MODULE 4

Physiological Processes Affecting Crop

Production (by Pacifico T. Vizmonte Jr. and Justo G. Canare Jr. as enriched
by Rosemarie T. Tapic)

Overview
This module presents the different physiological processes as occurs
in plants, the factors affecting the different processes and the plant
characteristics that differentiate the occurrence among the processes in
plants.

Time Allotment (1 Week)

Objectives
Upon the successful completion of this module, you are expected to:

1. Define photosynthesis, respiration, transpiration and translocation

2. Relate the physiological processes occurring in plants in the
formation/production of economic useful parts.

Pre-Assessment
Before you proceed to the discussion section of this module, I want you to first give your
initial thoughts on the following queries below.

1. Can you recall the end product of photosynthesis?

2. What are the factors that affect transpiration?

Discussion

This section begins with the discussion of the photosynthesis process and factors affecting
it. It then moves on the presentation of respiration and translocation. The last part of this
section provides the three types of transpiration and factors that affect transpiration.

PHOTOSYNTHESIS
A. The Process
Photosynthesis is a biochemical process that serves as the connecting link between
solar energy and life on earth. It is a constructive (anabolic) process by which food
of the plant is manufactured from simple inorganic materials through the agency
of chlorophyll; the energy needed is derived from sunlight. In this process, solar
energy is captured and converted into chemical energy stored in the form of
carbohydrates. It is the process where carbon dioxide (CO2) is fixed by the plants.

The products of photosynthesis are sources of energy and building materials for
plants and animals. They are also sources of industrial materials. Approximately
90% of the human population is engaged directly or indirectly in the production,
processing, and sale of the products and by-products of photosynthesis.

Photosynthesis is a complex process. The summary chemical equation is as

follows:

light
6CO2 + 6H2O C6H12O6 + 6O2
Chlorophyll

The process can be understood in terms of three partial steps as follows:

1. Diffusion of carbon dioxide to the chloroplast. Carbon dioxide in the air is

transported by turbulence or diffusion to the leaf stomata through which it diffuses
to the chloroplast.

2. Light reactions or photochemical reactions. In this reactions, light is

essential, while temperature is unimportant. They take place in the thylakoid
membranes of the chloroplast. Light energy is used to split water producing
molecular oxygen (O2), reduced nicotinamide adenine dinucleotide phosphate
(NADPH) and adenosine triphosphate (ATP). The light reactions can be divided
into three components:

a. Photolysis of water. Water is split into hydrogen ions (H+), electrons (e-)
and molecular oxygen (O2).

2H2O + light 4H+ + 4e- + O2

b. Reduction of NADP+ to NADPH. The electrons from water are transferred

through the photosynthetic electron transfer system to NADP+ to form
NADPH.
 NADP+ + H+ + 2e- + light NADPH

c. Photophosphorylation. Light and inorganic phosphate (Pi) are used to

convert adenosine diphosphate (ADP) to ATP.

ADP + Pi + light ATP

3. Dark reactions or biochemical reactions. In these reactions, light is not

required, but temperature plays an important role. The products of the light
reactions (NADPH and ATP) are used to fix carbon dioxide. The CO2 fixation can
occur by one of three pathways: (a) the Calvin-Benson or C3 pathway; (b) the
Hatch and Slack or C4 pathway; and (c) the Crassulacean Acid Metabolism or CAM
pathway.

In the C3 pathway, ribulose bisphosphate (RUBP) is the CO2 acceptor.

3-phosphoglyceric acid (PGA) is the first stable product (Figure IV-1). The name
of the pathway came from the fact that the first stable product (PGA) has three
carbon atoms. The Calvin-Benson cycle also occurs in C4 plants and CAM plants
as the last stage of the CO2-fixation process.

In the C4 pathway, phosphoenol pyruvic acid (PEP) is the initial CO2

acceptor and the first stable product is oxaloacetic acid (OAA), which is converted
to either malic acid or aspartic acid. OAA has four carbon atoms, hence the name
of the pathway. It is also called the Hatch and Slack pathway to honor the two
scientists who elucidated the pathway. The Calvin-Benson-cycle part of the
pathway occurs in the bundle-sheath cells, while the initial fixation of CO2 occurs
in the mesophyll cells.

