Crop Sci - Principles of Crop Production

UNIT
c 3

Factors affecting crop production

Lesson 1.0: Physiological processes (Photosynthesis)

Lesson Outcomes

At the end
At the of of
end thethelesson,
lesson,the
the students willbe
students will beable
ableto:to:
1. familiarize the process of photosynthesis, respiration, translocation and transpiration
s
2. evaluate the significance of physiological processes on crop production
3. compare and contrast respiration and photosynthesis
4. differentiate the transpiration from translocation

PHOTOSYNTHESIS
Photosynthesis is the manufacture of sugars and its precursors by green plants in the presence of
light and chlorophyll. The process is represented by the following chemical equation: a

From the above chemical equation of photosynthesis, two important conclusions have been shown
and supported with experimental evidence. (1) CO2 is fixed as glucose and must be reduced by a
reductant, which may either be H2O (for higher plants) and H2S in the case of the sulfur bacteria. (2) O2
gas formed in photosynthesis must arise from the H20 molecule not from CO2 as shown by the Ruben and
Kamen’s experiment:

Carbon dioxide is taken from the air through the stomata while water is absorbed from the soil by
the roots and transported in the xylem to sites of photosynthesis. Outdoors, light comes from the sun, but it
may be supplied by artificial lamps under experimental and greenhouse conditions.

About 90% of the total world’s photosynthetic output occurs in the oceans, while the remaining 10%
by land plants (Bonner and Galston, 1952). Among the land plants, about 7-8% is accounted for by forest
and grasslands, while the remaining 2-3% by cultivated crops.

The photosynthetic organ, tissues and organelles

Although photosynthesis can occur in any organ containing chlorophyll, the main organ for photosynthesis
is the leaf, while the main organelle involved is the chloroplastid. The features which make the leaf an ideal
organ for photosynthesis are: (1) its typically expanded form, (2) its usually perpendicular angle to incident
light, (3) its extensive internal surface with an efficient vascular system for channeling the various reactants
and end-products of photosynthesis, and (4) its pigments for light absorption. The anatomy of a leaf is
shown in Figure 7.
 Crop Sci - Principles of Crop Production

A mature mesophyll cell contains about 50 chloroplasts. The chloroplasts of higher plants are usually lens-
shaped bounded by a double membrane. The inner membrane invaginates parallel to the surface and
becomes organized into a specialized cytoplasmic body consisting of a stack of thylakoids or frets, called
the granum (plural: grana) embedded in a proteinaceous matrix called the stroma (Figures 8 and 9).
 Crop Sci - Principles of Crop Production

The drawing in Figure 9 shows two grana stacks interconnected by parallel sets of unstacked stroma
thylakoids that spiral up and around the stacks in a right-handed helical conformation. Where the stroma
thylakoids intersect with grana thylakoids, they are interconnected through neck-like membrane bridges.
Chlorophyll, the principal pigment in photosynthesis, is located in the partition between two adjacent
thylakoids. Chlorophyll a occurs in all higher plants, but other isomers like chlorophyll b, c, d, etc. are also
found. In higher plants, the two main isomers are a and b in a ratio of 3:1. While chlorophyll a is always
found in all plants, chlorophyll b may be replaced by some other isomers in some plant species.
Figure 10 shows the chemical structure of the chlorophyll molecule. Its basic unit is the porphyrin ring
system, a structure made up of four simpler pyrrole nuclei joined by carbon linkages. The center of the
porphyrin is occupied by a single magnesium atom. The phytol side chain gives chlorophyll a lipid character.
In ring b, chlorophyll a contains a methyl residue instead of the formyl residue in chlorophyll b (Heldt, 2005).

The accessory pigments closely associated with chlorophyll are the carotenoids, chiefly xanthophyll (lutein
and zeaxanthin) and carotene, (chiefly ß-carotene).
The similarity of the absorption spectrum of intact chloroplast to its action spectrum in terms of oxygen
evolution shows that light absorption by chlorophyll mediates oxygen evolution in plants, and this supports
chlorophyll as the principal pigment in photosynthesis (Figures 11). Before light can be utilized in
photosynthesis, it must first be absorbed by the plants. Light wavelengths readily absorbed by the pigment
are also the wavelengths where the intensity of photosynthesis is relatively higher. Figure 12 shows the
absorption spectrum of chlorophyll and other plant pigments. Absorption of wavelength in the green region
of the spectrum is low as green is typically reflected, and consequently photosynthesis is also relatively low.
 Crop Sci - Principles of Crop Production

Component reactions of photosynthesis

Photosynthesis is consisting of two component reactions which occur in sequence, namely, the light
reaction and the dark reaction (Blackman reaction). Light reaction is a photochemical process with an
absolute requirement for light, while dark reaction is a thermochemical process that can take place in both
light and dark.

