Ecophysiology

of
Vascular
Halophytes

Author
Irwin A. Ungar, Ph.D.
Professor
Department of Botany
Ohio University
Athens, Ohio

ypt CRC Press

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 PREFACE
The present volume represents the results of my research interests in the study of
halophytes over the last thirty years. Much of my personal research has been devoted to the
study of the responses of inland halophytes of North America to environmental stress. A
number of researchers throughout the world have made important contributions to our un­
derstanding of the physiological ecology of coastal halophytes and their research efforts have
contributed significantly in the preparation of this manuscript.
In recent years there has been an increase in research activity concerning the physiological
ecology of halophytes, which has been stimulated by a number of significant problems that
have developed because of the current usage of our land resources. The natural coastal and
inland salt marsh habitats have been strongly impacted by human activity and efforts are
being made to conserve existing salt marshes and in the creation of new salt marshes.
Knowledge of the physiology and ecology of marsh species is essential to carry out these
revegetation projects and for better management of salt marsh habitats. Irrigation of farmland
in drier regions of the world has caused the salinization of large areas, gas and oil well
drilling has produced large amounts of brine, and salting of roads in cooler climates has led
to the development of either saline soils or loss of vegetation. The activities of people living
in coastal areas of the world have caused major damage to salt marsh habitats and resulted
in the disappearance of salt marshes in many regions of the world.
The need for more food to feed our increasing population has stimulated a concerted
effort to find halophytes which are suitable for direct use as good for the human population
or as feed for livestock. Efforts have also been directed toward determining the physiological
and genetic mechanisms responsible for salt tolerance in plants. The more salt tolerant
halophytes could serve as donors for genetic material that would be utilized in the devel­
opment of new salt tolerant races for our most important agricultural plants. Although coastal
and inland salt marshes and salt deserts are currently used for grazing animals and some
harvesting of species is done for food, greater agricultural use may need to be made of these
lands in the future to meet the growing demands on currently used agricultural land to feed
the world's increasing population of people.
The main objective of this book is to review the literature which has been published
over the last two decades concerning the ecophysiology of halopytes. The topics covered
in this book range from the physiological responses of plants to salt stress and other edaphic
factors at different stages of development to the effect of competition on the growth and
distribution of halophytes. Other reviews in the past have either contained a very general
discussion of halophyte biology or emphasized one limited portion of the problem that
halophytes must respond to in adjusting to environmental conditions in saline habitats. In
this survey of the literature an attempt is made to discuss how the physical and biotic
environment may be influencing the physiological and ecological responses of halophytes.
I am sincerely indebted to my colleagues and graduate students who have worked
diligently in the area of halophyte ecology and provided much of the data reported on in
this book. My special thanks to Ohio University for providing me with a Faculty Fellowship
and John Baker research award to complete and revise the manuscript for this publication.
A number of researchers have provided me with encouragement and inspiration to continue
my research on the ecology of halophytes, including Paul Binet, Jean Boucaud, Jacques
Langlois, Olubukanla Ok us any a, Frank Drysdale, Stanley Woodell, Yoav Waisel, and Ve­
lentine Chapman. I wish to thank my former graduate students for their exchange of ideas
and the dedicated work they have carried out to make this publication more complete. A
number of the graduate students that I have worked with, including William Hogan, Allan
Macke, Karen McMahon, David Benner, David McGraw, David Loveland, Willa Ewing,
Barbara Wertis, Jeyarany Philipupillai, Donald Drake, Terrence Riehl, Hamid Karimi, Ajmal
Khan, Marlis Rahman, Kemuel Badger, Tonya Selby, Tira Katcmbe, and Barbara Ballard,
have stimulated my continuing interest in the ecology of halophytes and made significant
contributions to our understanding of the physiological ecology of halophytes.
The manuscript was completed while the author was a Visiting Fellow at Wolfson College
and Visiting Professor in the Laboratory of Plant Ecology, Department of Plant Sciences,
Oxford University. I thank Roger Hall for providing me space in his laboratory.
I am most sincerely appreciative of the support that my parents, wife and children have
given me through the years in my pursuit of research and during the preparation of the
manuscript for this book.
 AUTHOR
Irwin A. Ungar, Ph.D., is Professor of Botany in the Department of Botany, College
of Arts and Sciences, at Ohio University, Athens, Ohio. He received his Bachelor of Science
degree from the City College of New York in 1955 and his M.A. and Ph.D. degrees from
the University of Kansas in 1957 and 1961, respectively. In 1961 he was an Instructor in
the Department of Botany, University of Rhode Island and from 1962 to 1966 an Assistant
Professor in the Department of Biology at Quincy College. Since 1966, Dr. Ungar has held
successive appointments as Assistant Professor of Botany, Associate Professor of Botany,
and Professor of Botany in the Department of Botany at Ohio University. He was Chairman
of the Department of Botany from 1984 to 1989.
For the 1972 academic year, Dr. Ungar was a Visiting Professor and Chercheur Associe
in the Laboratoire de Physiologie Vegetale at the University of Caen and in the 1990 academic
year he was a Visiting Fellow at WoJfson College and a Visiting Professor in the Department
of Plant Sciences at Oxford University. Dr. Ungar is a fellow of the Ohio Academy of
Science and a member of the Ecological Society of America, Botanical Society of America,
and the American Association for the Advancement of Science. He is the author of over 80
publications and has presented his research findings at numerous national and international
meetings. His major research interest is in the effect of stress on the seed germination,
growth, and distribution of plants, with special emphasis in the ecology of halophytes. In
addition to his research interests, he is very active in both undergraduate and graduate student
teaching, Director of the Dysart Woods Laboratory since 1985, and other committee activities
at Ohio University.
 TABLE OF CONTENTS

Chapter 1
Introduction 1
I. Significance of Saline Habitats 1
II. Literature Review 1
III. Outline of Chapters Included in This Book 3
References 6

Chapter 2
Seed Germination 9
I. Introduction 9
II. Salt Tolerance 9
III. Population Variation 14
IV. Osmotic Effects 18
V. Specific Ion Toxicity 20
VI. Ionic Content of Seeds 23
VII. Dormancy 26
VIII. Growth Regulator — Salinity Interactions 28
IX. Seed Bank 31
X. Parental Effects 38
XL Conclusions 42
References 43

Chapter 3
Growth 49
I. Introduction 49
II. Effect of Salinity on Plant Growth 49
III. Growth Responses of Species of Atriplex to Salinity 51
IV. Salt Stimulation of Growth in Salicornia and Suaeda 53
V. Ionic Effects and Interactions with Other Factors 57
VI. Comparative Investigations with Halophyte Taxa 61
VIL Mycorrhizal Fungi 64
VIII. Phenology 65
IX. Conclusions 68
References 69

Chapter 4
Photosynthesis 73
I. Introduction 73
II. Effects of Salinity on Assimilation 73
III. Biochemical Inhibition 79
IV. Induction of Change in Carboxylation Pathway 80
V. Stable Carbon Isotope Ratios 81
VI. Field Investigations 83
VIL Conclusions 84
References 84

Chapter 5
Growth Regulators 87
L Introduction 87
II. Gibberellins.... 87
III. Cytokinins 91
IV. Auxin and Ethylene 91
V. Conclusions 92
References 92

Chapter 6
Plant Water Status 95
I. Introduction 95
II. Field Measurements of Plant Water Potential 95
III. Plant Water Status Responses to Salt Stress 101
IV. Conclusions 104
References 105

Chapter 7
Plant Ionic Content 107
I. Introduction 107
II. Ionic Content of Plants under Field Conditions 107
III. Seasonal and Daily Change in Ionic Content 114
IV. Ionic Content of Plants in Response to Salinity 120
V. Change in Ionic Content with Leaf Age 131
VI. Conclusions 134
References 134

Chapter 8
Waterlogging 139
I. Introduction 139
II. The Effect of Flooding on Plant Distribution and Growth 139
III. Tolerance to Waterlogging and Related Factors 142
IV. Conclusions 151
References 151

Chapter 9
Nitrogen 155
I. Introduction 155
II. Nitrogen Fertilization Experiments in Salt Marshes 155
III. Nitrogen and Halophyte Growth 159
IV. Nitrogen Fertilization Effects on Spartina alterniflora 163
V. Conclusions 169
References 169

Chapter 10
Genecology 171
I. Introduction 171
II. Ecotypic Differentiation 171
III. Height Forms in Spartina alterniflora 174
IV. Origin of Microspecies 180
V. Conclusions 182
References 182
Chapter 11
Competition 185
I. Introduction 185
II. Interspecific Competition 185
III. Intraspecific Competition 193
IV. Conclusions 198
References 198