In the CAM pathway, CO2 is fixed at night because the stomata of CAM
plants are closed during the day and open at night as a way of conserving
moisture. It has many reactions similar to those of the C4 pathway, but the Calvin-
Benson cycle is separated from the rest of the photosynthetic pathway in time – it
occurs during the day while the initial fixation of CO2 occurs during the night.

Based on the photosynthetic pathway followed, plants are classified as C3

plants, C4 plants, and CAM plants. Table 1 shows a comparison of the three types
of plants.
Table 1. Some characteristics of C3, C4 and CAM plants
Characteristics C3 plants C4 plants CAM plants
Typically temperate
species, e.g.,
spinach, wheat, Typically tropical or
potato, tobacco, semi-tropical Typically arid zone
sugar beet, soy species, e.g., corn, species, e.g., cacti,
1. Plant species
bean, sunflower, sorghum, sugarcane, Agave, bromeliads
following the
etc., and tropical amaranth, plants such as pineapple,
pathway
species, e.g. rice, adapted to high light most orchids, other
taro, sweet potato, intensity and succulents.
cassava, mungbean, temperature.
etc. Most plants are
C3 plants.
Mesophyll with
Mesophyll with no
distinct bundle Mesophyll with large
2. Leaf anatomy distinct bundle
sheath (Kranz vacuoles.
sheath
anatomy).
3. Energy
requirement 1:3:2 1:5:2 1:6.5:2
(CO2:ATP:NADPH)
4. Optimum
temperature for
15-25 25-40 35-45
photosynthesis
(oC)
5. CO2 fixation Two separated in Two separated in
One
pathways space time
6. Light saturation ¼ to ½ of full Unsaturated even at
1/10 of full sunlight
for photosynthesis sunlight full sunlight
7. CO2
compensation 30-70 0-10 0-5
point (l/l)
8. Maximum
values of net
15-30 35-45 1-5
photosynthesis
(mol CO2/m2/s)
9. Transpiration
ratio (H2O lost/ 450-950 250-350 18-125
CO2 fixed)
10. High (1/3 of total
Very low or absent Very low
Photorespiration photosynthesis)

The triosphosphate produced in the Calvin-Benson cycle (in C3, C4 and CAM
pathways) is the starting point for a wide range of biosynthetic pathways. The two
major end products of photosynthesis are sucrose and starch. However, their
syntheses are not parts of the Calvin-Benson cycle. Sucrose is the main form of
carbohydrates translocated in the plant. It is, therefore, an important raw material for
 the synthesis in different parts of the plant of many other organic molecules that are
necessary for growth, maintenance, and yield formation.

B. Factors Affecting Photosynthesis

Photosynthesis and consequently, crop productivity are influenced by a host of

variable factors. Included in these are both the internal and external or
environmental factors. The internal factors include nutritional status of the plant,
stomatal aperture and leaf characteristics such as age, morphology, leaf area index,
leaf angle and orientation. On the other hand, the external factors include light or
irradiance, temperature, carbon dioxide concentration, moisture and wind.

1. Internal Factors

a. Nutritional status of the plant. A deficiency of any of the essential element

will cause a decrease in the photosynthetic rate of the leaves. For example,
chlorophyll contains both nitrogen and magnesium; if they are limited in supply,
chlorophyll may not be formed. Precursor molecules for chlorophyll synthesis
include iron, and if it is not present, chlorophyll cannot be synthesized. In this
case, the deficiency of an element influenced the photosynthetic apparatus of
the plant.
In rice (O. sativa), nitrogen content per leaf area is closely related with
photosynthetic rate. Nitrogen is one of the most important nutrients and it is
often deficient in rice-cultivated areas. When nitrogen is limiting, both the leaf
area index (LAI) and the rate of photosynthesis per unit leaf surface is reduced.
The combination of these two factors results in a lowered rate of
photosynthesis.
b. Stomatal aperture. When water is abundant and nutrition is adequate, the
rate of photosynthesis is usually controlled by the stomatal aperture or opening
and the number of stomata per unit of surface area. Since carbon dioxide (CO2)
diffuses or moves into green tissues through the stomata, the stomatal number
and apertures must be great enough for passage of adequate levels of CO2. In
rice plants, stomata are found on both the upper and lower surfaces of the
leaf. Stomatal aperture is reduced by water stress, insufficient light, and high
carbon dioxide concentration.
c. Leaf age. The photosynthetic rate of young leaves is usually low but increases
as the leaf approaches full expansion. After reaching maturity, subsequent
photosynthetic activity varies widely with both species and environment.
Old, senescent leaves eventually become yellow and are unable to
photosynthesize because of chlorophyll breakdown and loss of functional
chloroplasts.
2. External Factors
1. Light (Irradiance). Under natural conditions, the process of
photosynthesis is “driven” by the visible portion of the spectrum or radiant
energy between 400 and 700 nanometer (nm) wavelength.