Light reaction
Light reaction takes place in two reaction centers that are embedded in the thylakoid membrane:
Photosystem I (PS I) and Photosystem II (PS II). The numbering of the photosystems corresponds only to
the sequence of their discovery. Localized in granal thylakoids, PS II has maximum absorption at 680 nm,
with P680 as the main pigment. It produces a very strong oxidant which can split water into hydrogen and
oxygen. On the other hand, PS I, found in stromal thylakoids, can utilize light with wavelengths up to 700
nm with P700 as the main pigment. It produces a strong reductant capable of reducing NADP+ to NADPH.
The non-cyclic flow of electrons during the light reaction is represented by the Z-scheme (Figure 13). In
summary, the electron flow in PS II and PS I results in the production of ATP and NADPH which are
subsequently used to fix CO2.
 Crop Sci - Principles of Crop Production

Dark reaction
The end-products of light reaction, namely ATP and NADPH, are subsequently used in CO2 fixation. The
fixation or reduction of CO2 into carbohydrates can occur via three pathways: (1) Calvin-Benson or C3
pathway, (2) Hatch-Slack or C4 pathway and the (3) Crassulacean acid metabolism or CAM pathway.

a. C3 or Calvin-Benson pathway
In the Calvin-Benson cycle, ribulose-1,5-bisphosphate (RuBP) is the CO2 acceptor and the enzyme
ribulose- 1,5 bisphosphate carboxylase/oxygenase (RubisCO) catalyzes the carboxylation reaction. The
first stable product, 3 phosphoglycerate (3-PGA), is a 3-carbon compound, hence the name C3
photosynthesis. The Calvin cycle proceeds in three stages: (1) carboxylation, during which CO2 reacts
with ribulose-1,5- bisphosphate (RuBP), in the presence of RubisCO; (2) reduction, during which 3-PGA
and eventually large carbohydrate units are produced at the expense of ATP and NADPH; and (3)
regeneration, during which RuBP re-forms for further CO2 fixation (Figures 14 and 15).

In the C3 pathway, fixation of one CO2 molecule requires a total of two molecules of NADPH and three
molecules of ATP. Moreover, five-sixths of the triose phosphate formed by photosynthesis is required for
the regeneration of RuBP. One molecule of triose-phosphate represents the net product and can be utilized
by the chloroplast or exported (check from new version).
 Crop Sci - Principles of Crop Production

Photorespiration and the C2 Pathway

Photosynthetic yield of C3 plants is reduced by the occurrence of photorespiration, which is a
consequence of the properties of RubisCO. RubisCO is a bifunctional enzyme in the C3 pathway that can
act either as a carboxylase or as an oxygenase. When RubisCO functions as a carboxylase, the C3 cycle
proceeds to yield two molecules of 3-PGA via carboxylation of RuBP. On the other hand, photorespiration
occurs when RubisCO behaves as an oxygenase, wherein oxidation of RuBP yields one molecule of 3-
PGA and one molecule of phosphoglycolate, a 2-carbon compound. For C3 plants, a more direct
contact of RubisCO to O2-rich environment favors photorespiration. At present atmosphere (i.e. 21% O2
and 0.038% CO2), RubisCO catalyzes the oxygenation of RuBP about 20-60% as rapidly as it catalyzes
RuBP carboxylation.
The C2 photorespiratory carbon oxidation cycle may operate to partially recover carbon lost
from photorespiration. This process involves the cooperative interaction of three organelles: chloroplast,
peroxisomes, and mitochondria (Figure 16). Photorespiration yields only one 3-PGA molecule in contrast to
two 3-PGA molecules in photosynthesis. Another 3-PGA is salvaged from phosphoglycolate, at the
expense of ATP, through a series of reactions in the peroxisome, mitochondria, and chloroplast via the C2
photorespiratory carbon oxidation cycle. In broad terms, photorespiration and the C2 cycle may be viewed
as a wasteful process. With the production of one phosphoglycolate (2- C) and one 3-PGA (3-C) instead of
two 3-PGA, 25% of the carbon is lost during photorespiration. It is also a metabolically- expensive reaction,
requiring five ATP and three NADPH for every oxygenation of RuBP. This is a significant energetic cost,
which can add up to approximately 50% of the amount of energy expended on carboxylation. Hence, plants
with higher rate of photorespiration (i.e. C3 plants) have lower rate of dry matter production because of
lower photosynthetic efficiency. Other photosynthetic pathways have evolved wherein photorespiration is
avoided. In plants with C4 or CAM physiology, photorespiration is almost entirely eliminated by
concentrating CO2 spatially (C4) or temporally (CAM) before CO2 is reduced to triose phosphates. In these
ways, the carboxylation function of RubisCO is favored over oxygenation.

