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EDITORIAL TEAM
Chief Editor : Dr.V.UMAMAHESH
Assistant Professor (Plant Physiology)
Agricultural College, Naira.
Co-Editors: Dr.K.Bala Krishna Reddy, Professor (Plant Physiology)
Dr.P.V. Rao, Professor (Plant Physiology)
Dr.A.Siva Sankar, Professor (Plant Physiology)
Dr.D.Vishnu Vardhan Reddy, Professor(Plant physiology)
Dr.K.L.Narasimha Rao, Professor (Plant Physiology)
Dr.V.Raja Rajeswari, Professor (Plant Physiology)
Dr. K. Ashoka Rani, Professor (Plant Physiology)
Dr. G. Rama Rao, Assoc.Professor(Plant Physiology)
Smt. V.Jayalalitha, Asst.professor (Plant physiology)
Dr.B.Srikanth, Asst.professor (Plant physiology)
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ACHARYA N.G. RANGA AGRICULTURAL UNIVERSITY

Department of Plant Physiology
Detailed Lecture Outlines
1. Course No : PPHY 161
2. Course title : CROP PHYSIOLOGY
3. Credit hours : 3 (2+1)
4. General objective : To impart knowledge to the students on different plant

metabolic processes and their functions in plants.
5. Specific objectives
a) Theory
By the end of the course, the student will be able to
i) Study the growth and development of plants
ii) Study the effect of nutrients and Plant growth regulators, and their
applications in agriculture.
iii) Understand the physiology of seeds and fruit ripening
b) Practical
By the end of the practical exercises, the students will be able to
i) Understand various plant metabolic processes, at different stages of plant

growth, which lead to development.
A) Theory Lecture Outlines
1. Introduction – definition of crop physiology – importance in agriculture and

horticulture.
2. Seed physiology – seed structures – development of embryo, endosperm,
perisperm and seed coat – morphological, physiological and biochemical
changes during seed development.
3. Seed physiology – physiological maturity – morphological and physiological
changes associated with physiological maturity in crops with examples –
harvestable maturity – seed viability and vigor – factors affecting seed viability
and vigor.
4. Seed physiology – methods of testing seed viability and vigor – germination –
utilization of seed reserves (carbohydrates, fats and proteins) during seed

germination – morphological, physiological and biochemical changes during

seed germination – factors affecting seed germination.
5. Growth and development – definition – types of growth – determinate and
indeterminate growth – monocarpic and polycarpic species with examples-
initiation and development of vegetative and reproductive structures.
6. Growth and development – measurement of growth – growth analysis –
growth characteristics – definitions and mathematical formulae.
7. Crop water relations – physiological importance of water to plants – water
potential and its components-Importance of water potential-Active and passive
uptake of water-measurement of water status in plants.
8. Crop water relations – transpiration – definition – significance – structure of
stomatal complex in monocots and dicots – role of stomata in transpiration –
transpiration in relation to crop productivity – Water Use Efficiency (WUE) –
WUE in C3, C4 and Crussulacean Acid Metabolism (CAM) plants – WUE of
major field crops – factors affecting WUE.
9. Photosynthesis – energy synthesis – Cyclic and Non cyclic photo
phosphorylation- carbon dioxide fixation – C3 pathway.
10. Photosynthesis – carbon dioxide fixation – C4 and CAM pathways – methods
of measuring photosynthesis.
11. Photosynthesis – photorespiration – factors affecting photosynthesis (light,
carbon dioxide, temperature, water stress, water logging, salinity, weeds /
weedicides etc.)
12. Photosynthesis – photosynthetic efficiency – significance of C3, C4 and CAM
pathways – relationship of photosynthesis and crop productivity.
13. Translocation of assimilates – phloem loading – apoplastic and symplastic
transport of assimilates – mechanism of phloem transport – phloem unloading
14. Source and sink concept – dry matter partitioning – harvest index of crops.
15. Respiration and its significance – importance of glycolysis, Tricarboxylic Acid
(TCA) cycle and Pentose Phosphate Pathway.
16. Respiration – interrelationship of respiration and photosynthesis – growth
respiration and maintenance respiration – alternate respiration – salt
respiration – wound respiration – measurement of respiration.
17. Nutriophysiology – definition – essential elements – criteria of essentiality of
elements – classification of plant nutrients based on their Biochemical role
and physiological function.
18. Nutriophysiology – physiology of nutrient uptake – active and passive uptake
of nutrients – functions of N, P, K, Ca and Mg.
19. Nutiophysiology – functions of Fe, Zn, Mn, Cu, B, Mo, Cl, Na and Si - their
mobility in phloem
20. Nutriophysiology – Deficiency and toxicity symptoms of plant nutrients
21. Nutiophysiology – foliar nutrition – hydroponics – solution and sand culture

22. Photoperiodism and flowering – importance of photoperiodism – classification

of plants based on photoperiodic responses – perception of photoperiodic
stimulus – Biological clock.
23. Photoperiodism – phytochrome – flowering hormones – vernalization and
flowering – importance of vernalization in relation to crop productivity.
24. Plant growth regulators – occurrence, biosynthesis, mode of action and
physiological role of auxins
physiological role of gibberellins
physiological role of cytokinins
physiological role of Abscisic Acid (ABA)
physiological role and ill effects of ethylene
29. Plant growth regulators – novel plant growth regulators – commercial
application of plant growth regulators in agriculture and horticulture
30. Senescence and abscission – definition – classification – theories of
mechanism and control of senescence – physiological and biochemical
changes and its significance – abscission and its relationship with
senescence.
31. Post harvest physiology – seed dormancy – definition – types of seed
dormancy – advantages and disadvantages of seed dormancy – causes and
remedial measures for breaking seed dormancy with examples – optimum
conditions of seed storage – factors influencing seed storage- International
Seed Testing Association (ISTA) standards
32. Post harvest physiology – fruit ripening – metabolic changes during fruit
ripening – climacteric and non-climacteric fruits – hormonal regulation of fruit
ripening (with etherel, Chloro Choline Chloride (CCC), polaris and
paclobutrazol) – use of hormones in increasing vase life of flowers
References:
1) Bidwell, R.G.S 1995, Plant Physiology, Macmillan Publishers Co., New

York .
2) Delvin, R.M. and Witham, F.H. 1986, Plant Physiology CBS Publishers &
Distributors, Delhi.
3) Frank, B. Salisbury and Cleon, W. Ross, 2005. Plant Physiology, CBS

Publishers & distributors, Delhi.

4) Gardener, P., Brent Pearce, R. and Roger, L. Mitchell, 1985. Physiology

of Crop Plants. Jodhpur Scientific Publications, Jodhpur.
5) Hopkins WG & Huner NPA. 2004. Introduction to Plant Physiology. John

Wiley & Sons
6) Lincoln Taiz and Eduardo Zeiger 2006, Plant Physiology, Sinauer

Associates, Inc. Publishers, Sunderland, Massachusetts.
7) Ray, G. Noggel and George, J. Fritz 1991. Introductory Plant Physiology.

Prentice Hall of India Pvt. Ltd., New Delhi.
*******

Lecture-1
IMPORTANCE OF CROP PHYSIOLOGY IN AGRICULTURE AND
HORTICULTURE
Genetic potential of a plant and its interaction with environmental factors
decides its growth and development by influencing or modifying certain internal
processes. Plant physiology studies about these internal processes and their
functional aspects. It helps to understand various biological processes of the plants
like Photosynthesis, respiration, transpiration, translocation, nutrient uptake, plant
growth regulation through hormones and such other processes which have
profound impact on crop yield.
Hereditary potential
Internal processes & Plant growth & Development

conditions
External environment
Crop: is a group of plants grown as a community in a specific locality and, for a

specific purpose.
1.1 Crop Physiology- Definition :

Crop physiology is the study of the ways in which plant physiological processes
are integrated to cause whole plant responses in communities.
The subject matter of crop physiology includes the ways in which the knowledge
of plant physiology is applied for better management of crops.
1.2 A BRIEF HISTORY OF CROP PHYSIOLOGY:
1771- Joseph priestly : Plants could regenerate oxygen in the atmosphere
1779- Ingenhouz : Studied the role of light in Photosyntheis
1804- De Saussure: Plant uptake of mineral nutrients and nitrates from soil.
1837- Boussingault: Nitrogen uptake by plants both from soil and atmosphere
1865- Julius Sachs :published ‘Experimentelle Pflanzenphysiologi’ . By this time

all branches of plant physiology were completely established, but their development
in the next fifty years showed little concern with crop productivity and yield, except in
the area of mineral nutrition. In this period essentiality and role of many mineral
nutrients was established. Towards the end of that period herbicide physiology also
became prominent.
W.LBalls (1915): Crop physiology, with the aim of understanding the dynamics of
yield development in crops, really began with the work of W.L.Balls. Along with

Holton he analysed the effects of plant spacing and sowing date on the
development and yield of Egyptian Cotton plants with in crop stands, not in
isolated plants. It was from his work the term ‘crop physiology’ came in to
existence. From then onwards, various scientists have started applying the
advances in physiological knowledge for better crop management.
1924- In England- a rapid development of the methods of growth and yield analysis
by different investigators (V.H.Blackman, F.G.Gregory, G.E.Briggs etc.) was started.
With the development of various methods of growth analysis, they started explaining
‘the physiology of crop yield’
1947: The concept of LAI (Leaf area index) was developed by D.J.Watson. This
index has provided a more meaningful way of analyzing growth in crops, and
stimulated renewed interest in crop physiology.
1950’s: Studies on photosynthetic rate of the leaf and the loss of photosynthates by
respiration was studied by the development of ‘Infra Red Gas Analysis
(IRGA)’method. This method has facilitated the estimation of short term rates of
Photosynthesis and respiration by crops in the field.
1953: Monsi and Saeki explained about the manner of light interception by the crop
canopy with their concept of light interception coefficient.
1963: Hesketh and Moss showed that photosynthesis by leaves of Maize,

Sugarcane and related tropical grasses could reach much higher rates, with less
marked light saturation, than leaves of other plants(This was the starting point for
research to find other photosynthetic CO2 fixation path ways like C4, and CAM
Mechanisms). The differences in pathway are associated with differences in
photosynthetic rate, in response to light intensity, temperature and oxygen level, in
photorespiration, in leaf anatomy and chloroplast morphology, in rate of
translocation, and in the efficiency of water use, which can have profound effects on
the physiology of yield determination.
Later on several research works were carried out to understand the

processes like translocation of food materials, their partitioning towards economic
yield, storage mechanisms, physiology of flowering, effect of stressful environmental
factors on crop growth and development, role of plant growth regulators in
increasing the crop productivity etc. All these areas have enriched the knowledge of
physiological processes and their role in deciding the crop yield.
1.3 IMPORTANCE OF CROP PHYSIOLOGY IN AGRICULTURE AND HORTICULTURE:
Many aspects of Agriculture and Horticulture can benefit from more

intensive research in plant physiology to provide practical solutions in agriculture
and horticulture. Understanding the physiological aspects of seed germination,

seedling growth, crop establishment, vegetative development, flowering, fruit and

seed setting and crop maturity provides a reasonable scientific base for effective
monitoring and beneficial manipulation of these phenomenon’s. Studying this
phenomenon with a view to develop better crop management practices forms the
subject matter of crop physiology. The importance of physiology in agriculture and
horticulture can be seen from the following examples;
 Seed is the most important input in agriculture. Germination of seed and

proper establishment of seedling depends upon various internal and external
factors. Knowledge in Seed physiology helps in understanding of different
physiological and morphological changes that occur during germination. Any
deviation in these processes causes Seed dormancy. The dormant condition
of the seed bars immediate use of harvested seed for next crop which is
important in intensive agriculture. By understanding the causes and effects of
this problem, Crop physiologists have come up with different methods of
breaking the seed dormancy.
Example: When ever Paddy is used as a seed material in the very next
season it is recommended to treat the seed either with HNO3 or with GA.
 The first prerequisite for higher yields in crops is high total dry matter
production per unit area. High dry matter production is a function of optimum
leaf area (Optimum leaf area Index) and Net Assimilation rate.
(CGR = LAI X NAR).
Example: Pruning operation in horticultural crops like Mango is done based
on this principle of proper canopy management for better photosynthesis.
 Total amount of dry matter produced less the photosynthates used in

respiration is the net product of photosynthesis. Economic yield depends on
how the dry matter is distributed among different organs of the plant. Partition
of total dry matter amongst the major plant organs is of interest to the farmers
as they are more interested in its partition towards economic yield.
Example: excessive vegetative growth period in Ground nut produces less
number of Pods as the reproductive period gets constricted. Thus, groundnut
varieties with relatively extended period of reproductive growth are desirable.
 The use of herbicides to kill unwanted plants is widespread in modern
agriculture. Majority of Herbicides -about half of the commercially important
compounds—act by interrupting photosynthetic electron flow.(Ex. Paraquat,
diuron). When the electron transport is blocked it virtually stops light reaction
of photosynthesis. When light reaction is stopped the dark reaction does not
happen and thus CO2 is not fixed as carbohydrate. Therefore the weed is
killed by starvation.

 Nutriophysiology is yet another important area to under stand crop physiology.

For the healthy growth of a crop around 16 essential elements are required.
Knowledge of nutriophysiology has helped in identification of essential
nutrients, ion uptake mechanisms, their deficiency symptoms and corrective
measures. It also helps to check the toxicity symptoms of various nutrients.
Examples:Zinc deficiency leads to Khaira disease in Rice. This can be

controlled either by soil or by foliar application of Zinc sulphate.
 Response of plant to the relative length of day and night is called as

photoperiodism. This concept was used to choose photo insensitive
varieties. The semi dwarf Rice varieties that have revolutionized Indian
agriculture, are lodging resistant, fertilizer responsive, high yielding and
photo insensitive. Photo insensitivity has allowed rice cultivation in non-
traditional areas like Punjab. Even in traditional areas rice-wheat rotation has
become possible only due to these varieties.
 Plants can regulate their growth through internal growth mechanisms involving
the action of extremely low concentrations of chemical substances called
Plant growth substances, phytohormones or Plant growth regulators. The
regulation of flowering, seed formation and fruit setting has been controlled
through the application of different hormones at the appropriate time of plant
height and age.
Example:Commercial preparations of rooting compounds are available

(Indole Buteric Acid @250 ppm) that promote callus and root formation which
can improve establishment from stem cuttings
 Indian agriculture being predominantly rain fed in nature, development of
drought resistant varieties is very important. Root zone depth, density of roots,
plant water potential, relative water content, water use efficiency, xerophytic
characters of leaves etc. are some of the characters helped to bred drought
tolerant varieties and to develop efficient irrigation management practices
(sprinkler and drip irrigation).
Among Several physiological approaches, transpiration efficiency or water

use efficiency is the most dependable trait, which is “the amount of dry
matter produced per unit amount of water transpired”. The importance of
water use efficiency (WUE) in influencing grain yield under water limited
conditions can be explained by the following model give by passioura.
Grain Yield = T x TE x HI
Where T = Total transpiration by the crop canopy
TE = Transpiration Efficiency or WUE
HI = Harvest Index (Economic Fraction of Dry matter)

This relationship provides an analytical tool to select the genotype with high
levels of T and TE.
 Post harvest losses of agriculture and horticulture are causing a great
distress to farming community. Moisture and temperature are the two
important factors causing physiological changes that reduce the post harvest
quality of grains. Control of moisture content and maintenance of low
temperatures have proved effective in storage of grains. Being perishable in
nature the magnitude of post harvest loss is comparatively higher in
horticultural crops.
Example: In recent years a method called ‘modified atmospheric storage’
was developed for prolonged post harvest life of fruits and vegetables.
Shelf life of cut flowers can be increased by application of kinetin

(cytokinin). This will reduce the burst of ethylene and thus reduces the rate of
senescence.
Thus, physiological understanding of crop plants provides the

fundamental scientific base about various aspects of metabolism, growth and
development. This is immensely important for crop improvement or
technology improvement in agriculture or horticulture.
*******
ANNEXURE
DCMU: Di chloro phenyl di -methyl urea (chemical name of Diuron)
10

Lecture-2
SEED PHYSIOLOGY
Much of the success on modern agriculture depends on the availability of high

quality seeds with good genetic potential and proven performance in germination,
emergence, and vigorous Vegetative growth.
Seed: Seed is a fertilized mature ovule containing an embryonic axis (embryo),

stored food material( endosperm) and a protective covering (seed coat or testa).
2.1 Seed structures:
Living embryo, the most important part of seed, consists of two structures (i)
embryonic axis and ( ii ) cotyledon(s). The embryonic axis is composed of three
parts namely ( i ) radicle (embryonic root), the hypocotyls (point of attachment of
cotyledons) and (iii) plumule (the shoot apex with the first true leaves). The three
parts of embryonic axis are easy to identify in dicots, but in monocots (especially in
the family Gramineae) their identification is difficult. In monocots, there is only one
cotyledon, which is reduced and modified to form the scutellum. The basal sheath
of cotyledon is elongated to take the shape of coleoptile, while in some cases (e.g.,
maize), the hypocotyls is modified to form mesocotyl. The base of hypocotyls
sheathing the radicle is termed as coleorhiza.(Figure.1)
Figure.1 Seeds of common bean (Phaseolus vulgaris ) (left), and Maize (Zea mays) (Right ), a
monocot, a dicot, showing the embryo.
*Figure.1 is from reference .5
11

2.2 Seed development: There are many variations in the pattern of seed
development when the entire plant kingdom is considered. However, the general
process of seed development is almost similar. This includes
 EMBRYO DEVELOPMENT
 ENDOSPERM DEVELOPMENT
 SEED COAT DEVELOPMENT
2.2.1 EMBRYO DEVELOPMENT:
Embryo is a connecting link between two generations of a plant and provides a

continuity of genetic material. In most of the Angiosperms, the embryo and
endosperm start their development as soon as double fertilization is over. Embryo is
diploid in its genetic constitution and is developed from the zygote formed by
fertilization between egg cell & one of the sperm nuclei. The embryo or embryonic
axis represents the life of a plant in miniature.
The size and shape of embryo varies among the plant species. In Monocots
(Ex.wheat) where the endosperm is well developed, the embryo occupies less space
of the seed than in dicot species. Exceptionally, the cotyledons of Castor are thin and
flattened (leaf like structure) since endosperm acts as a storage structure.
2.2.2 ENDOSPERM DEVELOPMENT:
Endosperm serves as the principal nutritive source (food material) for the
embryo during seed development and germination. Endosperm development
precedes embryo development. Endosperm is developed from primary endospermic
nucleus which is a resultant of fusion between two polar nuclei and one of the sperm
nuclei (which is formed from the division of generative nucleus).Thus, the endosperm
of Angiosperms is Triploid (3n) and its development follows any one of the following
three types.
A) Nuclear endosperm: In the nuclear type of endosperm development, the primary

endosperm nucleus divides by repeated mitotic free nuclear divisions without the
formation of walls. Ex. Wheat.
B) Cellular endosperm: In the cellular type of endosperm development, the first

nuclear division of the primary endosperm nucleus is followed by the formation of
either a longitudinal or transverse cell wall in the central cell. Subsequent nuclear
divisions and wall formations result in the formation of a cellular type of endosperm,
e.g., Petunia, Datura, Balsam, etc.
C) Helobial endosperm: It is an intermediate form of nuclear & cellular endosperm

development. Helobial type of endosperm development is prevalent in
monocotyledons.
12

In monocots development of endosperm reaches to its maximum by the time of

maturity. However, in dicots endosperm development is not so conspicuous because
of its continuous utilization by the embryo or it may remain as a small part of the
seed (e.g. Groundnut and Soyabean).
Seeds with well developed endosperm are called endospermic or albuminous

while those with small amounts of endosperm are non- endospermic or
exalbuminous.
2.2.2.1 EXAMPLES OF ENDOSPERMIC SEEDS:
Monocots: Rice, Wheat
Dicots: Castor, opium
Legumes: Fenugreek
NON ENDOSPERMIC SEEDS
Monocots: Orchids, Water plantain
Dicots : Grams, Peas, Beans
2.2.2.2 ALEURONE LAYER:
In cereals and endospermic legumes the majority of cells in endosperm are non-
living. However, the living cells are present in the outermost layer of endosperm and
this layer is known as aleurone layer. During the development of endosperm, cells in
aleurone layer are filled with protein granules and most of them are enzymatic in
nature. Consequently this layer functions both as a storage tissue and also secretes
enzymes of hydrolytic nature (like α-amylase) necessary for germination.
2.2.2.3 PERISPERM : In some of the plant species like coffee and Pig weed,
endosperm is absent and the perisperm acts as a storage tissue. Perisperm (2n) is
developed from maternal nucellus.
2.3 SEED COAT DEVELOPMENT:
During seed maturation the outer structure of ovules namely integuments

undergo marked recognition to form the seed coat (testa). The seed coat acts as a
protective barrier between the embryo and the external environment. The color and
the texture of the seed coats vary from species to species and with in the species.
Almost all seeds bear a scar like point called hilum. This is the point at which a
seed remains joined with the funicles. The micropyle, which is a small hole at one
end of hilum, is present in seed coats of many species. Sometimes hairs or wings
are also present in testa; these structures help in seed dispersal, e.g., in willow, lily
etc. Seed coat may also bear outgrowths of the hilum region, viz, strophiole which
13

controls movement of water into and out of seed, and the aril, which contain
chemicals. For example, the aril of nutmeg is used as a source of spice mace. In
castor bean, the aril, which is associated with micropyle, is called caruncle.
2.3 MORPHOLOGICAL AND PHYSIOLOGICAL CHANGES DURING SEED

DEVELOPMENT:
1. Soon after fertilization the embryo starts developing as soon as endosperm

completes its development. This phase of development is characterized by
cell division and differentiation.
2. Several sub-cellular organelles viz.,Plastids, Mitochondria, Ribosomes,
Golgi complex and Endoplasmic reticulum become recognizable immediately
after cell formation.
3. The sub-cellular organelles undergo changes in their arrangement and
activities. They also participate in the processes leading to the formation of
embryo.
4. The embryo shows variation in shape depending upon its location ( Folded
cotyledons in Cotton, Bent in Fabaceae, Spatulate in Lamiaceae and
peripheral in Caryophyllaceae)
5. Variation in the ratio of embryo and endosperm
- In dicots the developing embryo utilizes endosperm completely. Therefore
the ratio of embryo to endosperm is more.
- In monocots the ratio of embryo to endosperm is less.
2.4 BIOCHEMICAL CHANGES DURING SEED DEVELOPMENT:
1. The seed increases in its weight due to synthesis and deposition of seed
reserves like starch, fat, protein, nucleic acid and phytine.
2. Starch is the major carbohydrate that increases rapidly in the endosperm.
3. As development proceeds, the proteins (nitrogen) accumulate in the
cotyledons of the embryo.
4. The synthesis of proteins in the embryonic axis will be accompanied by
increase in RNA & DNA (nucleic acids) due to formation of new cells.
5. There will be simultaneous accumulation of storage lipids (lipid bodies) in the
cell membranes between phospholipid layers.
6. PHYTIN- a rich source of Phosphorus is deposited as myo-inositol hexa
phosphoric acid. This is found in the endosperm and aleurone cells of many
seeds. (During germination phytin is hydrolyzed by the enzyme phytase to
give inositol, which participates in numerous biosynthetic pathways essential
to the developing embryonic axis)
7. Seeds also synthesize minor constituents like alkaloids (caffeine-in
Coffee),Glycosides (Synigrin in Musturd), vitamins (Thiamin,Biotin and
14

ascorbic acid),inhibitors(Coumarin,ferulic acid and ABA)and hormones

(Auxins,gibberellins etc.,)
8. There will be a change in the pigmentation of seed coat of peas which
become colorless due to loss in the chlorophyll in chloroplasts.
9. Moisture content drops to 10-15% as a consequence of increase in depletion
of seed reserves.
10. Thus, the above biochemical changes of seed indicate the complexity of
development and maturity.
*******
15

Lecture-3
SEED PHYSIOLOGY
Crops are to be harvested at an optimum stage of maturity to get a

qualitative and quantitative yield. Seed yield and its quality depends on number of
factors .Time of sowing and harvesting stages are among the major considerations in
deciding the seed quality and productivity.
3.1 PHYSIOLOGICAL MATURITY: Identification of harvesting stage in terms of

easily measurable parameters is an important problem being faced by the farmers.
Physiological maturity or Biological maturity in terms of average moisture
content and maximum dry weight of seed is one such parameter on which farmers
can depend to harvest maximum.
Physiological maturity can be defined as “the stage at which the seed reaches
to its maximum dry weight, viability and vigor capability”.
3.1.1 MORPHOLOGICAL AND PHYSIOLOGICAL CHANGES ASSOCIATED WITH

PHYSIOLOGICAL MATURITY IN CROPS – SOME EXAMPLES:
Determination of physiological maturity in crops is very essential. Certain

morphological and physiological changes in different parts of the plants can serve as
a guideline to decide the physiological maturity. Effective and practicable guidelines
in this regard help the farmers to reap a good harvest.
- Formation of black layer in placenta region near the point of kernel

attachment in Maize and Sorghum.
- 35 to 40 days after anthesis in Sorghum.
- Change of internal pod walls in to brown color and attainment of pink
colored seed coat in groundnut.
- Change of color of involucre bracts from green to yellow in Sunflower.
- Measuring BRIX reading (Total soluble solids)in sugar cane by ‘Brix sugar
hydro meter’. If the reading is 17 or more the crop is ready for harvest.
Sucrose content of the cane and brix can be estimated with hand refracto
meter.
3.2 HARVESTABLE MATURITY:
Generally, farmers decide the maturity period for harvesting in terms of crop
duration from the time of sowing. However, this will change by the variety, soil fertility
and climatic conditions. Such harvest, which coincides with the ripeness of seed
beyond physiological maturity, is referred to as harvestable maturity. If the seeds
are retained on the mother plant beyond physiological maturity, several
morphological and physiological changes takes place in seeds and result in the
decline of seed quality and yield. For example,
16

- Rice crop harvested at 15% seed moisture content instead of 21%, results
in a yield reduction by 20 per cent.
- Harvesting the seed of Green gram crop after 4 weeks of pod initiation
reduces the yield compared to a harvest of 2-3 weeks pod initiation.
3.3 SEED VIABILITY AND VIGOR
Generally the terms viability and vigor are misused at different levels. To most
of the seed technologists viability means the capacity of a seed to germinate and
produce a normal seed ling. In another sense viability denotes the degree to which a
seed is active and possesses enzymes capable of catalyzing metabolic reactions
needed for germination and seedling growth. In this context, a given seed may
contain live and dead tissue and they may are may not be capable of germination i.e
they may be viable or non viable.
Vigor is usually defined as that condition of active good health and robustness
in seeds which upon planting, permits germination to proceed rapidly under a wide
range of environmental or field conditions. A seed is said to be vigorous when it
shows uniformity in germination and plant development under non-uniform
conditions.
In most of the cases vigor and viability are probably the highest at the time of
harvest and decrease gradually in storage. It is the vigor that declines first followed
by viability (Fig.2). The potential with which a seed produces a healthy seedling
depends on the vigor of seeds.
Figure.2 . Loss of viability and vigor during storage period
*Figure is from seed science and technology (by A.K.Joshi & B.D.Singh)
3.4 FACTORS AFFECTING VIABILITY AND VIGOR:

There are several factors affecting viability and vigor in seeds. Among them the
most important are
17

3.4.1. Genetic make-up
Vigorous seeds produce seedlings of good health and natural robustness. The
vigor of the seeds depends on the genome history of the individual seed and the
environment in which it is sown.
Hybrid seeds normally possess greater vigor due to their inherent genetic
make up. Genetic factors such as hard-seededness, resistance to diseases, and
seed chemical composition influence the expression of seed vigor.
Viability is also influenced by the genetic make up of the seed. Some kinds of
seeds are inherently short lived (e.g. onion, soybeans, ground nut etc.) and quickly
loose their viability.
3.4.2 Time of harvest
Generally, seed viability and vigor are maximum at the time of physiological
maturity. After physiological maturity seeds begin to deteriorate at varying rates
depending on genetic factor and on the conditions of storage environment
3.4.3 Environmental factors during seed development

a) Temperature
b) Moisture availability
c) Soil fertility
Fluctuating temperature during seed formation and maturity , Pre-harvest rain and
Pre harvest environment of high humidity and warm temperatures will affect seed
viability and vigor.
The amount of moisture in the seeds is the most important factor influencing seed
viability during storage. Generally if the seed moisture content increases storage life
decreases. If seeds are kept at high moisture content the losses could be very rapid
due to mould growth .very low moisture content below 4% may also damage seeds
due to extreme desiccation in some crops.
Seeds developed under moisture stress, nutrient deficiency, extreme

temperatures, etc. often result in light, shriveled seed or poor-vigor seeds.
3.4.4 Mechanical damage during threshing, Cleaning and transportation:

The rate of deterioration increases due to mechanical injury at the time of
harvesting and processing.
18

3.4.5 Infestation by micro organisms of seed in storage:
Storage fungi adversely affect the seed by bringing down the seed viability,
seedling vigor and also affect the chemical composition of seeds.
Aspergillus, Penicillium. Rhizopus were some most commonly occurring storage
fungi which reduces seed germination and seedling vigor and causes a variety of
symptoms on seedling.
The microflora activity is controlled by Relative Humidity temperature and
Moisture content of seed.
*******
19

Lecture-4
SEED PHYSIOLOGY
4.1 METHODS OF TESTING SEED VIABILITY AND VIGOR
There are several tests to evaluate viability and vigor in seeds. The tests
intended to determine viability and vigor must meet certain principles. The ideal test
should be
 Simple with out involving sophisticated equipment.
 Rapid- its procedure should take less time to give results.
 Effective in detecting minute as well as large differences.
 Equally useful for evaluating either individual seed or population of seeds.
4.1.1 VIABILITY TESTS:
A) Tetrazolium test: This test is often referred as a quick test since it can be
completed with in hours. The test is usually based on measuring the activity of
dehydrogenase enzyme in the tissues of embryo. It is conducted by using 2,3,5-
Triphenyl Tetrazolium Chloride (TTC)solution.
Principle: Any living tissue must respire. In the process of respiration the enzyme
dehydrogenase will be in a highly reduced state. When the seed is treated with
colorless tetrazolium solution, the living tissue of the seed by virtue of respiration and
having the dehydrogenase enzyme in a highly reduced state gives off hydrogen ions.
These hydrogen ions reduce the colorless tetrazolium solution in to red colored
formazan. Thus, the tetrazolium test distinguishes between viable and dead tissues
of the embryo on the basis of their relative rate of respiration in hydrated state.
2H+ + 2 e-
2,3,5 Triphenyl Tetrazolium Chloride Triphenyl formazan + Hcl
Dehydrogenase
(Colorless) ( Red color)
OXIDISED STATE NADH NAD REDUCED STATE
NADPH NADP
B) Sulphuric acid test:This is usually a non- enzymatic test. The principle involved
in this test is to distinguish the differential coloration of live versus dead tissue when
exposed to sulphuric acid. The living portion of the cut surface of the seed develops
deep rose color with in 5-10 minutes. Though this test takes less time, one must be
careful in handling concentrated sulphuric acid.
A comparison between the above tests clearly indicates that tetrazolium test is
more practicable, since it is quick and does not involve any risk.
20