Photosynthetic rate is partially a function of the level of irradiance. In total

darkness, no photosynthesis occurs and as irradiance increases, photosynthesis
increases until the light compensation point is reached, where photosynthetic fixation
of CO2 exactly equals respiratory release of CO2, i.e. there is no net movement of CO2
into or out of the leaf (net photosynthesis = 0). As irradiance further increases, net
photosynthesis (gross photosynthetic CO2 fixation minus respiration) increases rapidly
until the light saturation level is reached, at which point the photosynthetic rate levels
off.
The following are units or expressions of:
a. photosynthesis
1. mg CO2 /dm 2 leaf area/ hr-
2. mg CO2 /cm2 leaf area/sec
3. u mol CO2 / m2 leaf area/sec

b. irradiance
1. watt / m2
2. cal /cm 2 / min

Light intensity varies with the season, for instance in Los Baños, Laguna the light
intensity during the wet and dry seasons are:
a. wet season, 300 cal / cm2 / day
b. dry season, 500-600 cal / cm 2 / day

This means that there is a doubling of the available light during summer
and explains why light is often a limiting factor to high yields during the wet
season.

2. Temperature. The light dependent reaction of photosynthesis is little

affected by temperature but the dark reaction is very temperature dependent.
Within the temperature range of 20-35 0C, temperature has little effect on
photosynthesis. However, it does have a definite effect on respiration.
Enzymatically controlled reactions such as respiration occur in a temperature
range greater than 0oC and less than 50oC. Respiration increases by a factor of
2 or 3 for every increase in temperature of 10oC. This means that the high
tropical temperature has no effect on photosynthesis but will result in a higher
rate of respiration. This is one reason why yields are low in the tropics.

3. Carbon dioxide (CO2) concentration. Carbon dioxide is a gaseous

component of air. Dry air contains 78% nitrogen (N2); 21% oxygen (O2);
0.93% argon (Ar); 0.038% (380ppm) CO2 and traces of other gases. Although
carbon dioxide is at a low concentration, 85% to 92% of a plant’s dry weight
is derived from CO2 uptake in photosynthesis.
The average CO2 concentration of normal ambient (surrounding) air is
about 0.038% or about 380 ppm. Diurnal fluctuation of CO2 concentration
occur over the rice fields, increasing at night when only respiration is being
carried on and decreasing during the day when the rice plants are carrying out
photosynthesis.
Photosynthetic rates are enhanced by higher CO2 concentration, unless
stomata are closed by drought or water stress. The response of photosynthesis
to CO2 is similar to its response to irradiance.
There is a very strong interaction between CO2 level and light level. At low
irradiance, low levels of CO2 will saturate the photosynthetic mechanisms but
as light level increases, progressively higher concentrations of CO2 are required
to “use up” the NADPH and ATP produced in the light reactions. Another aspect
of CO2 levels is that within the canopy of a crop, CO2 levels are often well below
the general atmospheric level. This depletion depends on several factors
including canopy density, wind speed and irradiance.

4. Moisture. Water is a substrate for photosynthesis, but only about 0.1% of

the total water is used by the plant for photosynthesis. Transpiration accounts
for 99% of the water used by plants. Approximately 1% is used to hydrate the
plant, maintain turgor pressure and make growth possible.
The supply of water or moisture affects stomatal aperture or opening.
Stomata open because the guard cells take up water and swell. When water is
deficient, stomata close, hence the rate of photosynthesis is reduced.

