coverage, areas that do not water can be used for turfgrass irrigation and pro-
receive enough irrigation will have to be hand vide acceptable turf without detrimental effects
watered, or the system will have to be run longer to (Mancino and Pepper, 1992). However, irrigating
provide enough precipitation to those areas – with effluent has its own particular problems, espe-
resulting in too much precipitation on the areas cially during excessively dry periods (Hayes et al.,
that are adequately covered. A turfgrass manager 1990; Dean et al., 1996). When effluent water is
has to be intensely aware of the strengths and used a system of soil, plant and atmospheric moni-
weaknesses of the irrigation system that his/her toring is a firm requirement of irrigation program-
turfgrass depends on. Otherwise, normally small ming (Lockett et al., 2008). As you read and study
problems can become severe when conditions are this text you should come to the conclusion that a
extremely dry. monitoring system should be practiced at all levels
We all seem to realize that irrigation systems are and for all components of plant management.
likely to develop leaks over time. We realize that However, irrigation is perhaps the most important
wires and heads may be broken, that valves will of those components to be monitored. The more
require repair or replacement and that pumping wisely that we use our water, the more likely that
systems will require overhaul or replacement. we are going to have enough water to satisfy our
However, we tend to overlook the smaller things future needs. We have the means to manage irriga-
that can create big problems. Nozzles, for instance, tion precisely if we have the knowledge and experi-
wear out. As they wear, the irrigation pattern ence to use it.
becomes unbalanced and the heads use more water
with less uniform coverage. Nozzles can also get
Evapotranspiration
plugged occasionally and so block distribution.
Gears on rotors can wear out and stop working. The term evapotranspiration (ET) is a simple com-
When a rotor stops turning, the problem is usually bination of two familiar words, evaporation and

Irrigation and Water Management 115

transpiration. Factors that affect evaporation affect calculate ETo with accuracy, the altitude and lati-
ET and factors that affect transpiration also tude at the site and several weather factors have to
affect ET. Scientists like to use and study be known. Air temperature and wind speed 2 m
evapotranspiration because it can be calculated from above the soil surface must be known, as well as
a quantitative model. Most scientists, however, will humidity and solar radiation. These factors must
be quick to explain that it is best not to take eva- be recorded and converted, if necessary, to the
potranspiration too literally. If recent environmental proper units of measure before being used to deter-
conditions estimate evapotranspiration at 0.3 inches mine additional parameters, such as vapor pressure
(7.6 mm) over the past 72 hours that does not neces- and soil heat flux, that are needed to complete the
sarily mean that every turfgrass manager in the area calculation. Obviously, the calculation of ETo is
needs to apply 0.3 inches of irrigation to relieve the complicated and best left to on-site weather station
deficit. In fact, if you are managing a large site, your software or to local meteorologists or agronomists.
irrigation needs may be for 0.35 inches in one area, However, once ETo has been calculated the result is
0.20 inches in another area and 0.30 inches on the not complicated to use and can be an important
remainder. Irrigation is as much an art as it is a part of an irrigation program.
science and you need to use history, knowledge and The reference ET (ETo) is the result of a meteoro-
examination to help estimate how much water each logical calculation that is designed to calculate
area of your site really needs. evapotranspiration from a standard vegetated sur-
In most cases, ET does not have to be completely face. Fortunately for us, the standard vegetated
replaced to manage turfgrass at acceptable levels surface happens to be a grass cover crop. However,
(Qian and Engelke, 1999). Most turfgrasses have ET differs according to grass species and other
root systems extensive enough to acquire deep or important factors. In fact, ET differs by any
tightly held soil moisture remaining from earlier number of factors and that is why it is best used as
periods of rainfall that saturated the soil. For that a starting point and should be adjusted for your
reason, an intensively managed Kentucky bluegrass specific management needs.
(Poa pratensis) fairway might require as little as Once ETo is known, it can be adjusted for the
60% of ET replacement in areas where periods crop, in our case the particular grass, on-site
without rainfall last no longer than about 4 to 6 (Aronson et al., 1987b). The adjustment factor for
weeks (Shearman et al., 2005). Relatively drought- a particular crop is called its crop coefficient (Kc).
tolerant species such as tall fescue (Festuca arunidi- The Kc is a simple multiplier that customizes the
nacea) and bermudagrass (Cynodon dactylon) ETo to the transpiration attributes of a particular
generally do very well under irrigation practices of species. The ET rates of grasses vary substantially
60% ET in many locations (Fu et al., 2004). Even by season (Carrow, 1995). In the case of a turf-
poorly rooted species such as creeping bentgrass grass, the determination of Kc may also differ by
(Agrostis stolonifera), velvet bentgrass (Agrostis mowing height and other factors (Devitt et al.,
canina) and colonial bentgrass (Agrostis capillaris) 1992). The multiplication of ETo by Kc results in
may grow reasonably well at 60–80% ET during the reference ET standard for a particular crop,
the summer, and can nearly always handle such which is called ET c.
deficits in the spring and fall (DaCosta and Huang,
ETo × Kc = ETc
2006b). Obviously, deficit irrigation can be prac-
ticed at higher levels and longer periods in areas If you are familiar with the process, you can
with fairly frequent rainfall or during seasons when calculate your own crop coefficients specific to
rainfall is reasonably frequent (DaCosta and your site or you can simply adjust ET c, or even ETo,
Huang, 2006a). A good turfgrass manager adjusts based on history, knowledge and on-site experi-
the irrigation deficit by season and when condi- ence. The crop reference evapotranspiration, ET c,
tions vary from normal. is a standard, like ETo. The ET c is the ETo adjusted
The reference ET, called ETo, is the basic unit of for a specific crop. However, it is based on the crop
an ET calculation. It is most commonly derived when all environmental conditions are excellent for
from the Penman–Monteith scientific model plant growth and maintenance and no stress is
(Monteith and Unsworth, 1990). The Penman– present. The ET c is defined as the evapotranspira-
Monteith model is a combination of climatic fac- tion of a disease-free, well-fertilized crop grown in
tors that affect evapotranspiration. In order to a large field under optimum soil water conditions

116 Chapter 8
and achieving full production under the given cli-
matic conditions (Allen et al., 2004). Obviously,
conditions are rarely, if ever, perfect so the ET c
must be adjusted for weather and on-site condi-
tions before a truly accurate value of evapotranspi-
ration can be considered. We could continue this
discussion to include the adjusted ET c, appropri-
ately called ET c-adj, but instead, I will direct you to
the references at the end of this chapter for further
education in evapotranspiration and turn, in the
next section, to a more appropriate subject, preci-
sion agriculture, to further identify locally adjusted
irrigation needs. Once you obtain ET from your
weather station or any number of meteorological Fig. 8.5. A representation of a zoysiagrass (Japanese
sources, you can adjust it to determine irrigation lawngrass, Zoysia japonica) golf course fairway
need in small units of a large site using precision mapped for turf greenness using a set of reflectance
turfgrass management. sensors mounted on a spray boom. These areas are
Evapotranspiration adjusted for plant need is a believed to define differences in irrigation need, but
further inspection is required to determine if that is truly
very useful irrigation tool. It is also a good model
the case. A mobile unit that measures soil moisture
to study as a learning device because all of the would be a better choice for mapping irrigation need.
factors that have a major effect on the irrigation However, reflectance sensors are fast and can be
needs of your plants are included in the model. used as long as site inspections to determine that
There are several different methods for calculat- water deficit is truly the source of the difference in
ing ET and their accuracy among locations, greenness follows on from the spectral mapping. The
conditions and plant species is controversial darkest areas indicate the darkest green turf (the
(Allen et al., 2004). Rather than enter into a best watered turf) and the lightest areas indicate the
debate, I suggest that they are all relatively lightest green turf. The abbreviation ET can refer to
accurate and that you should use the one that is ETo, the reference ET, or to ETc, the crop-specific ET.
The number designations represent irrigation need. For
the most common in your region, and is the most
example, 0.7 ET identifies areas that require 70% of
readily available, and stick with it. Changing the calculated ET (ETo or ETc) that is consistently used
from one model to another will only make it at the site, while 1.1 ET refers to an irrigation need of
more difficult to assess your site and to customize 110% of the calculated ET.
the calculated ET to best satisfy your specific
needs. Once the model or source of information
is selected, your site must be divided into indi- irrigation requirements (Krum et al., 2010). The
vidual units based on irrigation need. maps are then used to section the area into indi-
vidual management units. Further observation will
facilitate the fine tuning of irrigation requirements
Precision irrigation management
over the entire site, resulting in a substantial con-
Irrigation need has to be constantly monitored. servation of water and an increase in positive
Individual areas should be identified and placed turfgrass response.
into management units that have common need Notice in Fig. 8.5 that individual irrigation pat-
(Carrow et al., 2010) (Fig. 8.5). The darkest areas terns caused by poor irrigation head spacing or
in Fig. 8.5 (designated ET 0.7), for instance, may system malfunctions are not visible on this fairway.
generally require 70% of the calculated ET (either Poor irrigation patterns are generally obvious in
ETo or ET c) that you choose to use, while areas precision mapping using spectral sensors. This par-
that have a designation of ET 1.1 require 110% of ticular system appears to be operating efficiently. It
your calculated ET. Observation and record keep- is unfortunate that areas of need are not distributed
ing are the activities most useful for identifying in such a manner that the irrigation system can be
areas of similar irrigation need. Large areas may adjusted to irrigate each management unit inde-
be mapped for management purposes using soil pendently. Although that is rarely the case, the
probes and/or reflectance sensors to estimate their irrigation system can normally be adjusted to

Irrigation and Water Management 117

provide more efficient irrigation of individual man- complicated than the simple condensation of water
agement units. Individual heads can be adjusted for vapor from warm air touching a cold object, the
arc or for nozzle size to accommodate more or less rapid cooling of the air is the primary principle
irrigation on areas where these needs have been behind the condensation. Dew on turf is mostly
identified. The adjustments will never be perfect, caused by warmer air containing sufficient water
but the savings in water and the increase in turf vapor coming in contact with colder grass leaves.
uniformity and value can be substantial (Carrow Dew usually forms shortly before sunrise when the
et al., 2010). difference between the temperature of the lower air
and the temperature of the turf is greatest, but it
may form at any time during the night. Dew is
Plant and environmental factors that affect
important as a form of precipitation. If a turfgrass
evapotranspiration and water use
is in need of water, it will absorb dew through its
The environmental factors previously identified as leaves, thus helping to avoid a stressful drought
affecting evapotranspiration, or ET – humidity, situation. Lack of morning dew on turfgrass in a
temperature, wind speed and radiation – are easily particular area when a general dew formation has
defined by their physical effects on evaporation. occurred is often an indication of drought stress.
However, these factors and others also affect tran- A turfgrass manager should be observant for such
spiration and turfgrass health. Because this text is a situation and investigate it if it occurs. Some spe-
primarily based on the physiological components cies, zoysiagrasses (Zoysia spp.) in particular, are
of photosynthesis, respiration and transpiration, it slow to form dew on leaves (Fig. 8.6). Consequently,
is necessary to discuss these four environmental in a mixed or side-by-side stand, one species may
factors in more detail to understand how they have dew on its leaves and another may not.
affect these processes. When relative humidity is high, evaporation is
low and transpirational cooling is negatively
affected. If the temperature is warm and sunny and
Humidity
humidity is high, photosynthesis is proceeding rap-
Humidity is a measure of the amount of water idly but cooling is inhibited by the reduction in
vapor in the air and is usually reported as relative transpiration. That creates a stressful situation for
humidity, the amount of water vapor in the air warm-season (C4) grasses and an extremely stress-
compared with the amount of water vapor that the ful situation for cool-season grasses. Under windy
air can potentially hold at current conditions. conditions, high humidity is less of a concern
Relative humidity is normally reported as a per- because wind increases evaporation and helps to
centage (%). Consequently, if the relative humidity counteract the high humidity. As always, there are
is 20% the air is dry, at 50% the air holds half the multiple factors that affect evaporation, hence
water it is capable of containing and at 90% the air transpiration. Humidity is only one.
is quite humid. Air temperature has a direct effect The need for irrigation is less during humid peri-
on relative humidity. As temperature increases, the ods. Declines in both evaporation and transpiration
air can hold more water. So when the air tempera- limit the need for additional water. The decline in
ture is 50 °F (10 °C) and relative humidity is 50%, irrigation need should be reflected in the reported ET.
the air is holding less water vapor than when the Consequently, the turfgrass manager has an accurate
air temperature is 90 °F (32 °C) and relative humid- estimate and should adjust irrigation accordingly.
ity is 50%. A combination of heat and humidity is However, the irrigation manager does not have a
a stressful condition for most turfgrasses, especially measure of heat buildup in the turfgrass due to high
cool-season (C3) grasses. photosynthesis, high respiration and poor transpira-
As warm air can hold more moisture than cold tion. An infrared canopy temperature device may be
air, water vapor in the air condenses on cold of use for comparing turfgrass temperature with the
objects. As warm, humid air comes in contact with surrounding air temperature (Throssell et al., 1987;
a cold object, the air cools rapidly. As the air cools Jiang et al., 2009). Often at midday the canopy tem-
it can no longer hold all of the water vapor that it perature will be higher than the air temperature and
was holding when it came into contact with the light misting or syringing may be required on impor-
cold object, so water condenses on the object, tant turfgrass in spite of the reduction in evaporation
forming dew. Although dew formation is more due to environmental conditions (Carrow, 1993).

118 Chapter 8
Fig. 8.6. The small rectangular plots in the middle of the photograph and the stand of turf in the foreground are
of zoysiagrass (Japanese lawngrass, Zoysia japonica). The square plots are bermudagrass (Cynodon dactylon).
The bermudagrass appears a lighter color because it is covered with dew. The zoysiagrass is damp but the dew
is not noticeable. The zoysiagrass is not drier than the bermudagrass, but it has a waxy coating on its leaves that
discourages the formation of large water droplets.

Cooling is particularly important on golf course increase during periods of high temperature unless
putting greens where localized dry spots, a condition it is restricted by water deficit or another plant
often observed on sand systems, occur (Tucker et al., stress. Although turfgrass plants may stop growing
1990). The most effective means of cooling turfgrass or turn yellow, they can generally sustain periods of
is to provide a combination of syringing with fans above normal temperature as long as humidity is
(Guertal et al., 2005). Misting and syringing increase not particularly high and winds are not extremely
humidity above the turf and add to the problems high or extremely low. Sufficient soil water is nor-
previously discussed. Therefore, water cooling by mally enough to maintain turfgrass health through
misting or syringing should only be practiced when it exceptionally warm weather (Perdomo et al.,
is imperative to sustain turf. A critical canopy tem- 1996). However, when extended drought condi-
perature is normally considered to be 104 °F (40 °C), tions occur, sunlight is intense and the air tempera-
the temperature at which enzymes begin to denature. ture is high, turf canopy temperatures may be as
Misting and syringing are labor intensive, therefore much as 40 °F (22 °C) above air temperatures
expensive, and their cooling effects are short lived, (Steinke et al., 2009).
less than an hour (DiPaola, 1984). Combining Evaporative demand is high and irrigation need
syringing with air movement caused by fans can increases when the weather is exceptionally warm.
extend the cooling period for up to an additional The ET rate will increase as temperature rises, indi-
hour (Guertal et al., 2005). Although their effects are cating the need for additional irrigation. Heat
short lived, misting or syringing may be necessary to stress also makes it more likely that a plant will
help turfgrass sustain itself through especially diffi- succumb to additional stresses such as traffic or
cult environmental conditions. As manager you disease. Sufficient irrigation is usually the key to
should be able to determine when misting or syring- resisting heat stress and related stress factors.
ing is required both by sight and by consideration of As you learned in earlier chapters, high tempera-
recent weather conditions. tures increase molecular activity. Consequently,
photosynthesis and respiration as well as all other
plant metabolic processes increase when the
Temperature
temperature increases. This rapid rate of less than
High temperature increases evaporation compared perfectly efficient plant metabolic processes
with low temperature. Transpiration will also produces substantial heat in addition to high air

Irrigation and Water Management 119

and soil temperatures. For that reason, transpira- relation to irrigation, extremely calm weather has
tion must increase to lower plant temperature. The the same effect on carbon dioxide availability.
plant root system must be sufficient to handle this Although the plant stomates may be open during
increased demand and water must be available. calm periods, little carbon dioxide is available
A grass that has been preconditioned to drought, because the air is not refreshed, causing photosyn-
meaning that it has already experienced mild thesis to slow and affecting respiration.
drought, is more likely to remain healthy through a For a particular temperature and humidity, an
sustained drought period or a period of exception- increase in wind speed causes an increase in evapora-
ally high temperatures than one that has not been tion and transpiration. These weather phenomena
preconditioned (Jiang and Huang, 2000a). cause an increase in ET. If high winds continue for
Preconditioning to drought encourages grasses to extended periods it will be necessary to increase irri-
grow deeper root systems (Jiang and Huang, gation. However, it must be remembered that irrigat-
2001). Therefore, over irrigation or frequent rain- ing in high wind is not particularly effective. Irrigation
fall during the spring encourages failure during a patterns are partially destroyed, causing the precipi-
hot, dry summer. Short periods of dry weather and tation to miss areas where it is needed. In addition,
no irrigation during the spring encourage drought much of the irrigation evaporates before it reaches
and heat tolerance, especially in cool-season grasses, the ground. If the nights are relatively calm, as often
during the summer. You should be aware of these occurs in some locations, irrigation can be performed
effects and manage spring irrigation accordingly. in the early morning before the next windy day.
However, if the nights are also windy, postponing
irrigation is advisable until calmer weather occurs or
Wind speed
until it can no longer wait. Such times may require
Environments that are extremely detrimental to hand watering of valuable turf. Hand watering
turfgrass growth include high wind combined with reduces the amount of irrigation water lost to evapo-
high temperature and low humidity. Under such ration and allows a competent technician to visibly
conditions, evaporation and transpiration are so assess and satisfy irrigation need.
rapid that turfgrass plants may enter water deficit
situations by late afternoon in spite of the presence
Radiation
of plentiful soil water early in the day. Plant wilting
may occur because plant roots cannot absorb water Solar radiation is a form of energy just like heat.
fast enough to meet transpiration demand or Consequently, intense solar radiation over a long day
because the soil water at the depth where the roots increases evaporation and transpiration. It also
are plentiful has been exhausted. More than likely, increases soil and plant warming, and encourages
transpiration demand is too fast for the roots to rapid and sustained photosynthesis. Consequently,
keep up. Consequently the plant cells near the sto- the need for irrigation increases when solar radiation
mates signal water deficit and the stomates close is high and days are long. Solar radiation is also part
either fully or partially, depending on evaporative of the ET calculation. It may seem that the energy
demand. increase in heat during high solar radiation and long
Assuming that light is available, the condition of days would be sufficient to account for the increase
wind-renewed continuous carbon dioxide in ET. However, such is not the case. Radiation is an
availability is perfect for rapid photosynthesis. The important component of evaporation and transpira-
warm conditions are perfect for rapid respiration tion; partly because of its important effect on photo-
of the carbon sequestered by photosynthesis and synthesis and partly because solar energy, like heat,
rapid plant growth should ultimately occur. encourages chemical reactions.
However, sufficient transpiration cannot be attained
to satisfy the system and the chance for rapid
Irrigation frequency
growth and development is lost. Because transpira-
tion cannot meet evaporative demand, the stomates If a means of irrigation is available and water is
close, effectively suppressing carbon uptake, and available for it, supplying sufficient water to your
photosynthesis slows. Unless substantial reserve turf is easy. Building a root system is not. Irrigation
carbohydrates are available, respiration and growth frequency is best determined by the seasonal health
also slow. Although it is rarely mentioned in of the turf and by the current strength of its root

120 Chapter 8
system. There are several factors that have been high and low rainfall and variable weather condi-
discussed that not only affect ET, they affect irriga- tions. The turfgrass manager always knows which
tion frequency. However, the extent of the root areas of the site are most likely to dry out quickly
system and the season of the year are the two most and have to be monitored closely. The manager
important factors that influence irrigation fre- also knows and has mapped the areas that tend to
quency. When ET is extremely high, and turfgrass hold water, where traffic should be avoided follow-
wilt occurs almost daily, it will be necessary to ing major precipitation events.
increase irrigation frequency. In that case, the Although environmental conditions influence
upper layer of soil, about 3 inches (7.6 cm), that irrigation frequency, the most important factors are
houses most of the turfgrass root system is drying root depth, root mass and season. In general, deep,
out too rapidly and irrigation frequency needs to infrequent irrigation is the most effective practice
be increased. That problem is likely to occur during for managing turfgrass health (Richie et al., 2002).
periods of high transpiration. Frequency adjust- Under most circumstances, poor root systems are
ments will normally be made on a seasonal basis the result of poor management preceding stressful
according to climate. However, they will also be periods. Frequent irrigation during active growth
made based on the extent of the turfgrass root sys- periods does not encourage adequate root growth.
tem and the plant’s ability to grow new roots. The daily irrigation often practiced on sand sys-
Like nearly all plants, turfgrasses lose cell turgor tems, for instance, is generally not required during
and wilt when they are seriously low in water. the spring and fall. Daily irrigation results in shal-
However, healthy grasses can survive a certain low soil wetting and may remain in the thatch
amount of wilt so it is not necessary to increase where it evaporates easily (Sass and Horgan,
irrigation frequency when wilt is noticed late in the 2006). Although winter desiccation is not uncom-
day on an occasional basis. Turfgrasses that are mon, the most important season for irrigation is
irrigated more frequently may maintain higher leaf summer. Warm-season grasses tend to have deeper
turgor pressures, but they are not necessarily more root systems than cool-season grasses (Christians,
healthy (Jordan et al., 2005). However, irrigation 2007). They also are better at performing photo-
should be applied more frequently when wilt is synthesis and use less water during hot weather
seen regularly. Wilt manifests itself as a grayish or (Feldhake et al., 1983). Consequently, they are bet-
bluish coloring that, with some experience, is easily ter adapted to summer stress and water deficit.
detected. The leaf blades loose their rigidity to wilt Cool-season grasses are more likely to experience
and do not spring back after traffic. Consequently, water deficit stress over the summer. In fact, the
footprints are deeper and easier to distinguish management practices of the fall and spring preced-
when the grass is wilting. Turfgrass managers look ing the summer may determine whether or not a
for wilted turf late in the day to assess irrigation cool-season grass will survive.
need. They also use other techniques to test soil All grasses, both cool-season and warm-season,
moisture before turf wilting. grow roots rapidly in the spring. Although warm-
Some turfgrass managers are quite proficient at season root growth continues to occur slowly over
testing moisture in soils they are familiar with by the summer, cool-season grass roots diminish
using a knife blade inserted into the soil. The ease (Fig. 8.7). Warm-season grass roots may diminish
with which the knife blade penetrates the soil tells slightly in the late summer into the fall but not at
the manager approximately how much soil water is anywhere near the rate that cool-season roots
present. Other managers may cut off a golf shaft diminish during the summer (Sartain, 2002).
then cut a vertical sighting slot in it for use as a During the summer, cool-season roots die faster
miniature soil probe. With experience, it is fairly than they can be replaced. A cool-season root sys-
easy to determine the moisture content of the soil tem cannot improve itself over the summer. Instead
by looking at its profile exposed in the vertical slot. it declines during the period. Therefore, trying to
More technical managers may use a soil moisture encourage cool-season grasses to grow deeper roots
probe to acquire more exacting measurements. during the summer only stresses the grass further.
Regardless, a good irrigation program includes The depth and mass of the root system as the plant
methods, means and protocols for continuous enters the summer often determines whether or not
monitoring of irrigation need. It also includes it will survive. Once the summer season begins, the
records of the site by specific location over years of root system continues to decline until the fall. It is

Irrigation and Water Management 121

 60 Rapid Rapid
50 spring summer

Root mass (mg)

growth decline
40
30 Winter
(5 months) Fall
20
Fall
10
0
Sept Oct Nov Apr May Jun Jul Aug Sep Oct
Month
3
Fig. 8.7. Root mass of a 1.5-cubic inch (25-cm ) section of creeping bentgrass (Agrostis stolonifera) roots throughout
a growing season in Columbus, Ohio. Observations were made on the 1st of each month.

highly unlikely that the turfgrass manager can do (7.6 cm) deep and the plants are under stress, it
anything to prevent this decline. In fact, any man- will do no good to water deeply. The deep water
agement practice that causes damage to the turf will be wasted. Stressed plants do not have suffi-
results in decline of cool-season grasses during cient energy to extend their root systems for
summer. If a plant’s root system is poor in May, by water. As manager you must adjust your practices
September the plant may be dead. to irrigate frequently with less water to keep the
Proper irrigation management during the spring short root systems supplied until conditions
and fall for cool-season grasses and during the improve and the plants are healthy enough to
spring and early summer for warm-season grasses grow more roots. An observant turfgrass manager
can be used to promote root growth. During these knows that cool-season plants provided with fre-
periods when the respective grasses are healthy and quent natural rainfall in the spring are more likely
growing, withholding irrigation temporarily will to need frequent irrigation during the summer.
encourage root growth (Fu et al., 2007). Frequent The manager also knows that these same plants
irrigation will discourage root growth. When will have to experience some light irrigation defi-
turfgrass plants are healthy they respond to water cits in the fall and the following spring to help
deficit by extending their root systems in search of encourage the root expansion that will get them
water. If sufficient water is always present, turf- through the next summer.
grasses may respond by allocating more energy to
shoot growth and stem growth and less energy to
Mowing and fertilization for low water use
root growth. Consequently, if the plants have not
experienced water stress before dry periods the Proper mowing and fertilization can help to reduce
plants will be poorly prepared to tolerate the dry water use. From a common sense standpoint, it
conditions. would seem that high mowing resulting in longer
During periods of rapid growth and generally leaf blades would cause a turfgrass to transpire
healthy environmental conditions, it is best to more water. Technically that is correct (Feldhake
withhold irrigation until at least the top 2 inches et al., 1983). A high-mown turf has more biomass,
(5 cm) of soil in a turfgrass stand is dry or until the a deeper root system and takes up more water
first signs of wilt appear. During those periods, (Madison and Hagan, 1962). The deeper root sys-
water deeply rather than frequently. A healthy tem of a higher mown turf, however, also helps it
turf will extend its root systems deeper into the to avoid dehydration and wilt. Because its roots go
soil to accumulate water. An unhealthy turf or a deeper, there is more water available to it. As men-
turf under stress, however, will not have the tioned earlier, short-mown turfgrass, because it has
energy to grow new or deeper roots. An unhealthy a shorter root system than higher mown turf,
or stressed turf will require close observation dur- requires more frequent irrigation. More frequent
ing dry periods. When necessary, adjust irrigation irrigation means more frequent evaporation of the
frequency to account for the depth of the root irrigation and more likelihood that a small portion
system. If your turfgrass roots are only 3 inches of the irrigation may run off or leach deeper into

122 Chapter 8
the soil profile than the short root system can program slightly low in N and slightly high in K
reach. A low-mown turf may not use quite as much will maximize water use efficiency.
water as a high-mown turf but turf quality declines
more rapidly between rainfall events (Feldhake
8.3 Species Adaptation
et al., 1984). Therefore, the relatively rapid decline
to Low Water Use
of low-mown turf may lead the manager to believe
that more irrigation is needed than is actually Any discussion of species adaptation to drought
required. The combination of all of these minor or water use must begin with a determination of
elements results in a larger irrigation requirement objectives. Are your objectives to manage the turf-
for low-mown turfgrass compared with the same grass site to remain green all through the growing
turf at a higher height of cut. The lower mown season or are you willing to accept some yellow-
plant uses slightly less water, but because of the ing and possible dormancy during dry periods?
multiple factors discussed requires more irrigation. Both cool-season grasses (Aronson et al., 1987a;
Higher mown turf may use slightly more water, but Fry and Butler, 1989) and warm-season grasses
will maintain turf quality longer between irrigation (Beard and Sifers, 1997; Huang et al., 1997) and
or rainfall events, thus reducing irrigation losses to their cultivars differ in propensity for surviving
factors other than plant uptake. Consequently, drought and for resisting leaf fire during dry con-
higher mown turf requires less irrigation. ditions. Scientists use leaf fire, a measure of the
In Chapter 7, you learned that high N fertiliza- browning of leaves as they dry during periods
tion leads to increased plant succulence. Increases without precipitation, as a measure of visual qual-
in N fertilizer cause increases in leaf turgor, but ity (Ebdon and Kopp, 2004). A plant’s tolerance
increases in K fertilizer can cause decreases in leaf to leaf fire is not the same as its propensity to
turgor (Carroll and Petrovic, 1991). Consequently, resist drought (Carrow, 1996). If your objectives
if the N fertilizer rate is relatively low and the K are to irrigate to maintain visual quality, leaf fire
fertilizer rate is relatively high, plant water use may resistance is most important to you. If you are
decline. However, if the N rate is relatively high looking for a species or cultivar that can survive
and the K rate is relatively low, plant water use may long periods without irrigation, drought tolerance
increase. If you use effluent water for irrigation, N is your primary concern. When researching spe-
fertilizer rates will be likely to need adjusting for cies and cultivars for drought or leaf fire resist-
the N in the effluent water (Devitt et al., 2008). ance in your particular area, you must keep your
Ebdon et al. (1999) found that as N fertility objectives firmly in mind. You must also consider
increases, water use generally increases in Kentucky the adaptability of the species for your site.
bluegrass; in the same study, they also found that Although tall fescue is a drought tolerant cool-
when N and P were high, increasing K increased season grass it would not be considered more
water use, but when N and P were normal, increas- resistant to wilt than hybrid bermudagrass
ing K lowered water use. It has also been found (Cynodon dactylon × Cynodon transvaalensis) or
that slow-release N, presumably because growth is buffalograss (Buchloe dactyloides). However,
more consistent, helps to lower water-use rate research suggests that where tall fescue is better
compared with quick-release N in both adapted to the environmental conditions than
St. Augustinegrass (Stenotaphrum secundatum) hybrid bermudagrass and buffalograss, it grows a
and some ornamental plants (Saha et al., 2005). deeper root system and is more tolerant of dry
Based on research and experience, the best mow- conditions (Qian et al., 1997). Consequently, a
ing height for reducing turfgrass water use is the species or cultivar will not demonstrate its poten-
one that maximizes root depth. Therefore, the tial for drought or leaf fire resistance in a location
highest mowing height will produce the best where it is not particularly well adapted.
results. The highest mowing height that meets your Water use rates, the speeds at which a turfgrass
customer expectations for aesthetic value and/or takes up water when sufficient water is present,
playability is the best choice for minimizing irriga- are presumably a measure of the amount of water
tion needs. The best fertilizer program is a bal- required by a particular turfgrass species. However,
anced one. Sufficient nutrition must be present for trying to determine the water use rate of any par-
turfgrass to grow most efficiently using the least ticular species is difficult. Again, it is likely that the
possible water. Within that realm of sufficiency, a best adapted species for the location will have an

Irrigation and Water Management 123

advantage in water use rate. Also, the water use Site inspection and drainage design
rate of a particular species does not necessarily
Natural drainage is, of course, a function of eleva-
mean that it is or is not drought tolerant. For
tion. However, water not only runs downhill on the
instance, perennial ryegrass (Lolium perenne),
surface of the soil, it can infiltrate the soil and sub-
colonial bentgrass (Agrostis capillaris) and tall
surface drain. A basin or depression surrounded by
fescue were found to use approximately equal
higher elevation cannot surface drain, it has to
amounts of water in a study in Oregon, but tall
subsurface drain. The water cannot move over the
fescue is clearly more drought tolerant than the
soil, it has to move through the soil. Unless the soil
other two species (Doty et al., 1990). However, in
in the basin is deep and sandy, water flowing into
a subirrigated study in Arizona, bermudagrass and
the basin during periods of runoff will take some
zoysiagrass (Japanese lawngrass, Zoysia japonica)
time to either evaporate or subsurface drain. As a
used less water than St. Augustinegrass and tall
turfgrass manager, you should be reluctant to
fescue, which is what would be expected at
depend on subsurface drainage, even in artificially
that location (Kneebone and Pepper, 1982).
constructed sand systems. Surface drainage is reli-
St. Augustinegrass is known to have poor drought
able. Subsurface drainage specifically constructed
tolerance, yet research in Florida suggested that it
for the purpose is also reliable if properly con-
could be grown on loam soil with as little as 0.5
structed, but has a finite lifetime (Murphy et al.,
inches (13 mm) of precipitation every 6 days
1993b). It may take 20 years or more, but properly
(Peacock and Dudeck, 1984). One reason that
constructed subsurface drainage will eventually
water use rates are so hard to define is because
fail. If you are the turfgrass manager at the end of
grasses are likely to use more water than necessary
that finite lifetime, you will not have a high opinion
when more water is present. Bermudagrass, for
of subsurface drainage. To you, subsurface drain-
instance, can survive on very little water, but it will
age will not be reliable. Surface drainage, however,
use a substantial amount of water when enough
is timeless unless construction practices nearby or
water is present (Kneebone and Pepper, 1984).
unlikely natural events destroy it.
Consequently, the best choice of grasses are those
Sports fields and golf course greens specially
that are best adapted to the conditions at your site
constructed of sand under United States Golf
and have the best drought tolerance or leaf fire
Association (USGA) or similar protocols provide
resistance, depending on your objectives. Those
excellent subsurface drainage. However, even these
are the grasses that you can grow best at the low-
systems can become ineffective if surface drainage
est water use.
is ignored during construction. Consider a golf
course bunker or sand trap. If you are a golfer, are
8.4 Managing Water Overload
interested in turfgrass management, or have ever
In most regions of the world, periods exist where constructed a bunker, you will realize that in most
the soil is saturated for a short time. Rainfall is cases these bunkers are not just holes in the
sometimes sufficient to saturate the upper layers of ground. They are highly engineered entities with
soil and produce substantial runoff, perhaps even more than adequate drainage, geotextile mats,
flooding, but these conditions rarely last for long. special sand and other custom features. Golf
Turfgrass cannot function when submerged or course bunkers are specially constructed holes in
when rooted in saturated soil. It has been found the ground designed to provide longevity over a
that saturated soil even 15 cm deep affects creeping typical hole in the ground. Nonetheless, they are
bentgrass (Jiang and Wang, 2006). However, turf- still holes in the ground and they still have the
grass is perfectly capable of tolerating such condi- attributes of holes in the ground. It may take them
tions for short periods. Depending on species, longer for drainage to fail compared with the typi-
submersion in cool water or residing in saturated cal hole in the ground but it will, in fact, fail.
soil for a few days is not particularly stressful to Bunkers collect all of the debris, silt, clay, and
turfgrass and it normally recovers from such condi- organic and inorganic material from the surround-
tions quickly. However, if these conditions are ing watershed that drains into them. A 10-year-old
recurring or if they last for extended periods, the bunker that still drains well is rare and bunkers
turf will decline and may not survive (Beard and that collect debris from a watershed of substantial
Martin, 1970). size fail relatively quickly.

124 Chapter 8
 A basin or depression in a turfgrass stand is like than a prevailing slope or slopes of sufficient fall to
a bunker. The basin collects debris from the sur- drain your site. The water must also have a place
rounding area that drains into it. At first, the water to go. As ridiculous as it seems, people seem to
in a sand system drains down through the sand ignore drainage in favor of mounds, beds or other
faster than it can surface drain into the basin. features in locations where they want them to be. It
Eventually, however, the healthy turfgrass begins to is common to see standing water or dead grass near
produce thatch and that thatch begins to break constructed mounds and other sloping features
down into mat that exists at various levels of because the natural drainage was destroyed.
organic degradation. The mat begins to interfere The first order of drainage is to pick a place for
with water infiltration. In addition, dust collects on the water to go. That place should be in the natural
the sand system. Sediments and salts from rainwa- drainage pattern. Otherwise, you will be making it
ter and irrigation enter the system. Eventually, these particularly difficult on yourself. If you have pock-
barriers to infiltration worsen to the point where ets, depressions, basins, ditches or perhaps an
runoff occurs and the basin begins to accumulate entire large site that has to be drained, follow the
debris on a regular basis. The accumulating debris natural drainage pattern or your time and labor is
soon forms a soil layer that holds water and the likely to be for naught. Basic surface drainage is
subsurface drainage, as well as the turfgrass in the from the highest point to the lowest point, and if
basin, begin to deteriorate. Some basins are con- there are ridges between those two points, a path
structed by design. Other, far less detrimental, must be constructed so that runoff can flow around
basins occur when attempts are made to make a the ridge and into the lowest point in the area. If
perfectly level surface. It is probably impossible to that lowest point is not a lake or a creek then a
build a perfectly level surface from sand but even if ditch swale or other feature must be present or be
it is not, the sand would eventually settle inconsist- constructed to carry the water into a creek, pond or
ently, forming minor hills and basins. There are possibly a sewer. The water has to have someplace
some cases, in the game that Americans call soccer, to go even if you have to pump it there.
for instance, where the slopes that provide surface
drainage are detrimental to the game. In that case, Constructing subsurface drainage
a flat playing field cannot be avoided but the expec-
Surface drainage is most often preferable to subsur-
tations for the life of the field should be reduced.
face drainage, but in some cases subsurface drain-
Otherwise, designs for turfgrass systems should,
age is necessary for cosmetic or functional reasons.
without exception, provide for adequate surface
Constructing subsurface drains to accommodate
drainage. As a turfgrass manager you should seek
water removal from small areas is fairly simple to
out the basins on your area of responsibility and
design and fairly easy to construct. As usual, the
either deal with them or closely observe them.
design requires a place for the water to go. To get
Eventually, declines in turfgrass growth will occur.
it there requires a ditch of sufficient depth and
slope containing a pipe of sufficient size and con-
struction. Corrugated plastic drain pipe is light,
Drainage
flexible and inexpensive, and is available with or
Providing drainage to your area of responsibility is without holes and in different sizes. A 4-inch
usually easier than irrigating it, but not always. (12-cm) plastic drain pipe is easy to handle and will
During construction, natural surface drainage is drain sufficient water for most circumstances. If the
sometimes destroyed by poor design and it can be pipe is expected to take up water, it has to have
difficult to reestablish. Fine soils and compacted holes in it. A pipe without holes carries water from
soils have high runoff potential and poor infiltra- one place to another but it does not take up water
tion. Low spots in these soils have to be properly from a saturated area. To understand how subsur-
drained because infiltration is slow and the soil in face drainage works, you need to remember some
a depression will remain saturated for long periods. of your basic soil principles.
Whether the site is a home lawn or a playing field, If you place a soaker hose on a loam soil on top
turfgrass systems are generally sites for recreation of sand on top of gravel containing a pipe, what
and standing water or saturated soil is detrimental happens (Fig. 8.8)? If you remember, fine particles
to use as well as to turfgrass health. It takes more soak up water from more coarse particles. The most

Irrigation and Water Management 125

 puddles on compacted soil. You see it happen all
Turf the time. The physics tells you that the fine com-
pacted soil should be soaking up the water and
actually it is. It just takes a long time (Box 8.2).
Soil There is actually pressure on the water to enter the
soil, but the compacted soil pores are small and few

Ditch edge
and fill up quickly. Once the compacted soil fills
Ditch edge

Sand the physics tells you that the water has no other
place to go except into the surrounding soil, which
is less compacted and therefore more coarse. So as
Gravel Pipe long as the puddle exists, it fills the compacted soil
around it with water, but the transfer of that water
through the tiny pores of the compacted soil into
the less compacted soil is very slow. Hence, the
Fig. 8.8. A ditch containing a drain pipe backfilled with water remains in the puddle for a long period of
gravel around the pipe, sand above the gravel and time as it drains through the compacted soil. If
soil above the sand. Subsurface ditches like this one water drains from sand to highly structured soil,
can be constructed to carry water from intermittently the soil will take it up fairly quickly until it is full,
saturated soil into a drain. When the soil on the top then, if possible, it will slowly drain into another
of the ditch becomes saturated water moves into the medium.
sand. When the sand becomes saturated water moves Because of coarse-to-fine soil physics, the person
into the gravel and then into the pipe for drainage. The who applies soil-based sod over sand ends up with
sand speeds the drainage process by bridging the
dead grass. Whatever water exists in the system
particle size gap between the soil and the gravel.
remains in the soil that came with the sod. Water
will not drain into the sand until the soil in the sod
difficult part of that principle to understand is the above it is saturated. Consequently, the roots in the
time factor. It is easy to remember that water saturated soil have no oxygen with which to per-
moves from coarse particles to fine particles until form respiration. As you learned earlier, a turfgrass
you are asked what happens when water drains in that situation dies of drought when, in fact, its
from sand into compacted soil, and you get roots are surrounded by water. For the same rea-
confused. You know from experience that water son, a pipe surrounded by gravel under sand under

Box 8.2. If you pour a bucket of water onto a layer of sand on top of a layer of gravel on top
of a layer of soil containing a pipe, does the water drain into the pipe?
In this system, the water will drain into the pipe if the flow downhill through a ditch filled with sand fairly
soil around it is saturated, but depending on soil rapidly and through a ditch filled with gravel even
type, the soil will probably drain too slowly for the faster. Consequently, a ditch like the one in Fig. 8.8
pipe to be useful as a drainage water conductor. If without the soil on top would drain very fast. Not only
the bucket of water is of sufficient volume to saturate would the pipe fill as rapidly as water could infiltrate
the system and cause water to flow into the pipe, the the sand, the sand and gravel would also carry
water will have to be poured at a rate slower than the drainage water, but more slowly, downhill toward the
infiltration rate of the soil around the pipe or the drain. The problem with such a system is that if the
system will overflow before water enters the pipe. strip of sand on top of the ditch is wider than 2 or 3
However, if such a system is used as a drainage inches (5–8 cm), the ditch will always be visible
ditch it will probably drain saturated soil rapidly because the turfgrass growing on the sand will look
enough to be adequate under most conditions with- different from the turfgrass growing on the soil. The
out any water ever entering the pipe. The water will sand would leach nutrients much faster than the soil
drain without entering the pipe because the sand and it would always be dry except for periods imme-
and the gravel are also drainage devices. Water will diately following rainfall.