Index 201
 1

Chapter 1

INTRODUCTION

I. SIGNIFICANCE OF SALINE HABITATS

The salt marsh habitat represents a very significant natural resource. Inland and coastal
salt marshes contain specific flora and fauna that are not found in other freshwater wetlands
or terrestrial plant communities.1*2 Coastal salt marshes play a significant role in food chains
of marine animals, and their disturbance could severely alter animal communities. Anthro­
pogenic activities have severely impacted these salt marsh ecosystems with pollutants and
damage from construction. Because of developmental activities, many natural salt marshes
have been disturbed or completely destroyed due to industrial development or other activities
of people living in coastal areas of the world. A number of efforts have been made to
preserve and reconstruct salt marsh communities, using the dominant halophytic species for
their revegetation.3-4
Because of the need for continually increasing our food supply to feed the expanding
world population, significant efforts have been made to develop species of plants that can
be grown for agricultural purposes in saline lands.5 8 First, attempts were made to select
cultivars of common crop species that demonstrated some degree of salt tolerance. 910 Second,
attempts have been made to use native halophytes either as a direct source of food or as
feed for farm animals. Several reviews have been written describing the multiple uses that
we are currenty making of halophytes for agricultural purposes. 1113 Many inland and coastal
salt marsh and salt desert habitats have been utilized for the grazing of cattle and sheep,
and the vegetation of some of these marshes has been greatly modified by the effects of
long-term grazing.14*16 In arid regions of the world, the continuous irrigation of agricultural
fields has led to the development of saline soils. Evaporation of irrigation water has produced
an accumulation of salts on the soil surface and made large areas which previously did not
have a soil salinity problem useless for farming.5"7

II. LITERATURE REVIEW

My primary goal for writing this book is to present a review of the current data about
the contributions of physiological ecology to our understanding of the ecology of halophytes.
A significant research effort, which has been made in recent years in the area of environmental
plant physiology, has contributed strongly to our understanding of how plants are responding
to saline environments. Both field research and laboratory investigations have been carried
out to ascertain the factors which are significantly affecting the seed germination, growth,
and reproductive responses of halophytes under environmental conditions that are prevalent
in coastal and inland salt marshes.
Earlier literature relating to the biology of halophytes has been described in considerable
detail by several researchers.17 2I A number of recent reviews are also available concerning
mangrove vegetation,22'27 and no attempt was made to review the literature concerning these
broadly distributed tropical coastal forest species in this book. Many publications have
described the distinct zonational patterns of plant species that occur in salt marsh habitats.2*28 *'
Efforts have been made to determine specifically which environmental factors are limiting
the growth and distribution of halophytes along soil salinity-flooding gradients.41"15 The
physiological and biochemical responses of plants to salinity stress have been reviewed by
a number of researchers in recent years.8-46-62
Several earlier comprehensive reviews have been written concerning the biology of
2 Ecophysiology of Vascular Halophytes

halophytes. 215 * 28 In recent years a number of reviews have concentrated on the functional
responses of plants to some aspect of salt stress.5960-63"66 The emphasis in the latter publi­
cations was on the physiological and biochemical responses of plants to salt stress. In my
analysis of physiological and ecological data, the main thrust will be to describe and clarify
some of the adaptations that species of plants have developed, which permit them to become
established and to grow and reproduce in saline environments. Successful establishment of
plants in salt marsh habits, which are subjected to a number of deleterious factors in their
physical environment such as flooding, salt stress, tidal abrasion and deposition, nutritional
ion imbalance, oxygen deficiencies, reducing environments, specific ion toxicities, and low
water potentials, is dependent on a number of physiological, morphological, and phenological
adaptations that make halophytic species successful in these stressful and unpredictable
habitats. Other abiotic and biotic factors, as well as combinations of the above-mentioned
factors, may also compound the problem that plant species must adjust to in order to become
successfully integrated in the salt marsh habitat.
The floristic composition of most plant communities that occur in salt marshes is low
when compared with adjacent vegetation from nonsaline habitats. Reduced diversity of plant
species in salt marsh and salt desert communities is correlated with the fact that many
terrestrial and aquatic vascular plant species do not have the necessary adaptations that would
provide them with a significant degree of salt tolerance.6*8 When we discuss the salt tolerance
of halophytes, we usually recognize that a group of biological characteristics have apparently
evolved in diverse groups of vascular plants that are necessary for their successful estab­
lishment in saline environments. No species may have developed all of the following char­
acteristics, but at least some of them are necessary adaptations for plants that are found
growing in these highly saline and unpredictable habitats: osmotic adjustment at the cellular
level, ion compartmentation, succulence, salt glands, salt hairs, ionic exclusion at the root,
tolerance to the dominant specific ions as well as tolerance to heavy metals, poorly aerated
soils, reducing environments, and low nitrogen availability — common problems that can
also be found in other nonsaline wetland ecosystems.
In this book I will use an ecophysiological approach to attempt to elucidate the factors
affecting growth at the different developmental stages and distribution of halophytes in saline
environments. It is clear that salt tolerance is one of the primary factors determining whether
or not plants can grow in salt marsh habitats. Intolerant glycophytes are not present in saline
environments, and most salt marsh and salt desert habitats generally contain a low diversity
of vascular plant species. Salt tolerance involves both cellular tolerance to salt stress and
whole organism phenomena, such as plant water balance and ion transport.2-50 The results
of investigations with salt tolerance of cell cultures often do not correlate well with the data
for salt tolerance of mature plants and their growth and distribution in field situations.67-68
A holistic approach is necessary to determine the nature of the responses of plants to
saline habitats. Besides the tolerance of species of plants to salinity, which is necessary for
the successful establishment of halophytes in saline habitats, a complex of biotic69*72 and
abiotic factors228-30'73"76 interacts to determine the level of biomass production, reproductive
output, and distribution of species along environmental gradients in salt marshes. Biotic
factors that are involved in limiting the growth of halophytes include herbivory, parasitism,
and both intraspecific and interspecific competition.69-72 The degree to which these biotic
factors are affecting the growth and distribution of halophytes varies in different zonal
communities and from one year to the next on any one marsh. Climatic and edaphic factors
also play a significant role in determining the success of halophytes in a particular habitat.
During a year of drought, mortality of plants is generally high because of the low soil-water
potentials that develop in marsh soils. 7 2 7 3 Seasonal changes in precipitation pattern must
also be adjusted to by those plants which are successful invaders of salt marsh environments.
Low soil-water potentials are produced in saline soils because of low levels of precipitation
 3

and high evaporation rates in areas of the world that are dominated by grassland and desert
plant communities. Unpredictable precipitation patterns from one year to the next can cause
dramatic changes in the zonational pattern of plants growing in inland salt marsh habitats.74*73
Edaphic conditions, such as soil-water potential, soil salinity, flooding, aeration, sulfide
accumulation, specific ion effects, and nutritional deficiencies, all play a role in determining
the physiological status of halophytes. The productivity and distribution of plants that are
found growing in saline environments are closely related to soil edaphic conditions. One of
the main tasks in the study of the ecophysiology of halophytes in the future will be to sort
out the relative significance of these solitary and interacting factors and the role they play
in determining the growth responses of halophytes at different stages of development. In
this book, I will attempt to describe from data available in the current literature some of the
general responses of halophytes to the various factors prevalent in saline habitats and how
they interact to determine the production and distribution patterns that currently exist in salt
marsh habitats.

III. OUTLINE OF CHAPTERS INCLUDED IN THIS BOOK

The chapters in this book contain a review of the recent literature concerning the eco­
physiology of vascular halophytes. Chapter 2 contains a discussion of the responses of seeds
to salinity and the variety of germination characteristics that halophytes have evolved to
survive the problem of salinity stress. Recent investigations with seed germination indicate
that the seeds of halophytes have developed a number of adaptations that distinguish them
from glycophytes. These include a general tolerance of seeds to low water potentials which
is greater than that for most glycophytes, various dormancy mechanisms which may serve
to avoid salt stress, and the capacity of seeds to resist extended periods of exposure to
hypersaline conditions and then continue to germinate when the salinity stress is removed.
Germination of halophytes often takes place when salinity levels are reduced under field
conditions. A delay of germination to periods of reduced soil salinity is of adaptive value
because it assures that at least some of the seedlings produced by a seed cohort will become
reproducing plants. Germination for most of the annual species of halophytes occurs over
an extended period of time, and some species form a persistent seed bank. Multiple ger­
mination periods play a significant role in preventing the local extinction of species in saline
environments, since they permit several periods for the establishment of plants in the highly
stressful and unpredictable salt marsh habitats.
Chapter 3 provides some details concerning the growth responses of halophytes to salinity
stress under both field and experimental laboratory conditions. Most halophytic species
examined vary in their degree of salt tolerance at different stages of development. Increases
in salinity from that in control nutrient solutions caused a reduction in growth for a number
of species of halophytes. However, the biomass production of some species is enhanced by
micronutrient levels of NaCl and other species have their growth stimulated by up to 200
mA/ NaCl. It is sometimes difficult to predict the distribution of halophytes based on lab­
oratory experiments that are used to determine their physiological limits of tolerance. Other
biotic and abiotic environmental factors may play a significant role in determining the biomass
production, fecundity, and survival of species in salt marsh habitats. Tolerance to salinity
may either represent the ability of a species to adapt to the general effects of low water
potentials or to the adjustment of plants to the presence of a specific ion. The occurrence
of specific ion toxicities, which are caused by some combination of specific ions, have been
reported in experimental investigations with halophytes.
Chapter 4 represents a discussion of the effect of salinity on the photosynthetic responses
of halophytes. Much of the early work concerning the carboxylation rates of halophytes was
done under laboratory conditions. However, in recent years with the development of portable
4 Ecophysiology of Vascular Halophytes