5. Wind. Some preliminary data need to be examined in order to understand

the role of wind in photosynthesis.
The maximum increase in dry weight per square meter per day observed in
rice in Los Baños, Laguna is about 30 g. If their mineral content accounts for
10% of the dry matter, the net increase in organic matter would be 27 g / m 2
/ day. To obtain 27 g of organic matter (dry weight /m2), it is necessary for the
plant to assimilate 39.6 g of CO2 / m2.
 The replacement of air above the rice crop with fresh air is important to the
photosynthetic efficiency of the crop. If the air were not replaced, it would
quickly become depleted of CO2. Air movement is accomplished primarily
through wind and turbulence. The movement of air through turbulence has
been computed 104 times faster than normal diffusion would be. Normally,
moderate winds and turbulence are important to the production of high yields.

RESPIRATION
Two of the most important processes carried out by green plants are
photosynthesis and respiration. Photosynthesis, which takes place in the chloroplasts, is
constructive and reductive, while respiration is degradative (destructive) and oxidative.
All humanity and animal life is dependent on the ability of plants to utilize solar energy
and to store it to provide material for respiration.

Every organism must extract energy from the organic fuel molecules that it either
manufactures or captures from the environment. These fuel molecules are transported
to all the cells of a complex organism, where they can be broken down to provide the
energy for cellular work.
Every plant cell then must extract energy from its long-term storage molecules –
the organic molecules it manufactures by photosynthesis. Within each plant cell, glucose
and other fuel molecules are broken down by the process of cellular respiration, a series
of chemical reactions that break apart molecules and transfer the energy stored in their
bonds to adenosine triphosphate (ATP) for use to carry out work in the cell.
Cells use different catabolic pathways to extract energy from the fuel molecules
they manufacture or ingest: aerobic respiration and anaerobic respiration. The type of
environment a cell inhabits may determine which catabolic pathway it uses to break down
fuel molecules. Cells that live in an environment where oxygen is plentiful use an aerobic
pathway, one that requires oxygen, whereas cells that inhabit waterlogged soils or
polluted water where oxygen is absent must use anaerobic pathways that do not require
oxygen.
A. The Process Of Aerobic Respiration

Plant cells extract energy from fuel molecules (e.g. glucose, fatty acids and other
organic compounds) by using aerobic respiration. This process involves a long sequence
of 30 or more chemical reactions, each regulated by a specific enzyme. During aerobic
respiration, energy is released as fuel molecules are catabolized to CO 2 and water. One
of the most common pathways of aerobic respiration involves breakdown of glucose.
Glucose is generally regarded as the starting point for the respiratory metabolism of
carbohydrates.
The overall process of respiration can be expressed by the equation below:
 C6H12O6 + 6O2 6CO2 + 6H2O + Energy
glucose oxygen carbon dioxide water

Chemically, respiration is oxidation. In oxidation, either oxygen is added to the

material being oxidized or hydrogen is removed from it. A substrate, or more specifically
a respiratory substrate, is any organic plant constituent oxidized partially (to more
oxidized compounds) or completely (to carbon dioxide and water) in respiratory
metabolism.

The chemical reactions of aerobic respiration are grouped into four stages, namely:
1. Glycolysis
2. Formation of acetyl coenzyme A
3. The citric acid cycle also known as Krebs cycle
4. Electron transport and chemiosmosis

Glycolysis occurs in the cytoplasm; all other parts of aerobic respiration occur in
the mitochondrion.

B. Importance Of Aerobic Respiration

1. It supplies the energy needed for the growth and maintenance of the plant.
2. It provides carbon skeletons needed for the synthesis of a large number of other
essential plant products. These products include amino acids for proteins, nucleotides
for nucleic acids, and carbon precursors for porphyrin pigments (such as chlorophyll
and cytochromes) and for fats, sterols, carotenoids, flavonoid pigments such as
anthocyanins, and certain other aromatic compounds such as lignin.

C. Factors Affecting The Rate Of Aerobic Respiration

1. Temperature. Temperature affects respiration rates. Within the range of 0 oC-

35oC, the rate increases about two to three times for each 10 oC rise. The effect of
temperature on respiration is an important factor in the storage of some crops. A
harvested plant part that is stored is often a living tissue and unless the product has
been cooked or processed, the enzymes are active and vital processes continue. Since
respiration is a degradative process, it should be retarded (slowed down/delayed) as
quickly and as completely as possible to prolong life. One way to retard respiration is
to refrigerate the product.