b. C4 pathway or Hatch-Slack
Leaves of C4 plants have a characteristic anatomy, known as Kranz anatomy, wherein vascular
bundles are surrounded by a sheath of chloroplast-rich parenchyma cells called the vascular bundle sheath,
which are surrounded by the spongy mesophyll. In C4 plants, the bundle sheath cells generally contain
more and/or larger chloroplasts than mesophyll cells The special anatomical feature of C4 leaves favors a
more efficient photosynthesis by assuming the following reaction steps. First, CO2 (in the form of HCO3-) is
initially fixed by phosphoenolpyruvate (PEP), in the presence of PEP carboxylase, in the mesophyll cells
forming the first stable product, oxaloacetate (OAA).
OAA is a 4-C compound, hence the name C4 photosynthesis. OAA is then converted to C4 acids
(malic acid or aspartic acid), which are subsequently transported to and decarboxylated (CO2 is released)
in the bundle sheath cell where RubisCO and RuBP are present in large quantities. Because of this CO2-
pumping mechanism, CO2 is generated in the bundle sheath cells resulting in its significantly higher CO2
concentration. This largely favors the carboxylation function of RubisCO such that CO2 in the bundle
sheath cells is refixed by RuBP via the C3 pathway. Apparently, the C3 cycle occurs in the bundle sheath
cells of C4 plants (Figure 19).
 Crop Sci - Principles of Crop Production

c. Crassulacean Acid Metabolism (CAM) pathway

The CAM pathway is found in succulent plants (e.g. cactus and members of the pineapple family).
While C4 plants eliminate photorespiration through a spatial CO2-concentrating mechanism, CAM plants
do this through temporal regulation of photosynthetic reaction steps. Like in the C4 pathway, PEP is the
initial CO2 acceptor and PEP carboxylase is the initial carboxylating enzyme. Malic acid, produced in the
initial carboxylation of PEP as in the C4 pathway, accumulates in the vacuole and acidifies the cells.
However, these processes occur in the dark which means that stomates of CAM plants are open at night.
During the light period (i.e. day), the malic acid is decarboxylated, yielding CO2 which is fixed through the
C3 pathway in the chloroplast. Since the stomates are closed at daytime, CAM plants maximizes water-use
efficiency which allow them to thrive even in arid environments (Figure 20).
 Crop Sci - Principles of Crop Production

Factors affecting photosynthesis

Operationally, three basic processes are evident during photosynthesis.

1) The diffusion of carbon dioxide from the air to the reaction sites in the leaves.
2) The light dependent reactions (photochemica);
3) The carbon dioxide fixation reactions (biochemical)

The main environmental factors affecting the above processes may be briefly stated as follows;

Diffusion. Carbon dioxide concentration in air and in the leaf, and the resistance to diffusion in the
pathway influence the diffusion rates.

Photochemical. Light intensity influences reaction rates. Light absorption is in turn affected by the
concentration of pigments, adequacy of nutrients and leaf morphology and position.

Biochemical. Temperature is the main factor, as well as the presence of inhibitors.

On a single leaf basis, net photosynthesis per leaf area of C4 plant species increases with light
intensity up to full sunlight, leaves of C3 plants become light- saturated at 1/4 - 1/3 full sunlight,
 Crop Sci - Principles of Crop Production

while shade leaves may be saturated at light intensities 1/10 of full sunlight. The efficiency of
photosynthesis in terms of light utilization at high light intensities is obviously greater in the C4
response.

On carbon dioxide concentration, photosynthesis is dependent on the ambient C02 concentration,

especially at high light intensities. The differential which exists in photosynthetic efficiency at high
light intensities between the C4 and C3 plants is greatly reduced at high carbon dioxide. Stomatal
aperture also has a significant influence on the diffusion resistance to carbon dioxide during periods
of moisture stress. Partial or complete stomatal closure during moisture stress to prevent loss of
water during transpiration serves as carbon dioxide diffusion resistance.