infrared gas analyzers, it has been possible to make diurnal and seasonal measurements of
the net photosynthetic rates of individual leaves, plants, and communities under field con­
ditions. Halophytes can generally tolerate higher levels of salinity than glycophytes, and for
some of the most salt tolerant species, there is apparently no significant reduction in their
net photosynthetic rates, even at the highest salinities examined. Several factors contribute
to the reduction in net photosynthetic rates of halophytes that are exposed to salinity stress.
These include reduced stomatal conductance, increased mesophyll resistance caused by a
decrease in carboxylating enzyme or its substrate, and inhibition of thylakoid membranes.
Induction of the CAM pathway has been reported for some species of succulents when they
are exposed to saline conditions, while other species are apparently not stimulated to change
their photosynthetic pathway when exposed to salt stress.
Chapter 5 contains a discussion of our current knowledgeregardingthe effect of salinity
on the growth regulator activity of halophytes. Several researchers have determined that
exogenous applications of gibberellic acid could overcome the inhibitory effects of salinity
on the growth of plants. However, growth regulator treatments were not found to be effective
at the highest salinities to which plants were exposed. Cytokinin levels have been reported
to decrease when plants were exposed to salt stress. Reductions in cytokinin levels could
negatively affect the rate of cell division and also cause closure of the stomatal apparatus,
inhibiting both the growth and net primary production of plants. Recently, it has been reported
that ethylene production is enhanced when plants are exposed to increases in salt stress and
that this effect is in response to low water potentials rather than to a specific ion toxicity.
Increases in the water potential of the growth medium caused a reversal in the level of
ethylene production.
Chapter 6 contains a review of the literature regarding the adjustment of plant water
status to salinity stress. Halophytes must be capable of adjusting the water potentials of their
tissues to that of the fluctuating soil water potentials in order to become successfully estab­
lished in saline environments. The development of the Scholander bomb and hygrometric
techniques for analyzing plant water status has allowedresearchersto make field as well as
laboratory measurements of plant water potentials and osmotic potentials. Halophytes are
able to adjust osmoticaJly to increasing levels of soil salinity, and the capacity to reduce
their tissue water potential is a determining factor in where they can grow and survive along
a soil salinity gradient. The two major mechanisms for the adjustment of water potentials
of plants arerepresentedeither by a reduction in tissue water content or by the accumulation
of inorganic ions. Other adaptations which control the level of salts in tissues include
succulence, salt glands, salt hairs, exclusion from roots, and abscission of organs.
Chapter 7 provides a review of the literature pertaining to field and laboratory investi­
gationsregardingthe ionic content of plants under a variety of soil salinity and experimental
conditions. Ash content of some species of halophytes reaches more than 50% of their dry
mass yields, indicating that a number of highly salt-tolerant species are able to accumulate
high ionic concentrations. A number of investigations have been made to determine the
specific ion content of plant organs. Some species of halophytes accumulate high concen­
trations of Na and Cl in their tissues, while other species are able torestrictthe uptake of
these ions. Experimental data indicate that most species of halophytic-dicotyledons respond
with an increase in tissue ion content when the soil salinity level increases. X-ray probe
microanalysis has been used to determine the relative content of elements in different tissue
regions and for different parts of plant cells. Data using these techniques indicate that plants
can compartmentalize ions among organs and tissues and within cells. Uptake of Na is
effected by the salinity of the medium, but it is also affected by the presence of Ca and
other cations in the soil solution. Salt tolerance may be in part regulated by the tolerance
level of species to high concentrations of salt, compartmentation of ions, and production of
compatible solutes to regulate the water potential of different cell compartments.
 5

Chapter 8 contains a discussion of the tolerance of halophytes to flooding conditions in

salt marshes. Tidal action effects plants by two major mechanisms: first, there is a mechanical
effect, where deposition and removal of silt by tides causes mechanical damage to plants
and uproots individuals. Second, there is the problem of tidal flooding at periodic intervals.
In inland salt marshes, plants are exposed to high water tables and periods of flooding after
heavy rainfall. Several soil factors are correlated with flooding, including low soil-oxygen
concentration, reducing environments, reduced nutrient availability, high sulfide content,
and heavy metal accumulation. Variation in the level of aeration of soils along a salt marsh
environmental gradient may play a significant role in determining the biomass production,
survival, and distribution of halophytes. A number of adaptations are necessary for plants
to carry out aerobic root respiration under flooding conditions. Some species of plants produce
large amounts of aerenchyma tissue, which allows oxygen to be transported from above­
ground organs to the roots. Transfer of oxygen from the roots to the soil may actually
increase the aeration level of soils and improve the availability of soil nutrients to plants.
Chapter 9 provides a review of the recent literature relating to the effect of nitrogen
availability on plant growth in saline habitats. Several researchers have reported that the
availability of nitrogen is one of the primary factors limiting the growth of halophytes in
salt marshes. It has been determined that the growth of halophytes could vary in different
areas of a salt marsh, even though the nitrogen content of the soil was similar in the two
areas investigated. A number of other soil factors, including high soil salinity, lack of
aeration, sulfide accumulation, and the presence of heavy metals, could inhibit the nitrogen
available to plants in salt marsh habitats, even when the soil nitrogen content did not vary.
Field fertilization experiments indicated that the fertilization of salt marsh soils with nitrogen
would increase the biomass production of plants growing in salt marsh habitats. Laboratory
studies have demonstrated that nitrogen fertilization could overcome some of the inhibitory
effects of salinity on the production of plant biomass.
Chapter 10 discusses the prevalence of ecotypes in salt marsh and salt desert habitats.
The occurrence of genetically distinct races has been reported for plants growing in saline
habitats. Greater salinity tolerance has been demonstrated for populations found in the more
saline part of a marsh gradient than for populations growing in areas of lower salinity. Two
major types of ecotypic differentiation appear to occur. First, broadly distributed species of
halophytes in coastal and inland habitats apparently develop genetically distinct ecotypes
along climatic gradients, in which the plants demonstrate different patterns of phenological
development. Second, one finds ecotypic differentiation and the development of microspecies
along salinity gradients in local marshes. Reciprocal transplant and uniform garden exper­
iments both indicate that genetic selection occurs in isolated plant populations on salt marshes.
In some cases, cleistogamy has developed as the primary breeding system, which further
increases inbreeding and the reproductive isolation of populations. Because of the repro­
ductive or geographic isolation of these populations from other populations on a salt marsh,
selection for microspecies will occur.
Chapter 11 contains a review of the available literature concerning the effect of com­
petition on the growth and distribution of halophytes. Both intraspecific and interspecific
competition have been reported to play a significant role in determining the biomass pro­
duction of plants, population size, and distribution of halophytes in saline habitats. Inter-
specific competition for space, nutrients, and light has been suggested to be the primary
factor limiting the growth of halophytes in nonsaline habitats. Competition also could restrict
species from zones on a salt marsh that are less saline than the habitat in which a species
dominates. Removal and transplant experiments in the field indicate that biomass production
of the target species increased when other species were removed from a plot. Intraspecific
competition has been shown to reduce plant biomass production in salt marsh habitats. In
saline environments, competition has been demonstrated to cause a decrease in yields and
6 Ecophysiology of Vascular Halophytes

to inhibit the reproductive capacity of species, but it has not been shown to effect survival.
This is probably because salt stress induces very high mortality in extreme habitats, and
competition plays little or no role in determining the level of survival of species in the most
stressful environments where salt stress is the prevailing factor controlling the mortality of
plants.
In this book I will review the literature on the ecophysiology of vascular halophytes.
Emphasis will be placed on the data available from publications concerning the inland and
coastal salt marsh vegetation over the past 20 years.