126 Chapter 8
a layer of soil with turfgrass on top drains slowly spray head used and other considerations. Each
and does not drain past soil saturation (Fig. 8.8). zone may be operated independently by a desig-
Nonetheless, it drains rapidly enough and com- nated valve and each head can be adjusted for
pletely enough to eliminate turfgrass decline result- output and for area covered. The most expensive
ing from saturated soil. Once the puddle effect is systems can be adjusted to automatically operate
removed, the soil drains naturally and it should be each head independently. Obviously, turfgrass irri-
rapid enough to prevent turfgrass damage. However, gation systems are complicated. However, the prin-
if periods of heavy rain intense enough to keep the ciples on which they are based are relatively simple
soil saturated for 3 or 4 days at a time are common, and easy to remember.
it would be better to fill the ditch with sand all the It takes pressure to move water through a pipe.
way to the surface in the area that tends to puddle. The size of the pipe and the pressure at the source
Just a small slit 3 to 4 inches wide is sufficient and determine how much water is available to the sys-
such a small slit is rarely noticeable in the turf. The tem. The highest point in the system will have the
small slit of sand will help the water move into the lowest pressure and the lowest point in the system
pipe faster. will have the highest pressure. Small pipes carry
Water drains from coarse particles to fine, but it less water and therefore must move water faster to
also drains faster from saturated fine soil into sand maintain the same flow rate as a larger pipe. The
than it does from saturated fine soil into gravel. In faster the water moves in the pipe, the greater the
the example (Fig. 8.8), under normal conditions it friction becomes and the harder it is to push the
is rarely necessary to cover the gravel with sand. water. Consequently, large pipes are preferable to
However, it will speed drainage if you do because smaller ones, but they are also more expensive, so
the particle size of the sand is more similar to the over-engineering the system is not always desira-
soil than the particle size of the gravel is to the soil. ble. The water window, the amount of time avail-
Therefore, when the soil becomes saturated, water able to irrigate the site, determines the rate at
will move faster from the soil to the sand than which water must be supplied to the system in
from the soil to the gravel. Nonetheless, the water order to accomplish the irrigation task. The
will move from saturated soil to gravel, it just amount of water required for a given irrigation
takes a little longer. The pipe should always be task is primarily determined by the plant material
surrounded by coarse sand or fine gravel to keep to be irrigated, by weather conditions and by con-
the soil from filling up the holes that allow the sumer expectations.
water to enter the pipe. If you happen to have Evaporation and transpiration combine to deter-
some old pipe that you are going to use for drain- mine how quickly water is used or lost from the
age and you intend to drill your own holes in it, turfgrass site. The combination of evaporation and
you only need to drill the holes on one side and the transpiration is called evapotranspiration (ET) and
holes need to go on the side that is down when the can be determined mathematically if the required
pipe is placed in the ditch. That is correct, the weather parameters are measured. There are many
holes go against the sand or gravel that you placed sources, including the Internet and government
in the bottom of the ditch before you placed the weather stations, available to determine the refer-
pipe on top of it. Consider air to be the largest of ence ET, which is referred to as ETo. The ETo can
particles. Water will not move into the air in the be adjusted by site-specific factors to determine the
pipe until it saturates the gravel or sand around the irrigation required fairly accurately. However, a
pipe. Therefore, it actually enters the pipe from the more accurate measure uses the ETo combined with
bottom, not from the top. a crop coefficient (Kc). The ETo multiplied by the Kc
for a particular species of turf, for instance, results
in ET c, a fairly accurate measure of water loss from
8.5 Chapter Summary
a turfgrass system containing that species.
Turfgrass irrigation systems are designed for and Although an ET c may be calculated from weather
are uniquely effective for uniform distribution of data accumulated from a station nearby, it is still
precipitation over large or small turfgrass areas. not completely accurate for estimating irrigation
They are normally divided into zones based on the need throughout an entire site. Site-specific man-
amount of water available, the species requiring agement practices can conserve water, save money
irrigation, management intensity, location, type of and improve the uniformity of the turfgrass on the

Irrigation and Water Management 127

site. Moisture data collection and/or spectral Puhalla, J., Krans, J. and Goatley, M. (1999) Sports
sensing of turf will result in designated areas called Fields: A Manual for Design, Construction and Main-
site-specific management units that require more tenance. Ann Arbor Press, Chelsea, Michigan.
or less irrigation than is indicated by the ET c.
Mapping and dividing the site into site-specific Suggested Websites
management units can result in substantial water
CIT (2009) Center for Irrigation Technology. Available at:
conservation and economic savings. In addition,
http://cit.cati.csufresno.edu/ (accessed 5 August
the knowledge and experience of a quality turf- 2009).
grass manager is necessary to make irrigation Hunter Irrigation Innovators (2009) Available at: http://www.
adjustments by observation. The use of site-spe- hunterindustries.com (accessed 7 October 2009).
cific irrigation management does not preclude the Hunter Irrigation Innovators (2009) Irrigation System Design:
monitoring of the site by a qualified manager or Student Workbook. Available at: http://www.hunter-
irrigation technician. industries.com/Resources/PDFs/Educational/
Too little water can seriously damage turf, but Domestic/ed_004_final.pdf (accessed 7 October 2009).
turfgrasses can stand longer periods of too little Hunter Irrigation Innovators (2009) Precipitation Rates and
water than they can stand periods of too much Sprinkler Irrigation: Instructor’s Manual. Available at:
http://www.hunterindustries.com/Resources/PDFs/
water. Standing water or saturated soil will usually
Educational/Domestic/precipitation_rates_and_
damage turf more quickly than drought. Although spr inkler_irr igation_wor kbook_instr uctor.pdf
proper irrigation is rarely ignored, proper drainage (accessed 7 October 2009).
is sometimes overlooked. Surface drainage is nearly Hunter Irrigation Innovators (2009) The Handbook of
always more important than subsurface drainage, Technical Irrigation Information: A Complete Refer-
but both can be used to properly manage water on ence Source for the Professional. Available at: http://
a turfgrass site. Regardless of which type of drain- www.hunterindustries.com/Resources/PDFs/Tech-
age is used – surface or subsurface drainage – the nical/Domestic/LIT194w.pdf (accessed 7 October
water must always have someplace to go. Natural 2009).
drainage patterns are best. It is much easier to fol- Irrigation Association (2009) Smart Practices. Sustain-
able Solutions. Available at: http://www.irrigation.org/
low the natural drainage than it is to create an
(accessed 7 October 2009).
artificial drainage pattern. Rain Bird (2009a) Manufacturer and provider of irrigation
products and services. Available at: http://www.
Suggested Reading rainbird.com (accessed 7 October 2009).
Rain Bird (2009b) Landscape Irrigation Design Manual.
Beard, J.B. (1989) Turfgrass water stress: drought resis- Available at: http://www.rainbird.com/pdf/turf/Irriga-
tance components, physiological mechanisms, and tionDesignManual.pdf (accessed 7 October 2009).
species-genotype diversity. Proceedings of the 6th Toro (2009a) Available at: http://www.toro.com (accessed
International Turfgrass Research Conference, Tokyo, 7 October 2009).
Japan, pp. 23–28. Toro (2009b) Do-It-Yourself: Sprinkler Planning and
Beard, J.B. (1993) The xeriscaping concept: what about Installation Guide. Available at: http://www.toro.com/
turfgrasses. International Turfgrass Society Research sprinklers/pig.pdf (accessed 7 October 2009).
Journal 7, 87–98. Toro (2009c) Sprinkler Replacement and System Trou-
McIntyre, K. and Jacobsen, B. (2000) Practical Drainage bleshooting: Toro Irrigation Design Service. Available
for Golf, Sportsturf, and Horticulture. Ann Arbor at:http://www.toro.com/sprinklers/repair.pdf (accessed
Press, Chelsea, Michigan. 7 October 2009).
Pira, E.S. (1997) A Guide to Golf Course Irrigation UC Davis (2009) Estimating Leaching Fraction Require-
System Design and Drainage. Ann Arbor Press, ments. Available at: http://ucce.ucdavis.edu/files/
Chelsea, Michigan. filelibrary/5049/773.pdf (accessed 11 June 2009).

128 Chapter 8
 Adjusting for Seasonal Conditions
9 and Temperature Stress

Key Terms
For purposes of this text, seasons are defined in the traditional sense. In the northern hemisphere, winter occurs
from 21 December to 20 March, spring from 21 March to 20 June, summer from 21 June to 20 September
and fall from 21 September to 20 December.
Lipid peroxidation occurs when free radicals take electrons from lipids, causing them to destabilize and degrade.
Because lipids are important components of cell membranes, lipid peroxidation can be very destructive to the
regulated passage of chemical species in and out of cells.
Enzymes and proteins denature, meaning that they unfold and become ineffective, at 104 °F (40 °C).
A heat shock protein is one of many proteins that may be synthesized in a plant to protect proteins and enzymes
from unfolding as a result of excessive heat. A heat shock protein is a form of chaperonin.
Chaperonins are a group of proteins that help to guide the correct folding of polypeptides (long chains of amino
acids) into proteins and enzymes without becoming part of the final structure.
Conduction is the transfer of energy from one molecule to another touching molecule. Warm grass leaves can
conduct heat to the air surrounding them.
Convection is a mass movement of energy, usually in liquids or gases. The bottom of a pool is always colder
than the surface because warm water rises by convection, and the ceiling in a room is always warmer
than the floor for the same reason. As warm grass leaves conduct heat to the surrounding air, the warmer
air rises.
Equilibrium is a balanced condition that occurs between two entities when they become equal in some respect.
When a grass blade is the same temperature as the air around it, the grass blade and the air have reached
equilibrium with respect to temperature.
Photorespiration occurs when Rubisco (ribulose bisphosphate carboxylase), the major enzyme in the
Calvin cycle of photosynthesis, binds oxygen instead of carbon dioxide. The process only occurs in
cool-season (C3) plants and is the main reason that C4 photosynthesis is more efficient than C3
photosynthesis when the weather is warm. It is called respiration because carbon dioxide is lost during
the process.
Colligative properties are the physical properties of solutions that are affected by the concentration of solutes
contained in them. Water containing a high concentration of solutes, such as the water in a plant cell, boils at
a higher temperature and freezes at a lower temperature than pure water.
A solute is a substance that is dissolved in water.
Intracellular water is the water inside the cell membrane. Intracellular water is higher in solutes than intercellular
water.
Intercellular water is the water contained in and around the cell walls of a plant.
Viscosity is the measure of the resistance of a liquid to flow. Liquids become more viscous as they cool and
approach their freezing temperature, and less viscous as they warm and approach their boiling temperature.
Photooxidation is the loss of an electron from a photoexcited chemical species, in this case, chlorophyll.
Chlorophyll photooxidation is most likely to occur on clear cold days when sunlight is intense but the
temperature is too cool to allow rapid chemical reactions.
Photoinhibition is a light-induced decrease in the activity of the photosynthetic process that can occur when
temperatures are either too hot or too cold for the pathway to operate efficiently.

©CAB International 2011. Turfgrass Physiology and Ecology (G. Bell) 129
9.1 Optimal Turfgrass Health is a Result 9.2 Seasonal Turfgrass Management
of Flexible Management Under Differing
We all know that C4 grasses do not grow particu-
Temperatures
larly well during periods of cool or cold weather
Air and soil temperatures are two of the few and that C3 grasses do not grow particularly well
environmental conditions that turfgrass managers during periods of warm or hot weather. We also
cannot realistically influence. As you probably know that turfgrasses are less green, sometimes
know, there are actually athletic fields and golf yellow or brown, during these stressful periods.
course greens that are artificially air-conditioned Consequently, we can estimate plant health by the
for temperature control during high-stress periods. plant’s level of greenness. As you have learned,
In these cases, perhaps we have carried plant man- grasses can sometimes be excessively green, an
agement a little too far, perhaps not; you can reach unhealthy situation, but usually, green is a good
your own conclusions. Regardless, it is very thing. In your experiences, you may have seen C4
unlikely that you will have the opportunity to grasses that are yellowish in early spring become
air-condition your turf, so we can assume for quite green in late spring and then less green in late
purposes of this chapter that you are going to have summer, followed by green again in early fall then
to manage turf during normal seasons and also yellow or brown over the winter. If you live in a
during extreme temperatures that are both normal temperate region, you have seen similar seasonal
and above or below normal for your climate. You changes in cool-season grasses. Although we knew
will need a seasonal plan for managing warm- that these seasonal effects occurred, until recently
season (C4) and/or cool-season (C3) grasses at your we did not have a means to accurately quantify
site and you will need to help prepare your plants them. However, the commercial availability of
for temperatures that are significantly above or spectral reflectance sensors has now made it rela-
below normal. tively easy to document the changes in turfgrass
A seasonal plan for managing your grass is one species influenced by season. Figure 9.1 demon-
that you develop through knowledge such as that strates the spectral reflectance response of two
contained in this text and revise according to warm- and two cool-season grasses averaged over
experience at your site. Each turfgrass species at least two growing seasons in Stillwater,
differs in its particular requirements within a set Oklahoma. Similar seasonal responses have been
of environmental variables. Consequently, each documented in scientific journals (Guertal and
turfgrass species, and sometimes cultivars within Shaw, 2004; Xiong et al., 2007). These grasses will
a species, require different management practices respond differently to management depending on
or different levels of management to grow their season.
best. Obviously, warm- and cool-season grasses
require different seasonal management, and indi-
Seasonal management of C3 and C4 grasses
vidual species within the C4 category or the C3
category also differ in their management needs. Warm-season (C4) grasses can be expected to
Heat stress is rarely of major importance in C4 respond to management inputs best when temper-
grasses and cold stress is rarely of importance in atures are relatively high. We recognize the opti-
C3 grasses, but they do occur. Prolonged extremely mum daytime temperatures for warm-season shoot
high temperatures can sometimes damage C4 growth to be from 80 °F to 95 °F (27 °C to 35 °C)
grasses and extremely low temperatures or ice (Beard, 1973). Consequently, temperatures
cover can damage C3 grasses. Nonetheless, the between 80 °F and 95 °F should be ideal for effi-
management of warm-season grasses is primarily cient use of fertilizer, water and other management
directed toward minimizing cold stress and the inputs for warm-season grasses. Warm-season
management of cool-season grasses is primarily grasses will use management inputs less efficiently
directed toward minimizing heat stress. when temperatures are lower than or greater than
Consequently, the major emphasis of this chapter the ideal range. The climatic temperatures (tem-
will be directed toward the seasonal management peratures averaged by day over the last 30 years)
of warm- and cool-season grasses, cold stress in should indicate when management inputs are best
warm-season grasses and heat stress in cool- utilized and when less management is better in
season grasses. your area. However, let us complicate this approach

130 Chapter 9
 Tall fescue Buffalograss
0.8 Creeping bentgrass Bermudagrass

0.7
NDVI

0.6

0.5
1 Jan 22 Feb 15 Apr 6 Jun 28 Jul 18 Sep 9 Nov 31 Dec
Day of year

Fig. 9.1. The seasonal spectral reflectance response trend calculated from data measured as the normalized
difference vegetation index (NDVI) of tall fescue (Festuca arundinacea), creeping bentgrass (Agrostis stolonifera),
buffalograss (Buchloe dactyloides) and bermudagrass (Cynodon dactylon) averaged over two growing seasons in
Stillwater, Oklahoma.

slightly and attempt to encourage root growth easy to adjust management based on temperature
with our inputs instead of shoot growth. The forecasts. If we are flexible enough to take
optimum temperatures for root growth in warm- advantage of unusually cool or warm periods
season grasses are between 75 °F and 85 °F (24 °C rather than trying to maintain our turf based only
and 29 °C). The optimum temperature range for on historical climate we can improve our turf’s
root growth in cool-season (C3) grasses is between ability to maintain reasonable health through
50 °F and 65 °F (10 °C and 18 °C) (Beard, 1973). stressful periods.
Soil usually heats and cools more slowly than air. Attempting to encourage growth in warm-season
Consequently, the daily maximum soil tempera- grasses in early spring and late fall or cool-season
ture is nearly always lower than the daily maxi- grasses during the summer when temperatures are
mum air temperature and the daily minimum soil not conducive for growth can be counterproduc-
temperature is almost always higher than the daily tive. During these periods, managing for increasing
minimum air temperature (Fig. 9.2). The daily density and greenness will probably harm your turf
average soil temperature tends to be slightly higher more than help it.
than the daily average air temperature during the
fall and winter but close to the same during the
9.3 High-temperature Stress in Grasses
spring and summer in most climates under most
soil conditions (Fig. 9.3). As you know, chemical reactions increase at an accel-
Management practices, fertilizer, water, cultiva- erating rate as temperature increases. That means
tion, etc. are most effective when your turf has that the metabolic processes in a plant and all of the
good-to-optimal growing conditions with which chemical reactions in its environment speed up with
to take greatest advantage of your inputs. Warm- increasing temperature. As temperature increases,
season grasses respond best to management dur- both photosynthesis and respiration increase.
ing late spring, early summer, late summer and However, when the leaf temperature reaches about
early fall. Cool-season grasses grow best in the 104 °F (40 °C) photosynthesis begins to decline
spring and fall and will continue to grow roots all (Berry and Björkman, 1980). Respiration also begins
winter when the soil is not frozen. It is reasonably to decline as temperatures approach approximately

Seasonal Conditions and Temperature Stress 131

 120 Daily maximum air temperature Daily minimum air temperature 35

Daily maximum soil temperature Daily minimum soil temperature

30
100
25

Temperature (°C)
Temperature (°F) 80
20

60 15

10
40
5
20
0

0 –5
1 Jan 22 Feb 15 Apr 6 Jun 28 Jul 18 Sep 9 Nov 31 Dec
Day of year

Fig. 9.2. The daily maximum air temperature, daily maximum soil temperature, daily minimum air temperature and
daily minimum soil temperature averaged over 13 years (1994–2006) in Stillwater, Oklahoma. Notice how the daily
maximum soil temperature is always lower than the daily maximum air temperature and how the daily minimum soil
temperature is always higher than the daily minimum air temperature.

Daily average air temperature

Daily average soil temperature
90 35

80 30
70
25

Temperature (°C)
Temperature (°F)

60
20
50
15
40
10
30
5
20

10 0

0 –5
1 Jan 22 Feb 15 Apr 6 Jun 28 Jul 18 Sep 9 Nov 31 Dec
Day of year

Fig. 9.3. The daily average air temperature (the average of a day’s maximum temperature and minimum temperature)
and daily average soil temperature in Stillwater, Oklahoma. Notice how the daily average soil temperature tends to be
higher than the daily average air temperature during the autumn and winter but about the same during the spring and
summer.

104 °F (40 °C) but the net balance favors respiration hydrate deficit, plants that have been healthy enough
(Huang and Gao, 2000). Consequently, photosyn- to have adequate stored carbohydrates before stress
thesis slows more rapidly than respiration, resulting are more likely to maintain quality during heat stress
in a net loss of carbohydrates. Because of this carbo- (Huang and Gao, 2000).

132 Chapter 9
 The decline in net photosynthesis during heat reason, grasses with high carbohydrate reserves
stress is believed to be related to a loss of integrity and sufficient light, water and carbon dioxide to
in cell membranes or in membranes contained in perform rapid photosynthesis are better prepared
chloroplasts and other organelles (DiPaola and to withstand high temperature stress than those
Beard, 1992). Cell membrane thermostability is an without (DiPaola and Beard, 1992).
important component of each plant’s ability to There is an old saying “starve a cold but feed a
tolerate heat (Marcum, 1998). As you will learn fever”. Whether or not “starve a cold but feed a
later, it is also an important component of cold fever” is good medical advice is unknown to this
tolerance in plants. Membranes are not only author. However, it is important to note that when
affected by the loosening of molecular bonds you have a cold and your system is not functioning
caused by heat, they may also be damaged by a well, you are not hungry, but when you have a
process called lipid peroxidation (Jiang and Huang, fever and you are burning a substantial amount of
2001). Lipid peroxidation occurs when free radi- calories, you do get hungry. Your fever is caused by
cals, usually oxygen, take electrons from a lipid, in the rapid respiration required for your body to
this case a membrane lipid, causing the lipid to manufacture the compounds needed to fight the
destabilize. Lipid peroxidation is more likely to pathogen that is causing your illness. Increasing
occur in polyunsaturated fatty acids than in satu- respiration causes high body temperature. When
rated fatty acids. For that reason, cool-season the temperature gets uncomfortable for us we drink
grasses, which have more unsaturated fatty acids in a lot of water and perspire to cool ourselves. Plants
their membranes than do warm-season grasses, are do the same thing but we don’t call it perspiration,
more readily affected by lipid peroxidation than we call it transpiration. In the last chapter, you
warm-season grasses. Polyunsaturated acids also learned that water is the best plant defense against
resist cold temperature degradation better than high temperature. However, if the temperature is
saturated acids, and this is one important reason high enough and lasts long enough, plants will
why cool-season grasses also resist cold tempera- eventually die, just as humans will eventually die
tures better than warm-season grasses. from a fever that is not controlled. Both humans
Free radicals are produced as an unintentional and plants use water as a defense against high-
by-product of many energetic biological reactions. temperature stress. The difference between the two
There are free-radical scavenging compounds gen- is that if water is not enough to relieve high-
erally referred to as antioxidants that quench or temperature stress in a human, that human can
stabilize free radicals, thus preventing much of the move to a cooler place, the plant cannot. If the
damage that could be caused by them. If you are temperature is too high, or the humidity is too high
familiar with human nutrition, you have probably and/or air movement is too low to allow rapid
heard the term antioxidant many times. High tem- transpiration of available water at the plant’s
perature depresses the synthesis of some of the current location, the plant will die.
more important antioxidants in grasses (Xu and
Huang, 2004). Consequently, lipid peroxidation
High-temperature stress
and the resulting degradation of membrane integ-
in warm-season grasses
rity is a major cause of high-temperature decline in
turfgrasses, especially cool-season grasses (Liu and Warm-season grasses are more capable of handling
Huang, 2000). There is some reason to believe that higher temperatures than cool-season grasses. The
applications of cytokinin(s), a plant hormone, may architecture of a warm-season grass, most notably
help to prevent lipid peroxidation in cool-season its extensive root system and efficient high-
grasses, but as yet this has not become a technique temperature photosynthetic pathway, allows it to
for practical management of grasses (Liu and thrive in temperatures that stress a cool-season
Huang, 2002a; Liu et al., 2002). grass. Warm-season grasses do not photorespire, so
A turfgrass plant’s rate of photosynthesis is also that particular source of heat, which occurs in cool-
limited by the amount of light, water or carbon season plants during high temperatures, is elimi-
dioxide available, but respiration has fewer restric- nated. Because they do not photorespire, warm-season
tions. At high temperatures, respiration continues plants use water more efficiently during high tem-
rapidly as long as sufficient carbohydrates, lipids peratures than cool-season plants. They are also
and proteins are available to respire. For that capable of obtaining more water than cool-season

Seasonal Conditions and Temperature Stress 133

grasses because they tend to have more extensive process, meaning that it is instigated or controlled
root systems. Finally, their high-temperature-efficient by plant response and that energy is required to
photosynthetic pathway enables more rapid replace- maintain it. Conduction and convection, however,
ment of the carbohydrates used during rapid respira- are passive processes, and are part of the natural
tion caused by high temperature. environment. We heat and cool our buildings by
Although warm-season plants are more capable conduction and convection. Air-conditioning, for
of cooling themselves and tolerating high tempera- instance, is an active process. We apply air condi-
ture than cool-season grasses, they are not immune tioning for the purpose of cooling a building and it
to it. When temperatures reach levels greater than works by a process of conduction and convection.
95 °F (35 °C), warm-season grasses begin to stress However, conduction and convection would occur
in a similar manner to cool-season grasses. At that whether the air-conditioner was present or not. We
point, temperatures are becoming dangerously simply use the air-conditioner to take advantage of
close to 104 °F (40 °C), the temperature at which conduction and convection processes for our own
enzymes and other proteins begin to denature comfort.
(meaning that they unfold and are no longer If you heat a cooking pan on a burner, then turn
useful). Both warm- and cool-season plants have the burner off, the pan eventually cools to the
physiological defenses that help prevent enzymes temperature of the air in the kitchen. It does that
and proteins from denaturing (DiPaola and Beard, by conduction and convection, two separate proc-
1992). These defenses are called heat shock proteins esses that combine to equilibrate the hot pan and
(DiMascio et al., 1994). the cooler kitchen air. We say that the pan cools to
the temperature of the kitchen, but that is not
entirely true. The pan and the kitchen air actually
Heat shock proteins
reach equilibrium (Box 9.1). The pan not only
Heat shock proteins are in a class of biological cools; it also warms the kitchen air. The effect on
compounds called chaperonins (Gething, 1997). In the air, however, is small compared with the cool-
a sense, the heat shock protein surrounds the target ing of the hot pan, so we tend to ignore it.
enzyme to prevent it from unfolding. Consequently, However, if we had several of these pans cooling,
the enzyme or protein maintains its original they would noticeably increase the temperature in
characteristics and continues to function in a useful the kitchen. All of those pans would be warming
manner. Genes that control heat shock protein the air around them.
synthesis are activated when plant temperature Conduction is the transfer of energy from one
approaches 104°F. The production of heat shock object or entity to another as the objects or enti-
proteins reduces the loss of enzymes to high- ties contact each other. Objects consisting of mol-
temperature denaturation. ecules at high relative energy transfer energy to
Heat shock proteins are most effective during the other objects or entities with molecules at lower
early stages of heat stress, so they perform best relative energy. In other words, hot objects trans-
when temperatures increase gradually rather than fer their heat to cold objects when the objects
abruptly (He and Huang, 2007). Slow temperature touch. Except for water, solids, liquids and gases
increases allow gene activation and adequate become less dense as they warm and are therefore
production of heat shock proteins as the plant’s lighter. Because the air around the pan becomes
temperature increases. Rapid temperature increases lighter as it warms, it rises and is replaced by the
overwhelm the plant’s heat shock response. When cooler air around it. As the pan warms the new air
temperatures rise rapidly, a plant cannot produce this also rises and is replaced. This process contin-
heat shock proteins quickly enough to overcome the ues until the pan is at the same temperature as the
denaturing of enzymes. In that case, fewer enzymes air around it. We call that process convection.
can be protected and more plant damage occurs. Once the pan is the same temperature as the air
around it, the conduction stops and convection no
longer occurs. The temperature of the pan is now
Conduction and convection
in equilibrium with the temperature of the air
Transpiration is not the only source of plant cool- around it. These conduction and convection
ing. Conduction and convection are also processes processes also occur in the case of warm grass
that help to cool a plant. Transpiration is an active leaves surrounded by cooler air. Conduction and

134 Chapter 9
 High-temperature stress
convection are not nearly as effective at cooling
in cool-season grasses
grass plants as is transpiration, especially when
air temperatures are high. However, the conduc- Although high-temperature stress can occur in
tion and convection processes provide some high- warm-season turf, it only occurs at extremely high
temperature relief, especially when limited water temperatures and it rarely kills grass. Cool-season
availability causes grass plants to close their grasses can be damaged by heat at much lower
stomates and thereby limit transpiration. temperatures than those that stress warm-season

Box. 9.1. A balance scale works by equilibrium.

Balance scales were one of the earliest measuring both sides of the scale. The weight on the scale will
devices and are still in use today. An object of change but it will still be in equilibrium. Equilibriums
unknown weight is placed in a tray on one side of the are maintained in living cells in the same way. When
scale and small weights are added to a tray on the too much of a compound is present on one side of
other side until the weights and the object counter- the cell membrane, the membrane allows a portion
balance. When the weight of both trays is equal, the of it to cross to the other side so that a new equilib-
balance has reached equilibrium. The amount of rium can be achieved.
weight required to counterbalance the object is Equilibrium in a living cell is considerably more
exactly what the object weighs. complicated than equilibrium in a simple balance
The equilibrium is considered stable because it scale. The inside and the outside of the cell may
does not change unless subjected to an outside harbor unequal amounts of a particular compound
influence. If you touch one side of the scale with when equilibrium is achieved because many other
your finger the equilibrium is disrupted. However, compounds also influence the equilibrium simultane-
once you remove your finger the scale will quickly ously. However, the complicated balance achieved by
return to equilibrium. The equilibrium can be main- the cell works in the same manner as the simple
tained if the same amount of weight is added to balance achieved by this scale.

Seasonal Conditions and Temperature Stress 135

grasses. Cool-season turfgrass shoots prefer of Kentucky bluegrass (Poa pratensis) at 95 °F
temperatures between 60 °F and 75 °F (16 °F and (35 °C) (Watschke et al., 1972). When photorespi-
24 °C) (Beard, 1973). Temperatures greater than ration occurs, heat is generated but carbon is lost
75 °F begin to cause stress in cool-season plants. rather than gained. Photorespiration does not
High air temperatures and high soil temperatures occur in warm-season plants, but it occurs
extending through both days and nights are most progressively more often in cool-season plants as
detrimental to turfgrass growth. However, it is the temperature increases.
high soil temperatures (Xu and Huang, 2000) and In high temperatures, photorespiration results in
high night temperatures (Xu et al., 2003) that cause a substantial decrease in carbohydrate production
the most damage. If nights remain cool and/or soil from that which would occur in a cool-season
temperatures can be reduced high temperature plant in lower temperatures where C3 photosynthe-
stress is minimized (Fu and Huang, 2003). High- sis is more efficient. It is this reduction in carbohy-
temperature stress can cause reduced plant density, drate production that often causes the loss of
tiller density, root number and root fresh weight in cool-season plants, either directly or indirectly,
cool-season grasses (Xu and Huang, 2001). It is when exposed to high temperature for extended
likely that shoot growth cessation is actually periods. Low carbohydrate production is the rea-
caused by a net loss of carbohydrates in the roots son that cool-season plant root systems decrease
(Aldous and Kaufmann, 1979). Increases in soil during warm summers. The reduction in nutrient
temperatures during the summer season cause uptake, and more importantly, in water uptake, by
declines in the production of new roots in cool- the weakening root system can cause plant death.
season grasses. By the end of the summer, the ratio The loss of energy to photorespiration may also
of dead roots to live roots increases to an annual weaken plants sufficiently that they become more
high (Huang and Liu, 2003). Root depth is also sensitive to insect or disease damage.
affected as it tends to become less with increasing It should come as no surprise that heat stress and
soil temperature (Martin and Wehner, 1987). At drought stress in cool-season grasses are related
the end of the summer, cool-season grasses tend to (Wang and Huang, 2004). Even warm-season
be in their poorest, most vulnerable condition of grasses under drought exhibit canopy temperatures
the year. much higher than air temperatures (Steinke et al.,
Much of the high temperature stress that occurs 2009). Drought stress predisposes cool-season
in cool-season grasses is directly related to their grasses to heat stress (DiPaola and Beard, 1992).
propensity for photorespiration when temperatures Drought-stressed cool-season grasses succumb to
are high. You learned about photorespiration in heat stress at lower temperatures than plants with
Chapter 3. Photorespiration occurs when Rubisco, adequate water. In contrast, cool-season grasses
the enzyme that binds carbon dioxide in the Calvin that have had the opportunity to acclimate to mild
cycle, binds oxygen instead. Photorespiration is not drought and higher temperatures, and have been
a useful process in terms of carbon acquisition. In managed properly in the spring and fall preceding
fact it is called respiration because it results in a the summer can withstand a hot summer in a tem-
loss of carbon. As you know, the occurrence of perate climate or a mild subtropical climate.
photorespiration increases as temperature rises
because oxygen is more soluble in water than is
Cool-season grass acclimation
carbon dioxide at high temperature, and also
to summer seasons
because, especially when temperature is high, light
is intense and days are long, photosynthesis is Annual cool-season grass species avoid summer
occurring so rapidly that the air around the heat by producing abundant seed then dying in late
turfgrass leaves becomes extremely high in oxygen spring or summer. When temperatures cool in the
and low in carbon dioxide. In experiments where fall, the seeds germinate and the species prospers.
oxygen availability was seriously restricted, the Some of these annuals, such as annual bluegrass
photosynthesis of cool-season grasses approached (Poa annua) make nice turfgrasses for certain loca-
that of bermudagrass (Cynodon dactylon) even tions and uses. However, in spite of the ubiquitous
when temperatures were high (Watschke et al., nature of annual bluegrass, its general adaptation
1973). In another experiment, low oxygen enabled to most regions as a weed species and its propensity
a nearly two-fold increase in the net photosynthesis to act as a perennial in some locations under

136 Chapter 9
certain management, it still tends to make a poor for long periods during the day (Bell et al., 2000).
turfgrass over the summer. We need perennial cool- Because rapid photosynthesis is not necessary, both
season turfgrasses that can withstand the rigors of cool-season grasses and warm-season grasses also
summer and maintain reasonable health during reduce leaf chlorophyll levels during summer, as
that season. We have several cool-season turf- evidenced by reductions in greenness (Xiong et al.,
grasses that can survive summer seasons, but they 2007). In addition, carotenoid pigment levels
perform best if they have an opportunity to accli- increase (Demmig-Adams, 1990). Carotenoids act
mate to or are forced to acclimate to summer stress as receptors of excess light energy that might oth-
before its actual arrival. erwise damage plants. These adjustments in leaf
pigment concentrations cause summer turfgrasses
to become less green and more yellow (Fig. 9.1).
Turfgrass response to longer day length
As turfgrass leaves grow more upright and less
As day length increases, cool-season grasses accli- chlorophyll is synthesized, light interception is
mate by growing more upright. Prostrate grasses, reduced and the leaves remain cooler. If tempera-
such as creeping bentgrass (Agrostis stolonifera) tures warm gradually as summer approaches, cool-
and bermudagrasses (Cynodon spp.), tend to toler- season grasses have sufficient time to acclimate to
ate exceptionally low mowing because they expose the upcoming stressful season. However, if the tem-
a large amount of leaf area to sunlight. Consequently, perature warms abruptly and remains high, as
low mowing does not affect a prostrate grass as sometimes happens, our grasses are unprepared
negatively as it does an upright growing grass. For and stress increases. We can help to avoid this situ-
that reason, cool-season grasses acclimate to long ation by subtly forcing our grasses to prepare for
days by growing more upright and reducing the summer. It is possible to encourage upright growth
amount of light interception by each leaf. Less light by mowing in multiple directions by rotation. Each
interception results in less heat in the leaf and successive mowing should be performed in a differ-
upright growth exposes more leaf surface area to ent direction than the last. For instance, mow north
the air, thereby enhancing both evaporation and and south for this event then east and west for the
conduction. When days are long, rapid photosyn- next one. Upright growth can be further encour-
thesis is not necessary. As days lengthen, light satu- aged on intensively managed turf, such as golf
ration occurs for longer and longer periods, course putting greens or bowling greens, by prac-
meaning that photosynthesis is at a maximum rate ticing shallow grooming or brushing (Fig. 9.4).

Fig. 9.4. A reel-type mower fitted with a grooved roller and a grooming attachment, devices that improve mowing
consistency, encourage plant density and help to encourage upright growth.

Seasonal Conditions and Temperature Stress 137

 Fertilizing cool-season grasses with nitrogen imme- Cool-season grasses under summer stress are
diately before or during summer is not recommended more susceptible to disease and insect damage than
except for light applications on intensively managed the same grasses during other times of the year.
turf such as golf courses or athletic fields where rapid Frequent monitoring of turf status is required dur-
recovery from damage is desired. Nitrogen fertiliza- ing the summer. Cool-season grasses can sustain
tion encourages chlorophyll synthesis. Encouraging damage quickly during this period and will heal
chlorophyll synthesis during a period when your very slowly. Insect or disease pressure must be
grasses are naturally reducing their chlorophyll con- monitored frequently and dealt with immediately.
tent is counterproductive. Withholding nitrogen fer- Damage can spread quickly on stressed turf. Most
tilization in late spring helps the grasses to acclimate insecticides and fungicides can be applied safely to
to the upcoming summer season. cool-season grasses when temperatures are high.
However, you must be careful with these applica-
tions. There are a few examples of these products,
Managing cool-season grasses
such as sterol inhibitor fungicides, that can cause
to survive summer heat
damage during hot weather. Some products include
Being proactive and attempting to help your cool- carriers like emulsifiers or oils that can be harmful
season grasses through a stressful summer often in hot weather. Read the label, and check with your
increases rather than decreases stress. It is the man- supplier before application. It is also important to
agement processes executed in the previous fall and stay within the range of recommended rates.
spring that get cool-season grasses through the sum- Although insecticides and fungicides are usually
mer with the least amount of stress. Water is a safe for summer applications, many herbicides can
plant’s best defense against summer stress and, as damage grasses during periods of high tempera-
you learned earlier, irrigation management is ture. Some herbicides that are routinely applied
paramount during the summer. Many other man- without incident during cooler times of the year
agement practices harm the turf rather than help it. can cause damage to desirable grasses during sum-
mer. Depending on the herbicide, damage may
occur on cool-season grasses, warm-season grasses
Management practices to avoid during
or both. Sometimes the damage is particular to
summer stress
certain species. Many herbicides that are safe on
It was mentioned earlier that except for light appli- one cool-season grass in the summer may not be
cations to certain types of turf, nitrogen fertiliza- safe on others. The same is often true of warm-
tion should be avoided on cool-season grasses season grasses. You should always be careful with
during the summer. Conversely, potassium fertiliza- chemicals, but summer is a time to be exceptionally
tion may help, especially if soil potassium is low. careful.
Practices that damage turf, no matter how slight, Low-mown turf is more susceptible to heat stress
should be avoided. An exception to that statement than high-mown turf. High-mown turf has more
is small-tine aerification or spiking when required leaf area above the soil surface where the turfgrass
to penetrate a thatch or soil layer and promote crowns reside. The greater leaf area of the high-
water infiltration and soil oxygenation. Turf on mown turf shades and insulates the soil better than
poorly aerated soil is at greater risk of heat-related does low-mown turf. A close-mown turfgrass can-
damage than turf on well-aerated soil (Huang opy can easily exceed the air temperature around it
et al., 1998). As increasing soil temperature causes during the heat of an afternoon on a clear summer
increasing respiration, oxygen is removed from the day (Throssell et al., 1987). Golf course fairways,
soil and replaced by carbon dioxide. When this tees and putting greens are especially vulnerable to
process occurs in poorly aerated soil, cool-season high temperature stress because they are closely
grasses may seriously decline (Rodriquez et al., mown. They also sustain traffic damage, which can
2005b). Thatch or layering problems are best be especially problematic during stress periods
addressed during the fall or spring, but sometimes when recovery is slow.
these problems are not noticed until symptoms of Efforts should be made during the summer to
stress appear during the summer. Turfgrass roots channel traffic to little used areas, allowing the high-
need an ample supply of water and oxygen even if traffic areas to recover. You should make an attempt
minor damage has to occur to get it to them. to route traffic so that the entire turfgrass area is

138 Chapter 9
used intermittently, thus giving each small region the and only occurs in warm-season grasses (Hale and
greatest possible time to recover before traffic is Orcutt, 1987).
applied there again. The alternative is to construct
mulch, gravel or concrete pathways that can be used
Freezing stress in grasses
to channel traffic away from stressed turf.
Sand topdressing is another practice that should Do you remember what colligative properties are?
be avoided during high temperature except in Colligative properties were introduced in Chapter 4
amounts small enough that the sand can be and are part of most introductory chemistry classes.
watered in or lightly brushed. Sand readily reflects There are four colligative properties of solutions.
light and heat, and when left in a pile for an hour They are that the higher the concentration of sol-
during lunch on a hot sunny day will almost utes in a liquid the higher the boiling point will be,
surely cause the turf around it to wilt and possibly the lower the freezing point will be, the lower the
die. Sand also has sharp edges. If topdressing is vapor pressure will be and the greater the osmotic
applied at high enough rates that dragging is pressure will be. Colligative properties are very
required to facilitate canopy infiltration, the sharp important to plants because they keep plant cells
edges of the sand will cut the turfgrass leaves as from freezing until air or soil temperatures get very
dragging occurs, creating damage that can only low; far below the freezing temperature of pure
heal slowly. Each time the surface receives equip- water. Plant cells contain many different solutes,
ment or foot traffic, the leaves will be pressed including sugars, amino acids, nutrient ions and
against the sharp sand, causing more damage. many other elements and compounds.
Topdressing with sand in the summer should be To understand why freezing and chilling stresses
executed with restraint. occur in plants, you have to remember some simple
A combination of surface fans and light misting plant anatomy. Plants have a cellular structure that
of water can effectively reduce canopy temperatures is slightly different from animal structure in that
on high-value turf such as golf course putting each cell has not only a cell membrane, but also a
greens. However, this practice also increases can- cell wall. The cell walls add support and act in a
opy humidity and should be closely monitored for manner similar to how a skeleton works in verte-
disease activity (Rodriquez et al., 2005a). brates like us. However, because plants don’t move
Although summer can be a stressful time for turf- they do not require joints in their support struc-
grasses, especially cool-season turfgrasses, successful ture, so their anatomy is basically a bunch of semi-
managers can minimize damage through prepara- rigid cells stacked on top of each other, with each
tion and responsible management. Water is the most cell wall filled with and surrounded by water. Small
important component of a plan for high-temperature imperfections or gaps between cells or between the
stress relief during summer. Proper management membrane and the cell wall are filled with water.
during the spring and fall growing seasons preceding Because water does not compress, its presence in
summer will minimize stress. Applying tactics to the small amounts around and in the cell walls pro-
management program that help the turf acclimate to vides rigidity, but its propensity to flow if space is
the coming summer can also help. available also encourages flexibility. The water
inside a plant cell is called intracellular water and
the water in the cell walls and surrounding them is
9.4 Cold-temperature Stress in Grasses
called intercellular water (Fig. 9.5). If the intracel-
Although cool-season grasses are generally adapted lular water freezes the cell nearly always dies.
to very cold temperatures, they are occasionally However, it is more likely that the intercellular
damaged by subfreezing weather and sometimes by water will freeze, damaging membranes and cell
ice cover. A cool-season grass that gradually accli- walls or, more often, causing cell dehydration.
mates to cold weather, however, is rarely damaged The intracellular, inside space contains the living
by it. Conversely, warm-season grasses are quite parts of the plant. All of the plant’s metabolic proc-
sensitive to cold. Temperatures do not have to esses occur in the inside space in water. Consequently,
reach freezing before some warm-season turfgrasses the inside water contains a high concentration of
begin to experience stress. This cold temperature solutes necessary to perform plant growth and
stress that may occur when temperatures are below maintenance. Activity is always occurring in the
about 50 °F (10 °C) is referred to as chilling stress inside space. However, activity, but not living

Seasonal Conditions and Temperature Stress 139

Fig. 9.5. The structure of plant cells. Living protoplasm is surrounded by a membrane which is encased in a cell wall
filled with and surrounded by water.

activity, is also occurring in the intercellular, out- in water, the higher its osmotic pressure becomes,
side space. The outside space provides water and meaning that when water of a high solute
nutrients to the inside space. The outside space is in concentration is separated by a water-permeable
contact with the xylem, the transpirational plant membrane from water with an even higher solute
veins that furnish the water and nutrients for plant concentration, water will move from the high con-
metabolism and the water for transpiration. The centration of solute to the higher concentration of
outside water also contains a high concentration of solute until the concentrations are equal. Solutes do
nutrients and other solutes, but its concentration is not dissolve in ice, so for the purposes of solute
not as high as that of the inside space. Consequently, concentrations, ice is not the same as liquid water.
the outside space freezes before the inside space. When the intercellular space begins to freeze the
Water is the only known compound that amount of water becomes less and less, but the
expands when it cools. All other compounds amount of solute remains the same, so the solute
shrink as they cool. Consequently, when intercel- concentration increases. When the solute concen-
lular water freezes, the expanding water can cause tration in the intercellular space becomes great
damage to cell walls. Ice crystals are sharp, so the enough, water begins to flow from the intracellular
freezing of intercellular water can also pierce cell space to the intercellular space by osmosis. This
membranes. However, it is dehydration that most flow dehydrates the intracellular space and if the
often causes a cell to fail following the freezing of dehydration becomes excessive can cause the cell to
intercellular water. die. If water does not flow from the cell to the
To understand the cell dehydration process you intercellular space, its solute concentration will not
must remember the colligative property called increase and it will probably freeze and die. For
osmosis. The higher the concentration of solutes that reason, the dehydration process is more likely

140 Chapter 9
to save the life of the cell than to kill it. The inter- (Zoysia spp.) inner membranes following chilling
cellular and intracellular spaces of a warm-season stress (Rogers et al., 1977).
grass are maintained at lower solute concentrations If the temperature increases, metabolism may
than those of cool-season grass and the warm- return to normal, but if the chilling temperature per-
season grass is therefore more likely to freeze. sists or becomes lower, cell membranes may become
disrupted badly enough that they begin to leak pro-
toplasm noticeably. Water-soaked lesions will form
Chilling stress in warm-season grasses
on turfgrass leaves, and the leaves and possibly the
Chilling stress is a condition that can occur in plants will probably die (DiPaola and Beard, 1992).
warm-season plants when temperatures fall below
50 °F (10 °C). It is defined as low-temperature stress
Chlorophyll photooxidation
in the absence of freezing (Levitt, 1980). Chilling
stress results from the loss of selective permeability Photooxidation is a destructive event (Niyogi,
of cell membranes. It can also occur because of 1999). As temperature declines, photosynthesis,
changes in metabolism caused by cool weather, or respiration and transpiration decline. In this case,
may occur from a combination of these. the decline in photosynthesis is most detrimental.
When temperature declines, photosynthesis slows.
The chemical reactions in both the z-scheme and the
Cell membrane permeability
Calvin cycle are affected. However, light energy
Cell membranes are primarily comprised of phos- interception is not affected. Chlorophyll molecules
pholipids. I am sure that you are aware of the continue to intercept light energy, but because the
effects of cold on lipids. Lard, for instance, is a liq- photosynthetic system is slow, this excitation energy
uid at high temperature but a solid at low tempera- has nowhere to go. Eventually, some chlorophyll
ture. Vegetable oil does not become a solid until its molecules reach an energy level high enough to pass
temperature gets extremely low, but its viscosity (its an electron directly to the PSII (photosystem II)
thickness or resistance to flow) becomes greater as receptor. This excess energy passing into the photo-
its temperature declines. Plants with cell membranes synthetic system causes damage that has to be
containing greater amounts of polyunsaturated repaired. The chlorophyll that passes the electron to
fatty acids are less likely to experience cold damage PSII has become oxidized, hence the name pho-
than those containing mostly saturated fatty acids tooxidation. The oxidized chlorophyll molecules
(Cyril et al., 2002). Warm-season turfgrass cell become free radicals seeking electrons from other
membranes contain lipids closer to lard than to compounds, and damaging membranes and other
vegetable oil and cool-season cell membranes con- metabolic processes. Chlorophyll photooxidation is
tain more lipids like vegetable oil. Consequently, most likely to occur on clear cold days when intense
cool-season cell membranes are not seriously light is present, but chemical processes are exces-
affected by cold temperature until the temperature sively slow because of cool temperatures.
gets extremely low, but warm-season membranes
can be seriously affected when the weather is cool.
Dehydration
In plants, especially warm-season plants, there is a
membrane transition from flexible to rigid as the Motor oil is rated by viscosity because rapidly
temperature is reduced. You probably remember that moving close-fitting metal mechanical systems like
cell membranes are like selective sieves. They allow gasoline engines require different grades of oil for
certain molecules to penetrate but not others. If the lubrication according to temperature. A gasoline
membrane becomes rigid, however, its selectivity is engine containing an oil of high viscosity and close-
affected and it begins to permit unwanted com- fitting parts may be damaged shortly after start-up
pounds to penetrate both into and out of the proto- if its oil is too viscous to squeeze between its
plasm. That, in turn, makes it very difficult for the close-fitting parts without creating excessive pres-
cell to maintain efficient metabolism, and the cell’s sure. Once the engine warms, the oil viscosity will
controls on metabolic processes may be compromised. be reduced and damage will not occur. However,
Researchers have found extensive damage to inner when the oil is cold and viscosity is high, damage is
chloroplast membranes of bermudagrass, but little likely. Therefore, gasoline engines require
damage to the more cold-tolerant zoysiagrass low-viscosity oils when temperatures are cold.