4.1.2 VIGOR TESTS

a) Speed of Germination:
The speed of germination is an important aspect of vigor and provides a
reasonably good index of vigor of any seed lot. Mathematically speed of germination
is expressed as “co-efficient of Germination” (CG). Higher the value for speed of
germination greater will be the vigor.
100 (A1+A2 ----------- +Ax)
CG = --------------------------------
A1T1+ ------------Ax Tx
A: Number of seeds germinated
T: Time corresponding to A
X: No of days to final Count.
b) Exhaustion Test:
This test is more suitable for cereal seeds. The moist paper toweling or crape
craft paper used in this test has to be rolled loosely along with the seeds placed on
the printed line of the paper. Those seeds which exhaust their reserve food material
and produce seedlings with roots and shoots extending more than two inches are
said to be vigorous.
c) Respiration Test :
Respiration is one of the important processes common to all living organism.
In seeds the rate of respiration and vigor are closely related. Hence, it is natural
assumption that seeds with high vigor would have higher respiration rates . The
differences in respiration rates have been used to distinguish between high, medium
and low vigor seeds in maize, wheat and soybean. This test can be used to detect
loss of vigor due to mechanical damage, chilling, desiccation and other causes.
Other tests like accelerated aging test (AAT), Brick gravel test, Electrical
Conductivity test are also used to estimate seed vigor.
4.2 Mobilization and Utilization of Seed Reserves
Once seeds have imbibed water and are hydrated, metabolic functions begin to
accelerate if dormancy was previously broken. Enzymes to become active upon
hydration and are involved in mobilization of seed storage reserves. Once storage
reserves are breaking down, they are moved as small molecules from the source to
those cells or organs that have a need for the product. In general meristematic and
actively growing cells are energy and nutrient sinks. In germinating seeds,
cotyledons and endosperm are sources.
21

Stored sources of energy and nutrients include:
Starch: Starch is found in amyloplasts. Starch serves as a source of reduced carbon

for respiration and metabolism. Enzymes in involved in starch degradation are
β-amylase, α-amylase and Starch phosphorylase.
Fats (oils): They are Stored in fat bodies. The high energy content of fats is broken
down by β-oxidation in glyoxysomes and used for respiration and metabolism.
Storage proteins: Stored in protein bodies .Storage proteins are rich in amino acids
with abundant nitrogen and sulfur. Amino acids rich in N are asparagine, glutamine
and lysine. Sulfur-containing amino acids are methionine and cysteine.
Storage proteins are broken down by aminopeptidases, carboxypeptidases, and
endopeptidases.
4.3 MORPHOLOGICAL, PHYSIOLOGICAL AND BIOCHEMICAL CHANGES DURING

SEED GERMINATION:
Seed germination is the resumption of active growth of the embryo that results
in the rupture of the seed coat and emergence of the young plant. During the
process of germination seeds undergo several metabolic changes. These changes
can be summarized as follows.
 Initially the seeds imbibe water through the openings in the seed coat. Upon
imbibition the cells become turgid, the seed increases in its size/volume and
the seed coat becomes more permeable to diffusion of gases (O 2 and CO2).
 There will be a change in the sub cellular organization of the embryo,
endosperm and aleurone layer.
 Phytochrome becomes biologically active and triggers the process of
germination in some plant species.
 The tissue of rehydrated seeds activate the enzymes which help in
- Hydrolysis or break down of storage food material (starch, proteins and
fats)
- Transfer of nutrients from cotyledons and endosperm to the growing
points
- Utilization of hydrolyzed products in the synthesis of new material
 Following the activation of enzymes, the embryo initiates growth in terms of
cell division and elongation
 The hormones (GA, Auxins and Cytokinins) necessary for germination will be
synthesized or activated.
 Growth of the root-shoot axis occurs at the expense of the storage tissue.
22

 The enlarging root-shoot axis exerts internal pressure on the seed coat and
results in its rupture. In case of dicots the pressure developed between the
cotyledons helps in rupturing of the seed coat and emergence of growing
point.
 It is primary root that emerges first and supports in the establishment of
seedling
 Finally, the seedling establishes and becomes autotrophic.
4.4 FACTORS AFFECTING GERMINATION:
The effects of the environment on seed germination is quite complex. A few

of the major environmental factors are discussed here.
a. water : Water is the basic requirement for seed germination. It is essential for
enzyme activation. Thus, permits breakdown, hydrolysis, translocation and use of
reserve food material. How ever, extreme moisture may inhibit germination.
b. Air(O2 & CO2) :Most of the seeds germination well under normal concentration of
gases ( 20% O2, 0.03% CO2 and 80% nitrogen gas). Among them adequate supply
of O2 must be available for oxidative process like breakdown of inhibitors.
Rice seeds can germinate even in the absence of O2 (anaerobic conditions),
although the seedlings are weak and abnormal. However, N2 gas has no influence.
c. Temperature: The effect of temperature on germination can be expressed in

terms of cardinal temperatures that are minimum, optimum and maximum
temperature at which germination occurs. The optimum temperature may be defined
as the temperature at which greatest percentage of germination is recorded with in
the shortest period of time. Some of the seeds like wheat and barely can tolerate and
germinate at minimum temperature of 3 – 5o c. The optimum temperature for most
seeds is between 15oC (wheat) to 30o C (rice). Maximum temperatures at which
some species can tolerate and germinate is 350 C (groundnut), 400C (sorghum) and
500C (cantaloupe). [However, some species germinate at temperatures approaching
the freezing point e.g. alpine seeds.]
The above examples provide a clue that seeds differ in their requirement of
temperature for germination. The low temperature pre – treatment before
germination, usually called stratification, and high temperature pre – treatment
before germination, usually called high temperature pre – treatment are to change
the macromolecular structure of the seeds which in its original form in some way
prevents germination.
23

d. Light :The greater promotion of light on germination occurs in red region (660nm)
followed by an inhibition zone in the far red region (730 nm). Such promotary and
inhibitory effects of red and far-red light respectively on germination is mediated by
phytochrome. It is referred to as biologically active in pfr form (far red absorbing
form) and germination can proceed. Exposure to far – red light (730) reconverts
phytochrome to the red absorbing form (pr) and germination is blocked.
e. Soil conditions : Saline conditions inhibit seed germination. High salt

concentration prevents imbibition of water by the seed. The same osmotic effect is
also observed when the fertilizers are placed in close contact with seeds.
********
24

Lecture-5
GROWTH AND DEVELOPMENT
5.1 ‘Growth’ - definition:
Growth and Development are the most fundamental and conspicuous

characteristics of all living organisms. According to dictionary, growth is the
advancement towards maturity and development is a gradual increase in size. The
plant physiological definition of growth is ‘an irreversible increase in mass, weight or
volume of a living organism, organ or cell.
5.2 Growth Curve:
Typical growth pattern of an annual plant is represented in figure.3. This can

be divided into three phases.
1. Lag period of growth:
During this period the growth rate is quite slow because it is the initial stage of
growth.
2. Log period of Growth
During this period, the growth rate is maximum and reaches the top because
at this stage the cell division and physiological processes are quite fast.
3. “Senescence period or steady state period :
During this period the growth is almost complete and become static. Thus the
growth rate becomes zero.
FIG.3 A TYPICAL ‘S’ SHAPED GROWTH CURVE

*Figure 3 is from reference 6
25

If the growth rate is plotted against time, an ‘S’ shaped curve is obtained
which is called sigmoid curve or grand period curve.
The growth curve described above is seen in most cases, although there is a
considerable difference due to variations in plant species as well as environmental
factors. It is also apparent that growth in all parts of a multi cellular plant is not
uniform. In higher plants, it is restricted only to the meristematic zones which are
found near the root and the shoot tips, in the vascular cambium and in certain parts
of young leaves.
Growth pattern of annual crops:
Here the first phase relates to the seed germination and seedling growth. Seeds
germinating below ground are dependent on stored material in the cotyledons until
the seedling emerge in to light and start photosynthesis. Hence, initial increase in
weight is negligible. (LAG PHASE)
The second phase of growth is characterized by a rapid and often linear increase
in dry matter production and terminates with flowering(anthesis). (LOG PHASE)
It is associated with tillering, stem elongation and leaf expansion in cereals. In case
of indeterminate crops such as Cotton, Pigeon pea etc. it is characterized by
formation of branches with large number of leaves.
Initiation of flower buds siginifies end of rapid growth phase (grand period of
growth) and indicates onset of flowering.
The third phase of growth is marked by a reduction in growth rate until growth
ceases at maturity. Assimilates stored in leaves and stems are translocated to
partially sustain seed growth. At the end of this growth period, water is lost from
aerial plant parts, photosynthesis stops and crop ripens (STATIONARY PHASE
AND DEATH).
5.3 TYPES OF GROWTH:

a) Determinate Organs :
Those organs that grow to certain size and then stop growing are called
determinate organs. After their growth is completed they eventually senesce and die.
Examples of such organs are leaves, flowers and fruits etc.
Determinate Growth:
If, reproductive growth starts only after completion of vegetative growth it is called
as determinate growth habit.
Eg. Maize.
26

Indeterminate Organs:
Those organs which grow continuously with the activity of meristems are
indeterminate organs. Examples are roots and vegetative stems of perennials.
These structures always remain youthful, because of the meristematic activity.
Indeterminate Growth:
Here, vegetative and reproductive growth overlaps. This is shown in plants that have
a capacity for both vegetative growth and flowering over an extended period.
Eg. Redgram, Soybean etc.,
5.4 Monocarpic and polycarpic species :
Monocorpic species flower only once and then die. Thus, in a sense mononcarpic
species are determinate as for as the entire plant is concerned.
Ex.Rice, Maize,Sunflower,Sugarcane,sorghum etc.
Polycarpic species flower more than once in life cycle. Here, the vegetative and
reproductive periods overlap each other. This is seen in most of the tree species.
Most monocrapic species are annuals. However, some of them are biennials
and perennials also. Many varieties of bamboos may grow and live for over 50 years
and then they flower and die. Thus bamboos are perennial but monocarpic. All
polycarpic plants are perennials.
5.5 Development :
Growth leads to the development. Development is defined as ordered change or

progress often (but not always) towards a higher, more ordered or more complex
state. However, these two processes are often linked together and occur in
sequence. Growth is a quantitative change in contrast to development which is more
of a qualitative change.
5.5.1 INITIATION AND DEVELOPMENT OF VEGETATIVE STRUCTURES:
5.5.1.1 Root growth: Radicle is the embryonic root. During the seed germination
and seedling formation, it grows to form primary root of the seedlings. A growing
root usually has 4 distinct regions,
1. Root cap
2. Meristematic region
3. The region of cell elongation and
4. The region of differentiation and maturation.
27

The root cap protects the root tip. The meristematic region in young root is
situated just below the root tip. The cells in this region are responsible for growth in
the root. The meristematic region consists of numerous small, compactly arranged
thin walled cells almost completely filled with cytoplasm. They have very small
vacuoles and comparatively large nuclei. Inter cellular spaces are absent. Only a
few cells in the meristem may actually be involved in the longitudinal growth of the
root.
F.A.L. Clowes and B.E. Jupiner (1968) demonstrated that there is a quiescent
center in the meristematic region, where no cell division takes place. This center is
located just above the root tip. It is surrounded by a group of actively dividing cells,
which give rise to the column of cells forming roots.
The region of cell elongation is made up of column of newly derived cells. It is

the elongation of these cells, which causes the root tip to project forward and push
through. Most cells in this region elongate at least 15 folds and increase in diameter
which results in the development of considerable pressure by the elongation of root.
The region of differentiation and maturation:
The cells in the region of differentiation and maturation differentiate into various
tissues, characteristic to the mature root; the epidermis, cortex and stele.
In roots xylem and phloem differentiate only acropetally and as continuation of

the older xylem and phloem in the more basal part of the root.
During differentiation most cells increase in size and vacuolation.
5.5.1.2 Stem growth : The life of stem starts as a plumule. It grows to form the
shoot of the seedling. The longitudinal growth of stem and formation of various
organs like branches, leaves, flowers is the function of stem meristem.
Tunica Corpus Theory:
To explain the cellular organizations of stem meristem, A.Schmidt (1924) first

proposed a tunica corpus theory. Accordingly , most apical meristems contain two
zones, an outer tunica and an inner Corpus. Tunica consists of one to several layers
of cells at the surface of the merisem while corpus cells are beneath the tunica layer.
The cells in tunica divide by anticlinal division i.e in a plane perpendicular to the
surface of the stem, whereas the corpus cells divide in many different ways.
The formation of branches leaves or outer appendages on the stem are initiated
in the formation of a primordium or out growth at the surface of the meristem, just
below the tip. In the formation of aerial organs both tunica as well as corpus layers
are involved. The tunica normally forms the epidermis of the organ derived from the
28

meristem, while corpus cells produce majority of the internal tissues of the new
organ.
Auxins normally promote the elongation of stem. They induce the elongation of
cells. Gibberellins also promote stem elongation and they do this by promoting cell
division as well as cell elongation.
5.5.1.3 Leaf initiation and Growth: Elevations appear on the periphery of the
meristem in a regular pattern. Leaf primordia appear as dome shaped on the
periphery of the stem. They appear at nodal positions of the stem, which have an
intercalary meristem when the leaves are to be produced in pairs; each pair usually
appears to right angle to the preceding pair, the two leaves in a pair generally
opposite to each other.
The growth of individual leaf also follows the typical sigmoidal pattern, just like the
growth of the entire plant. In most plants, the shape and form of leaves are fixed and
little variation found among them.
However, many plants have leaves of different shape. The phenomenon is termed as
heterophylly, which is quite common in aquatic plants.
5.5.2 INITATION AND DEVELOPMENT OF REPRODUCTIVE STRUSCTURES

5.5.2.1 Initiation and Development of Flower:
Once the biochemical requirements for evocation of flowering are completed and
the meristem has reached the point of no return, it develops either into an
inflorescence (a cluster of flowers) or solitary flowers. In most plants, the pattern of
flower initiation and development is almost similar. As an example of flower initiation
in Capsicum annum(Green pepper) the first microscopically visible change in the
shoot apex is the change in its shape. The apex almost becomes flat from conical,
due to the inhibition of growth in the central portion of the meristem. Some
protuberances develop from this meristem in a whorled manner. Floral parts (sepals,
petals etc) are formed due to the development of the protuberances. The outermost
whorl of the protuberances forms the sepal and next to it forms petals and so on.
Most plants produce bisexual flowers containing functional male (stamens) and
female (pistils) parts. Other species contain staminate (male) and pistillate (female)
flowers only on different individual plants.
Auxins and Ethylene stimulates the formation of female flowers, where as

gibberellins increase the ratio of male to female flowers in the cucumber.
Initially, the floral parts are tightly enclosed with in the outer most part, the sepals,
constituting a floral bud. Subsequently expansion of the flower bud in to an open
flower occurs. The cause of the flower opening is usually due to the differential
growth of the inner and outer sides of the sepals and petals.
29

5.5.2.2 Fruit and Seed Development:
The first stage in fruit and seed development is rapid cell division without much
enlargement due to cytokinin production by the endosperm which is growing at this
stage. Various tissues of the parent plant viz, the ovary, receptacle and sometimes
parts of the floral tube may be involved in the formation of fruits.
Following the cell division stage cell enlargement phase of growth proceeds
and this is by auxins produced in the seeds. If the seeds are removed from a
developing fruit, development stops, however it can be restarted again by the
application of auxins.
It was observed that fruit development in cucumber is dependent on auxins
which originate from the ovule, while some fruits respond rather to gibberellins than
to auxin treatment.
At this stage in the development of fruits the concentration of organic acids and
sugars begin to increase followed by decrease in osmotic potential. This is related to
the increasing absorption of water and growth by enlargement of cells.
ANNEXURE
*Figures are from reference 6
30

Lecture-6
GROWTH AND DEVELOPMENT
6.1 MEASUREMENT OF GROWTH: Growth can be measured by a variety of

parameters as follows
A. Fresh Weight
Determination of Fresh weight is an easy and convenient method of
measuring growth. For measuring fresh weight, the entire plant is harvested,
cleaned for dirt particles if any and then weighed.
B. Dry Weight
The dry weight of the plant organs is usually obtained by drying the materials
for 21 to 48 h at 70 to 80oC and then weighing it. The measurements of dry weight
may give a more valid and meaningful estimation of growth than fresh weight.
However, in measuring the growth of dark grown seedling it is desirable to take fresh
weight.
C. Length
Measurement of length is a suitable indication of growth for those organs
which grow in one direction with almost uniform diameter such as roots and shoots.
The length can be measured by a scale. The advantage of measuring length is that
it can be done on the same organ over a period of time without destroying it.
D. Area
It is used for measuring growth of plant organs like leaf. The area can be
measured by a graph paper or by a suitable mechanical device. Nowadays modern
laboratories use a photoelectric device (digital leaf area meter) which reads leaf area
directly as the individual leaves are fed into it.
6.2 GROWTH ANALYSIS:

Growth analysis is a mathematical expression of environmental effects on
growth and development of crop plants. This is a useful tool in studying the complex
interactions between the plant growth and the environment. Growth analysis in crop
plants was first studied by British Scientists (Blackman 1919, Briggs, Kidd and west
1920, William 1964, Watson 1952 and Blackman, (1968). This analysis depends
mainly on primary values (Dry weights) and they can be easily obtained without great
demand on modern laboratory equipment.
The basic principle that underlie in growth analysis depends on two values (1)
total dry weight of whole plant material per unit area of ground (w) and (2) the total
leaf area of the plant per unit area of ground (A).
The total dry weight (w) is usually measured as the dry weight of various
plant parts Viz, leaves, stems and reproductive structures. The measure of leaf area
(A) includes the area of other organs viz, stem petioles, flower bracts, awns and
31

pods that contain chlorophyll and contribute substantially to the over all
photosynthesis of the plants
According to the purpose of the data, leaf area and dry weights of component
plant parts have to be collected at weekly, fortnightly or monthly intervals. This data
are to be used to calculate various indices and characteristics that describe the
growth of plants and of their parts grown in different environments and the
relationship between assimilatory apparatus and dry matter production. These
indices and characteristics are together called as growth parameters. Some of the
parameters that are usually calculated in growth analysis are crop growth rate
(CGR), relative growth rate (RGR), net assimilation rate (NAR), Leaf area ratio
(LAR), Leaf weight ratio (LWR). Specific Leaf Area (SLA), Leaf area index (LAI) and
Leaf area duration (LAD). Accuracy in calculations of these parameters and their
correct interpretation are essential aspect in growth analysis.
6.2.1 Advantages of growth analysis
a) We can study the growth of the population or plant community in a precise
way with the availability of raw data on different growth parameters.
b) These studies involve an assessment of the primary production of vegetation
in the field i.e. at the ecosystem level (at crop level) of organization.
c) The primary production plays an important role in the energetics of the whole
ecosystem.
d) The studies also provide precise information on the nature of the plant and
environment interaction in a particular habitat.
e) It provides accurate measurements of whole plant growth performance in an
integrated manner at different intervals of time.
6.2.2 Drawbacks of Growth Analysis

In classical growth analysis sampling for primary values consist of harvesting
(destructively) representative sets of plants or plots and it is impossible to follow the
same plants or plots through out whole experiment.
6.3 Growth Characteristics – Definition and Mathematical Formulae

The following data are required to calculate different growth parameters in
order to express the instantaneous values and mean values over a time interval. In
the following discussion W, W L, W S and W R are used to represent the dry weights of
total plant(w), dry leaves(wL), stem(W S) and roots(W R) respectively. Whereas A is
the leaf area and P is the land area.
6.3.1 Crop Growth Rate (CGR):C

D.J. Watson coined the term Crop growth rate. It is defined as the increase of
dry matter in grams per unit area per unit time. The mean CGR over an interval of
time T1 and T2 is usually calculated as show in the following formula
32

I W2 – W 1
CGR = -------- X ---------------- (g m-2 day -1)
P T2 – T1
Where CGR is the mean crop growth rate, W 1 and W 2 are the dry weights at
two sampling times T1 and T2 respectively.
6.3.2 Relative Growth Rate (RGR):R

The term RGR was coined by Blackman. It is defined as the rate of increase
in dry matter per unit of dry matter already present. This is also referred as
Efficiency index since the rate of growth is expressed as the rate of interest on the
capital. It provides a valuable overall index of plant growth. The mean relative
growth rate over a time interval is given below.
Loge W 2 – Loge W 1
RGR = ---------------------- (g g-1 day -1)
T2 – T1
6.3.3 Net Assimilation Rate (NAR):E:

The NAR is a measure of the amount of photosynthetic product going into
plant material i.e. it is the estimate of net photosynthetic carbon assimilated by
photosynthesis minus the carbon lost by respiration. The NAR can be determined by
measuring plant dry weight and leaf area periodically during growth and is commonly
reported as grams of dry weight increase per square centimeter of leaf surface per
week. This is also called as Unit leaf rate because the assimilatory area includes
only the active leaf area in measuring the rate of dry matter production.
The mean NAR over a time interval from T1 to T2 is given by
W 2-W 1 Log e A2 – Log e A1

NAR = --------- X ------------------------ (g cm-2 wk-1)
T2-T1 A2 – A1
6.3.4 Leaf Area Ratio (LAR)
The LAR is a measure of the proportion of the plant which is engaged in
photosynthetic process. It gives the relative size of the assimilatory apparatus. It is
also called as capacity factor. It is defined as the ratio between leaf area in square
centimeters and total plant dry weight. It represents leafiness character of crop
plants on area basis.
A
LAR = ------- (cm2g-1)
W
33

6.3.5 Leaf Weight Ratio (LWR)

It is one of the components of LAR and is defined as the ratio between grams
of dry matter in leaves and total dry matter in plants (g). Since the numerator and
denominator are on dry weight basis LWR is dimensionless. It is the index of
leafiness of the plant on weight basis.
WL
LWR = -------
W
6.3.6 Specific Leaf Area (SLA)

It is another component of LAR and defined as the ratio between leaf area in
cm2 and total leaf dry weight in grams. This is used as a measure of leaf density.
The mean SLA can be calculated as
A
SLA = -------- (cm2g-1)
WL
6.3.7 Specific Leaf Weight (SLW)

The reversal of SLA is called as SLW. It is defined as the ratio between total
leaf dry weight n gms and leaf area in cm2. It indicates the relative thickness of the
leaf of different genotypes.
WL
SLW = ------- (g cm-2)
A
6.3.8 Leaf area index (LAI):
D.J. Watson coined this term. It is defined as the functional leaf area over unit
land area. It represents the leafiness in relation to land area. At an instant time (T)
the LAI can be calculated as
LAI = A/P unit less
For maximum production of dry matter of most crops, LAI of 4-6 is usually
necessary. The leaf area index at which the maximum CGR is recorded is called as
‘optimum leaf area index’.
6.3.9 Leaf Area Duration (LAD): d: It is usually expressed as a measure of leaf

area integrated over a time period. Some takes into account both the magnitude of
leaf area and its persistence in time;
it represents the leafiness of the crop growing period. Thus the unit of measurement
of LAD may be in days or weeks or months.
34

LA1+LA2(T2 – T1)
LAD (Leaf area basis) = -----------------------
LA1 + LA2 (T2 – T1)
LAD (LAI basis) = -----------------
This is expressed as cm2 d -1
L
A
I
TIME 1 2
Leaf area duration (Shaded area)
For how many days/weeks functional leaf area is present on the plant can be estimated by
this LAD.
*******
35

Lecture-7
CROP WATER RELATIONS
Almost every plant process is affected directly or indirectly by the water

supply. Decreasing water content is accompanied by loss of turgor and wilting,
cessation of cell enlargement, closure of stomata, reduction in photosynthesis and
interference with many basic metabolic processes. Eventually, continued dehydration
causes disorganization of the protoplasm and death of most organisms.
7.1 PHYSIOLOGICAL IMPORTANCE OF WATER:
The importance of water can be summarized under the following general

headings.
a) Constituent of Protoplasm
Water is important quantitatively and qualitatively, constituting 80 to 90
percent of the fresh weight of most herbaceous plant parts and over 50 percent of
the fresh weight of woody plants. Water is important as a part of the protoplasm as
the protein molecules which constitute the protoplasm framework, changes in
structure if the water content is dropped below certain level. This ultimately leads to
death of plants. It is true that a few plants and organs can be dehydrated to the air-
dry condition or even to the oven-dry condition as in the case of some kinds of seeds
and spores, without loss of viability, but a marked decrease in physiological activity
always accompanies decrease in water content.
b) Solvent
A second essential function of water in plants is as the solvent in which gases,
minerals and other solutes enter plant cell and move from cell to cell and organ to
organ. The permeability of most cell walls and membranes to water in a continuous
liquid phase extending throughout the plant in which translocation of solutes of all
kinds occurs.
c) Reagent
Water is a reactant or reagent in many important processes, including
photosynthesis and hydrolytic process such as the hydrolysis of starch to sugar. It is
also essential for fixation of carbon dioxide or nitrate.
d) Maintenance of Turgidity
Another essential role of water is the maintenance of the turgidity which is
essential for cell enlargement and growth, and for maintaining the form of
herbaceous plants. Turgor is also important in the opening of stomata and the
movements of leaves, flower petals and various specialized plant structures.
Inadequate water to maintain turgor results in an immediate reduction of vegetation
growth.
36

7.2 WATER POTENTIAL AND ITS COMPONENTS:
Water potential or chemical potential of water is a quantitative expression of

the free energy associated with water.
Water potential is symbolized by the Greek letter Ψ (psi) and is defined

relative to the water potential of pure water, which is zero. Hence the value of Ψ is
always negative. The units of water potential are mega Pascal (MPa). It is a relative
quantity and depends on concentration, pressure and gravity at the same
temperature.
Water potential as the sum of component potentials which may be written as

Ψ = Ψs + Ψm + Ψp + Ψg
Where Ψs = Solute or osmotic potential (symbol π)
Ψm = Matric potential (Symbol T)
Ψp = Pressure potential (Symbol P)
Ψg = Gravitational potential (Symbol G)
Osmotic potential
The osmotic potential, Ψs (or π) is the component produced by solutes
dissolved in the cell sap, chiefly vacuolar sap.
Matric Potential
The matric potential, Ψm (or T) refers to water held in micro capillaries or
bound on surfaces of the cell walls and other cell components.
Pressure potential
The pressure potential Ψp (or P) is the turgor pressure produced by diffusion
of water into protoplasts enclosed in walls which resist expansion. In the xylem of
transpiring plants Ψp is usually negative and in guttating plants it is positive as a
result of root pressure.
Gravitational Potential
The effect of gravity, Ψg (or G) is a term of negligible importance within root or
a leaf but becomes important in comparing potentials in leaves at different heights on
trees and in soils.
Upward movement of water in a tree trunk must overcome a gravitational
force of 0.01 Mpa/m and gravity causes drainage of water downward in soil.
The volume of matric water is very small compared the volume of vacuolar
water in parenchyma, therefore Ψm has a negligible effect on the total water
potential Ψ. However, in developing seeds or thick walled cells where the vacuolar
37

water constitutes a small fraction of the total water, matric potential can control the
cell water potential.
Thus for herbaceous plants and annual field crops of a short vertical height
(less than 10 m) the values of the matric potential and gravitational potential are
small and are commonly omitted. Thus
Ψ = Ψs + Ψp or P-π
Water always moves from less negative water potential to more negative water
potential.(Figure.4)
*Figure.4 is from reference 5
7.3 IMPORTANCE OF WATER POTENTIAL:

Water potential is a diagnostic tool that enables the plant scientist to assign a
precise value to the water status in plant cells and tissues.
The lower the water potential in a plant cell or tissue, the greater is its ability to
absorb water. Conversely, the higher the water potential, the greater is the ability of
the tissue to supply water to other more desiccated cells and tissues.
Thus water potential is used to measure water deficit and water stress in plant cells
and tissues.
As a general rule, leaves of most plants rooted in well watered soils are likely to
have water potentials between about -2 and -8 bars. With decreasing soil moisture
supply, leaf water potential will become more negative than -8 bars and leaf growth
rates will decline. Most plant tissues will cease growth completely ( i.e., will not
enlarge) when water potential drops to about -15bars.
7.4 UPTAKE OF WATER

The way in which water is entered in to the root hair and the precise mechanism of
water absorption is has been explained by two different approaches.
(a)Active uptake:
Water is absorbed as a result of activities in the root itself and does not concern with
any process in shoot.
(b)Passive uptake of water:
The governing force of water absorption originates in the cells of transpiring shoots
rather than in root itself.
38

Although the absorption of water by roots is believed to be a passive,

pressure driven process, it is nonetheless dependent on respiration. Respiratory
inhibitors (such as cyanide),anaerobic conditions (waterlogged condition) decrease
in the hydraulic conductance of most roots .These are some supporting points for
active absorption of water. How ever the exact role of respiration and active uptake is
not clear.
Barring few exceptions, it is now believed that uptake of water is a Passive
process. Tension or negative pressure originating at the actively transpiring leaf
surface creates a pulling force for water movement in xylem.(Cohesion-tension
theory of Dixon and Jolly)
The movement of water inside the plant is driven by a reduction in free energy,
and water may move by diffusion, by bulk flow or by a combination of these
fundamental transport mechanisms. Water diffuses because molecules are in a
constant thermal agitation, which tends to even out concentration differences. Water
moves by bulk flow in response to a pressure difference, when ever there is a
suitable path way for bulk movement of water. Thus, water potential difference ( i.e
solute potential and pressure potential) across the cells starting from root hairs to
xylem plays an important role in uptake and transport of water.
Figure is from reference 5
39

7.5 METHODS OF MEASURING WATER STATUS IN PLANTS:
There are two general ways to describe the water status or internal water
balance of plant and plant tissue. The first one is based on the energy associated
with water in the plant tissue. Water potential is considered by most plant
physiologists to be the most useful and significant way to describe the water status of
plant tissues. In terms of water potential, water deficit exists in a tissue when ever its
water potential is less i.e., more negative than zero mega Pascal (Mpa). The water
potential is measured by (1) liquid immersion method (dye method) (2)vapor
equilibration method (Thermocouple Psychrometer) and (3)pressure chamber
method.
7.5.1 Liquid immersion method or dye method or Chardakov’s Falling Drop

Method
Tow graded series of sucrose solutions (ranging from 0.15 to 0.50 molal in
increments of 0.5molality) are placed in test tubes set up in duplicate. Homogeneous
plant tissue is placed into each test tube of one of the series (test series). Only a
drop of methylene blue is mixed into each solution of the second series (control
series) Plant tissue is not added to the control series and the dye does not
appreciably change the osmotic potentials.
After the tissue is incubated for 15 to 30 minutes, It is removed from each

tube. The actual time of incubation can be just long enough for osmosis to proceed
and change the concentration of each solution in the test series; the attainment of
equilibrium is not necessary. After the tissue is removed, a small drop of the
respective control series solutions is introduced below the surface of its
corresponding test solution. If the drop rises in the test solution, it means that the
drop is lighter and that the tissue incubation solution is more concentrated – an
indication that water from the solution entered the tissue. Conversely, if the drop
falls, it means that the test solution is lighter-an indication that water has left the
tissue and diluted the solution. In this latter instance, the water potential of the
solution initially is more negative than that of the tissue. Accordingly, if the density of
the drop from the methylene blue solution is the same as that of the test solution, the
drop will diffuse into the solution uniformly. At this point (called the null point), the
water potential of the tissue and solution is equal.
40

7.5.2 Vapour equilibration (Thermocouple Psychrometer) Method

Psychrometry (the prefix "psychro-" comes from the Greek word psychein,
"to cool") is based on the fact that the vapor pressure of water is lowered as its water
potential is reduced. Psychrometers measure the water vapor pressure of a solution
or plant sample, on the basis of the principle that evaporation of water from a surface
cools the surface.
41