REFERENCES
1. Chapman, V. J., Ecosystems of the World 7, West Coastal Ecosystems, Elsevier, Amsterdam, 1977.
2. Waisel, Y., Biology of Halophytes, Academic Press, New York, 1972.
3. Cole, D. P., The restoration of coastal vegetation in Florida, Proc. SthAnnu. Conf., Tampa, Florida, 1978.
4. Niesen, T. and Josselyn, M., The Hayward regional shoreline marsh restoration: biological succession
during the first year following dike removal, Tiburon Center Environ. Stud. Tech. Rep., 1, 1, 1981.
5. Boyko, H., Salinity and Aridity, Junk Publishers, The Hague, 1966.
6. Hoftlaender, A., Aller, J. C , Epstein, E., San Pietro, A., and Zaborsky, O. R., The Biosaline Concept,
Plenum Press. New York, 1979.
7. San Pietro, A., Biosaline Research. A Look to The Future, Plenum Press, New York, 1982.
8. Staples, R. C. and Toenniessen, G. H., Salinity Tolerance in Plants, John Wiley & Sons, New York,
1984.
9. Shannon, M. C , Breeding, selection, and the genetics of salt tolerance, in Salinity Tolerance in Plants,
Staples, R. C. and Toenniessen, G. H., Eds., John Wiley & Sons, New York, 1984, Chap. 13.
10. Ponnamperuma, F. N., Role of cultivar tolerance in increasing rice production on saline lands, in Salinity
Tolerance in Plants, Staples, R. C. and Toenniessen, G. H., Eds., John Wiley & Sons, New York, 1984,
Chap. 14.
IK Mudie, P. J., The potential economic uses of halophytes, in Ecology of Halophytes, Reimold, R. J. and
Queen, W. H., Eds., Academic Press, New York, 1974, 465.
12. Somers, G. F., Food and economic plants: a review, in Biosaline Research, A Look to the Future, San
Pietro, A., Ed., Plenum Press, New York, 1982, 127,
13. O'Leary, J. W., The role of halophytes in irrigated agriculture, in Salinity Tolerance in Plants, Staples,
R. C. and Toenniessen, G. H., Eds., John Wiley & Sons, New York, 1984, Chap. 16.
14. Zahran, M. A. and Abdel Wahid, A* A., Halophytes and human welfare, in Contributions to the Ecology
of Halophytes, Sen, D. N. and Rajpurohit, K. S., Eds., Junk Publishers, The Hague, 1982, 233.
15. Chapman, V. J., Salt Marshes and Salt Deserts of the World, Interscience, New York, 1960, Chap. 12.
16. Tiedemann, A. R.( McArthur, E. D., Stutz, H. C , Stevens, R., and Johnson, K. L., Eds., Symposium
on the Biology of Atriplcx and Related Chenopods, Tech. Rep. I NT-172, Forest Service, U.S. Department
of Agriculture, Ogden, UT, 1984.
17. UphorT, J. C. T., Halophytes, Bot. Rev., 7, 1, 1941.
18. Stacker, O., Das Halophytenproblem, Ergebnisse BioL. 3, 26S, 1928.
19. Chapman, V. J., The new perspective in the halophytes, Q. Rev. Btoi, 17, 291, 1942.
20. Adrian!, M. J., Der Wasserhaushalt der Halophyten, in Handbuch der Pflanzenphysiologie, Band III
Pflanze und Wasser, Ruhland. W., Ed., Springer-Vertag, Berlin, 1956, 902.
21. Adrian!, M. J., Halophyten, in Encyclopedia of Plant Physiology, Vol. 4, Ruhland. W., Ed., Springcr-
Verlag, Berlin. 1958, 709.
22. Chapman, V. J., Mangrove Vegetation, J. Cramer, Vaduz, Liechtenstein, 1976.
23. Teas, H. J., Physiology and Management of Mangroves, Junk Publishers, The Hague, 1984.
24. Tomlinson, P. B., The Botany of Mangroves. Cambridge University Press, Cambridge, 1986.
25. Hutchings, P. and Saenger, P., Ecology of Mangroves, University of Queensland Press, St. Lucia, 1987.
26. Walsh, G. E., Mangroves: a review, in Ecology of Halophytes, Reimold, R. J. and Queen, W. H., Eds.,
Academic Press, New York, 1974. 51.
27. Barth, H., The biogeography of mangroves, in Contributions to the Ecology of Halophytes, Sen, D. N.
and Rajpurohit, K. S , Eds., Junk Publishers, The Hague, 1982. Chap. 3.
28. Chapman, V. J., Salt Marshes and Salt Deserts of the World, J. Cramer, Lehrc, 1974.
 7

29. Ward, J. M., Studies in ecology on a shell barrier beach. I. Physiography and vegetation of shell barrier
beaches, Vegetation. 14, 240, 1967.
30. Ranwell, D. S., Ecology of Salt Marshes and Sand Dunes, Chapman and Hall, London, 1972.
31. Ungar, I. A., Inland halophytes of the United States, in Ecology ofHalophytes, Reimold, R. J. and Queen,
W. H., Eds., Academic Press, New York, 1974, 235.
32. Macdonald, K. and Barbour, M. G., Beach and salt marsh vegetation of the North American Pacific
coast, in Ecology of Halophytes. Reimold, R. J. and Queen, W. H., Eds., Academic Press, New York,
1974, 175.
33. Ungar, I. A., The vegetation of inland saline marshes of North America, north of Mexico, in Grundfragen
und Methoden in der Pflanzensoziologie, Tuxen, R., Ed,, Junk Publishers, The Hague, 1972, 397.
34. Barbour, M. G., Craig, R. B., Drysdale, F. R., and Ghiselin, M. T., Coastal Ecology, Bodega Head.
University of California, Berkeley, 1973.
35. Eleuterius, L. N. and Eleuterius, C. K., Tide levels and salt marsh zonation, Bull. Mar, Sci.f 29, 394,
1979.
36. Earle, J. C. and Kershaw, K. A., Vegetation patterns in James Bay coastal marshes. 111. Salinity and
elevation as factors influencing plant zonations, Can. J. Bot., 67, 2967, 1989.
37. Kolbek, J., Dostalek, J., Jarolimek, I., Ostry, I., and Sek-Ha, L., On salt marsh vegetation in North
Korea, Folia Geobotan. Phytotaxonom., 24, 113, 1989.
38. Ellenberg, H., Sea marshes and inland saline habitats, in Vegetation Ecology of Central Europe, Cambridge
University Press, Cambridge, 1988, 348.
39. Binet, PM La Flore du Littoral, Penn Ar Bed, 25, 33, 1961.
40. Beeftink, W. G., The coastal salt marshes of western and northern Europe: an ecological and phytoso­
ciological approach, in Wet Coastal Ecosystems, Chapman, V. J., Ed., Elsevier, Amsterdam. 1977,
Chap. 6.
41. Chapman, V. J., Coastal Vegetation, Pergamon Press, Oxford, 1964.
42. Steiner, M., Zur Okologie der Salzmarschen der Nordostlichen Vereinigtcn Staaten von Nordamerika,
Jahrb. Wiss. Bot., 81, 94, 1934.
43. Johnson, D. S. and York, H. H., The relation of plants to tide-levels, Carnegie Inst. Wash. Pub., 206,
1, 1915.
44. Kurz, H. and Wagner, K., Tidal marshes of the Gulf and Atlantic coasts of northern Florida and Charleston,
South Carolina, Florida State Univ. Studies. 24, 1, 1957.
45. Miller, W. R. and Egler, F. E., Vegetation of the Wequetequock-Pawcatuck tidat marshes, Ecol. Monogr.,
20, 143, 1950.
46. Scrogonov, B. P., Structure and Function of Plant Cells in Saline Habitats, John Wiley & Sons, New
York, 1973.
47. Strogonov, B. P., Physiological Basis of Salt Tolerance of Plants, Israel Program for Scientific Translations,
Jerusalem, 1964.
48. PoUakoft-Mayber, A. and Gale, J., Plants in Saline Environments. Springer-Verlag, Berlin, 1975.
49. CaJdwell, M. M., Physiology of desert halophytes, in Ecology ofHalophytes, Reimold, R. J. and Queen,
W. H., Eds., Academic Press, New York, 1974, 355.
50. Green way, H. and Munns, R., Mechanisms of salt tolerance in nonhalophytes, Annu. Rev. Plant Physiol..
31, 149, 1980.
51. Albert, R., Halophyten, in Pflanzenokohgie und Mineralstoffwechsel. Kinzel, E., Ed., Verlag Eugen
Ulmer, Stuttgart, 1982, Chap. 3.
52. Bernstein, L. f Effects of salinity and sodicity on plant growth, Annu. Rev. Plant, Phytopathology, 13,
295, 1975.
53. Brun, AM Mise au point bibliographique concernant I'etude des effects de la salinite sur les vegetaux, Ann,
Fac. ScL Yaounde, 28, 59, 1981.
54. Binet, P., Aspects physiologiques de Thalophyte et de la resistance aux sels, Soc. Bot. Fr. Actualites Bot..
3/4, 1, 1978.
55. Flowers, T. J., Physiology of halophytes, Plant Soil. 89, 41, 1985.
56. Rowers, T. J., Hajibagheri, M. A., and Clipson, J. W., Halophytes, Q. Rev. BioL, 61, 313, 1986.
57. Flowers, T. J., Halophytes, in Ion Transport in Plant Cells and Tissues. Baker, D. A. and Hall, J. L.,
Eds., North-Holland, New York, 1975, Chap. 10.
58. Flowers, T. JM Troke, P. F., and Yeo, A. R., The mechanism of salt tolerance of halophytes, Annu.
Rev, Plant. Physiol,, 28, 89, 1977.
59. Munns, R., Green way, H., and Kirst, G. O., Halotolerant eukaryotes, in Encyclopedia of Plant Phys­
iology, Vol. 12C, Lange, 0 . L., Nobel, P. S., Osmond, C. B. t and Ziegler. H . Eds., Springer-Verlag,
Berlin, 1983, 59.
60. JefTeries, R. L., Osmotic adjustment and the response of halophytic plants to salinity, Bioscience, 31, 42,
1981.
8 Ecophysioiogy of Vascular Halophytes