Seasonal Conditions and Temperature Stress 141

As temperature cools, oil viscosity increases. most of the soil is frozen. Consequently, the very
Temperature has the same effect on water. top layer of soil where the turfgrass crowns reside
As soil temperatures cool, soil water becomes becomes saturated. At night, when this water
more viscous. The more viscous the soil water freezes, it expands and crushes turfgrass crowns.
becomes, the more difficult it is for plant roots to Slushy, partially frozen conditions such as slowly
absorb it. Consequently cool temperatures slow melting ice or partially frozen wet snow can also
water uptake, water translocation and transpira- cause damage. During such conditions, ice crystals
tion. In temperate and cooler climates, soil water can be pushed into and pierce turfgrass crowns as
may be frozen for much of the winter. If the soil traffic or another pressure is applied overhead.
water freezes when our turfgrasses are still green Ice encasement basically eliminates air exchange
and active the plants may dehydrate. As photosyn- from the living parts of the plants to the atmos-
thesis, respiration and other metabolic processes phere. Although the turfgrass may be dormant or
occur, water is used. This water, however, cannot partially quiescent, its roots and crowns are still
be replaced when the soil water is frozen. The same performing respiration at reduced rates.
situation can occur when our plants need water but Consequently, the respiration process is removing
the soil water is too viscous to be absorbed rapidly oxygen from the air and replacing it with carbon
enough to satisfy plant needs. It is likely that much dioxide. Eventually the plants may die of anoxia, a
of the winter turfgrass damage attributed to freez- lack of oxygen. However, low oxygen and high
ing is actually caused by dehydration. carbon dioxide is a more severe condition than low
oxygen alone (Castonguay et al., 2009). If you
have areas of high-value turf covered in ice for
Turfgrass damage by ice encasement
more than 20–30 days, it would probably be worth
In many regions, ice encasement of grasses for your effort to remove the ice. It could make the
short periods during the winter is common. difference between live and dead turf next spring.
Creeping bentgrass has been known to survive
under ice cover for up to 90 days (Tompkins et al.,
Cold acclimation in turfgrass plants
2004). For the most part, cool-season grasses can
tolerate this ice encasement for short periods of 2 As temperatures cool in late fall, turfgrasses begin
or 3 weeks. However, longer periods or periods of to “harden off” for the winter, which means that
freezing and thawing of the ice can be very detri- they begin to prepare themselves to withstand tem-
mental to the turfgrass below it. Warm-season peratures that would normally be damaging.
grasses are more susceptible to ice damage than As winter approaches, both warm- and cool-
cool-season grasses. season turfgrasses begin to translocate carbohydrates
Ice can cause damage to grasses in three ways. It into their roots and stems (McKell et al., 1969; Dunn
can crush or pierce crowns and roots as it freezes, and Nelson, 1974). At least in some grasses, these
it can restrict air exchange for a long enough long-chain carbohydrates may be quickly converted
period that the air under the ice becomes of poor to sucrose, which accumulates in the turfgrass
enough quality to damage turf, or it can be pushed crowns and stems when necessary to prevent freezing
into crowns or roots by pressure from traffic or (Fry et al., 1993; Dionne et al., 2001a). As tempera-
other sources. tures cool, shoot growth slows, but photosynthesis
Periods of freezing nights and thawing days are continues as long as the plants are green. The carbo-
likely to cause more damage to turfgrass than con- hydrates produced by photosynthesis and stored in
stant freezing temperatures. When ice encasement belowground roots and stems for the winter also
first occurs, it usually does not crush or pierce provide fuel to allow them to continue slow metabo-
enough crowns to cause substantial damage. lism over the winter after their leaves die. Some
However, if the sun or warm temperature melts the grasses of course, depending on species and location,
ice daily then it freezes overnight repeatedly, dam- remain green all winter and continue to produce
age is likely to be extensive. As the ice melts, water carbohydrates throughout the period. Carbohydrates
gathers on the soil surface. Under ice cover, the stored but not used during the winter may provide
upper soil is usually frozen. Water from the melting for rapid new growth in the spring.
ice helps to melt the very top layer of soil and infil- During cold acclimation, turfgrasses also accu-
trates it slowly, but it cannot penetrate far because mulate proteins and other nitrogen-rich compounds

142 Chapter 9
that decrease in the spring as the plants de-acclimate A frozen wet soil is more damaging to turfgrass
(Dionne et al., 2001b). This protein accumulation crowns than a frozen dry soil. A wet soil expands
has been implicated in the greater freeze tolerance more when it freezes than a dry soil, and is more
of some bermudagrasses (Gatschet et al., 1996) and likely to crush crowns. However, wet soils are less
some zoysiagrasses (Patton et al., 2007a,b). So it likely to freeze and resist rapid temperature declines
seems that warm-season grasses that rapidly accu- better than dry ones. Consequently, light fall and
mulate carbohydrates and proteins during cold winter irrigation can help to prevent freeze damage
acclimation are likely to be more cold tolerant to turfgrass. The moist soil resists the rapid tem-
(Zhang et al., 2006). perature declines that are most detrimental to turf-
During acclimation, grasses also go through a grass plants but does not contain enough water to
dehydration process. Less water is stored in tissues crush crowns as it freezes. Soil moisture is especially
as winter approaches. This dehydration is perhaps important during early cold snaps that occur before
the most important process for plant protection the turf has had sufficient time to acclimate to cold
against freezing. Cool-season grasses dehydrate temperatures. However, it is important not to apply
more completely than warm-season grasses and soaking irrigation. The dryness of the crown is an
this is believed to be a major reason that cool- important component of resistance to freezing. We
season grasses are more resistant to freezing. Dry want the crown to remain relatively dry.
tissue is less likely to freeze than succulent tissue. Proper fall fertilization can also help turfgrasses
Another reason that cool-season grasses are better to resist chilling and freezing. It is widely believed
at resisting freeze damage than warm-season grasses that warm-season plants should not be fertilized
is because fructans, the storage carbohydrate that with nitrogen for a month or more before freezing
you know to be used by cool-season grasses, is temperatures are likely to occur. The succulence
soluble, and the starch stored by warm-season normally encouraged by nitrogen fertilization could
grasses is not. Consequently, carbohydrate storage counteract the dehydration of the plant tissue as
in cool-season grasses has colligative properties, the grasses harden off for winter. This no- or low-
but in warm-season grasses it does not. That also nitrogen protocol is not supported by scientific
helps cool-season grasses to survive cold tempera- research (Reeves et al., 1970; Richardson, 2002).
tures that kill warm-season grasses. However, it is based in logic and seems to make
In the spring, grasses de-acclimate quickly with sense, so most managers are reluctant to use nitro-
warming soil temperatures (Davis and Gilbert, gen fertilizer on warm-season grasses as winter
1970). Sudden cold temperatures following approaches. Cool-season grasses are not detrimen-
de-acclimation can be stressful. De-acclimated tally affected by late-season nitrogen fertilization.
plants, however, are able to partially re-acclimate In fact, as stated in earlier chapters, late fall is an
when soil temperatures drop, thus helping to pre- excellent time to promote root growth with nitro-
vent severe damage (Tompkins et al., 2000). gen fertilization of cool-season grasses without
encouraging shoot growth.
When cell membranes begin to increase in rigid-
Managing cold-temperature stress
ity with the chilling of warm-season grasses, one of
When we consider temperature, we normally think the first membrane-regulated processes affected is
of air temperature. Although cold air affects our potassium exchange. Some of the potassium neces-
turf, it is soil temperature that is most important. sary for cell functions is lost to intercellular spaces
Dead turfgrass leaves in the winter are common to when the cell membranes are disrupted. Potassium
warm-season grasses north of tropical regions and helps to increase turfgrass resistance to many
even to cool-season grasses in regions where win- stresses, including cold. Replacing late-season
ters are exceptionally cold. However, it is the nitrogen applications to warm-season grasses with
crowns (the thick, whitish part of the turfgrass that late-season potassium applications helps warm-
grows at soil level where grass shoots and roots season grasses to resist chilling stress and freezing
meet) that determine the life or death, and even the stress (Juska and Murray, 1974).
current health of a turfgrass plant, and the crowns Research suggests that high phosphorus levels may
are in the soil. Soil temperatures are ultimately interfere with the cold-protection qualities enhanced
more important than air temperatures for the cold by potassium (Reeves et al., 1970). In addition,
survival of turfgrass plants. potassium may not be utilized fully unless sufficient

Seasonal Conditions and Temperature Stress 143

nitrogen is also present (Reeves and McBee, 1972). should be familiar with. It is not unique to warm- or
Consequently, light applications of nitrogen should cool-season grasses, and is usually the reason that
accompany late-season potassium applications for warm-season grasses yellow when the temperature
best results. Nitrogen fertilization in the fall has the is excessively high in the summer and that cool-
added advantage of encouraging warm-season grasses season grasses yellow as the temperature declines in
to stay green longer in the fall and green up earlier in the winter. As in photooxidation, photoinhibition
the spring, thereby extending the growing season for occurs when light energy is absorbed but cannot
about 20–25 days (Reeves et al., 1970; Richardson, normally proceed down the photosynthetic path-
2002). Greenhouse research found that a fertilizer way. When the temperature is high, photoinhibition
ratio of 4:1:6 nitrogen:phosphorus:potassium encour- occurs because of the disruption to protein struc-
aged the best cold tolerance in two bermudagrasses tures that can occur at temperatures greater than
(Gilbert and Davis, 1971). Depending on soil tests, 104 °F (40 °C). Cold-temperature inhibition occurs
the phosphorus in this formulation may be avoided to when the PSII (photosystem II) reaction center, spe-
reduce potential nutrient runoff that could enhance cifically the D1 protein, is damaged by excess
surface water eutrophication. energy. The damaged proteins are constantly
Iron applications can also be beneficial to warm- repaired and replaced to allow some photosynthesis
season grasses approaching winter. Iron applica- to proceed even during normal metabolism
tion encourages chlorophyll synthesis which, in (Tyystjärvi, 2008). Consequently, photoinhibition
turn, encourages carbohydrate production and usually does not permanently damage photosyn-
storage. In addition, the iron promotes aesthetic thetic pathways and they return to normal when
color without causing an increase in succulence light becomes less intense or temperatures return to
(Goatley et al., 2005). levels more conducive for chemical activity.
Some ice damage and dehydration may be
avoided by encouraging grasses to grow deeper
9.5 Chapter Summary
roots. In summer, the soil gets cooler as you dig
deeper because the finer minerals that accumulate The best management practice available for main-
deeper hold more water and because the deeper soil taining turfgrass under the extreme hot and cold
is protected from the sun’s radiation. In the winter, temperatures common to your area is choosing the
the deeper soil stays warmer again because it holds best species and cultivar for your site(s). Warm-
more water and because air temperature has less season species and cultivars differ in their tolerance
effect on it. The upper soil may warm from the sun of cold conditions, and cool-season species and cul-
but freeze again at night. Deep soil, however, resists tivars differ in their tolerance of heat (Sifers and
freezing. Builders and construction personnel are Beard, 1993; Dunn et al., 1999; Anderson et al.,
always aware of the “frost line” in their area. The 2003). In some cases, you never have the opportu-
frost line is the depth of soil below which the nity to establish or reestablish the turfgrass that you
ground never freezes and, although it has nothing are working with, but in many cases you do. Be sure
to do with turfgrass, is a good example of how the to choose wisely. In addition to extension services,
soil reacts to freezing temperatures. The frost line local agricultural universities, local growers and
changes with climate. In warm regions it is quite local practitioners, selection information is available
shallow, but in cold regions it can be quite deep. at the website of the National Turfgrass Evaluation
Although the frost line is too deep for turfgrass Program (NTEP). If you live in the USA, you can
roots in many areas, a turfgrass plant with a deep find locations near you that have completed turf-
root system is more likely to find water in the win- grass quality trials for the NTEP. If not, you can
ter than one with a shallow root system. compare your climatic conditions with those in the
USA to find the locations that are similar to yours.
There is little that we can do about changes in
Photoinhibition
temperature. Temperature is one of the very few
Photoinhibition, the disruption of the photosynthetic things that affects our turf that we have very little
process can detrimentally affect photosynthesis control over. Consequently, a seasonal management
when temperatures are either too hot or too cold plan needs to be formulated to take advantage of
(Long et al., 1994). Photoinhibition is not some- the best growing conditions in your region to pre-
thing that we can control, but it is a term that you pare your turf for the worst conditions. In addition,

144 Chapter 9
seasons differ from year to year and we have to be Suggested Reading
prepared to adjust our management practices so as
DiPaola, J.M. and Beard, J.B. (1992) Physiological effects
to best affect our turf under unusual temperature of temperature stress. In: Waddington, D.V., Carrow,
conditions as well as under those that are relatively R.N. and Shearman, R.C. (eds) Turfgrass. ASA-CSSA-
normal for our region. A flexible management SSSA, Madison, Wisconsin, pp. 231–268.
regime is necessary for best management.
Knowledge of turfgrass responses to weather and
the overall environment is required to make man- Suggested Websites
agement decisions. Because we cannot seriously NTEP (National Turfgrass Evaluation Program) (2009)
affect the temperature at our site, we have to be Available at: http://www.ntep.org (accessed 17
able to react to it in the most positive manner. December 2009).

Seasonal Conditions and Temperature Stress 145

 Growing Grass on Soil, Sand and Salt
10

Key Terms
Sand, for purposes of this chapter, refers to root-zone mixtures of sand with or without amendments specifically
designed as root-zone material for turfgrass systems.
Native soil, or simply soil, for the purposes of this chapter, refers to disturbed or undisturbed mixtures of sand,
silt, clay and organic material commonly found in natural settings.
Black layer is an anaerobic soil layer where sulfur-reducing bacteria have turned the soil black. Black layer occurs
when a soil pocket or layer remains saturated with water most of the time.
Localized dry spot is a condition where the soil has become hydrophobic and restricts water infiltration.
Nutrient leaching occurs when a nutrient, usually nitrogen, moves downward through the soil past the root zone
with percolating water before plants can take it up.
Spoon feeding is a series of frequent spray applications containing light rates of nutrients, perhaps in the range
of 10 pounds of nitrogen per acre (11 kg ha−1) or less.
A foliar application is a spray application of solution at less than 20 gallons per acre (190 l ha−1) in very fine
droplets meant to attach to the turfgrass leaves for absorption rather than to the soil or thatch.
A typical spray application of nutrients is commonly applied at 20 to 40 gallons per acre (190–380 l ha−1) and
contains 20 to 40 pounds of nitrogen per acre (22–44 kg ha−1) and possibly other nutrients.
Saline in the traditional sense refers to a solution high in salts. In this text, it refers to a solution high in salts other
than sodium.
Sodic refers to a solution, including a soil water solution, containing a high proportion of sodium ions.
Saline+sodic in this text refers to a solution, including a soil solution, high in both sodium and other salts.
Electrical conductivity (EC) is defined as the reciprocal of electrical resistance and refers to a material’s ability to
carry electricity. It is measured in Siemens. The EC of soil solution is measured on a saturated soil extract
and is designated ECe. The EC of water is designated ECw.
ESP, the exchangeable sodium percentage, is the proportion of exchange sites on a soil’s cation exchange capac-
ity (CEC) occupied by sodium reported as a percentage.
SAR stands for sodium adsorption ratio and is a measure of sodium hazard that is calculated by comparing the
amount of sodium in solution with the amount of calcium plus magnesium.
A leaching fraction is the proportion of irrigation and rainfall that carries salts with it through the root zone.

vehicle traffic would soon compact and contain too

10.1 Different Media Require
much water and too little air. For that reason,
Different Management
many turfgrass sites, especially those designed spe-
In Chapter 7, a review of basic soil properties was cifically to provide characteristics conducive to
provided because of its importance for nutrient playing sports, are built on sand. Properly designed
retention. In Chapter 8, soil was discussed for its and constructed sand surfaces are highly resistant
importance in water retention. The perfect soil for to compaction, but they require different manage-
plant growth would contain 45% mineral solids, ment from most natural soils.
5% organic solids, 25% water and 25% air. Many native soils have high sand contents, but
However, that particular soil on a turfgrass site few have a sand content as high as those designed
that was required to sustain high pedestrian or for turfgrass systems and probably none have sands

146 ©CAB International 2011. Turfgrass Physiology and Ecology (G. Bell)
of such uniform particle distribution as those slicing the soil, drilling it or disrupting it with high-
selected for turfgrass. Turfgrass sand systems are pressure water or air can also be effective (Wiecko
designed to resist compaction, but they are not a et al, 1993; Praemassing et al., 2009). Natural
particularly good media for growing grass. It takes freezing and thawing is an excellent aerifier but one
a knowledgeable individual to grow turfgrass on that we have little control over. Natural aerification
sand, especially on a sand surface that is under and mechanical aerification offer the same benefits
constant stress. However, knowledgeable turfgrass and usually have the same drawbacks.
managers tend to prefer working with sand systems Aerification is one of our most beneficial turf-
because the manager can control nearly all of the grass management practices (Jaabak, 1993). High-
inputs. use areas such as golf course tees, greens and
fairways, sports fields, grass tennis courts and
bowling greens need to be aerified regularly.
10.2 Problems With Soil
However, we hesitate to aerify as often as we
As we discussed in Chapter 7, natural soil contains should because the practice is so disruptive to play-
both large and small pores called macropores and ing surfaces. Hollow-tine aerification, the process
micropores. Natural soils develop structure that of punching holes and removing cores, is the most
consists of aggregates. The micropores inside the effective aerification process that we practice
aggregates hold water and the macropores that (Baker, 1994). However, it temporarily results in a
form between the aggregates hold air. This situa- very poor playing surface, especially for a golf
tion is perfect for plant growth but is easily dis- course or bowling green. For that reason, the play-
rupted if the soil is compressed by traffic. Traffic ers who use the turf surface dislike aerification and
compression crushes the aggregates and forces only allow it because they have been educated to
them closer together. In a simple sense, traffic com- realize its necessity. As managers, we try to balance
pression of the soil turns the macropores into the agronomic needs of the turf with the needs of
micropores. Consequently, a soil that was once our customers. Sometimes the needs of our turf
25% air and 25% water at field capacity is now conflict with those of our customers so we have to
something like 10% air and 40% water or worse. be creative to satisfy our goals with as little disrup-
Following traffic compression, there is still plenty tion as possible. After all, if it wasn’t for the players
of water but not enough air to properly sustain or the customers, we would not have a job.
root respiration. We call traffic compression, or Hollow-tine aerification needs to be practiced on
any other type of soil compression, compaction most playing surfaces at least twice per year and
(see Chapter 5) and we deal with it primarily more often if possible (McCarty, 2001). Agronomically
through aerification (aeration), a process that was speaking, the best time to aerify is when the turfgrass
also introduced in Chapter 5. The term “aerifica- roots are growing the fastest and need the most oxy-
tion” is preferred in this text so as not to confuse gen. The fastest root growth occurs in the spring, so
the cultivation practice with the aeration of water from a plant management standpoint, spring is the
and other media. best time to aerify. From a customer standpoint,
however, that may be a poor choice. Winter is often
the best time to aerify home lawns, commercial
Improving soil aeration
grounds or parks because few people use them at
Plants, including turfgrasses, are highly adaptable that time of year. If the site is not being used, the soil
and able to alter their physiology to improve their cores can be left on the surface and allowed to rein-
performance in compacted soils (Agnew and corporate naturally. By spring, the cores should be
Carrow, 1985). However, the constant traffic and gone but the holes will not necessarily be filled.
rate of compaction on many turfgrass sites is too Macropores will still exist to promote oxygenation
much stress for the plant to adapt to and the turf and root growth during the spring.
deteriorates (O’Neil and Carrow, 1983). Aerification Aerification of sporting sites should be accom-
is a mechanical soil disruption process that is plished when the turf can heal most rapidly
designed to relieve compaction, improve water (Puhalla et al., 1999). For cool-season grasses that
infiltration and promote oxygen retention on high- is usually in mid-spring and mid-fall. For warm-
traffic soils (Aldous et al., 2001). It is usually season grasses the best times to heal occur in
accomplished by punching holes in the soil, but mid-to-late spring and mid-to-late summer

Growing Grass on Soil, Sand and Salt 147

depending on normal temperatures and conditions. Core aerification appears to be a very disruptive
Core aerification timing has to be adjusted using process for a playing surface (Fig. 10.1). However,
common sense and climate. The needs of your close observation reveals that the process only
grasses and those of your customers will differ by affects a small portion of the playing surface
location and use (Baker et al., 1999a). directly (Box 10.1). For instance, core aerification
using 3⁄8-inch (9.5-mm) diameter tines on 1 by 2
inch (2.5 × 5.1 cm) centers only affects about 5.5%
Aggressive aerification
of the playing surface. However, if the surface is
Although compaction is a situation that normally aerified in three different directions about 16% of
occurs on native soil, core aerification should also the surface is affected. That calculation is made
be practiced on sand, although turfgrass managers with the assumption that no two holes touch when,
who work on sand media need not be as aggressive in fact, this is highly unlikely. Nevertheless, the
with their aerification practices as those who work result is a very close estimate. Consequently, an
on soil (Praemassing et al., 2009). Some turfgrass aggressive aerification event such as the one just
managers who have to grow grass on native soil for described can be almost as effective as aerifying
sporting operations, including golf, practice aggres- three times. A general recommendation for core
sive aerification. By aggressive I mean that they aerification would be to remove about 20% of the
aerify in multiple directions for a single event, pull surface area each year (McCarty and Miller, 2002),
relatively large cores, and/or set the spacing between although the actual amount removed depends on
cores as small as possible. In order to provide their the conditions and use of the site, and should
customers with the best playing surfaces possible, always be subject to adjustment depending on
they aerify less often but they aerify aggressively recent events and conditions.
when they do. Although the turf requires more
time to heal following aggressive aerification, it
Aerification processes
disrupts the playing surface for a shorter period of
time than would multiple aerification events during Core aerification of turfgrass areas is usually
the season. Aggressive aerification is not quite as practiced at depths of approximately 3 inches
effective a management tool as multiple aerifica- (7.6 cm). If you practice a 3-inch aerification two
tion events, but it is nearly as good. to four times per year, it will not be long before a

Fig. 10.1. The result of an aerification event on a research putting green using 3⁄8-inch (9.5-mm) tines on 2 by 2 inch
(5.1 × 5.1 cm) centers. The cores have been removed from the area in the foreground. This operation only removes
about 3% of the putting surface but it looks as if it is more disruptive than that.

148 Chapter 10
 Box 10.1. A formula for calculating the surface disruption of an aerification event.
Calculating the surface area removed by a core (length of area/length of tine spacing) × (width of
aerification event is fairly simple when you have a area/width of tine spacing)
background in basic mathematics. You have to
know the diameter of the hole left following core Consequently, if you use 3⁄8-in. (0.375-in.) tines at 2 ×
removal so that you can calculate its area. You also 2 in. spacing:
need to know the spacing of the tines so that you Hole area = 3.14 × (0.375/2)2 = 0.110 sq. in.
can calculate the number of holes that will be
pierced in a given area. The overall formula is The number of holes per sq. ft. = (12 in./2 in.) ×
this: (12 in./2 in.) = 36
So the surface disruption is:
Surface area affected in percent =
Hole area in inches × no. of holes per sq. ft./144 sq. 0.110 sq. in. × 36 holes per sq. ft./144 sq. in. per sq.
in. per sq. ft. × 100 ft. × 100 = 2.75%
or If you use 10 mm tines at 50 × 50 mm spacing:

Hole area in mm × no. of holes per m2/1,000,000 mm2 Hole area = 3.14 × (10/2)2 = 78.5 mm2
per m2 × 100
The number of holes per m2 = (1000 mm/50 mm) ×
The area of the holes is calculated from their radius (1000 mm/50 mm) = 400
(r ) by the formula for the area of a circle:
So the surface disruption is:
πr 2 = 3.14 × (hole diameter/2)2
78.5 mm2 × 400 holes per m2/1,000,000 mm2 per
The number of holes per unit area is calculated by: m2 × 100 = 3.14%

compaction layer builds up at the base of the tine tines that slice the turf as they are pulled across it
penetration (Fig. 10.2). Every 3 to 5 years that (Fig. 10.3). Slicing tines may also be mounted on
compaction layer will have to be disrupted. the typical small aerifiers normally used for golf
Deep-tine aerification is the process used to break course greens. The slits left in the green barely
up shallow compaction layers and improve deep affect the putting surface.
water penetration and soil aerification. It is very
effective (Morgan et al., 1965). Deep-tine
aerification is performed using deep coring tines
or solid spikes, verti-drain units whose tines
penetrate on an angle to cause additional soil
disruption, or deep drills that can be used alone
or with drill-and-fill techniques where an amend-
ment, perhaps sand, is injected into the holes as
they are drilled. Regardless of the technique used,
deep-tine aerification is usually more disruptive
than 3-inch coring. Deep-tine procedures lift and Compaction layer

displace more soil, make deeper holes that are

harder to fill, require more cleanup and take
longer to heal. Soil
Core aerification and deep-tine operations are
disruptive, but there are additional procedures that
are less severe. Slicing is a process that uses thin, Fig. 10.2. Repeated aerification to a consistent depth
triangular-shaped tines to penetrate the soil, leav- is likely to cause a layer of compaction to occur at
ing long but very thin slits. These units rarely lift the base of the tine penetration over time. Deep-tine
soil and result in minor disruption. Slicing units are aerification should be practiced occasionally to break
usually pull-behind units with a series of disk-type up that layer.

Growing Grass on Soil, Sand and Salt 149

Fig. 10.3. Slicing units are usually pulled behind a tractor or utility vehicle. The blades on this unit are used for slicing
golf course fairways and athletic fields. Much finer, less disruptive slicing blades are also available for this unit, as well
as shatter core blades, spikers and core aerifiers. Fine slicing blades cause very little disruption of playing surfaces,
are fast to use and require almost no cleanup. Although slicing provides very positive aeration, the effects are
generally short-lived compared with core aerification.

Water injection is also a method that creates little turfgrass systems, such as golf course putting
disruption to a playing surface (Karcher and Rieke, greens, bowling greens, tennis courts and athletic
2005). Extremely small high-pressure nozzles are fields are built on sand. Consequently, you may
used to inject small amounts of water through the spend considerable time managing relatively small
turf surface, fracturing the soil as they pass over it areas of turfgrass to perform up to extreme
(Toro, 2010). Once the unit passes, you can hardly expectations on sand. It is important to note that
tell that it has been there. Air injection units are managing turf on sand is quite different from
also available but, at this time of writing, little is managing turf on most soils. Those differences
known about their use and effectiveness. will be discussed later in the chapter. For now, let
It is tempting to develop an aeration program me assure you that sand is not a perfect system.
that does not include the disruptive process of core There are problems that occur on sand that
aerification. Slicing or water injection can be seldom occur on soil.
accomplished with little or no disruption to playing
surfaces. In addition, they require less labor and
Black layer
almost no cleanup. Slicing is considerably faster as
well. However, a quality aerification program has Black layer is not a problem unique to sand, but it
to include punching holes and removing cores. is rarely found on soil. Black layer appears as a
Although slicing and water injection can have posi- fairly thin layer of black sand usually close to the
tive aeration effects, those effects are usually short- surface of a sand root zone (Fig. 10.4). In extreme
lived, and these techniques are best used as conditions, such as when soil-bound sod is laid on
supplemental positive practices to a well-designed top of a sand root zone (see Chapter 8), the turf can
core aerification program. be pealed back to expose the black layer below.
The black layer will smell like rotting eggs, the
smell that occurs when sulfur is reduced by bacte-
10.3 Problems With Sand
ria. Black layer smells because it is caused by,
In terms of adequate root respiration, sand is encouraged by or occupied by (and we don’t really
superior to most native soils. Sand does not com- know for sure) sulfur-reducing bacteria such as
pact to a point that deters root respiration in a Desulfovibrio desulfuricans (Hodges, 1992a).
turfgrass system unless the sand is extremely fine Black layers are anaerobic, meaning that they con-
or it contains fine materials such as silt, clay or tain no oxygen. Consequently, they are believed to
organic material. Most turfgrass is managed on be associated with pockets of water-attracting clay,
soil. However, some of the most highly managed silt or organic material. These fine materials will

150 Chapter 10
Fig. 10.4. A black layer in a sand root zone. This area smells like rotting eggs.

hold water and will not release it to the sand until lower soil pH can help to reduce the potential for
they are completely saturated. Consequently, a black layer to occur provided that it is not already
layer of fine material in sand is filled with water present (Adams and Smith, 1993).
most of the time. In most cases, black layer is not a particularly
Although black layer can be severe enough that difficult situation to handle. Aggressive core aerifi-
turfgrass roots will not penetrate it, this is not cation with cores removed followed by topdressing
always the case. Turfgrass roots are discouraged by with clean sand that closely matches the original
black layer, but they can sometimes grow through root zone will usually improve the situation. Deep-
it, resulting in turfgrass decline rather than death tine aerification is advisable to make sure that the
(Cullimore et al., 1990). It is believed that to some black layer is broken and that it is infiltrated by
extent plant roots can oxygenate the soil as they columns of clean sand. The more often the aerifica-
pass through it. Consequently, at least in some tion is practiced and the more aggressively that it is
cases, turfgrass plants can break through, or even performed, the sooner the layer disappears.
break up, a black layer. Nevertheless, black layer
always causes temporary-to-terminal decline and is
Localized dry spots
always a serious problem. In some cases, black
layer or the circumstances that cause it can result in Localized dry spots (LDS) are another detrimental
a weakening of the turf to the point that it becomes condition usually only experienced in sand systems.
fatally sensitive to other stresses (Hodges, 1992b). Although water-repellent native soils are known to
Black layer is often the result of poor sand con- exist, water repellency is usually only a serious
struction or poor maintenance practices. Improperly problem for turfgrasses grown on sand (York and
mixed organic amendments added to sand systems Baldwin, 1992). As sand became popular as a
during construction can lead to pockets of poten- growing medium for golf course putting greens, it
tially dangerous anaerobic conditions. Black layer became apparent that certain spots in some greens
can also be caused by improper topdressing with repelled water. It was soon discovered that the sand
highly organic materials, soil or soil-containing particles in these spots were coated with a water-
sand. Sulfur-containing fertilizers or other materi- repellent material that was believed to be fulvic
als added to a system with an existing black layer acid (Wilkinson and Miller, 1978). Further research
can make the black layer worse (Berndt and and observation tended to confirm that some
Vargas, 1992). However, because sulfur-reducing organic material, possibly fulvic acid, was coating
bacteria are not particularly tolerant of low pH the sand in LDS causing them to repel water
soil, additions of sulfur or sulfate materials that (Tucker et al., 1990).

Growing Grass on Soil, Sand and Salt 151

 Sand is a drought-prone root-zone medium so it depended on the number of consecutive applica-
is not unusual for grass grown on sand to exhibit a tions and the air temperature. Little or no discol-
drought response when conditions are dry. High oration was observed when the air temperature
spots on the site, south-facing slopes, poor irriga- was less than 75 °F (24 °C), but injury was severe
tion patterns or areas of heavy thatch can look like when the air temperature exceeded 95 °F (35 °C).
LDS when irrigation is needed. However, if suffi- As a turfgrass manager, you will be faced with
cient irrigation is applied, these areas do not making important decisions about which manage-
decline. However, LDS are noticeable even under ment practices you apply. If you intend to use a
adequate irrigation. They may be identified as pud- potentially dangerous, but potentially effective,
dles during rainfall or irrigation, or as dry turf on tool make sure that you test it thoroughly on turf
an otherwise dew-covered surface. When water is with little value, such as a nursery or practice
applied to LDS, it does not penetrate the soil, it green, before you apply it to an area of high
simply runs away. Consequently, turfgrass root importance.
systems are stunted, the soil holds little water and There has been some indication that topdressing
the turf declines rapidly when rainfall is less than with porous ceramic clay can be effective for
substantial. reducing or eliminating LDS but it has not become
The problem of LDS is more widespread and a standard practice (Minner et al., 1997). Currently,
harder to handle than black layer. York (1993) the application of wetting agents and frequent core
conducted a survey of golf course superintendents aerification have become the management prac-
in the UK to determine how many were experienc- tices most commonly employed for the relief of
ing LDS. A total of 112 superintendents were sur- LDS (Leinauer et al., 2007; Lyons et al., 2009).
veyed, and 86% of them indicated that LDS was a Wetting agent applications are made early in the
problem at their course. season before symptoms appear, and at regular
Because the hydrophobic coating on sand parti- intervals throughout the season, depending on the
cles in LDS is organic, it can be removed with product.
hydrogen peroxide (H2O2). However, this is a dan-
gerous procedure for a golf course putting green,
10.4 Sand System Design
bowling green, tennis court or athletic field.
and Construction
Applying hydrogen peroxide is similar to applying
light rates of glyphosate (Roundup) to control The United States Golf Association (USGA) Green
annual bluegrass (Poa annua) in creeping bentgrass Section Recommendations for Putting Green
(Agrostis stolonifera) turf. The light rate of glypho- Construction (USGA Green Section Staff, 2004)
sate may work, but it may kill all of the grass on are by far the most commonly used methods for
the surface as well. Hydrogen peroxide has been constructing turfgrass sand systems for all kinds of
used successfully many times in recent years to sporting applications. The University of California
control algae in creeping bentgrass on putting Sand Putting Green Construction and Management
greens with no detrimental affect to the turf. These (Davis et al., 1990) method is also popular and a
can be dangerous applications, however. At higher third method, the Airfield System (Airfield Systems,
rates hydrogen peroxide can kill almost anything, 2009), is lesser known but appears to be gaining in
including turf. popularity (Xiong et al., 2006). Each of these sys-
High pH treatments, another potentially danger- tems has specific construction recommendations
ous type of application, may also be effective for that must be followed or the final product may not
controlling LDS (Karnok et al., 1993). This is meet expectations (Gibbs et al., 1993).
another treatment that is hard to recommend Turfgrass sand systems are constructed not only
because of its potential detrimental effects on the for compaction avoidance but also to facilitate
turf. Karnok et al. (1993) demonstrated that an drainage. Subsurface drainage is extremely slow in
application of 0.1 molar sodium hydroxide native soil compared with pure sand. It is not
(0.1 mol/l NaOH) sufficient to saturate the upper 2 uncommon for a new pure sand system to drain at
inches (50 mm) of soil, followed by a water flush of 30 inches (76 cm) per hour or more. As a result, the
one pore volume, significantly reduced hydropho- surface can maintain play shortly following a
bicity if repeated three times. The researchers severe storm or after a week of rainy weather. Such
reported that the severity of turfgrass injury conditions provide immense benefits to players and

152 Chapter 10
spectators. However, these benefits do not last for- recommended as well. The particle size of the
ever (Baker et al., 1999b). sand in the root zone is selected to match the
Turfgrass sand systems have a long but finite drainage of the system. A USGA system is con-
lifetime (Murphy et al., 1993b). They gradually structed to maintain a perched water table – a
become compacted and the drainage eventually zone of nearly saturated sand above a gravel layer
slows (Baker et al., 1999b). However, they can that resides above the soil base (Fig. 10.5). The
provide superior playing conditions for 25 years or California system recommends a root-zone sand
more before they need to be replaced. As you have that sits directly on the soil base. The USGA sys-
learned, quality turfgrass systems produce substan- tem recommends at least 60% coarse-to-medium
tial thatch. The thatch is then degraded by soil size sand and no more than 20% fine sand because
microorganisms to humus. Therefore, healthy turf- the system retains water by design, and the finer
grass systems produce their own soil organic mate- sands are not required. The California system
rials that will eventually promote compaction and drains slowly into the subsoil and does not retain
discourage drainage in a sand system. In addition water. So the California system recommends that
the system is constantly bombarded by silt and clay at least 90% of the root-zone material consists of
carried by wind, rainfall and irrigation. Nature is coarse, medium and fine sand combined, and is
nearly always victorious over man and the sand more tolerant of fine sand because the smaller
system eventually becomes more like soil than particles hold more water in the root-zone mate-
sand. For that reason, and because these aging rial. Neither system recommends that more than a
properties can encourage surface compaction, sur- minor fraction of the root-zone material be of
face drainage should not be ignored in sand sys- very coarse or very fine sand, and silt and clay
tems. It is not wise to rely solely on subsurface should make up not more than a very small pro-
drainage, even on a USGA, California or Airfield portion. Sands that have a broad particle size
System constructed sand system. It is interesting to distribution (meaning that they have a substantial
note that in spite of superior subsurface drainage, percentage (> 5%) of each size category from very
all three of these systems retain more water at the fine to very coarse) are easier to compact than
bottom of small slopes than they do at the top sand with a narrow particle size distribution
(Prettyman and McCoy, 2003; Xiong et al., 2006). (Davis et al., 1990) (Fig. 10.6). Both the USGA
Consequently, if a client is specifically interested in and California systems have proved effective but
a sand system because it drains uniformly, that they require slightly different irrigation manage-
does not appear to be an advantage. Sand systems ment. The Airfield System has not been time
drain quickly, but they do not necessarily drain tested but early indications are promising (McInnes
uniformly. and Thomas, 2008).
Turfgrass sand systems require sands of a spe- Amendments are often added to sand systems to
cific particle size distribution in the root zone, and improve water and nutrient retention (Bigelow
in the USGA system a particular size of gravel is et al., 2004; McClellan et al., 2009). The

USGA California Airfield

30 cm 12 in. Root zone Root zone Root zone

Geotextile

8 cm 3 in. Intermediate
Liner
Gravel Drain
10 cm 4 in.
Drain pipe
Composite superstructure
(mostly air)
Drain pipe

Fig. 10.5. Cross-sectional views demonstrating the general characteristics of the USGA (United States Golf
Association), California and Airfield sand systems.

Growing Grass on Soil, Sand and Salt 153

 (a) Pore size distribution (b) Pore size distribution

Fig. 10.6. Sands with a broad particle size distribution compact more readily than sands with a narrow distribution.
Notice how small some of the pores are in sand A compared with sand B. Sand B is already compacted about as
much as possible but sand A can be compacted further as the small particles continue to settle into the gaps left by
the larger ones.

amendments could be organic or inorganic. Organic distribution range, does not compact like soil but it
amendments have been used historically but does not hold water and nutrients like soil does
inorganic amendments such as calcined clay, either.
diatomaceous earth, zeolite and others show prom-
ise. Organic amendments such as rice hulls and
10.5 Managing Turfgrass
various composts have been used successfully but
on Soil and Sand
the most common amendment is peat. The more
peat contained in the mix, the tighter the medium Would you rather manage turfgrass on soil or
will hold water (Li et al., 2005). An 85:15 by vol- sand? Without a doubt, managing turfgrass on
ume sand:peat mix is common. The sand:peat mix sand is more difficult than managing turfgrass on
encourages rapid establishment compared with soil unless the site is expected to maintain heavy
pure sand (Bigelow et al., 2001; Waltz and McCarty, traffic. Where heavy traffic is expected, a turfgrass
2005). The USGA system tends to encourage the on sand may be sustainable but a turfgrass on soil
use of amendments, but the California system tends very likely is not. If you are a knowledgeable and
to discourage them. Given time, a turfgrass forms experienced turfgrass manager you may prefer to
its own organic soil layer. Some believe that the manage grass on sand regardless of the level of traf-
addition of organic material into a pure sand root fic expected. However, managing grass on sand will
zone can encourage layering problems and incon- require more labor, and therefore more expense,
sistencies in the media. If rapid establishment is than managing on soil where little traffic is
required, the best course of action is to add organic expected. So the expense of growing on sand has to
material. If time is not an important factor, perhaps be justified by a noticeable increase in turfgrass
it is best to use an unamended root zone. Unless a quality and your customer has to be convinced that
consensus is reached or research is performed that the increase in quality is worth the expense.
clearly identifies the best course of action, with a
little knowledge and experience you will have the
Differences in irrigation management
opportunity to form your own opinion.
The important message in this section is this: In general, an uncompacted loam soil has a near
sand, especially sand in a narrow particle size 50:50 split of micropores and macropores.