Investigators make a measurement by placing a piece of tissue sealed inside a small

chamber that contains a temperature sensor (in this case, a thermocouple) in contact
with a small droplet of a standard solution of known solute concentration (known Ψs
and thus known Ψw). If the tissue has a lower water potential than that of the droplet,
water evaporates from the droplet, diffuses through the air, and is absorbed by the
tissue. This slight evaporation of water cools the drop. The larger the difference in
water potential between the tissue and the droplet, the higher the rate of water
transfer and hence the cooler the droplet. If the standard solution has a lower water
potential than that of the sample to be measured, water will diffuse from the tissue to
the droplet, causing warming of the droplet. Measuring the change in temperature of
the droplet for several solutions of known Ψw makes it possible to calculate the water
potential of a solution for which the net movement of water between the droplet and
the tissue would be zero signifying that the droplet and the tissue have the same
water potential.
Psychrometers can be used to measure the water potentials of both excised and
intact plant tissue. Moreover, the method can be used to measure the Ψs of
solutions. This can be particularly useful with plant tissues.
A major difficulty with this approach is the extreme sensitivity of the

measurement to temperature fluctuations. For example, a change in temperature of
0.01°C corresponds to a change in water potential of about 0.1 MPa. Thus,
psychrometers must be operated under constant temperature conditions. For this
reason, the method is used primarily in laboratory settings.
7.5.3 Pressure chamber method
A relatively quick method for estimating the water potential of large pieces of
tissues, such as leaves and small shoots, is by use of the pressure chamber. This
method was popularized by P. Scholander and coworkers.The pressure bomb is a
device that is used to determine the plant moisture stress and the water potential of a
leafy shoot and in based on the assumption that the water column in a plant is
almost always under tension because of the pull exerted by the osmotic influences
(water potential) of the living cells of the leaves. If the tension is high, the water
potential of the leaf cells is very negative. When a stem is cut, the water column (in
xylem) is disrupted and because the water column is under tension, it will recede
back into the stem toward the leaves. The shoot is placed in a chamber, with the cut
end protruding through an airtight hole. Pressure is increased within the chamber
and the water column with in the twig are forced back to the cut surface. The
pressure in the chamber is then carefully recorded.
42

The pressure required to force the water to appear at the cut surface is equal
to the tension (but with the opposite sign) of the water column at the time the shoot
was cut. If low pressure is sufficient to force water to the cut surface of the shoot, the
shoot is under relatively low moisture stress. But if high pressure is required to force
to the cut surface the moisture stress (tension) is relatively high due to very negative
water potential of the leaf cells.
7.5.2 The second way to describe water status is to measure the quantity of water
in a tissue i.e. its water content and to express it in relation to a selected references.
Three of these methods are
-fresh weight method

- dry weight method and
- relative water content (RWC) method.
*******
43

Lecture-8
CROP WATER RELATIONS
8.1 TRANSPIRATION:
The loss of water from aerial parts of plants in the form of vapor is known as
transpiration. The loose arrangement of the living thin walled mesophyll cells, which
results in an abundance of inter cellular space provides an ideal condition for the
evaporation of water from internal leaf surface. Part of the epidermal surface of the
leaf is made up of a great number of microscopic pores called stomata. Water vapor
collected in the intercellular spaces of leaf mesophyll diffuse into the atmosphere
through the open stomata. This form of transpiration is termed as stomatal
transpiration.
In addition to stomatal transpiration, water is lost as vapor directly from leaf
surfaces and through lenticels (small opening in the corky tissue covering stems and
twigs). The former is called cuticular transpiration and the later lenticular
transpiration.
Stomatal Transpiration
The stomatal transpiration accounts to 80 to 98% of the total transpiration loss
from plants (tree to herbaceous plants). Under very dry conditions, stomata are
closed and water loss occurs through the cuticle and lenticels.
Cuticular Transpiration
The cuticular transpiration accounts to 2 to 20% of the total transpiration loss
from plant (xerophytes to mesophytes). The cuticle although retards water loss, is
some what permeable to water vapor. In plants with thick cuticles, this form of
transpiration is insignificant.
Lenticular Transpiration
The lenticular transpiration amount to 0.1 to 1% of the total transpiration loss
from plants which is insignificant when compared to stomatal transpiration. However,
lenticular transpiration may cause some desiccation in those trees that shed their
leaves at the on set of the winter. During cold winter water absorption by roots is at a
minimum, thus the importance of lenticular transpiration is increased.
8.2 SIGNIFICANCE OF TRANSPIRATION
Transpiration is advantageous because.
 It creates suction force and help in the ascent of sap.
 It helps in the absorption of water and minerals by roots.
 It helps in evaporating excess amount of water from moist soil.
 It plays a role in translocation of food from one part of the plant to the
other.
 It brings opening and closure of stomata which indirectly influences the
gaseous exchange for the processes of photosynthesis and respiration.
44

 It helps in dissipating the excess energy absorbed from the sun, which will
otherwise raise the leaf temperature.
 It maintains suitable temperature of leaves by imparting a cooling effect.
Transpiration is regarded as an unavoidable (necessary) evil. It is unavoidable

because leaf structure (stomata) favorable for uptake of Co 2 and O2 necessary for
photosynthesis and respiration is also favorable for the loss of water through
transpiration.
Transpiration is an ‘evil’ because often it causes injury by dehydration due to
heavy transpiration loss when the atmospheric conditions are aggressive such as
high light intensity, hot winds, depleted soil moisture and poor water retentive
capacity of soil.
8.4 STRUCTURE OF STOMATAL COMPLEX IN MONOCOT AND DICOT

SPECIES
The appearance of the guard cells differs characteristically from the surrounding
epidermal cells. The guard cells of some plant species, particularly grasses
(monocots), are dumbbell shaped and are associated with epidermal cells that also
differ in appearance from the rest of the epidermal cells. These epidermal cells are
called subsidiary cells or accessory cells. In case of dicots, the guard cells are
generally bean shaped. (Fig.5)
One other distinguishing feature of guard cell is the presence of chloroplasts.
Epidermal cells do not possess chloroplasts.
8.4.1 INVOLVEMENT OF STOMATA IN TRANSPIRATION :

The stomatal movement is generally understood as a direct response to
increase or decrease in the osmotic potential that result from osmotic changes that
cause water to move in or out of the guard cells. If water moves in, the cells expand
(become turgid); if water moves out, they go flaccid. When the guard cells are
turgid, the stoma is open; when the guard cells are flaccid, the stoma is closed. To
effect this movement of water, an exchange must takes place between the guard
45

cells and the surrounding mesophyll and epidermal cells. The development of a
more negative osmotic potential in the guard cells would cause of water potential
gradient to develop between the guard cells and their neighboring cells. Water would
diffuse in to the guard cell, causing them to become more turgid. The development of
a less negative osmotic potential in the guard cells would of course, cause a water
potential gradient to develop in the opposite direction, and water would flow out of
the guard cells into the neighboring cells. (see the diagram in the annexure)
8.5 TRANSPIRATION IN RELATION TO PRODUCTIVITY:
The importance of water use efficiency (WUE) in influencing grain yield under
water limited conditions can be explained by the following model give by
passioura.
Grain Yield = T x TE x HI
Where T = Total transpiration by the crop canopy
TE = Transpiration Efficiency or WUE
HI = Harvest Index (Economic Fraction of Dry matter)
This relationship provides an analytical tool to select the genotype with high levels of
T and TE.
8.6 WATER USE EFFICIENCY:

The water use efficiency (WUE) of field crops is defined as follows:
‘It is the amount of dry matter produced per unit amount of water transpired’
Dry matter production (DM)
WUE = ---------------------------------------------
Evapotranspiration
This is expressed as g DM kg-1 water WUE measurements can be made on
plants in containers, on individual plants, and on crop communities. They can be
used for economic yield as well as total dry matter calculations.
A related term, water requirement is the reciprocal of WUE. Water requirements
is usually expressed in weights of equal magnitude, such as g water (g DM)-1
8.7 Water use efficiency in C3 & C4 plants.

Field data for WUE, when regrouped into C3 and C4 species, illustrate a two fold
increase for C4 species when calculated for either grasses or dicots
Water use Efficiency (g DM (kgH2O)-1) for C4 and C3 species.
Species Grasses Dicots
C3 1.49 1.59
C4 3.14 3.44
Large differences in WUE occur when species are categorized by Co2 fixation
pathway. It is now accepted that the WUE of C4 species is generally higher than C3
species. The higher WUE of C4 species is a result of higher photosynthetic rate
under high light and temperature and lower transpiration rates under low light.
46

The WUE values for both C3 and C4 species are low compared with CAM
plants. One CAM species, pineapple (Ananas comosis), has shown a WUE of
20g, DM kg-1 water. Use of crop species with CAM is limited because the Co2
fixation and overall productivity of CAM plants is low (CAM is only a survival
mechanism but not a productive mechanism).
8.8 WUE of major field crops :

Crop Co2 Fixation pathway water requirement water Use Efficiency
(gH2O, gDM-1) (g DM kg-1 H2O)
Maize C4 388 2.58
Sorghum C4 402 2.49
Potato C3 532 1.88
Sugar beet C3 606 1.65
Wheat C3 613 1.63
Soybean C3 704 1.42
Alfalfa C3 993 1.01
8.9 FACTORS INFLUENCING THE WATER USE EFFICIENCY:
1. Climatic factors
WUE has almost an inverse relationship with Relative humidity (R.H), lower
R.H. increases evapotranspiration (ET) without a corresponding increase in crop
yield. On the other hand, factors such as sunlight and temperature that affect ET,
rate of photosynthesis and dry matter production will either decrease or increase
WUE.
2. Agronomic practices and crop management

Early sown crops will escape the moisture stress while the delayed sowing
favors heavy weed growth which creates severe competition for water, light, nutrients
etc. the crop should have an optimum crop canopy (4-6 LAI for most crops) for
proper light interception. Plant Population and plant architecture influence WUE by
influencing the interception and utilization of solar energy. Depth of sowing also
influences water availability and there by seedling emergence, vigor and final yield.
3. Antitranspirants
WUE can also be improved by using antitranspirants which reduces
transpiration. These antitranspirant may influence stomatal closure (Phenyl mercuric
acetate, ABA, CCC Salicylic acid etc) or form a film (Hexadecanol, cetyl alcohol etc.)
on the leaf surface or increase plant reflectivity (eg. Kaolinite) and reduce leaf
temperatures.
47

4. Use of mulches
Mulches are beneficial in conserving or economizing water use by plant
ranging from 10-50%. It depends upon the crop in which it is used, rainfall, wind
velocity and temperature of both air and soil. Organic mulches (straw, rice dust, saw
dust etc.) light colored and light reflecting mulches reduce soil temperature. But black
colored mulches such as black, grey, transparent polythene sheets and petroleum
products increase temperature up to 5-80C. Plastic mulches are costly and suitable
to areas where soil temperatures are low and unfavorable to crop growth.
5. Use of shelter belts

Shelter belts decrease the damaging effects of winds on crops and modify the
micro climate. Higher humidity and lower vapor pressure deficit prevail in shelter belt
that reduces heat. This will help ultimately for higher WUE. Increased yields due to
shelter belts are reported in case of cotton, onion, sweet potato, tomato and wheat.
6. Method and quantities of water application.

Frequent light irrigations keep the soil surface wet for a longer period but
consequently greater loss due to evaporation. Heavy application of water causes
heavy deep percolation losses.
Selection of proper method of irrigation is important based on soil and crop
characteristics. WUE is generally higher with sprinkler irrigation than surface method
of irrigation. Drip irrigation also increases production and decrease the water use by
a crop.
Among surface irrigation methods, WUE of crops increases in the order of
wild flooding, border strip, check basin, basin and furrow irrigation.
7. Fertilizer application
WUE of crops invariably increases with the application of fertilizers on
deficient soils under adequate soil moisture conditions. This is particularly with high
yielding varieties and hybrids.
8. Weed control
Weeds due to their early establishment and a better root system are able to
exhaust soil moisture more effectively than crop plants. Therefore, both yield and
WUE are reduced. Controlling weeds is essential for higher WUE of crops.
*******
48

ANNEXURE
Movement of water towards more negative water potential
49

Lecture-9
PHOTOSYNTHESIS
9.1 DEFINITION: Photosynthesis is the process by which organisms convert light

energy into chemical energy in the form of reducing power as NADPH and ATP. This
reducing power is used to fix carbon dioxide as carbohydrates (sugars). In oxygenic
photosynthetic organisms, including higher plants, the source of reducing equivalents
is H2O, releasing O2 as a by product.
Thus, the overall reaction of oxygenic photosynthesis can be represented as.

LIGHT
CHLOROPHYLL
This equation is frequently represented by the simplified form:
CO2 +2 H2O (CH2O) + H2O + O2
Reactions of Photosynthesis:
During the normal functioning of the photosynthetic system, light serves to

reduce nicotinamide-adenine dinucleotide phosphate (NADP). This in turn serves as
a reducing agent for carbon fixation in the carbon reduction cycle. ATP is also
formed during the electron flow from water to NADP, and it too used in carbon
reduction. The chemical reactions in which water is oxidized to oxygen, NADP is
reduced, and ATP is formed are generally known as the light reactions while the
carbon reduction reactions are called the dark reactions.
All most all reactions up to NADP reduction takes place with in the thylakoids, while
the carbon reduction rections take place in the aqueous region of chloroplast, the
stroma. The following figure explains about the link between light reaction and dark
reaction.
50

9.2 ENERGY SYNTHESIS:
Energy synthesis is the prime function of photosynthesis. It involves the role of

various aspects like chloroplast and its structure, pigments present in chloroplast,
Electro magnetic spectrum, photosynthetically active radiation, light interception by
canopy, Capturing of light by pigment molecules, Light reaction (photolysis of water,
electron transport between photo systems, generation of reducing power) and
utilization of this reducing power in fixation of CO2 to carbohydrates (dark reaction).
Each of this aspect is briefly explained here.
9.2.1 CHLOROPLAST AND ITS STRUCTURE:
In photosynthetic eukaryotes, photosynthesis takes place in the sub cellular

organelle known as chloroplast.(Fig.6)
The most striking aspect of the structure of the chloroplast is this extensive
system of the internal membranes known as thylakoids, (On these thylakoid bodies
some round shaped photosynthetic units called quantasomes are presnt.) All the
chlorophyll is contained with in this membrane system, which is the site of the light
reactions of photosynthesis. The carbon reduction reactions or dark reactions
which are catalyzed by water soluble enzymes, takes place in the stroma, the region
of the chloroplast outside the thylakoids. Stroma forms the matrix of the chloroplast.
In this portion of chloroplast, lamellae are loosely arranged. The lamellae which are
found in this region are called stroma lamellae.
9.2.2 PHOTOSYNTHETIC PIGMENTS:
There are four different pigments in higher plants- two chlorophylls (Chl. A and
Chl. B) and two carotenoids (carotene and xanthophyII). The chlorophylls are green
in color whereas carotene and xanthophylls are orange and yellow, respectively.
Magnesium present in chlorophyll molecule is crucial to the capture of light energy.
51

All organisms actually contain a mixture of more than one kind of pigment, each
serving a specific function. As the energy of light absorbed by carotenoids is rapidly
transferred to chlorophylls, carotenoids are termed as accessory pigments.
9.2.3 ELECTRO MAGNETIC SPECTRUM:
In the electromagnetic spectrum the wavelengths between 400-700nm is

called as PHOTOSYNTHETICALLY ACTIVE RADIATION (PAR).
9.2.4 PRINCIPLE OF LIGHT ABSORPTION BY PLANTS:
The depression of light intensity within the plant community is mainly due to
interception of light by leaves. The absorption of radiant energy by a plant
community would follow the Lambert-Beer’s Law.
Beer’s Law states that the absorption of light by a solution is proportional to

the concentration of the solution and the distance through which the light travels
Here the absorbance is defined as A = log I0 / I where I0 is the intensity of light that is
incident on the sample and I is the intensity of light that is transmitted by the sample.
52

Lambert expressed this relationship mathematically as
I = I0e-KX
Where I0 = Incident light intensity
I = Intensity of the light after passing through the solution
K = Absorption coefficient.
X = Distance or path length .
Davidson and Philip modified this equation for absorption of energy by a plant
canopy. They substituted LAI for x (i.e the path length). The hypothesized equation
for absorption of radiant energy by a plant canopy then becomes
I = I0 e –KA
(or) K = log e (Io/I) / A
The proportion of incident light that is intercepted by the canopy does not
depend on leaf area index alone, but also on the architecture of the canopy. In a crop
the leaves are not isolated, but are arranged in a canopy. The way they are arranged
will affect the proportion of incident light that is intercepted by the crop canopy as a
whole. Thus, the extinction coefficient (K) is a measure of the light intercepting
efficiency of the leaf area. Leaf characters of importance in this respect are:
leafEangle, leaf area, continuity of leaf layers etc.
9.2.5 THE TWO PHOTO SYSTEMS:
9.2.5.1 Evidence for the existence of PSI and PSII. (Red drop & emerson
enhancement effect)
Experiments measuring the quantum yield (i.e. the number of O 2 molecules

released for each Quanta absorbed) is approximately 0.1. The reciprocal of
quantum yield is quantum requirement i.e. the number of quanta required for each
O2 molecule evolution. It is 10). If the quantum yield of photosynthesis is measured
at different ranges of wave lengths most of the ranges are remarkably constant.
However at the extreme red edge of the chlorophyll absorption (>680nm), the yield
drops drastically. The phenomenon is known as RED DROP.
Conclusion: It is proposed that at this wave length only one photosystem remains in
operation. As another photosystem is not able function at this wavelength quantum
yield drops.
53

Another puzzling experimental result was the enhancement effect, discovered

by Emerson. The rate of photosynthesis was measured separately with light of two
different wave lengths and then the two beams were used simultaneously. When
exposed to a wavelength more than 680 nm (far red region) a specific rate of
photosynthesis was observed. Likewise when the exposure was given at
wavelengths less than 680 nm some other effect was observed. When the system
was exposed to the light of both wavelengths simultaneously, the effect on
photosynthesis exceeded the sum of the two effects caused separately. This
provided evidence that the two pigment systems worked in co-operation with each
other and the increase in photosynthesis was due to synergism. This phenomenon
is known as Emerson Enhancement effect. This experiment also explained that,
there exist two photo systems (PS-I & PS-II)
Based on these two observations Hill and Fay Bendall (1960) proposed that
light reactions of photosynthesis involve two photochemical events.
9.2.5.2 PHOTOSYSTEM I & II:
The reaction centre chlorophyll of photo system I absorbs maximally at 700

nm in its reduced state. Accordingly, it is named P 700 (the P stands for Pigments),
the reaction centre chlorophyll of photo system II absorbs maximum at 680 nm (P
680 )so its reaction centre chlorophylls and associated electron transport proteins is
located predominantly in the stacked regions of the grana lamellae. Whereas the
photo system I, its associated antenna pigments, ATP synthase enzymes are found
exclusively in the stroma lamella and at the edges of the grana lamellae. (At a
wavelength grater than 680 nm PS-II can not operate. Therefore, quantum yield
decreased beyond 680 nm. This is what red drop is.)
Photo system I contains large amount of chlorophyll a, a small amount of

chlorophyll b and some B carotene. Photo system II also contains chlorophyll a, B
carotene but a large amount of chlorophyll b.
9.2.6 LIGHT REACTIONS:
Almost all the chemical processes that make up the light reactions of
photosynthesis are carried out by four protein complexes; photo system II (P 680),
the cytochrome b-f complexes, photo system I (P 700) and the ATP synthase.
SEQUENCE OF LIGHT REACTIONS:
9.2.6.1 Water is exidized to oxygen by Photo system II:
The chemical reaction by which water is oxidized is given by the following

equation.
2H2O O2 + 4H+ + 4e-
54

This equation indicates that four electrons are removed from two water
molecules, generating an oxygen molecule and four hydrogen ions.
Robert Hill demonstrated that isolated chloroplasts evolved Oxygen when

they were illuminated in the presence of suitable electron acceptor, such as
ferricyanide. The ferricyanide is reduced to ferrocyanide by photolysis of water .This
reaction is now called as HILL REACTION and it explains that water is used as a
source of electrons for CO2 fixation and Oxygen is evolved as a by product.
The protons produced by oxidation of water is released into the lumen of the
thylakoid, not directly into the stromal compartment. These protons are eventually
released from the lumen to the stroma through the process of ATP synthesis. In this
way the electro chemical potential formed by the release of protons during oxidation
of water contributes to ATP formation.
Water Oxidation System: Manganese (Mn) is an essential cofactor in the water

oxidizing process. Analytical experiments indicate that four Mn ions are associated
with oxygen evolving complex. Other experiments have shown that cl and Ca ions
are also essential for O2 evolution.
9.2.6.2 ELECTRON TRANSPORT AND PHOSPHORYLATION:
The Z scheme (named because of its shape) illustrates electron transport and
the production of NADPH and ATP in chloroplasts.
9.2.6.2.1 NON CYCLIC PHOTOPHOSPORYLATION:
Non cyclic photophosphorylation involves integration of two photo systems

(PS-I and PS-II). This is one of the means of ATP production in chloroplasts.
Non-Cyclic : Electrons follows a non cyclic track.
Photo : it is the light energy that drives electrons along
this Track.
Phosphorylation : As electrons are driven along the track ADP is
phosphorylated to yield ATP.
This can also be termed as non-cyclic electron transport to refer to the

manner of electron flow during the process.
MECHANISM:
The primary flow of electrons within a given granum thylakoid may be initiated
almost simultaneously for each photo system (PS I and PS II).
55

After excitation of P700 the electrons are passed on to a chlorophyll molecule

(Ao). The electrons are then passed to series of Iron-sulfur proteins (Fe-S) and
finally to ferridoxin (Fd). The ferridoxin-NADP reductase serves to reduce NADP to
NADPH which is used in the Calvin cycle to reduce Co2.(Fig. 7)
The transfer of electrons to NADP creates debit (commonly referred to as a

hole) in photo system I. However this deficit is made up by the excitation of P 680 of
photo system II. The excited P680 of PS II transfers electrons to pheophytin,
plastoquinones, and cyt b6-f complex. Cytochrome b6-f complex transfers electrons
to plastocyanin (PC), which in turn reduces P700* (excited P700). The hole created in
photo system II is filled by-electrons that are derived from the oxidation of water.
In addition to the energy stored as redox equivalents (NADPH) by the light

reactions a portion of the photons energy is utilized for the synthesis of ATP during
the transfer of electrons between plaotoquinone and cyt b6-f complex. This
phenomenon of synthesis of ATP in light reactions of photosynthesis is known as
photophosphorylation. It is now widely accepted that photophosphorylation works
via the chemi-osmotic mechanism first proposed in 1960 by Peter Mitchell.
56

The basic principle of chemi-osmosis is that in concentration differences and

electrical potential differences across the membranes are a source of free energy
that can be utilized by the cell for the synthesis of ATP. In the light reactions electron
flow is coupled to proton translocation, creating transmembrane proton motive force
(pmf). The energy in the proton motive force is then used for synthesis of ATP by the
enzyme called ATP synthase.
This synthesis of ATP coupled to a linear flow of electrons from water to

NADP is called as non cyclic photophosphorylation. The precise number of ATP
molecules formed in non-cyclic photophorylation is unclear although it is generally
thought two per O2 evolved.
9.2.6.2.2 CYCLIC PHOTOPHOSPHORYLATION:
The cyclic photophosphorylation operates when chloroplasts are illuminated

with wave lengths of light greater than 680nm. Under these circumstances only
photo system I is activated and electrons are not removed from H 2O. When the flow
of electrons from H2O is stopped, non cyclic assimilation retarded, oxidized NADP is
no longer available as an electron acceptor. Activation of photo system I by wave
lengths of light greater than 680 nm causes electron to flow from P 700 to chlorophyll
molecule and Ferridoxin. Then the electrons instead of pass on to NADP return back
to P700 via cyt b6-f complex, plastoquinone and plastocyanin.
Cyclic transport system is likely to result in the synthesis of ATP at two

locations. One is between Fe-s protein and cyt-b6 complex and another between
cyt-b6 and cytochrome f.
Significance
Evidence for the operation of cyclic electron transport in C3 plants in vivo is

limited but it has been demonstrated under physiological conditions in vivo in C4
plants where there is an additional ATP requirement in their carbon fixation pathway.
It may also play an important role in the synthesis of ATP required for protein
synthesis during PS II repair following photo inhibition.
57

9.2.6.2.3 PSEUDO CYCLIC PHOSPHORYLATION:
Another source of generation of ATP is that electrons might be transferred

from ferridoxin back to oxygen reducing it to water. It is possible that this process
might also involve an electron transport chain and produce ATP. Here the electron
that is cycled back to reduce molecular oxygen to water is not the same that is
released from the water. Hence it is called as pseudo cyclic phosphorylation.
9.2.7 CARBON DIOXIDE FIXATION /DARK REACTIONS OF PHOTOSYNTHESIS:
During the light reactions of photosynthesis, the photochemical oxidation of

water to molecular oxygen is coupled to the generation of reduced pyridine
nucleotide (NADPH) and ATP. The reactions associated with the reduction of CO 2 to
carbohydrate are coupled to the consumption of NADPH and ATP. These reactions
are referred to as the Dark reactions of the photosynthesis.
9.2.7.1 The C3 Cycle (C3 Photosynthetic Carbon Reduction Cycle)
The PCR cycle is sometimes referred to as the Calvin cycle in honor of its
discoverer, the American biochemist Melvin Calvin, other pathways associated with
the photosynthetic fixation of CO2, such as the C4 photosynthetic carbon assimilation
(PCA) cycle and the C2 photo respiratory carbon oxidation cycle (PCO), are either
auxiliary to or dependent on the basis PCR cycle.
In the C3, PCR cycle, carbon dioxide from the atmosphere and water are
enzymatically combined with a five-carbon acceptor molecule to generate two
molecules of a three carbon intermediate. These intermediates are reduced to
carbohydrate using the photo chemically generated ATP and NADPH in the light
reactions. The cycle is completed by the generation of five-carbon acceptor.(Fig.8)
The C3 PCR cycle proceeds in three stages:
1. Carboxylation of the CO2 acceptor, ribulose 1,5 – bisphosphate, to form

2 molecules of 3. Phosphoglycerate, the first stable intermediate of the
PCR cycle.
2. Reduction of this carboxylic acid to a carbohydrate in the form
glyceraldehydes 3- phosphate.
3. Regeneration of CO2 acceptor, ribulose 1.5-biophosphate from
glyceraldehydes 3-phosphate.
The Carboxylation of Ribulose Bisphosphate:
CO2 enters the PCR cycle by reacting with ribulose 1,5-bisphosphate to yield
two molecules of 3-phosphoglycerate, a reaction that is catalyzed by the chloroplast
enzyme ribulose bisphosphate carboxylase/oxygenase, referred to by the acronym
RUBISCO.
58

The reduction step of the C3 PCR cycle:
In this stage, the 3-phosphoglycerate formed as a result of the carboxylation

of RuBP (Ribulose 1,5-bisphosphate) is first phosphorylated to 1,3-bis phospho
glycerate by the ATP generated in the light reactions and is then reduced to
glyceraldehyde 3-phosphate, using the NADPH generated by the light reactions.
The chloroplast enzyme NADP-glyceraldehyde 3-phosphate dehydrogenase
catalyzes this step.
Regeneration of Ribulose 1, 5 -Bisphosphate:
One molecule of glyceraldehyde 3-phosphate is converted to dihydroxy

acetone 3-phosphate (DHAP). The DHAP then undergoes aldol condensation with a
molecule of glyceraldehydes 3-phosphate to form fructose 1,6-bis phosphate. This
product is hydrolyzed to fructose 6 phosphate. This fructose 6 phosphate combines
with third molecule of glyceraldehyde 3-phosphate to give erythrose 4-phosphate
and xylulose 5-phosphate. This reaction is catalyzed by transketolase.
Erythrose 4-phosphate then combines with DHAP to yield a seven

carbon sugar, sedoheptulose 1,7-bisphosphate which is further hydrolyzed to give
sedoheptulose 7-phosphate. Sedoheptulose 7-phosphate donates a two carbon unit
to the fifth molecule of glyceraldehyde 3-phosphate and produce ribose 5-phosphate
59

and xylulose 5-phosphate as its products. Two molecules of xylulose 5-phosphate

are epimerized to give ribulose 5-phosphate. The third molecule of ribulose 5-
phosphate is formed by the isomerization of ribose 5-phosphate. Finally, ribulose 5-
phosphate is phosphorylated with ATP. Thus generating CO2 acceptor ribulose 1,5-
bisphosphate.
Energy requirement: In order to synthesize the equivalent of 1 molecule of hexose

sugar, 6 molecules of CO2 are fixed at the expense of 18 ATP and 12 NADPH. In
other words, the PCR cycle consumes 2 molecules of NADPH and 3 molecules of
ATP for every CO2 fixed.
6CO2+11H2O+12 NADPH + 18ATP Fructose- 6- P +12 NADP+6 H+ + 18 ADP + 17 Pi
*******
ANNEXURE
Apart from stroma lamellae ribosomes serving as sites for the protein synthesis are also
found scattered in this region. Lipid, protein and nucleic acid metabolisms are also found in
this region.
Chlorophyll molecule is a typical porphyrin derivative possessing a cyclic tetrapyrollic

structure in which one pyrrole ring is partially reduced. The tetrapyrorolic nucleus contains a
non-ionic magnesium atom held by two covalent and two coordinate bonds in the centre of
the molecule. Magnesium is crucial to the capture of light energy. The presence of methyl
group (CH3) at position 3 in the second pyrrole ring makes chlorophyll-a and when it is
replaced by formyl group (-CHO) it forms chlorophyll-b.
Chemical formula Absorption peaks

1 Cholorophyll-a C55H72O5N4Mg 662, 430nm
2 Cholorophyll-b C55H70O6N4Mg 642, 453nm
An action spectrum is measured by

plotting a response to light such as oxygen
evolution, as a function of wavelength. If
the pigments used to obtain the absorption
spectrum are the same as those that cause
the response, the absorption and action
spectra will match. In the example shown
here, the action spectrum for oxygen
evolution matches the absorption spectrum
of intact chloroplasts quite well, indicating
that light absorption by the chlorophylls
mediates oxygen evolution.
*For further information see reference.6
60

Lecture-10
PHOTOSYNTHESIS
10.1 THE C4 PHOTOSYNTHETIC CARBON ASSIMILATION (PCA) CYCLE
Hal Hatch and Roger Slack elucidated what is now known as the C 4 PCA
cycle. Using sugarcane leaves, they established that the C4 acids malic acid and
aspartic acid were the first stable detectable intermediates of photosynthesis in
leaves of C4 plants. The primary carboxylation in these leaves was not catalyzed by
Rubisco but by PEP carboxylase.
The basic C4 PCA cycle consists of four stages.
1. Assimilation of CO2 involving carboxylation of phosphoenol pyruvate (PEP) in

the mesophyll cells to form C4 acids (Malate and / or aspartate)
2. Transport of C4 acids to the bundle sheath cells.
3. Decarboxylation of C4 acids within in the bundle sheath cells and generation
of CO2 which is reduced to carbohydrate via the C3 PCR cycle.
4. Transport of the C3 acid formed by the decarboxylation (pyruvate or alanine)
back to the mesophyll cell and regeneration of the CO2 acceptor,
phosphoenol pyruvate.
Anatomical differences
A cross section of a typical C3 leaf reveals essentially one type of

photosynthetic, chloroplast containing cell, the mesophyll. In contrast, a typical
C4 leaf has two distinct chloroplast containing cell types, the mesophyll and the
bundle sheath cells. The extensive network of Plasmodesmata connects
mesophyll and bundle sheath cells, providing a pathway for the flow of
metabolites between cells. This is called as KRANZ ANATOMY. (Fig.9)
A B
61