61. Lerner, H. R., Adaptations to salinity at the plant cell level, Plans Soil. 89, 3, 1985.
62. Wyn-Jones, R. G., Salt tolerance, in Physiological Processes Limiting Plans Productivity, Johnson,
C. B. t Ed., Butterworths, London, 1981, Chap. 15.
63. Jefferies, R. L., Aspects of salt-marsh ecology with particular reference to inorganic mineral nutrition, in
The Estuarine Environment, Barnes, R S. K and Green, J., Eds., Applied Science Publishers, London,
1972. Chap. 5.
64. Jennings, D. H., The effects of sodium chloride on higher plants, Biol. Rtv . 51, 453, 1976.
65. Lauchli, A., Regulation des Salztransportcs und Salzausschiessung in Glykophyten und Halophyten, Ber.
Deutsch. Bot. Ges., 92, 87, 1987.
66. Jennings, D. H., Halophytes, succulence and sodium—a unified theory, New Phytoi, 67, 899, 1968.
67. Smith, M. K. and McComb, J . A., Effect of NaCl on the growth of whole plants and their corresponding
callus cultures, Ausu 7. Plant Physioi, 8, 267, 1981.
68. Hedenstrom, H. and BreckJe, S.-W., Obligate halophytes? A test with tissue culture methods, Z. Pflan­
zenphysioi, 74, 183, 1974.
69. Ellison, A. M., Effects of competition, disturbance, and herbivory on Salicomia europaea, Ecology. 68,
576, 1987.
70. Bertness, M. D., Wise, S., and Ellison, A. M., Consumer pressure and seed set in a salt marsh perennial
plant community, Oecologut, 71 t 190, 1987.
71. Bazely, D. R. and Jefferies, R. L., Changes in the composition and standing crop of salt-marsh communities
in response to the removal of a grazer, / . EcoL, 74, 693, 1986.
72. Ungar, I. A., Bcnner, D. K., and McGraw, D. C M The distribution and growth of Salicomia europaea
on an inland salt pan, Ecology, 60, 329, 1979.
73. Ungar, I. A., Population ecology of halophytc seeds, Bot. Rev., 53, 301, 1987.
74. Ungar, I. AM Population dynamics of inland halophytic communities, Bu//. Soc. Bot. Fr., 121,287, 1974.
75. Ungar, I. A., Population characteristics, growth, and survival of the halophyte Salicomia europaea.
Ecology, 68, 569, 1987.
76. Abdei-Razik, M. S. and Ismail, A. M. A., Vegetation composition of a maritime salt marsh in Qatar in
relation to edaphic features, J. Vegetation Scr, 1, 85, 1990.
Introduction
Chapman, V. J. , Ecosystems of the World 1. West Coastal Ecosystems, Elsevier, Amsterdam, 1977.
Waisel, Y. , Biology of Halophytes. Academic Press, New York, 1972.
Cole, D. P. , The restoration of coastal vegetation in Florida, Proc. 5th Annu. Conf., Tampa, Florida, 1978.
Niesen, T. and Josselyn, M. , The Hayward regional shoreline marsh restoration: biological succession during
the first year following dike removal, Tiburon Center Environ. Stud. Tech. Rep., 1, 1, 1981.
Boyko, H. , Salinity and Aridity, Junk Publishers, The Hague, 1966.
Hollaender, A. , Aller, J. C. , Epstein, E. , San Pietro, A. , and Zaborsky, O. R. , The Biosaline Concept, Plenum
Press, New York, 1979.
San Pietro, A. , Biosaline Research, A Look to The Future, Plenum Press, New York, 1982.
Staples, R. C. and Toenniessen, G. H. , Salinity Tolerance in Plants, John Wiley & Sons, New York, 1984.
Shannon, M. C. , Breeding, selection, and the genetics of salt tolerance, in Salinity Tolerance in Plants, Staples,
R. C. and Toenniessen, G. H. , Eds., John Wiley & Sons, New York, 1984, Chap. 13.
Ponnamperuma, F. N. , Role of cultivar tolerance in increasing rice production on saline lands, in Salinity
Tolerance in Plants, Staples, R. C. and Toenniessen, G. H. , Eds., John Wiley & Sons, New York, 1984, Chap.
14.
Mudie, P. J. , The potential economic uses of halophytes, in Ecology of Halophytes, Reimold, R. J. and Queen,
W. H. , Eds., Academic Press, New York, 1974, 465.
Somers, G. F. , Food and economic plants: a review, in Biosaline Research, A Look to the Future, San Pietro,
A. , Ed., Plenum Press, New York, 1982, 127.
O'Leary, J. W. , The role of halophytes in irrigated agriculture, in Salinity Tolerance in Plants, Staples, R. C. and
Toenniessen, G. H. , Eds., John Wiley & Sons, New York, 1984, Chap. 16.
Zahran, M. A. and Abdel Wahid, A. A. , Halophytes and human welfare, in Contributions to the Ecology of
Halophytes, Sen, D. N. and Rajpurohit, K. S. , Eds., Junk Publishers, The Hague, 1982, 233.
Chapman, V. J. , Salt Marshes and Salt Deserts of the World, Interscience, New York, 1960, Chap. 12.
Tiedemann, A. R. , McArthur, E. D. )., Stutz, H. C. , Stevens, R. , and Johnson, K. L. , Eds., Symposium on the
Biology of Atriplex and Related Chenopods, Tech. Rep. INT-172, Forest Service, U.S. Department of
Agriculture, Ogden, UT, 1984.
Uphoff, J. C. T. , Halophytes , Bot. Rev., 7, 1, 1941.
Stocker, O. , Das Halophytenproblem, Ergebnisse Biol., 3, 265, 1928.
Chapman, V. J. , The new perspective in the halophytes, Q. Rev. Biol., 17, 291, 1942.
Adriani, M. J. , Der Wasserhaushalt der Halophyten, in Handbuch der Pflanzenphysiologie, Band III. Pflanze
und Wasser, Ruhland, W. , Ed., Springer-Verlag, Berlin, 1956, 902.
Adriani, M. J. , Halophyten , in Encyclopedia of Plant Physiology, Vol. 4, Ruhland, W. , Ed., Springer-Verlag,
Berlin, 1938. 709.
Chapman, V. J. , Mangrove Vegetation, J. Cramer, Vaduz, Liechtenstein, 1976.
Teas, H. J. , Physiology and Management of Mangroves, Junk Publishers, The Hague, 1984.
Tomlinson, P. B. , The Botany of Mangroves, Cambridge University Press, Cambridge, 1986.
Hutchings, P. and Saenger, P. , Ecology of Mangroves, University of Queensland Press, St. Lucia, 1987.
Walsh, G. E. , Mangroves: a review, in Ecology of Halophytes, Reimold, R. J. and Queen, W. H. , Eds.,
Academic Press, New York, 1974, 51.
Barth, H. , The biogeography of mangroves, in Contributions to the Ecology of Halophytes, Sen, D. N. and
Rajpurohit, K. S. , Eds., Junk Publishers, The Hague, 1982, Chap. 3.
Chapman, V. J. , Salt Marshes and Salt Deserts of the World, J. Cramer, Lehre, 1974.
Ward, J. M. , Studies in ecology on a shell barrier beach. I. Physiography and vegetation of shell barrier
beaches, Vegetation. 14, 240, 1967.
Ranwell, D. S. , Ecology of Salt Marshes and Sand Dunes, Chapman and Hall, London, 1972.
Ungar, I. A. , Inland halophytes of the United States, in Ecology of Halophytes, Reimold, R.J. and Queen, W. H.
, Eds., Academic Press, New York, 1974, 235.
Macdonald, K. and Barbour, M. G. , Beach and salt marsh vegetation of the North American Pacific coast, in
Ecology of Halophytes, Reimold, R. J. and Queen, W. H. , Eds., Academic Press, New York, 1974, 175.
Ungar, I. A. , The vegetation of inland saline marshes of North America, north of Mexico, in Grundfragen und
Methoden in der Pflanzensoziologie, Tuxen, R. , Ed., Junk Publishers, The Hague, 1972, 397.
Barbour, M. G. , Craig, R. B. , Drysdale, F. R. , and Ghiselin, M. T. , Coastal Ecology, Bodega Head. University
of California, Berkeley, 1973.
Eleuterius, L. N. and Eleuterius, C. K. , Tide levels and salt marsh zonation, Bull. Mar. Sci., 29, 394, 1979.
Earle, J. C. and Kershaw, K. A. , Vegetation patterns in James Bay coastal marshes. III. Salinity and elevation
as factors influencing plant zonations, Can. J. Bot., 67, 2967, 1989.
Kolbek, J. , Dostalek, J. , Jarolimek, I. , Ostry, I. , and Sek-Ha, L. , On salt marsh vegetation in North Korea,
Folia Geobotan. Phytotaxonom., 24, 113, 1989.
Ellenberg, H. , Sea marshes and inland saline habitats, in Vegetation Ecology of Central Europe, Cambridge
University Press, Cambridge, 1988, 348.
Binet, P. , La Flore du Littoral, Penn Ar Bed, 25, 33, 1961.
Beeftink, W. G. , The coastal salt marshes of western and northern Europe: an ecological and phytoso-
ciological approach, in Wet Coastal Ecosystems, Chapman, V. J. , Ed., Elsevier, Amsterdam, 1977, Chap. 6.
Chapman, V. J. , Coastal Vegetation, Pergamon Press, Oxford, 1964.
Steiner, M. , Zur Okologie der Salzmarschen der Nordostlichen Vereinigten Staaten von Nordamerika, Jahrb.
Wiss. Bot., 81, 94, 1934.
Johnson, D. S. and York, H. H. , The relation of plants to tide-levels, Carnegie Inst. Wash. Pub., 206, 1, 1915.
Kurz, H. and Wagner, K. , Tidal marshes of the Gulf and Atlantic coasts of northern Florida and Charleston,
South Carolina, Florida State Univ. Studies, 24, 1, 1957.
Miller, W. R. and Egler, F. E. , Vegetation of the Wequetequock-Pawcatuck tidal marshes, Ecol. Monogr., 20,
143, 1950.
Scrogonov, B. P. , Structure and Function of Plant Cells in Saline Habitats, John Wiley & Sons, New York,
1973.
Strogonov, B. P. , Physiological Basis of Salt Tolerance of Plants, Israel Program for Scientific Translations,
Jerusalem, 1964.
Poljakoff-Mayber, A. and Gale, J. , Plants in Saline Environments, Springer-Verlag, Berlin, 1975.
Caldwell, M. M. , Physiology of desert halophytes, in Ecology of Halophytes, Reimold, R. J. and Queen, W. H. ,
Eds., Academic Press, New York, 1974, 355.
Greenway, H. and Munns, R. , Mechanisms of salt tolerance in nonhalophytes, Annu. Rev. Plant Physiol., 31,
149, 1980.
Albert, R. , Halophyten, in Pflanzenokologie und Mineralstoffwechsel, Kinzel, E. , Ed., Verlag Eugen Ulmer,
Stuttgart, 1982, Chap. 3.
Bernstein, L. , Effects of salinity and sodicity on plant growth, Annu. Rev. Plant. Phytopathology, 13, 295, 1975.
Brun, A. , Mise au point bibliographique concemant l’etude des effects de la salinite sur les vegetaux, Ann. Fac.
Sci. Yaounde, 28, 59, 1981.
Binet, P. , Aspects physiologiques de l’halophyte et de la resistance aux sels, Soc. Bot. Fr. Actualites Bot., 3/4,
1, 1978.
Flowers, T. J. , Physiology of halophytes, Plant Soil, 89, 41, 1985.
Flowers, T. J. , Hajibagheri, M. A. , and Clipson, J. W. , Halophytes , Q. Rev. Biol., 61, 313, 1986.
Flowers, T. J. , Halophytes , in Ion Transport in Plant Cells and Tissues, Baker, D. A. and Hall. J. L. , Eds.,
North-Holland, New York, 1975, Chap. 10.
Flowers, T. J. , Troke, P. F. , and Yeo, A. R. , The mechanism of salt tolerance of halophytes, Annu. Rev. Plant.
Physiol., 28, 89, 1977.
Munns, R. , Greenway, H. , and Kirst, G. O. , Halotolerant eukaryotes, in Encyclopedia of Plant Physiology, Vol.
12C, Lange, O. L. , Nobel, P. S. , Osmond, C. B. , and Ziegler, H. , Eds., Springer-Verlag, Berlin, 1983, 59.
Jefferies, R. L. , Osmotic adjustment and the response of halophytic plants to salinity, Bioscience, 31, 42, 1981.
Lerner, H. R. , Adaptations to salinity at the plant cell level, Plani Soil, 89, 3, 1985.
Wyn-Jones, R. G. , Salt tolerance, in Physiological Processes Limiting Plant Productivity, Johnson, C. B. , Ed.,
Butterworths, London, 1981, Chap. 15.
Jefferies, R. L. , Aspects of salt-marsh ecology with particular reference to inorganic mineral nutrition, in The
Estuarine Environment, Barnes, R. S. K. and Green, J. , Eds., Applied Science Publishers, London, 1972,
Chap. 5.
Jennings, D. H. , The effects of sodium chloride on higher plants, Biol. Rev., 51, 453, 1976.
Lauchli, A. , Regulation des Salztransportes und Salzausschiessung in Glykophyten und Halophyten, Ber.
Deutsch. Bot. Ges., 92, 87, 1987.
Jennings, D. H. , Halophytes, succulence and sodium—a unified theory, New Phytol., 67, 899, 1968.
Smith, M. K. and McComb, J. A. , Effect of NaCl on the growth of whole plants and their corresponding callus
cultures, Aust. J. Plant Physiol., 8, 267, 1981.
Hedenstrom, H. and Breckle, S.-W. , Obligate halophytes? A test with tissue culture methods, Z. Pflan-
zenphysiol, 74, 183, 1974.
Ellison, A. M. , Effects of competition, disturbance, and herbivory on Salicornia europaea, Ecology, 68, 576,
1987.
Bertness, M. D. , Wise, S. , and Ellison, A. M. , Consumer pressure and seed set in a salt marsh perennial plant
community, Oecologia, 71, 190, 1987.
Bazely, D. R. and Jefferies, R. L. , Changes in the composition and standing crop of salt-marsh communities in
response to the removal of a grazer, J. Ecol., 74, 693, 1986.
Ungar, I. A. , Benner, D. K. , and McGraw, D. C. , The distribution and growth of Salicornia europaea on an
inland salt pan, Ecology, 60, 329, 1979.
Ungar, I. A. , Population ecology of halophyte seeds, Bot. Rev., 53, 301, 1987.
Ungar, I. A. , Population dynamics of inland halophytic communities, Bull. Soc. Bot. Fr., 121, 287, 1974.
Ungar, I. A. , Population characteristics, growth, and survival of the halophyte Salicornia europaea, Ecology, 68,
569, 1987.
Abdel-Razik, M. S. and Ismail, A. M. A. , Vegetation composition of a maritime salt marsh in Qatar in relation to
edaphic features, J. Vegetation Sci., 1, 85, 1990.