154 Chapter 10
The micropores hold water and the macropores one of the first industries rationed. Sand systems
hold air. Sand with a narrow particle size distribu- require more frequent irrigation than soil systems.
tion such as that normally recommended for turf-
grass system construction has very few micropores
Differences in nutrient management
and lots of macropores. The sand holds very little
water but contains lots of air. In most climates, Nutrient leaching is an event that you are probably
natural rainfall is sufficient to grow reasonably familiar with. Nutrients are lost through leaching
dense, uniform turfgrass on soil but not on sand. In when their soluble forms percolate downward
most climates, artificial irrigation will be required through the soil in water. Leaching of nitrogen is
for turfgrass grown on sand. common in all types of media, especially sand, but
The primary reason that sand is used in turfgrass most nutrients are held tightly enough by the soil
sand systems is because it does not compact appre- that they resist leaching (Petrovic, 1993). Frequent
ciably until finer materials become part of the fertilization in small amounts or the application of
medium and because it has large pores that drain nitrogen and potassium in slow-release forms is
readily allowing rapid use of the surface following necessary for turfgrass fertilization in sand
rainfall. Because it drains so well, sand is not a systems.
good growing medium unless you have access to As you know, the cation exchange capacity
water. Turfgrass on sand does not necessarily (CEC) holds nutrients in soil. The higher the CEC,
require more water than turfgrass on soil, but it the more nutrients are retained by the soil and the
requires water more frequently. For instance, in lower the likelihood that nutrients other than nitro-
most climates it is not necessary to irrigate a home gen will be lost to leaching. A typical loam soil has
lawn more than once a week during dry periods, a CEC of about 7 to 16 cmol kg−1 of soil (Carrow
and a very general recommendation would be to et al., 2001). A cmol kg−1 (centimole per kilogram)
apply 1 inch (25 mm) of precipitation each week. If is equal to 1 meq. (milliequivalent) per 100 grams
that lawn was on a sand system, it would require at of soil, and these are the two units most commonly
least two irrigations a week at 0.5 inch (13 mm) per used to report soil CEC. It is not necessary to
irrigation. It might even require three applications understand the units as long as you realize what
a week to prevent wilt, depending on the species constitutes a high CEC and what constitutes a low
present, the depth of the root system and the water CEC. A fertile clay loam soil may have a CEC from
retention of the medium. 20 to 50 cmol kg−1 and organic matter is likely to
One advantage of a sand system is that it can have a CEC as high as 140 to 250 cmol kg−1. Sand,
take more water than necessary with no detrimen- in contrast, has a very low CEC. Pure sand is likely
tal effects. Heavy prolonged rainfall can be quite to have a CEC of less than 1 cmol kg−1 and an
difficult to deal with on soil but not on sand. 85:15 peat-amended sand will probably have a
Turfgrass can not sustain itself in saturated soil for CEC ranging from 1 to 3 cmol kg−1. Obviously,
more than a few days before it declines for lack of sand is not a growing medium that retains many
root oxygen. Sand systems drain readily and are nutrients.
rarely saturated for more than an hour or two. Nutrient deficiencies are common in sand sys-
Once saturated, water in a sand system drains tems. For that reason, soil testing of sand systems
through the subsurface irrigation. Consequently, should be done more frequently (two to three times
sufficient root oxygen is normally always available. per year) than it is for native soil. In addition,
For that reason, managers of sand systems tend to nutrient management has to be adjusted to accom-
over-water rather than under-water. Although that modate the poor nutrient retention of sand systems.
may not be harmful to the turf in most cases, it Nutrients should be applied to close-mown sand
quickly becomes expensive if you have to purchase systems like bowling and putting greens weekly to
water or purchase energy to operate your pumping maintain consistent color and growth. Higher
stations. So unless you work on one of those mown turf, athletic fields for instance, may be fer-
extremely rare sites where you can irrigate from a tilized once every 2 weeks, but weekly is better.
natural aquifer by gravity flow, irrigation is expen- Sand systems generally do not require more ferti-
sive and over-irrigation is a waste. In addition, lizer than soil but they do require smaller amounts
water and energy are in limited supply and should per application more often. Although soil rarely
be conserved. If we don’t conserve them we will be requires additional micronutrients as long as the

Growing Grass on Soil, Sand and Salt 155

pH is close to normal, sand systems usually respond sand systems so that nutrients are available as the
best to one or two broad-spectrum micronutrient plants need them but not in an excessive amount
applications annually. which will leach away with the next major precipi-
A spray application of nutrients to sand systems tation event. An alternative is the use of slow-
weekly is often called “spoon feeding”, indicating release fertilizers that release small amounts of
that only a small amount of nutrients is applied. nitrogen and potassium over time. As you learned
The term “foliar application” or “foliar feeding” is in Chapter 7, slow-release fertilizers are more
often used interchangeably with spoon feeding. expensive than quick-release sources but can be
Spoon feeding, however, implies a light application applied less often, thereby providing a savings in
of nutrients, whereas foliar application implies a labor expense.
light application of water that usually contains
only small amounts of nutrients to prevent foliar
10.6 Managing Turf on Soils High
burn. A typical spray application applies a larger
in Salts
amount of nutrients than a spoon-feeding applica-
tion and a larger amount of water than a foliar Many ornamental plants, including trees, grow
application. My objective here is not to correct the very poorly in salty soils. Turfgrasses in general, are
use of these terms but to make certain that you some of the most resistant species to poor soil con-
understand what is meant when they are used in ditions. Nonetheless, turfgrasses can be difficult to
this text. There are cases of calcareous sands that maintain in soils high in salts, especially at the aes-
have to be fertilized by foliar application because thetic and/or functional levels that many of your
the pH of the medium is so high that many nutri- customers have come to expect. The purpose of this
ents precipitate from the soil solution and are no section of the text is to supply you with general
longer plant available once they enter the sand. information needed to manage turfgrass on soils
There are some alkaline soils that work the same high in salts. You will learn how salts affect grasses
way. Phosphorus, iron and manganese are the most and techniques that can help you manage grasses
likely nutrients to be deficient in the case of calcar- on salty soil. However, if you are tasked with man-
eous sand or alkaline soil (Carrow et al., 2001). In aging a turfgrass on a soil high in salts, especially
many cases trying to adjust the pH of calcareous sodium, you will need to seek considerably more
sands and some alkaline soils is not possible or not education than can be supplied here. Additional
practical. It could take tons of sulfur over 100 sources of information are recommended at the end
years to affect the pH in many of these sands and of the chapter. In addition, professional or expert
soils. Consequently, foliar fertilization becomes the consultation should be considered before attempt-
only available option for growing good grass. In ing to develop a program for the management of
most cases, alkaline sands and soils are manageable turfgrass on a salty site. Each site has unique
if foliar fertilization is used. characteristics, requiring techniques that range
Generally, the nutrients of major concern in sand from simple leaching solutions to complex pro-
systems are nitrogen and potassium. Not only are grams of drainage reparation, salt remediation,
these the two fertilizer nutrients needed in the foliar feeding, chemical irrigation management and
greatest amount, they are also the two nutrients perpetual leaching practices.
most likely to be lost to leaching. In a sand system,
potassium is not held tightly as it is in soil and it
Dealing with salts on sand-based systems
leaches readily just as nitrogen does unless suffi-
cient organic material is present to bind it (Petri In addition to the differences that we have dis-
and Petrovic, 2001). As a sand system ages, organic cussed concerning the management of soil versus
material increases in the system and more nutrients sand systems, one more component requires atten-
are retained (McClellan et al., 2007). So frequent tion. Irrigated sand systems tend to accumulate
fertilization of an old system (10 years old or more) salts. As was indicated earlier, sand systems require
may not be as critical as that of a new system. frequent, light irrigation in comparison with soil
Regardless, sand systems always need more fre- because the sand systems retain little water. As you
quent fertilization to grow their best turf than do irrigate under these light, frequent restrictions the
soil systems, provided all other factors are the soil is only wet for a short distance into the profile
same. Spoon feeding is the best process to use on and the improved aeration of a sand system, rapid

156 Chapter 10
water use by turfgrass and shallow wetting add to
Classifications of salt-affected soils
rapid drying of the irrigation applied. An irrigation
source that does not contain salts of one type or Up to this point, I have avoided the use of the term
another is rare. In fact, as will be discussed later in “saline”. Saline traditionally means a solution high
this chapter, irrigation water that is too pure is in salts. However, in the study of plants and plant
undesirable. As the water in the soil evaporates the management, sodium has a special classification.
salts in the water are left behind. These salts build Consequently, saline as it refers to irrigation and
up with frequent irrigation and drying until main- soil water solutions becomes confusing because the
taining adequate turf becomes difficult. So they reader is not sure whether the author is referring to
need to be removed periodically. all salts or salts other than sodium. For that reason,
It is generally not difficult to remove salts from in this text, saline means salts other than sodium.
sand systems. It usually only requires a period of A solution high in sodium will be called sodic and
heavy rainfall or an irrigation event specifically a solution high in both sodium and other salts will
designed for salt removal. The most common mis- be called saline+sodic so that there is no mistake.
take managers make concerning designed-to-leach The United States Salinity Laboratory (USSL) uses
irrigation events is not applying enough water these classifications to identify categories of
(Fig. 10.7). salt-affected soils (USSL Staff, 1954). The USSL
In typical sand used for USGA system classifications are the most widely used in the
construction, up to 30% of the profile consists of world but there are countries or regions and at
macropores. Because the profile is 12 inches (30 cm) least one continent, Australia for instance, that
deep, the macropores make up almost 4 inches because of the soils that are most prevalent, may
(10 cm) of the profile. If you want to fill all of those have slightly different classifications.
pores with water, you have to apply 4 inches The soil solution of a saline soil has an electrical
(10 cm) of irrigation. Once the pores are filled, you conductivity (ECe) > 4 dS m−1 and an exchangeable
can then begin to apply more irrigation to leach the sodium percentage (ESP) of below 15. The unit
soluble salts. Consequently, about a 5-inch (13-cm) designation “dS m−1” is an abbreviation for deciSie-
irrigation event is required to leach the system. On mens per meter. A Siemen is a measure of electrical
a 12-inch profile that is approximately 3.1 gallons conductivity defined as the reciprocal of electrical
of water per square foot (127 l m−2), a tremendous resistance. Conductivity refers to the ability of a
amount of water. On a USGA system, you will material to carry electricity. There are relatively
know when leaching begins. The main drain will simple electrical meters used to measure solution
suddenly fill and water will come pouring out of conductivity as an Ohm meter or multimeter meas-
the entire pipe. Make sure that the drainage pipe is ures resistance. A conductivity of 4 dS m−1 is the
clear before you apply a leaching event and remem- conductivity designated by the USSL to distinguish
ber that many of your nutrients will be leached between a saline soil solution and a typical soil
along with the other salts. solution.
The unit “ESP”, the exchangeable sodium per-
centage, is the proportion of exchange sites on the
10 cm 4 in.
Precipitation needed to fill the root zone cation exchange capacity of the soil occupied by
before leaching can begin
sodium reported as a percentage. A measure of 15
ESP means that 15% of the exchange sites in the
30 cm12 in. Root zone
soil are occupied by sodium. The USSL classifies
sodic soils as those soils with an ESP of 15% or
Gravel
greater.
10 cm 4 in.
A saline soil is usually easier to manage than a
sodic soil. A saline soil will usually, but not always,
Drain pipe have a basic pH, but this is normally less than 8.5.
Saline soils are often called “white alkali” because
Fig. 10.7. A designed-to-leach irrigation event requires
a considerable amount of water. A 12-inch (30 cm) root they are usually basic and they often have a white
zone that contains 30% air-filled pore space requires crust on them, especially on the low spots where
an irrigation event of almost 4 inches (10 cm) just to fill water collects and then dries. Conversely, sodic
the soil pores before leaching can begin. soils are called “black alkali”. Sodic soils have an

Growing Grass on Soil, Sand and Salt 157

ECe < 4 dS m−1 and an ESP > 15. By definition, growth when they experience minor saline or sodic
sodic soils contain considerable sodium but no conditions (Qian et al., 2001). Bermudagrass
more than a typical amount of other salts. Sodic (Cynodon spp.) species and cultivars differ in their
soils are nearly always basic, with a pH usually tolerance to salt (Youngner and Lunt, 1967).
greater than 8.5. These are the most difficult salt- However, they all appear to respond to high-salt
affected soils to manage. media by decreasing shoot growth and increasing
Saline+sodic soils are those with ECe > 4 dS m−1 root growth, which results in increasing root-to-
and ESP > 15. Although this combination sounds shoot ratios (Dudeck et al., 1983). Most warm-
devastating, the resulting soil is often easier to season turfgrasses respond to salt media in the
manage than a sodic soil. The pH is usually lower same way (Marcum, 1999), and some cool-season
than 8.5 and the soil tends to take on the charac- grasses are believed to respond similarly.
teristics of a saline soil rather than a sodic soil. In As salinity or sodicity increases in a soil solution,
a way, the saline components have a positive buff- its water potential decreases. Consequently, it
ering effect on the negative sodium component. becomes harder and harder for grasses to take up
soil water. Salt-tolerant grasses respond to this
decline in soil water potential by extending their
The effects of salty soil on turfgrass
root systems. Their roots grow both deeper and
You probably remember from chemistry class that with more mass (Marcum, 2008). Most salt-tolerant
salts dissociate easily into cations and anions in warm-season turfgrasses also have the ability to
water. Salt-affected soils usually contain cations of lower the water potential within their roots and
sodium, potassium, calcium and magnesium and shoots to encourage the absorption and transloca-
anions of chloride, sulfate, carbonate and bicarbo- tion of water and nutrients into the roots and
nate that dominate the soil solution. All of these throughout the plant (Marcum and Murdoch,
components are found in and, with the possible 1990). The lower the water potential of the media,
exception of sodium, are necessary for plant func- the lower the plant’s water potential becomes
tion. It is not the existence but the proportion of (Peacock and Dudeck, 1985; Dudeck et al., 1993).
these components in the soil solution that deter- At least some turfgrass species practice a salt
mines a soil’s plant support characteristics. avoidance process called ion exclusion (Volkmar
Most, but not all, salt-affected soils are found in et al., 1998). Ion exclusion occurs when salts are
arid regions. Salts are easily leached by rainfall, but compartmentalized and excreted through salt
in dry regions rainfall does not occur in a sufficient glands on the leaves. Sodium and chlorine are
amount to perform this leaching function. If rainfall known to be excreted from leaf salt glands on sea-
equals evapotranspiration, salts usually leach away. shore paspalum (Paspalum vaginatum), manila-
Otherwise they remain in the upper soil layers and grass (Zoysia matrella), mascarenegrass (Zoysia
cause problems for plant growth. Excess salinity tenuifolia), St. Augustinegrass (Stenotaphrum
affects grasses and other plants by making it more secundatum), bermudagrasses, Japanese lawn grass
difficult for them to access water and nutrients. (Zoysia japonica), centipedegrass (Eremochloa
Saline or sodic conditions result in poor growth ophiuroides) and Kentucky bluegrass (Poa praten-
with additional symptoms similar to drought stress sis) (Marcum and Murdoch, 1994; Qian et al.,
(Harivandi et al., 1992). Leaf blades are likely to be 2001). The efficiency of salt gland excretion and
narrow and stiff with a blue-green coloring that the density of salt glands on the leaves are known
eventually leads to wilt. As the soil solution becomes to influence the salt tolerance of zoysiagrasses
more saline its water potential becomes lower, caus- (Zoysia spp.). The salt tolerance of bermudagrass
ing it to hold water more tightly. Plants have more cultivars is also determined by the efficiency with
difficulty absorbing this tightly held water, which which they excrete sodium through their salt
causes drought-like symptoms. The plants are, in glands (Marcum and Pessarakli, 2006). As we learn
fact, experiencing a physiological drought. more about these mechanisms, it may be possible
True salt-tolerant plants may experience higher to design management practices to specifically
rates of shoot growth under minor saline or sodic encourage reductions in water potential and
conditions (Marcum, 1999). However, accelerated efficient ion exclusion.
root growth is more likely. In fact, even some mod- When the soil medium is high in sodium, tissue
erately salt-tolerant grasses will increase root concentrations of nutrients such as calcium,

158 Chapter 10
potassium, magnesium, manganese and phosphorus sites can be expensive but are critical for the devel-
are affected (Dudeck and Peacock, 1993). As sodium opment of an effective management program
levels increase in the soil and in the plant these nutri- (Carrow et al., 2001). The conditions on salt-
ent levels decline. Thankfully, nitrogen and iron do affected sites can vary widely, often based on eleva-
not seem to be affected (Ackerson and Youngner, tion, and a sufficient number of samples should be
1975; Dudeck and Peacock, 1985a). However, high taken to form a truly representative composite. On
sodium in the soil solution interferes with calcium highly problematic sites, the advice of a profes-
and potassium uptake, and these are two very sional or expert should be sought. Qualified special-
important nutrients. Sodium also tends to replace ists may be found as commercial consultants, soil
calcium in root membranes, leading to loss of mem- and water laboratory analysts, agricultural univer-
brane integrity, and it may replace potassium in sity personnel and others in your particular field of
some reactions, thus disrupting metabolism. turfgrass management. These advisors may suggest
Sodic soils are especially detrimental to plant plans that include foliar application of nutrients,
growth because the high sodium content causes the aggressive plans for cultural improvement, major
soil to lose its structure. The resulting soil system is drainage renovations, leaching programs and con-
similar to soils that have been highly compacted. version to more salt-tolerant grasses.
Soils are normally very high in calcium and magne-
sium, and those two elements usually occupy most
Irrigation concerns on salt-affected sites
of the cation CEC. However, in sodic soils, sodium
replaces some of the calcium and magnesium in the In our industry and others, poor-quality irrigation
CEC, causing soil structure to deteriorate. The water is a potential source of salts. Clean water is
replacement of calcium with sodium in the soil CEC always in high demand. Because lawns, parks, golf
causes the bonds within soil aggregates to weaken, courses and even athletic fields are not considered
and the aggregates eventually break up into fine particularly important in comparison with food
particles of clay and humus, destroying soil struc- crop irrigation and other human uses for water, we
ture. Sodic soils not only hold a lot of water and may have to use poor-quality water or be content
hold it tightly, they become impermeable as the with no irrigation at all. Therefore, irrigation and
aggregates fall apart and the macropores disappear. leaching with water containing high salt concentra-
Although sodicity is, in most cases, the most tions is becoming more common for turfgrass
detrimental condition that exists in salt-affected managers. The use of this water for irrigation and
soils, saline soil solutions can sometimes contain the amount of leaching that is appropriate follow-
toxic amounts of certain salts. While chlorine, ing each use has to be considered.
bicarbonates or hydroxides can reach toxic levels The USSL lists salinity hazards in water as low
in saline or sodic soils, boron is the element most (ECw < 0.25 dS m−1), medium (0.25 £ ECw < 0.75 dS
likely to be found in amounts toxic to turf (Carrow m−1), high (0.75 £ ECw < 2.25 dS m−1) and very high
et al., 2001). Boron toxicity most commonly occurs (ECw ³ 2.25 dS m−1). Sodicity is measured in SAR,
in arid regions where irrigation water is high in the sodium adsorption ratio, rather than as the
boron. Its symptoms are necrosis of leaf tips, and exchangeable sodium percentage (ESP) introduced
regular mowing tends to temporarily relieve the earlier. The SAR is a comparison of sodium con-
stress as the boron that accumulates in the tips is centration with calcium and magnesium concentra-
mowed off. According to Carrow et al. (2001), tion. It is calculated as the number of CEC sites
grasses can tolerate soil boron accumulations as occupied by sodium, divided by the square root of
high or higher than 10 parts per million (ppm) half of the sum of the sites occupied by calcium
depending on species and cultivar. However, the plus magnesium:
authors recommend levels below 1 ppm (1 mg kg−1)
for sensitive grasses and other plants.
SAR =
[Na+ ]
Developing a management program
1
([Ca2+ ] + [Mg2+ ])
2

The first step in developing a management program

for turfgrasses grown on salt-affected soil is proper The USSL classifies sodicity as low (SAR < 10),
assessment. Chemical assessments of salt-affected medium (10 £ SAR < 18), high (18 £ SAR < 26) and

Growing Grass on Soil, Sand and Salt 159

very high (SAR ³ 26). Furthermore, because salinity salt-affected site but will not be able to achieve the
has a positive effect on the negative sodicity of a aesthetic or functional value expected. In that case,
solution, a more powerful analysis results from a salt-tolerant species (and cultivar) must be selected
considering the salinity and sodicity in combina- to meet site requirements. In some cases, a less salt-
tion. Even a soil with low sodicity can become tolerant turfgrass may be invaded by more salt-
more difficult to manage if very pure irrigation tolerant weeds on a salt-affected site. However,
water is used. Pure water can cause a high propor- selection of a salt-tolerant turfgrass species and
tion of all the soil salts, including calcium, to leach, cultivar gives the turfgrass the opportunity to out-
thereby causing a disruption in surface stability compete most weeds.
that affects permeability. In the USA, the deicing of highways with
sodium chloride has led to the need for and to the
selection of salt-resistant turfgrass species and
Leaching
cultivars. For instance, alkaligrass (Puccinellia
The most common method for improving salt- spp.) was first considered for use as a salt-tolerant
affected sites is to develop a leaching program for turfgrass for low-maintenance sites after discov-
the downward removal of salts from the root zone. ery along salt-affected roadsides in parts of the
For this program to work, however, the water and country where it was not a particularly common
salt must have someplace to go and the soil must be species (Butler et al., 1974). Alkaligrass is widely
permeable enough to allow adequate infiltration. considered the most salt tolerant of the cool-sea-
The portion of irrigation water and natural precipi- son grasses suitable for low maintenance turfgrass
tation that moves through the root zone carrying (Marcum, 2008). Inland saltgrass is probably the
salts with it is called the leaching fraction, or some- most salt tolerant of the warm-season grasses suit-
times, the leaching requirement. The higher the able for low-maintenance turf (Marcum, 2008).
leaching fraction, the more salts removed. In a Species, however, differ by cultivar in their adapt-
saline+sodic soil, another salt, usually gypsum, ability to salt-affected sites. The most popular
must be applied to help leach the sodium in prefer- turfgrass in the northern USA, Kentucky blue-
ence to other less detrimental salts. Gypsum is cal- grass, is not particularly salt tolerant (Torello and
cium sulfate and is used to replace sodium ions Symington, 1984), but there are cultivars of the
with calcium ions. Leaching is possible with salty species that demonstrate moderate salt tolerance
irrigation water, but the leaching fraction must be (Qian et al., 2001). Seashore paspalum is gener-
substantially greater to have a desirable effect. ally considered the most salt tolerant of the turf-
To permit leaching, irrigation water must be grass species suitable for highly maintained
applied to soil saturation and move through the turfgrass, but salt-tolerance differs by cultivar
root zone carrying salts with it. Consequently, irri- within the species (Lee et al., 2004a). There could
gating past the point where water runs off is not be cases where a highly salt tolerant cultivar of
productive. It is best to irrigate for the length of manilagrass, mascarenegrass, St. Augustinegrass
time required to saturate the soil and stop before or bermudagrass performed better on a salt-
runoff. Following a short period, perhaps 1 to 3 affected site than a poorly salt-tolerant seashore
hours depending on how rapidly the soil drains, paspalum (Dudeck and Peacock, 1993; Lee et al.,
irrigation can be resumed again to saturation. This 2004b). Take care to select the best cultivars
process can be continued, if necessary, until the within the species when salt tolerance is a major
desired leaching fraction is reached. Leaching frac- consideration.
tions are best calculated by a qualified laboratory Certain cultivars of creeping bentgrass (Agrostis
which will advise you on what information is stolonifera) are reasonably salt tolerant (Marcum,
required and how to gather it. 2001). However, in general, this cool-season species
can not compete with many warm-season species
for salt tolerance. Particular cultivars of tall fescue
Salt-tolerant grasses
(Festuca arundinacea; synonym: Schedonorus
Compared with most plants, grasses are relatively phoenix) (Lunt et al., 1961; Greub et al., 1985)
tolerant of salinity. However, there are some grasses creeping red fescue (Festuca rubra) (Harivandi
that will grow on salt-affected sites where others et al., 1982; Greub et al., 1985; Torello and Rice,
won’t. More often, turfgrasses will grow on a 1986), and perennial ryegrass (Lolium perenne)

160 Chapter 10
(Dudeck and Peacock, 1985b; Greub et al., 1985) applications more often. Because sand is a basically
exhibit moderately good salt tolerance. Salt toler- inert system, a knowledgeable turfgrass manager
ance of species and cultivars may also be affected has the opportunity to control water and nutrients
by growth stage (Harivandi et al., 1982; Dudeck more precisely and to vary inputs more easily when
and Peacock, 1985b; Wu and Lin, 1993) with ger- circumstances change. Soil, however, is more for-
mination and seedling response considered more giving of mistakes and easier for a novice to han-
sensitive to salt than mature growth (Harivandi dle. Where compaction is a serious problem, sand
et al., 1992). Choosing a salt tolerant species and is advantageous, but if a site is vast or only sustains
cultivar is obviously the easiest way to help avoid light-to-moderate use, soil is superior. Root respira-
poor turfgrass performance during construction on tion that provides for adequate root growth and
a salt-affected site. Renovation to a salt tolerant plant transpiration determine which medium
species and cultivar is also an important considera- performs best under the requirements of the site.
tion on established turf. Soil salinity and sodicity affect turfgrass by
osmotic inhibition of water uptake, by creating ion
imbalances in plants and, in some cases, by ion
10.7 Chapter Summary
toxicity. Although turfgrasses respond to osmotic
Native soil mixtures differ substantially from arti- stress by increasing their root growth and root and
ficially constructed sand root zones, not only in shoot water potential, they can be overcome by
their constitution but in their management. physiological drought in salt-affected soils. The
Although sand is a far better system for many turf- ability of turfgrass species to tolerate salt-affected
grass surfaces used for sporting purposes than is soil ranges from those that are salt sensitive to
soil, sand requires a more knowledgeable manager those that are highly tolerant. Sensitivity to
and is not without its problems. Compaction is not salt-affected soils also differs substantially among
a problem on sand systems, but black layer and cultivars of a particular species. Most of the salt-
localized dry spots are fairly common. Sand systems tolerant turfgrasses are warm-season species, but
are expensive to construct and have a finite life- some cool-season species demonstrate moderate
time. A poorly constructed sand system can be very tolerance.
difficult to manage. Nonetheless, a properly con- Salt-affected soils are classified as saline, sodic or
structed sand system is far superior for sites that saline+sodic. In addition to osmotic inhibition, ion
have to sustain regular traffic than a soil system imbalance and ion toxicity, sodic soils can destroy
would be. Not only are sand systems highly com- soil structure, thereby seriously affecting permea-
paction resistant, they also drain rapidly, providing bility. For that reason, sodic soils are hardest to
firm playing surfaces following rainfall when soil manage and require chemical treatment with gyp-
sites could not be used. sum or other means to positively affect soil
Aeration is the main factor that prevents native structure.
soils from providing excellent playing surfaces. Leaching is the most common practice for turf-
Most turfgrass areas are required to sustain moder- grass improvement on salt-affected soil. A leaching
ate traffic. For that reason, aerification practices fraction at a problematic site is determined after
are required to promote enough rooting activity for critical assessment of the soils and irrigation. On
a turf on a soil site to maintain excellent health and sand systems, leaching is practiced regularly, in
aesthetic value. However, soil has an advantage most cases annually, according to need. Salts tend
over sand in nutrient and water retention. For the to build up near the surface of sand systems
most part, nutrients are tightly bound and gener- because the practice of light, frequent irrigation
ally available in soil, but are poorly held and easily does not allow for leaching. Salt-affected soils and
leached in sand. Soil remains damp following rain- poor irrigation water negatively affect respiration
fall, providing water for the turf, but sand drains and transpiration.
quickly, providing mostly air. For that reason, the
systems require different management.
Sand requires frequent applications of nutrients Suggested Reading
and frequent irrigation compared with soil. Beard, J.B. and the United States Golf Association
Although the total input of water and nutrients are (2002) Turf Management for Golf Courses, 2nd edn.
approximately the same, sand requires smaller Ann Arbor Press, Chelsea, Michigan.

Growing Grass on Soil, Sand and Salt 161

Carrow, R.N. and Duncan, R.R. (1998) Salt Affected Turf- Puhalla, J., Krans, J. and Goatley, M. (1999) Sports Fields:
grass Sites – Assessment and Management. Ann A Manual for Design, Construction and Maintenance of
Arbor Press, Chelsea, Michigan. Sports Fields. Ann Arbor Press, Chelsea, Michigan.
Carrow, R.N., Waddington, D.V. and Rieke, P.E. (2001) White, C.B. (2000) Turf Manager’s Handbook for Golf
Turfgrass Soil Fertility and Chemical Problems: Course Construction, Renovation, and Grow-in.
Assessment and Management. Ann Arbor Press, Sleeping Bear Press, Chelsea, Michigan.
Chelsea, Michigan. Witteveen, G. and Bavier, M. (1998) Practical Golf Course
Cockerham, S.T. (2008) Culture of natural turf athletic Maintenance: The Magic of Greenkeeping. Sleeping
fields. In: Pessarakli, M. (ed.) Handbook of Turfgrass Bear Press, Chelsea, Michigan.
Management and Physiology. CRC Press, Boca
Raton, Florida, pp. 151–170.
Suggested Websites
Harivandi, M.A., Butler, J.D. and Wu, L. (1992) Salinity
and turfgrass cuture. In: Waddington, D.V., Carrow, Airfield Systems (2009) Home page. Available at: http://
R.N. and Shearman, R.C. (eds) Turfgrass. ASA- www.airfieldsystems.com (accessed 19 Jan 2010).
CSSA-SSSA (American Society of Agronomy-Crop FAO (Food and Agriculture Organization of the United
Science Society of America-Soil Science Society of Nations) (1992) The use of saline waters for crop pro-
America), Madison, Wisconsin, pp. 207–230. duction. FAO Irrigation and Drainage Paper No. 48,
Marcum, K.B. (2008a) Relative salinity tolerance of turf- T0667/E (Rhoades, J.D., Kandiah, A. and Mashali,
grass species and cultivars. In: Pessarakli, M. (ed.) A.M.). Available at: http://www.fao.org/docrep/
Handbook of Turfgrass Management and Physiology. T0667E/t0667e01.htm#TopOfPage (accessed 3 Feb
CRC Press, Boca Raton, Florida, pp. 389–406. 2009).
Marcum, K.B. (2008b) Physiological adaptations of turf- USGA (United States Golf Association) (2004) USGA
grasses to salinity stress. In: Pessarakli, M. (ed.) Recommendations for a Method of Putting Green
Handbook of Turfgrass Management and Physiology. Construction: 2004 revision (Green Section Staff).
CRC Press, Boca Raton, Florida, pp. 407–418. Available at: http://www.usga.org/course_care/
McCarty, L.B. (2001) Best Golf Course Management ar ticles/constr uction/greens/Green-Section-
Practices. Prentice Hall, Upper Saddle River, New Recommendations-For-A-Method-Of-Putting-Green-
Jersey. Construction/ (accessed 6 Jan 2010).
McCarty, L.B. and Miller, G.L. (2002) Managing Ber- USSL (United States Salinity Laboratory) (2010) Home
mudagrass Turf: Selection, Construction, Cultural page. Available at: http://www.ars.usda.gov/Main/
Practices and Pest Management Strategies. Ann site_main.htm?modecode=53-10-20-00 (accessed 1
Arbor Press, Chelsea, Michigan. Feb 2010).

162 Chapter 10
 The Ecology of Turfgrass
11 Management

Key Terms
Ecology is the interaction of an organism with its own species, with other species and with its environment.
An environment is the sum of all natural and artificial conditions under which a particular organism is expected to
live and grow.
Applied ecology is the use of theories and models to study and understand human impact on the human
environment.
Physiological ecology, also called ecophysiology, refers to the response of an organism to environmental
factors such as light, temperature, nutrients and others.
A population is the sum of individuals of a particular species occupying a particular space at the same
time.
The carrying capacity of an ecosystem is the number of individuals within a population that can be supported by
the current environment and with the resources available. Carrying capacity can also be considered the
greatest density possible at the resource level available.
The law of the minimum states that population growth is limited by the amount of the most limiting resource
during the most limiting period of the year.
Density is the number of individual plants in a given area. In turfgrass, however, we measure density by the
number of tillers, also called shoots, instead of by the number of plants.
The law of limiting factors states that population growth is not only limited by too little of a resource but is also
limited by too much of a resource.
A community is a collection of populations – plant, animal, microorganism, etc. that interact with each other either
directly or indirectly.
Interspecific competition occurs among different species.
A niche is a resource or set of resources and a microenvironment within a community for which a population is
uniquely adapted and for which it can successfully compete.
An ecosystem is the sum of activities and interactions within a biotic community as influenced by its abiotic
environment.
Landscape ecology is a form of applied ecology that recognizes human activity as a factor that combines with
natural disturbances to determine the spatial patterns that constitute a landscape.
Intraspecific competition occurs when individuals of a population compete with each other for resources.
A blend is a combination of turfgrass cultivars of the same species.
A mixture is a combination of turfgrass species.
Biomass is the amount of plant material by weight in a given area. Clipping yield is a measure of biomass but it
does not account for the total plant biomass in a given area of land; clipping yield only measures the biomass
over a particular height above that land.
The −3/2 power law of self thinning, also called the −1.5 self thinning law, is a mathematical description of
the natural balance that occurs during intraspecific competition between plant biomass and density. In
mown turfgrass, the slope of the self-thinning line is different from that in unmown plants and is believed to
be −1/2 = −0.5.
Turfgrass ecology could be explained as the use of natural means to maintain a managed turfgrass system to a
particular level of human expectation with the fewest inputs and the least environmental impact.

©CAB International 2011. Turfgrass Physiology and Ecology (G. Bell) 163
 time and use as important factors are identified
11.1 How to Make Nature Work For You
and incorporated.
Ecology is a very old science and an even older Applied ecology was not meant to be used by turf-
philosophy. It is deeply grounded in the workings grass managers and other agronomists. As with other
of nature and nature’s effects on life, but it does not sciences, it was meant to be a means of gathering
depend on evolution and the controversies com- knowledge of the natural world and its many interac-
mon to evolutionary philosophy. We can see and tions with us and other organisms. Some ecologists,
study ecological activity at any time within the in fact, believe that managed turfgrass should be
natural environment around us. Because ecological eliminated. They argue that the inputs required to
principles have a profound effect on plants, or vice properly manage turf exceed its environmental ben-
versa, it benefits us to stay informed of advances in efits. It would not be wise for us to overlook those
ecological science. beliefs. Indeed, these ecologists may be correct.
Although ecology is a very old endeavor, it did However, turf has many properties that positively
not become a popular science until the 1960s or influence our environment(s). Instead of criticizing
1970s, when a majority of people in many countries our use of turf, perhaps these ecologists could recog-
began to realize that they were having a critical, nize its benefits and teach us how to manage it using
often negative, effect on their environment(s). At environmentally sound practices. In truth, they have
that time, our environmental impact began to already, but unknowingly, done that for us to some
become a subject of great interest. Ecology was extent. We can use what they have taught us not only
soon recognized as a means to study and change to affect our environment(s) more positively but also
environmental impact and society began to notice to manage our turf more effectively. As scientists and
and embrace it. practitioners we can build on those lessons to con-
In very simple terms, ecology is the study of how tinue learning better techniques.
environments influence the organisms that live in
them and how the organisms affect each other.
11.2 Introduction to Ecological Theory
Ecology, however, like nature from whence it is
derived, is extremely complex and at least for the If you are to use ecology to your advantage, it is
present, beyond our realm of total understanding. best to consider it a concept rather than a science
Nonetheless, many of the principles of ecology that or a theory. It is a way of thinking, not a means of
others have studied and developed are sound and addressing a practical problem. If you can imagine
can be very useful to us as turfgrass managers. always using nature first and synthetic or industrial
The human desire to understand our environ- means of management second, you are thinking
ment and limit our negative impact upon it led to ecologically. If you can manage your turf to out-
the development of applied ecology, one of the compete weeds to the point that you can physically
many branches of ecological study. Applied ecol- remove them more cheaply than you can apply a
ogy is the use of theories and models to study pesticide, you are practicing ecological manage-
and understand human impact on the human ment. If you can convince your end user or cus-
environment. You could say that applied ecology tomer that a 1 or 2% weed encroachment is
is in its infancy. Moreover, there are so many acceptable, you are saving them money and you are
major and minor principles that affect human also practicing ecological management. Ecological
interaction with our environment(s) and our management not only reduces our environmental
potential for detrimental influence that the study impact, it nearly always saves money. The major
of applied ecology will probably never end. problem is that we have not identified the manage-
Practitioners of applied ecology endeavor to ment practices or management systems necessary
devise mathematical models that explain human to meet our customer expectations using ecological
impacts on our environment(s). Such endeavors means exclusively. It would benefit us to do so.
are ridiculously difficult but nevertheless impor-
tant. Models can be used to explain what hap-
Ecological concepts
pens for a given human input and how that input
affects the local environment. Currently, most of According to ecology, we live in and are affected by
the models that have been developed are not an environment which includes all natural and
completely accurate but they will improve with artificial factors around us. We can think of this

164 Chapter 11
environment as huge, incorporating the environ- is the number of individual plants that can be
ment surrounding all humans, or we can think of it supported by the amount of resources available.
as very small – the environment that only affects us At that point no new plants can be added. As the
personally. Thinking too big is far too complex and population nears the carrying capacity, growth
too vague for any practical purpose. Therefore, slows and eventually levels off as plants die and
think small, think about the environment that are replaced in a sort of dynamic equilibrium
affects you and you can learn how to affect those (Fig. 11.1).
factors that influence it. Think about the environ- The idea of carrying capacity began in early
ment that affects a particular species and cultivar of ecological study with a proposal called the law of
your turf – in a particular location, under a particu- the minimum (Leibig, 1840). The law of the mini-
lar mowing height and frequency, with or without mum states that a population’s growth is limited
irrigation – and at what level it needs to be man- by the amount of its most limiting resource during
aged, and you can make progress in ecological its most limiting period of the year. Hence, the law
management. of the minimum not only considers the resources
We have come a long way in this text, and up to available to the population but also the environ-
this point everything that we have studied has ment. Population growth may fluctuate with
been a form of ecology. That particular branch of season but the population can only temporarily
ecology is called physiological ecology. grow beyond the resources available during its
Physiological ecology, also called ecophysiology,
refers to the response of an organism to environ-
mental factors such as light, temperature, water, 50
Carrying
soil and nutrients. We have not only studied the capacity
effects of light, temperature, water, soil and nutri- 45
ents, we have studied the physiological plant
activities that occur in response to the factors dis- 40
cussed. We have actually been working within the
realm of ecology all along. Now, instead of physi- 35
ological ecology, we are going to concentrate on
applied ecology, a lesser used science for plant 30
management.
Time

25
An ecological population
20
A population is a group of individuals of the
wth
same species living in a particular location at the n gro
same time. The size of the location is determined 15 u latio
Pop
by the observer. The population of interest could
be all of the Kentucky bluegrass (Poa pratensis) 10
plants growing in a home lawn or it could be all
of the creeping bentgrass (Agrostis stolonifera) 5
plants growing on a golf course putting green.
The individuals of a population compete with 0
each other and with other organisms. However, if 0 10 20 30 40 50 60 70
a plant has a minimal amount of everything that Density (shoots per unit area)
it needs it can continue to grow and reproduce
throughout its lifetime. Therefore, if the plants in Fig. 11.1. As a population of plants reaches its
carrying capacity, population growth slows and
the population have all of the resources that they
eventually levels off in a sort of dynamic equilibrium.
need, significant competition does not occur. The When the carrying capacity has been reached, a
population continues to grow as offspring are new plant cannot be added unless an old plant dies.
produced more rapidly than aged plants die. The only way to increase the population is to supply
Theoretically, the population will continue to sufficient new resources to increase the carrying
grow until it reaches its carrying capacity, which capacity.