The figure ‘A’ shows that in a C4 leaf both mesophyll and bundle sheath cells
possess chloroplasts.
Discovered in tropical grasses (e.g., sugarcane and maize), the C4 cycle is

now known to occur in 16 families of both monocortyledons and dicotyledons,
and it is particularly prominent in Gramineae (sugarcane, com, sorghum),
Chenopodiaceae (Atriplex), and Cyperaceae (sedges). About 1% of the
characterized species have C4 metabolism.
The primary carboxylatiion reaction, catalyzed by PEP carboxylase, which is

common to all three variants, occurs in the cytosol of the mesoophyll chloroplasts
by NADPH using NADP malate dehydrogenase. The malate formed enters the
chloroplast of the bundle sheath cell and there undergoes oxidative
decarboxylation, yielding pyruvate. The CO2 released within the bundle sheath
cells is converted to carbohydrate by the Calvin cycle. ( Fig.10)
The residual C3 acid is transported back to the mesophyll as pyruvate and

converted to phosphoenolpyruvate in the mesophyll chloroplast.
62

10.1.1 There are three variants in PCR cycle of C4 plants:
Variant Principal C4 acid Decarboxylating Examples.

transported to the enzyme
Name
bundle sheath cells
NADP – ME Malate NADP-dependent Malic Maize, Sugarcane,

enzyme Sorghum
NAD-ME Aspartate NAD-dependent Malic Millet, Pigweed,

enzyme Panicum milliaceum
(variga) Amaranthus
PEP-CK Aspartate Phosphoenol pyruvate Guinea grass

carboxy kinase (Panicum maximum),
chloris gayana
These variants differ principally in the C4 acid transported into the bundle sheath
cells and decarboxylating enzyme.(Fig.11)
63

A) NADP-ME Type
The primary carboxylation reaction occurs in the cytosol of mesophyll cells

and is catalyzed by phosphoenol pyruvate carboxylase (PEP carboxylase) using
HCO3– rather than CO2 as a substrate.
Oxalo acetate formed in this reaction rapidly reduced to malate in the

mesophyll chloroplasts by NADPH. The C4 acid (Malate) then transported to the
chloroplasts of bundle sheath cells and undergoes oxidative decarboxylation by
NADP-ME. The CO2 released within the bundle sheath cells is reduced to
carbohydrate by the PCR cycle. The C3 acid (Pyruvate) formed in the
decarboxylation is transported back to the mesophyll. In the final step of PCA cycle,
the pyruvate is converted to phosphoenol pyruvate with in the mesophyll
chloroplast.
B) NAD-ME Type :The primary carboxylation reaction occurs in the cytosol of the
mesophyll cells and is catalyzed by PEP carboxylase using HCO3-.
Oxalo acetate formed undergoes transamination in the cytosol, with glutamate

as the amino donor and forms into Aspartate. The C4 acid (Aspartate) then
transported to the bundle sheath cells. In the bundle sheath cells, the asparatate is
first reconverted into oxalo acetate by transamination in the mitochondria. The
oxalo acetate in the mitochondria is then reduced and decarboxylated by the NAD-
Malic enzyme. The CO2 released within the bundles sheath cells is reduced to
carbohydrate by PCR cycle. The C3 acid pyruvate undergoes transamination and
formed in to alanine which will be transported back to mesophyll cells.In the
measophyll cytosol alanine appears to be converted to pyruvate via transamination.
In the final step the pyruvate is converted to phosphoenol pyruvate within the
mesophyll chloroplast.
C)PEP-CK type
The primary carboxylation reaction occurs in cytosol of mesophyll cells and is

catalyzed by PEP carboxylase using HCO3-. The Oxalo acetate undergoes
transamination and formed into aspartate a C4 acid. The C4 acid then transported
to the bundle sheath cell and is reconverted into oxalo acetate. The oxalo acetate
is decarboxylated by PEP carboxy knase and forms into phosphoenol pyruvate.
The PEP then transported to mesophyll cells.
64

10.1.2 Significance of C4 Photosynthesis
C4 PCA cycle effectively shuttles CO2 from the atmosphere into the bundle
sheath cells. This transport process generates a much higher concentration of CO 2
in the bundle sheath cells than would occur in equilibrium with the external
atmosphere. This elevated concentration of CO2 at the site of carboxylation of
ribulose 1,5 biphosphate by RUBISCO results in the suppression of ribulose
biphosphate oxygenation or Photorespiration. This is called as CO2 concentration
mechanism.
10.1.3 Energy Requirement
One interesting feature of the cycle is that the enzyme that catalyzes the
regeneration of phosho enol pyruvate by the enzyme pyruvate-orthophosphate
dikinase, consumes two “high-energy” phosphate bonds through the conversion of
ATP to AMP. There by, explaining the high energy requirement of C4 cycle.
Total energy requirement for fixing one molecule of CO2 in C4 plants is 5 ATP
plus 2 NADPH. This includes the cost of concentrating CO2 with in the bundle
sheath cell i.e. 2ATP per CO2
10.2 CRASSULACEAN ACID METABOLISM (CAM)
CAM is an acronym for crassulacean acid metabolism, but the mechanism is

not restricted to the family crassulaceae alone, like the C4 PCA cycle, it is found in
many angiosperm families.
The CAM mechanism is similar in many respects to the C4 PCA cycle, but
differs from it in two important features.
1) In C4 plants the formation of C4 acids is spatially, but not temporally, separated

from the decarboxylation of the C4 acids and re fixation of the resulting CO2 by the
PCR cycle. In CAM plants, the formation of C4 acids is temporally but not spatially,
separated from decarboxylation and refixation. CAM plants lack the specialized leaf
anatomy kranz leaf anatomy typical of C4 plants.
2) CAM plants open their stomata during the cool, desert nights and closed during
the hot, dry days. This minimizes water loss. The CO2 is assimilated at night.
(Fig.12)
65

The CO2 assimilation is accomplished by carboxylation of phosphoenol

pyruvate to oxalo acetate, which is then reduced to malate. The PEP originates from
the breakdown of starch and other sugars by the glycolytic pathway. The C 4 acid
accumulates as malic acid in large vacuoles. The accumulation of substantial
amounts of malic acid has long been recognized as dark acidification of leaf.
With the onset of day, the stomata close, preventing loss of water and further
acquisition of CO2. The leaf cells become deacidified as the reserves of vacuolar
malic acid are consumed. Decarboxylation is achieved by the action of NADP malic
enzyme on malate. Because the stomata are closed, the internally released CO 2
cannot escape from the leaf and instead is reduced to carbohydrate by operation of
the C3 PCR cycle. The C3 acid remaining after decarboxylation is thought to be
converted into starch or sucrose thus recovering the original starting material.
10.2.1 Significance:
The CAM mechanism enables plants to maximize their water – use efficiency
due to skoto active opening of stomata. Typically a CAM plant loses 50-100 g of
water for every gram of CO2 gained, compared with the values of 250-300 and 400-
500g for C4 and C3 plants respectively. Thus CAM plants are specially adapted to
arid environment. Like in C4 plants, the elevated internal concentration of CO2
effectively suppresses the photo respiratory oxygenation of ribulose biphosphate and
favors carboxylation.
66

10.3 MEASUREMENT OF PHOTOSYNTHESIS:
There are different methods by which photosynthesis can be measured.
A. Assay of Chloroplast activity (Hill reaction)
Either intact or isolated chloroplasts of leaf fragments evolve oxygen and

electrons during photosynthesis. Photosynthesis hence is measured by the changes
in the absorption spectra of 2,6-di chloro phenol indo phenol (DCPIP) which is
reduced by the evolved electrons. It represents chloroplast activity. This method is
convenient to identify photosynthetic inhibitors.
B. Gas exchange Measurement (Infra – red Gas Analysis IRGA)
Photosynthetic gas exchange refers to the fluxes or flows of CO2 between the
plant and environment.
Infrared gas analysis is the most popular, accurate and simple technique
determining photosynthetic or respiratory CO2 exchange technique in air. It is based
on the principle that the hetero atomic gas molecules typically absorb radiation at
specific infrared wave bands and that each gas has got a characteristic absorption
spectrum and not interfered by gas molecules of identical atoms. Hence IRGA can
be used for accurate determination of concentrations of CO2 uptake by small leaves.
C. Monitoring with 14CO2 : Photosynthesis true to the field, is measured by
14
a) Exposing Photosynthetic tissue to CO2
14
In this photosynthetic tissue is exposed to a gas of CO2. By killing the
14
tissue after a definite time CO2 fixed is determined and the rate of Photosynthesis
is estimated.
b) Exposing leaf to a gas mixture in a closed space:
In this method the leaf is exposed in a closed space to a gas mixture containing
the carbon isotopes and the rate of decrease of radioactivity of the gas is measured
by a built in β- counter.
*******
67

Lecture-11
PHOTOSYNTHESIS
11.1 THE C2 PHOTO RESPIRATORY CARBON OXIDATION (PCO) CYCLE
All land plants have an additional metabolic activity and loose a considerable
amounts of their photosynthates as CO2 with in a few seconds of its being fixed.It
occurs when plants are illuminated. This process of CO2 release is light dependent
and because of its otherwise superficial resemblance to respiration, it has been given
the name Photorespiration. During the process O2 is consumed and CO2 is
generated in light. It is independent of mitochondrial respiration but occurs in
chloroplasts and peroxisomes and only a few reactions occur in mitochondria.
However, unlike true respiration it performs no useful function in making energy
available to the plant. Instead its occurrence leads to loss of energy and this is
responsible for a considerable reduction in the yield of many of the crops.
Definition: Photorespiration means evolution of CO2 by green leaves in light.
11.1.1 Mechanism:
RuBISCO is a by bifunctional enzyme capable of carboxylation as well as

oxygenation of RuBP. The operation of the PCO cycle involves cooperative
interactions between three organelles, Chloroplasts, mitochiondria and
peroxisomes.(Fig.13)
Figure.13
68

The oxygenation of RuBP results in the formation of 2-phosphoglycolate and

3-phosphoglycerate.The phospho glycolate is rapidly hydrolyzed to glycolate by a
specific enzyme phosphatase. Glycolate leaves the chloroplast and diffuses to
peroxisome. There it is oxidized by glycolate oxidase to glyoxylate and hydrogen
peroxide. The peroxide is destroyed by the action of catalase and glyoxylate
undergoes transamination and the product is the amino acid glycine.
Glycine leaves the peroxisomes and enters mitochondria, where two

molecules of glycine are converted to serine. Glycine is thus, the immediate
source of the photo respiratory CO2. Serine leaves the mitochondrion and enters
peroxisome, where it is converted first by transamination to hydroxy pyruvate and
then by reduction to glycerate.
Finally, glycerate reenters the chloroplast, where it is phosphorylated to yield

3-phosphoglycerate.
11.1.2 Significance:
Photorespiration is a response to the low CO2/O2 ratios prevalent in the

present day atmosphere and has no functional role. Another possible explanation is
that photorespiration is necessary under conditions of high light intensities and low
CO2 concentration ( e.g. when stomata are closed because of water stress ) to
dissipate excess ATP and reducing power from the light reactions, thus preventing
damage to the photosynthetic apparatus. Today, there is no conclusive evidence for
the role of photorespiration in the carbon metabolism of the leaf.
Many plants do not photo respire, this is not because that RUBISCO have
different properties, rather, it is a consequence of interesting mechanisms that
concentrate CO2 in the RUBISCO environment is in the case of C4 and CAM plants.
(CO2 concentration mechanism)
11.1.3 Energy requirement
Oxygenation of ribulose bis phosphate and operation of PCO cycle consumes

2ATP and 2.5 NADH for each ribulose bisphosphate oxygenated.
11.2 FACTORS AFFECTING PHOTOSYNTHESIS
Many plant and environmental factors influence photosynthesis such as light,

CO2, temperature, availability of water as well as plant age and genetics.
11.2.1 LIGHT
The liner portion of light response curves of photosynthesis represent the part of the
process in which photosynthesis is strictly light limited. Under this condition, each
increment in light elicits a proportional increase in photosynthetic rate, resulting in a
69

linear relationship between photosynthetic rates and photon flux densities. At higher
photon flux densities, the photosynthetic response to light starts to level off and
reaches saturation. Once saturation is reached, further increases in photon flux
densities no longer affect photosynthetic rates, indicating other factors such as
RUBISCO activity or the metabolism of triose phosphates, have become limiting.
In the Dark, CO2 fixation is negative and there is net CO2 evolution from the
leaf because of respiration. As the photo flux density (availability of light energy)
increases net CO2 evolution decreases, after it reaches Zero, the plants shifts to net
CO2 fixation. The Photon flux density at which net CO2 exchange in the leaf is zero
is called light compensation point. The light compensation points of sun plants are
high in the range of 10-20 u mol m-2 S-1 where as corresponding values for shade
plants are low in the range of 1-5 u mol m-2S-1. The values for shade plants are
lower because respiration rates in shade plants are very low, so little photosynthesis
suffices to bring the rates of CO2 evolution to zero.
LIGHT RESPONSE CURVE
11.2.2 CARBON DIOXIDE: Carbon dioxide is a trace gas in the atmosphere

constituting approximately 390ppm.
CO2 RESPONSE
CURVES OF C3 & C4
SPECIES
Classical net photosynthetic curve for C3 and C4 species.Dashed vertical lines at 350 and 700 µ lit/lit
70

mark the current CO2 level and the doubled concentration predicted to be reached some time late in
the next century.Arrows indicate incremental rise in net photosynthesis due to the CO2 doubling.
In plants with CO2 concentrating mechanisms which include C4 and

CAM plants, the CO2 concentrations at the carboxylation sites are often saturating.
So a C4 plant such as corn cannot increase its photosynthetic performance with the
increase in atmosphere CO2 concentrations.In the C3 Plants, on the other hand,
increasing Ci levels (Ci partial pressure of CO2 in the inter cellular spaces of leaf)
continue to stimulate photosynthesis over much broader range.
Also markedly different in C3 and C4 plants is the CO2 compensation point, the
CO2 concentration at which CO2 fixation by photosynthesis balances CO2 loss by
respiration and net CO2 is zero. Plants with C4 metabolism have a CO 2
compensation point of zero or nearly zero (CO2 compensation point of C3 plants is 35
– 50 ppm, while for C4 plants 0 – 10 ppm.)
In C3 plants additional CO2 decreases photorespiration by increasing the ratio

of CO2 to O2 ratio, which leads to faster net photosynthesis.
11.2.3 TEMPERATURE :
When photosynthetic rates are plotted as a function of temperature, the

curves have a characteristic bell shape. The ascending arm of the curve represents
a temperature dependent stimulation of photosynthesis up to an optimum, the
descending arm associated with deleterious effects.
When photosynthetic rates are measured in air at normal and at high CO 2

concentrations: at high CO2 there is ample supply of CO2 at the carboxylation sites
and the rate of photosynthesis is limited primarily by biochemical reactions. At
ambient (normal) CO2 concentrations photosynthesis is limited by the activity of
RUBISCO.
Temperature rise and CO2 fixation are conflicting processes. An increase in

carboxylation rate with temperature and a decrease in the affinity of the RUBISCO
for CO2 is observed as the temperature rise. These opposite effects slow down the
temperature response of Photosynthesis at ambient CO2 concentrations.
Respiration rates increase as a function of temperature and the interaction

between photorespiration and photosynthesis becomes apparent in temperature
responses. The quantum yield of photosynthetic carbon fixation in a C4 plant remains
constant with temperature, reflecting typical low rates of photorespiration. In the C3
plant, the quantum yield decreases sharply with temperature, reflecting stimulation of
photorespiration by temperature and an ensuring higher energy demand per net CO 2
fixed.
71

Optimal temperature is the point at which the capacities of various steps of

photosynthesis are optimally balanced. Although there are exceptions, C4 plants
generally have higher temperature optima than C3 plants and this difference is
controlled largely by lower rates of photorespiration in C4 plants.
TEMPERATURE
RESPONSE
CURVES
11.2.4 Water
When water becomes limiting, cell expansion is first retarded so that growth is
reduced. With only a little more water stress, stomata begin to close and CO 2 uptake
is restricted. Photosynthesis is then limited by water because of retarded leaf
expansion and because of restricted CO2 absorption.
11.2.5 Other factor:
Factors like weedicides, salinity and water logging also affect the rate of
photosynthesis. Majority of Herbicides -about half of the commercially important
compounds—act by interrupting photosynthetic electron flow.(Ex. Paraquat, diuron).
When the electron transport is blocked it virtually stops light reaction of
photosynthesis. When light reaction is stopped the dark reaction does not happen
and thus CO2 is not fixed as carbohydrate. Therefore the weed is killed by starvation.
*******
72

Lecture-12
PHOTOSYNTHESIS
12.1 PHOTOSYNTHETIC EFFEICIENCY
Photosynthetic efficiency is the amount of dry matter fixed by the crop in a unit
area per unit time .Photosynthetic efficiency (Eµ) of a crop can be estimated with
the following formula
Chemical energy captured by a crop

Eµ =
Solar energy received
Thus, photosynthetic efficiency can be understood as net gain of chemical

energy per total incident solar radiation in a unit area.
Considering supplies of total PAR (Photosynthetically active radiation) to

land area on a growing crop, overall biomass production efficiencies are always
much below 18 percent. How ever this 18% is only a theoretical estimation which
is calculated as follows. Assume that 12 moles of photons represent the
maximum number of photons needed to fix one mole of CO2 and that an average
photon in the PAR region (400-700nm) has a wavelength of 550nm. From
Planck’s equation (E= h‫ )ﬠ‬relating photon energy and wavelength we can
calculate that one mole of such photons has an energy of 217,000 J (51,900 cal).
Twelve moles of photons would therefore have an energy of 2.6 X 10 6 J. This is
the input energy. The output energy, one mole of fixed carbon in carbohydrate,
has energy of about 0.48 X 106 J. Efficiency equals output energy / input energy
or 18 percent.
Furthermore, only 40-45 percent of the sun’s energy is in the PAR

region, so the theoretical maximum efficiency from all the sun’s energy is only
about 8 percent (45 percent of 18 percent). However, experimentally, the
efficiency of energy conversion in plants-the ratio of energy stored as organic
substance to the amount of energy available from the sun varies between 0.1 to
1.0% under conventional agricultural practices to between 6 and 10% under
conditions of intensive agriculture. Many crops including forest trees and
herbaceous species, convert only 1 to 2 percent of the PAR striking the field
during the growing season into stored carbohydrates. Much PAR is wasted by
striking the bare ground between young plants before leaves have grown enough
to absorb it.
Optimizing the interception of solar radiation by the canopy is an important

component of biomass production. This is influenced by the rate of development
of leaf area, leaf area duration and canopy architecture.
73

12.1.1 Photosynthetic efficiency of C3 & C4 crops:
In normal air, increasing temperatures gradually decrease efficiencies

of C3plants, with efficiencies of C4plants remaining constant. As temperatures
rise above 30oC efficiencies of the most C3plants become lower than those of C4.
This efficiency crossover with increased temperature results from lower net
photosynthesis in C3 plants because of faster CO2 loss by photorespiration. The
absence of detectable photorespiration in C4 plants even above 30oC gives them
a substantial efficiency advantage at high but not at low temperatures and
especially in non-shaded conditions
The relative photosynthetic efficiencies of C3 &C4 crops can be

calculated by considering the energetics of these processes occurring in plants.
Among C3, C4 and CAM pathways, it is clear that in C3 plants occurrence of
photorespiration leads to loss of Carbon through glycine decarboxylation. Further,
such oxygenase function of RuBISCO increase under abiotic stress such as
higher temperature, water stress etc. due to reduced levels of CO2 in sub-
stomatal cavity.
In course of evolution a relatively more efficient alternate C fixation

mechanism under water limited and high temperature environments was evolved
in plants i.e., “C4 pathways” as seen in C4 plants.
In terms of energy requirements:
 C3 photosynthesis under normal conditions requires 3 ATP +2NADPH for

fixation of one molecule of CO2.This is calculated from the following
equation.
6CO2 + 18 ATP + 12NADPH + 12 H+ C6H12O6 +18 ADP +18Pi + 12 NADP + 6H2O
 Under ambient CO2 concentration / and O2 conc. in the air, with

occurrence of photorespiration simultaneously in C3 plants this
requirement increases to approximately. 5ATP + 3.2 NADPH
On the Other hand
 In C4 plants the energy requirement for fixation of one CO2 molecule is 5 ATP
+ 2 NADPH
6CO2 + 30ATP +12NADPH+12H+ C6H12O6 + 30 ADP + 30 Pi +12NADP+ 6 H2O
 In C4 plants Photorespiration is negligible due to the CO2 concentration

mechanism .However, for concentrating CO2 in bundle sheath cells it needs
two additional ATPs. Thus, it needs 5 ATP.
74

 Any photo respiratory CO2 evolution from bundle sheath cells is refixed by
PEP carboxylase ase due to its high affinity for CO2
Thus, on the basis of

1. Energetics in lines with C loss due to photorespiration and
2. Affinity for CO2
It can be concluded that under ambient conditions compared to C3 plants, C4

are “Photosynthetically Efficient”.
12.2 SIGNIFICANCE OF C3, C4 & CAM PATHWAYS- A COMPARISON
C3 C4 CAM
Typically temperate species Typically tropical or semitropical Typically arid zone species
e.g. rice, wheat, barley, species e.g. Maize, sugarcane, e.g. cacti, orchids. Agave,
sugar beet, soybean, sorghum, Amarathus Plants succulent plants. Pine apple
sunflower, spinach, potato, adapted to high light, high is the only cultivated crop
tobacco, , oats and rye. temperature and also semiarid among this group.
Moderately productive, Highly productive, 80t per hectare Usually very poorly productive
yields of 30 tones dry for sugarcane is possible (However, Pineapple is highly
weight per hectare possible productive).
(sunflower is highly
productive)
Mesophyll Cells containing Kranz-type anatomy is essential Lack Kranz anatomy. Only
chloroplasts do not show feature. Often have two distinct one type of Chlroplast.
Kranz-type anatomy. Only types of chloroplast (both in Budle
one type of chloroplast. sheath and mesophyll cells)
Initial CO2 acceptor is Initial CO2 acceptor is phosphor CO2 acceptor is PEP in the
ribulose bisphosphate enol pyruvate (PEP) (a 3-carbon dark and RuBP in the light
(RuBP) a 5 carbon sugar. acid)
Initial CO2 fixation product Initial CO2 fixation product is the 4- CO2 fixation products are
is the 3 – carbon acid carbon acid oxalo acetate. oxalo acetate in the dark and
phosphoglycerate phospho glycerate in light.
Only one CO2 fixation Two CO2 fixation pathways Two CO2 fixation pathways
pathway separated in space. separated in time.
High rates of glycolate Low rates of glycolate synthesis, Low rates of glycolate
synthesis and no photorespiration. synthesis; no photorespiration
photorespiration
75

Low water use efficiency High water use efficiency and High water use efficiency and
and low salinity (ion) salinity tolerance. salinity tolerance.
tolerance
Photosynthesis saturates at Do not readily photo saturate at Do not readily photo saturate
1/5 full sunlight highlight, at high light.
High CO2 Compensation Low CO2 compensation point High affinity for CO2by night
point (50-150ppm) (0 – 10-ppm)
Open stomata by day Open stomata by day Open stomata by night

(photo active) (Photoactive)
( (Skoto active)
12.3 PHOTOSYNTHESIS AND CROP PRODUCTIVITY
Crop plants grow almost entirely by photosynthesis. Thus, plant productivity

interms of primary production of biomass is simply a measure of the total
photosynthesis of the plants less respiration, which has occurred during its growth.
DRY MATTER YIELD
CANOPY PHOTOSYNTHESIS
CANOPY RESPIRATION
PHOTO DARK
LEAF IRRADIANCE RESPIRATION RESPIRATIO
PHOTOSYNTHESIS N
LAI LEAF EXTENT OF

ANGLE LEAF DISTRIBUTION IRRADIANCE
PLANTING IN LEAF ARRANGEMENT ON PLANT

FIELD
76

Total Dry matter yield or total biomass is a consequence of crop

canopy efficiency in intercepting and utilizing the solar radiation variable during
the growing season. Leaves are the main plant organs which intercept solar
energy. For maximum crop growth rates, sufficient leaf area must be available in
the canopy to intercept most of the solar radiation incident on the crop canopy.
This is called as LAI. Mere interception of solar energy is not sufficient, it has to
be converted in to the biomass and it depends on the Net Assimilation Rate
(NAR) Thus, total dry matter production or productivity of a crop is a factor of how
long the crop can maintain an active, green leaf canopy that can produce
photosynthates.
CROP GROWTH RATE = LAI X NAR
*******
77

Lecture-13
ASSIMILATE TRANSLOCATION IN HIGHER PLANTS
13.1 TRANSLOCATION OF ASSIMILATES:
The assimilate transport is a process of exchange of metabolites among the
functionally specialized organs and tissues as a coordination of activities of plants as
whole.
This is essentially required to know
1. The distribution of photosynthetic products in plants
2. Their accumulation in storage organs
3. Their mobilization during resumed growth and
4. The effect of climatic factors and farming practices on the above process
13.1.1 Definition
The process of sugar movement from source to sink in sieve tube is called as
translocation.
13.1.2 Types of assimilate transport
The assimilate transport is of two types depending on their distance of transport.

A) Short distance transport: Movement of sugar from mesophyll cell to the vicinity
of sieve elements in the smallest veins of the leaf involving a distance of only
one or three cells diameter.
B) Long distance transport: Translocation of sucrose and other solutes from
source to sink inside the sieve elements is referred as long distance transport.
The two pathways function simultaneously and intimately inter twined

forming a continuous process encompassing the entire plant. These are extending in
parallel and pressed against each other resulting in the exchange of mobile
substance.
13.1.3 Anatomy of phloem tissue
Phloem is a complex tissue than the xylem. The main components of phloem
are sieve tubes. These are the longitudinally arranged individual cells called sieve
elements with perforated end walls called sieve plates. They are living cells. The
mature sieve elements appear to contain living protoplasm, although devoid of
nucleus, posses differentially permeably membrane but do not have a tonoplast.
Much of the protoplasm is in the form of P-Protein (phloem protein) a fibrilliar
protein.
78

*Figure is from reference.6

13.1.4 ASSIMILATE TRANSLOCATION IN RELATION TO “SOURCE” AND
“SINK”
Assimilate partitioning involves the production of assimilates in photosynthetic
organs (source) loading into sieve tubes, and translocation to the growing parts
(sink), where unloading takes place. An intact plant consists of multiple “Sources”
and “sink”
SOURCE
A source is any plant part that export carbon. Leaves are the principles sources of
assimilates.
SINK
The centers of storage or consumption of assimilates are the “Sinks”. A sink is an
organ that has a net import of assimilates which would be used for growth or storage.
All actively growing or metabolizing tissues are sinks.
13.2 PATHWAYS OF ASSIMILATE TRANSLOCATION
Assimilates after they are formed in the photosynthetic cells must traverse a
complex short distance of 3 to 4 parenchymatous cells (tenths of a millimeter) and
get concentrated in the phloem against the concentration gradient. This involves
additional energy expenditure.
In this route assimilate pass across the mesophyll cells in two pathways
separated or simultaneously. They are
13.2.1. Symplastic transport of assimilates (via plasmodesmata) (living part of
the plant)
Along this route, the mesophyll cells are interconnected with thin strands of
cytoplasm penetrating the cell walls in many places called plasmodesmata. Munch
79

(1930) termed this continuous protoplast in an aggregation of cells in complex

organisms as symplast.
Thus, symplastic transport can be defined as Translocation of substances
from one cell to another via plasmodesmata.
13.2.2 Apoplastic transport of assimilates (via free space)
In this pathway, assimilates leave the cytoplasm across its outer membrane,
on to the surface of the assimilating cells, (i.e. the apoplast) where in the solution
forming a film around the free intercellular space between cells.
Thus, apoplast includes all non living cells, cell walls and intercellular spaces
in stems and leaves.
13.3 ASSIMILATE TRANSLOCATION CAN BE UNDERSTOOD FROM THE
FOLLOWING HEADINGS
 Phloem loading
 Mechanism of phloem transport
 Phloem unloading
13.3.1 PHOLOEM LOADING
The site of phloem loading is the sieve cell- companion cell complex of
source. In source leaves, sugars become more concentrated in the sieve elements
and companion cells than in the mesophyll cells. So in phloem loading the sucrose is
transported against its chemical potential gradient and is evidence for active
transport of this solute.
Sugars might move entirely through the symplast via the plasmodesmata or,
alternatively might enter the apoplast at some point enroute to the phloem. In the
later case, the sugars could be actively loaded from the apoplast into the sieve
elements and companion cells by an energy driven carrier located in the plasma
membranes of these cells.
So the uphill transport of sucrose from the apoplast into the sieve cells
requires energy in the term of ATP. Giaquinta (1977) proposed sucrose proton
transport model involving energy. According to this an ATP ase builds up a proton
gradient across the plasma membrane by splitting ATP into ADP + Pi and excreting
the resulting H+ into the apoplast. The sucrose is taken up by sucrose proton and is
transported across membrane mediated by a carrier which is activated by
protonation. High ATP ase activity at the surface of the sieve element cell justified
this assumption.
80

ATP –dependent sucrose transport in sieve element loading
+
H ATP ase
ATP
+
H H
+
ADP+ Pi
+
H +
H
SUCROSE SUCROSE
+
+
Low H concentration
High H concentration
13.3.2 MECHANISM OF PHLOEM TRANSPORT (Munch pressure Flow

Hypothesis)
Munch pressure flow system is the only one understandable mechanism for
translocation of assimilates in plants. The workable hypothesis of Munch system is
mass flow or pressure flow. It is driven by metabolic processes of sources and sink
tissues. Phloem loading by the sources tissue exerts push and unloading by the sink
tissue pull, both driving the mass flow in the phloem. A sequence of physiological
steps occur in this system are shown in the following figure.
The assimilates from the mesophyll cells of source when actively loading up
in phloem, water potential of the phloem sap decreases. Consequently water is
drawn from xylem tissues into the sieve cells raising the hydrostatic pressure. This
pushes assimilates to flow enmasse, from one sieve tube to the other uninterruptedly
through open sieve pores. About 70% of sieve plate pores are freely kept open in a
number of other angiosperm species. Only on wounding callose, a carbohydrate is
synthesized which plugs sieve pores to protect from a major loss of photosynthates.
In the physiological sink, the assimilates are removed from the sieve cells thus
the water potential increased and water moves out of sieve cells into xylem. Both
osmotic water up take in the sieve cells of the source and water loss from the sieve
cells of sink, result in the circulation of water under pressure flow from the source to
the sink (translocation) in the phloem carrying assimilates, down to the sink cell and
water from sink cells to source cells (Transpiration) in the xylem.
81

Composition of phloem sap:

Sucrose is the most important mobile organic solute (200-300 mM) followed
by amino acids (30-200mM) and organic acids. Among inorganic solutes it is
potassium that dominates.
13.3.3 PHLOEM UNLOADING:
The process occurs at the site of sink. In many ways the events in sink
tissues are simply the reverse of the events in sources. Transport in to sink organs,
such as developing roots, tubers and reproductive structures, is termed import. The
following steps are involved in the import of sugars in to sink cells.
1. Sieve element unloading: This is the process by which imported sugars leave
the sieve elements of sink tissues
2. Short distance transport: After sieve element unloading, the sugars are
transported to cells in the sink by means of a short distance transport pathway.
This pathway has also been called post sieve element transport.
3. Storage and metabolism: In the final step, sugars are stored or metabolized in
sink cells.
82