Seed Germination
Ungar, I. A. , Population ecology of halophyte seeds, Bot. Rev., 53, 301, 1987.
Riehl, T. E. and Ungar, I. A. , Growth and ion accumulation in Salicornia europaea under saline field conditions,
Oecologia, 54, 193, 1982.
Riehl, T. E. and Ungar, I. A. , Growth, water potential, and ion accumulation in the inland halophyte Atriplex
triangularis under saline field conditions, Acta Oecol. Oecol.Plant., 4, 27, 1983.
Wertis, B. and Ungar, I. A. , Seed demography and seedling survival in a population of Atriplex triangularis
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Ungar, I. A. , Halophyte seed germination. Bot. Rev., 44, 233, 1978.
Ungar, I. A. , Germination ecology of halophytes, in Contributions to the Ecology of Halophytes, Sen, D. N. and
Rajpurohit, K. , Eds., Junk, The Hague, 1982, 143.
Workman, J. P. and West, N. E. , Germination of Eurotia lanata in relation to temperature and salinity, Ecology,
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Strogonov, B. P. , Physiological Basis of Salt Tolerance of Plants, Israel Program for SClentific Translation,
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Strogonov, B. P. , Structure and Function of Plant Cells in Saline Habitats, John Wiley & Sons, New York, 1973.
Ungar, I. A. , Inland halophytes of the United States, in Ecology of Halophytes, Reimold, R.J. and Queen, W. H.
, Eds., Academic Press, New York, 1974, 235.
Rozema, J. , The influence of salinity, inundation and temperature on the germination of some halophytes and
non-halophytes, Oecol. Plant., 10, 341, 1975.
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Scheele in a Georgia salt marsh, Am. J. Bot., 65, 502, 1978.
Blum, U. , Seneca, E. D. , and Stroud, L. M. , Photosynthesis and respiration of Spartina and Juncus salt
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Drake, B. G. , Light response characteristics of net C02 exchange in brackish wetland plant communities,
Oecologia. 63, 263, 1984.
Eckardt, F. E. , Dynamique de l’ecosysteme, strategie des vegetaux, et echanges gazeux: cas des enganes a
Salicornia fruticosa, Oecol. Plant., 7, 333, 1972.
Drake, B. G. , Photosynthesis of salt marsh species, Aquatic Bot., 34, 167, 1989.
McNulty, I. , The effect of salt concentration on the growth and metabolism of a succulent halophyte, in
Physiological Systems in Semiarid Environment, Hoff, C. and Riedesel, M. , Eds., University of New Mexico
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Martin, C. E. , Lubbers, A. E. , and Teeri, J. A. , Variability in crassulacean acid metabolism: a survey of North
Carolina succulent species, Bot. Gaz., 143, 491, 1982.