2. Oxygen concentration. Oxygen is an essential component of respiration. With

other factors being constant and not limiting, the rate of respiration decreases as
oxygen concentration decreases. Lowering the oxygen concentration by increasing
 the concentration of carbon dioxide or nitrogen is useful in storing certain fruit and
vegetable crops. This modified storage atmosphere slows down rapid respiration.

3. Soil condition. Compacted or water logged soils are generally poorly aerated.
This condition reduces respiration in the roots resulting in poor plant growth. Mineral
nutrient deficiencies affect the respiratory enzymes, indirectly causing a reduction in
respiration.

4. Light. Plants that grow in low light intensities exhibit lower respiration rates. Low
light reduces photosynthesis, thus decreasing the amount of carbohydrates available
for respiration.

Table 2. A comparison of photosynthesis and aerobic respiration.

Items Photosynthesis Aerobic respiration
Raw materials CO2 , H2O C6H12O6, O2
End products C6H12O6, O2 CO2 , H2O
Plant cells that Plant cells that contain All plant cells
have these process chlorophyll
Parts of cell Chloroplast Cytoplasm, mitochondrion
involved
Pathway of energy Light energy NADPH/ATP Energy stored in fuel
energy stored in molecules  NADH/ATP
carbohydrate molecules  energy for work in cell

TRANSLOCATION

To maintain its metabolic activity, the plant needs to circulate to the various plant
parts a wide variety of compounds. In multicellular plants, the water and inorganic
substances absorbed by the roots have to be transported to the leaves. The sugars and
other organic compounds synthesized there have to be distributed to all parts of the
plant, where they are used for growth and maintenance, or stored. These parts include
the parts that we harvest.

To meet these needs, higher plants have developed two translocation systems,
the xylem and the phloem. The xylem provides the pathway by which the water and
mineral nutrients absorbed by the roots are distributed to all parts of the plant, including
the highest leaves. The phloem, on the other hand, provides the pathway by which the
sugars and other organic compounds produced in the leaves and some inorganic ions
move to the different parts of the plant.

Sugars, which are synthesized during photosynthesis, move throughout the plant,
principally through the phloem tissue. The movement can be downward, from leaves to
roots, but lateral and upward movement from leaves to buds or to fruits or to storage
organs also occurs. Translocation takes place in the long sieve elements connected end
to end to form sieve tubes.

The rate of translocation of sugars in the phloem is rapid, in some instances more
than a thousand times faster than simple diffusion of sugar through water. The rate of
translocation has been measured in many plants and average values of 1 to 6 g/cm 2/hr
have been found in developing fruits and tubers.

Much of the carbohydrate translocated within plants is sucrose. This disaccharide

is formed by the linkage of glucose and fructose accompanied by the removal of a
molecule of water.

Movement of materials in living plants has been observed to occur in different

ways, namely:
1. ordinary diffusion, which transports ions and molecules slowly;
2. cytoplasmic streaming, which transports molecules and ions within the cytoplasm
at a considerably faster rate than diffusion;
3. mass flow translocation of material in the phloem;
4. very rapid upward movement of water and mineral nutrients through the xylem;
lateral transport of materials along the vascular rays radially from sieve tubes into
the cambium tissue and xylem.

Translocation to a plant part can be reduced or completely stopped by pests, diseases,

or mechanical breakage (as in lodging).

TRANSPIRATION

Transpiration is the loss of water in the form of water vapor from aerial parts of
plants. It is basically an evaporative process. It involves two stages: (1) the evaporation
of water from cell surfaces (dependent on energy, the latent heat of vaporization, which
is equal to 539 cal/gram), (2) the diffusion of water vapor out of the plant, mainly through
the leaves. The driving force of transpiration is the vapor pressure gradient between the
leaf interior and the atmosphere.