This three transport steps together constitute phloem unloading, the

movement of photosynthates from the sieve elements and their distribution to the
sink cells that store or metabolize them.
PHOLEM UNLOADING IS BOTH SYMPLASTIC AND APOPLASTIC:
In vegetative sinks that are growing such as roots and young leaves,
phloem unloading and transport into receiver cells are usually symplastic. In other
sinks unloading is apoplastic.
Ex. Apoplastic unloading is required in developing seeds because there are no
symplastic connections between the maternal tissues and the tissues of the embryo.
When unloading is symplastic, transport of sugars occurs through
plasmodesmata to the receiver cells where it can be metabolized in the cytosol
vacuole i.e., Passive unloading occurs from a high concentration sieve elements to
low concentration in sink cell.
APOPLASTIC TRANSPORT IS ENERGY DEPENDENT:
When unloading is apoplastic, the transported sugars can partially be
metabolized in the apoplast (free space) it self. Ex. In sugarcane sucrose splits into
glucose and fructose by an enzyme invertase in the apoplast itself and glucose or
fructose that is taken up by the receiver cell.
In case of apoplastic unloading, sugars must cross at least two membranes
1. Membrane of sieve element and companion complex
2. Membrane of the receiver cell.
This transport of sugars across the membrane is shown to be active Thus
apoplastic transport is an energy dependent process. Phloem unloading is strictly
controlled by sink metabolism. Faster the metabolism greater will be the sink
demand.
It also depends upon the activity of phytohormones .In case of legumes, number
of nodules and nodule activity also controls phloem unloading.
*******
83

Lecture-14 SOURCE AND SINK CONCEPT
14.1 SOURCE AND SINK CONCEPT
Determination of factors that influence yield is a large and intensively studied

area of crop physiology. Yield is the manifestation of all physiological processes
occurring in plants. There are different types of yield among the domesticated crops.
They are as follows.
Crop Types of yield / Storage organ
Cereals Seed / grains mainly for CH2OS
Pulses Proteins
Oilseeds Seed mainly for lipids / oils / waxes
Sugarcane Cane for sugar
Fibers Bolls for lint especially in cotton
Tubers Tubers for CH2Os
Fodders Fresh or dry weight of stalk
Since yield is the resultant of dry matter partitioned between the different parts
of plant, possibilities of changing the distribution of assimilates in crops by
physiological manipulation is one of the most promising ways of increasing
agricultural productivity. To achieve this, knowledge of source – sink concept is
important to understand the direction of transport of assimilates.
The movement of assimilates from source to sink is currently believed to occur as
follows
1) The photosynthetic source cell produces the sugars, which can move
symplastically or apoplastically to the sieve tubes.
2) Phloem loading increases the sugar concentration of sieve tubes above that
of the apoplast.
3) At the sink, carbohydrates are being absorbed and either actively partitioned
into cell constituents (eg. Starch) or changed to other carbohydrates. Phloem
unloading lowers the concentration of sugars in sieve tubes.
4) The buildup of sugars at the source and the removal of sugar at the sink
establish a hydrostatic pressure gradient, which moves water and sugar from
sources to sinks.
Thus, higher production of assimilates (source strength), their rapid
translocation and utilization in growth and development (sink strength) are some key
factors of concern to increase crop yield.
14.2 DRY MATTER PARTITIONING IS DECIDED BY SOURCE STRENGTH AND
SINK STRENGTH:
14.2.1 SOURCE STRENGTH:
Source strength is a function of source size and source activity.
84

Source strength = Source size X source activity

The total amount of photosynthates produced in a crop is decided by the
optimum crop canopy size (actively photosynthesizing leaf area or source size) and
the rate of assimilation per leaf area (photosynthetic efficiency or source activity) of
crops. The estimates of efficiency of utilization of photo synthetically active radiation
(PAR) some crops is given below.
Crop Utilization of PAR (%)
Maize (Zea mays) 6.2
Sugarcane (Saccharum spp) 7.7
Rice (Oryza sativa) 6.2
Sugar beet (Beta vulgaris) 8.8
Soybean (Glycine max) 7.7
14.2.2 SINK STRENGTH:

Sink strength is a function of sink size and sink activity.
Sink Strength= sink size X sink activity
Sink size or sink capacity is the maximum space available for accumulation of
photosynthetic products.In grain crops it is expressed as number and size of grains.
Sink activity is the inherent capacity of the sink to create a translocation gradient for
photosynthates assimilated at source.
According to the mass flow hypothesis, any increase in photosynthesis
increases hydrostatic pressure and translocation rate. However, this is true only if
sinks have the ability to utilize more assimilates. If they are unable to utilize the
increased production there would be a steady build up of sugars in the system
causing a feedback inhibition resulting in reduced photosynthesis. Presumably,
photosynthetic rate would be altered to the rate at which sinks could accept
assimilates.
Thus, for leaf photosynthesis to be at maximum potential rates, sinks must
be able to rapidly utilize the assimilates produced by the source. Under these
conditions partitioning would be controlled by sink strength that is sink availability
and the rate at which available sinks can utilize assimilate (sink demand).
Role Of Hormones In Alteration Of Source-Sink Relationship: The effect of
hormones on enzymatic activity and the elasticity of sink cells can have a dramatic
effect on partitioning. Indoleacetic (IAA), cytokinins, ethylene and gibberllic acid,
when applied to cut stem surfaces causes assimilates to accumulate in the region of
application In bean seedlings, the main control over the distribution of sucrose
between root and shoot sinks can be attributed to auxin and cytokinin concentrations
in various sinks. Hormonal influences on initiation, development and abortion of
flowers and seeds have a significant effect on the source – sink relationships in
crops.
85

14.3 HARVEST INDEX
Two useful terms used to describe partitioning of dry matter by the plant are
Biological yield & economic yield. The term biological yield was proposed by
Nichiporovich (1960) to represent the total dry matter accumulation of a plant’s
system. Economic yield and agricultural yield have been used to refer to the volume
or weight of those plant organs that constitute the product of economic or agricultural
value. The proportion of biological yield represented by economic yield has been
called the harvest index or the migration coefficient. These terms characterize the
movement of dry matter to the harvested part of the plant. The harvest index, the
most widely used tem, is defined as follows.
Economic yield
Harvest index = ----------------------- X 100
Biological yield
(It must be remembered that the biological yield total, often does not include root
weight because of the difficulty in obtaining those values)
Crop yield can be increased either by increasing the total dry matter produced
in the filed or by increasing the proportion of economic yield (the harvest index) or
both. There is potential for increasing yields by both methods.
The economic fraction of biological yield usually referred to harvest index
varies among the species.
Crop Harvest Index (%)
Cereals 50 – 60
Pulses 30 – 45
Oilseeds 45 – 50
Sugarcane 35
Yield of a crop is a function of biomass X Harvest index .Grain yield in
cereals is a product of dry matter accumulation and partitioning.
Thus, grain yield can be understood as product of a number of subtractions called
yield components which can be expressed as follows.
Y = Nr Ng Wg
Where, Y = grain yield
N = number of reproductive units (e.g heads, ears, panicles per
unit of ground areas)
Ng= the number of grains per reproductive units and
Wg- the average weight per grain
Thus, the economic yield (Harvest index) depends upon the total dry matter
produced and the amount of which is partitioned towards reproductive units in the
post anthesis period. Yield components are affected by management, genotype and
environment.
*******
86

Lecture-15
RESPIRATION
15.1 INTRODUCTION
The word respiration is derived from the Latin word ’respirare’ (literally, to
breathe). Aerobic (oxygen requiring) respiration is common to nearly all eukaryotic
organisms. It is a biological process by which reduced organic compounds are
mobilized and subsequently oxidized in a controlled manner. During respiration,free
energy is released and transiently stored in a compound, ATP, which can be readily
utilized for the maintenance and development of plants.
Respiration can be defined as ‘an oxidation-reduction process’ in which
respirable substrates such as Carbohydrates, proteins, fats etc. are oxidized to CO2
and O2 absorbed is reduced to water, with the release of chemical energy stored in
the bonds of the complex molecules (ATP).
The overall respiration process may be represented as
C6H12O6 + 6 O2 6CO2+ 6H2O + 686 K.Cal.
Mitochondria and respiration:

Various enzymes of oxidation of pyruvic acid (TCA Cycle) and electron transport
chain (which is a series of oxidation-reduction reactions) are located in matrix of
Mitochondria .The mitochondrion is made of double unit membrane. The inner layer
of this membrane is deeply folded inward to form the cristae (transverse membranes)
that lie more or less cross wise in the mitochondrion. The membrances appear to
have small knob like structures about 70 A0 in diameter called F1 – ATP ase
attached to their inner surface by stalks about 30 A0 long called Fo particles. These
structures are concerned with the synthesis of ATP, the cells energy mobilization
compound.
*Figure. is from reference 6
87

15.2 SIGNIFICANCE OF RESPIRATION:

Respiration is an important process because
1. It releases energy which is consumed in various metabolic processes essential for
plant life and activates cell division.
2. It brings about the formation of other necessary compounds participating as
important cell constituents.
3. It converts insoluble food into soluble form.
4. It liberates CO2 and plays a part actively in maintaining the balance of carbon cycle in
nature.
5. It converts stored energy (potential energy) into usable form (kinetic energy).
Respiration is complex process which includes:
i. Absorption of oxygen.
ii. Conversion of carbohydrate (complex) to CO2 and water
(simpler substances) i.e., oxidation of food.
iii. Release of energy.
iv. Formation of intermediate products playing different roles in
metabolism
v. Loss of weight in plants as a result of oxidation.
15.3 Out line of Respiratory Metabolism:

The breakdown of a respiratory substrate in respiration, proceed
through a series of reactions each catalyzed by a specific enzyme. The sequence of
reactions is generally referred to as respiratory metabolism. Which Includes the
following steps.
STEP I : Glycolysis:
This step operates in the cytoplasm and is common to both aerobic
and anaerobic respiration. Pyruvic acid (CH3COCOOH) – a key respiratory
metabolite – is formed from oxidation of glucose (starting point for respiratory
metabolism in plants) in a series of reactions collectively called as Glycolysis.
Oxidative Pentose Phosphate Pathway
It is another sequence of reactions where in glucose molecule after its
phosphorylation as Glucose – 6 – Phosphate is directly oxidized to phosphogluconic
acid and then to ribulose – 5 – phosphate. Electrons released in between these two
steps reduce NADP to NADPH2 , each molecule of which after passing through ETS
system produces 3 molecules of ATP like in glycolysis. This is another way of
breakdown of glucose and production of energy.
STEP-II: Tricarboxylic Acid Cycle (TCA cycle/Krebs Cycle/ Citric Acid Cycle)
Pyruvic acid produced by glycolysis is oxidized in a sequence of
reactions referred to as tricarboxylic acid cycle. Several intermediates of the tri
carboxylic acid cycle under go decarboxylations so that CO2 is produced. Other
88

intermediates are oxidized through the agency of the co – enzyme NAD+ and hence
NADH (reducing power) is generated in the process.
STEP III: Respiratory chain / Electron Transport Chain
NADH is oxidized via, the respiratory chain, so that NAD+ is
regenerated. Electron Transport Chain transfers electrons from NADH – produced
during glycolysis, the pentose phosphate pathway and the TCA cycle – to Oxygen.
O2 is the final electron acceptor in the respiratory chain and is reduced to water.
This electron transfer releases a large amount of free energy much of which is
conserved through the synthesis of ATP from ADP and Pi catalyzed by the enzyme
ATP Synthase. Collectively the redox reactions of the electron transport chain and
the synthesis of ATP are called Oxidative Phosphorylation. (synthesis of ATP in
light reaction of photosynthesis is called as photophosphorylation).
*Figure is from reference 6
15.4 GLYCOLYSIS:
The reactions of glycolysis process may be divided into two phases:

Preparatory phase and Oxidative phase. Preparatory phase consists of 4 steps
and oxidative phase of 5 steps. During preparatory phase glucose is converted into
fructose – 1,6 diphosphate and during oxidative phase fructose 1,6 diphosphate split
into two three carbon compounds which are converted to pyruvic acid.
89

REACTIONS OF GLYCOLYSIS
Preparatory phase
In this step glucose is phosphorylated with ATP in the presence of the

enzyme hexokinase to produce glucose-6- phosphate and ADP. Then glucose 6
Phosphate is converted to its isomer fructose-6- Phosphate in the presence of
phospho gluco isomerase enzyme. Fructose 6 phosphate in the presence of ATP
molecule and the enzyme phopho hexokinase forms fructose 1-6 diphosphate and
ADP.
Oxidative phase
In this step splitting of fructose 1,6 – diphosphate into two three – carbon
compounds – 3 phopho glyceraldehyde and dihydroxy acetone phosphate in the
presence of enzyme aldolase occurs. These two sugars are inter converted through
the enzyme phopho triose – isomerase.
90

If 3 – Phosphoglycerladehyde is depleted, additional amounts will be formed by

the isomerization of dihydroxyacetone phosphate, continuing with glycolysis, 3-
phosphoglyceraldehyde is converted to 1,3 – diphosphoglycerate. This reaction
involves the incorporation or addition of inorganic phosphate to first carbon of 3
phosphoglyceraldehyde and the reduction of NAD+ to NADH and is catalyzed by the
enzyme phosphoglyceraldehyde dehydrogenase.
The consumption of incorgnic phosphate in the oxidation of 3 –

phosphoglyceraldehyde is important to the plant because this phosphate is involved
in the synthesis of ATP in the next reaction of the glycolytic sequence. In the
presence of ADP and the enzyme phosphoglycero kinase, 1,3-diphosphoglycerate is
converted to 3 - phospho-glycerate and ATP. The process of ATP formation by the
transfer of phosphate from one of the intermediates of the pathway (in this case; 1,3
– diphosphoglycerate ) to ADP is termed as substrate level phosphorylation .
This process represents the major way in which ATP is generated from bond energy
under anaerobic conditions and is particularly important to fermentation.
The 3 – Phosphoglycerate that is formed in the above reaction is

transformed to 2 – phosphoglycerate by the activity of the enzyme
Phosphoglyceromutase. The elimination of the elements of water (dehydration) from
2 – Phosphoglycerate in the presence of enolase results in the formation of
phosphoenolpyruvate. In the presence of ADP and pyruvate kinase,
phosphoenolpyruvate is transferred to ADP to form ATP, another example of
substrate level Phosphorylation,
5.4.1 Significance of Glycolysis:
The EMP pathway( or glycolysis) , which is also referred to as the

hexose diphosphate pathway, is the chief pathway in which glucose or intermediates
are converted to pyruvate (Pyruvic acid). It involves the inter conversion of sugars
and transfer of phosphate groups and the ultimate conversion of one six – carbon
compound to two three – carbon molecules. It is an anaerobic pathway in which
some NADH and ATP is generated. The ATP is generated by substrate level
phosphorylation. The overall reaction sequence for the EMP pathway is based on
the following:-
Overall Reaction Sequence
1 Glucose + 4 ADP + 2 ATP + 2 Pi + 2NAD 2 Pyruvate + 2 ADP + 4 ATP + 2 NADH
Balanced Reaction
1 Glucose + 2 ADP + 2Pi + 2NAD 2 Pyruvate + 2 ATP + 2 NADH
91

In the first phase, the conversion of glucose to fructose – 1,6 –

diphosphate, there is no energy gain. Indeed, two ATP molecules are consumed for
every glucose molecule phosphorylated. However, in the second phase, the
conversion of fructose – 1,6 – diphosphate to two molecules of pyruvate, four ATP
molecules are formed – two for each triose split off from fructose – 1,6 –
diphosphate. If we consider the complete EMP pathway, the conversion of one
molecule of glucose to two molecules of pyruvate results in a net gain of two ATP
and two NADH molecules.
In the sequence of reactions of glycolysis there is a net gain of two molecules of

ATP. These steps may be summarized as follows:
Glucose + 2ATP Fructose 1,6-diphosphate + 2ADP
2 ATP molecules are used
2(1,3 – DiPhosphoglyceric acid) + 2ADP (3- phosphoglyceric acid) +2 ATP
2 (2- phosphoenolpyruvic acid) + 2ADP 2 Puruvic acid + 2ATP
4 ATP molecules produced
(Since 2 ATP are used, there is a net gain of 2 ATP only)
However, Two NADH2 molecules are also produced in the process and
from its each molecule 3 ATP molecules are produced. As a result 6 more ATP
molecules are formed, though these are from ETS chain. In eukaryotes, NADH2 from
Glycolysis enters one step late in ETS (due to loss of energy or utilization of one ATP
in transfer), hence it yields only 2 ATP molecules, but in prokaryotes, it yields 3 ATP
molecules.
15.5 FATE OF PYRUVATE TO CO2 AND H2O: (Aerobic Respiration)
Breakdown of Pyruvate is of considerable importance and significance

from the stand point of energy production. In the absence of oxygen it may be
degraded to Ethyl alcohol and CO2 or Lactic acid or Alanine but in presence of
Oxygen the usual fate of Pyruvate is to degrade up to CO2 and H2O level through
Krebs Cycle and respiratory chain in which energy is trapped in the form of ATP
molecules. Mitochondria have all the enzymes necessary for complete oxidation of
pyruvic acid to CO2 and H2O. All enzymes of TCA cycle are found in the
mitochondrial matrix. A summary representation of the scheme can be Krebs Cycle
Pyruvate CO2 + electrons + Protons
Respiratory Chain (electron transport chain)
Pi + ADP + electrons + protons + Oxygen Water + ATP
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15.6 FORMATION OF ACETYL COA
Pyruvic acid cannot directly enter in to the krebs cycle and therefore
converts in to Acetyl Co-A. The Oxidative decarboxylation of Pyruavte into Acetyl
CoA involves the presence of at least five essential co factors and a complex
enzyme. The Cofactors involved are Mg ions, thiamine pyrophosphate (TPP), NAD +,
Co enzyme A (CoA) and Lipoic Acid.
15.7 KREBS CYCLE (CITRIC ACID CYCLE, TRICARBOXYLIC ACID CYCLE)
The first reaction of the krebs cycle is the condensation of acetyl CoA with
oxaloacetate to form citric acid and release CoA. The result of this reaction,
catalyzed by condensing enzyme, is that a four carbon dicarboxylic acid is converted
to a six carbon tri carboxylic acid.(Fig. )
Figure is from reference 6

Through a series of reactions involving four oxidation steps and three
molecules of H2O (one utilized in the condensation reaction), oxaloacetic acid is
regenerated from citric acid. In the process two molecules of CO2 and 8 H atoms are
released. We should note that these acids and other organic acids of the Krebs cycle
are displayed in the ionized form (R – COO-) and are labeled accordingly (i.e.,
Citrate, oxaloacetate, and so on.)
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Acetyl CoA is the connecting link between glycolysis , and the Krebs Cycle (citric
acid cycle or tricarboxylic acid cycle) so named because of the cyclic manner in
which the starting compound, oxaloacetate is regenerated. The cycle is named after
an English biochemist Krebs who played a major role in its discovery.
15.8 SIGNIFICANCE OF TCA CYCLE:
In the four oxidation steps four pairs of H+ ions and four pairs of electrons are
removed from the intermediates of the cycle. Three of these pairs of H+ ions and
electrons are utilized in the reduction of pyridine nucleotides (NAD). The one
remaining pair of H+ ions and electrons is taken up in the reduction of the FAD
prosthetic group of succinic dehydrogenase.
The NADH and FADH produced by the reactions of the Krebs cycle represent
reducing power that can be harnessed in the presence of oxygen to produce ATP.
The production of ATP is accomplished by a series of oxidation-reduction reactions
involving chemicals that comprise the electron transport system (ETS). The ETS is
intimately associated with the mitochondrial membranes and the Krebs cycle
apparatus.
By means of the Krebs cycle, and the electron transport system, pyruvate is
oxidized to CO2 and H2O. Thus the complete oxidation of glucose to CO2 and H2O
may occur through glycolysis, the Krebs Cycle, and the electron transport system.
Through its association with the electron transport system, the oxidations of the
krebs cycle can account for the formation of 24 ATP molecules. Thus the krebs
cycle is far more efficient in the release of energy than are either glycolysis or
fermentation. The reactions of the krebs cycle and the electron transport system
require the presence of oxygen and are confined to the mitochondria.
15.9 ELECTRON TRANSPORT SYSTEM AND PHOSPHORYLATION:
For aerobic organisms it is essential that the enzymes and reduction products
of the Krebs cycle be associated with the Electron Transport System. Through this
association, the reduced pyrimidine nucleotide NADH, FADH, and (rarely) NADPH
are reoxidized. The energy released from these oxidations is utilized in the synthesis
of ATP. The synthesis of ATP via electron flow through the ETS, with oxygen as the
terminal electron acceptor, is known as oxidative phosphorylation and takes place in
mitochondria.
In essence, the ETS is a chain of carriers consisting of nicotinamide adenine

dinucleotide (NAD) flavin nucleotides (FAD, sometimes FMN,) Co enzyme Q (CoQ) ,
and the cytochromes (cyt b, c, a, a3) and non-heme iron proteins also seem to be
involved but their role is not known exactly.
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Most important to the living plant is the fact that each step in the system is
characterized by a decrease in the energy level. In other words, the carriers
presumably operate in order of an increasing tendency to undergo reduction
(reducing potential becomes increasingly positive from NADH through cytochrome
a3.
The electron will flow from a higher to lower energy level. Thus with each step in
the system the energy level of the electron is lowered and the energy difference is
transformed into phosphate bond energy by the conversion of ADP to ATP.
Hydrogen ions are released in the oxidation of reduced co enzyme Q (CoQ).
These hydrogen ions may play an important role in ATP production. Only the
electrons are passed along the series of cytochromes.
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15.9 SIGNIFICANCE OF ELECTRON TRANSPORT CHAIN:
A further study will show that for every pair of electrons passed along this system,
three ATP are formed. The synthesis of ATP occurs in the oxidation of
NADH, in the oxidation two cytochrome b’s and in the oxidation of two cytochrome
a’s. At their lowest energy level, the electrons are passed to oxygen from reduced
cytochrome a3 thereby activating the oxygen. In this state, oxygen will accept free
hydrogen ions to form water.
15.10 ENERGY BALANCE SHEET OF RESPIRATORY METABOLISM:
If we now consider the complete degradation of a molecule of glucose, first to

two pyruvic acid molecules via the EMP pathway and then to the two acetyl CoA
molecules through the oxygen-requiring Krebs cycle to CO2 and water, it is possible
to have thirty-eight ATP molecules generated.
Pathway NADH (3ATP) FADH (2ATP) ATP Total ATP
EMP (yield) 2 (6) 0 2 8
Pyruvic acid to 2 (6) 0 0 6

Acetyl CoA
Krebs Cycle 6 (18) 2 (4) 2 24

(yield)
Total ATP 10 X 3 = 30 2X 2 = 4 4 38 ATP
15.11 THE PENTOSE PHOSPHATE PATHWAY PRODUCES NADPH AND
BIOSYNTHETIC INTERMEDIATES:
The glycolytic pathway is not the only route available for the oxidation of
sugars in plant cells. Sharing common metabolites, the oxidative pentose
phosphate pathway (also known as the hexose monophosphate shunt) can also
accomplish this task. The reactions are carried out by soluble enzymes present in
the cytosol and in plastids. Generally, the pathway in plastids predominates over the
cytosolic pathway.
The First two reactions of this pathway involve the Oxidative events that
convert the six – carbon glucose – 6 – phosphate to a five – carbon sugar, ribulose 5
– phosphate, with loss of a CO2 molecule and generation of two molecules of
NADPH (not NADH). The remaining reactions of the pathway convert ribulose 5
phosphate to the glycolytic intermediates glyceraldehyde,3- phosphate and fructose-
6- phosphate. Because glucose 6 phosphate can be regenerated from
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glyceraldehyde- 3- phosphate and fructose- 6- phosphate by glycolytic enzymes, for

six turns of the cycle we can write the reactions as follows:
6 Glucose-6-P +12NADP+ +7H2O 5 Glucose-6-P+ 6 CO2 + Pi + 12NADPH + 12H+
The net result is the complete oxidation of one glucose – 6- phosphate molecule to
CO2 with the concomitant synthesis of 12 NADPH molecules.
14
Studies of the release of CO2 from isotopic ally labeled glucose
indicate that glycolysis is the moiré dominant breakdown pathway, accounting for 80
to 95% of the total carbon flux in most plant tissues. However the pentose
phosphate pathway does contribute to the flux and developmental studies indicate
that its contribution increases as plant cells develop from a meristematic to more
differentiated state.
The oxidative pentose phosphate pathway plays several roles in plant

metabolism:
 The Product of two oxidative steps is NADPH, and this NADPH is thought to
drive reductive steps associated with various biosynthetic reactions that occur
in the cytosol such as lipid biosynthesis and nitrogen assimilation.
 Some of the reducing power generated by this pathway may contribute to
cellular energy metabolism; that is, electrons from NADPH may end up
reducing O2 and generating ATP.
 The pathway produces ribose 5 phosphate, a precursor of the ribose and
deoxyribose needed in the synthesis of RNA and DNA respectively.
 Another intermediate in this pathway, the four carbon erythrose – 4 –
phosphate, combines with PEP in the initial reaction that produces plant
phenolic compounds including the aromatic amino acids and the precursors
of lignin, flavonoids and phytoalexins.
 During the early stages of greening, before leaf tissues become fully
photoautotrophic, the oxidative pentose phosphate pathway is thought to be
involved in generating Calvin Cycle intermediates.
*******
97

Lecture-16
INTER RELATIONSHIP OF RESPIRATION AND PHOTOSYNTHESIS
Total dry matter produced by a crop is a function of canopy photosynthesis

and canopy respiration. The difference between these two factors gives us is the
amount of net photosynthates that are actually available for the growth and
development of the plant. Canopy respiration includes both dark respiration and
photorespiration.
Survival of any plant is decided by its metabolic activities. Plant needs energy in
the form of ATP for these metabolic activities to run. This ATP is generated by the
oxidation of the stored food materials like carbohydrates, proteins and lipids. The
stored food materials are however produced by photosynthesis. (Fig)
*Figure is from reference 6
Thus, for its survival the plant must always maintain a photosynthetic –
respiratory ratio exceeding 1. Otherwise, plants are starved to death. A greater ratio
may result from a daily change between relatively cool night temperatures and
moderately high day temperatures.
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This interrelationship can be better understood from the concept of growth and
maintenance respiration.
16.2 GROWTH AND MAINTENANCE OF RESPIRATION:
Respiration provides opportunity for two different sorts of processes in

plants: maintenance and growth.
Maintenance is primarily concerned with repair and restoration and the

operation of all the metabolic systems necessary for the normal functioning of the
plant. These include turnover, transport, maintenance of gradients of all sorts and
the requirements for operating controls and signal system. Growth is mainly the
synthesis and accumulation of all the material and operational systems that
constitute the plant.
Growth and maintenance are two quite different processes. American

agriculturist K I MeCree has shown how they can be quantified independently.
Maintenance respiration is clearly a function of plant size (at least in herbaceous
plants those do not possess large masses of metabolically inert tissue) so it can be
represent as cW. Where “c “is a constant, W is the dry weight of the plant. Growth
respiration, however, depends only on the actual new growth being made by the
plant. Which is best measured as the net rate of photosynthesis. So growth
respiration may be represented of “kP” Where “k” is another constant and P is the
rate of Photosynthesis.
Thus the equation for respiration is the R= kP + cW.
Therefore, the rate of photosynthesis of a plant should always surpass the total
value of growth and maintenance respirations putting together. This is very important
for achieving higher yields particularly in root crops and tuber crops where large
amount of photosynthates are needed for storage.
16.3 Respiratory Substrates:
A respiratory substrate is any organic plant constituent oxidized

partially or completely(to CO2 and water)in respiratory metabolism. Carbohydrates
are the principal respiratory substances in cells of higher plants. The most important
respiratory substrates among carbohydrates are Sucrose & starch. In addition of
Carbohydrates, other substances also some times serve as respiratory substrates in
some plant tissues.
Eg: Castor Lipid reserves in endosperm.
CAM plants Organic acids ( Malic acid, Glycolic acid etc.)
Detached Leaves Proteins
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16.4 Respiratory quotient: (R.Q)
Respiratory quotient may be defined as the ratio between the volume of

carbon dioxide given out and oxygen taken in simultaneously by a given weight of
the tissue in a given period of time at standard temperature and pressure.
Volume of CO2 evolved
Respiratory quotient = -----------------------------------
Volume of O2 absorbed
For example C6H12O6 + 6O2 6CO2 + 6H2 O
In this reaction volume of CO2 equals to volume of O2. Thus the RQ of

carbohydrates (for example Glucose ) is 1.
The following table enumerates R.Q. values of different substrates
Substrates R.Q.
Carbohydrates 1.00
Proteins when ammonia is 0.99

produced in oxidation
Proteins with amide formation 0.80
Fats 0.70
Organic Acids 1.33
16.5 ALTERNATE RESPIRATION/CYANIDE RESISTANT RESPIRATION:
The Alternative Oxidase

If cyanide ( 1 m/M ) is added to actively respiring animal tissues, cytochrome c
oxidase is inhibited and the respiration rate quickly drops to less than1 % of its initial
level. However, most plant tissues display a level of cyanide- resistant respiration
that can represent 10 to 25 %, and in some tissues up to 100 %, of the uninhibited
control rate. The enzyme responsible for this oxygen uptake been identified as a
cyanide – resistant oxidase component of the plant mitochondrial electron transport
chain called the alternate oxidase.
It appears probable that mitochondria in tissues exhibiting the alternate pathway
have two parallel electron transport chains from substrate ( e.g., NADH ) to O2 .
One is the cyanide – sensitive respiratory chain terminated by cytochrome oxidase.
The second (alternate) pathway is terminated by the alternate oxidase. Electrons
feed off the main electron transport chain into the alternative pathway at the level of
the ubiquinone pool. The alternate oxidase, the only component of the alternative
pathway catalyzes a four – electron reduction of oxygen to water.
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ADP + Pi ATP
NADH FLAVOPROTEINS CO-Q ALTERNATE O2 H2O

OXIDASE
Alternative pathway contribute to plant metabolism:

One example of the functional usefulness of the alternate oxidase is its activity
during floral development in certain members of the Araceae (the arum family) – for
example, the voodoo lilly.
Just before pollination tissues of the club like inflorescence, called the appendix
which bears male and female flowers, exhibit a dramatic increase in the rate of
respiration via the alternative pathway. As a result, the temperature of the upper
appendix increases by as much as 25O C over the ambient temperature for a period
of about 7 hours.
During this extraordinary burst of heat production, certain amines, indoles,
and terpenes are volatilized, and the plant therefore gives off a putrid odor that
attracts insect pollinators.
It has been suggested that the alternative pathway can function as an “energy
over flow” pathway, oxidizing respiratory substrates that accumulate in excess of
those needed for the growth, storage, or ATP synthesis.
16.6 SALT RESPIRATION: When roots are absorbing salts, the respiration rate
rises. This rise has been linked to the fact that energy is spent in absorbing salts or
ions, and that required energy is supplied by increased respiration. This
phenomenon called as salt respiration.
The inference is that respiration represents the increased metabolism

needed to generate energy for the active transport of ions. Unfortunately the
relationship is not always linear, and salt respiration may persist after the salts are
removed. Consequently, salt respiration does not offer many useful clues to the
nature of the coupling of respiration and ion transport.
16.7 WOUND RESPIRATION:

Wounding of a plant organ stimulates respiration in that organ. It initiates
meristematic activity in the region of the wound resulting in the development of
‘wound callus’. It has been shown that wounding is correlated with an increase in the
sugar content of the half cut potato tuber. Perhaps the increase in respiration is due
to an increased availability of respiratory substrate in wounded tissues.
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16.8 MEASUREMENT OF RESPIRATION:

Methods for measuring the rates of respiration involve quantitative
determination of the CO2 evolved or the oxygen consumed. An individual plant or a
plant part or tissue slices, or cell suspension or tissue homogenate or mitochondria
forms the sample
DIFFERENT METHODS OF MESUREMENT OF RESPIRATION:
There are five methods of measurement of respiration
A. Weight method: The CO2 produced during respiration by a plant tissue is
trapped in a barium hydroxide solution. Weight of the barium carbonate formed
indicates the amount of CO2 produced.
B. Titration Method: The CO2 produced is made to dissolve in NaOH and the
amount of CO2 absorbed is determined by titration.
C. Quantitative method: Release of CO2 can be measured by passing an air
stream through a sealed chamber containing plant tissue into an alkaline solution
where changes in PH or E.C. are recorded. From these recorded changes the
quantity of CO2 released by tissue is calculated.
D. Mano metric method: The rates of O2 uptake or CO2 release by plant tissue are
measured with a mano meter. In this method, sliced samples of plant tissue are
suspended in water in a flask. Changes in the amount of respiratory gases with in
the flask are reflected in changes in its pressure. These pressure changes can be
measured with a manometer.
E. Infra red CO2 analyzer or paramagnetic O2 analyzer method:
This is a Sophisticated instrument capable of measuring changes in CO2 or O2 of the
atmosphere and chambers in which plants, or bulky tissues such as large fruits are
sealed.
In addition, a membrane bound polarographic electrode sensitive to O2 can
measure changes in the concentration of O2 in a solution. It should also be
mentioned that mass spectrometric methods are often used in the recent years
to study the gas exchange between plant tissues and the external atmosphere.
*******
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Lecture: 17
NUTRIO PHYSIOLOGY
17.1 DEFINITIONS:
Nutriophysiology deals with metabolic and biochemical functions of the
chemical elements and their interaction with other aspects of plant physiology and
plant biochemistry. It deals with initial acquisition of chemical elements, their
distribution with in the plant and interactions of the plant with its chemical media.
The supply and absorption of chemical compounds needed for growth and
metabolism of plants may be defined as Plant Nutrition and the chemical
compounds required by the plants are termed as Nutrients.
Higher plants are autotrophic organisms that can synthesize their organic
molecular components out of inorganic nutrients obtained from their surroundings.
For many mineral nutrients, this process involves absorption from the soil by roots
and incorporation in to the organic compounds that are essential for growth and
development. This incorporation of mineral nutrients into organic substances such as
pigments, enzyme cofactors, lipids, nucleic acids and amino acids is termed as
nutrient assimilation. Nutrition and metabolism are thus very closely related.
17.2 ESSENTIAL PLANT NUTRIENTS AND CRITERIA OF ESSENTIALITY:

Plants contain about 80 to 90 percent of water by weight and the remaining 10
to 20 percent is the dry weight. The dry matter consists of a number of organic
compounds such as carbohydrates, proteins, lipids and others. Nearly 90 percent of
the dry weight of the plant consists of carbon, hydrogen and oxygen. All the mineral
elements together contribute only about 6 percent of the dry weight of the plant.
The finding of certain element in plant does not signify that this element is
essential for the growth of the plant. For finding out whether an element is essential
or not, Arnon and Stout (1939) proposed the criteria of essentially. These criteria
are as follows.
A. A deficiency of the element makes it impossible for a plant to complete its life
cycle.
B. The functions of an element cannot be replaced by another element.
However, recent studies have shown that some of these elements can be
partially replaced by others for example magnesium (Mg) by manganese (Mn)
and Potassium (K) by rubidium (Rb)
C. The essential element must be directly involved in the metabolism of plant or it
may be required for the activation of an enzyme system.
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Based on the above criteria the following elements are now known to be
essential for higher plants.
Carbon C Magnesium Mg
Hydrogen H Iron Fe
Oxygen O Manganese Mn
Nitrogen N Copper Cu
Phosphorus P Zinc Zn
Sulphur S Molybdenum Mo
Potassium K Boron B
Calcium Ca Chlorine Cl
Sodium Na
Silicon Si
Cobalt Co
17.3 MACRO AND MICRO NUTRIENTS:

Based on the element concentration in plant material, the essential plant
nutrients may be divided into macronutrients and micronutrients. Macronutrients
are found or needed in relatively higher amounts than micronutrients. For example,
the content of the macronutrient N is thousand times grater than the content of
micronutrient Zn in plant tissue. Following this classification
C,H,O,N,P,K,Ca,S,Mg,(Na,Si) may be defined as macronutrients. The micronutrients
are Fe,Mn,Cu,Zn,Mo,B,Cl.
BENEFICIAL ELEMENTS:
Sodium has beneficial effect and in some cases it is essential. There are
some plant species, particularly the chenopodiaceae plants and species adapted to
saline conditions that take up this element in relatively high amounts. Na is also
required for turnips, sugar beets and celery. The same is true for Si, which is an
essential nutrient for rice. Cobalt is an essential element for the growth of the blue-
green algae, but it has not been shown to be essential for other algae or for higher
plants. It is also required by certain legumes to fix atmospheric nitrogen. Here,
however the cobalt ion is necessary for the symbiotic bacteria present in the nodules
associated with the roots.
This division of the plant nutrients into macro and micro nutrients is some
what arbitrary and in many cases differences between the contents of macronutrients
and micronutrients are considerably lower than the example cited above. For
example, the Fe or Mn content of plant tissues is some times nearly as high as the
content of S or Mg. The content of micronutrients is often far in excess of their
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physiological requirements. In many plant species, chloride also occurs

comparatively in high concentrations. Yet it is only needed in minute quantities in
photosynthesis. This particular example demonstrates clearly that the content of
plant nutrient in plant organs (Leaves, stems, fruits, roots) does not give any
indication of the quantity effectively needed for physiological and biochemical
process.
17.4 CLASSIFICATION OF PLANT NUTRIENTS BASED ON THEIR
BIOCHEMICAL ROLE AND PHYSIOLOGICAL FUNCTION: Essential elements are
now classified according to their biochemical role and physiological function. Based
on the biochemical behavior and physiological functions, plant nutrients may be
divided into four groups.
Nutrient
Uptake Biochemical function
elements
1st group In the form of CO2, HCO-3, Major constituents of the organic
+ 2-
C,H,O,N,S H2O, O2, NO3, NH 4, N2SO4 compounds of the plant. Essential
, SO2. The ions from the soil elements of atomic groups which are
solution, the gases from the involved in enzymatic processes.
atmosphere. Assimilation by oxidation reduction
reactions.
2nd Group In the form of phosphates, They are important in energy storage
P, B, Si boric acid or borate, silicate reactions or in maintaining structural
from the soil solution. integrity. Elements in this group are often
present in plant tissues as phosphate,
Borate and silicate esters in which the
elemental group is bound to the hydroxyl
group of an organic molecule
(i.e sugar-phosphates) (Esterification*).
The phosphate esters are involved in
energy transfer reactions.
rd
3 Group In the form of cations from Present in plant tissues as either free ions
K, Na, Mg, the soil solution except or ions bound to substances such as the
Ca, Mn, Cl chlorine pectic acids present in the plant cell wall.
Of particular importance of their roles as
enzyme cofactors and in regulation of
osmotic potentials.
4th Group In the form of ions or Present predominantly in a chelated form
Fe, Cu, Zn, chelates from the soil Incorporated in prosthetic groups. Enable
Mo solution electron transport by valency change.
*
Esterification:
Compounds formed by condensation of an acid and alcohol with elimination of water ADP + Pi = ATP
(Source : Taiz and Zeiger 2002)
*******
105

Lecture-18
NUTRIOPHYSIOLOGY
18.1 PHYSIOLOGY OF NUTRIENT UPTAKE
Mineral nutrients are found either as soluble fractions of soil solution or as adsorbed
ions on the surface of colloidal particles. Various theories proposed to explain the
mechanism of mineral salt absorption can be placed in two broad categories:
I) Passive Absorption
II) Active Absorption
ION UPTAKE IS BOTH ACTIVE AND PASSIVE:
After several decades of research on this process of ion uptake it is now
believed that the process involves both passive and active uptake mechanisms.
Whether a molecule or ion is transported actively or passively across a
membrane (casparian band, plasma membrane or tonoplast) depends on the
concentration and charge of the ion or molecule, which in combination represent the
electrochemical driving force.
Passive transport across the plasma membrane, occurs along with the electrochemical
potential. In this process Ions and molecules diffuse from areas of high to low
concentrations. It does not require the plant to expend energy.
Active transport: In contrast, to passive transport energy is required for ions diffusing
against the concentration gradient (electro chemical potential). Thus active transport
requires the cell to expend energy.
18.1.1 PASSIVE TRANSPORT MECHANISM:

A) Diffusion: Simple diffusion to membranes occurs with small, non polar molecules (i.e
O2, CO2). In this process ions or molecules move from the place of higher concentration to
lower concentration. It needs no energy.
B) Facilitated diffusion: For small polar species (i.e H2O, Ions and amino acids)
specific proteins in the membrane facilitate the diffusion down the electrochemical
gradient. This mechanism is referred to as facilitated diffusion.
a) Channel proteins: The specific proteins in the membrane form channels (channel
proteins), which can open and close, and through which ions or H2O molecules pass in
single file at very rapid rates.
A K+ and NH4+ channel also operates by the same process of facilitated diffusion. In
addition Na+ can also enter the cell by this process.
b) Transporters or Contransporters: Another mechanism involves transporters or
contransporters responsible for the transport of ions and molecules across membranes.
Transporter proteins, in contrast to channel proteins, bind only one or a few substrate
molecules at a time. After binding a molecule or ion, the transporter undergoes a
structural change specific to a specific ion or molecule. As a result the transport rate
across a membrane is slower than that associated with channel proteins.
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Three types of transporters have been identified.
 Uniporters transport one molecule (i.e. glucose, amino acids) at a time down a
concentration gradient.
 Antiproters : catalyze movement of one type of ion or molecule against its
concentration gradient. This is coupled with the movement of a different ion or
molecule in the opposite direction. Examples of antiporter transport are H+ - Na+
and H+ - Ca+2 transport into the vacuole.
 symporters catalyze movement of one type of ion or molecule against its
concentration gradient coupled to movement of a different ion or molecule down its
concentration gradient in the same direction. The high H+ concentration in the
apoplast provides the energy for symporter transport of NO3- and the other anions.
Therefore, the energy for antiporter and symporter transport originates from the
electric potential and/or chemical gradient of a secondary ion or molecule, which is
often H+.
*Figures are from reference 6
18.1.2 ACTIVE TRANSPORT MECHANISM: Larger or more-charged molecules have

great difficulty in moving across a membrane, requiring active transport mechanisms (i.e.,
sugars, amino acids, DNA, ATP, ions, phosphate, proteins, etc.) Active transport across a
selectively permeable membrane occurs through ATP-powered pumps that transport
ions against their concentration gradients. This mechanism utilizes energy released by
107

hydrolysis of ATP. The Na+ - K+ ATP pump transports K+ into the cell and Na+ out of the
cell, another example is the Ca +2 ATP pump.
Thus, it can be understood from the above discussion that the ion transport
mechanisms operate both actively and passively. For some of the ions the uptake
mechanism is active and for some others it is passive.
18.2 FUNCTIONS OF ESSENTIAL ELEMENTS

The functional roles of essential elements in plants vary greatly from element
to element.
A) Nitrogen
Nitrogen is the fourth most abundant element in plants following C, O and H.
1. N is a major structural constituent of the cell. The cytoplasm and the cell
organelles contains varying amount of nitrogen largely in combination with C,
H, O, P and S.
2. It is an essential constituent of the different types of metabolically active
compounds, like amino acids, proteins, nucleic acids, prophyrins, flavins,
enzymes and co-enzymes.
3. Being essential for the formation of protoplasm, the deficiency of N inhibits cell
enlargement
4. As much as 70 Per cent of the total leaf N may be in chloroplast. Thus the
early symptoms of N deficiency are general yellowing or chlorosis.
5. Plants contain about 1 to 3 per cent of N on dry weight basis.
B) Phosphorus
1. It is a structural component of the membrane systems of the cell and the
mitochondria.
2. It is an essential constituent of nucleoproteins, organic molecules (ATP, ADP
etc) which play an important role in the energy transfer reactions of cell
metabolism, nucleic acids, and coenzymes like NADP.
3. Phosphorus in the phytin of seeds is regarded as a reserve.
4. The unique functions of phosphate in metabolism are its formation of
pyrophosphate bonds which allow energy transfer, Uridine triphosphate (UTP)
Cytidine triphosphate (CTP) and guanosine triphosphate (GTP) are involved
in the synthesis of ribonucleic acids (RNA).
5. Being a constituent of ADP, Phosphoglyceradehyde and ribulose phosphate,
P is involved in the basic reactions of photosynthesis.
6. Phosphorus is relatively more abundant in the growing an storage organs.
C) Potassium:
1. K plays a significant role in stomatal opening and closing. The mechanism
of stomatal closure and opening depends entirely on the K flux.
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2. K+ enhances the translocation of assimilates and promotes rate of Co 2

assimilation.
3. K activates the number of enzymes involved in incorporation of amino acids
into proteins and the synthesis of peptide bonds.
4. Potassium regulates the membrane permeability and keeps the protoplasm in
proper degree of hydration.
5. Potassium is known to increase the resistance of plants to moisture stress, to
heat and to diseases caused by pathogenic fungi and other micro organisms.
6. Inadequate K restricts the formation of xylem and phloem tissue. Lignification
of the vascular bundles is generally impaired by K+ deficiency. This effect
probably makes K deficient crops more prone to lodging.
D) Calcium
1. Calcium is required for cell elongation and cell division
2. Calcium plays an essential role in biological membranes. Calcium is deposited
in the cell wall as calcium pectate. Ca deficiency obviously impairs membrane
permeability and membranes become more leaky.
3. Germination and growth of pollen as well as the growth of rhizobium root
nodules is affected under low levels of Ca2+ supply.
4. Ca appears to play a role in the inhibition of abscission and delays leaf
senescence.
5. Calcium is an essential co-factor or an activator of a number of enzymes
concerned with hydrolysis like lipase and α-amylase
6. Ca is a structural component of the chromosomes. Where in it possibly binds
the DNA to protein. Ca deficiency is known to result in chromosomal
abnormality.
7. Calcium plays an important role in neutralizing acids. Particularly citric acid,
malic acid, oxalic acid which may become injurious to plants.
E) Magnesium
1. The most well known role of magnesium is its occurrence at the centre of
the chlorophyll molecule. It is therefore essential for photosynthesis.
2. Mg is a constituent part of the chromosomes and also essential constituent
of poly ribosomes, the key organelle concerned in protein synthesis.
3. Magnesium is known to play a catalytic role as an activator of a number of
enzymes, most of which are concerned in carbohydrate metabolism,
phosphate transfer and decarboxylations. The enzyme inorganic
pyrophosphatase, as such is inactive. It becomes functional only in the
presence of Mg.
4. It is also required for the activation of the enzyme RuBP carboxylase.
5. Nitrogen metabolism is also influenced by Mg nutrition.
F)Sulphur
Plants mainly absorb S in the form of SO42-
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1. Sulphusr is a constituent of amino acids,cystine, cysteine, and Methionine.

2. The characteristic odour of cuciferous plants , onion and garlic is due to the
presence of sulphur as a constituent of volatile oils.
3. Several other biological active compounds like vitamins (Thiamine and biotin),
lipoic acid, acetyl co-enzyme A, ferredoxin and glutathione contain sulphur as
an essential part.
4. The active adenosine-5-phospho sulphate (APS) is an important sulphate
donor which is involved in the synthesis of glycosides in mustard oil. Being
involved in the activation of number of enzymes, participating in the dark
reactions of photosynthesis, sulphur is involved in carbohydrate metabolism of
the plants.
5. The total S content in plant tissues is in the order of 0.2 to 0.5 percent in the
dry matter.
*******
110

Lecture:19
NUTRIOPHYSIOLOGY
19.1 FUNCTIONS OF PLANT MICRO NUTRIENTS
A. IRON
1. Iron is a constituent of cytochromes, ferredoxin, catalase and peroxidase. The
cytochromes(cyt b6 and cyt-f) play an important role in electron transport
process of oxidative phosphorylation (respiration) and photo phosphorylation
(photosynthesis). The iron containing protein ferredoxin plays an important
role in the reduction of CO2, atmospheric nitrogen and of sulphate.
2. The leghaemoglobin present in the root nodules of leguminous crops contains
iron as an essential constituent.
3. Although iron is not a component of the chlorophyll molecule, it is essential for
its synthesis. It has some role in the synthesis of the chlorophyll precursor
protoporphyrin –IX. Most of the iron (60-80% of the iron content of the leaves)
is found in the chloroplasts.
4. The important iron containing enzymes of higher plants is succinic
dehydrogenase, cytochrome oxidase, catalase, peroxidase and aconitase.
B. MANGANESE
1. Manganese is believed to be specific activator of some enzyme like oxidases,
peroxidases, dehydrogenases and kinases.
2. Manganese is known to be a constituent of nitrite reductase and
hydroxylamine reductase, both of which are concerned in nitrogen
assimilation. In the absence of manganese, nitrite accumulates and leaves
show symptoms of nitrogen deficiency. Manganese regulates the reduction of
nitrite into hydroxylamine and subsequently into ammonia by influencing
hydroxylamine reductase.
3. Manganese plays a key role in carbohydrate metabolism and also affects the
absorption of calcium and potassium ions.
4. Manganese is involved in the oxygen evolving step in photosynthesis –of PSII
(water oxidizing enzyme complex)
C. COPPER
1. The copper containing compounds plastoquinones and plastocyanins are
involved in the electron transport from chlorophyll to NADP during the primary
reactions in photosynthesis. Thus copper stress in plants has been shown to
result in a decreased rate of photosynthesis.
2. Copper is a constituent part of several enzymes like nitrite reductase,
cytochrome oxidase and ascorbic oxidase.
111

3. Relatively, high concentrations of Cu occur in chlorophasts. About 70 percent

of the total Cu in the leaf is found in these organelles.
D) BORON
1. Boron is associated with the reproductive phase in plants and its deficiency is
often found to associate with sterility and malformation of reproductive
organs. Boron is thus required for the germination of pollen and growth of
pollen tube.
2. Boron plays an important role in carbohydrate metabolism, particularly in the
translocation of photosynthates. Boron was considered to be involved in
the translocation of sugars in the form of sugar borate complexes. These
complexes pass more rapidly through cell membranes than the free sugars.
3. Boron is required for the proper development and differentiation of vascular
elements.
E) ZINC
1. Zinc is a metal component of a number of metallo enzymes like alcohol
dehygenase and lactic dehydrogenase.
2. Zinc is essential for the synthesis of the amino acid tryptophan, a precursor
of an important plant growth hormone indole acetic acid (IAA).
3. Zinc is closely involved in N-metabolism of the plant.
4. Zinc plays a role in plant metabolism involved in starch formation. Zinc along
with Cu has been shown to be a constituent of the enzyme super oxide
dismutase.
F) MOLYBDENUM
1. Mo is an essential component of two major enzymes in plants viz.,nitrate
reductase and nitrogenase. Nitrate reductase concerned with the reduction of
nitrate to nitrite in both microorganism and higher plants.
Nitrogenase consists of two enzyme protein complexes, the bigger of which
contains Fe and Mo in a ration of about 9:1
2. By virtue of being a constituent of nitrate reductase, Mo plays direct role in
nitrogen metabolism of plants.
3. Mo is known to be a specific inhibitor of acid phosphatase.
G) CHLORINE
1. Chlorine has been shown to be involved in the oxygen evolution in photo
system II in photosynthesis ( Cl and Mn are important for this reaction)
2. It raises the cell osmotic pressure.
3. Chlorine accelerates the activation of amylase which converts starch into
soluble sugars.
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19.2 FUNCTIONS OF BENEFICIAL NUTRIENTS

a) COBALT
1. Cobalt is required by rhizobia for the fixation of elemental nitrogen both by
leguminous and non-leguminous symbionts.
2. It is a structural component of vitamin B12 (cyanocobalamine).
3. B12 is essential for the formation of hemoglobin concerned in nitrogen fixation.
b) SODIUM
1. In higher plants, sodium has so far been shown to be essential only for two
halophytic species Atriplex vasicaria and Hologeton glomeratus.
2. The best known role of sodium is in the maintenance of osmotic relations of
the cell.
3. Sodium has beneficial effect on growth and water relations of sugar beet.
c) SILICON
1. The effect of Si is especially important in the yield and quantity of the rice
crop.
2. Recent studies have shown that, Silicon imparts disease resistance and
lodging resistance in paddy
3. The grain yield of the plants with Si is twice more than the plants without Si.
4. The concentration of Si in rice will be around 100 mg g-1.
19.3 MOBILITY (PHLOEM TRANSPORT) OF INORGANIC SOLUTES
The mineral nutrients initially acquired by the roots move up ward in xylem.
Many of them are then subjected to redistribution via the phloem but a few are not.
Immobility in the phloem presumably is caused by failure of these elements to enter
the sieve tube.
Bukovac and Wittwer (1957) studied the mobility of many radio actively
labeled mineral elements applied to leaves of bean plants and classified them into
three groups based on the mobility in phloem.
Mobile Intermediate Immobile
Nitrogen Iron Calcium
Phosphorus Manganese Boron
Potassium Zinc
Magnesium Copper
Chlorine Molybdenum
Rubidium
Sodium
Sulphur
*******
113

Lecture-20
NUTRIOPHYSIOLOGY
20.1 DEFICIENCY SYMPTOMS OF PLANT NUTRIENTS
A) NITROGEN
Nitrogen being a mobile element within a plant, it’s deficiency results in
movement of N from older to younger (upper) leaves. As a result the older
leaves turn yellow in colour.
1. In young plants, growth is stunted with yellowish green leaves. Yellowing of
leaves is due to the collapse of chloroplasts resulting in decrease of
chlorophyll content.
2. Yellowing always starts from the older leaves and spreads to young ones.
3. Severe N deficiency affects the leaf tissue to become dry and necrotic.
4. Older leaves are shed prematurely.
5. Root growth is affected and branching is restricted.
6. Shoot become short, thin with up right growth and spindly appearance.
7. Flowering is reduced.
8. In cereals, tillering is poor, number of ears per unit area and number of grains
per ear head is reduced.
B) PHOSPHORUS
Generally the symptoms of P deficiency appear in the older leaves.
1. Young plants are stunted with dark blue green leaves.
2. Stems become slender
3. Root system is limited. Primary and secondary roots elongate in length with
short tertiary roots.
4. In many plant species P deficiency induces the formation of anthocyanin
pigment and the leaves acquire purple color primarily along margins and on
the lower stalk (eg. Maize)
5. The formation of fruits and seeds is depressed and ripen slowly.
6. Plant often dwarfed at maturity.
7. In potato, tubers may develop rusty lesions in the flesh.
C) POTASSIUM
1. The most characteristic symptom of K deficiency is tip and marginal
scorching of most recently matured leaves.
2. In barley which is the most susceptible of the cereals, numerous small brown
areas develop in the areas between the veins.
3. Roots are slender and poorly developed in sugar beet.
4. In certain temperate fruit trees, K deficiency may result in severe “die-back”
5. Resistance of plants to infection by bacterial and fungal pathogens is reduced.
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6. Putriscine, (a Diamine) accumulates under K+ deficiency.
D) CALCIUM
1. Symptoms of Ca deficiency appear earliest and severely in meristametic
regions and young leaves because it is immobile one.
2. Breakdown of meristemetic tissues in stems and roots occur.
3. Roots poorly developed, lack fiber and may appear gelatinous.
4. The apical bud “dies off” and thus small branches arise from lower leaf axils
giving the plant bushy appearance.
5. Little or no fruiting.
6. Calcium deficiency causes “bitter pit” disease in apple and “blossom end
rot” in tomato and watermelon.
E) MAGNESIUM
1. Interveinal yellowing or chlorosis occurs in older leaves and under severe
deficiency the areas become necrotic.
2. In cotton, Mg deficiency induces formation of anthocyanin pigments and a
reddish coloration of leaves during winter.
3. In citrus, the chlorosis commences at the tips and margins of the leaf and
spread in ward towards midrib by leaving inverted V shape at the bottom of
the leaf.
4. In general fruit trees are susceptible to Mg deficiency and show varied
patterns of chlorosis, necrosis and pigmentation of old leaves which shed pre
maturely.
F) SULPHUR
1. Chlorotic symptoms first appear in younger most recently formed leaves.
2. Shoot growth is more affected than root growth.
3. Leaves become narrow and chlorotic in cruciferae.
4. Stems often become slender.
G) IRON
1. Being relatively immobile, Fe deficiency appears first on the younger leaves of
the plant.
2. In most species chlorosis is interveinal and a fine reticulate pattern can be
observed in the newly formed leaves, the darker green veins contrasting
markedly against a lighter green or yellow background. The youngest leaves
may often be completely white and totally devoid of chlorophyll.
3. In the leaves of cereals the deficiency is shown by alternate yellow and green
strips along the length of the leaf.
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4. In fruit trees, iron deficiency is associated with high leaves of Ca Co 3 in the

soil, where it is called “lime induced iron chlorosis”
H) MANGANESE
1. Mottled chlorosis with veins green which appears first on younger leaves.
2. Stem becomes yellowish green, often hard and woody.
3. In groundnut leaves, the veins are surrounded by a definite zone of non
chlorotic interveinal tissue.
4. Characteristic symptoms of Manganese deficiency in certain crops have been
given specific names. They are grey speck of oats, speckled yellow of
sugar beet, marshy spot of peas, pahala blight of sugarcane etc.
I) BORON
1. B is immobile in most plant species and symptoms appear first at tops and
roots.
2. The youngest leaves are misshaped, wrinkled and are thicker and of a dark
bluish green color. Shed prematurely.
3. Plants dwarfed, stunted, flower development and seed production usually
impaired.
4. In fruit trees, the bark of the stem may become rough and show splitting.
Fruits often develop corky areas internally.
5. Roots exhibits swollen root tips.
6. Characteristic symptom of B deficiency in certain crops are
Heart rot in sugar beet Hollow stem in cauliflower
Brown heart of turnip Corky pith of apples
Hard fruit of citrus
J) COPPER
1. Young leaves show chlorosis
2. Terminal leaves, shoots are wilted and distorted, frequently followed by death
resulting in development of several auxillary buds.
3. Cereals show bushy growth with white twisted tips and reduced panicle
formation.
4. Poor or no heading in cabbage and lettuce.
5. In citrus, Cu deficiency leads to exanthema or die-back.
K) ZINC
1. Little leaf and rosette are the common symptoms of Zn deficiency.
2. In maize, Zn deficiency results in white or yellow emerging leaves, a condition
called white bud.
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3. Leaves chlorotic and necrotic, young growth first affected, resulting,

premature shedding.
4. In Rice Zn deficiency causes ‘khaira disease’
5. In monocots and particularly in maize, chlorotic bands form on either side
of the midrib of the leaf.
L) MOLYBDENUM
1. Symptoms of Mo deficiency are similar to those of N deficiency because in the
absence of Mo, nitrate is not metabolized.
2. Leaves show marginal scorching and rolling or cupping.
3. In extreme deficiency, the leaf lamina is not formed and only the rib is present
in cauliflower which is called whiptail disorder.
M) CHLORINE
1. Chlorosis of younger leaves and an overall wilting of the plant.
2. In some plant species, like tomato, leaves show chlorotic mottling, bronzing
and tissue necrosis.
N) SILICON
1. Withering of leaves and wilting of plants, which are called weeping willow
habit of growth.
2. In cereals and grasses, necrotic spots develop on leaves.
20.2 NUTRIENT DEFICIENCIES AND SOME SPECIFIC NAMES
Specific names have been given to symptoms of certain nutrient deficiencies. They
are:
Calcium Bitter pit in apple, Blossom end rot in tomato
Manganese Grey speck of oats, speckled yellow of sugar beat, Marsh
spot of pea pahala blight of sugarcane, Frenching of tung
tree.
Boron Heat rot in sugar beat, Browning or hollow stem in
cauliflower, Top sickness of tobacco, Brown heart of turnip,
Cracked stem of celery, Corky pith of apple, hard fruit of
citrus.
Copper Exanthema or dieback in citrus
Zinc White bud in maize, khaira disease in Rice.
Molybdenum Whiptail in cauliflower
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20.3 TOXICITY SYMPTOMS OF PLANT NUTRIENTS

As excess supply of one or more of the essential elements, particularly the
heavy metals brings about a reduction in growth and often leads to the production of
visual symptoms which at times are characteristic of the elements supplied in
excess. The visual effects of excess supply of elements may be due to a direct effect
of toxicity of elements or by inducing deficiency of another element. The excess
supply of particulars elements reduces the uptake and utilization of other essential
elements. Eg. Excess phosphorus supply may retard the uptake and translocation of
Cu and Zn, thus shows Cu and Zn deficiency symptoms.
Under field conditions N, S and Al toxicities are common. Mn toxicity is
observed in acid soils.
A) NITROGEN
1. As a Nutrient – excessive nitrogen commonly produces plants that vegetative
so that yield is reduced. In sugarcane both juice quality and sugar recovery is
reduced. Fruit quality is impaired in organs and peach. Cereals receiving
excess N have been shown to exhibit lodging as a result of the decreased
thickness of the cell walls or due to the poor development of the mechanical
tissues of the stem or due to both. In tomato, plants are highly vegetative and
fruit production is reduced.
2. As a salt a high level of N application increases the salinity of the soil. Also the
form of nitrogen (NH4+ or NO3--) and associated ions (SO4-, NA+ Ca++) may
markedly affect soil structure and plant response.
B) SULFUR
Excess sulfur causes interveinal yellowing in sorghum and lemon leaves. No
symptoms of S toxicity is observed in leaves of sugar beet, tomato, cotton and
alfalfa, but growth and leaf size was considerably reduced in all the crops
Excessive concentrations of sulfur dioxide in the atmosphere also cause
toxicity in the plants. The symptoms produced by excessive So2 was divided into (a)
acute injury (b) Chlorotic injury.
The acute injury consists of collapsed marginal or interveinal areas which at
first have a dull, water soaked appearance, later drying and bleaching to an ivory
color in most species. These lesions are caused by rather sudden absorption of
enough gas to kill the tissue.
Chlorotic injury is a yellowing of the leaf which may progress slowly and
bleaches completely. bleaching most of the chlorophyll and carotenoids . This is due
to absorption of an amount of gas somewhat insufficient to cause acute injury or it
may be caused by absorption over a long period of time of sublethal amounts of gas.
Another type of S toxicity is Hydrogen sulfide toxicity (sulfide injury). In
poorly drained rice soils, the H2S toxicity is common. This disorder is commonly
called as “akagare” or “akiochi”. This anaerobic disorder is characterized by
inhibited root development and browning (or blackened) and death of roots which
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precede the stunting of shoots. Field observation shows that the roots of affected
plants are blackened from the accumulation of iron sulfides (and possibly
manganese sulfides) formed under reducing conditions. This black coloring
disappears upon exposure to the atmosphere after several hours due to oxidation, a
confirmation of sulfide formation. The H2S toxicity also depresses the uptake and
translocation of P and other nutrients.
C) ALUMINUM
Aluminum toxicity is wide spread in most of the acidic soils. In general, roots
are affected first, more severely than the tops under aluminum excess. In most
species, roots appear thick and brown and the tips of the root lets often appear
enlarged. In barely, the general growth of the plants and tillering are severely
restricted. Leaf tips often turn brown and wither. The stem becomes dark brown, no
ears are formed and plants often die prematurely. The root growth is severely
restricted and the root tips appear dark brown and enlarged.
*******
119