Ecology of Turfgrass Management 165

most limiting season. For example, the density of not always the dominant one. In our systems we
a cool-season grass lawn may improve in the have chosen the dominant species, the turfgrass,
spring and the fall but its ability to sustain that but we also have to be able to maintain that domi-
density will always be limited by the summer sea- nance. Our management asserts turfgrass domi-
son. We already knew that. Ecology did not iden- nance, but maintaining that dominance is not
tify that occurrence for us it simply gave us a always easy. Nature likes diversity and, without
principle with which to define it. Now let us use our interference, what was once a perfect (in our
that principle and others to help us make good eyes) turfgrass system will quickly turn into a
decisions concerning our turf. struggle for dominance among the species present
The law of the minimum was replaced by the law and among the current species and invaders.
of limiting factors (Blackman, 1905). The law of Ecologists call that succession and it is typical of
limiting factors states that population growth is nature. Our job is to create an environment where
limited by too little of a resource but is also limited succession does not occur. We want our community
by too much of a resource. In turfgrass management to remain at a particular stage of succession in
nothing could be truer; water is good but too much perpetuity. However, we want to do that with as
water is not good, nitrogen is good but too much little disruption as possible.
nitrogen is not good. Warm temperatures are nice A turfgrass that is adapted to the climate where
but hot temperatures are not good; cool is nice but it is grown and mowed on a regular basis is likely
cold is not. With rare exception, in everything that to proliferate and may be the dominant species in
we do, there is a low limit and a high limit between a community (Box 11.1). Therefore, growing
which we must always operate to achieve our turfgrass where it is naturally adapted and mow-
greatest success. ing it regularly to remove most of its competitors
allows the grass to flourish. In that case, it is rea-
sonably easy to manage in the monoculture that
The ecological community
we prefer. If turfgrass is not adapted to the cli-
An ecological community is a collection of popu- mate where we want to grow it management
lations that interact with each other. Communities becomes complex. In that case, managing our
are not only defined by the populations that they monoculture becomes very expensive, both in
contain but often by physical characteristics or, economic terms and in environmental terms. We
in our case, management levels. A forest and a have to supply what the turfgrass needs even
meadow would normally be considered to be two though it is not naturally available at our location.
different communities. A tennis court would be In most cases, the missing component is water.
considered as in a different community from the Because of that, our industry has become unde-
soccer pitch beside it, which would be in a differ- servedly notorious for luxury water use. If society
ent community from the park surrounding it. does not want us to grow turf where it is not
Within this complex, there may also be commu- naturally adapted, so be it, but if we are asked to
nities of bedding plants that require care too. do it, we can. All we have to do is to synthetically
The idea of a community including grasses, supply the natural components that are missing.
weeds, birds, insects, fungi, bacteria, humans The missing component could be water, or it
and other organisms gives considerable depth to could be a temperature conducive to growth, a
the concept of interspecific competition. growing medium, a nutrient or a collection of
Ecologically speaking, a human playing tennis on factors. We don’t always apply synthetic
a grass court is not using the turf for sporting components; we usually supply natural
purposes: the human is competing with the turf components synthetically (e.g. irrigation and
for space. If the human travels across the court nitrate fertilizer).
repeatedly in the same place for a long enough There are usually only a few species that prolifer-
period of time, that human may not win the ate in a community but many other species that
match with her opponent but she will win the occur in small numbers. For instance, a forest
match with the turf for space. She will destroy could contain 24 different species of trees but two
the turf and someone will have to fix it. of those populations (a population is only one spe-
Within a community, there is a struggle among cies) might make up 44% of the total number of
species for dominance. The largest population is trees (Smith and Smith, 2001). Although the trees

166 Chapter 11
 Box 11.1. Mowing is a means we use to limit the number of species that can live in a
community.
When we mow, we affect a turfgrass community by would probably call it practically sterile. We would
preventing the growth of most weeds. Trees and both be right.
shrubs are automatically eliminated from the site. If we were to allow a turfgrass site to naturalize, the
Most grasses and many broadleaved weeds repro- turfgrass, if it was chosen properly for the site, would
duce by producing stalks that rise above the existing probably be one of the most abundant species pro-
canopy before producing seed. If those plants can viding that the natural environment was best suited
only reproduce by seed, they are highly discouraged for grassland. However, it might not dominate a
by mowing. We also eliminate any plant that cannot grassland community and most certainly would not
produce leaves close to the soil surface and any dominate if the area was one where trees proliferate.
herbaceous plant that, like trees and shrubs, main- In contrast, if we were to allow the site to naturalize
tains a leading growth structure (an apical meristem) but continued mowing on a regular basis, the turf-
above the soil surface. The lower we mow the more grass would probably be able to maintain its domi-
plant species we eliminate. In doing so, we also nance. Turfgrasses are extremely competitive plants.
eliminate those animals and microbes that feed on If the turfgrass could not dominate in a natural but
those plants or have symbiotic relationships with mowed environment, then it is not adapted to the site
them. We call this community perfect; a naturalist and we are probably the only thing keeping it alive.

make the forest there are hundreds of other species how humans can exist within those ecosystems
also living in that community. Each of those species with the least environmental impact. Our goals are
has found a niche, a resource(s) in a conducive similar yet different. We hope to manage turfgrass
microenvironment for which they are uniquely systems to meet the expectations and demands of
adapted. A niche allows a population to live and our customers in the most natural way and with
successfully compete within a community. Each the least environmental impact possible. Depending
population within the community has a niche that on our customers’ expectations, we may have to
allows it to be a part of the community. If a species use synthetic means to achieve satisfactory results.
cannot find a niche within a community it cannot However, our objectives are always to use the least
compete with other populations in the community synthetic inputs possible and to affect our sur-
and therefore cannot survive. It would be nice if we roundings little if at all. In order to do that, we
could eliminate the niche that allows turfgrass must study the ecology of our ecosystem and for
insect and disease pathogens to live within a turf- that purpose we should be thinking small. Our
grass community. However, the turfgrass is one of ecosystem is the turfgrass community and its envi-
the resources that enable the existence of a niche ronment. Our management should affect that eco-
for turfgrass pathogens. In most cases, we would system and no other. That may seem like an
have to eliminate the turfgrass in order to destroy impossible task and it probably is, but that is the
the niche for its pathogens. ideal that we should strive for.

An ecosystem Ecosystem stability

A system is a complex in which the parts interact A natural ecosystem is in constant flux. It is in a
to produce the behavior of the whole. Ecosystems constant mode of succession. Succession can be
can be very large or very small. Ecosystems tend to cyclic, occurring over periods that we can study
take on the size of their environment. An ecosystem and predict, but it is normally a very slow process
is a combination of a biotic community of species involving changes in population dominance within
and its abiotic environment. Ecologists tend to the ecosystem’s community over extended peri-
study large ecosystems and use energy-dependent ods. Over such extended periods an ecosystem’s
models to define the most efficient way to conserve environment also changes, causing flux in the
those systems in their natural state or to determine ecosystem.

Ecology of Turfgrass Management 167

 Cyclic changes in predator/prey relationships can 50 Carrying
also cause flux. This flux is similar to changes capacity
caused by season and is called oscillation. As a 45
population builds, its predator numbers also build
as the predator’s resource (the population) increases. 40
Eventually, predator numbers increase enough to
cause a decline in the population. As the popula- 35
tion declines, predator numbers also decline, thus
completing the cycle which, assuming that the
30
population has sufficient resources, begins again.
Disturbances also cause ecosystem flux.

Time
25
Natural disturbances that cause changes to an
ecosystem could include fire, flooding, wind dam-
20
age, ice damage or drought. A relatively stable
th
ecosystem will return to its pre-stress level follow- grow
15 tion
ing a disturbance. If the disturbance is severe, how- ula
Pop
ever, recovery may take considerable time. In the
turfgrass world, we would renovate following a 10
severe natural disturbance. Some man-made distur-
bances can also be severe. Mining, logging, cultiva- 5
tion and the ever-present expanding presence of the
human population cause serious disturbances. 0
For turf-management purposes, disturbances are 0 10 20 30 40 50 60 70
minor but always present, often at varying levels of Density (shoots per unit area)
severity. We usually have traffic damage that dis-
Fig. 11.2. Because turfgrass populations fluctuate in
rupts the system. Cultivation practices regularly
response to many factors, a turfgrass population never
affect the quality of the turfgrass population and achieves its carrying capacity for more than a very
the ecosystem in general. There are seasonal factors short time. Instead, turfgrass numbers fluctuate around
to consider, and damage caused by invading weeds, carrying capacity.
pathogens and pests. Management mistakes are
particularly common causes of flux in the turfgrass
unit of time. In other words, it is a system that can
population. Because of all these potential fluctua-
be expected to yield a certain average amount of
tions, it is difficult to manage turfgrass at the den-
product over a given period of time, usually a year,
sity of its carrying capacity with any consistency.
and maintain that yield perpetually. The concept of
The population always fluctuates around carrying
sustainability in crop production is based on har-
capacity, sometimes even exceeding it temporarily
vest. In a turfgrass system, sustainability is based
(Fig. 11.2). Part of your job is to manage carrying
on aesthetic value, functional use or both.
capacity and minimize disturbances. We will dis-
A more socially oriented definition of sustaina-
cuss that later.
bility may include a clause that the sustainable
system should not interfere with other sustainable
Sustainability systems and should not have a detrimental effect
on its ecosystem. Those are also terms that we
Sustainability has become a favorite word in politi-
should strive to meet effectively. In doing so, we
cal and social circles. Consequently, it tends to take
can maintain better turf, or at least acceptable turf
on multiple meanings which depend on the person
with less social and political pressure.
using the term. The concept of sustainability is
vague, but the simple definition of the term is not
vague and it is most certainly something that we
Landscape ecology
should strive for. A sustainable system is one that
perpetuates under proper management. In crop Landscape ecology is a relatively new branch of
production, a sustainable system is one whose yield ecology. It can be described as natural ecology that
per unit of time equals its expected production per recognizes human activity as a factor which

168 Chapter 11
combines with natural disturbances to determine let us study the intraspecific competition that
the spatial patterns that constitute a landscape. occurs within a population.
Obviously, managed turfgrass surfaces are part of Intraspecific competition explains many things
landscape ecology. Because of our ability to man- that we know to be true of turfgrass. For instance,
age turfgrass ecosystems to near perfection, our mowing a turfgrass lower without exceeding the
industry – lawns and golf courses especially – has limit of its adaptation causes turf density to
come under scrutiny. In some instances, poor alter- increase. Density can not increase beyond carrying
natives to lawns have been suggested as landscapes capacity; by definition, carrying capacity is the
that use water and nutrients more conservatively. maximum density that can be supported by the
That may be true in climates where turfgrass is ecosystem. So by reducing the mowing height of
marginally adapted, but it is usually not true where the turf, we were able to increase the carrying
turfgrass is adapted (Erickson et al., 2008). capacity of the ecosystem. How can that be?
Turfgrass has a positive influence in residential When mowing height is lowered biomass is
applications (Beard, 1973). As the study of land- reduced. Biomass is the weight of plant material in
scape ecology matures, the advantages of turfgrass a given area. Biomass and density have an inverse
in low-input landscapes will probably be recog- relationship within available resources (Fig. 11.3).
nized. Whether or not that is true, landscape ecol- Consequently, if we lower mowing height, thereby
ogy may provide us with additional information reducing the biomass of a mature turfgrass popula-
that helps us to manage turf more conservatively tion, fewer resources are needed to maintain bio-
without loss of quality. mass and the surplus resources can be used to
increase density. As a mature turf is mowed lower,
reducing the biomass to a point below the full use
Intraspecific competition
of the resources available, the plants will allo-
Populations not only have to compete with other cate the excess resources to producing more tillers
populations for space and resources, the individu- and/or daughter plants, causing an increase in den-
als of a population have to compete with each sity. Eventually, enough tillers and/or daughter
other. As the population reaches carrying capacity, plants are produced that resources again become
individuals within it compete with each other for limiting. At that point, intraspecific competition
resources. We call that intraspecific competition. occurs. The new plants have to compete with the
Intraspecific competition occurs among individuals other new plants and with their parents for space
of a species and cultivar and among cultivars in and resources. Some new plants will not survive
blends. A blend is a combination of cultivars of the and a small number of less competitive parents may
same species. A mixture is a combination of turf- die. Soon the population will become fairly stable
grass species. Competition among cultivars of the around the new carrying capacity.
same species is intraspecific and competition among Intraspecific competition is a means of competi-
species is interspecific. We will consider interspe- tive selection. In all likelihood, the strongest plants
cific competition in Chapters 12 and 13. For now, for the environment will survive. Intraspecific

(a) High biomass (b) High density

Density Density

Biomass Biomass

Total resources Total resources

Fig. 11.3. As the plant biomass and density of a mature turfgrass population have a relationship within the available
resources, when biomass is reduced, a surplus of resources is available to fuel an increase in density. The pie charts
represent the resources available to the population, and the segments within them represent the portion of those resources
allocated to biomass and to density. (a) When biomass is high, density is low. (b) When biomass is low, density is high.

Ecology of Turfgrass Management 169

competition occurs among plants of the same spe- out-compete the lesser adapted ones. Overall growth
cies. The plants share a common niche. Consequently, in biomass would continue, for the most part, until
competition is normally severe. In a seeded popula- the growth of older plants slowed and more
tion, where minor genetic variation exists, selection resources were available for the birth of new plants.
will occur. However, in a vegetatively propagated As a population matures in nature, biomass is
cultivar, the plants are genetically identical so elimi- added at a rate of approximately three parts bio-
nation happens at random and may occur as a mass at the expense of two parts density, i.e.
result of interference or damage from an outside increasing biomass has a negative effect on density
source. Because turfgrass plants are so highly com- (Fig. 11.4). The relationship has come to be known
petitive with each other, increases in density usually as the −3/2 power law of self thinning or the −1.5
result in stands of immature plants struggling with self thinning law (Yoda et al., 1963; White and
each other for available resources. Immature plants Harper, 1970). Mown turfgrass, however, does not
are difficult to manage because they are more sus- respond as a natural plant and its self-thinning line
ceptible to unfavorable changes in environment, is believed to have a slope of −0.5 (Lush, 1990).
they are more easily damaged and they are more
susceptible to pest problems.
11.3 Turfgrass Ecology
Resource allocations between plant biomass and
density are ecologically described by the law of self Turfgrass ecology has never really been defined,
thinning. The density of a population will continue but it could be explained as the use of natural
to increase until it reaches the carrying capacity of means to maintain a managed turfgrass system to a
its ecosystem. At carrying capacity, no new indi- particular level of human expectation with the
viduals can be added unless an individual dies. In a fewest inputs and the least environmental impact.
natural situation, once the population reached the The idea behind turfgrass ecology is to allow
carrying capacity, intraspecific competition would nature to work for you. The only way to do that is
result in some individuals being eliminated as others to understand the natural boundaries of your eco-
increased in biomass. Selection would occur as the system and to work within them. The law of limit-
plants best adapted to the environment were able to ing factors tells us that more than a minimum of a

50 50
Carrying
capacity
45 45

40 40

35 35

30 30
Biomass
Time

25 25

20 20

15 Natural 15
self-thinning line
10 10

5 Turfgrass 5
self-thinning line
0 0
0 10 20 30 40 50 60 70 80 90 100 110 120
Density (shoots per unit area)

Fig. 11.4. Natural self thinning occurs along a balance between biomass and density that results in a line with a
slope equal to −1.5. Mown turf however, self thins along a line with a slope of −0.5.

170 Chapter 11
resource is good but too much of a resource is not rhizomes or both. Their propensity to form daugh-
good. We need to know our boundaries and we ter plants is a characteristic of their biotype
need to know how those boundaries are likely to (Karcher et al., 2005a,b). Consequently, the most
change with changing environments and sudden aggressive of these biotypes has a propensity to
disturbances. dominate the stand. Although a dominant cultivar
Sexual reproduction results in genetic differentia- may increase its share of the stand with time, it is
tion among offspring of the same parents. It follows highly unlikely that it will eliminate the other culti-
then that any seeded cultivar, with the possible vars (Brede, 2004). Nor is it likely that a dominant
exception of Kentucky bluegrass, an apomictic individual plant would claim more that a small
(asexually reproducing) species, results in a stable portion of a mature turfgrass stand. That is to our
but wide selection of genetically different individu- advantage.
als. So a synthetic cultivar, as a seeded variety is Turfgrasses are blended or mixed for a reason.
called, is a group of very similar but slightly different Blends and mixes confer genetic diversity. Genetic
plants. It would seem that, under a given set of cir- diversity makes a stand stronger by increasing sta-
cumstances, some of these individuals would be bility. A genetically diverse stand of forage grasses,
more competitive than others. This appears to be legumes or turfgrass is potentially better able to
true of turfgrass, because seeded populations, espe- resist disease, insect damage and even weed
cially in golf course greens where competition is encroachment (Chen et al., 1997; Picasso et al.,
extremely fierce, become patchy, and what appear to 2008). That said, however, weak cultivars or spe-
be vegetative offspring of the same parent form iden- cies do little to improve blends and mixtures
tical patches that are easy to differentiate. If indi- (Dernoeden et al., 1998). Each cultivar or species
vidual selection occurs within a population, then in the blend or mixture must be reasonably well
cultivar selection must occur within cultivar blends adapted to the site or the genetic diversity that it
(Lickfeldt et al., 2002a). In a cultivar blend, compe- adds to the stand has little effect. If we were to quit
tition is occurring on two levels, one among indi- managing a turfgrass site it would quickly convert
viduals and a second among cultivars. Fortunately, to a natural stand and within 2 to 3 years it would
however, cultivars or biotypes are so similar that be a mix of many plant species. That ecosystem
unless they differ by a particularly important charac- would be quite stable, meaning that it would resist
teristic, such as mowing height, or particularly change. Each of many plant species would have
widely by drought tolerance, they can be managed found a niche in the community and established a
in the same way (McGuan et al., 2004). population. Other organisms, including microor-
The competitive advantages of a cultivar are ganisms and insects, would also have diversified
probably more important than management prac- and would be in competition for resources and
tices for determining cultivar dominance (Lickfeldt space within the community. Many pathogens
et al., 2002b). Cultivar dominance or species domi- would exist, but in all likelihood each would be
nance in a seed mixture may be determined during specific to a different plant species. Consequently,
establishment. A cultivar that germinates more if one plant species contracted disease, the proxim-
quickly than the others, or grows more vigorously ity of like plants, separated by non-susceptible
as a seedling, has a distinct advantage. That par- plant species, would slow the spread of disease to
ticular cultivar may not be the best suited to the the point that little damage would be likely to
environment but it establishes more rapidly and occur and it would hardly be noticed in the com-
has a competitive advantage over the smaller seed- munity. Although we cannot reach that extent of
lings of the other cultivars. If it reaches maturity diversity in a turfgrass monoculture, we can take a
first, it shades its competitors and easily wins the lesson from nature and diversify as much as possi-
battle for space. Once the blend is established the ble. Hence, blends and mixes are usually
best performing cultivar for that particular envi- advantageous.
ronment would be expected to dominate. However, Ecological study reminds us that stability is a
that may not be the case. Again, the most aggres- natural characteristic that a turfgrass monoculture
sive cultivar has the advantage. Spreading grasses, is lacking. Therefore our systems are prone to rapid
those that grow by stems, differ in their ability to fluctuation. Rapid fluctuations are detrimental to
spread rapidly. Most warm-season (C4) grasses and the health of a community and to the sustainability
many cool-season (C3) grasses spread by stolons, of an ecosystem. Nature teaches us that stability is

Ecology of Turfgrass Management 171

preferred in a biotic community. Consequently, it is When you add a limiting resource to a turfgrass
beneficial for us to maintain stability in all things system you increase carrying capacity. The law of
turf related and we have the tools to do so. We self thinning, however, does not change, although
have integrated pest management plans that will be the self-thinning line now begins at a higher maxi-
presented in the next two chapters, and we plan mum density (Fig. 11.5). Turfgrass management
and schedule management such as mowing and objectives nearly always include achieving the
fertilization to reduce rapid changes in growth and maximum turf density that the ecosystem can sus-
promote stability. Changes in season can have a tain. Once you have achieved the maximum carry-
profound effect on plant communities (Lush, ing capacity attainable, you need to determine the
1988a). We need to pay more attention to what is minimum acceptable biomass.
happening around us and be flexible to changes in Ecological doctrine tells us that the greatest
weather and other environmental conditions. We biomass is achieved at relatively low density
can predict how plants will respond to changes in (Kandel et al., 2004). Relatively low density also
season (Koski et al., 1988; Xiong et al., 2007). We encourages seed production. Plants are more
can formulate management plans that are easily likely to reach full maturity if intraspecific compe-
adjusted to current conditions. We have discussed tition is minimal (Bednarz et al., 2006). The rela-
how to do that with fertilizer, irrigation and cul- tionship between turfgrass biomass and density is
tural practices, and we have determined what governed by self thinning (Lush, 1990). However,
inputs are required to positively affect photosyn- we have a great deal of control over biomass
thesis, respiration and transpiration. There is no through mowing (Lush and Rogers, 1992).
prescription for turfgrass management. You have Therefore, we have a great deal of control over
to accept what nature requires from you and density as well. Within a turfgrass species’ range
remain alert to what nature offers you. of adaptation we can choose its mowing height, in
Although it is difficult to manage a turfgrass this way controlling its biomass and determining
system that nature is happy to destroy, we do have its density. It would seem advantageous then to
some positive things working for us. For one thing, lower mowing height to the lowest acceptable
we can adjust carrying capacity and we can also height, thereby increasing density to the maxi-
affect density and biomass. Anything that we can mum. However, mowing low is rarely a wise
do to encourage photosynthesis, respiration and choice because as biomass decreases and density
transpiration increases carrying capacity. increases, maturity declines and the stand becomes
Consequently, we have already discussed many more easily damaged or disrupted (Spak et al.,
methods that we can use to affect carrying capacity. 1993) (Fig. 11.6). Turfgrass mown at low mowing
We need light, water and carbon dioxide to pro- heights is always in a juvenile state.
mote photosynthesis. Soil oxygen promotes respi- Mature plants have a competitive advantage
ration. Sufficient irrigation is used to satisfy over immature plants (Kendrick and Danneberger,
transpiration needs and we can add nutrients that 2002). Immature plants are easily damaged.
supply the elements needed for vegetative and Therefore, populations of immature plants are
reproductive growth. In order to increase carrying unstable. Rapid fluctuations occur in the turf-
capacity we simply have to determine which of the grass population and the overall health of the
physiological processes is limiting further growth system is difficult to sustain when the population
of tillers (vegetative growth) and daughter plants consists of immature plants. A particularly knowl-
(reproductive growth), and which factors are edgeable turfgrass manager is required to grow
needed to encourage it. We can do that through grass at low mowing heights. In addition, the
knowledge, experience, or trial and error. Of system requires an exceptionally high amount of
course, there comes a time when inputs become labor, energy and resource inputs. Low-mown
excessive. The law of limiting factors states that systems such as putting and bowling greens are
population growth is limited not only by too little sometimes required. Otherwise low mowing
of a resource but also by too much. Therefore, we should be avoided. Low mowing is not economi-
must always be looking for that upper limit of cally or environmentally sound. If you want to
nitrogen, water and other resources that causes a make your job easier and your turf system more
decline in carrying capacity when an increase is sustainable, mow as high as your end-user
expected. expectations will allow.

172 Chapter 11
 25 50

20 Old carrying New carrying 45

capacity capacity
15 40

10 35

5 30

Biomass
Additional resources
Time

0 increase carrying capacity 25

5 20

10 15

15 10

20 Turfgrass 5
self-thinning line
25 0
0 10 20 30 40 50 60 70 80 90 100 110 120
Density (shoots per unit area)

Fig. 11.5. Carrying capacity increases and density increases when a limiting resource is added to the turfgrass
system. With an increase in carrying capacity, the self-thinning line moves to a new maximum density but the
relationship between biomass and density does not change.

25 50

20 45

15 40
Matu

Biom

10 35
rity

ass

5 30 Biomass
Time

0 25
Den
sity

5 20

10 15

15 New 10
carrying
20 capacity 5
Turfgrass
self-thinning line
25 0
0 10 20 30 40 50 60 70 80 90 100 110 120
Density (shoots per unit area)

Fig. 11.6. Mowing low to improve density reduces biomass and results in a turfgrass stand that is easily damaged or
disrupted. Compare with Fig. 11.5, which shows a stand with the same increased carrying capacity (resulting from the
addition of a limiting resource).

Ecology of Turfgrass Management 173

 11.4 Chapter Summary competitive ability. We try to maintain stability and
consistency so that the turfgrass can flourish and
Ecology is the science of natural systems. It can
maintain its dominance in the community. One
instruct us on what to expect from our turfgrass
practice that encourages stability is to maintain the
under differing environments and management. By
turfgrass at the oldest maturity possible. We can
following ecological principles as closely as possi-
affect turfgrass maturity by adjusting mowing
ble, we can take advantage of natural phenomena
height higher to favor biomass over density.
and reduce our economic and environmental
Although density is normally a condition that we
impact.
encourage, we can manage more economically with
According to ecology, a turfgrass of the same
the least environmental impact and the highest
species living in a stand at a particular site consti-
stability when biomass is favored. Therefore, ecol-
tutes a population. If it is a mixed stand of turf-
ogy encourages us to maintain at the highest mow-
grass species, each species constitutes a population
ing height that is acceptable to our end user and
and the populations compete for space and
our purpose. By taking advantage of natural phe-
resources. The turfgrass and other organisms at the
nomena, we can promote sustainability in our
site and those that may visit the site make a com-
turfgrass system, thus reducing the cost of our
munity. Each population is a part of the commu-
labor and minimizing our effect on surrounding
nity because it is able to occupy a particular space
ecosystems.
and can compete for a particular set of resources.
The combination of resources and microenviron-
ment that allows a species to compete within the Suggested Reading
community is called a niche. The combination of
Barbour, M.G., Burk, J.H., Pitts, W.D., Gilliam, F.S. and
the biotic community and abiotic environment is
Schwartz, M.W. (1999) Terrestrial Plant Ecology, 3rd
called an ecosystem. We manage the turfgrass and edn. Benjamin Cummings, Menlo Park, California.
its competitors in such a way as to make the Danneberger, T.K. (1993) Turfgrass Ecology and Man-
turfgrass(es) the dominant population(s) in the agement. Franzak & Foster, Cleveland, Ohio.
ecosystem. In many respects, we can also manage Park, D.M., Cisar, J.L., Erickson, J.E. and Snyder, G.H.
the environment. (2008) Influence of landscape and percolation on P
To promote the turf as the dominant species, we and K losses over four years. In: Nett, M.T., Carroll,
provide resources that help it compete with other M.J., Horgan, B.P. and Petrovic, A.M. (eds) The Fate
populations. We provide light, water and carbon of Nutrients and Pesticides in the Urban Environment.
ACS Symposium Series 997, American Chemical
dioxide to promote photosynthesis, soil oxygen for
Society, Washington, D.C., pp. 107–132.
respiration and irrigation for transpiration. We
Sachs, P.D. and Luff, R.T. (2002) Ecological Golf Course
provide nutrients for consistent growth and cul- Management. Ann Arbor Press, Chelsea, Michigan.
tural management that affects the turf positively Schulze, E.D., Beck, E. and Müller-Hohenstein, K. (2002)
and/or its competitors negatively. Plant Ecology. Springer-Berlin, Heidelberg.
Rapid fluctuations caused by disturbances are Smith, R.L. and Smith, T.M. (2001) Ecology and Field
detrimental to the turfgrass because they affect its Biology, 6th edn. Benjamin Cummings, New York.

174 Chapter 11
 Managing Competition Among
12 Plant Species

Key Terms
A habitat is the place where an organism lives. In this case, it refers to the place occupied by a turfgrass com-
munity. The habitat is defined by its location and by its environment.
Habitat management is the use of management practices that alter the environment of a turfgrass ecosystem to
favor turfgrass persistence and discourage its competitors and antagonists.
Spot spraying is the use of herbicide by locating and spraying individual weeds in the turfgrass canopy. This is
only feasible when the weed population is low.
Broadcast spraying is the typical practice of spraying a pesticide over an entire area in an attempt to remove all
of the weeds from the site without having to locate them.
Allelopathy is the production by plants of toxic chemicals that selectively inhibit the growth or reproduction of other
plants.
A competitive plant strategy is the sum of genetically fixed physiological and morphological adaptations required
to conquer a habitat and to persist with optimal use of resources.

12.1 Manage Your Sites to Favor

with no pesticide use. If we manage our systems to
Turfgrass and Discourage Other Plant
encourage turfgrasses and discourage other plant
Species
species we can potentially learn to manage turf
In many parts of the world, the use of herbicide with minimal weed encroachment with no herbi-
applications for managing turfgrass nearly weed cide use. For the present, we can most certainly use
free is discouraged. Although herbicide use for our knowledge of ecology to develop better plans
producing food crops is tolerated, the use of herbi- for permanent weed reductions.
cides for managing urban crops such as turf that If we were to choose exactly the right species or
are primarily produced for aesthetic value is dis- combination of species for a particular turfgrass
couraged socially and sometimes governmentally. site and use all of the techniques at our disposal to
Although the pesticides that we use are very much reduce weed pressure and encourage turfgrass
like and sometimes exactly the same as the medi- growth, we could probably meet the expectations
cines that we take, they are usually deemed more for a home lawn, athletic field, or perhaps even a
dangerous. Their use is also considered, rightfully golf course fairway, by using mechanical weed
so, to be less important and more likely to have a removal. However, we would also want to balance
detrimental effect on natural environments. In the economic cost of herbicide application with the
many, perhaps most, instances pesticides applied to economic cost of the hand labor needed for
turfgrass are considered undesirable. We have to mechanical removal. That would be more difficult
accept that and limit or eliminate pesticide use. on a large property like a park or a golf course, but
Under pesticide restrictions, perfect or near-perfect might be possible on a home lawn or athletic field.
turfgrass is not possible at our current level of Ecological management is all about using natural
knowledge. However, we are good at what we do factors to manage turfgrass at a particular level of
and we can achieve near-perfect turf with limited human expectations. As you learned in Chapter 11,
pesticide use and, in most cases, acceptable turf natural communities and human expectations are

©CAB International 2011. Turfgrass Physiology and Ecology (G. Bell) 175
often not compatible. However, we can learn to use differentially adapted to various sites along the
particular natural factors to encourage turf domi- Italian peninsula and Italy’s major islands
nance within a natural community. For reasons (Annicchiarico et al., 2006). In a study in Canada,
that you will learn in Chapter 13, especially con- the proportion of each species in mixtures of the
cerning predator/prey relationships, we will prob- same four species (Kentucky bluegrass, red fescue,
ably not be able to manage today’s turfgrass tall fescue and perennial ryegrass), including mul-
monocultures on a sustainable basis without the tiple cultivars of each, changed in proportion
use of fungicides and insecticides. For that reason, between that seeded in 1988 and that in the
we may have to lower our expectations. However, mature stands 4 years later (Hsiang et al., 1997).
it is quite likely that we can learn to manage desir- In this study, perennial ryegrass was the dominant
able turf at high expectations with little or no use species in mature stands, possibly because of its
of pesticide. rapid germination and superior seedling vigor
compared with the other species. Kentucky blue-
grass and creeping red fescue were competitive
12.2 Competition Among Turfgrass
with the perennial ryegrass, but the proportion of
Species
tall fescue in stands was much smaller than that
Competition among different turfgrass species which had been seeded. Tall fescue has better heat
within a community is by definition a form of tolerance but poor cold tolerance compared with
interspecific competition. In many instances, how- the other three species and might not have been
ever, it does not matter to us which species survives able to compete well for that reason. Three of the
or which dominates the stand as long as the stand four species and many of their cultivars appeared
meets our expectations for color, density, uniform- to be relatively well adapted to the site and able to
ity and the other factors that our customer deems survive and compete for resources. Such a mixed
important. In fact, species mixtures improve the stand would normally be able to reduce the com-
biological diversity in a community. Diversity was petitive ability of weed species better than a single
introduced in Chapter 11 as a positive factor for stand of one individual turfgrass species. It is also
the stability and sustainability of an ecosystem. unlikely that any of these turfgrass species would
Consequently, in the ecological management of a be completely eradicated by natural competition in
turfgrass system, cultivar blends and species mix- any of these mixtures. They would always be
tures are preferred. The blends that were discussed present to some extent. Although perennial rye-
in Chapter 11 and the mixtures to be discussed in grass was dominant at the Canadian site, Kentucky
this chapter are more naturally resistant to distur- bluegrass dominated perennial ryegrass in a study
bances than single-cultivar or single-species in Pennsylvania (Brede and Duich, 1986). In
monocultures. warmer regions of temperate zones, tall fescue is
the most competitive of cool-season (C3) turfgrass
species and tends to dominate mixed stands
Interspecific turfgrass mixtures
(Bremer et al., 2006). In species mixes in northern
Turfgrass mixtures and blends are ecologically Missouri, tall fescue dominated Kentucky blue-
favored over single species and cultivar monocul- grass but not perennial ryegrass (Dunn et al.,
tures. However, if the mixed community is to have 2002). In a slightly warmer climate, such as that of
an ecological advantage compared with other plant southern Missouri, tall fescue would probably
species all component grasses must be well adapted dominate perennial ryegrass as well, but both spe-
to the site. Each turfgrass must be able to persist cies would persist creating a stronger stand than
and sustain itself in competition with other plant either would alone if they were both reasonably
species. In a turfgrass system, the most important well adapted to the environmental conditions.
plant competitor of each turfgrass should be Cool-season grasses are often overseeded into
another turfgrass. In such an ecosystem weed warm-season (C4) grasses in the USA to provide a
encroachment is discouraged. green cover in winter. The two species compete with
Research has indicated that cultivars of Kentucky each other during spring and fall but they dominate
bluegrass (Poa pratensis), creeping red fescue during the seasons, summer and winter, for which
(Festuca rubra), tall fescue (Festuca arundinacea) they are best adapted. The most difficult portion of
and perennial ryegrass (Lolium perenne) were the overseeding process is the transition from winter

176 Chapter 12
cover (cool-season) to summer cover (warm-season). dominance. In the UK, in mixtures with perennial
In many instances, portions of the cool-season spe- ryegrass, creeping red fescue and colonial bentgrass
cies will persist in the warm-season species through (Agrostis capillaris) had poor cover compared with
much of the summer, demonstrating how difficult it perennial ryegrass when subjected to wear (Gore
is to eradicate even a poorly adapted species from a et al., 1979). However, both species did very well
community. In cooler regions, where the cool-season when wear was not present. A species’ ability to
species is poorly adapted to the summer heat but resist wear or some other human influence affects
also somewhat tolerant, it may be extremely difficult the dominance of a turfgrass community (Sorochan
to remove with cultural practices alone during sum- et al., 2001; Samaranayake et al., 2008). Mowing
mers when temperature and humidity are lower height and nutrient input can also affect species
than normal (Horgan and Yelverton, 2001). dominance in a mixed stand (Carrow and Troll,
Persistent mixtures of cool- and warm-season turf- 1977). If we use management practices to assert
grasses have also been attempted where both species turfgrass dominance we will probably not elimi-
are reasonably well adapted. nate weed encroachment but we can reduce it
In Utah, research was attempted to mix fine fes- substantially.
cues (Festuca spp.) with buffalograss (Buchloe
dactyloides), a native species, for sustained, year-
12.3 Competition between turfgrass
long green color in low-maintenance situations
and other plants
(Johnson, 2003). Both types of grass are well
adapted to low maintenance and performed rea- In a turfgrass system, plants other than turfgrasses
sonably well in monoculture. The mixed stands are considered weeds. In fact, some types of turf-
also performed reasonably well, but the fine fescues grass in another turfgrass may be considered weeds
dominated the buffalograss to an extent that sum- in some locations under certain circumstances. Our
mer greenness was marginal and the experiment propensity for uniform single-species turf makes
was not deemed successful. In Missouri, a similar management, especially weed management, diffi-
project was attempted to promote year-long green- cult. Because all turfgrasses share a similar niche,
ness, primarily in athletic fields (Dunn et al., 1994). interspecific competition among them is fierce.
The use of common bermudagrass (Cynodon dac- However, for that same reason, they tend to
tylon) was tested in separate combination with exclude other plants from that particular niche.
Kentucky bluegrass, perennial ryegrass and two Nonetheless, weeds are persistent and opportunis-
fine fescues. Because athletic field use was to be tic colonizers that make the most of available
tested, wear was applied to the plots regularly, resources or disturbances in the turfgrass canopy
which reduced the effectiveness of the fine fescues (Fig. 12.1). Some weeds, dandelion (Taraxacum
in the fescue/bermudagrass mix. However, the officinale) for instance, are so good at utilizing
Kentucky bluegrass/bermudagrass mix and the resources and exist in a niche different enough
perennial ryegrass/bermudagrass mix persisted. from turfgrass that they can invade even the most
Bermudagrass would be expected to be the most dense turfgrass canopies and be highly competitive.
wear tolerant and to recover most rapidly from Most weeds, however, are excluded from a dense
injury compared with the other two species. None- turfgrass canopy and we can use management
theless, the bermudagrass was the weakest compo- practices to discourage others.
nent of the mixtures because it was the most poorly It is not possible to completely eradicate weeds in
adapted species for the region. The bermudagrass turfgrass. We can come very close to eradicating a
had poor quality in monoculture and was domi- particular weed species using herbicides, but we are
nated by both Kentucky bluegrass and perennial unlikely to ever remove it completely. Pesticides are
ryegrass in mixtures. In most cases, the species best a part of most turfgrass management plans although
adapted to the natural environment dominates the they should not be the first alternative employed;
community, but human activity also constitutes a they should be the last. We can come close, in some
portion of the turfgrass environment and human cases close enough, to customer expectations with-
use can also determine species dominance. out pesticide use. If we strive to minimize weed
Where two species are more or less equally populations through habitat management we may
adapted to the natural environment, it is human be able to meet customer expectations without
activity and management that determine species pesticides or to meet their expectations by spot

Managing Competition Among Plant Species 177

 than the collective biomass of the turfgrass
plants that can fit onto the surface of its root zone.
The tree requires more water and has a deep root
system to retrieve water when necessary. Within the
turfgrass root zone, the grass may be competitive
with the tree for water, but deeper in the soil, the
tree has the advantage. Consequently, as the soil
surface dries between rainfall events, turf must
tolerate the dry conditions while the tree can reach
deeper into the soil to find water. It seems that not
only are the shoots of the tree more competitive
than the shoots of the turf for light, but the roots
of the tree are also better competitors for water.
However, there is more to be considered.
Fig. 12.1. Hail damage is an unlikely but potential Tall fescue and Kentucky bluegrass provide
source of disturbance. This damage opens the serious competition for commercially produced
turfgrass canopy and makes a good seed bed for an eastern redbud (Cercis canadensis) and pecan
opportunistic weed. Photo by Craig Evans, Stillwater, (Carya illinoinensis) trees (Griffin et al., 2007).
Oklahoma. Two other tree species, silver maple (Acer sacchari-
num) and honeylocust (Gleditsia triacanthos),
spraying herbicides rather than broadcast spraying reduced the quality of creeping red fescue (Festuca
them. Spot spraying is environmentally friendly in rubra), roughstalk bluegrass (Poa trivialis) and
that it requires very little use of pesticide compared Kentucky bluegrass in spite of sufficient water and
with a broadcast spray that is applied over an entire nutrients (Whitcomb, 1972). Research indicates
area. Spot spraying is the act of locating weeds and that turfgrasses are able to compete with or out-
spraying them individually wherever they are found. compete trees for nitrogen and potassium uptake
Habitat management is the use of management (Nielsen and Wakefield, 1978; Tworkoski and
practices that alter the environment in a turfgrass Glenn, 2001). However, in the production manage-
habitat to favor the turf and exclude weeds and ment of peach (Prunus persica) trees, some
other competitors or antagonists. Environmental turfgrasses may provide desirable competition
factors both alone and in combination can affect because they provide erosion protection while out-
the population density of desirable and non- competing other damaging weeds (Tworkoski and
desirable species (Gaussoin and Branham, 1989). Glenn, 2001). Some weeds damage trees and some
Turfgrasses and trees are an example of plant trees damage turf, and vice versa, by producing
types that do not exist well together. They nega- chemicals that are toxic to other species.
tively affect each other wherever they are found, Many species are known to produce chemicals
usually to the eventual exclusion of the turf wher- that are selectively toxic to other species (Barbour
ever the competing trees are reasonably well et al., 1999). These chemicals are called allochemi-
adapted. Earlier in the text, you learned that car- cals, and they may be produced as root exudates or
bon, hydrogen and oxygen are the most important may be present throughout the plant. The produc-
basic nutrients for plants and that no life exists tion of chemicals by one plant that selectively
without photosynthesis. For photosynthesis to inhibit the growth or reproduction of another plant
occur, light and two of the basic nutrients, water is called allelopathy. Some cool-season turfgrasses,
and carbon dioxide, must be present. Turfgrass creeping red fescue, perennial ryegrass and
eventually loses the ecological competition with Kentucky bluegrass have been shown to produce
trees because the trees capture most of the light. root exudates that inhibit the growth of the flower-
Turfgrasses are very competitive with trees in most ing dogwood (Cornus florida) tree and forsythia
other respects though, and there is much that we (Forsythia intermedia) (Fales and Wakefield, 1981).
can learn from that competition. Bermudagrass and tall fescue are known to pro-
In addition to light, trees may be more competitive duce root exudates that affect the growth of young
for water than turfgrass (Fales and Wakefield, pecan trees (Smith et al., 2001). Conversely, pine
1981). The biomass of a tree is considerably greater (Pinus halepensis) needles have an allelopathic

178 Chapter 12
effect on bermudagrass and tall fescue (Nektarios The K- and r-strategy theory has been criticized
et al., 2005). Allelopathy can sometimes be useful because it does not take disturbances and stress
for the suppression of weeds in turfgrass. into account. For that reason, Grime (1974, 1977)
Researchers in Michigan found that mulched proposed a three-strategy model called the C-S-R
maple (Acer spp.) and oak (Quercus spp.) leaves model. The “C” stands for competitor and consti-
reduced dandelion counts in Kentucky bluegrass by tutes long-lived perennial plants that populate sites
at least 53% (Kowalewski et al., 2009). Researchers with adequate resources and have to endure little
in Iowa found that corn (Zea mays) gluten meal stress. The “S” stands for stress-tolerant plants and
inhibits root growth of many monocotyledonous refers to C-type plants but those that can tolerate
and dicotyledonous weed species (McDade and sites with few resources and severe environmental
Christians, 2001). Corn gluten meal is commer- stresses. The “R” plants are basically the same
cially available for pre-emergence treatment of short-lived rapidly reproducing plants identified as
some weeds, such as crabgrass (Digitaria spp.). r in the K- and r-strategy. The C-S-R model also
Future research is likely to produce more naturally proposes that many plants are not pure C, S or R
occurring allochemicals for use as alternatives to strategists, but employ combinations of two or of
synthetic herbicides. all three strategies.
A third model, the Resource Ratio Model, sug-
gested that all plant species are limited in their
The ecology of weed competition
distribution by resources (Tilman, 1988). Some of
Ecologists have attempted to define plant competi- these resources are scarce and the plants that are
tion by hypothesizing a number of different but successful in competing for these limiting resources
similar competitive plant strategies. They define the will prevail. These limiting resources are most often
term “strategy” as the sum of genetically fixed light and nutrients, specifically nitrogen. There is
physiological and morphological adaptations also a fourth model, the Facilitation–Tolerance–
required to conquer a habitat and to persist with Inhibition Model, proposed by Connell and Slatyer
optimal use of resources (Schulze et al., 2002). In (1977). Facilitation, tolerance and inhibition are
this case, a strategy is not a plan but a means of not plant strategies but pathways of succession.
adapting to the conditions of a habitat in such a Each requires that changes or disturbances occur in
manner that a plant can tolerate the environment the ecosystem before succession can proceed.
and exist and reproduce using the resources avail- All of these models have merit and are probably
able. The first published competitive plant strategy far more right than wrong. However, none of them
was that of Initial Floristic Competition based on a can fully predict the consequences of plant compe-
model by Egler (1954). This early model assumes tition. Environments are constantly changing or
that a large number of species are present at germi- evolving and disturbances occur without warning.
nation but that as the long-lived slowly developing Disturbances are rarely positive occurrences for a
species mature the short-lived species gradually die turfgrass manager, but they often are positive or
off. We can control the plants that live longer than neutral in a natural environment. Fires, for instance,
turf by mowing, so we could only hope that succes- rejuvenate grasslands or high winds topple trees in
sion was that simple. forests making way for new growth and creating
A more recent theory is that of K- and r-strategies diverse habitat that is particularly attractive to
(MacArthur and Wilson, 1967). The K classifica- many species. For us, however, change is something
tion refers to plants that are long lived, grow to be avoided unless it makes the grass greener or
slowly and accumulate large amounts of biomass. denser or the surface more consistent.
An r classification refers to plants that are short
lived and reproduce rapidly. A large amount of an
Managing weed competition
r-strategy plant’s energy goes into seed production
rather than biomass. Presumably, as the K-strategy Some turfgrass weeds are K-strategists and some
plants mature, they will eventually out-compete the are r-strategists. For basic identification, the ones
r-strategy plants, but that is not the case in turf- we refer to as perennials are K-strategists and the
grass management. In fact, the theory is a good one ones we call annuals are r-strategists. Turfgrasses
but nature is considerably more complicated than cannot completely replace either of these weed
these two simple categories would imply. types, so both weed types are consistently

Managing Competition Among Plant Species 179

antagonistic. Some of these weeds are “S” or
stress-resistant strategists. No matter where in
the world you are located, there are weeds that
take advantage of compacted soil, poor fertility,
shade, saturated soil or some other environmen-
tal or human-induced stress to colonize a niche
in a turfgrass community. Within that particular
habitat, they are more competitive than turf-
grass. Consequently, we have to change that
microenvironment at that specific location to
make it more conducive to turf and less condu-
cive to the stress-tolerant weed. Spraying the
weed with a herbicide is not the answer. That is
only a short-term fix. Changing the environment Fig. 12.2. This turfgrass damage was caused by drought.
to favor turf at the expense of the weed is a long- If the turf is not renovated quickly, weeds will soon take
term solution. over the damaged areas and, once established, they
will be difficult to remove.
The Resource Ratio Model suggests that some
resources are likely to limit the population growth
of our turf and that those resources are likely to be In order to do battle with a weed you must know
sunlight and nitrogen. That is nothing new to us. what it is. You must at least know whether it is a
We know that tree removal is often necessary for summer annual, a winter annual or a perennial.
adequate light with which to grow turfgrass and The more you know about the weed, its environ-
we are well aware that nitrogen is our most impor- mental preferences, limiting resources and life
tant nutrient. Our goal is to facilitate the growth of cycle, the better you will be able to minimize its
the turfgrass population by providing all of the encroachment. The first thing you need to do is
ingredients most conducive for the turfgrass and find a good weed identification site for your area
least conducive for its competitors. By doing that, on the Internet, locate a local weed expert and buy
we create an ecosystem that is intolerant of most a good weed identification and management book
invaders and we grow strong turfgrass plants that or books that cover the weeds most common in
inhibit the growth of invading populations. We your region. Be prepared to identify the weed and
attempt to predict disturbances and avoid them if study it so that you know how best your turf can
possible. When they occur, we should rapidly move compete with it.
to repair damaged turf before invasion occurs. Annual weeds live less than a year. They
Invasion of an open space is inevitable (Fig. 12.2). germinate, grow rapidly, reproduce (usually by
It is unlikely that spreading turfgrass plants will producing abundant seed) and then die before or
cover the damage before interloping weed seeds during their most stressful season. Biennial weeds
germinate and grow. Such are the basics of control- are similar to annual plants but they grow
ling weeds in turf. vegetatively during one year, then produce seed or
For many years, we have divided weeds into tubers and die the next year. Annual weeds are very
three categories to facilitate study and control. We opportunistic. Dormant seeds of annual plants
call them summer annuals, winter annuals and usually have to experience a period of cold tem-
perennials. So for turfgrass purposes we subdivide perature (summer annuals) or warm temperature
the r-strategy weeds into two categories based on (winter annuals) before they can break dormancy
their season of strongest vegetative growth. We and germinate. For that reason, summer annuals
recognize one category of K-strategy weeds, but we germinate in the spring and winter annuals
could consider two. It might be a good idea to germinate in the fall (Fig. 12.3). In addition to the
divide the perennial weeds into categories of cool- temperature change, there are normally other fac-
and warm-season plants to define their strongest tors typical for each species that must happen, or
seasons. However, based on what we have learned factors that must be present, before dormancy is
about ecology, our predecessors seemed to know broken and germination occurs. If all other factors
what they were doing. So let us continue using the are satisfied, all seeds must imbibe (take up) water
traditional groupings for study. before they can germinate.