Growth Regulators
Waisel, Y. , Biology of Halophytes. Academic Press, New York, 1972.
Ungar, I. A. , Salt tolerance of plants growing in saline areas of Kansas and Oklahoma, Ecology. 47, 154, 1966.
Barbour, M. G. , Is any angiosperm an obligate halophyte?, Am. Midi. Nat., 84, 105, 1970.
Prisco, J. T. and O'Leary, J. W. , Enhancement of intact bean leaf senescence by NaCl salinity, Physiol. Plant.
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17, 187, 1968.
Boucaud, J. and Ungar, I. A. , Influence of hormonal treatments on the growth of two halophytic species of
Suaeda, Am. J. Bot., 63, 694, 1976.
Kirkham, M. B. , Gardner, W. R. , and Gerloff, G. C. , Internal water status of kinetin-treated plants, Plant
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Mizrahi, Y. , Blumenfeld, A. , and Richmond, A. E. , Abscisic acid and transpiration in leaves in relation to
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Ungar, I. A. , The effects of salinity and hormonal treatments on the growth and ion uptake of Salicornia
europaea, Soc. Bot. Fr. Actualites Bot., 3-4, 95, 1978.
Nieman, R. H. and Bernstein, L. , Interactive effects of gibberellic acid and salinity on the growth of beans, Am.
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O'Leary, J. W. and Prisco, J. T. , Response of osmotically stessed plants to growth regulators, Adv. Front. Plant
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Wochok, Z. S. and Sluis, C. J. , Gibberellic aCld promotes Atriplex shoot multiplication and elongation, Plant
Sci., Lett., 17, 363, 1980.
Kefu, Z. , Mingliang, L. , and Jiayao, L. , Reduction by GA, of NaCl-induced inhibition of growth and
development in Suaeda ussuriensis, Aust. J. Plant Physiol., 13, 547, 1986.
Huber, W. and Sankhla, N. , Eco-physiological studies on Indian arid zone plants. II, Effect of salinity and
gibberellin on the activity of the enzyme of amino-aCld metabolism in leaves of Pennisetum typhoides,
Oecologia. 13, 271. 1973.
Itai, C. , Response of Eucalyptus ocCldentalis to water stress induced by NaCl, Physiol. Plant. 43, 377, 1978.
Khan, M. I. , Khan, M. A. , and Khizar, T. , Plant growth regulators from species differing in salt tolerance as
affected by soil salinity. Plant Soil. 45, 267, 1976.
Khizar, T. and Khan, M. I. , Effect of temperature on the activity of growth inhibitor-A isolated from Suaeda
leaves, Biol. Plant., 19, 231, 1977.
Chrominski, A. , Weber, D. J. , and Smith, B. N. , NaCl-salinity-dependent conversion of ACC to ethylene in the
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Chrominski, A. , Bhat, R. B. , Weber, D. J. , and Smith, B. N. , Osmotic stress-dependent conversion of 1-
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Chrominski, A. , Khan, M. A. , Weber, D. J. , and Smith, B. N. , Ethylene and ethane production in response to
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Collins, J. C. and Kerrigan, A. P. , Hormonal control of ion movements in the plant root?, in Ion Transport in
Plants, Anderson, W. P. , Ed., Academic Press, New York, 1973, 589.

Plant Water Status

Berger, A. , L’ alimentation en eau en milieu sale, Soc. Bot. Fr. Actualites Bot., 3-4, 159, 1978.
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Munns, R. , Why measure osmotic adjustment?, Aust. J. Plant Physiol., 15, 717, 1988.
Jefferies, R. L. , Rudmik, T. , and Dillon, E. M. , Response of halophytes to high salinities and low water
potentials, Plant Physiol., 64, 989, 1979.
Glenn, E. P. and O'Leary J. W. , Relationship between salt accumulation and water content of dicotyledonous
halophytes. Plant Cell Environ., 7, 253, 1984.
Glenn, E. P. , Relationship between cation accumulation and water content of salt-tolerant grasses and a
sedge. Plant Cell Environ., 10, 205, 1987.
Chapman, V. J. , Salt Marshes and Salt Deserts of the World, J. Cramer, Bremerhaven, 1974.
Waisel, Y. , Biology of Halophytes, Academic Press, New York, 1972.
Osmond, C. B. , Bjorkman, O. , and Anderson, D. J. , Physiological Processes in Plant Ecology, Springer-
Verlag, Berlin, 1980.
Karimi, S. H. and Ungar, I. A. , Development of epidermal salt hairs in Atriplex triangularis Willd., in response to
salinity, light intensity, and aeration, Bot. Gaz., 150, 68, 1988.
Scholander, P. F. , Bradstreet, E. I. , Hammel, H. Τ. , and Hemmingsen, E. A. , Sap concentrations in
halophytes and some other plants. Plant Physiol., 41, 529, 1966.
Jefferies, R. L. , The ionic relations of seedlings of the halophyte Triglochin maritima L., In Ion Transport in
Plants, Anderson, W. P., Ed., Academic Press, New York, 1972, 297.
Kramer, D. , The possible role of transfer cells in the adaptation of plants to salinity, Physiol. Plant, 58, 549,
1983.
Dawson, Τ. E. and Bliss, L. C. , Species patterns, edaphic characteristics, and plant water potential in a high-
arctic brackish marsh, Can. J. Bot., 65, 863, 1987.
Langlois, J. , Evolution de la pression osmotique et des teneurs en eau, en Cl et Na’ au tours de la croissance
de l’axe principal chez Salicornia stricta Dumort, Rev. Gen. Bot., 75, 377, 1968.
Naidoo, G. , Seasonal plant water relations in a South African mangrove swamp. Aquatic Bot., 33, 87, 1989.
Romo, J. T. and Haferkamp, M. R. , Water relations of Artemisia tridentata spp. wyomingensis and Sarcobatus
vermiculatus in the steppe of southeastern Oregon, Am. Midi. Nat., 121, 155, 1989.
Drake, B. G. and Gallagher, J. L. , Osmotic potential and turgor maintenance in Spartina alterniflora Loisel.,
Oecologia. 62, 368, 1984.
Scholander, P. F. , Hammel, H. T. , Bradstreet, E. D. , and Hemmingsen, E. A. , Salt balance in mangroves,
Plant Physiol., 37, 722, 1962.
Scholander, P. F. , Hammel, H. T. , Bradstreet, E. D. , and Hemmingsen, E. A. , Sap pressure in vascular
plants, SClence, 148, 339, 1965.
Scholander, P. F. , How mangroves desalinate seawater, Physiol. Plant, 21, 251, 1968.
Ungar, I. A. , The relationship between soil water potential and plant water potential in two inland halophytes
under field conditions, Bot. Gaz., 138, 498, 1977.
Ungar, I. A. , Autecological studies with Atriplex triangularis in Symposium on the Biology of Atriplex and
Related Chenopods, Tiedemann, A. R. , McArthur, E. D. , Stutz, H. C. , Stevens, R. , and Johnson, K. L. , Eds.,
Gen. Tech. Rep. INT-172, Forest Service, U.S. Department of Agriculture, Ogden, UT, 1984, 40.
Riehl, T. E. and Ungar, I. A. , Growth and ion accumulation in Salicornia europaea under saline field conditions,
Oecologia, 54, 193, 1982.
Riehl, T. E. and Ungar, I. A. , Growth, water potential and ion accumulation in the inland halophyte Atriplex
triangularis under saline field conditions, Acta Oecol. Oecol. Plant, 4, 27, 1983.
Kaplan, A. and Gale, J. , Modification of the pressure-bomb technique for measurement of osmotic potential in
halophytes, /. Exp. Bot., 25, 663, 1974.
Ustin, S. L. , Pearcy, R. W. , and Bayer, D. E. , Plant water relations in a San FranClsco Bay salt marsh, Bot.
Gaz., 143, 368, 1982.
Antlfinger, A. E. and Dunn, E. L. , Water use and salt balance in three salt marsh succulents. Am. J. Bot., 70,
561, 1983.
Tyndall, R. W. , Teramura, A. H. , and Douglass, L. W. , Potential role of soil moisture defiClt in the distribution
of Cakile edentula. Can. J. Bot., 64, 2789, 1986.
Tyerman, S. D. , Hatcher, A. I. , West, R. J. , and Larkum, A. W. D. , Posidonia australis growing in altered
salinities: leaf growth, regulation of turgor and the development of osmotic gradients. Aust. J. Plant Physiol., 11,
35, 1984.
Karimi, S. H. and Ungar, I. A. , The effect of salinity on the ionic content and water relations of Atriplex
triangularis, in Symposium on the Biology of Atriplex and Related Chenopods, Tiedemann, A. R. , McArthur, E.
D. , Stutz, H. C. , Stevens, R. , and Johnson, K. , Eds., Gen. Tech. Rep. INT-172, Forestry Service, U.S.
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Ewing, K. , Earle, J. C. , PicClnin, B. , and Kershaw, K. A. , Vegetation patterns in James Bay coastal marshes.
II. Physiological adaptation to salt-induced water stress in three halophytic graminoids, Can. J. Bot., 67, 521,
1989.
Pammenter, N. W. and Smith, V. R. , The effect of salinity on leaf water relations and chemical composition in
the subantarctic tussock grass Poa cookii Hook F., Ν. Phytol., 94, 585, 1983.
Matthews, M. A. , Volkenburgh, E. V. , and Boyer, J. S. , Acclimation of leaf growth to low water potentials in
sunflower, Plant Cell Environ., 7, 199, 1984.
Houchi, R. , Morant-Avice, A. , Ferard, G. , and Coudret, A. , Effects of NaCl and NajS04 on water relations of
Plantago maritima (L.) and Plantago lanceolata (L.). Biol. Plant, 30, 457, 1988.
Ferard, G. , Coudret, A. , and Lasceve, G. , Action d’un choc osmotique sur Ies echanges d’eau des Plantago
maritima L. et des Plantago lanceolata L., C. R. Acad. SCl. Paris. 294, 775, 1982.
Coudret, A. , Action du NaCl les contraintes et les relations hydriques dans les parties aeriennes de Plantago
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Ferard, G. and Coudret, A. , Relations entre Γ absorption, le transport de I’ion sulfate et les contraintes
hydriques chez Plantago lanceolata et Plantago maritima soumis a des variations de potentiel osmotique,
Physiol. Veg., 20, 703. 1982.
Bolanes, J. A. and Longstreth, D. J. , Salinity effects on water potential components and bulk elastic modulus of
Alternanthera philoxeroides Griseb., Plant Physiol., 75, 281, 1984.