Lecture-21
NUTRIOPHYSIOLOGY
21.1 FOLIAR NUTRITION
Foliar nutrition of mineral nutrients by means of sprays offers methods of
supplying nutrients to higher plants more rapidly than methods involving root
application. Foliar nutrition consists of spraying dilute nutrient solutions on foliage in
order to feed the plants with required nutrients in short time.
21.1.1 Mechanism of uptake: In terrestrial plants the uptake of solutes by the
surfaces of leaves and other aerial parts is severely restricted by the external wall of
the epidermal cells. This wall is covered by a layer of wax and cutin and contains
pectin, hemi cellulose and cellulose. The direct penetration of solutes from the leaf
surface through open stomata into the leaf tissue is unlikely because a cuticular layer
also covers the surface of guard cells in stomatal cavities. Further more ion uptake
rates from foliar sprays are usually higher at night, when the stomata are closed,
than during the days, when the stomata are open.
The movement of solutes across the cuticular layer takes place in cavities
or channels called ectodesmata by process of diffusion. Ectodesmata extend
through the outer epidermal cell walls from the inner surface of the cuticle to the
plasma membrane. When a substance reaches the plasma membrane of an
epidermal cell it will be absorbed by mechanisms similar to those which operate in
root cells.
21.1.2 Advantage of foliar nutrition: The advantage of foliar nutrition is high
recovery rate while the limitation is the smaller amounts of nutrients that can be
supplied at a given time. It is very essential not to excess the specified concentration
of fertilizer solutions, for the foliar spray to avoid the risk of leaf scorch. Fertilizers
when used in excess concentration damage the leaf tissue causing necrosis.
The technique has great practical utility under certain conditions.
a) Correction of visual symptoms: Nutrient deficiencies observed during early
stages of crop growth can be corrected by foliar sprays.
b) Low nutrient availability in soils: in calcareous soils, for example, iron
availability is very low and iron deficiency (Lime induced iron chlorosis) is
widespread. Foliar spray is much more efficient than the soil application of
expensive iron chelates.
c) Decrease in root activity during reproductive stage: As a result of sink
competition for carbohydrates, root activity and thus nutrient uptake by the
roots decline with the onset of the reproductive stage. Foliar sprays containing
nutrients can compensate for this decline.
Usually micronutrients, which are required in low rates, are foliar sprayed. Urea
sprays are used for supplying nitrogen. Ammonium phosphate fertilizer is also
used to spray to improve seed setting in multicut fodder legumes like Berseem.
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21.2 HYDROPONICS
The method of growing plants in aqueous nutrient solutions is known as
hydroponics culture or hydroponics. The growing of higher plants with their roots in
dilute solutions of mineral salts instead of in soils has led to a vastly increased
understanding of plant nutrition. The use of water culture technique for growing
plants permits precise control of the supply of nutrient ions in the root environment.
The first recorded use of water culture technique was in the year 1699 by John
Woodward. Latter the process of growing plants in water culute was named as
hydroponics by Gericke 1930.
The water culture technique is extensively used to grow certain high value crops
(eg.Tomato, lettuce and strawberry) in glass houses during off seasons in
metropolitan areas of developed countries. In addition, vegetables are grown in
plastic houses in a few coastal desert regions, where sea water is desalinated and
supplied to roots in precisely the amounts required by the growing plants.
21.2.1 TYPES OF SOLUTIONS CULTURE:

The three types of solution culture are
1. Static 2. Flowing 3. Mist system
In static system, the plant is supported in such a way that its roots are
immersed in culture solutions containing nutrient elements. The solution is kept
oxygenated by bubbling air through it from a compressed air line. This system is
mostly used for experimental purpose. In flowing systems, the solution move
continuously allowing its composition to remain relatively constant at the root
surface. The nutrient film technique, where roots grow in films of solution constantly
moving down inclined troughs is a commercial version of the flowing system.
In mist system, roots hang in the air and are sprayed intermittently with a
nutrient solution.
21.2.2 SAND CULTURE:

In a modification of the solution culture technique, plants are grown with their
roots anchored in a solid inert aggregate, such as sand culture technique. The added
advantage of this solid medium culture is that the roots grow in a natural medium and
no other means of support needs to be provided.
Most important form of solid culture techniques is sand culture. Sand culture is
sometimes referred to as slop culture, drip culture, or intermittent renewal. Plant
nutrients are supplied with solutions, as in water culture. The major difference is that
the plants are grown in silica sand or other inert material (perlite, vermiculite, peat
etc.). The solution is applied to the sand and then allowed to drain off. In this way
plant nutrients are supplied, aeration is provided, and the roots of the plant are
supported. Additional support may be required for the aerial portions of the plant,
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especially tomatoes and cucumbers. Sand should be medium to coarse in size (.0.25
mm-2mm).
Perhaps the most limiting aspect of solid media is that access to roots is achieved
only with major damage.
21.2.2.1 Types of solid medium culture
Based on the addition of nutrient solutions to the solid culture the following
three systems are categorized.
1. Slop culture:- Nutrient solutions are poured over the surface of the solid
medium.
2. Drip culture:-Nutrient solutions are dripped on the surface.
3. Sub irrigation:- Nutrient solutions are forced from the bottom of the container
to come up to the surface of the solid medium.
Methodology
Stock solutions are prepared separately from inorganic salts containing the
essential elements for normal plant growth. Hoagland and Arnon (1950) formulated
two nutrient solutions which have been very widely used and the term “Hoagland
solution” has become a household word in laboratories devoted to plant nutrition all
over the world. Hoagland solution II contains ammonium ions as well as nitrate and
as a result it is a better buffered one. pH of the nutrient solutions will be in the range
of 5 to 7.The final nutrient solution is prepared by adding the correct proportion of
stalk solutions and diluted with deionised water. The containers are filled with
nutrient solution and covered with aluminum foil or thick non transparent paper in
order to eliminate light incidence and algal growth in containers.
Week or ten days old seedling are taken, the roots are submerged in the
nutrient solution and the stem projected through an opening cut in the container lid.
To ensure the stem to be held rigid, the opening is stuffed with some inert padding
material such as cotton. The nutrient solution is well aerated periodically by
employing an air-pump and change once in few days depending on undesirable pH
changes and the growth rate of the plants.
For commercial cultivation, large shallow tanks (nutrient film technique) are
used. The nutrient solutions are replenished often enough to maintain their
concentrations. Nutrient solutions are made to flow over the plant roots from a
recirculating tank .
Use in Agriculture
1. To grow plants in the laboratory for experimental purpose such as studies
involving nutrio physiology, herbicide physiology, radio tracer technique for ion
uptake and movement etc.
2. Commercial cultivation of high value crops in glass houses during off seasons
in metropolitan areas.
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3. Deficiency and toxic symptoms of mineral elements can be developed and

studied with different crop plants.
*******
123

Lecture-22
PHOTOPERIODISM AND FLOWERING
Light controls many physiological processes in the plant among them the
process of flowering is important even in the life cycle of the plant. In case of
annuals, floral induction signals the end of vegetative growth. In perennials, floral
initiation is not accompanied by such changes and the vegetative and
reproduction growth occur simultaneously. Reproductive growth is a complex
process which includes anatomical, morphological, physiological and biochemical
process. All these changes are finally manifested into transformation of
vegetative to reproductive bud.
Before floral primordial can be initiated, the plant must complete a period
of vegetative growth of attain some minimal leaf number. When this condition is
attained the plant is said to be ripe-to-flower. In most plants, ripeness to-flower
is attained after the plant has produced several leaves. In some annual grasses a
minimum of 7 leaves must be developed before the plant is ripe to flower. On the
other hand, a plant such as pharabtis nil is ripe to flower within a day after the
cotyledons have emerged. The cotyledons presumably contain enough stored
food to support subsequent reproductive development.
However, most of the commonly cultivated plants and weedy

annuals attain the ripe-to-flower condition 2-3 weeks after seedling emergence
when a few leaves have fully developed.
22.1IMPORTANCE OF PHOTOPERIODISM IN AGRICULTURE:
The knowledge of photoperiodism and its application has great economical potential
as follows
 The yield of tubers, corns, bulbs and rhizomes can be increased substantially
by increasing or decreasing the duration of day or night.
 Annuals may be grown twice or thrice in a year.
 Perennials might flower throughout the year.
 Hybridization experiments have got a fillip because different varieties growing
in different areas with different durations and flowering at different times are
made to grow and flower side by side by artificially controlling their
photoperiods.
 Understanding the concept of photoperiodism has helped to choose photo
insensitive varieties and cultivars. These cultivars are best suited in intensive
agriculture.
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22.2 CLASSIFICATION OF PLANTS BASED ON PHOTOPERIODIC RESPONSES
Garner and Allard (1920) observed that flowering in Maryland mammoth

variety of tobacco (Nicotiana tabaacum) and Biloxi soybean (Glycine max) can be
controlled by artificially increasing or decreasing the natural duration of sunlight.
They introduced the term photoperiodsm and defined that as the response of
plant to the relative length of day and night. However, the duration of dark
period is also much important to understand the concept.
Garner and Allard classified the plants into 5 types based as their
photoperiod response of flowering
A. Short day plants (SDP) B. Long day plant (LDP) C. Day neutral plant (DNP)
D. Short –long- day plants (SLDP) E. Long- short- day plants (LSDP)
A)Short day plants (SDP)
Plants that readily flower when the day length is less than its critical day
length (photoperiod required to induce flowering is critical day length). For
example, critical day length for xanthium (SDP) is about 15.5 h and it flowers only
when day length is 15.5 h or less It was also shown that even a brief exposure to
light during dark period inhibits flowering and therefore it requires a continuous
and uninterrupted dark period for its flowering. On the other hand, a brief period
of darkness during day time had no effect on flowering. Because of the
importance of dark period in flowering, these are also called as Long Night
Plants.
B)Long day plants (LDP)
These plants flowers readily only when the photoperiods are longer than the
critical photoperiod. For example annual Hyoscyamus has a critical photoperiod
less than 10h.30m. As a result, this plant does not flower with a photoperiod less
than 10h.30m but it promptly flower under any length of photoperiod above this
limit and even within 24h of light. These plants are also called as short night
plants.
C)Indeterminate or Day Neutral Plants
Indeterminate or DNP flowers readily over a wide range of day length from
relatively SDL to continuous illumination.
D) SLDP
These plants require first short photoperiod and then long photoperiod for
flowering.
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E) LSDP
These plants require first long days and then short photoperiod for flowering
These plants do not flower when exposed to either SD or LD and need both
LD and SD for its flowering
22.2.1 EXAMPLES OF PLANTS WITH DIFFERENT PHOTOPERIOD

REQUIREMENT ARE GIVEN BELOW
1. Short day plants
A. Qualitative SDP (Plants specifically requiring SD)
1. Chrysanthemum 2. Tobacco 3.Xanthium 4. Rice (Oryza sativa)
5.Japanese morning glory (Pharbatis nil)
B. Quantitative SDP (Plants promoted by SD)
1. Cosmos 2. Cotton 3. Hemp
2. Long day Plants
A. Qualitative LDP (plant specifically requiring LD)
1.Barley (winter barley) 2.Wheat 3.Henbane (Hyoscyamus Niger)
4.Spinach 5.Sugar beet
B. Quantitative LDP (plants promoted by LD)
1.Lettuce 2.Blue grass 3.Clover
3. Day neutral plants
1.Cucumber 2.Balsam 3.maize 4.Tomato
4. LDSDP
1.Bryophllum 2.Cestrum Nocturnum (Night blooming Jasmine)
5. SDLDP
1.Winter rye 2. Candy tuft
22.3 PERCEPTION OF PHOTOPERIODIC STIMULUS
Knott (1934) first observe that photoperiodic induction is perceived by young

expanded leaves. Later Knott’s work was confirmed by Chailakhyan (1936) he
divided chrysanthemum plants into four groups and varied the regime of light
126

and dark cycles in all the four groups by using light proof cases. The four groups
of plants are as follows:
Group A Entire plant continuously received long day treatment
Group B Lower leaf portion received short day treatment while upper
defoliated portion received long day treatment
Group C Lower leafy portion received long day treatment while upper
defoliated portion received short day treatment
Group D Entire plant continuously received short day treatment
He observed that flowering occurred in those plants where the leaves

received short day treatment (Group B&D) but it failed in those plants where the
leaves received long day treatment (Group A&C). On the basis of above
observation he concluded that short day stimulus is perceived by the leaves.
Floral stimulus
The flowering stimulus produced in the photo induced leaves is translocated

to the shoot apices for flower evocation. It has been observed that floral
stimulus is similar in long and short day plants by grafting experiments. If a leaf
from a photo induced plant is removed and then grafted on a non induced plant,
then this plant flowers. Apparently a chemical substance is produced during
photo inductive cycle, which is transmitted during grafting to non induced plants
and evokes flowering. Chailakhyan (1936) named this flower inducing chemical
substance as Florigen. The flowering stimulus travels through phloem but
independent to the transport of photosynthates.
22.4 Biological clocks:
Study of Biological clocks is also known as chronobiology. The best understood

biological clock is the circadian rhythm.
The various physiological processes do not occur at a constant rate during the
different times of the day and night. The rate fluctuates showing peaks and dips at
regular intervals. In other words the occurrence of the process is rhythmic. At first
glance it may appear that such rhythms may be because of different external factors
like temperature, light and humidity which fluctuate during the different times of the
24 hrs day. However, the plants continue to exhibit such rhythms even when held at
constant conditions. It means that the occurrence of such rhythms is not because of
variations in the external conditions (not exogenous) but appears to be controlled by
the plant itself (endogenous). They are called as endogenous rhythms.
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Most of the endogenous rhythms that occur in plants have a periodicity of

frequency of about 24 hrs. That is, this process is repeated once in about 24 hrs and
called circadian rhythms. Some properties of these diurnal rhythms are
demonstrated by the sleep movement of the primary leaves of bean (Phaseolus
multiflorus).(Fig.)
During the day the leaves are horizontal, while during the night the leaves
assume a more vertical position. The plant is exposed to normal light and dark period
for 2 days and was subsequently kept under light. The primary leaves continue to
open and close once in about 24 hrs even continuous light.
The other examples of circadian movement in higher plants are cell mitosis,
respiration, CO2 fixation in Bryophyllum, enzymes activity, nectar odor and flower
petal movement.
Experiments conducted by De Mairan, Pfeffer, Bunning and Stern and

others have clearly shown the existence of an internal (endogenous) time
measurement system in plants. The biological clock, as it is called, is believed to be
very accurate as it measures the rhythms of a small period as one minute to as large
a period as one year. Though the nature of the biological clock or the endogenous
factor controlling it is not known, it is suggested to be located within the cell
apparatus of even unicellular organisms.
*******
128

Lecture-23
PHOTOPERIODISM AND FLOWERING
23.1 PHYTOCHOROME
Phyrochrome is a blue proteinaceous pigment composed of two components:

a protein and a chromophore - which give its light absorbing properties.
Borthwick and Hendricks (1946) suggested that phytochrome exists in two
forms: a red light absorbing form (pr), blue green in color and a far red light
absorbing from (pfr), light green in color. These two forms are inter convertible
and it is located within plasma membrane. Phytochrome is produced in light
grown tissues:
P660 P730
It shows on absorbing red light, Pr form is converted to pfr during day time,
while pfr form gradually converted into pr form in dark on absorbing far red light.
23.1.1 Role of phytochrome in short day plants
At the end of light period the phytochrome is predominantly present in the pfr
form and the ratio of pfr to pr is such that, the formation of the flowering stimulus
is prevented. During the long dark period, spontaneous conversion of pfr to pr
takes place or pfr is destroyed, and the ratio of pfr to pr finally drops to a level
where metabolic processes are triggered which leads to the formation of
flowering stimulus. This duration represents the critical dark period requirement
for the flowering of SDP. It varies from plant to plant. If the dark period is
interrupted by flash of red light, pr is converted to pfr. And the pfr to pr level is
such that the formation of the flower stimulus is prevented.
23.1.2 Role of phytochrome in long day plants
Long day plants require a high ratio of pfr to pr for the formation of flowering
stimulus. Such a high ratio of pfr to pr is attained at the end of a long day. If the
night is too long, pfr reverts to pr or is destroyed and the flowering stimulus is
prevented from being formed. When the night is interrupted by a flash of red
light, pr is converted to pfr, there by raising the ratio of pfr to pr to a level which
allows the flower stimulus to form.
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23.2 FLOWERING HORMONES:
Gibberlic acid (GA) seems to play an important role in flowering. It may be

directly involved in the formation of florigen.
GA’s can substitute for long day requirements and also the cold treatment
requirement in several species for its flowering. Auxins are also known to induce
flowering in short day pine apple and they are used commercially for this
purpose. They have also been found to be effective on long day plant like winter
barley and Hyoscyamus Niger. Exogenous application of cytokinins is also
known to induce flowering in many species, under non-inductive photoperiods
including chrysanthemum and Phorbitis nil.
23.3 VERNALIZATION AND FLOWERING
It has been noticed that certain plants in addition to an appropriate

photoperiod, require low temperature treatment during their early stages of life period
for subsequent flowering in the later stages. It was found that winter wheat which is
usually sown in winter and flowers in summer can be converted into spring wheat
which is sown in spring and flower in summer, if slightly germinated seed are given
artificial cold treatment. This conversion of winter variety of wheat into spring variety
by low temperature or chilling treatment was termed as Vernalization by Lysenki
(1928). Due to low temperature treatment the period of vegetative growth of the plant
is reduced resulting into early flowering.
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Chourad (1960) defined vernalization as acquisition or acceleration of the

ability to flower by a chilling treatment. The optimum temperature for vernalisation is
0 to 50C.
The phenomenon of vernalization was studied in several winter annuals Eg.

Petkus winter rye (Secale cereale), some biennials Eg. Henbane (Hyoscyamus
niger) and also in certain perennials Eg. Apples (Pyrus malus). All the above plants
require a low temperature treatment for subsequent flowering. However certain
plants such as Petkus winter rye, do not have an absolute cold requirement for
flowering. In such cases, the cold temperature only shortens the time of flowering,
where as in biennial strain of Henbane the vernalization treatment is absolute and
they cannot flower without vernalization.
Perception of Cold Stimulus:
Apical meristem, embryos and actively dividing cells are the potential sites of
vernalization. The common feature for vernalization is the presence of actively
dividing apical meristems. Hence, vernalization treatment requires.
1. Actively dividing apical meristems.

2. Respiration substances such as carbohydrate.
3. Presence of oxygen.
4. Water (40 to 80%)
Presence of Floral Hormone:
The German botanist G. Melcher’s (1939) suggested that low temperature

induces the formation of a hormone, vernalin a hypothetical substance and has not
been isolated so far. The cold stimulus can be transmitted from one plant to another
by a graft union Eg. Henbane. Here, if a vernalized plant is grafted to an
unvernalized plant the later also shows flowering.
Mechanism of vernalization: Hormonal Theory.
Lang and Melcher’s (1947) suggested that a precursor A is converted into a

thermolabile compound B during cold treatment. Under normal conditions B changes
into a stable product (C Vernalin) which causes flowering. But at higher temperature
B is converted into D and flowering does not take place due to devernalization.
(fig.3). further experiments have shown that vernalin acts in association with the
flower inducing stimulus, florigen in order to induce flowering.
A B C Flowering
Precursor Thermolabile Vernalin
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Devernalization:
The effect of low temperature treatment for flowering is nullified if the plants
are immediately given high temperature treatment. This phenomenon is called
devernalization. The degree of devernalization decreases if the duration of the cold
treatment has been longer. However, the devernalized plant can be vernalized again
by subsequent cold treatment.
Vernalization and gibberellins:
Lang et al (1952) demonstrated that application of GA’s can replace the low
temperature requirement for vernalization in many plants. Eg. Henbane. This plant
remains vegetative and retains its rosette habit during first growing season after
passing through the winter period flowers in the next season. The GA’s cause such
plant to flower even during the first year.
23.4 IMPORTANCE OF VERNALIZATION:
1. Vernalization increases cold resistance in plants.

2. It reduces the vegetative period of development of plants and induces early
flowering.
3. Winter varieties of crop plants can be converted into spring varieties by
vernalisation.
*******
132

Lecture: 24
PLANT GROWTH REGULATORS
Growth of the plant has for long been believed to be due to the minerals
absorbed from the soil and the food materials synthesized by the plant. It is now
however recognized that the growth of the plant is very much regulated by certain
chemical substances known as growth regulators. These substances are formed in
one tissue or organ of the plant and are then transported to other sires where they
produce specific effects on growth and development.
Philips (1971) defined growth hormones as a substance which is synthesized

and is transported to other cells where in extremely small quantities influences
development processes.
The plants are known to produce 5 classes of hormones namely auxins,

gibberellins, cytokinins, abscisic acid and ethylene. A plant tissue may contain more
than one of these growth regulators at the same time. The leaf primordial and
developing seeds contain both auxin and gibberellins and in some plants ABA also.
Both auxins and gibberellins cause stem elongation by different mechanisms while
ABA and ethylene inhibits stem growth.Thus, two or more growth regulators may be
similar in their action. When the effect is more than the sum of their individual effects
it is called synergistic and when the action of two growth regulators is opposite it is
called antagonistic. The final growth and development of the plant is the sum of total
interactions of different growth regulators that are present in the plant.
24.1 AUXINS:
The idea of the existence of auxins in plants was for the first time conceived by
Charles Darwin in 1881. He showed that coleoptile of canary grass could bend
towards light when it is unilaterally illuminated. However the coleoptile failed to bend
when its tip was covered with an opaque cap. Most of the knowledge about auxins
comes from the work on oat (Avena sativa) coleoptile. Went (1926) demonstrated
that coleoptile tips contain a substance capable of elongation of decapitated
coleoptiles. He placed several freshly cut coleoptile tips on an agar block which was
kept on a piece of inert material like glass. After several hours he cut the agar block
into small cubes. He placed the agar cubes accentrically on decapitated coleoptile
stumps for 2 hours in the dark. The effect of agar cube was similar to that of the tip
as was shown by curvature of the coleoptiles( Avena coliaptile curvature test)
The first higher plant from which auxin could be extracted was maize kernels.
It was identified as IAA. Indole Acetic Acid is the major auxin occurring in plants.
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24.1.1OCCURRENCE:
All the parts of the plant body produce auxin. However the major sites of auxin
production are the shoot tip, developing seeds and buds. The amount of auxin
present in different parts of the plant varies greatly. The amount is highest in the
stem tip and coleoptile tip and decreased gradually down words.
Synthetic auxins:
There are a number of synthetic chemicals which are similar to IAA in their
biological activity. However they do not occur in any plant. The important synthetic
auxins are IBA (Indole Buteric Acid), NAA (Naphthalene Acetic Acid), 2, 4-D (2, 4
Dichloro Phenoxy Acetic Acid) and 2,4,5-T (2,4,5-Trichloro Phenoxy Acetic Acid).
24.1.2 BIOSYNTHESIS OF AUXINS:
Indole acetic acid (IAA) is synthesized from an amino acid Tryptophan in 3

different pathways. These 3 pathways have different intermediate compounds and
the pathway is named after the intermediate compound produced as follows
(a) Indole Pyruvic Acid Pathway (b) Tryptamine Pathway and (c) Indole Acetol
Doxime Pathway.
Indole Pyruvic Acid Pathway:
The first reaction involves transmission of Tryptophan to Indole Pyruvic Acid.

The enzyme tryptophan amino transferase transfers amino groups (NH2) from
Tryptophan resulting in the formation of Indole Pyruvic Acid.
In second step Indole pyruvic acid is decarboxylated to form Indole

acetaldehyde. The enzyme involved is Indole pyruvic decarboxylase. A molecule of
CO2 is removed. In the final step Indole acetaldehyde is oxidized to IAA by 2
enzymes namely Indole acetaldehyde dehydrogenase and Indole acedaldehyde
oxidase.
Tryptamine Pathway:
This pathway is of rare occurrence and was shown in tomato. Here tryptophan
is first decarboxylated by the enzyme tryptophan decarboxylase forming tryptamine.
Tryptamine then undergoes deamination forming indole acetaldehyde. The enzyme
responsible for this reaction is amine oxidase. In the final step indole acetaldehyde is
oxidized to IAA.
Indole Acetaldoxime Pathway:
This pathway is characteristic of the members of the family Brassicaceae. In

this pathway tryptophan is first converted to indole acetal doxime which in turn is
134

further converted to indole aceto nitrile directly or through an intermediary compound

glucobrassicin. Indole aceto nitrile is converted to indole acetic acid. Zinc is essential
for biosynthesis of auxin since it activates the enzyme tryptophan synthatese.
BIOSYNTHESIS OF AUXINS:
TRYPTOPHAN
Deamination
Tryptophane decarboxylase
Tryptophane transaminase NH3 CO2
INDOLE 3 PYRUVIC ACID TRYPTAMINE
Indole pyruviate decarboxylase CO2 NH3 Amine Oxidase
Indole 3 acetaldehyde
Indole acetaldehyde dehydrogenase
Indole acetaldehyde oxidase
INDOLE 3 ACETIC ACID
II. Indole Acetal Doxime Pathway:
Tryptophan
Indole acetal doxime
Glucobrassicin
Indole acetonitrile
Indole acetic acid.
TRANSPORT:
Auxin moves towards morphological basal end in stem cuttings. The

movement of auxin is polar and basipetal. In roots it is polar but acropetal. In xylem it
moves along the transpiration stream.
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24.1.3 MODE OF ACTION:
It was observed that whenever auxin causes elongation of the coleoptile, the
external medium in which the coleoptile was flooded, becomes acidic. The pH which
was originally near neutral becomes decreased to 4.5. On this basis Hager, Menzel
and Krans (1971) proposed acid growth hypothesis of auxin action. In this it was
explained that the H+ ions decreases PH value and presumed to break the acid
labile bonds or activate wall hydrolyzing enzymes and render the cell was soft. This
will create suitable conditions for cell elongation.
Auxin and cell elongation:
The primary physiological effect of auxin is to promote the elongation of cells

which may be due to increasing osmotic pressure and permeability of cytoplasm to
water and decreasing cell wall pressure.
Auxin stimulates the production of hydrolyzing enzymes like B-1, 3-gluconase,

pectin methyl esterase and cellulase which soften cell wall and increase the plasticity
resulting in reduction of wall pressure and cell elongation. Under the influence of
auxin, cellulose synthetase increases and new wall material is synthesized within the
cell wall resulting in extension or growth of the cell.
24.1.4 PHYSIOLOGICAL ROLE OF AUXINS:
A. Cell division:
Auxin has been found to be responsible for initiating and promoting cell
division in certain tissues eg. Cambium. When ever wound is caused in the plant a
swelling called callus is developed because of the proliferation of the parenchyma
cells stimulated by auxin and a chemical substance traumatic acid. This can be put
to practical use in grafting where the callus plays an important role in strengthening
the union between stock and sion. Hence, during grafting of grapes, it was found that
immersion of stock and sion in 0.1 percent of IAA resulted in quick growth of callus
and success of graft union.
B. Root initiation:
The stem cuttings of some plants readily form adventitious roots when put in
the soil. Adventitious root formation will takes place at the basal end of stem cuttings.
Cuttings of such plants which do not readily root, form abundant roots when treated
with auxins. Synthetic auxins like IBA, NAA are particularly very effective. In general
cuttings of herbaceous plants readily respond to auxins while those of woody
perennials like eucalyptus, mango and others fail to respond to auxin application.
In air layering of guava auxins like IBA at 500 ppm is used for root initiation.
136

C. Apical dominance:
The presence of apical bud causes a complete or partial inhibition of lateral

buds. This is due to the presence of higher concentration of auxin at the apical bud,
which causes a preferential movement of nutrients towards it.
In plants like sunflower, the main stem continues to grow and the lateral buds
in the axils of leaves do not emerge and grow into branches. However when the tip
of the main stem is cutoff or when it terminates into an inflorescence, the lateral buds
emerge into branches. In either case the influence of the shoot apex on the lateral
bud is lost. This phenomenon of inhibition of laterals by the shoot apex is termed as
apical dominance.
In plants like potato and tomato apical dominance is weak consequently the
apical growing point of the main stem fails to suppress the emergence of lateral
buds. Such plants are therefore extensively branched and bushy.
D. Inhibition of abscission layer:
Abscission is a process of dissolution of the middle lamella and primary walls

of the cells at the base of the petiole, pedicle or peduncle. Abscission refers to
detachment of plant organs. It is a balance between the inhibition of auxin and
promotion of substances like ABA, Ethylene etc several auxins (2,4-D, IAA, NAA)
inhibits the abscission of both leaves and fruits.
Ex: application of NAA at 20 to 30 ppm twice at 15 days interval reduces flower and
fruit drop in chillies and cotton, 2,4-D prevents defoliation in cabbage and cauliflower
that often occurs during harvest.
E. Flower initiation.
Application of auxin inhibits flowering in several photoperiodically sensitive

plants such as xanthium, soybean and others. But the exception is pineapple
(Ananas comosus) a day neutral plant. This plant can be made to bloom promptly
with the application of NAA or 2,4-D. However in this plant the effect of auxin is not
direct but is mediated through ethylene formation.
Application of auxin also alters the sex ratio of flowers. In monoecious

cucurbits increase the number of female flowers but the decrease the number of
male flowers. Similarly in dioecious plants like cannabis, the male plants start
producing female flowers when auxin is applied. Here again the effect of auxin is not
shown to be direct but is mediated through ethylene formation.
137

F. Production of parthenocarpic fruits:
It is a general observation that in the absence of pollination and fertilization

the ovary of the flower does not develop into the fruit, but the flower abscises and
falls. However application of auxin causes development of ovary into the fruit in
several plants such as tomato, brinjal and others. Such fruits are seedless as these
have developed without the normal process of fertilization these are known as
parthenocarpic fruits. The presence of large number of seeds in a fruit lowers its
commercial value in the canning industry.
G. Eradication of weeds:
Plant roots are extremely sensitive to auxins. Very high concentration of

auxins over stimulates growth promoting activities of root cells resulting in distorted
roots with blocked sieve tubes. The roots ultimately decay and the plant is killed. 2,4-
D and 2,4,5-T are effective weedicides at higher concentration of 1 to 3 percent . 2,4-
D is selective weed killer. It is highly toxic to broad leaved plants or dicotyledons
while relatively non toxic to narrow leaved plants or monocot.
H. Growth in thickness:
The stem of dicots and gymnosperms not increase in length but also in
thickness. Increase in thickness of the stem and root is due to radial growth. The
process is termed as secondary growth.
I. Vascular differentiation:
Not only the activation of cambial rings but also the differentiation of cambial
derivatives into xylem and phloem is also under the control of hormones. Interaction
of both auxin and GA is involved in this.
J. Prevention of lodging:
Naphthyl acetamide when applied on the base of oats and flax, they grow stiff,
woody and erect. Thus lodging in these crop plants is prevented.
*******
138

Lecture:25
GIBBERELLINS
The discovery of Gibberellins was quite accidental. Japanese worker