180 Chapter 12
 Temperature
Jan 2009

Winter annual seeds deposited

Summer annual seeds germinate
Warm temperature period satisfied Jul 2009

Summer annual seeds deposited

Winter annual seeds germinate

Jan 2010
Cold temperature period satisfied

Winter annual seeds deposited

Summer annual seeds germinate
Warm temperature period satisfied Jul 2010

Summer annual seeds deposited

Winter annual seeds germinate

Jan 2011

Fig. 12.3. Winter annual weeds flower and seed in the spring. Once the dormant seeds have experienced the
extended warmth of summer, they are ready to germinate in the fall. Summer annuals flower and seed in the fall,
break dormancy by experiencing the extended cold temperatures of winter and are ready to germinate in the spring.

Dormant seeds of an annual species can remain species are present, the most pervasive annual spe-
in the soil for a single year or can be there for sev- cies will dominate the initial stand. If you prepared
eral years, depending on the species. Consequently, this site as a turfgrass seedbed and applied sufficient
seeds of an annual weed may accumulate in the soil seed, the turfgrass should dominate, but weeds,
for quite some time waiting for the right conditions mostly annual weeds, will also be present. Annual
to germinate (Fig. 12.4). We call that accumulation weeds are very opportunistic and tend to invade any
a soil bank. Annual species deposit great numbers open area in a turfgrass stand (Lush, 1988b). If you
of seeds in the soil bank each year. Perennial species prepare a seedbed and treat the area like a seedbed,
also add seeds to the soil bank, but most perennial weeds will germinate and, in fertile soil, will reach
species produce small numbers of seeds compared full cover rapidly. Weed seed does not have to be
with annual species so they have fewer seeds in the applied; weed seed is already present in the soil
bank. When conditions are right, the seeds bank. Consequently, if you intend to grow turf, the
germinate. turf must out-compete the weeds for space and for
If you use a cultivator of some kind to turn over resources. Weed competition can be highly detri-
or disrupt the soil at a particular location in the mental to the establishment of a turfgrass stand
spring or fall then add water regularly for a few (Maddox et al., 2007). Nonetheless, some weed
days, seeds begin to germinate. Although perennial encroachment is inevitable. The encroachment can

Managing Competition Among Plant Species 181

 annual bluegrass. Consequently, unless the site is
completely sterilized, annual bluegrass often occurs
in seedbeds meant for creeping bentgrass. Even
though creeping bentgrass is competitive with
annual bluegrass, some cultivars are more competi-
tive than others (Beard et al., 2001; Henry et al.,
2005). For that reason, if annual bluegrass compe-
tition is expected, a highly competitive cultivar of
creeping bentgrass should be seeded, which must
also be adapted to the environment and resistant to
the stresses common to the site. Choosing the right
species and cultivar or the right mix or blend can
save considerable maintenance following establish-
ment, regardless of the site or the grass that you are
Fig. 12.4. These winter annual weeds (henbit, Lamium working with.
amplexicaule) germinated in this warm-season A dense, mature turfgrass stand is weed resistant.
turfgrass (bermudagrass, Cynodon dactylon) as the The full canopy cover of dense turf shades the soil,
grass was nearing dormancy in the previous fall. The thus preventing some weed species from germinat-
weeds could not compete with the turf if the grass was ing (Arrieta et al., 2009). Management practices
growing vigorously. such as irrigation and aeration may also restrict
some weed species (Henry et al., 2009). It is possi-
be reduced by applying more turfgrass seed at high ble that some turfgrasses may produce allochemi-
rates (Busey, 2003). However, as you learned in cals that affect certain weeds. Allelopathy is
Chapter 11, intraspecific competition will occur as suspected between perennial ryegrass and zoysia-
the stand matures. The more seedlings in the stand, grass (Japanese lawngrass, Zoysia japonica) but as
the longer it will take to reach maturity, and the yet has not been found between turfgrasses and
more juvenile the plants, the easier they are to dam- weeds (Lickfeldt et al., 2001; Zuk and Fry, 2006).
age or infect. By increasing the seeding rate beyond Although many management practices are effective
the normal requirement you are trading one risk for for reducing weed competition, the most effective
another. The decision should depend on the amount practice is mowing, specifically mowing height.
of weed encroachment expected, and that is most In a study in the state of Washington, weed
affected by time of year. encroachment was reduced in perennial ryegrass
When you plan to establish or renovate a site, when mowing height was increased from 1.5 to 2.5
choosing a grass that germinates quickly and grows inches (3.8–5.1 cm) (Miltner et al., 2005). In
and spreads rapidly is desirable. In fact, overseed- Maryland, a tall fescue mowing height of 3.5
ing regularly with a rapidly germinating and vigor- inches (8.8 cm) had less smooth crabgrass (Digitaria
ously growing species such as perennial ryegrass ischaemum) encroachment than tall fescue mowed
can be an effective method for reducing the oppor- at 1.3 or 2.2 inches (3.2 or 5.5 cm) (Dernoeden
tunistic weed encroachment that often occurs on et al., 1993). In many instances, fertilizer also
athletic fields (Elford et al., 2008). However, rapid affects weed encroachment into turfgrass. A combi-
germination and establishment does not necessarily nation of fertilizer and high (4 inches versus 2
indicate the best adapted plants (Gardner and inches) mowing height reduced dandelion and
Taylor, 2002). The best adapted species are those white clover (Trifolium repens) encroachment into
that can dominate the community at maturity. a mixture of Kentucky bluegrass, perennial rye-
Creeping bentgrass (Agrostis stolonifera), the most grass and creeping red fescue by 75% (Calhoun
widely used species for golf course putting greens et al., 2005). After removing the dandelion and
in the USA, is often invaded by annual bluegrass white clover with herbicide, the fertilized plots also
(Poa annua). As both creeping bentgrass and had a slower re-infestation than the non-fertilized
annual bluegrass are cool-season grasses that can plots. As you learned in Chapter 11, a mature plant
grow at low mowing heights they are very competi- nearly always has a competitive advantage over a
tive with each other. The best periods for the estab- seedling. You also learned that low mowing does
lishment of creeping bentgrass are also the best for not allow turfgrasses to mature to their greatest

182 Chapter 12
potential. Consequently, high mowing still removes
weeds that cannot sustain mowing stress, increases
the maturity of the turf so that it is more competi-
tive and creates canopy cover and shade that fur-
ther discourage weed growth and, in some cases,
weed germination.
It is generally believed that nitrogen fertilization
favors grasses, especially turfgrasses, over most
other plants. However, because turfgrass does not
require high amounts of phosphorus fertilizer, high
phosphorus fertilization is likely to favor weeds
rather than turfgrass. Phosphorus is also the most
likely element to encourage eutrophication of sur-
face water, and over-fertilization with phosphorus Fig. 12.5. These perennial weeds (dallisgrass, Paspalum
should be avoided (Soldat and Petrovic, 2008). dilatatum) in a warm-season turfgrass (bermudagrass,
Many plants, including grassy weeds such as large Cynodon dactylon) have been allowed to mature and
spread. They are considerably more difficult to remove
crabgrass (Digitaria sanguinalis), crowfootgrass
now than they were when they were young.
(Dactyloctenium aegyptium) and annual bluegrass
respond well to high soil phosphorus levels
(Hoveland et al., 1975). Other plants, including weed emergence have been defined (Danneberger
large crabgrass and crowfootgrass, respond posi- and Vargas, 1984; Fidanza et al., 1996; Kaminski
tively to high soil potassium levels (Hoveland et al., and Dernoeden, 2007). All too often disruptive
1975). Low potassium has also been used to reduce cultivation is scheduled to coincide with the best
dandelion populations (Tilman et al., 1999). periods for weed germination and development or
Turfgrasses, in general, respond well to high potas- the worst periods for turfgrass recovery. Either sit-
sium levels, but care should be taken and adjust- uation is detrimental. Choosing the best possible
ments should be made if weeds appear to respond procedures to enhance turfgrass activity is the pri-
as well as your turf to a particular potassium ferti- mary focus of an effective long-term weed control
lization plan. There are so many combinations of plan. However, all plans have alternatives. It is not
mowing height and fertilizer that could be prac- necessary to conduct cultural procedures that open
ticed for weed reduction that it is difficult to a turfgrass canopy during times when weed germi-
choose a course of action. You should begin with nation is most likely. If the turfgrass is a spreading
the program most likely to encourage your turf- type and it has maturity, it can fill in rapidly during
grass species’ growth and density, and make small periods that are not quite as conducive to weed
adjustments by observation to discourage weed germination as others. Such periods may include
encroachment. Every situation is likely to require early spring for both warm- and cool-season
constant monitoring. There are so many different grasses, before the soil is warm enough for most
weeds and so many possible habitats that on-site summer annual weeds to germinate, or late fall for
observation and trial and error decision-making is cool-season grasses, after the primary germination
the best course of action. Weed reduction without period for winter annuals has passed. These peri-
herbicides takes considerable time and knowledge. ods are also better for seeding (Box 12.1). The high
That is why most people are reluctant to work with percentage of turfgrass seeds present in the seedbed
it when so many reliable herbicides are available. have a better competitive advantage with weed
An ecological plan, however, is a permanent seeds either before or after the period in which
improvement. A herbicide is only a temporary fix. nature has determined that most weed seeds should
Constant monitoring also gives you the opportu- germinate.
nity to spot spray or to mechanically remove weeds
when they are young and much easier to control
12.4 Biological Herbicides
(Fig. 12.5).
The timing of cultural practices can be very Biological herbicides could fit into the predator/
important for weed reduction. We know the life prey relationships that will be presented in
cycles of most weeds and, in some cases, models for Chapter 13. However, they are not entirely natural.

Managing Competition Among Plant Species 183

 Box 12.1. When are conditions best for seeding a cool-season turfgrass?
Let us assume that you want to establish a home Research demonstrates that annual bluegrass can
lawn with a cool-season (C3) grass and that you germinate at nearly any time during the fall and
can choose the time most conducive to do it. We spring, but that its germination is greatest in the early
normally recommend seeding a cool-season grass fall (Kaminski and Dernoeden, 2007). Therefore, if
during the fall rather than the spring because the we seed in the early fall and we have a seed bank
competition with summer annual weeds in the spring relatively high in annual bluegrass our grass will
can be sufficient to slow development of the experience severe competition from the weed. We
cool-season turfgrass until summer. During the need an alternative method.
summer, annual weeds have a particularly strong Instead of seeding the turfgrass in the early fall, we
advantage over cool-season turf. In the fall, there is can prepare the seedbed exactly as we normally
no competition with summer annuals, only with would, keep it moist just as we would to germinate
winter annuals and perennials, with which the turf- turfgrass and wait for the annual bluegrass to germi-
grass is highly competitive. Consequently, you nate. Because you have created and maintained a
would probably suppose that the best time to seed perfect seedbed, most of the weed seed present will
would be as soon as summer temperatures began germinate. After 3 to 4 weeks you kill the weeds with
to cool into the range conducive for cool-season a nonselective herbicide and seed your turf. By that
turfgrass growth, so that the grass would have as time, you have removed most of the weed competi-
much time as possible to mature before winter. tion and there is still plenty of time for the turfgrass to
Although that plan is intuitively correct, there is grow and develop before the winter.
actually a better time. If using a nonselective herbicide is not possible,
Let us assume that your major weed competition in hand removal of weeds is great, or tilling the weeds
the fall will be from annual bluegrass (Poa annua). under will work reasonably well.

Biological herbicides are formulations of living would not be used as biological control agents.
organisms that prey on weed species in a turfgrass Consequently, the use of a biological control agent
community. Biological weed control is a synthetic is considered by most to be preferable to the use of
application of these organisms to the community a synthetic herbicide. For that reason, recent
with the intention of causing harm to the plants research has begun to target the use of biological
they selectively use for food or, we could say, the herbicides as an environmentally sound means of
plants that they selectively infect. Biological herbi- weed control.
cides are usually disease-causing pathogens. The organisms in biological herbicides may be
Natural toxins and biological herbicides should present in the turfgrass community before an appli-
not be confused with one another. The allochemi- cation is made. However, they are rarely present in
cals already presented in Chapter 11 are toxins. sufficient numbers to infect and kill weeds until
However, we are more familiar with toxins that environmental conditions are exactly right for
potentially affect us. Plant products such as caf- infection. Actually, if the pathogen was not present
feine and nicotine are toxins that kill insects and, in before application, it is likely that the organisms
great enough concentrations, also kill us. Although applied will not survive long enough to cause any
caffeine is considered to be a safe food product, a real damage to the target weeds. The greatest
concentrated dose could be fatal. Many fruit seeds advantage of a biological herbicide is that it is a
contain cyanogenic glycosides that can be fatal in naturally occurring organism. That advantage is
high enough doses. Some people believe that natu- also a detriment. If the environment was conducive
rally occurring toxins make harmless pesticides to support pathogen numbers great enough to
when used as directed. Others believe that natural cause sufficient weed damage, the organisms would
toxins are tolerable but synthetic toxins are not. already be present in high numbers. Consequently,
Some think that we should ban them all. This is a the herbicide has to be applied when the environ-
politically charged issue, as is the use of disease- ment is sufficient to sustain it for a period long
causing agents. However, pathogens that attack enough for it to infect and kill the target weeds to
plants rarely attack humans and those that did which it was applied. For that reason, biological

184 Chapter 12
herbicides often fail. Synthetic herbicides rarely reason, we are happy to see them die after they kill
fail. Turfgrass managers are reluctant to spend the weeds.
money on a biological herbicide that could fail
when they have a synthetic herbicide in which they
12.5 Chapter Summary
are confident. They make that decision, not because
they are not concerned for our natural environment Ecological weed control is not easy. It is not some-
but because they have little choice from an eco- thing that you think about, develop a plan for and
nomic viewpoint. Reliable biological herbicides are proceed with successfully. Instead, it is something
needed so that we have economically as well as that you think about, develop a plan for, and con-
environmentally sound alternatives to synthetic stantly monitor, customize and adjust. It is neces-
herbicides. Applications of biological herbicides sary to identify the weed species at your site and to
when conditions are conducive, and of synthetic study them. They should be classified into winter
herbicides if they are not, is another possible man- annuals, summer annuals and perennials. You
agement strategy (Bewick, 1996). should learn how those weeds differ from your
The fact that the organisms used as weed patho- turf. Which nutrients do they need in the greatest
gens in biological herbicides can only exist in dam- amounts compared with the turf? How much water
aging populations when environmental conditions do they require? Can they stand low mowing? Do
in the habitat are particularly conducive for disease they need light to germinate? Is there some factor
is not only a restriction, it is also an advantage. such as compaction or low nutrition for which they
Many of the pathogens that are used as biological are better suited than the turf? These questions and
herbicides are classified in the same genus as the others need to be answered before you can provide
pathogens that cause disease in turf. Sclerotinia the situations that favor the turfgrass and discour-
sclerotiorum, for instance, has been successfully age the weed. Obviously, using herbicides is easier,
tested in controlled environments as a biological and is necessary when near-perfect turfgrass sur-
herbicide for dandelion (Riddle et al., 1991). faces are required. However, habitat management
Sclerotinia homeocarpa causes dollar spot, a dis- is far more permanent than herbicide applications.
ease of many turfgrasses, with particular severity Choosing the right turfgrasses for the environ-
on creeping bentgrass and Kentucky bluegrass. mental conditions and customer requirements is
Puccinia spp., the same genus of organisms that necessary for successful habitat management.
causes rust on perennial ryegrass, Japanese lawn- Blends and mixtures are normally more competi-
grass and other turfgrasses, has demonstrated tive than monocultures, but each species and culti-
potential for the control of nutsedge species var must be adapted to the environment for the
(Cyperus spp.) (McCarty and Tucker, 2005). combination to be successful. Rapidly germinating
Bipolaris spp., from the genus of pathogens that and vigorously spreading species are desirable, but
causes leaf spot on turfgrasses, and Pyricularia the species that are most competitive at maturity
spp., from the genus that causes gray leaf spot on are more important for long-lasting weed suppres-
turfgrasses, both have potential for use in control sion. Certain weeds can often survive conditions
of goosegrass (Eleusine indica) (Figliola et al., that are not conducive to vigorous turfgrass growth.
1988). Finally, a biotype of Xanthomonas Such conditions may be compaction, poor drainage,
campestris, the bacterial pathogen that succeeded low fertility or soil salts. If the turfgrass is expected
in eliminating ‘Toronto’ creeping bentgrass as an to compete, these conditions will have to be perma-
economically viable cultivar, has been suggested for nently repaired. Otherwise, the weed will always
use in reducing populations of annual bluegrass in have the upper hand.
bermudagrass (Johnson, 1994). With the exception Most weeds cannot persist under regular mow-
of X. campestris, these pathogens are all fungi. ing. However, there are enough that can to make
However, fungi, and especially bacteria, are well our job very difficult. Weeds are opportunistic and
known for their ability to develop resistance to tenacious. They are considerably easier to remove
fungicides and antibiotics, and for their ability to when they are young than after they mature and
rapidly evolve to infect alternative hosts. Therefore, spread. Higher mowing heights tend to discourage
if these pathogens were able to exist in the ecosys- most weeds and provide a competitive advantage
tem in great numbers for a long period, there is a for the turf. A dense turfgrass canopy shades the
chance that they could adapt to infect turf. For that soil and makes it difficult for some weeds to

Managing Competition Among Plant Species 185

germinate and difficult for other weeds to survive McCarty, L.B. and Tudker, B.J. (2005) Prospects for man-
turfgrass competition. Nitrogen fertilization usu- aging turf weeds without protective chemicals. Interna-
ally favors the turfgrass, but phosphorus fertilizer tional Turfgrass Society Research Journal 10, 34–41.
often favors weeds. Potassium is highly desirable Soldat, D.J. and Petrovic, A.M. (2008) The fate and trans-
port of phosphorus in turfgrass ecosystems. Crop
for turfgrass growth, but too much potassium may
Science 48, 2051–2065.
lead to weed encroachment. Finally, turfgrass dam-
age needs to be addressed quickly before weeds
have an opportunity to get established in damaged Suggested Websites
areas. Georgia Turf (2010) Available at: http://commodities.
For the most part, ecological weed control is a caes.uga.edu/turfgrass/georgiaturf/index/ (accessed
constant process of addressing problems and devis- 1 March 2010).
ing solutions. It requires perpetual study, observa- University of California (2010) UC IPM Online: Statewide
tion and problem solving but the long-lasting Integrated Pest Management Program. ‘How to
benefits are particularly rewarding. Manage Pests.’ Identification: Weed Photo Gallery.
Available at: http://www.ipm.ucdavis.edu/PMG/
weeds_intro.html (accessed 10 August 2010).
Virginia Cooperative Extension (2010) Virginia Tech
Suggested Reading
Weed Identification Guide. Available at: http://www.
Busey, P. (2003) Cultural management of weeds in turf- ppws.vt.edu/scott/weed_id/rightsid.htm (accessed 1
grass: a review. Crop Science 43, 1899–1911. March 2010).

186 Chapter 12
 Managing Competition Between
13 Turf and its Pests

Key Terms
Integrated pest management (IPM) is a plan that combines different pest-management strategies to manage
pest damage below a threshold level.
A threshold level is the level of turfgrass damage that your customer is willing to accept. It is a turfgrass manager’s
job to maintain the turfgrass stand below the threshold level determined by the customer.
A single tactic system is the use of a single strategy to control a turfgrass predator. Single tactic systems can be
very reliable in the short term but they are always temporary. The most common single tactic systems for
predator control are breeding for genetic resistance and pesticide applications.
Predation is the consumption of one organism by another and in its broadest sense includes parasitism.

13.1 How Can We Affect Relationships more maintenance. As turfgrass density increases,
Between Predator and Prey? more intraspecific competition occurs. The severe
competition among grass plants causes each plant
For many organisms, turfgrass is food. The grasses
to become weaker. The weaker plants are more
from which turfgrasses were derived were, and in
easily damaged. In addition, the presence of a sub-
many cases still are, forages for cattle, sheep and
stantial source of food, the nice dense turf, attracts
other mammals. In fact, cattle, sheep, goats, horses
more predators. For that reason, the most perfect
and a few other animals served as the first mowing
turfgrass stand is the stand most likely to crash.
devices. That is not the case today. We rarely com-
bine turfgrass and forage grass for aesthetic or
13.2 Predator/Prey Relationships
functional turf. However, turfgrass is still food for
insects and microorganisms. Therefore, turfgrass is The natural environment is one huge predator/prey
a resource for those organisms and, as you learned relationship. All organisms, including humans,
earlier, an increase in resources usually means an compete within their species for space and resources.
increase in the organisms that use them. When turf- The individuals best adapted to the local environ-
grass is lush, green and dense, as we like it, turfgrass ment survive and multiply. However, because we,
pests become numerous, sometimes lethal. For that as humans, are at the top of the food chain, we
and other reasons that you should already tend to forget that everything else is prey. Although
understand, as turfgrass becomes more perfect, its humans may be attacked and eaten by large ani-
maintenance becomes increasingly difficult. As it mals, such a case is rare and generally only occurs
reaches its carrying capacity, especially a carrying because the animal has lost its fear of us or was
capacity that has been artificially induced by our starving. Most large animals are hunted, and as
management, it becomes quite vulnerable to dam- long as that continues they will fear us and avoid
age. Management inputs beyond those that occur us instead of considering us as food. Every other
naturally increase the maintenance required of us plant, animal or microorganism, though, is consid-
severalfold. Turfgrass management and its mainte- ered food for something. Every organism has to
nance do not sustain a 1:1 linear relationship. Every either photosynthesize or eat, and there are a lot of
positive management input requires severalfold insects and microorganisms that eat turf.

©CAB International 2011. Turfgrass Physiology and Ecology (G. Bell) 187
 The typical predator/prey relationship is based worms may use turfgrass as an alternative food
on random encounters. As a prey population source. Regardless, the random predator/prey
grows, the likelihood of its predators encountering model generally fits managed turfgrass sites very
prey increases. As predators encounter more prey well, mostly because sites managed for turf are
the predator populations become healthier, fewer nearly all turfgrass. Therefore, alternative prey or
die and more reproduction occurs. The predator predators of other species are only present in the
populations grow because there is an abundance of turfgrass habitat in very small numbers.
prey. However, as the predator populations increase
and more predators encounter prey, the prey popu-
Human impacts on predator/prey
lation becomes unhealthy, many die and reproduc-
relationships
tion slows. The decline in prey is then followed by
a decline in predators and the cycle begins again Humans have a tendency to focus their crop pro-
(Fig. 13.1). duction efforts on a very narrow scale instead of
Of course we know that the predator/prey rela- considering the crop, in our case turf, as part of a
tionship is not random. Predators seek prey and are much larger ecosystem. We forget that there is
only found in habitats where prey exist. However, a multitude of activity occurring on our site. Even
the random model is a good illustration of how the a somewhat inert pure sand system has a huge
relationship works. In addition, predators may microbiological community (Bigelow et al., 2002;
prey on more than one species and concentrate Elliott et al., 2004). These microbial populations
their efforts on an alternative species when a pri- are affected by local environments in the same way
mary species declines. In that case, although the as turfgrass (Giesler et al., 2000). They live,
population of primary prey has declined, predator reproduce and die in a turfgrass community. They
numbers may remain the same, but the predators compete with each other, prey on each other, and
will be more concentrated in habitats conducive to sometimes prey on turf. There are a number of
the alternative prey species than in habitats condu- animals and birds, including humans, which include
cive to the primary prey. Such a response is com- turfgrass sites as part of their habitat. A turfgrass
mon in fall army worms (Spodoptera frugiperda) in site is part of a much larger ecosystem and is
regions where wheat (Triticum aestivum) is plenti- affected by that system. Turfgrass is part of the
ful. Once the wheat crop is harvested, the army human community. Turfgrass, in fact, does not
exist in the wild. Forages exist in the wild but
turfgrass, by definition, only exists because of
human intervention.
Humans have a huge impact on turf. Human
Prey activities on managed turf determine at least a
Predators portion of the predator/prey relationship in the
Population

community. The more highly managed the turf,

hence the most desirable turf, the more impact that
humans have on it. In a natural environment, the
predator/prey relationship exists in relative balance.
Time
However, we can use knowledge, experience and
creativity to influence the predator/prey relation-
Fig. 13.1. In a random predator/prey relationship, ship on managed turf, and that is what we are
predators increase after a sufficient lag time following about to consider in the rest of this chapter.
a prey increase. However, as the predators feed on
the prey, the prey population begins to decline. As
the prey resource declines, the predator populations 13.3 Managing Turfgrass Predators
decline and the cycle begins again. The random model
There are many organisms that we consider to be
of predator/prey encounters fits turfgrass populations
fairly well because where management is practiced,
turfgrass pests. Some organisms are competitors,
turf is nearly all turfgrass. Consequently, the turfgrass is like weeds, and others are predators, like insects
the only prey present in large numbers and most of the and disease pathogens. In Chapter 12, we discussed
predators present are predators of turf or predators of interspecific competition primarily from other
turfgrass predators. plants such as dandelions (Taraxacum officinale)

188 Chapter 13
and crabgrass (Digitaria spp.). In this section of expected to be perfect. We can’t do perfect but we
Chapter 13, I will briefly introduce integrated pest can come awfully close if we use every strategy
management (IPM), a system of pest control strate- available to us, including pesticides.
gies that you are probably familiar with. Turfgrass
IPM is designed to guide the planning of pest con-
Turfgrass IPM
trol programs for both competitors and predators.
Its use is desirable for designing weed control plans As mentioned earlier, IPM is a collection of strate-
as well as plans for predator control. The ecologi- gies incorporated into a pest management plan.
cal management of interspecific competition is a One of the philosophies of IPM is that all alterna-
basic part of IPM. Both management systems take tive strategies should be employed before pesticides
similar approaches to IPM, but IPM has a much are applied. At our current state of knowledge,
broader scope. It would be valuable for you to maintaining turfgrass at a 0% damage threshold or
consider how the ecological management of inter- close to it is not possible without pesticides.
specific competition presented in Chapter 12 fits Therefore, pesticides are always part of the plan
into a complete IPM program. However, in this where perfection is desired. If our customers
chapter we are primarily concerned about how demand perfection then we have no choice but to
predator/turf relationships work. In the next employ pesticides. We can encourage our custom-
section, you will see how the ecological principles ers to accept something less than perfection but we
of turfgrass/predator relationships can affect man- are not the ones who determine the threshold, they
agement plans that enhance an IPM program. are. So if we are to maintain our profession indefi-
In the USA and many other countries, IPM, or nitely we need to strive to manage perfection with-
something similar, is the accepted method for out pesticides.
managing turfgrass pests. Most turfgrass managers According to most pest management experts,
practice IPM, whether they realize it or not. The there are six basic strategies always considered
processes of IPM are multifaceted and cover when developing an IPM plan. They are:
practically every technique for pest control. Our
1. Regulatory strategies: government intervention
study will be more specific to natural factors that
to prevent damaging pests from being introduced
influence turfgrass pests and how to use them
into an area, for the removal of noxious weeds, or
effectively. However, the techniques presented
to restrict products or procedures that encourage
would probably be the most important part of an
pest population growth.
IPM plan.
2. Genetic strategies: the development of cultivars
When developing or revising an IPM plan it is
or species with genetic resistance or other adapta-
not only the technology that makes a difference, it
tions that make the target organism, our grass,
is also the philosophy. The IPM process is a combi-
more competitive with its pests.
nation of strategies used to maintain a particular
3. Cultural strategies: these include mowing, irri-
turfgrass site under a desired threshold level of pest
gation, fertilization and soil cultivation to help our
damage. The threshold level is determined by the
plants be more competitive or to remove conditions
end user, your customer. Those thresholds are less
that encourage pests.
likely to be based on nature conservation than they
4. Physical strategies: the actual capture of insects
are on economics. Sadly, most people are 100% in
or animals, hand removal of weeds, dew removal,
favor of nature conservation as long as it does not
equipment cleaning and other labor-intensive
affect what they want to do or cost them money.
practices to discourage the spread of disease.
Fortunately, managing turfgrass to near perfection
5. Biological strategies: the use of one organism to
is expensive in both time and money. Therefore,
affect another, such as a natural predator or a
most customers are happy to endure some weed
competitor. Some would include naturally occurring
encroachment and a little insect or disease damage
toxins in this category, but I will not.
rather than pay for perfection. The exceptions are
6. Chemical strategies: the use of naturally occur-
bowling greens, putting greens and tennis courts,
ring or manufactured chemicals to kill pests.
where the condition of the turf can drastically
influence play. On those areas, the damage thresh- Many of these strategies can be employed effec-
old is 0%, meaning that no more than 0% damage tively to discourage a pest. By combining strategies,
will be tolerated. In other words, the turf is the pest control program is more continuous, more

Managing Competition Between Turf and Pests 189

reliable and more effective. A good IPM program is viral diseases of turfgrass. St. Augustinegrass decline
a long-term solution, not a temporary treatment. is generally considered to be the only economically
The opposite of an IPM program is a single-tactic important viral disease of turf. The St. Augustinegrass
system. decline virus also attacks centipedegrass (Eremochloa
ophiuroides), resulting in the disease called centipe-
degrass mosaic. In the USA, gray leaf spot was a
Single-tactic systems
disease of St. Augustinegrass and tall fescue, but it
A single-tactic system is the use of a single strategy was not a major disease on perennial ryegrass until
to control a turfgrass predator. Single-tactic sys- recently (in the last 15 years or so). The gray leaf
tems can be very reliable in the short term, but they spot pathogen had to adapt to prey on ryegrass.
are always temporary. The most common single- Breeding and genetics may be our strongest line of
tactic systems for predator control are breeding for defense against predators, but because predators
genetic resistance and pesticide application. Both of readily adapt to alternative hosts, it is not always
these strategies are included in IPM planning effective as a single-tactic system.
because they can provide excellent control. Insects and microorganisms are highly adaptable.
However, neither strategy is reliable long term. For that reason, predator resistance to a single-
Turfgrass species are attacked by many different tactic system is likely to occur over time. Chemical
organisms, depending on climate and use. However, strategies are extremely effective for predator con-
a turfgrass species in a given region usually only trol but, as was mentioned in Chapter 12, their
has major problems with a small number of par- effects are quite temporary. One application must
ticular insects or pathogens. Bermudagrass be followed by another and another. If, after a
(Cynodon dactylon), for instance, is somewhat period of intense chemical control, it appears that
resistant to most diseases but is highly susceptible you have eliminated the problem, just wait a year
to spring dead spot (Ophiosphaerella herpotrica or two and it will probably reappear. It is highly
and others) disease in regions where winters are unlikely that you will ever be successful in com-
fairly cold. Tall fescue (Festuca arundinacea) is pletely eliminating a turfgrass predator. They will
highly susceptible to brown patch (Rhizoctonia nearly always come back, and when they do, they
solani) disease and St. Augustinegrass (Stenotaphrum may have developed resistance to your previous
secundatum) and fine fescues (Festuca spp.) tend to practices. That is not an uncommon occurrence.
attract chinch bugs (Blissus insularis). Kentucky Multiple chemical applications encourage
bluegrass (Poa pratensis) is often damaged by blue- predator resistance. Because every individual of a
grass billbug (Sphenophorus parvulus) and peren- species usually has a unique genetic composition,
nial ryegrass (Lolium perenne) by gray leaf spot the likelihood that a few individuals will be resistant
disease (Pyricularia grisea). Breeders have or may to a pesticide application is high. With insects and
be able to develop cultivars resistant to these highly especially with microorganisms that breed rapidly, a
important diseases and insects but, over time, the resistant population can occur. A resistant predator
insects and diseases may adapt to feed on the resist- population is most likely to occur following multi-
ant cultivars, or the old diseases or insects may be ple applications of a single pesticide or different
traded for new ones. Therefore, the breeding of pesticides that use the same mode of action.
resistant cultivars requires continuous attention, Following the first pesticide application a small
and breeding alone is not completely effective for portion of predators survive. Survival includes a
predator control. portion of the susceptible individuals and all of the
When Kentucky bluegrasses, for instance, resistant ones. The remaining predators reproduce,
were developed to resist melting out disease resulting in more resistant individuals. As more
(Helminthosporium spp.), the most important susceptible individuals are eliminated by recurring
disease of the species at the time, other diseases, pesticide applications, the population comes to
summer patch (Magnaporthe poae) and necrotic comprise mostly resistant individuals (Fig. 13.2).
ring spot (Leptosphaeria korrae), went from being By using multiple applications of a chemical with
minor predators (diseases) to major ones. For some the same mode of action you have almost eliminated
unknown reason, St. Augustinegrasses that are intraspecific competition among resistant and sus-
resistant to chinch bug damage are more susceptible ceptible predators. Because of your actions, only
to St. Augustinegrass decline, one of the very few resistant predators exist. Continuing applications

190 Chapter 13
 Resistant pathogen carefully. Do not encourage a resistant predator
Susceptible pathogen population. Make sure that you kill what you
intended to kill and do not kill unrelated organ-
isms, your turf for instance, or your neighbor’s
garden. Remember that insecticide and fungicide
applications kill not only predators, but related
beneficial organisms. These products are broad-
Pesticide A Reproduction Pesticide A Reproduction based to kill multiple insect predators and fungi.
When you kill the pathogen, you also kill some of
its competing beneficial organisms. Be careful what
you apply and when you apply it.

Knowledge is a weapon
Pathogen killed by pesticide In order to effectively manage a pest, information
and knowledge are required. Knowledge is a
Fig. 13.2. Predator resistance. With the first application
valuable weapon and, in this case, is even more
of a pesticide, much of the predator population will be
important than experience. If you intend to do
killed, but many are likely to survive. A few of those that
survive may be resistant to the mode of action of the your best to manage a turfgrass pest, predator or
pesticide. These resistant individuals grow and breed competitor, you must have an intimate knowledge
and the resistant population builds. With repeated of that organism. Books, libraries, conferences,
applications of pesticide with the same mode of action, newsletters, magazines, Internet sources and
most of the predators remaining will be resistant and personal contacts are all sources of information
a pesticide with a different mode of action will be and knowledge. You may like to do things your-
required to kill them. self, make your own decisions and avoid large
gatherings, but if you intend to do the best possi-
will not only reduce intraspecific competition, they ble job with your turf, you have to have sources
will also kill beneficial organisms in the same class of information, including conferences and personal
as the predators, thus inhibiting interspecific com- contacts (Box 13.1).
petition as well. The result will be a very high Every turfgrass predator has a history. More
population of resistant predators. Continuing importantly, it has a life cycle, conditions that it
applications of the same products will further likes and those that it dislikes, susceptibility to
increase the resistant predator population and certain other predators, and symptoms and signs
make the damage worse. If you are in a situation that identify its presence. You must be familiar
where you have to use pesticides to meet customer with those characteristics in order to control it.
expectations, you must use them wisely and Disease and insect damage do not occur randomly,

Box 13.1 Building a “network” of information sources.

You can’t know everything, and conversations with of industry developments and investigate many other
people experiencing the same problems and those situations using their network. A network is an
who have solved them or done research on them is extremely valuable resource. A good network takes
invaluable. Business people call an individual’s list of years to develop, but it grows rapidly following your
personal contacts a network and they consider first few contacts as others refer contacts to you.
networks to be extremely valuable sources of Developing a network is not the least bit difficult, you
information. Those with the best networks can gather simply have to be willing to meet new people and to
information about any number of subjects from all listen to new ideas. Most importantly, you have to
over a country, perhaps the world, with a few phone belong to organizations and attend the meetings
calls. They can effectively screen potential employ- where you are most likely to meet other people in
ees, learn about new laws, find a new job, keep track your profession.

Managing Competition Between Turf and Pests 191

they occur for a reason, and they are, in most cases, symptoms and signs of impending predator
predictable. We do not know everything that we problems, photographs can be a very valuable
need to know about turfgrass predators, partially source of instruction. More importantly, a wide-
because they are so easy to kill using pesticides and angle photograph is an instant record of location
partially because they are biotic organisms that eas- and the photograph can be labeled easily for date,
ily adapt to multiple environments. However, we either automatically by the camera or digital file or
are always learning more about turfgrass pests and by hand. Close-up photographs can be included to
if you want to or are forced to use fewer pesticides, show the intensity of the damage and to help with
you have to learn everything that you can about a identification. A digital photograph is an instant
particular pest to manage it effectively. source of information that can pinpoint the loca-
tion where an assistant found a potential problem
that you need to observe, and it is a record that
Keep watch on your site
may help you with early detection of similar occur-
You may be a parent or an older brother or sister. rences in the future. With photograph in hand, you
You may have been a baby-sitter. During your life do not need the assistant to guide you to the site.
you probably have had at least some small respon- You will probably want to keep a map of loca-
sibility for looking after young children. When you tions where problems have occurred so that you
are responsible for a young child, you have to keep know where to look for similar problems in the
a close watch on the child and the environment future. You may also want to keep a journal of
around that child. Potentially dangerous situations specific techniques that you used to attempt a cure.
happen fast and without warning. Record those attempts that worked and those that
Managing turfgrass is similar but easier than didn’t for future reference.
having responsibility for a small child. There is Watch for the environmental conditions that
usually some warning before a predator attack, but precede predator problems. Insect problems gener-
you must be paying attention to see it. When ally occur by life cycle. Usually there are only one
watching a child you try to place dangerous objects or two life stages of insects that are damaging to
out of reach and provide a harmless environment. turfgrass and they only occur during certain peri-
You do the same with turf, but as with a child, you ods of the year. Therefore, monitoring and some
realize that there is no such thing as a completely control techniques are only needed for a few weeks
harmless environment. You have to keep watch. or less each season. You have to know the life cycle
Where there is turfgrass there will always be turf- of the insect to determine when these periods
grass predators. occur.
As a manager, you must develop a system for A good turfgrass manager is always aware of
monitoring your entire site. The site, large or small, changing environmental conditions and what those
requires daily inspection by someone knowledgea- conditions mean for possibilities of predator dam-
ble in the symptoms and signs of invading preda- age. Turfgrass pathogens are always present in
tors. As with any human disease, early detection is some form or another. However, turfgrass diseases
the best possible means of correction. Every large only occur when the environment is more condu-
turfgrass area has predator “hot spots”. A hot spot cive for the pathogen population growth than it is
is an area that is most vulnerable, often for reasons for turfgrass growth. There are usually multiple
unknown, to a particular predator or pest. Specific environmental factors that affect disease pressure
diseases or insect damage will usually occur in a (Vargas et al., 1993). The period of likely disease
hot spot before they occur on the rest of the site. occurrence is extended when the turfgrass is weak-
Therefore, the hot spots are the places you monitor ened by poor management or other stress (Vargas,
most closely. They act as early warning devices 1994). Consequently, turfgrass health is an impor-
once you identify them. tant part of disease management as it is of weed
Part of identifying hot spots for certain diseases management. That is usually not the case where
and insects is record keeping. Records can be jour- insects are concerned. Some damaging insects may
nals, maps or photographs. Photographs are under- seek out the healthiest turf for laying their eggs and
utilized in turfgrass management by all but causing problems. Because insects attack according
instructors. If a turfgrass manager is to train assist- to stage of life cycle and pathogens attack during
ants and laborers to be on the lookout for certain certain environmental conditions, there is some

192 Chapter 13
warning as to when to expect insect and disease
problems. That is one reason why record keeping,
especially recording the dates of occurrence, is so
important. A good plan for predator control hinges
on a good monitoring program and good record
keeping.