There are three types of transpiration, namely:

1. Cuticular transpiration – loss of water takes place directly through the cuticle of
the leaf epidermis. In some plants, this contributes about 5-10% of the water loss.
2. Lenticular transpiration - loss of water takes place through lenticels, which are
found in stems of trees and in some fruits.
3. Stomatal transpiration - loss of water takes place through stomata, which are
found on leaves. The leaves of most plants have stomata on the lower surface,
although certain species, especially grasses, have them on both upper and lower
 surfaces. Stomatal transpiration can account for more than 90% of the water loss
from plants.

Transpiration is often called a “necessary evil”. Evil because it can result in excessive
loss of water from plants, as in periods of low relative humidity and high temperatures.
It is estimated that up to 99% of the water absorbed by plants is lost through
transpiration. For example, a corn plant absorbs about 200 liters of water during its
growth from the seedling stage to maturity, but only about 2 liters are present in a mature
plant. Internal water stress can reduce the rate of growth and the yield of crops. Worse,
it can kill the plant.

However, transpiration has benefits for crops. Transpiration helps in the mobilization
of soil nutrients toward the roots. It aids in the translocation of mineral nutrients absorbed
by the roots. It cools the plant, thereby maintaining a favorable plant temperature for
growth and development.

The factors that affect transpiration are:

1. solar radiation – it is the main source of energy for the evaporation of water; visible
radiation (light) can directly or indirectly induce the opening of stomata. That’s why
transpiration is high during the day, except in CAM plants.
2. temperature – an increase in temperature increases the capacity of the air to absorb
water vapor.
3. relative humidity – as relative humidity becomes lower, transpiration increases.
4. wind – a gentle breeze increases the rate of transpiration compared to still air.
5. soil moisture availability – when there is lack of water in the soil, transpiration is
reduced.
6. carbon dioxide concentration – an increase in carbon dioxide concentration induces
partial closure of stomata. Thus, transpiration is reduced.
plant adaptations such as modification of leaves into scales or spines, degree of cuticular
deposition, sunken stomata, closing of stomata during the day (in CAM plants), presence
of hairs on the leaf, solar tracking in which the leaf is oriented such that its lamina is
almost parallel to the rays of the sun thereby reducing its heat load, etc.

Summary
 Photosynthesis and respiration are two of the most important processes carried
out by green plants.
 Translocation is the transport of water and inorganic substances to the leaves.
 Transpiration could be evil to plants when it is too much.
Post-Assessment
Problem Solving. Identify the physiological process that affects the plant and how to
solve it. Download the form in the post assessment part to answer the following
questions.

1. The green grass in the lawn covered with coconut shell for one week turned yellow
green in color. Why? How can the yellow green colored grass recover?

2. The harvested carrots stored under 30oC (room temperature) changes it weights every
day. Why? How to solve/minimize this problem?

RUBRICS: (Content = 6pts; Punctuality = 4pts)

CRITERIA 6 points 4 points O point
CONTENT Write atleast two Write atleast one All the answers
sentences or sentence and one were incorrect or
more and two correct answer write only phrase
correct answers using correct with incorrect
using correct grammar. grammar.
grammar.
PUNCTUALITY - Submit the output Submit the output
on or before the after the deadline
deadline (day and time)

Reference
Alfonso-Alejar, A.M., and M.L. Dionisio-Sese. 1999. Fundamentals of Plant Physiology.
Plant Physiology Society of the Philippines, Pasig City.

Gardner, F.P., R.B. Pearce and R.L. Mitchell. 1985. Physiology of Crop Plants. The Iowa
State University Press.

Halfacre, R.G and J.A. Barden. 1979. Horticulture. McGraw Hill Book Co. New York.

Hartmann, H.T., A.M. Kofranek, V.E. Rubatzky and W.J. Flocker. 1988. Plant Science.
Prentice Hall Inc., New Jersey, U.S.A.

International Rice Research Institute (IRRI). Photosynthesis and respiration of the rice
plant. A slide tape instructional unit. GM-4. Rice Production Training Series. IRRI.
Los Baños, Laguna.

Rimando, T.J. 2001. Lecture Syllabus in Crop Science I. Department of Horticulture, U.P.
Los Baños, Laguna.
Salisbury, F.B. and C.W. Ross. 1992. Plant physiology. 4th ed. Wardsworth Publishing Co.
California.

Yoshida, S. 1981. Fundamentals of Rice Crop Science. Intenational Rice Research

Institute, Los Baños, Laguna.