Plant Ionic Content

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Breckle, S.-W. , Ionengehalte halophiler Pflanzen Spaniens, Decheniana, 127, 221, 1975.
Sharma, Μ. L. and Tongway, D. J. , Plant induced soil salinity patterns in two saltbush (Atriplex spp.)
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Bjerregaard, R. S. , West, Ν. E. , Caldwell, Μ. M. , and Mayland, H. F. , Standing crops and dynamics of
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Breckle, S.-W. , Studies on halophytes from Iran and Afghanistan. II. Ecology of halophytes along salt
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Oecologia, 27, 157, 1977.
Guy, R. D. and Reid, D. M. , Factors affecting 13C/12C ratios of inland halophytes. II. Ecophysiological
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Gorham, J. , Hughes, L. L. , and Wyn Jones, R. G. , Chemical composition of salt-marsh plants from Ynys Mon
(Anglesey): the concept of physiotypes, Plant Cell Environ., 3, 309, 1980.
Stienstra, A. W. , Mineral composition of Halimione portulacoides (L.) Aellen leaves, Hydrobiologia, 52, 89,
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87, 241, 1972.
Winter, K. , Troughton, J. , Evenari, M. , Lauchli, A. , and Luttge, U. , Mineral ion composition and occurrence of
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25, 125, 1976.
Poulin, G. , Bourque, I. , Eid, S. , and Jankowski, K. , Composition chimique de Salicornia europaea L., Nat.
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Goodin, J. R. and McKell, C. M. , Atriplex spp. as a potential forage crop in marginal agricultural areas, Proc.
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Beadle, N. C. W. , Whalley, R. D. B. , and Gibson, J. B. , Studies in halophytes. II. Analytic data on the mineral
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Breckle, S.-W. , Zur Okologie und zu dem Mineralstoffverhaltnissen absalzender und nicht absalzender
Xerohalophyten, Diss. Bot., 35, 1, 1976.
Ernst, W. H. O. , Okologische Anpassungsstrategien an Bodenfaktoren, Ber. Dtsch. Bot. Ges., 96, 49, 1983.
Stumpf, D. K. and O'Leary, J. W. , The distribution of Na*, K+ and glycinebetaine in Salicornia bigelovii, J. Exp.
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Weber, D. J. , Rasmussen, H. P. , and Hess, W. M. , Electron microprobe analysis of salt distribution in the
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Eshel, A. and Waisel, Y. , Distribution of sodium and chloride in leaves of Suaeda monoica, Physiol. Plant. 46,
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Khan, Μ. A. , Weber, D. J. , and Hess, W. M. , Elemental distribution in shoots of Salicornia pacifica var.
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Lanning, F. C. and Eleuterius, L. N. , Silica and ash in tissues of some plants growing in the coastal area of
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Joshi, A. J. , Amino aClds and mineral constituents in Sesuvium portulacastrum L., a salt marsh halophyte,
Aquatic. Bot., 10, 69, 1981.
Waisel, Y. , Biology of Halophytes, Academic Press, New York, 1972.
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Goldstein, G. , Rada, F. , Sternberg, L. , Burguera, J. L. , Burguera, M. , Orozco, A. , Montilla, M. , Zabala, O. ,
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Mert, H. H. and Vantar, Y. , Salinity, osmotic pressure and transpiration relationships of Salicornia herbacea in
its natural habitat, Phyton, 18, 71, 1977.
Rajpurohit, K. S. and Sen, D. N. , Seasonal variation in chloride ion percentage of plants and soils of Pachpadra
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Riehl, Τ. E. and Ungar, I. A. , Growth and ion accumulation in Salicornia europaea under field conditions,
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Riehl, Τ. E. and Ungar, I. A. , Growth, water potential and ion accumulation in the inland halophyte Atriplex
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Joshi, A. J. , Effects of seawater in amino aClds and mineral ions composition in Salicornia brachiata Roxb, J.
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Bennert, H. W. and Schmidt, B. , On the osmoregulation in Atriplex hymemelytra (Torr.) Wats. (Chen-
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Cooper, A. , The effects of salinity and waterlogging on the growth and cation uptake of salt marsh plants, N.
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Cheeseman, J. M. , Bloebaum, P. , Enkoji, C. , and Wickens, L. K. , Salinity tolerance in Spergularia marina.
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Cheeseman, J. M. and Wickens, L. K. , Control of Na* and K* transport in Spergularia marina. III. Relationship
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Cheeseman, J. M. and Wickens, L. K. , Control of Na* and K* transport in Spergularia marina. I. Transpiration
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Cheeseman, J. M. and Wickens, L. K. , Control of Na* and K* transport in Spergularia marina. II. Effects of plant
size, tissue ion contents and root-shoot ratio at moderate salinity, Physiol. Plant, 67, 7, 1986.
Okusanya, Ο. T. and Ungar, I. A. , The growth and mineral composition of three species of Spergularia as
affected by salinity and nutrients at high salinity, Am. J. Bot., 71, 439, 1984.
Stelzer, R. and Lauchli, A. , Salt- and flooding tolerance of PucClnellia peisonis. III. Distribution and localization
of ions in the plant, Z. Pflanzenphysiol., 88, 437, 1978.
Storey, R. and Wyn Jones, R. G. , Salt stress and comparative physiology in the Gramineae. III. Effect of
salinity upon ion relations and glycinebetaine and proline levels in Spartina X townsendii, Aust. J. Plant Physiol.,
6, 831, 1978.
Allen, E. B. and Cunningham, G. L. , Effects of vesicular-arbuscular mycorrhizae on Distichlis spicata under
three salinity levels, N. Phytol., 93, 227, 1983.
Robinson, S. P. and Downton, W. J. S. , Potassium, sodium and chloride ion concentrations in leaves and
isolated chloroplasts of the halophyte Suaeda australis R. Br., Ausi. J. Plant Physiol., 12, 471, 1985.
Kappen, L. , Widerstandsfahigkeit von Halophyten gegenuber Gefrieren and Salzstress und ihre moglichen
biochemischen Ursachen, Ber. Dtsch. Bot. Ges., 92, 55, 1979.
Demming, B. and Winter, K. , Sodium , potassium, chloride and proline concentrations of chloroplasts isolated
from a halophyte, Mesembryanthemum crystallinum L., Planta, 168, 421, 1986.
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concentrations in salt-sensitive and halophytic species, Aust. J. Biol. SCl., 19, 741, 1966.
Greenway, H. , Growth stimulation by high chloride concentrations in halophytes, Isr. J. Bot., 17, 169, 1968.
Braun, Y. , Hassidim, Μ. , Lerner, H. R. , and Reinhold, L. , Studies on H * -translocating ATPase in plants of
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Jeschke, W. D. , Aslam, Z. , and Greenway, H. , Effects of NaCl on ion relations and carbohydrate status of
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Matoh, T. , Watanabe, J. , and Takahashi, E. , Sodium, potassium, chloride, and betaine ne concentrations in
isolated vacuoles from salt-grown Atriplex gmelini leaves, Plant Physiol., 84, 173, 1987.
Aswathappa, N. and Bachelard, E. P. , Ion regulation in the organs of Casuarina species differing in salt
tolerance, Aust. J. Plant Physiol., 13, 533, 1986.
Gorham, J. , Photosynthesis, transpiration and salt fluxes through leaves of Leptochloa fusca L. Kunth, Plant
Cell Environ., 10, 191, 1987.
Hajibagheri, M. A. and Flowers, T. J. , X-ray microanalysis of ion distribution within root cortical cells of the
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Bai, A. R. and Dutt, S. Κ. , Salt tolerance mechanism in Acanthus iliClfolius 1., Indian J. Plant Physiol., 30, 170,
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Austenfield, F. A. , Untersuchungen zum Ionenhaushalt von Salicornia europaea, 1. unter besonderer
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1974.
Stumpf, D. K. , Prisco, J. T. , Weeks, J. R. , Lindley, V. A. , and O'Leary, J. W. , Salinity and Salicornia bigelovii
Ton. seedling establishment. Water relations, J. Exp. Bot., 37, 160, 1986.
Ownbey, R. S. and Mahall, B. E. , Salinity and root conductivity: differential responses of coastal succulent
halophyte, Salicornia virginica, and a weedy glycophyte, Raphanus sativus, Physiol. Plant., 57, 189, 1983.
Mahall, B. E. and Park, R. B. , The ecotone between Spartina foliosa Trin. and Salicornia virginica L. in salt
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1985.
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Genecology
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