13.4 Affecting Relationships Between

Predators and Turf
The ecological concept of predator control is to use
the predator’s natural enemies against it. With that
statement, the first thing that probably comes to
mind is biological control. We could use nematodes
to control white grubs (Phyllophaga spp., Fig. 13.3. Spring dead spot is a common disease
Cyclocephala spp., Popilia japonica and other bee- of bermudagrass (Cynodon dactylon) in temperate
tles of the order Coleoptera) or bacteria to control climates where winters are cold. The damage that you
fungi but, as you learned in Chapter 12, biological see on cultivars 3–6 is severe. The turf in the patches
control is not particularly reliable. Although biologi- is dead and will not recover. Cultivars 1 and 2 are
cal control may be part of your pest control pro- resistant to the disease, but if conditions are exactly
gram, it is the abiotic enemies of a predator that are right they will also be infected. However, with proper
most reliable in facilitating its demise. Just as we can management, the occurrence of the disease in cultivars
1 and 2 can be maintained below a low threshold level
influence the local environment to favor photosyn-
with no pesticide applied.
thesis, respiration and transpiration, we can influ-
ence the local environment to reduce predation.
However, throughout any discussion of predator to help relieve spring dead spot pressure compared
management, it must be remembered that an increase with higher heights (1.5 inches, 38 mm) of cut
in turfgrass is an increase in its predators’ resources. (Martin et al., 2001). The spring dead spot patho-
More turf attracts more turfgrass predators and gen seems to prefer a relatively basic soil pH, so N
healthier turf feeds more predators. Therefore, it fertilization with an acidifying fertilizer such as
might seem that from an ecological standpoint we ammonium sulfate tends to discourage the disease.
cannot have healthy dense turf without high preda- However, N applications late in the growing season
tor numbers. Remember, though, that turfgrass and are likely to encourage the disease (Dernoeden
its predators do not share the same niche. Turfgrass et al., 1991). Vertical mowing and aeration for
may be a source of nutrition for the predators, but thatch reduction also reduces the susceptibility of
there are other biotic and abiotic factors that deter- bermudagrass to spring dead spot (Tisserat and
mine whether or not a turfgrass predator can live in Fry, 1997). None of these practices alone is suffi-
a turfgrass community. cient to provide protection, but together and in
combination with the proper irrigation to help
mask symptoms, bermudagrass turf can often be
Cultural practices that affect predators
managed below damage thresholds with a combi-
Mowing height alone, irrigation alone, nutrition nation of cultural practices and no pesticide.
alone, thatch reduction alone or other cultural In rare cases, a single cultural practice can
practices alone will probably not have a significant substantially reduce predator damage. Bermudagrass
impact on turfgrass predators but, in combination, may contract dollar spot (Sclerotinia homeocarpa)
management of cultural practices for specifically in late spring, but an application of N fertilizer is
discouraging targeted pests can provide adequate often enough to relieve disease pressure (Carrow
protection in many cases. Spring dead spot disease et al., 1987). Nitrogen fertilizer also helps to reduce
pathogens, for instance, are a major predator of dollar spot in Kentucky bluegrass, hybrid bluegrass
bermudagrass in temperate regions (Fig. 13.3). The (Poa pratensis × Poa arachnifera) and perennial
disease is difficult to control with fungicides, ryegrass (Treadway et al., 2001; Teuton et al.,
although low mowing, 0.5 inches (13 mm), seems 2007). It has also been known to reduce anthracnose

Managing Competition Between Turf and Pests 193

on annual bluegrass (Poa annua) and is recom- best on your site. Make sure to monitor your pro-
mended as a control measure for rust disease grams, especially when changes in activities are
(Puccinia spp.) on multiple grasses (Vargas, 1994; made. There may be conditions on your site that
Inguagiato et al., 2008). Nitrogen fertilization require different management from that which is
affects many turfgrass diseases. Sometimes, the normally recommended.
source (organic or inorganic) of N fertilizer has an
effect on disease (Landschoot and McNitt, 1997).
Genetic resistance to predators
In most cases, N fertilization discourages disease,
but in the case of insects, the opposite may be true. One means of enhancing turfgrass resistance to
Fertilized St. Augustinegrass, for instance, is more predators is through the use of mixes and blends.
susceptible to chinch bug than non-fertilized Many predators, especially disease pathogens, are
St. Augustinegrass (Busey and Snyder, 1993). quite specific to the cultivars and species of turf-
Although N fertilization is often a positive means grass that they prefer (Rose-Fricker et al., 1997). In
for reducing disease pressure, it has also been addition, cultivars of turfgrasses and turfgrass spe-
known to encourage gray leaf spot disease on per- cies differ in their level of susceptibility to predator
ennial ryegrass (Williams et al., 2001). attack. By mixing or blending we can provide vari-
Some insects, such as black turfgrass aetenius able predator preferences and plant resistance to
(Aetenius spretulus), prefer irrigated turf over dry help slow the spread of disease or insect damage
turf (Jo and Smitely, 2006). In some cases, a (Fig. 13.4). Many diseases spread plant to plant
micronutrient has an effect on a predator, as in Mn through extended hyphae. If all of the surrounding
suppression of take-all patch disease plants are resistant, the disease has nowhere to go.
(Gaeumannomyces graminis) in creeping bentgrass In addition, if only a portion of the stand is affected,
(Agrostis stolonifera), but that is rarely the case the visual quality and playability of the turf will be
(Heckman et al., 2003). Then again, soil pH often better than that of turf in a stand of uniformly sus-
affects diseases such as creeping bentgrass dead ceptible plants (Abernathy et al., 2001; Brede,
spot (Ophiosphaerella agrostis), spring dead spot 2005). Presumably, predation by common insects or
and take-all patch (Couch, 1995; Kaminski and pathogens would remove the susceptible species or
Dernoeden, 2005). The point of all of this is that cultivars from the mixed stand, but that is not
random assumptions are not a part of predator always the case (Golembiewski et al., 2001). Often,
management. Predators and their prey must be the predator damage is not severe enough to kill the
managed specifically. You also must be familiar plants affected. If the damage is slowed sufficiently
with how your management of one predator affects by the mixed grasses, many susceptible plants may
another. Irrigation management, for instance, that
has a positive effect on the control of one predator
may have a negative effect on the control of Resistant host
Susceptible host
another (Jiang et al., 1998). Also, you must moni-
tor and record your efforts and their effects. For
many years, chinch bug damage was considered
to be mostly a problem in drought-stressed
St. Augustinegrass but recently that has been ques-
Disease
tioned (Vazquez and Buss, 2006). Researchers who
found no difference in chinch bug damage on irri-
Disease
gated and non-irrigated St. Augustinegrass suggest
that chinch bug damage may simply look like
drought damage or that the chinch bug populations
breed more rapidly in the warmth of drought
Fig. 13.4. Turfgrass mixes and blends are likely to
stressed turf. Acidifying fertilizers help to prevent
contain species or cultivars with variable resistance to
earthworms and microarthropods in some instances predators. Because susceptible plants are interspersed
(Potter et al., 1985), but not others (Backman with resistant ones, damage spreads more slowly,
et al., 2001). Scientific or professional advice is a fewer plants are damaged and visual quality and
great source of good information, but you are playability is less affected than it would be in a single
responsible for developing the programs that work species, single cultivar stand.

194 Chapter 13
escape and reproduce, thereby maintaining the et al., 2000). As little as 40% of endophyte-infected
mixed genetics in the stand. perennial ryegrass overseeded into a stand of
Turfgrass breeding programs are constantly striv- Kentucky bluegrass can significantly reduce dam-
ing to develop new varieties that are resistant to a age by bluegrass webworm (Parapediasia teter-
species’ most common predators. Much progress rella) (Richmond and Shetlar, 1999). Endophytes
has been made toward both disease and insect are generally capable of surviving the same condi-
resistance in grasses, although many predator tions as their turfgrass hosts, but some may be
problems remain a significant challenge. affected by certain conditions, such as severe cold
(Rochefort et al., 2007). In some cases, it may be
found that a particular endophyte confers other
Biological control of predators
positive characteristics in addition to insect resist-
Biological control of predators is deemed prefer- ance. For instance, research has demonstrated that
able to applications of synthetic pesticides by most at least one endophyte-infected tall fescue cultivar
people. Whether or not a synthetic (that is, an arti- maintains higher leaf turgor pressure under drought
ficial) application of a biological organism is really stress than its uninfected counterpart (Richardson
any different from a synthetic application of a et al., 1993), while other cultivars may not (White
manufactured pesticide is a subject for debate. et al., 1992). Researchers in New Jersey found that
However, society appears to believe that biological endophyte-infected perennial ryegrass was not only
control mechanisms are less dangerous to the resistant to insect damage but also to red thread
human environment than manufactured pesticides, disease (Laetisaria fuciformis) (Bonos et al., 2005).
and public perception is important. Consequently, To this date, seed-borne endophyte infected turf-
development of biological mechanisms for turf- grasses are our greatest success for biological pest
grass management has become a subject of interest. management in grasses. Unfortunately, endophyte
The greatest success of biological control methods infection has only been successful in perennial rye-
in turfgrass has not been in the application of bio- grass and the tall and fine fescues; it is currently not
logical agents but in the use of seed-borne fungi commercially possible in most turfgrass species.
called endophytes.
Other potential methods for biocontrol
Endophytes of turfgrass predators
Endophytic fungi are a group of microorganisms There have been favorable reports concerning com-
that populate a turfgrass plant and can be borne by posting and the application of compost for the
a turfgrass seed. Consequently, the fungi in the seed suppression of turfgrass fungal pathogens. The
grow with the plant. These fungi form a symbiotic application of organic composts derived from some
relationship with turfgrass and other plants much materials is believed to alter the turfgrass commu-
like the relationship between plants and mycor- nity enough to suppress pathogens. The bacteria in
rhizae that you probably studied in biology. In the some types of compost are believed to compete
case of endophytes and turfgrass, the fungi live in directly with parasitic fungi that attack turf, and
the plant and facilitate a resistance to some insect many bacteria that exist in compost have been
predators. Although white grubs, the most com- identified as antagonistic toward turfgrass fungal
mon insect predators of turfgrasses, do not appear pathogens (Boulter et al., 2002). However, as yet,
to be affected by endophytes, other insects are actual successes in disease reduction in applications
(Potter et al., 1992). Chinch bug damage in peren- of compost in the turfgrass industry have not been
nial ryegrass is directly related to the number of reported. This idea, though, is a good example of
plants infected with endophytes (Richmond and using ecological tactics to benefit turfgrass manage-
Shetlar, 2000). Endophytes are also known to dis- ment and is likely to produce positive commercial
courage feeding by billbugs, most caterpillars and products with further study.
various types of webworms. In mixtures of Certain waste products or extracts derived from
Kentucky bluegrass and perennial ryegrass, blue- the processing or tilling of crop plants have been
grass billbug (Sphenophorus parvulus) damage was tested as sources of biocontrol agents. A crop of
directly related to the proportion of endophyte- oilseed rape (Brassica napus), for instance, tilled
infected perennial ryegrass in the stand (Richmond into a sod farm infected with sting nematodes

Managing Competition Between Turf and Pests 195

(Belonolaimus longicaudatus) was effective in important. These discriminating factors, with spe-
reducing nematode populations compared with cial preference for customer expectations, will be
sites under continuous bermudagrass (Walker used to determine pest damage thresholds for each
et al., 2005). Extracts of poinsettia (Euphorbia category. An athletic field complex might have
pulcherrima) and spotted spurge (Chamaesyce three or four pest management categories. The pri-
maculata) have also demonstrated promise in the mary field might have a very low pest management
control of sting nematodes (Cox et al., 2006). threshold such as 2 or 3% damage. Practice fields
Other uses of crop plants, their extracts or post- might have a higher damage threshold, 10%, and
process waste products have also been tested for the surrounds may be acceptable at 20%. Home
antagonistic activity on turfgrass predators. None lawns and commercial property would be managed
have yet reached a stage of widespread use. to thresholds primarily determined by the home or
Another possibility for biological control of preda- business owner. A home lawn would have one
tors is the application of nematodes or fungi that threshold value but a commercial property might
prey on insect pests, or bacteria that feed on have two or three. The idea is to manage without
pathogenic fungi. allowing damage to exceed the threshold levels but
Many of these biocides have been tested and a to set the levels as high as possible. A golf course
few have been found effective enough to reach com- provides an excellent example.
mercial production in limited amounts. None has A golf course, depending on manager and
reached the general industry acceptability of BT membership, would probably be divided into at least
(Bacillus thuringiensis), a bio-insecticide commonly three pest management categories. The highest
used in ornamental horticulture. A bio-fungicide or priority units, the putting greens, would be Category
bio-insecticide must not only be effective for pest 1. Category 1 might have a zero tolerance or 0%
control, it must also be able to live in a turfgrass pest damage threshold. The tees and fairways,
environment under differing environmental condi- Category 2, might have a 10% damage threshold
tions. Strains of these organisms differ in their abil- and the rough, Category 3, a 20% damage thresh-
ity to infect the target pest and also in their viability old. A 20% damage threshold is high. A 2 or 3%
and persistence in a turfgrass community (Thompson weed encroachment, for instance, is quite noticeable.
et al., 2005). Because they are living organisms, Insect and disease damage, however, are not particu-
they require special care during manufacture and larly noticeable until the grass actually turns brown.
are expensive to produce. However, many turfgrass If 20% of your turf is brown, that is a lot of brown
managers would pay a high price for them if they turf. Obviously, you also have to consider what you
knew that they would be effective. Some of these call damage. Does the turf have to be brown or just
products may normally be effective but may fail in yellow to be considered damaged? For our purposes,
a particular season for unknown reasons (Tomaso- let us go with brown as damage and let us call this
Peterson and Perry, 2007). Such is the fate of bio- golf course Any Valley Country Club.
logical organisms. They require certain resources Any Valley CC is located in a temperate region
that we may not realize are missing in any particu- and has creeping bentgrass putting greens, Kentucky
lar situation. Biocides and other biological predator bluegrass fairways and tall fescue rough. The
controls will probably be commonplace for turf- putting greens are susceptible to a number of fun-
grass use in the future. At present, though, turfgrass gal diseases, primarily dollar spot, and insects,
managers rely on pesticides, where possible, not primarily black cutworm (Agrotis ipsilon). The
because they do not care about environmental man- major problems on the fairways are also fungal
agement but because pesticides hardly ever fail diseases, mostly summer patch, and the primary
(Wang et al., 2001). insect problem is bluegrass billbug. Problems in the
rough are brown patch and white grubs.
Let us consider how you are going to manage the
13.5 Making a Management Plan
greens at Any Valley CC to resist pest damage.
A pest management plan begins with the criteria Because the Category 1 threshold is 0%, you are
that you will use to divide your site(s) into pest going to have to include pesticides on the putting
management categories. Categories will differ by greens but you are going to use them wisely. If we
turfgrass species, mowing height, customer do everything that we can ecologically with our
expectations and any other factors that you deem fairways (Category 2) and rough (Category 3) our

196 Chapter 13
pesticide use will be extremely limited. We will
rarely have to apply pesticides to maintain fairway
and rough at threshold damage levels, but we will
use them if necessary.
Before you develop a pest management plan for
the putting greens, you have to do some research
on dollar spot disease and black cutworm. You find
that, like most fungal diseases, dollar spot prefers
damp conditions. Consequently, sun is one of its
natural enemies, as well as dew removal, air move-
ment and temperatures warm enough to facilitate
drying but not high enough to stress the turf. We
can use these natural enemies against the pathogen.
In the absence of rainfall, the primary time for Fig. 13.5. Although this putting green looks like all one
dampness is following morning dew. Dew contains cultivar, it is not. In fact there are several cultivars in
guttation fluid that you learned about earlier. this photograph, all in areas of the same size as the
rectangular areas severely infected with dollar spot
Guttation fluid contains sugars, proteins and nutri-
(Sclerotinia homeocarpa). All cultivars of creeping
ents that fungi feed on, and increases their resources.
bentgrass (Agrostis stolonifera) can be infected with
Consequently, dew removal is an important process dollar spot, and all of the cultivars in the photograph
for dollar spot reduction (Williams et al., 1996). show symptoms. However, some cultivars are severely
Dew can be removed immediately after sunrise by infected while others have minor infection.
mowing or poling (also called whipping) but mow-
ing works best (Ellram et al., 2007). You will also
find that the dollar spot pathogen can exist as dor- renovation for which you can choose a cultivar or
mant hyphae (sclerotia) in the thatch and repro- blend with some resistance to dollar spot.
duces by spores, both of which can be carried by Unfortunately, as far as we know at present, there
mowing equipment and other tools that touch the are no cultivars of creeping bentgrass with resist-
putting surface. So hosing off mowers after mow- ance to cutworm damage (Williamson and Potter,
ing one putting surface and before mowing another 2001). Although renovating patches of putting
can help to delay the spread of the pathogen, and greens is not a practice to be considered, it would be
cultural practices to properly manage thatch can a practice that you would want to consider in dam-
help to reduce the disease. aged sections of fairway and rough.
Delaying the spread of a pathogen is usually syn- According to Couch (1995) the dollar spot
onymous with managing it. As was mentioned ear- pathogen grows slowly when daytime temperatures
lier, diseases occur when the environment favors the reach 60 °F (16 °C) and rapidly when daytime tem-
growth of the pathogen over the growth of the turf. peratures are consistently between 70 and 80 °F
For that reason, if you can delay the spread of dis- (21–27 °C). The nighttime humidity must also
ease until the environmental conditions change, you reach 85% before dollar spot becomes a significant
are exercising some control over it. Therefore, problem. It is interesting to note that even reason-
healthy turf is an enemy of disease. Dollar spot is ably dry climates can have a nighttime humidity
more likely to occur on dry turf than it is on turf that reaches 85%. Cold air holds less water than
that is irrigated properly, so irrigation is a weapon. warm air, so as warm air cools overnight the rela-
Nitrogen fertilization is also an enemy of dollar tive humidity increases and is normally highest
spot. Dollar spot spreads from one plant to another when the night’s temperature is lowest. The night
most rapidly by colonizing wounds. The rapid temperature is usually lowest about an hour before
growth and healing encouraged by adequate N sunrise. Early-morning humidity should be moni-
fertilization helps to delay the spread of dollar spot. tored closely. We can’t adjust the temperature or
Finally, there are cultivars of creeping bentgrass that humidity but we can use them as indicators for
exhibit low levels of dollar spot resistance (Fig. 13.5). pesticide applications.
Although you will not be renovating sections of Pesticides should be applied only when neces-
your putting greens if you maintain near 0% dam- sary. If you develop a good ecological management
age, there may be an opportunity for a large-scale plan for the common predators on your Category 2

Managing Competition Between Turf and Pests 197

and Category 3 turf, few pesticides will be needed for disease occurrence and end them when the
and you will only apply them when the environ- danger has past. You should have a good reason
mental conditions are exactly right for disease for applying preventative pesticide measures on
infection or when disease or insect damage is your Category 1 turf instead of just applying gen-
present and approaching the damage threshold. On eral blanket protections. You will probably need
your Category 1 turf, however, you will probably insecticide to prevent cutworm damage as well,
be more effective, more efficient and use less but those applications can be made at the first sign
pesticide by applying disease applications preventa- of insects or symptoms. Constant monitoring dur-
tively (before you see symptoms) than you will be ing the most likely damage periods is required.
by applying curatively (after you see symptoms) Visual inspections should occur daily and a simple
(Box 13.2). Most fungicides require applications soap flush, 1 ounce of lemon-scented dishwashing
threefold or fourfold greater to cure a disease than liquid in 2 gallons of water sprinkled over 1 square
to prevent one. Consequently, using preventative yard of turf (30 ml of liquid in 7.6 l of water over
applications when disease pressure is known to be 1 m2), should be performed weekly (Niemczyk and
high is a conservative measure for areas where Shetlar, 2000). At the first sign of insects or dam-
near-zero infection is required. age, an insecticide application is warranted.
You will begin those preventative applications Monitoring should continue, as multiple genera-
when the environmental conditions are conducive tions usually occur. You should also study the

Box 13.2. Prevent or cure?

Because the surfaces have to be so precise, if you conducive for disease. Consequently, to be sure to
are managing areas such as golf course greens or prevent disease, we would make three applications
bowling greens you will have to set your threshold in the late spring 14 days apart and three in the early
damage level very low, usually near 0%. That kind of fall on the same interval. That would require a total of
intense management is not possible without the use 10.8 ounces of fungicide per 1000 square feet
of pesticides. In most disease cases (not insects), it (3.4 g m−2) of greens. Suppose that we wait until we
is usually safer and cheaper to apply preventative observe disease to apply fungicide. The best case
(before symptoms appear) applications instead of scenario is that no disease would occur or that it
curative (after symptoms are seen) applications. That would only occur once in the spring or once in the fall.
is because the fungicide rate required to cure a dis- That is unlikely, however. The most likely best case
ease outbreak is much greater than the rate required scenario would be disease observation followed by
to prevent one. Consider this example: two applications of fungicide at 3.7 ounces per 1000
We set our threshold level for disease damage on square feet 14 days apart in late spring and two
the putting greens at Any Valley Country Club at 0%. applications in early fall to cure the disease. That
The greens are creeping bentgrass (Agrostis stolon- would require a total commitment of 14.8 ounces per
ifera) and the most common disease that occurs is 1000 square feet (4.5 g m−2), or about 1.3 times more
dollar spot (Sclerotinia homeocarpa). Chlorothalonil pesticide than the preventative program. A worse
is a very popular contact fungicide used to control case curative scenario would be three applications in
dollar spot on creeping bentgrass putting greens. It the spring and three in the fall at either 3.7 ounces
is sold under the trade name of Daconil Ultrex (and per 1000 square feet, or a worse case still, 5.0
others). A preventative application of Daconil Ultrex ounces per 1000 square feet, with those applications
is 1.8 ounces per 1000 square feet (0.55 g m−2) at totaling 22.2 or 30.0 ounces of pesticide per 1000
7–21 day intervals, depending on the environmental square feet (6.8 or 9.2 g m−2), respectively. In the two
conditions. A curative rate is from 3.7 to 5.0 ounces worse cases, the curative program would require two
per 1000 square feet on a 14-day schedule. So the to three times more pesticide than the preventative
curative rate is two to three times greater than the program. So although a preventative fungicide pro-
preventative rate. gram appears to be a poor economic and environ-
Let us assume that we can usually manage dollar mental choice, in cases where a very low threshold
spot preventatively with applications at 14-day level is required, it is often a better alternative, both
intervals and that we have 6 weeks in the late spring economically and environmentally, than a curative
and 6 weeks in the early fall when conditions are program.

198 Chapter 13
insect to develop a program like the one you devel- highly conducive to the growth and reproduction
oped for dollar spot. Cutworms, for instance, like of the pathogen, disease will occur and spread
to live in burrows in the thatch so thatch control is rapidly.
a preventative measure. They also like to live in Single-tactic systems are not very effective for
aerification holes, so if aerification absolutely has controlling turfgrass predators. If single-tactic sys-
to be performed during the season that is condu- tems are effective, it is only on a temporary basis.
cive to cutworm damage, an insecticide applica- For that reason, pesticides only control the preda-
tion afterward will probably be required. If tor until the next outbreak, and genetic resistance
possible, postpone the aerification and save an only controls the predator until the predator
application. adapts. Fortunately, in the case of genetic resistance
The best time to apply an insecticide for cut- to a predator, it may take a considerable time for
worm control is in the late afternoon as the insects the predator to adapt. In other cases, the predator
hide during the day and feed at night. The fresh may adapt quickly or other predators may become
insecticide on the leaves when the cutworms begin dominant and more damaging. A combination of
to feed is very effective. The insects also tend to live effective management strategies is the best course
in surrounding areas and crawl onto the greens at of action for reducing predator damage. Adjusting
night. Therefore, if the green is treated, the green environmental conditions to favor the turfgrass
surrounds should also be treated. As I stated earlier, over the predator is a permanent improvement.
if you must use a pesticide, use it wisely. Mixing control strategies makes it substantially less
As the superintendent of Any Valley CC, you will likely that a predator will become resistant to a
also have to learn about summer patch and billbug, management plan.
and brown patch and white grubs. There will also Devising an integrated management plan that
be other predators of primary concern in your area permanently deters predator damage requires in
that you will have to deal with on Category 1, 2 or depth knowledge of both predator and turfgrass
3 turf. You will have to learn about them and species. It also requires constant observation and
develop economic and environmentally sound pro- good record keeping. Turfgrass managers can
grams to discourage them. Learning and devising improve a turfgrass site for characteristics such as
procedures to reduce predator damage takes time light and air movement. The manager can use
and effort. However, once the protocols have been resistant mixtures or blends of resistant turfgrass
determined, their execution becomes normal and species and cultivars. He/she can adjust irrigation,
reasonably easy. Good turfgrass attracts high num- fertilizer, mowing height, soil pH and many other
bers of predators. You must be knowledgeable and factors to manage the situation best for turfgrass
you must be vigilant. growth and least for predator growth. In order to
do so, he/she must be willing to learn about the
grass and the pest, to be especially vigilant and, in
13.6 Chapter Summary
some cases, very creative.
In order to survive, organisms either have to per-
form photosynthesis or obtain nourishment directly
or indirectly from an organism that does. Because Suggested Reading
they perform photosynthesis, turfgrass plants pro- Boulter, J.I., Boland, G.J. and Trevors, J.T. (2000) Com-
vide energy to a variety of different species, includ- post: a study of the development process and end-
ing insects and diseases. As a turfgrass stand gets product potential for suppression of turfgrass disease.
more dense and healthy, it provides more resources World Journal of Microbiology and Biotechnology 16,
for its predators. Insects sometimes seek dense, 115–134.
green turfgrass in which to lay their eggs so that Burpee, L.L. (1993) Integrated control of turfgrass dis-
their offspring can obtain the best nourishment. eases: research and reality. International Turfgrass
Society Research Journal 7, 80–86.
Healthy turf is usually more resistant to disease
Couch, H.B. (1995) Diseases of Turfgrasses, 3rd edn.
than unhealthy turf, but that does not necessarily Krieger Publishing, Malabar, Florida.
mean that a high pathogen population is not Fidanza, M.A. and Dernoeden, P.H. (1997) A review of
present. In most cases, it means that the turfgrass brown patch forecasting, pathogen detection, and
can resist disease in spite of a fairly high pathogen management strategies for turfgrasses. International
population. If environmental conditions become Turfgrass Society Research Journal 8, 863–874.

Managing Competition Between Turf and Pests 199

Mann, R.L. and Newell, A.J. (2005) A survey to deter- Watschke, T.L., Dernoeden, P.H. and Shetlar, D.J. (1994)
mine the incidence and severity of pests and dis- Managing Turfgrass Pests. CRC Press, Boca Raton,
eases on golf course putting greens in England, Florida.
Ireland, Scotland, and Wales. International Turfgrass
Society Research Journal 10, 224–229. Suggested Websites
Nelson, E.B. (1997) Microbial inoculants for the control of
turfgrass diseases. International Turfgrass Society CENTERE (2010) TurfFiles. Center for Turfgrass Envi-
Research Journal 8, 791–811. ronmental Research and Education (CENTERE),
Niemczyk, H.D. and Shetlar, D.J. (2000) Destructive Turf North Carolina State University Turfgrass Manage-
Insects. H.D.N. Books, Wooster, Ohio. ment Group. Available at: http://www.turffiles.ncsu.
Potter, D.A. (1991) Ecology and management of turfgrass edu/Default.aspx (accessed: 2 April 2010).
insects. Annual Review of Entomology 36, 383–406. EDIS (2010) Turf Diseases. EDIS (Electronic Data
Potter, D.A. (1993) Integrated insect management in Information Source), UF (University of Florida) IFAS
turfgrasses: prospects and problems. International (Institute of Food and Agricultural Sciences) Exten-
Turfgrass Society Research Journal 7, 69–79. sion. Available at: http://edis.ifas.ufl.edu/topic_turf_
Potter, D.A. (1998) Destructive Turfgrass Insects: Biology, diseases (accessed 2 April 2010).
Diagnosis, and Control. Ann Arbor Press, Chelsea, Iowa Waste Reduction Center (2010) Golf Course Pollu-
Michigan. tion Prevention Guide. Available as a pdf at: http://
Potter, D.A. (2005) Prospects for managing destructive www.iwrc.org/resources/index.cfm (accessed 12
turfgrass insects without protective chemicals. Interna- August 2010).
tional Turfgrass Society Research Journal 10, 42–54. Landscape America (2010) Lawn and Landscape Prob-
Raikes, C., Lepp, N.W. and Canaway, P.M. (1996) An lems. Available at: http://www.landscape-america.
integrated disease management (IDM) strategy for com/problems/problems_index.html (accessed: 2
winter sports turf. Journal of the Sports Turf Research April 2010).
Institute 72, 72–82. University of Georgia Cooperative Extension (2010)
Sachs, P.D. and Luff, R.T. (2002) Ecological Golf Course Turfgrass Diseases in Georgia: Identification and
Management. Ann Arbor Press, Chelsea, Michigan. Control. Available at: http://pubs.caes.uga.edu/
Vargas, J.M. Jr. (1994) Management of Turfgrass Dis- caespubs/pubcd/B1233/B1233.html (accessed: 2
eases, 2nd edn. CRC Press, Boca Raton, Florida. April 2010)

200 Chapter 13
 Making the Right Decisions
14

Key Terms
A patio is an outdoor sitting or dining area that is usually paved.
A pergola is an open-roof structure of rafters and cross members or latticework supported by columns.
The Turfgrass Transition Zone describes a geographic band in the USA where cool-season (C3) grass adaptation
meets warm-season (C4) grass adaptation. The zone is characterized as a climate suited to heat tolerant
cool-season grasses and cold tolerant warm-season grasses.

14.1 Learn to Use Your Knowledge

constructed to define it from the lawn. The existing
to Your Advantage
lawn was a cold-tolerant common bermudagrass
What if you were a lawn or landscape professional (Cynodon dactylon), which is used for most home
and you were asked to design a small turfgrass area lawns in the area. The home was in Stillwater,
purely for ornamental purposes? You would begin Oklahoma, which is in the United States Turfgrass
by asking questions of the customer to find out Transition Zone. The Turfgrass Transition Zone is
exactly what their preferences were. You would a geographic band that extends across the mid-
consider your options based on those customer southern USA at the border of the temperate cli-
expectations and formulate a plan for customer mate zone and the subtropical climate zone. Both
approval. As questions arose, you would answer warm-season (C4) grasses and cool-season (C3)
them. As problems were presented, you would solve grasses may survive the transition zone climate.
them. When the site was finished, you would either However, the cool-season grasses should be fairly
assume the maintenance or provide the customer well adapted to warm summers and the warm-
with a management plan to maintain the turfgrass season species have to be fairly well adapted to
at the expected level of aesthetic value. If the job cold winters. In addition to cold tolerant bermuda-
was done well, you would receive praise and refer- grass, good choices for home lawns in Stillwater
rals from your customer that could amount to a and in the transition zone are zoysiagrass (Japanese
significant amount of business. So let us see how lawngrass, Zoysia japonica) and tall fescue (Festuca
you might do that using an actual example of a arundinacea). However, those species are most
homeowner’s back lawn site. A partner and I seeded commonly used in light shade or partial shade in
the lawn for this project in the fall of 2006. Stillwater, and bermudagrass is by far the most
The assignment was to build a patio constructed common grass in the area. The bermudagrass culti-
of pavers covered by a pergola connected to an var or biotype used has to be more cold tolerant
existing wooden deck attached to the house by a than most bermudagrasses to survive the winters in
walkway also constructed of pavers. The patio was reasonably good condition. In this case, bermuda-
to be used as an entry to the small shed on the lawn grass was not an option. Because the species grows
and as a covered but open sitting area for use by so aggressively, it is a very tenacious weed in flower
the homeowners and visitors during nice weather. beds, and beds bordered by bermudagrass require
The existing small planting bed was to be extended considerable effort to be maintained bermudagrass
the full width of the backyard and a simple border free. For that reason, our customer insisted that we

©CAB International 2011. Turfgrass Physiology and Ecology (G. Bell) 201
use a species other than bermudagrass. Although the house to accumulate approximately in the mid-
zoysiagrass, tall fescue and other species are less dle of the lawn with nowhere to go. The bermuda-
aggressive and a better choice near flower beds, grass was surprisingly healthy in spite of the
they are rarely used because they are readily puddling that followed any major rain event. That
invaded by aggressive indigenous bermudagrass. was only because the climate in Stillwater is rea-
Consequently, our most difficult decision in this sonably dry at about 34 inches (86 cm) of precipita-
project was choosing an appropriate turfgrass for tion per year. Rainfall events are often short but
the small turfgrass area in the back lawn. We intense, sometimes with long periods (2 to 4 weeks)
needed a species that would not aggressively invade of no rain at all.
the planting beds but one that would resist ber- After thoroughly inspecting the site, recording
mudagrass encroachment. measurements and observing deficiencies and
potential problems, we determined a plan and com-
pleted a simple drawing (Fig. 14.2). The homeown-
14.2 Planning the Project
ers selected the materials to be used and the design
The back lawn of the residence was an area meas- for the pergola from several different options and
uring 2556 square feet (237 m2) and was com- construction began in the summer of 2006. We
pletely fenced with a 5-foot (1.5-m) high wood decided that the small section of lawn between the
privacy fence. The area consisted of a wood deck patio and the north fence would remain in
attached to the back of the house and two small bermudagrass because it did not border on any
planting beds near the back fence (Fig. 14.1).
A storage shed stood on the north side of the prop-
erty supported on treated wood timbers placed Before
over pea gravel about 8 inches (20 cm) deep. The
lawn consisted of a cold tolerant common ber- Planting beds
mudagrass. Drainage on the site was poor. Runoff Lawn
Wood deck
drained away from the west fence and away from
South

House
Shed

New lawn design

Planting beds
Lawn
Patio
South

Walkway
Wood deck
House
Shed

1 block = 1 foot = 30 cm
Fig. 14.1. Photo of the homeowner’s back lawn as
it looked before reconstruction. The photographer is Fig. 14.2. A drawing of the site before construction
standing near the south fence facing north. Notice the (above) and a drawing of the proposed changes
shed on the north side, the small planting bed on the (below). In the new design the back lawn has been
west and a small portion of the wood deck attached reduced from 2556 sf (237 m2) to 720 sf (67 m2). The
to the house supporting the hose caddy. A second small lawn will be easy to manage at near aesthetic
planting bed in the southwest corner cannot be seen. perfection with minimal inputs and labor-intensive hand
The visible planting bed is in the approximate middle of weeding practices. Drainage occurs under the borders
the west fence line. The lawn is common bermudagrass of the planting beds and the walkway. Water from
(Cynodon dactylon). Runoff drains away from the these drains and from lawn runoff collects under the
house and away from the west fence to accumulate patio and drains through a subsurface pipe along the
near the middle of the lawn. north fence east into the sewers at the street.

202 Chapter 14
planting beds except the area immediately behind We completed the drainage systems before mov-
the shed. That area was expected to remain in ing on with the project so that we would not have
mulch deficient of any plant material. However, we to contend with standing water during the rest of
needed to select a species other than bermudagrass the construction (Fig. 14.3). We used the same
for the center lawn that was now limited to a space principles to determine how to drain the lawn that
of 720 square feet (67 m2). were presented in Chapter 8. We made no attempt
to change the natural drainage of the site. We stud-
ied the situation and made decisions based on the
Facilitating drainage
simplest way to meet our objective. We designed
Drainage was improved by supporting all of the the system to capture water and channel it to the
new design elements on a bed of at least 12 inches lowest part of the site. We then determined a place
(30 cm) of pea gravel that drained into a subsurface for the water to go based on the natural drainage
drain pipe. The drain pipe extended along the and the options available to us. The street being
north fence line east and emptied into the sewers in adjacent to the lowest part of the homeowner’s
the concrete street at the front of the house. property was the natural area in which to drain the
Consequently, the large bed on the back fence water. The street also contained an elaborate sewer
drained into the gravel under its border then system to carry the water away. The only difficult
drained toward and under the patio. Water collect- part of the drainage construction was the ditch
ing in the middle of the lawn drained toward and that had to be dug to the road. There was enough
under the patio and water draining from the bed of a rise in the land between the back yard and the
near the house drained under the walkway toward slope to the street that the water would not drain
the patio. The gravel under the shed also drained naturally. Consequently, the ditch was the only
into the gravel under the patio and the patio gravel possibility for removing water from the lawn.
drained into the pipe and flowed toward the street. Notice also that we considered drainage a top pri-
Since the reconstruction of the back lawn, drainage ority for the site. Without proper drainage, the
is no longer a problem. backyard would never have been the site that the
We had some concern about using pea gravel homeowners wished it to be and much of our
as a base for the patio and walkway pavers. That effort would have been wasted. Drainage is always
is something that we had never attempted before a top priority.
and is not recommended. Normally a highly
compactible fine gravel mixture is used under
pavers to provide a solid base. That type of base,
however, has attributes much like compacted soil
has when compared with natural soil. Drainage
from highly compactible gravel is faster than
from compacted soil but the compactible gravel
holds more water than other types of gravel and
drains more slowly. Because the patio would be
in the lowest part of the yard, we wanted it to be
able to collect water underneath without heaving
resulting from swelling and contracting during
extremely wet or dry periods. We had to choose
between attempting the pea gravel option, which
we felt would be successful, or a more intricate
and complicated drainage plan. We chose the pea Fig. 14.3. The drainage system was the first part of
gravel and have not been disappointed. Drainage the project completed. Here the borders for the planting
beds have been installed over a deep gravel base.
in the lawn is excellent and there have been no
The excavation is complete for the patio, its borders
problems with the patio or walkway. We placed are installed and the site has been backfilled with pea
a geotextile mat across the top of the gravel gravel. The pergola has also been constructed and
before applying the sand in which we set the work on the lawn is just beginning. Cultivation has also
pavers. The mat keeps the sand from migrating begun and the results are noticeable in a strip beside
into the pea gravel. the large planting bed.