Optimum temperatures for net photosynthesis of C3 species range from 25-30 °C. Above this
temperature range, photorespiration increases more rapidly than photosynthesis, resulting in net
photosynthesis decline. By contrast C4 plants, which lack photorespiration, have temperature
optima far higher than C3 plants, and can thus survive in high temperature and high light intensity
situations where C3 plants are typically photosynthetically inactive.

As regards leaf age, maximum photosynthetic efficiency is reached about full leaf expansion and
thereafter, there may be some decline in efficiency. However, the capability of the leaf for
photosynthesis during its life span depends largely on the environment to which it is exposed to.
Older shaded leaf may lose the ability to photosynthezise efficiently at high light intensities but
remain photosynthetically efficient at lower light intensities which occur at the lower canopy. One
reason for reduced photosynthetic efficiency in older leaves may be a relative increase in their
respiration rate with age.

Unlike those for individual leaves, the over-all photosynthetic efficiency of a crop canopy is quite
different because it involves the aggregate response of many leaves which occupy different
positions, and, therefore, environments, in the canopy.

Light interception by the canopy. The percentage of the incident light intercepted by the canopy
is a function largely of the leaf surface area of the crop. In the early stages of growth, there is
sufficient leaf area available to intercept radiation, and much is wasted by striking the soil. At some
stage, however, a closed canopy should be formed such that almost all the incident radiation is
absorbed by the crop. A convenient measure of the “leafiness” of a crop is the leaf area index (LAI)
which is defined as the ratio between leaf area of a crop, and the area of ground occupied by
the crop. In some crops like the soybean, the percentage light intercepted is directly proportional to
the LAI up to the point when 95% of the light was intercepted (defined as the critical LAI). The
production of LAI above the critical value represents unnecessary leafiness and a waste of
photosynthates. However, the critical LAI for a crop is not absolute but depends on the intensity of
the incident radiation. Similarly, when comparison are made across canopies formed by different
planting arrangements, crop growth rate depends on the amount of light intercepted by each canopy
arrangement, at least until a closed canopy is formed and all the incident light is intercepted. For
soybeans, the rate of dry matter production remains constant for LAI greater than the critical LAI
(referred to as the critical LAI response). For some crops like kale and rice, however, crop growth
rate decline when LAI exceeds the critical value, due to the `parasitic’ effects of lower shaded
leaves in the canopy, which are not photosynthesizing sufficiently to make up for respiration. This is
termed an optimum LAI response.

Light distribution through the canopy. Incident light striking a leaf surface is either reflected by
the leaf, transmitted through it, or absorbed. Transmission in the visible range is greater at
wavelengths in the 520-640 nm region (i.e. green). Consequently, there is a change in the quality of
light reaching the leaves in the lower canopy. Suggestions for improving light distribution through
the canopy include more diffuse spatial arrangement of leaves, more erect leaves and smaller leaf
size to increase the amount of `flecking’ of direct radiation onto lower leaves.

a. Photosynthesis and crop yield.

There are a number of possible approaches to improving the photosynthetic efficiency of the
plant canopy in order to increase total output or yield. Among the options include: 1)
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improvement of the photosynthetic efficiency of the individual leaves, and 2) the manipulation of
the canopy structure so as to optimize the light distribution over the canopy leaf surface.
However, there is a considerable debate whether higher grain yield potential could be achieved
by increasing total canopy photosynthesis. There is no doubt that increasing canopy
photosynthesis may have a direct effect on yield in such crops where the economic yield/product
is in the vegetative growth (e.g. sugarcane, fibers, leafy vegetables and turf grass). In many
grain species, however, total biological dry matter production is already more than adequate and
it is apparent that further increases will not necessarily increase grain yield. The problem thus
becomes a matter of changing the relative partitioning of the photosynthates by the plant
into vegetative and economic grain components. Consideration of the relative partitioning of
photosynthates between vegetative and reproductive components by the plant has led to the
concept of harvest index, which is defined as the proportion of the total biological yield
which is recovered as economic yield (Donald, 1962, 1968). Harvest index of important grain
crops is given in Table 9.

Table 9. Harvest index of some grain crops.

Crop Harvest Index

Wheat 0.40-0.55
Corn 0.40-0.55
Sunflower 0.30-0.35
Dry beans 0.45-0.55
Lentils 0.45-0.55
Soybean 0.25-0.35
Sorghum 0.40-0.55

This problem is particularly evident in indeterminate crops such as grain legumes, in which
continued vegetative growth occurs during pod development. This vegetative dry matter
production represents photosynthates which could have been diverted into grain.