Making the Right Decisions 203

 Once the drainage was complete, we were free to Because of its excellent heat tolerance and its
begin the renovation of the lawn. During the reno- ability to form a deep root system compared with
vation, because of the way we planned the eleva- other cool-season grasses, tall fescue is the most
tion of the beds, patio and walkway, we were able popular shade grass in the Turfgrass Transition
to further facilitate good drainage by smoothing Zone. Therefore, many sod growers have a consist-
the lawn surface to eliminate small collection pock- ent demand for tall fescue sod. However, because
ets and by maintaining a slight but consistent slope tall fescue is, for all practical purposes, a bunch-
to the patio. There was a small amount of soil type grass, it does not form a good sod and gener-
redistribution during this project, but because the ally falls apart during cutting or during installation.
design improved upon existing patterns rather than Sod growers in the area must support tall fescue
changing them, no soil was added to or removed sod either by using nylon mesh netting or by
from the site. As we graded the lawn we were care- mixing it with Kentucky bluegrass. Most growers
ful not to allow small collection pockets to develop prefer the Kentucky bluegrass mix to nylon netting.
as the soil settled back into place. Normally we A small amount, 5 to 10%, of Kentucky bluegrass
would have rolled the surface to firm it and encour- is sufficient to provide good tall fescue sod han-
age settling, but in this case, it was not necessary. dling characteristics and the bluegrass is expected
Because bermudagrass is such a difficult species to to die during its first summer. However, in some
kill we had enough time between the initial cultiva- cases, the bluegrass does not die and in others actu-
tion of the site and seeding to allow the soil to settle ally it spreads in competition with the tall fescue.
sufficiently and to make fine adjustments to elimi- In Chapter 9 you learned about the NTEP (the
nate low spots. National Turfgrass Evaluation Program) in the
The turfgrass species that we chose for the back USA. Kentucky bluegrasses are tested in areas of
lawn was Kentucky bluegrass (Poa pratensis). This the transition zone as well as in areas where they
is normally a poor choice for a lawn in Stillwater are better adapted for their ability to tolerate heat,
but, in this case, it met the objectives of the home- drought and other conditions stressful for cool-
owner perfectly and provided what we believe was season grasses. By studying the NTEP results from
an excellent choice for the site. After 4 years in transition-zone locations, it was clear that a few
existence, the grass has exceeded our expectations. recently developed Kentucky bluegrass cultivars
When making choices for turfgrass selection or for tolerated transition-zone conditions very well. We
turfgrass maintenance, knowledge is very valuable. selectively studied both the Kentucky bluegrass
Consider all of your options, not just the popular NTEP trials in Stillwater and those that had been
ones. You might just know more than you think performed close by in Wichita, Kansas, and found
you know and you might just succeed where others three cultivars that demonstrated good potential
fail. Let us consider the thought process that we for use at our site. We ruled out other cool-season
used to select our grass. grasses except for tall fescue and all warm-season
grasses except for zoysiagrass. After screening, we
were left with three possible choices for our lawn,
Selecting the turfgrass
Kentucky bluegrass, tall fescue or zoysiagrass.
Kentucky bluegrass makes a beautiful ornamental Because of its persistence in tall fescue sod and the
grass, especially in a location like Stillwater positive results of some cultivars in recent NTEP
where the warm-season grasses are dormant for trials nearby, we considered Kentucky bluegrass to
5 to 6 months. If irrigated and fertilized suffi- be an equally viable choice compared with zoysia-
ciently, Kentucky bluegrass will remain green all grass and tall fescue. A description of our selection
year long in Stillwater and will be exceptionally process is presented later in this section.
pleasing during the springs and falls. Most As you may have noticed earlier, we designed the
Kentucky bluegrasses, however, do not tolerate site so that the small ornamental lawn was com-
the hot, sometimes dry, summers in Stillwater and pletely enclosed by planting beds or structures
except on sod farms, are simply not used. Because except for the end bordered by the south fence only
it spreads by rhizomes, Kentucky bluegrass is an (Figure 14.2). For that reason, bermudagrass
excellent sod-forming turfgrass and is used by sod encroachment could not occur unless seed or sprigs
farms in mixtures with tall fescue to improve sod were carried into the lawn by humans, birds or the
strength. wind. Although the neighbor’s lawn to the south

204 Chapter 14
was bermudagrass we did not fear encroachment bluegrass for the site. We would have preferred to
from that direction because the fence shaded a seed a blend of the three cultivars that we identified
small area on its north side for nearly the entire as having positive potential, but a blend was not
day. Most bermudagrasses have no adaptation for available at that time. Consequently, we seeded the
shade and require full sun for more than half of most conveniently available cultivar of the three. In
each day to grow and spread. So we felt that it was reality, the perfect plan is often not achievable. In
unlikely that bermudagrass would encroach from this case, a second, single cultivar option worked
the south neighbor’s lawn, but that if it did, it could out very well.
easily be removed by hand and a strip of mulch
placed along the fence to prevent further encroach-
14.3 Turfgrass Establishment
ment. We used one of its natural enemies, shade,
against the bermudagrass to restrict its encroach- Before the Kentucky bluegrass could be established,
ment. As you can see, the complete enclosure of the the bermudagrass on the main (center) lawn had to
lawn by other structures also had purpose. be removed. Simply removing the sod was not an
Once the attributes of the site and species adapta- option as the bermudagrass would readily grow
tion to the transition-zone location had been con- back from rhizomes. The existing lawn had to be
sidered to eliminate all but three candidates, it was killed, and killing bermudagrass is difficult.
time to consider the use and expectations of the site The first step in preparation for planting
to choose the best grass. The lawn was to be prima- Kentucky bluegrass was to apply a full rate of
rily ornamental. Therefore, we wanted the best glyphosate, a nonselective herbicide, to the existing
looking grass year-round. As zoysiagrass is dormant bermudagrass lawn. Glyphosate is sold as Roundup
for nearly half the year, it was eliminated, but not and under several other trade names. It is nonselec-
without discussion. Zoysiagrass has a very attrac- tive, meaning that it kills anything green and is
tive gold color in dormancy and could have been a available for direct sale to homeowners in the USA;
very nice choice. However, the homeowners pre- it is also a herbicide that is widely considered to be
ferred a green grass year-round to a gold one and very safe to use. However, a single application of
were willing to mow early in the spring and late in glyphosate or any other herbicide will not kill ber-
the fall to maintain a green grass, another impor- mudagrass completely, and as a 0% threshold for
tant consideration. Tall fescue is also a very attrac- bermudagrass encroachment was all that could be
tive grass but it has a more coarse texture and does tolerated, we had to make sure that the existing
not get quite as dense as Kentucky bluegrass. In stand was completely eliminated. For a second time
addition, it gets brown patch (Rhizoctonia solani) in this project, we used the natural adaptation of
disease every summer in Stillwater and develops a bermudagrass against itself. Bermudagrass grows
brownish tint for a few weeks of the year because of by both stolons and rhizomes. Although the stolons
the disease. If the brown patch infection is severe, it are usually killed by glyphosate, some of the rhi-
could require fungicide treatments, a possibility that zomes survive. Consequently, surviving rhizomes
we would like to eliminate. Kentucky bluegrass can eventually produce new plants and, with time, can
get rust (Puccinia spp.) in the spring and fall in completely replenish the stand. For that reason, at
Stillwater that gives it a brownish tint but N ferti- least two applications of glyphosate, and often
lizer reduces the damage and pesticides are not three, are needed to completely eliminate a ber-
necessary. It also gets a dangerous disease called mudagrass stand. Knowledge of the pest, in this
summer patch (Magnaporthe poae) which could kill case a turfgrass, is required to control it.
it, but our adapted cultivars appeared to have some Following the first glyphosate application, we
resistance to the disease. Other diseases are possible, continued a program to eliminate the existing ber-
but dangerous infection is rare and the possibilities mudagrass completely. We could have waited until
of infection were about the same for both spe- all of the bermudagrass was brown following the
cies. Insect damage is also a possibility for both first glyphosate application, but that was not neces-
species, but is fairly rare and does not differ between sary. Once the initial glyphosate application was
the two grasses. Because of the likelihood of occa- applied, we waited only long enough to observe the
sional fungicide applications (every 2 to 3 years) on first signs of decline to begin cultivation. Because
brown patch, but mostly because Kentucky blue- the site was small we used a simple garden cultiva-
grass has slightly more visual appeal, we chose tor to till the soil. If soil tests had revealed a need

Making the Right Decisions 205

for a pH adjustment or a need for macronutrients After mowing, the lawn was fertilized lightly and
we would have fertilized and tilled the nutrients, the fertilizer was irrigated in. After about three
sulfur or lime into the soil at this point. In this case, more weeks, the bluegrass was approaching full
however, the soil was in good condition and adjust- cover and we began a standard fertilizer mainte-
ments were not required. After tilling, we raked nance program.
and prepared the site just as we would for a seed
bed, but we did not apply seed. Instead we applied
Assessment of the establishment procedures
irrigation in the same manner that we would use on
a new seed bed. The homeowner applied a very Throughout the establishment process we did eve-
small amount of irrigation two to three times per rything “by the book”, so to speak, including time
day to keep the soil damp. Soon weeds began to of establishment. It is usually best to establish
germinate and bermudagrass began to sprout. cool-season grasses in the fall instead of the spring
A second application of glyphosate about 5 weeks because there is less weed competition. We seeded
after the first application killed the weeds and the our grass in mid September, which is a perfect time
bermudagrass. to establish a cool-season grass in the Stillwater
Once the weeds had died completely following area. However, we forgot to anticipate annual
the second glyphosate application (about 2 weeks bluegrass (Poa annua) germination and competi-
later) the site was raked a second time to fill small tion with the Kentucky bluegrass. Because these
depressions that had occurred during soil settling, two grasses are in the same genus they share
and the irrigation treatments were begun again. almost the same niche. Consequently, annual blue-
Following 3 weeks of irrigation treatments, few grass is a major competitor of Kentucky bluegrass.
weeds had germinated and no bermudagrass plants It seems that this particular lawn had a serious
were found. So glyphosate was applied a third amount of annual bluegrass seed just waiting for
time, this time as a spot spray. We waited 3 days to the right conditions to germinate, and we provided
be sure that glyphosate absorption had occurred, those conditions for it. Out of curiosity, we
and the seeding process began. checked the Kentucky bluegrass seed for annual
A final raking was applied to the site, the bluegrass contamination and found absolutely
Kentucky bluegrass seed was scattered, and the site none. The annual bluegrass was resident in the
lightly raked again to partially cover the seed. soil. We should have known.
Irrigation was applied as before, and about 2 Encroachment by weeds other than annual blue-
weeks later, bluegrass germination began to occur. grass was minor and the individual weeds were
We advised the homeowner to continue irrigation easily removed by hand. Once full cover had been
three times daily for a week after germination, and reached that fall, however, annual bluegrass
then to start applying less frequently, but in a accounted for about 30% of the stand. Except for
slightly greater amount. Irrigation was applied the visible presence of annual bluegrass the lawn
twice daily for the next week and once daily for was beautiful in the following spring. However, the
two more weeks. By that time, the plants had annual bluegrass died during the summer, leaving
developed reasonably good root systems so irriga- some unsightly holes in the lawn (Fig. 14.4). In
tion was applied on alternate days for 2 weeks and order to provide the Kentucky bluegrass with an
then twice a week for as long into the fall as mow- opportunity to fill in over the fall, winter and spring
ing was required. After that irrigation was prac- without annual bluegrass competition we chose to
ticed only during very dry periods until spring. apply a pre-emergence herbicide before annual
We wanted the mowing height to be 3 inches for bluegrass germination in the fall. As the following
the first season to provide sufficient photosynthetic summer approached, the Kentucky bluegrass had
leaf area for the turf to develop a strong root sys- filled in all but the small area where runoff drained
tem. Consequently, the first mowing occurred under the patio, and full cover was achieved
when the bluegrass reached a height of 4 inches (Fig. 14.5). At present, fall pre-emergence applica-
(10 cm). The mower blade was sharpened in prepa- tions are no longer necessary, and the Kentucky
ration for the first mowing to facilitate as clean a bluegrass has been quite reliable in successfully
cut as possible and to minimize the possibility that competing with the annual bluegrass. Only a small
the mower blade would grab young plants and pull amount of annual bluegrass encroachment occurs
them from the ground owing to their weak roots. each fall and this is easily hand weeded.

206 Chapter 14
 important component of cool-season grass estab-
lishment in the area.
We know that annual bluegrass germinates as
temperatures cool in the fall and the soil is suffi-
ciently damp for seeds to imbibe water. Therefore,
annual bluegrass germination is likely following
the sufficient amount of rainfall that occurs after
about mid September. In this case, we provided
perfect conditions for germination of annual blue-
grass as well as for our Kentucky bluegrass. The
only reason that we had more Kentucky bluegrass
than annual bluegrass germination is that there was
more Kentucky bluegrass seed present. How could
Fig. 14.4. The Kentucky bluegrass (Poa pratensis) we have done this differently?
lawn as it appeared in October 2007, one year after We know when annual bluegrass germinates. We
seeding. A pre-emergence herbicide has been applied also know approximately how long it will take for
to prevent the fall establishment of annual bluegrass
Kentucky bluegrass to germinate and establish.
(Poa annua). The open areas in the otherwise dense
Consequently, because there is a reasonably mild
canopy are where annual bluegrass existed and died
over the summer. fall season in the Stillwater area, we can establish
cool-season grasses in October and often in
November. In that case, we should have used the
same procedures that we did to remove the possi-
bility of bermudagrass encroachment. We should
have treated the area like a seed bed in mid-
September, allowed the annual bluegrass to germi-
nate, removed that annual bluegrass and then
seeded the Kentucky bluegrass. If we had followed
that procedure, we would have had full Kentucky
bluegrass cover in the first fall or early in the first
spring with very minimal annual bluegrass
encroachment. If we had thought things through a
little better, we would have known how to proceed.
Nonetheless, we learned from the mistake and we
won’t make that mistake again. In combination,
Fig. 14.5. The Kentucky bluegrass (Poa pratensis) knowledge and experience are great partners for
lawn in June 2008. The only area that has yet to fill in is
learning and for the retention of what we learn.
the area where runoff water drains under the patio. Fall
pre-emergence herbicides are no longer necessary to
control annual bluegrass (Poa annua), as the Kentucky 14.4 Making a Management Plan
bluegrass has proved to be an excellent competitor and
little encroachment of annual bluegrass has occurred By its third year, our Kentucky bluegrass lawn was
since full cover was achieved. outstanding and had exceeded our expectations. It
still looked great after surviving its third summer of
100°F (38°C) and higher temperatures (Fig. 14.6).
A lesson learned
As expected, the bluegrass remained green during
Although our procedures for establishment were the winter seasons, offering a nice contrast with the
generally correct we could have saved ourselves otherwise mostly brown and leafless landscape
some labor and a pesticide application by adjusting (Fig. 14.7). However, we did not achieve this
those procedures to minimize annual bluegrass success without a good management plan. It is espe-
encroachment. As you have been reminded cially difficult to manage a bermudagrass lawn in
throughout this text, establishment procedures and the front of a house and a Kentucky bluegrass lawn
maintenance procedures need to be finely adjusted at the back. The management procedures are very
to the target site. In this case, we overlooked an different for the two grasses and good protocols for

Making the Right Decisions 207

 front lawn to the back lawn, but what if the home-
owner forgets to do it? It seems as if it would be a
simple thing to remember but you have to think
that someday the homeowner will mow the front
lawn at 1.5 inches (38 mm), forget to raise the
height to 3 inches (76 mm) in the back and scalp
the bluegrass. That could result in severe damage
to the back lawn. To prevent that from happening
we recommended that the homeowner compromise
and use a consistent mowing height of 2.5 inches
(64 mm), the lowest height that we felt was com-
fortable for the Kentucky bluegrass. At that height,
the bluegrass has remained in good condition and
Fig. 14.6. The Kentucky bluegrass (Poa pratensis) the bermudagrass looks reasonably good. To prove
lawn as it looked near the end of the summer in August our point, the homeowner has already scalped the
2009. The turf has fully matured and it has developed a back lawn once while attempting to maintain at
strong root system. Without such a root system it could
two mowing heights. Fortunately, it was not in the
not survive the extended 100 °F (38 °C) Oklahoma
middle of the summer.
summers and still look like this.

Irrigation
We did not design the backyard to include an auto-
matic irrigation system. However, after irrigating
all of the separate beds and lawns at the back, as
well as those at the front, one at a time, for a year,
an irrigation system was installed. The homeown-
ers had always irrigated their bermudagrass lawn
once a week when necessary at about 1 inch
(25 mm) of irrigation, depending on the condi-
tions. We recommended that the same amount of
irrigation be applied to the Kentucky bluegrass
lawn, but to apply it twice a week instead of once
a week. We recommended that both grasses be
Fig. 14.7. The lawn as it looked in December 2009. irrigated twice a week at approximately 0.5 inches
The grass is the only thing that is green in this photo. (13 mm) of precipitation each event to maintain
The rest of the ornamental plants have turned brown consistency. We would not recommend that ber-
and the leaves fell from the trees long ago. The mudagrass be irrigated more often than twice a
Kentucky bluegrass (Poa pratensis) adds a nice splash week, but for bluegrass in Oklahoma, twice per
of green color to the otherwise brown landscape.
week is a requirement. In cooler climates, Kentucky
bluegrass will go dormant to protect itself from
both have to be considered and executed. Even drought. It can remain dormant for several weeks
practices as simple as mowing become complicated under cool conditions and survive. However, we
enough to cause a decline in one grass or the other. have found that when it goes dormant in our
region, it can only survive for a short time, 1 or 2
weeks, before it dies. After experiencing a long,
Mowing
hot summer, there is very little root system surviv-
Because Kentucky bluegrass is poorly adapted to ing. Consequently, frequent irrigation is required
the summer heat of Stillwater, we need to be careful to keep the bluegrass from wilting severely. As you
not to mow the turf too low. Neither do we really recall, we irrigate to the depth of the root system
want to mow the bermudagrass too high or it will on a cool-season grass in the summer. The grass is
look ragged and wavy. It is really no problem to not healthy enough to grow deep roots to search
adjust the mowing height on the mower from the for water at that time.

208 Chapter 14
 By installing an automatic irrigation system and 1000 square feet (24–36 kg ha−1) depending on
using it wisely the homeowners were able to save recent temperatures. Even warm-season grasses use
water. It was no longer necessary to irrigate during less fertilizer during hot summers. The front lawn
the day when the temperatures were higher, the was to receive less fertilizer when temperatures were
radiation was more intense and the wind was blow- extremely warm and the higher rate if the tempera-
ing. By setting the system to automatically irrigate ture had been more mild or close to normal. The
before sunrise when it was cool and calm, less irri- final fertilization in September was to be applied at
gation evaporated and less water was needed. The 1.0 pound N per 1000 square feet (49 kg ha−1) and
homeowners turn the irrigation system off follow- an optional application of 0.5 pounds N per 1000
ing a rainfall and do not activate it again unless no square feet (24 kg ha−1) and 1.0–2.0 pounds K per
precipitation occurs for several days. They also 1000 square feet (49–98 kg ha−1) could be applied in
adjust to using less water during cloudy periods and October to help avoid winter kill if desired. Based on
to using more water during windy periods. They soil tests, monthly fertilizer is applied in a 17:3:5
can also adjust by ET c (crop evapotranspiration) if ratio of N-P-K during normal monthly applications.
they desire, as the calculations for their area are The October fertilizer high in K has never been
available on the internet. They monitor their system applied, and although some winter kill occurs each
to make sure that the heads are adjusted properly so year, the lawn recovers rapidly. The original recom-
that they are not irrigating the street, driveway or mendations have proved to be acceptable and the
walkways and they observe and sometimes check front lawn is normally in very good condition. By
their distribution patterns periodically. The proper customizing applications by season instead of apply-
installation and use of an automatic irrigation sys- ing a straight 1.0 pound N per 1000 square feet
tem can most definitely save water compared with (49 kg ha−1) every month the homeowner saves 1.0
the use of manually operated sprinklers. Most pound N per 1000 square feet a year (49 kg ha−1)
homeowners and commercial landowners, however, with no loss of turfgrass quality.
require education in the adjustment and use of their The back lawn is fertilized differently from the
systems. You can help. front. The Kentucky bluegrass in the back is fer-
tilized specifically to encourage root growth.
Fertilizer is applied at 0.5 pounds N per 1000
Fertilization and cultivation
square feet (24 kg ha−1) in late February or early
Fertilization of a warm-season grass like bermuda- March following the first mowing, when the tem-
grass and of a cool-season grass like Kentucky perature has warmed and the grass has begun to
bluegrass differ substantially by season. grow. An application at that time when the tem-
Consequently, we had to develop different fertilizer perature is still too cool for rapid shoot growth
programs for the front lawn and the back lawn. tends to encourage root growth. Fertilizer is not
The bluegrass required less fertilizer than the ber- applied again until October when the air tem-
mudagrass and fewer applications. Bermudagrass perature cools and shoot growth begins to slow.
is a nitrogen-loving plant. It grows and performs The October application is followed by a
best when fertilized with substantial amounts of N. November application and a December applica-
Rates of 0.75 to 1.00 pounds N per 1000 square tion to encourage root growth during the winter.
feet per month (37–49 kg ha−1) are common fertili- Nitrogen is applied at 1.0 pound N per 1000
zation rates for bermudagrass in the Stillwater square feet (49 kg ha−1) in October and November
area. We begin fertilizing at greenup and stop and at 0.5 pounds N per 1000 square feet in
about 6 weeks before dormancy is expected. December. Air temperatures usually remain warm
Generally, fertilization would begin on approxi- enough for active root growth in Stillwater until
mately 1 April and end on 1 September, depending late December, and roots will continue to grow
on annual conditions. slowly over most of the winter as long as the
To improve efficient use of fertilizer we suggested grass remains green. Consequently, the fertilizer
that front lawn fertilization begin with 0.5 pounds schedule encourages healthy deep roots as the
N per 1000 square feet (24 kg ha−1) at greenup fol- bluegrass goes into summer.
lowed by 1.0 pound N per 1000 square feet (49 kg By managing the fertilizer and irrigation properly,
ha−1) in May and June. In July and August, fertilizer neither the front nor the back lawn requires aerifi-
was to be applied at 0.50 to 0.75 pounds N per cation, vertical mowing or additional cultivation.

Making the Right Decisions 209

Pre-emergence herbicides are applied to the ber- front or back are insignificant. This would proba-
mudagrass in the spring and fall, and yellow bly not be the case if the turf were managed at a
nutsedge (Cyperus esculentus) control and broad- high level like a golf course fairway or professional
leaf weed control are usually applied once per year athletic field would be.
typically as a spot spray instead of a broadcast
application. Yellow nutsedge is the major problem
14.4 Chapter Summary
in the back lawn and has required a herbicide
application, but weeds are generally removed by Hopefully, you have found this final chapter inter-
hand. By nature a Kentucky bluegrass canopy is esting as I described the thought process for the
extremely dense and severely limits weed encroach- establishment and maintenance of a particularly
ment. Once the annual bluegrass pressure was unusual turfgrass for my region. Considering alter-
relieved, the yellow nutsedge was the only weed natives and thinking through problems is reward-
that required herbicide and it was applied as a spot ing, not only in finding solutions but in practical
spray rather than a broadcast spray. The bluegrass and economic efficiency. Successful planning now
sometimes gets rust disease in the spring or fall often saves considerable time later.
when the weather remains cloudy and damp for In truth, I was very familiar with this particular
several days, a condition that rarely occurs. The job in all details because it was my lawn. My wife
infections have not been severe and could probably and I are the homeowners and my wife was my
be easily controlled with an additional fertilizer partner. Now we are working on the flower beds
application if desired. Other predator problems and I have become the partner.

210 Chapter 14
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226 References
Index

Page numbers in bold refer to illustrations and tables

abscisic acid (ABA) 55, 57 bacteria 150, 151, 185

absorbance 77, 78, 79 see also pathogens
acclimation 137, 139, 142–143 basins 125
adaptation 24–25, 33, 34, 123–124, 166, 176 bentgrass (Agrostis spp.) 25, 59, 72
adenosine diphosphate (ADP) 18, 19, 48, 49 creeping bentgrass (A. stolonifera)
adenosine triphosphate (ATP) color measurement 7
glycolysis role 45 differential adaptation 176
production 22, 47, 57 ecological population 165
proton pump fuelling 42 performance observation 6
solute pumping role 51 root mass 86, 122
synthesis 18, 48, 49, 50 seasonal spectral reflectance response 131
see also energy shade effect 84–85
aerification (aeration) spring health appearance 41
aggressive 148 bermudagrass (Cynodon dactylon)
explained 9, 72, 147 aggressiveness trait 35
methods 71, 72, 147–150, 151, 152 color measurement 7
requirement reasons 161 construction project use 201
seasonal 138 control 205
surface disruption calculation formula 149 dew formation 119
timing 147–148 dormancy 34
see also air; compaction fertilization 209
aggregates 70, 92–93, 147 management practices 64
air performance observation 6
injection 150 seasonal spectral reflectance response 131
movement species 25
improvement 21, 23 best looking 3, 4
insufficiency 86 best management practices 2, 3
maintenance 32 bicarbonate 31–32, 33
restriction 20, 21, 50–51, 82–83, 87 biocides 195–196
temperature 132 biology 2, 3, 4, 5, 7, 10
see also aerification; boundary layer biomass 63, 70, 169, 170, 172, 173, 174
Airfield System construction method 152, 153 black layer 150–151, 161
alkali 157–158 blends 171, 176, 194
allelopathy 178–179, 182, 184 see also mixtures
amino acids 48, 49 bluegrass (Poa spp.) 25
amendments 153–154 annual bluegrass (P. annua) 40, 41, 63,
see also fertilizers; nutrition 136–137, 182, 207
amylase 38 Kentucky bluegrass (P. pratensis) 34, 165, 204,
anatomy 30, 42 205, 207–208, 209
anions 89–90, 158 boron 159
anoxia 142 boundary layer 23, 27, 50–51
antibiotics, resistance 185 breeding programs 195
antioxidants 133 brown patch disease 63, 205
Any Valley Country Club 196–197 brushing 137
apoplast 42, 51, 52 buffalograss (Buchloe dactyloides) 25, 33, 34, 78, 131

227
buildings 21, 81 production 16, 105
see also air, movement, restriction; shade synthesis 16, 137, 138
bunkers 124 chloroplast 26, 27, 39
cholesterol precursors 48, 49
citric acid (Krebs) cycle 48, 49
C-S-R model, three-strategy model 179 clay 89, 90, 91, 92
calcium (Ca) 97, 104, 106, 159 climate 98, 166
calcium sulfate (Gypsum) 95 cohesion-tension theory 55
Calvin cycle (dark reaction) cold 33, 130, 139–144, 176
bundle sheath cell role 30–31, 35, 53 see also temperature
carbon dioxide importance 23 colligative properties 38, 52, 139, 140, 143
compounds 45, 46–47 see also osmosis
fuelling 16, 17, 22–23, 31 colloids 90, 94
process 19–20 color
Rubisco interaction 25–26 enzyme affinity 15, 77
canopies 77, 79, 86, 139 health indicator 77
see also shade; trees measurement 7
capillary action 10, 54, 72 photons 15
carbohydrates photosynthetically active radiation 22
conversion to sucrose 142 turf 77, 78, 101, 102, 130
efficient use encouragement 37 see also greenness; light
energy provision 57 communities, ecological 166–167
partitioning 43–44 compaction (compression)
production 136 causes 72, 90–91, 92, 154
storage 37, 132, 143 explained 9, 51, 91, 147
see also starch; sugars layer 149
carbon 22, 26, 31–33, 35, 93–94, 136 remedies 60, 72, 93, 94, 161
carbon dioxide (CO2) water drainage problems 126
availability 10, 57, 120 see also aerification
binding 26, 35 competition
compensation point 37 control 63
concentrations 31, 32–33, 35 inhibition 191
conversion to sugar 18 interspecific 176, 177–179, 191
deficiency 20 intraspecific 6, 82, 169–170, 182
fixing 37 management 175–186, 187–199
forms in water 31 maturity advantage 171
loss 33 pests 187–199
production 9 root 84, 86
seasonal requirement 17 species selection role 169–170, 171
water ratio 29–30 strategies 179–183
carotenoids 16, 137 see also trees; vegetation control; weeds
carpetgrass (Axonopus spp.) 25 compression see compaction
carrying capacity 165, 168, 170, 172, 173, 187 conduction 134–135
cation exchange capacity (CEC) 70, 89–90, 94, 155 conductivity, electrical 157, 158
cations, components 158 construction 2–3, 152–154, 157, 201–210
centipedegrass (Eremochla ophiuroides) 25 convection 134–135
chaperonins (heat shock proteins) 134 cooling 21, 57, 119, 134, 139
checklists 66, 67 see also conduction; convection; transpiration
chlorine excretion 158 crabgrass (Digitaria spp.) 63
chlorophyll crop coefficient (Kc) 116, 127
degradation 17 crowns 19, 143
energy carrier 16–17, 31 cryptochrome 80
light energy receptor 16 cultivars 68, 144, 171
photon absorption 15, 22 cultivation 183, 209–210
photon color affinity 15, 77 cutworm 197, 198, 199
photooxidation 141 cytokinins (plant hormone) 133

228 Index
dallisgrass (Paspalum dilatatum) 183 requirement 40
dark reaction see Calvin cycle sequestration processes 20
day length response 137–138 sinks 19, 30, 37, 38–40, 57
de-acclimation 143 see also root systems; seeds
decision-making 7, 11, 65, 95, 201–210 sources 19, 30, 38–40, 44, 57
dehydration 62, 140-1, 141–142, 143, 144 see also leaves; sucrose
denaturation 51, 134 storage 8, 40
density 169, 172, 173, 174, 178 transfer 9
deoxyribonucleic acid (DNA) 47, 106 transport 57
dew removal 197 see also adenosine triphosphate; glycolysis;
disease heat; light; photosynthesis
cause 63 environment 2–3, 95, 103, 105, 180
control 193–194, 197, 205 enzymes 7, 15, 26, 31, 38
damage 196 see also Rubisco
hot spots recording 192 equilibrium, dynamic 28–29, 134, 135, 165, 173
pressure 192 equipment 65, 67, 71, 112–115, 127, 137
research requirement 197 see also mowers
resistance 190 establishment 205–207
spread 171 evaporation 9, 10, 62, 119, 120
susceptibility 86, 138, 190, 193 see also evapotranspiration
see also pathogens evapotranspiration (ET)
disturbances 168–169, 174, 179 calculation 116, 120, 127–128, 209
see also drought; ice; mowing; traffic; weather; factors 118
wind speed irrigation frequency factor 121
ditches, sub-surface 126 percentage replacement 116
diversity 171, 176 process 115–117
dollar spot (Sclerotinia homeocarpa) 185, 193, 197 rate 119, 120
dominance 166, 171, 174, 176–177, 182 reference standard 116–117, 127
dormancy 34, 51, 180, 181 exchangeable sodium percentage (ESP) 157, 158
drainage 107–108, 124–127, 128, 152–153
drought
damage 180 Facilitation-Tolerance-Inhibition Model 179
heat stress relationship 136 fans 21, 139
physiological 158, 161 fatty acids 48, 49, 133, 141
preconditioning 120 feeding 95, 156
protection 57 see also fertilizers; nutrition
tolerance 34, 35, 116, 123, 124 fermentation 45
fertilizers
acidifying 95, 194
ecology 2, 3, 4, 6, 163–174, 175 applications frequency 103
ecophysiology 165 color response 101, 102
ecosystems 6, 167–168 decision-making 95
electron transport system 47, 48–51 degradation 100
elimination 170 effects 70, 71
see also self-thinning formulations 97, 144
endophytes 195 guidelines 98
see also fungi management 88–89
energy rates 209–210
cells 48, 106 recommendations 95, 98
components 37–38 release rates 100–102, 156
conversion 8, 12 seasonal 143, 209
extraction 44–45 stability promotion 172
loss 17, 51, 57 timing 99, 138, 144, 209–210
molecules 102 see also nitrogen; nutrients; nutrition; phosphorus;
photons 22 potassium; sulfur
redistribution 81–82 fescue (Fescue spp.) 25, 34, 63, 131, 201
release 57 fluid systems 51–52, 54, 197

Index 229
frost line 144 hydrophobicity 69, 152
fructans 8, 37, 143 hydroponics 89
fungi 62, 185, 195, 196, 197
see also pathogens
fungicides 176, 185, 198, 205 ice damage 142, 144
immaturity 172
information gathering 11, 191
genetics 7, 76, 174, 176, 194–195 Initial Floristic Competition model 179
germination 180, 181, 207 injury 60–61, 62, 66
glucose 8, 23, 37, 38, 49 see also healing; mowing
glycolysis 44–46, 47, 48 inoculation 68
glyphosate (Roundup) 152, 205–206 insecticides 176, 198
golf courses 2, 117, 124, 196 insects 138, 190, 192–193, 194, 196
see also putting greens inspections 124–125, 198
grasses integrated pest management
cool-season (C3) 24–35, 85, 130–131, (IPM) 10–11, 189–190, 199
135–137, 138–139 see also pesticides
see also bentgrass; bluegrass; ion exclusion 158
fescue; ryegrass iron (Fe) 16, 95, 104, 105, 144
warm-season (C4) 24–35, 130–131, irradiance 76–81
133–135, 141, 144 irrigation
see also bermudagrass; buffalograss; concerns, salt-affected sites 159–160
centipedegrass; St. Augustinegrass; design 108–115, 157
zoysiagrasses efficiency 115
gravel 153, 203 effluent water use 115, 123
grazing 59 frequency 84, 120–122, 206
greenness 130, 177 management 108–115, 117–118, 154–155
see also color recommendations 208–209
growth 12–13, 101, 169, 170, 172 seasonal 121, 122
see also root systems; shoot growth spray heads 112–115, 127
gypsum (calcium sulfate) 95, 160 system faults 115
wetting agents applications 152
withholding 122
habitat management 178 zones 111, 112, 113, 127
hail damage 178 see also water
healing 61–62, 86 isobutylidine diurea (IBDU) 86, 100
see also injury
health 41, 77, 84, 130, 192, 199
heat Japanese lawngrass (Zoysia japonica)
dissipation need 9 see zoysiagrasses
generation 136
production 17
shock proteins 134 knowledge 191–192, 201–202
sources 76 see also information
stress 130, 136, 138 Kranz anatomy 30–33, 35
tolerance 133, 176 Krebs cycle (citric acid cycle) 48, 49
water energy source 29
see also temperature
heme synthesis pathway 105 laboratories 96, 97, 157, 159–160
henbit (Lamium amplexicaule) 182 landscape 66, 69, 168–169
herbicides 40, 138, 175, 183–185, 205, 210 law of limiting factors 166, 170–171, 172
human activity 59, 168–169 law of minimum 165
see also traffic law of self-thinning 170, 172
human expectations 175–176 lawns 3, 202
humidity 55, 100, 105, 118–119 layering problems 138
hydathode 54 leaching 101, 155, 156, 157, 158, 160, 161
hydrogen peroxide 152 leaf fire measurement 123

230 Index
leaf spot 190 minerals 89, 92
leaves mistakes, management 168
energy sinks/sources 38, 39 mitochondria 26, 27, 47
salt glands 158 mixtures 171, 176–177, 194
tip damage 62, 63, 68 moisture data collection 128
vascular system 30 molecules, binding opportunity 15
visual quality 123 monitoring 115, 117, 138, 192, 194
light morphology 80
adaptation 33, 34 mowers 65, 71, 137, 197
bands 22, 75–76, 79 see also equipment
colors 14, 22, 76, 78, 79, 80 mowing
day length response 137–138 damage 59, 60–66, 208
distribution 14, 79 effect 63, 64, 82, 166, 167
energy 13, 17, 22 facilitation 69
interception reduction 137 frequency 65
management 93 height
penetration restriction 79 adjustment 87, 174
performance effect 35 comparison 82
quality 79, 80, 81, 85 effect 94, 138
see also shade low, avoidance 172, 173
reaction (z-scheme) 16–17, 22, 31, 35 variation 206
reflected 76–77, 78, 79 injury shock period 81–82
saturation 27–28 management 64–66
spectrum 13 methods 137
transmission 77, 79 reasons 71, 169, 172
wavelengths 13, 14, 22, 76 specialized 60
see also photosynthesis timing 197
lime 95 tolerance 73, 137
lipids 133, 141 mutation, genetic 76
loam 90, 93, 154–155
localized dry spots (LDS) 151–152, 161
NADPH (reduced form of nicotinamide adenine
dinucleotide phosphate) 17–18, 22,
macromolecules 41–42 45, 47, 48, 49
see also proteins; ribonucleic acid nanometer (nm) 13
macronutrients 88–89, 97–104 National Turfgrass Evaluation Program (NTEP) 204
see also nitrogen; phosphorus; potassium nematodes control 195–196
macropores 92, 93, 94 niche 167, 170, 171, 174, 177, 206
magnesium (Mg) 16, 97, 104, 106, 159 nitrogen (N)
malate 31, 48, 49 detrimental 85
management practices 58–73, 130–131, 144, 145 fertilization 97–102, 106, 183, 186, 209
see also aerification; irrigation; mowing; leaching 155, 156
topdressing loss propensity 100
manganese (Mn) 104–105 overapplication hazard 95
mapping 117, 128, 192 potassium interaction 105
mat 70 recommendations 34–35
maturity 172, 174, 183 release rates 100–102
membrane, cell 133, 141, 143 soil content reports 95
metabolism 7, 119–120, 139–140, 141 sources 97–98, 102
see also photosynthesis; physiology; respiration use efficiency 34–35
metals 89 see also fertilizers
see also micronutrients; sodium normalized difference vegetation index (NDVI) 6, 131
microbes 62, 68, 70, 188 nutrients
micronutrients 89, 95, 104 application 155–156, 161
see also calcium ; magnesium; sulfur availability limiting 95
microorganisms 54, 62, 190 deficiencies 97, 155, 156
micropores 92, 93, 94 interactions 105

Index 231
nutrients (continued) phosphoenolpyruvate (PEP) carboxylase 31, 32, 33, 45
loss 155 phosphoglycerate (PGA) 20, 22, 23, 26, 27, 31
management 93 phosphoglycolate (PGL) 26
overapplication hazard 95 phosphorus (P)
rankings 88–89 cold-protection qualities interference 143–144
retention 91, 155 fertilizer 102–103, 106, 144, 186
soil content reports 95, 97, 98 overapplication hazard 95, 183
solubility 93 recommendations 103
tissue concentrations 158–159 soil reports 97
see also oxygen; solutes photochrome 80–81, 82
nutrition 88–106, 156 see also pigments
see also fertilizers; nutrients photographs 192, 202
nutsedge 210 photoinhibition 144
photon flux 13, 14, 15, 22, 28, 79
photooxidation 141
observation 6–7, 117, 192–193 photorespiration 25–30, 35, 44, 57, 99, 136
organelles 26, 27, 37 photosynthates 37, 39
osmosis 38, 43, 52, 94, 140, 161 photosynthesis
overseeding 176–177 components 13–22
oxaloacetate 31, 48, 49 see also Calvin cycle; carbon dioxide; light; water
oxidation, chemical 18 decline 132, 133, 141
oxygen defined 7, 12
air percentage 25 disruption 62
assimilation equation 26 encouragement 12–13, 105
consumption 8 energy provision 57, 199
electron transport role 50 improvement 75–76
encouragement 106 light role 13–16, 78, 105
lack 142 limiting factors 133
nutrition role 94 maintenance 32–33
in water 29–30 material removal 60, 62
oxygen-evolving complex 18 pathways 22, 24, 33, 35
potential 63
process 9, 12–23
paints 15, 16, 78–79 promotion factors 172, 174
pathogens rate 5, 40, 137
as biological herbicides 184, 185 reduction 20
control 195 requirements 8, 33, 78, 86, 93, 178
population growth 6, 192 sinks 38, 40
prey adaptation 190 sources 37, 39
specificity 171 stalling 10
spread delay 197 types 8
see also bacteria; disease; fungi wind speed effect 120
Penman-Monteith model 116 see also Calvin cycle; light; z-scheme
pentose phosphate pathway 46–47, 57 photosynthetically active radiation (PAR) 13, 14, 22
perennial ryegrass (Lolium perenne) 25 photosystems 17, 22, 141
performance factors 6–7, 29–30, 33, 35 physiology 2, 3, 4, 5–6, 7, 80
peroxisome 26, 27 see also, metabolism; photosynthesis; respiration;
personnel management 66 transpiration; temperature
see also training phytochrome 79, 80–81
pesticides 175, 176, 177–178, 196–198 see also pigments
see also integrated pest management pigments 16, 77, 79, 80–81
pests 187–199 see also carotenoids
Pfr state 80, 81 pipes 110, 111–112, 125, 126, 127
pH treatments 152 planning 130, 199, 207
phloem 30, 40–43, 44, 51, 53, 57 plants
phosphates 45, 47, 48 characteristics 33–35, 59, 60, 62
see also NADPH classifications 179–180

232 Index
 hormones 133 growing medium performance 150, 161
needs diagnosis 1–11 oxygen role 5, 94, 105–106
populations 34, 165, 168, 171, 174 plant/animal differences 45
soil structure facilitators 92 poor, symptoms 5
see also grasses; leaves; vascular system process 37–51, 57
potassium (K) promotion 172, 174
application rates 103–104 seasonal 40
DNA/RNA synthesis 106 soil compaction role 72
fertilization timing 138 types 44, 48–51, 57
leaching 156 water uptake energy provision 70
membrane-regulated exchange 143 see also photorespiration
nitrogen interaction 105 Rhizoctonia solani 63
osmosis management 38 ribonucleic acid (RNA) 40–41, 47, 106
overapplication hazard 183 ribulose bisphosphate carboxylase oxygenase
pumps 56 see Rubisco
purposes 105 ribulose bisphosphate (RuBP) 20, 22, 26
sodium uptake interference 159 root systems
soil reports 97 building 120–121
stress relief qualities 103 competition 84, 86
utilization sufficiency 143–144 encouragement 87, 131
precipitation 115, 127 growing medium effect 150, 158
see also irrigation; rainfall growth 87, 121–122, 131
predators mass 122
characteristics 191 pressure 53–54
control 190–191, 193, 194, 195–196 state of health 99
management 188–195 temperature effect 85–86
/prey relationships 168, 183–184, 187–188 water absorption 53–54
resistance 190, 191, 194–195 water uptake 57
proteins 18–19, 40, 51, 134, 142–143 Rubisco (ribulose bisphosphate carboxylase oxygenase)
protons 18–19, 22, 42, 48, 51, 56 deficiencies 33
protoplasm 51, 52 role 20, 22, 25–26, 27, 29, 31, 35
pumps 18–19, 42, 51, 56 runoff 99, 100
putting greens 2–3, 69, 82, 83, 152 rust 194, 205
see also golf courses ryegrass (Lolium spp.) 25
pyruvate 31, 45, 46, 49

safety 66
radiance 13, 15, 17 St. Augustinegrass (Stenophrum
see also light secundatum) 25, 34, 124
radiation 13, 14, 22, 76, 118, 120 salt 89, 94–95, 156–161
rainfall 99, 107, 108 sand systems
see also precipitation advantages 92, 161
random model 188 characteristics 14–17, 91, 152, 153
reaction centres see photosystems construction 152–154, 157
record keeping 117, 194 core aerification 148
recuperative ability 73 design 152–154
reduction, chemical 18 drainage 124–125
reflectance 15, 77, 79, 117, 131 fertilization 155
reflection 76–77 particle size 89, 90, 153, 154
reproduction 171 problems 150–152
research 4–5 sustainability 154
Resource Ratio Model 179, 180 water inputs 161
respiration scales, balance, equilibrium 135
defined 7 science 3–4
encouragement 105 seashore paspalum (Paspalum
equation 8–9 vaginatum Swartz) 25, 160
explained 37 seasons 40, 41, 130, 144, 145, 172

Index 233
seeds 181, 182, 183, 184 sprinklers 112, 114–115
selection see also irrigation
competitive 169–170, 171, 182 stability 167–168, 171, 174
decisions 182, 185 starch 8, 37–38, 39, 143
information 144 stems 19, 40
knowledge importance 204–205 stolons 38
traits 73 stomates 38, 55–57
self-thinning 170, 172, 173 stress
sensors 117, 130 human interventions 59
shade measurement 44
adaptation 82 relief qualities, potassium 103
differences 81 resistant strategists 180
effect 81, 83, 84–86 seasonal 27, 29–30
elimination 93–94 susceptibility 63
light penetration 77 temperature 130, 131–136, 139–141
management 82, 83–85, 87 water deficit 121
problems 10 see also cold; compaction; drought; heat;
responses 80, 81 mowing; shade; soil,
stress, measurement 44 saturation
tolerance 85, 86 stroma 18
shoot growth 63, 81, 85, 86, 99 succulence 72
Siemen 157, 158 sucrose 23, 37, 38, 40, 42, 43
silt 89, 91, 92 see also sugars
single-tactic systems 10–11, 190–191, 199 sugars 18, 19, 37, 40, 46–47
sodium 94, 157, 158–159 see also glucose; sucrose
soil sulfur 95, 104, 106, 150, 151
acidity 70 summer 28, 29, 99, 122, 138–139
aeration 50–51, 91, 147–150 summer patch disease 205
attributes 89 survival 59, 61
classifications 91, 157–158, 161 sustainability 168, 174
fertile, appearance 69–70 symplast 42, 51, 52
moisture 121, 143
pH 95, 152, 194
problems 147 techniques journal 192
profile, sand putting green 69 temperature
saturation 9, 97, 98, 124, 127, 155 chemical reaction speed effect 70
sodicity 157–158, 159–160, 161, 204 climatic 130
structure 92–93, 159 daily 131, 132
temperature 132, 143 increase 29
test reports 95–97 influence 98–99, 100, 102
texture 90–92 management practices adjustment
water-holding capacity 91 need 19–20, 118, 144
see also aerification; clay; compaction; physiological effects 35, 99, 119,
sand systems; silt 131–132, 133, 136
solar spectrum categories 75–76 reduction, canopies 139
solutes 9, 19, 37, 38, 51, 140 tolerance 33–34, 133–134
see also potassium thatch 66–72, 92, 138, 153
species thykaloid 18, 22, 42
list 25 tillers (vegetative growth) 169, 172
number limitation 166, 167 tissue 42, 97, 158–159
spreading methods 169, 171 topdressing 60, 70, 71–72, 139, 152
see also grasses; plants; selection toxins 45, 159, 178–179, 184
spectral sensing 128, 130, 131 tracheids 52, 53
sports fields 124 traffic
spraying 6, 178 control 94, 138–139
see also irrigation damage 69, 72, 73, 138, 168

234 Index
 problems 59–60 water
sustainability effect 154 adhesion 10, 52, 55
tolerance 58, 86 amounts 128
see also compaction availability 110
training 65–66 cellular/intracellular 139–140
transpiration cohesion 10, 52, 55
activity 5 effluent 115, 123
components 51–57 energy source 29
control 62 expansion 140
cooling role 9, 51 flow friction factor 110, 111–112
defined 7 frequency 155
disruption 62 loss 62, 65
encouragement 105, 172, 174 management 107–128
explained 37 molecules motion 29
irrigation adjustment factor 121 movement 9–10, 52–53
likened to perspiration 133 oxidation 50
process 9–10 pH 31, 32
pumping mechanisms 51 potential 10, 42–43, 52, 55, 158
reduction 20, 63, 83 pressure 110, 127
regulation 55 sources 107, 109–110
water uptake effect 9–10, 70 splitting 18, 22
wind speed effect 120 transport regulation 55
see also evaoptranspiration uptake 9–10, 57, 70, 158
trees 21, 23, 84, 86, 178 use 34, 35, 37, 118–124
turf/turfgrasses, functional viscosity 141–142
characteristics 59, 60, 62 window 109, 127
turfgrass, defined 58–59 see also irrigation; soil, saturation
Turfgrass Transition Zone, United States wear 72–73, 177
of America 201, 204 see also traffic
turgor (hydrostatic) pressure 38, 121, 123 weather 98
see also climate; temperature; wind speed
weeds
ubiquinol (UQH2) 47, 48, 49 accumulation (soil bank) 181
United States Golf Association (USGA) 2–3, 152, categories 63, 136–137, 179–180, 181, 182
153, 154, 157 competition management 40, 63, 179–183
United States Salinity Laboratory control 63, 64, 179–186, 204–205, 206, 210
(USSL) 157, 159–160 eradication 177–178
University of California Sand Putting Green identification 180
Construction and Management strategists 179–180
method 152, 153, 154 see also bluegrass (Poa spp.), annual
wilt 121, 123, 155
wind speed 21, 118, 120
vacuoles 37, 38, 51, 52
see also organelles
vapor pressure 28–29 xanthophylls 16, 77
vascular system 30, 30, 51, 52–53, 55, 140 xeriscape 61
see also phloem xylem 30, 51, 52–53, 55, 140
vegetation control 21, 23
see also shade; trees
vessels 52, 53 z-scheme (light reaction) 16–17, 22, 31, 35
viruses 190 see also electron transport system
volatilization 100 zoysiagrasses 25, 33, 34, 117, 119, 124, 201

Index 235