Kurosawa (1926) in Japan while conducting experiments on rice disease caused by
Gibberella fujikuroi (causal organism for foolish seedling of rice or bakane disease)
observed that the fungus caused excessive growth in rice. He applied the fungal
extracts to intact healthy plants and observed enhanced growth. Later Yabuta and
Sumuki (1938) named the active principle as gibberellin. Further it was purified,
crystallized and named as gibberellic acid (Curtis and Cross 1954). Now gibberellins
are designated as GA1, GA2 and so on. The common gibberellic acid is GA3. At
present 112 types of gibberellins are known.
25.1 Occurrence and Site of synthesis:
Gibberellins are synthesized in the young leaves (major site), shoot tip, root
tip and the developing seeds.
25.2 Bio synthesis:
3 M. Acetylyl CoA
Mavalonate
+ATP+Mn+Mg
4M. Isopentenyl pyrophosphate
Trans geranyl pyrophosphate.
Tran’s farnesyl pyrophosphate.
Geranyl geranyl pyrophosphate (GGPP)
Copalyl pyrophosphate
(kaurene synthatase)
Kaurene
Oxidation
GA12 aldehyde
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Acetyl co A is the precursor for the biosynthesis of gibberellins. Three

molecules of acetyl co A are linked together to form a molecule of mevalonic acid.
Mevalonic acid in turn is activated in the presence of ATP, Mn and Mg and is
converted to isopentenyl phosphate (IPP). This is a 5 carbon compound. Four
molecules of IPP undergo stepwise condensation, first to Trans geranayl
pyrophosphate (GPP) then to trans farnesyl pyrophosphate (FPP) and finally to form
a diterpene called geranyl geranyl pyrophosphate (GGPP). This is 20 carbon
compound. This GGPP is converted to kaurene. The conversion of GGPP to kaurene
is carried out by the enzyme kaurene synthetase in two steps.
First GGPP is converted to copalyl pyrophosphate. In the second step copalyl

pyrophosphate in turn is converted to kaurene. Kaurene undergoes oxidation and a
series of reactions resulting in the formation of gibberellins.
Transport
Transport of gibberellins is passive and non polar. Gibberellins move both in

xylem and phloem and vice versa through vascular ray cells.
25.3 Mode of action:
There are several hypotheses to explain the mechanism of GA in the plants.
1. Increase in the endogenous auxin content:
Whenever GA causes cell enlargement, the effect is not considered to be

direct. The effect is indirectly mediated through formation of auxin, in turns is held
responsible for the cell elongation.
Gibberellin has been shown to cause synthesis of amylase in barley

aleurone cells. This enzyme converts starch to reducing sugars resulting in an
increase of osmotic pressure, causing entry of water into the cells and cell
enlargement.
25.4 PHYSIOLOGICAL EFFECTS:
A. Stimulation of stem growth:
The most important effect of GA is the stem elongation when GA is applied

the stem elongates markedly. As a result such plants grow taller. GA caused stem
elongation has the following characteristic features (1) enhanced stem growth is not
due to increased formation of nodes and internodes but results from rapid elongation
of internodes. Therefore GA treated plants do not differ from control plants in the
number of nodes and internodes. Elongation of internodes is due to both cell division
and cell elongation. Younger internodes respond better than older ones and plants
grown in light respond better to GA than those grown in the dark. However, not all
140

plants respond equally to GA application. It is only the genetic dwarf and rosette
plants which show marked stem elongation.
Genetic dwarfs:
From the field grown maize four mutants were isolated. These mutants grow
only up to about one fourth the heights of normal tall varieties. When gibberellin is
applied, the dwarf plants respond readily and show marked stem elongation. These
become comparable in height to corresponding tall varieties. Here the degree of
stem elongation is proportional to the concentration of GA that is applied.
B. Bolting:
Production of floral axis is called bolting. Bolting and flowering are induced
normally after photo induction or vernalisation. Bolting however can be induced
without vernalisation by the treatment of the plant with gibberellins.
Many plants require a period of low temperature for flowering. Application of GA

replaces the vernalization (0-50C) requirement for the flowering of carrot, beetroot,
chicory and others. Vernalization or low temperature requirement is usually met with
when the plants pass through natural winter. However this low temperature
requirement can be completely overcome and plants can be made to flower in high
temperatures by applying GA.
Therefore low temperature requirement of plants can be replaced with GA.
C. Flowering in long day plants:
Gibberellins promote flowering in long day plants under un favorable SD

conditions. Ex: Niger.
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D. Parthenocarpic fruits:
Gibberellins have been found to be more effective than auxins in causing

parthenocarpic development of fruits in plants like tomatoes, apples, pears and stone
fruits.
Gibberellin application promotes panicle exertion. Generally 30% of the

panicle is covered by leaf sheath. Application of GA + Brassinosteroids is practically
used in commercial seed production of Rice.
E. Breaking of dormancy:
Gibberellins are effective in breaking the dormancy in potato tubers and in

tree buds in winter.
In potato the tubers remain dormant for weeks after harvest. However when GA
is applied the buds sprout soon after the tubers are harvested. This will be useful to
use the freshly harvested tuber for sowing. The seed material has to be dipped in 0.5
to 1.0 g of GA /lit of water.
ANNEXURE
Gibberellin has been shown to cause synthesis of amylase in barley aleurone

cells. This enzyme converts starch to reducing sugars resulting in an increase of
osmotic pressure, causing entry of water into the cells and cell enlargement. GA3 is
also known to increase permeability of aleurone cells to sucrose. It increases the
activity of membrane synthesizing enzymes. Synthesis of phospholipids is also
increases due to GA application. The gearing up of all these metabolic activities
results in cell elongation.
*******
142

Lecture:26
CYTOKININS
Skoog and his coworkers discovered cytokinins when they were trying to
identify a compound to initiate and sustain the proliferation of cultured tobacco pith
tissue. Crystals of a cell division inducing substance was later isolated for the first
time by Miller, from an autoclaved herring sperm DNA in 1951 and named it as
Kinetin. The liquid endosperm of coconut (coconut milk) is also found to be rich in
cell division causing factors. Letham (1963) extracted, purified and crystallized
cytokinin from immature kernels of maize and named it as zeatin.
26.1 OCCURRENCE:
Naturally occurring cytokinins are N6- substituted adenine derivatives. Usually

Zeatin is the most abundant naturally occurring free cytokinin. There are also
synthetic cytokinin compounds that have not been identified in plants, most notably
of which are the diphenyl urea type cytokinins, such as thidiazuron, which is used
commercially as a defoliant and an herbicide.
Cytokinins, occur freely and also as a component of RNA of plants,

microorganisms and animals. In higher plants root tips, shoot tips, developing fruits,
xylem sap and germinating seeds are rich sources of cytokinins. Root tips synthesize
cytokinins and transport them through the xylem to all parts of the plants. This might
explain their accumulation in young leaves, fruits and seeds in to which xylem
transport occurs.
26.2 BIOSYNTHESIS:
The biosynthetic pathway of free cytokines is not completely understood.

There are two methods in which they may be produced. The first is the direct
pathway, involving formation of Isopentenyladenosine-5-monophosphate (IPAMP)
from AMP and dimethylallyl pyrophosphate (DMAPP), to form zeatin-type
compounds.
143

AMP (Adinosine dimethylallyl

mono phosphate) pyrophosphate (DMAPP)
isopentenyladenosine -5-
monophosphate (IPAMP)
zeatin
Another possibility is that they may be released by the hydrolysis of tRNA,

first to mono nucleotides and then to free cytokinins. In spite of extensive effort
having been focused on these pathways information is still highly fragmented.
TRANSPORT:
When cytokinin is applied to leaves and stems, the hormone does not move
and the effect is localized. Cytokinin is carried passively along the transpiration
stream in xylem from root. It moves in phloem in a basipetal polar direction in very
small quantities.
26.3 MODE OF ACTION:
Cytokinin is a structural component of transfer RNA molecule. They may help

in binding of mRNA with tRNA - amino acid complex during protein synthesis.
Cytokinins increase the synthesis of nucleic acid by increasing the enzyme t RNA
synthatase and decrease the degradation by reducing the activity of ribonuclease.
Cytokinin increases the incorporation of phosphorous in to nucleic acids and adenine
into RNA.
26.4 PHYSIOLOGICAL ROLE:
 Cell division
Cytokinins are known to be regulators of cell division in mature cells. The

most important effect of cytokinins is stimulation of cell division in excised tissues.
The number of cell divisions increases proportionally to the concentration of added
cytokinin when auxin is not limiting. Cytokinins alone does not promote cell division.
When both auxin and cytokinins are added together, cells divide rapidly and the
callus tissue grows.
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 Morphogenesis
Root and bud differentiation
Cytokinins in interaction with auxins control morphogenesis. The cells of

tobacco pith do not either grow or differentiate when only auxin or only cytokinin is
added to the medium. However when the medium contains both auxin and kinetin in
the ratio of 10:1 pith cells grow and forms a mass of unorganized cells (callus).
If the ratio of auxin to cytokinin is more in the medium, a number of roots are
initiated from the callus. If the ratio is less (which means more cytokinins than
Auxins) a number of shoot buds are initiated.
 Anti Senescence hormone (Richmond - Lang effect)
Cytokinins delay senescence. Generally, protein and chlorophyll content of

the leaf decreases with the increase in age. Thus, when leaf becomes old, it turns in
to yellow, become senescent and finally shed of. Senescence of leaves can be
delayed by application of kinetin. Cytokinins delay senescence by increased
synthesis of proteins.
The delay of senescence of leaves and other organs of the plants by

cytokinins is called as Richmond - Lang effect.
In an experiment, one of the two primarily opposite leaves of a bean plant was
treated with Benzyl adenine. This treatment accelerated senescence of untreated
leaf. This is because of mobilization of organic metabolites and minerals from
untreated leaf to cytokinin treated leaf because of cytokinin acts as mobilizing
centers.
Retardation of senescence of vegetables can be achieved by cytokinins. Green

vegetables like cabbage, lettuce and celery deteriorate rapidly after harvest. Post
harvest spray of Benzyl adenine at 10 to 40 ppm or post harvest dip of 10 ppm
increase shelf life of these vegetables.
 Promotion of lateral bud growth
Application of cytotinins reduces apical dominance. The action of cytokinin is

antagonistic to that of auxin in apical dominance. The lateral buds of intact plants
which otherwise remain arrested; can be made to grow by applying kinetin. It may be
due to the differentiation of vascular tissue in the presence of cytokinins.
The pathogen Corynebactrerium facians causes a disease called Witches

broom in many plants. This symptom is characterized by loss of apical dominance
and emergence of numerous lateral branches which give the appearance of a
145

broom. This effect is due to the secretion of cytokinin namely isopentenyl adenine by
the pathogen.
 Breaking of dormancy
Cytokinins can replace the red light (660 nm) requirement in seed germination
of lettuce and tobacco. Lettuce seeds require the presence of red light for
germination in addition to moisture, air and suitable temperature. However the seeds
can be made to germinate in the dark by applying Kinetin. Thus, Kinetin replaces the
red light requirement for germination.
In cocklebur, each fruit (but) contains two seeds which are of unequal in size.
The lower one in largest and germinates while the upper seed is dormant. Here the
dormancy is due to the presence of germination inhibitors. This dormancy is over
come by the application of kinetin.
 Cell enlargement
Cortical cells of tobacco root were observed to enlarge four times of their
normal size in the presence of kinetin.
*******
146

Lecture:27
ABSCISIC ACID (STRESS HORMONE)
27.1 Occurrence: the plant growth regulator ABA is one of the wide spread and
naturally occurring inhibitor found in plants. Addicot and his colleagues (1964)
isolated this abscission causing compound from cotton bolls and named it as
abscisin I & abscisin II. It is now known that ethylene is the hormone that triggers
abscission and that ABA induced abscission of cotton bolls is due to ABA’s ability to
stimulate ethylene production. In higher plants ABA occurs in all parts of the plant
body. It has been reported from the leaves of birch (Betula sps) and tubers of potato.
ABA is found in all parts of the seed namely the seed coat, embryonic axis,
cotyledons and endosperm.
27.2 Biosynthesis:
It is a sesquiterpenoid (15-carbon) which is partially produced via the mevalonic

pathway in chloroplasts and other plastids.
ABA is synthesis in plants involves two pathways (1) carotenoid pathway (2)
Mevalonic acid pathway or Isoprenoid pathway.
Mevalonate
Isopentenyl diphosphate Isopreniod pathway
Farnesyl diphosphate
Zeaxanthin
Violaxanthin
Neoxanthin Carotenoid pathway
Xanthaxin
ABA aldehyde
ABA
Site of synthesis
All parts of the plants such as stem, root and leaves. Fruits and seeds are
also capable of ABA synthesis.
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Transport
ABA is transported by both the xylem and phloem, but it is normally much
more abundant in the phloem sap.
27.3 Mode of action
ABA is involved in the short term physiological effects (e.g. stomatal closure),
as well as long term developmental processes (e.g. seed maturation).Rapid
physiological responses of ABA frequently involve alteration in the fluxes of ions
across membranes and may involve some gene regulation as well.
Mode of action of ABA in causing various physiological effects can be seen in

three different ways
1) ABA brings about changes in membrane permeability for different ions (Like
K+, Ca++ etc.) . It plays a role in stomatal closure.
2) ABA inhibits DNA & RNA synthesis (transcription) finally leads to senescence.
3) It inhibits translation (Protein synthesis) thus formation of enzymes is blocked.
Germination process is affected ultimately leads to dormancy.
27.4 PHYSIOLOGICAL ROLE
A) Seed dormancy:
Application of ABA inhibits seed germination in several species.
80%
70%
60%
Germination 50%
40%
30%
20%
0 0.2 2.0 3.0
Conc ABA mg/l
Similarly seeds which are dormant are shown to contain ABA.
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Seeds of apple remain dormant and fail to germinate till they are exposed to a
period of stratification. such seed show the presence of ABA. When the seeds are
stratified the ABA content falls with a corresponding increase in GA content. Thus, it
can be concluded that the seed dormancy is controlled by GA-ABA balance at least
in some species.
ABA helps in inhibiting precocious germination and vivipary. This is very important
because dormancy caused by ABA do not allow the seed to germinate while it is still
on its mother plant.
B. Bud dormancy
In woody species, dormancy is an important adoptive feature in cold

climates. When a tree is exposed to very low temperatures in winter it protects its
meristems with bud scales and temporarily stops bud growth. Bud dormancy in Acer,
betula and other temperate tree sps. This is accompanied by build up of ABA within
these plants.
C. Effect of stomata
Application of ABA causes rapid closure of stomata. The stomatal aperture
progressively decreases with the concentration of ABA.
*FIGURE IS FROM REFERENCE 6

ABA accumulates in higher concentration in wilting leaves. This accumulated
ABA closes stomata. ABA might inhibit the formation of enzymes which are
responsible for the conversion of starch into sugar and formation of organic acids. It
reduces the osmotic concentration and causes closure of stomata.
149

D. Senescence
ABA is quite effective in promoting senescence of excised leaf disks of both
monocots and dicots. Although leaf disks are affected, when sprayed on the
corresponding intact leaf, ABA is not effective even at higher doses in inducing
senescence.
E. Flower initiation
ABA induces flowering in SD plants and inhibits the same in LD plants. Here
the hormone inhibits vegetative growth and causes apical bud dormancy. In such
plants onset of dormancy precedes flowering. Therefore, effect of ABA on flowering
is indirect.
F. Antagonism
ABA inhibits GA stimulated growth in various forms. Therefore ABA is known
as Antigibberellin.
*******
150

Lecture:28
ETHYLENE
Neljubow (1901) a Russian plant physiologist was the first to show the
importance of ethylene present in the illuminating gas as a growth regulator of plants.
He observed that dark grown pea seed lings growing in the laboratory (illuminated
with coal gas) exhibited symptoms that were later termed as triple response: reduced
stem elongation, increased lateral growth, and abnormal horizontal growth. Denny
(1924), reported that ethylene is highly effective in inducing fruit ripening. Gane
(1934) established that ethylene is produced by the apple fruits during ripening in
storage.
28.1 Occurrence
In higher plants, all most the parts of the plant body produce ethylene.In
general meristematic regions and nodal regions are most active in ethylene
biosynthesis. However, ethylene production also increases during leaf abscission
and flower senescence, as well as during fruit ripening. This is otherwise called as
phytogerontological hormone .
28.2 Bio-synthesis
The amino acid Methionine is the precursor of ethylene, and ACC (1-amino
cyclo propane 1-carboxylic acid) serves as an intermediate in the conversion of
methionine to ethylene. Methionine activated by ATP gives rise to S-adenosyl
Methionine (SAM). This reaction is mediated by enzyme Methionione adenosyl
transferase. In the next step, SAM breaks into 5’-methyl thio adenosine (MTA) and
amino cyclo propane carboxylic acid (ACC). This reaction is carried by the enzyme
ACC synthase. ACC is oxidized to ethylene with the release of HCN (Hydrogen
cyanide) and Co2.
METHIONINE
ATP
ACC
ACC OXIDASE
ADP + Pi SYNTHASE
ACC (1-amino cyclo ETHYLENE
S- ADENOSYL
propane 1-carboxylic acid) HCN + Co2
METHIONINE (SAM)
MTA
Amino ethoxy vinyl glycine (AVG), and Amino oxy acetic acid (AOA) block
the conversion of SAM to ACC. Silver Nitrate and Silver thio sulphate are the specific
inhibitors of ethylene action.
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Ethylene Transport
Ethylene being a gas can easily diffuse into plant tissues through the
intercellular spaces. From ripening fruits, ethylene diffuses out into the atmosphere
through the cut end of pedicel and fruit surface.
28.3 MODE OF ACTION:
There are several theories to explain the mechanism of action of ethylene.
Membrane permeability
Ethylene is considered to dissolve in cell membranes altering their

permeability. Ethylene is highly soluble in lipids which are constituents of cell
membranes.
Another hypothesis is by regulating auxin metabolism. Ethylene treatment

results in a reduction in the content of diffusible (free) auxin. This may result from (1)
decreased synthesis (2) decreased transport and (3) increased binding. Ethylene
has been shown to inhibit transport of auxin from the site of production to the site of
action.
28.4 PHYSIOLOGICAL ROLES (Includes both positive and negative effects)

A. Fruit ripening
Broadly fruits can be classified into two types on the basis of their respiratory
pattern during ripening. In some fruits like apple and banana as the fruit matures
and attains its maximum size, the rate of respiration decreases and becomes very
low. After the fruit is harvested and stored for ripening, there is a great increase in
the rate of respiration and the rise continues till it attains a sharp peak. This is called
climacteric peak and the fruits are called climacteric fruits. In climacteric fruits
ripening occurs even after harvesting. The climacteric rise is soon followed by a
sharp decline.
The non climacteric fruits like grapes and lemon, the respiratory rate gradually
decrease after the fruit is harvested without showing any abrupt rise. The peak
respiratory rate in climacteric fruits usually corresponds to peak ethylene production.
Application of ethylene hastens ripening of climacteric fruits such as banana,

mango, apple and tomato. This is being commercially employed. In non climacteric
fruits such as lemon and orange ethylene application does not hasten ripening
however rate of respiration increases greatly.
B. Abscission and senescence
Ethylene promotes both abscission and senescence of flowers. The flowers

of orchids and roses the most sensitive to externally applied ethylene. Ethylene also
152

promotes leaf abscission. In general older leaves are more sensitive to ethylene and
abscise faster than younger ones. Older leaves produce more ethylene than
younger ones. This is probably responsible for abscission of older leaves.
C. Roots on stem cuttings
Application of ethylene promotes callus formation and initiation of adventitious

roots on the stem cuttings. Some times adventitious roots may arise on the stem of
intact plant as well.
D. Root and shoot growth
Ethylene inhibits linear growth of the stem and root of dicotylidons. The effect
increases with increasing concentration.
E. Flowering and sex expression
Application of ethylene causes flowering in pine apple and shift the sex ratio of
flowers towards femaleness in several cucurbits and cannabis.
F. Epinasty
Ethylene causes swelling of cells on the upper part of the petiole of the leaf
resulting in drooping of leaves (down ward curvature). This is termed as epinastry.
It is best exhibited in leaves of tomato, potato and pea etc.
G. Thinning in apple
Thinning of fruits in apple eliminates biennial bearing and also improves fruit
size and quality. Application of ethephon at 100 to 300 ppm reduces fruit set. In
cotton also ethylene induces this fruit thinning.
H. Exudation of sap and latex
When ethereal is applied to rubber plants, flower of latex continues for a

longer duration Etherel probably prevents coagulation of late and consequent
blocking of laticiterious ducts.
*******
153

Lecture:29 PLANT GROWTH REGULATORS
29.1 NOVEL PLANT GROWTH REGULATORS
A) JASMONATES: Jasmonic acid (JA) and its methyl ester (MEJA) occur in several
plants and also in the oil of jasmine.
a) JA and MeJA inhibit the germination of non dormant seeds and stimulate the
germination of dormant seeds.
b) JA plays a role in the formation of flowers, fruit and seed. It is suggested by

the relatively high levels of this compound in developing plant reproductive
tissues.
c) Reported a role in insect and disease resistance of plant.
d) JA stimulates tomato and apple fruit ripening.
B) BRASSINO STEROIDS (BRs) : Brassinolide, a potent plant growth stimulator,

was the first BR isolated; it was discovered in rape (Brassica napus) pollen in
1979.So far, more than 40 brassinolide analogues, collectively known as
brassinosteroids, have been identified and characterized from many different plant
species.
 Immature seeds and pollen contain the highest concentrations of

brassinosteroids.
 When applied exogenously to intact plants at micromaolar or nanomolar
concentrations BRs can induce a variety of physiological responses,
including seed germination, pollen tube growth, stem elongation, leaf
unrolling and bending, ,vascular differentiation, induction of ethylene
biosynthesis, altered gene expression, and stress response modulation.
 They help to Increase flowering fruit set and yield
 Foliar spray of 0.3 ppm BR at panicle initiation and flowering stage
increases yield in rice
C.SALICYLIC ACID: Salicylic or ortho hydroxy benzoic acid belongs to a diverse

group of plant phenolics. The interest in salicylic acid began with the discovery that
this compound is a natural trigger for the metabolic explosion which raises the
temperature of the thermogenic inflorescences of Arum Lillie.
 SA increase flower longevity by inhibiting ethylene biosynthesis by blocking

the conversion of ACC to ethylene.
 SA regulates some aspects of disease resistance.
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D. TRIACONTANOL: Saturated primary alchohol isolated from shoots of alfalfa.

Response is very rapid in increasing growth. Enhanced growth in rice and maize is
reported.
29.2 GROWTH RETARDANTS
The term growth retardant refer to the chemicals that slow down cell division and
cell elongation of shoot tissue and regulate plant height physiologically without
formative effects. They do not occur naturally in plants.
Examples:
 AMO 1618 ,
 CCC (Chloro choline chloride ) (2-Chloroethyl – trimethyl ammonium chloride)
 Chlormequat chloride ( Cycocel)
 Alar or B9
 paclobutrazol
 Mepiquat chloride,
29.3 GROWTH INHIBITORS
Growth inhibitors suppress the growth of plants. ABA and ethylene are called as
natural growth inhibitors. They bring about certain formative changes in plants. There
are synthetic growth inhibitors also.
Examples:
 Malichydrazide (MH)
 2,3,5-T or Triiodo benzoic acid (TIBA)
29.4 COMMERCIAL APPLICATION OF PLANT GROWTH REGULATORS IN

AGRICULTURE AND HORTICULTURE:
A) AUXINS
a) IBA (@250 ppm) and NAA were found to increase root development in the
propagation of stem cuttings.
b) 2,4-dichlorophenoxy acetic acid (2,4-D) stimulates excessive uncontrolled

growth in broad leaf plants for which it is used as a herbicide.
c) Application of NAA (Napthalene Acetic Acid) reduces flower and fruit drop in
Mango.
d) NAA application brings uniform flowering and fruit set by inducing ethylene
formation in Pineapple.
e) NAA application @ 10-100 ppm during fruit setting period controls boll
shedding in cotton crop.
155

B.GIBBERELLINS:
a) GA is used extensively on seedless grape varieties to increase the size and

quality of the fruit. Pre- bloom spray of 20 ppm induces rachis of the fruit
cluster to elongate. This creates looser clusters that are less susceptible to
disease during the growing season.
b) GA is used to increase the yield of barley malt and to decrease the time
required for this process to occur. Application of GA to germinating barley
supplements the endogenous content of this hormone and accelerates the
production and release of hydrolytic enzymes. They can easily degrade the
stored carbohydrates.
c) Foliar spray of GA3, at 100 ppm during panicle initiation stage enhances the
panicle exertion and increases seed weight and yield in hybrid rice.
d) GA has also has been used to control flower sex expression in cucumbers
and squash. GA application tends to promote maleness in these plants.
e) Gibberellic acid is also applied to citrus crops, though the actual use depends
on the particular crop. For example GA3 is sprayed onto oranges and
tangerines to delay or prevent rind-aging, so that fruit can be harvested later
without adverse effects on rind quality and appearance. For lemons and limes,
GA3 synchronizes ripening and enhances fruit size.
f) Gibberellic acid is used extensively to increase the sucrose yield of

sugarcane. Sugarcane, a normally fast-growing C4 member of the Poaceae
(grass) family, is sensitive to cooler winter temperatures, which reduce
internode elongation and subsequent sucrose yield. The adverse effects of
cooler temperatures can be counteracted by the application of GA3.
C.ETHYLENE:
a) Because ethylene regulates so many physiological processes in the plant

development it is the most widely used plant hormones in agriculture. Auxins
and ACC can trigger the natural biosynthesis of ethylene and in several cases
are used in agricultural practice.
b) Because of its high diffusion rate, ethylene is very difficult to apply in the field
as a gas, but this limitation can be over come if an ethylene releasing
compound is used. The most widely used such compound is ethephon or 2-
chloro ethyl phosphonic acid (CEPA)( trade name: ethrel).
156

c) Ethrel @ 100-250 ppm sprayed at 2-3 leaf stage induce femaleness in

cucumber and melons.
d) It helps in degreening of citrus and banana which increases its market

acceptability.
e) Storage facilities developed to inhibit the ethylene production and promote

preservation of fruits have a controlled atmosphere of low 02 concentration
and low temperature that inhibits ethylene biosynthesis. A relatively
concentration of C02 (3-5%) prevents ethylene action as a ripening promoter.
D) OTHER GROWTH REGULATORS:
 AMO 1618 (a quaternary ammonium salt) is used in the cultivation of

ornamental plants and causes a bushy shape and a sturdy growth of the
treated plants.
 paclobutrazol : Reduces the problem of biennial bearing in Mango
 Mepiquat chloride, Chlormequat chloride ( Cycocel) : used in ornamental
plants for shorter internodes and thicker stems (used in poinsettias), it also
prevents lodging and increases tillering in cereals.
 Malichydrazide (MH): prevents premature sprouting of onion and potato
 2,3,5-T or Triiodo benzoic acid (TIBA): Increases flowering in chrysanthemum
*******
157

Lecture-30
SENESCENCE
30.1 Definition: The term senescence’ is derived from a Latin word “Senescere”
which means to grow old’. The terms Senescence, Programmed Cell Death (PCD)’,
Apoptosis’ and Ageing’ are often used synonymously in plant or animal systems. All
these terms generally refer to death of cells, organs or organisms.
In the life cycle of higher plants when they have reached a certain stage of
maturity, they senesce and die. It should be clarified that the ageing is not the same
as senescence. ‘A degenerative and irreversible change in a plant which leads
to death’ is called senescence where as ageing is ‘the process of attaining
maturity with the passage of time’.
30.2 CLASSIFICATION OF SENESCENCE (TYPES OF SENESCENCE):

Depending upon the part of the plant in senescence, Leopold (1961) has classified
senescence in to following four categories.
A. Overall senescence: (whole plant senescence): In this kind of senescence, there

is senescence and death of the entire plant, which usually takes place at the end
of reproductive phase.
Eg: Cereal crops like maize, wheat and rice and also in mustard and cabbage
B. Progressive senescence: In normal development of most annual plants, there is
progressive senescence, where the oldest leaves senesce and die first. The
senescence moves from leaves to the stem and then to under ground parts.
Eg: Tobacco.
C. Top senescence (Shoot senescence) Here senescence and death of all above
ground parts occurs, while the under ground root portions survive and give rise to
new buds in the next season.
Eg: Sugar beet, Banana, Ginger
D. Deciduous senescence (simultaneous senescence): In this kind of senescence,
all leaves senesce and die, leaving the stem and roots alive as in gulmohar
(Delonix regia) and Raavi (Ficus religiosa), Eucalyptus.
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The occurrence of senescence differs both in their causes and in its nature.
Senescence of the entire plant after a single reproductive cycle is called
monocarpic senescence. Senescence may be delayed when flowers and fruits are
removed. Some monocarpic plants like Agave americana generally live and grow for
many years before flowering. They die when they produce fruits. Thus senescence is
an irreversible one. On the other hand in tobacco plant older leaves senesce as the
plant grows. But the senescence can be reversed when the apical meristem of the
plant is removed.
30.3 THEORIES OF MECHANISM OF SENESCENCE:
30.3.1 Nutritional theory: Molisch (1920) suggested that senescence was caused
by nutritional deficiency. Different parts of the plants compete for nutrition. Fruit or
growing tip for example, might form stronger sinks for translocation and thus
accumulate much of nutrients, that the older leaves would be starved. This triggers
off the senescence process in the older leaves and they start acting as a nutrient
source for younger structures. One of his observations is that, if the fruit, seed or
growing tip is removed, senescence is greatly delayed in the leaves. However,
nutritional theory is not applicable to many situations. It is not possible to retard
senescence in annual plants that have flowered or fruited even by application of
fertilizers. Dioecious plants, which bear only male flowers and require no additional
nutrients for fruit formation also undergo senescence.
30.3.2 Hormonal theory: It is observed that senescence is initiated as a result of

change in hormonal content of the organ. Some believe that a hormonal signal is
sent from developing fruits to leaves and other vegetative parts, where it triggers
senescence. It is now well known that ABA produced in the developing fruits and
seeds induce senescence in the leaves.
Nooden and Leopold (1978) proposed a term “Death hormone” for the
causative agent of senescence in monocarpic species. The death hormone is a
chemical substance which is supposed to be produced in developing seeds and
translocated to vegetative parts through xylem and initiates senescence in leaves.
This hypothesis is based on the observations that preventing flowering or fruiting
retarded senescence in many cases.
30.3.3 Alternations in nucleic acid and protein contents with senescence and
usually it will decline in senescing plants (Carr and Pate, 1967)
30.3.4 Increase in enzymatic activities of ribonuclease and proteases with the

onset of senescence in plant (Martin and Thimann, 1972)
159

30.3.5 Structural alterations particularly evidence of deteriorative changes in

membranes and organelles (Poovaiah and Leopold , 1973)
30.3.6 Increased accumulation of free radicals with ageing (Choudhri, 1988)

No hypothesis is fully satisfactory or complete and various criticisms can be
put forward for and against of these hypothesis.
30.4 METHODS OF PREVENTING LEAF AND FLOWER SENESCENCE:
All the plants do not exhibit same response to growth hormones. Cytokinins
appear to be more effective in delaying senescence in many herbaceous plants.
GA’s are effective in preventing senescence of ash (Fraxinus) because the
endogenous GA content of the leaves decrease during leaf senescence. Auxins (IAA
and 2,4-D) have been found to retard senescence in certain trees. Delaying the leaf
senescence is a desirable phenomenon in annual crops like rice for better grain
filling. Premature senescence of panicle also affects the grain yields in rice. By the
use of growth regulators like Kinetin and triactontenol, the panicle senescence can
be retarded by maintaining high succinic dehydrogenase activity (SDH) in the panicle
components. High rate of N application at booting stage will maintain High SDH
activity and retard panicle senescence.
30.5 PHYSIOLOGICAL AND BIOCHEMICAL CHANGES DURING SENESCENCE:
Physiological Biochemical
1. Yellowing of leaves because 1. Decrease in starch synthesis
Of decline in chlorophyll content
2. Decrease in photosynthetic rate 2. Loss of ATP synthesis
This may be due to
a. Ultra structural changes in chloroplasts 3. Decrease in DNA and RNA
b. Decrease in chlorophyll content Synthesis
c. Increase in stomata resistance and
d. Decrease in the activity of Rubisco enzyme 4. Decrease in protein
3. Decrease in rate of respiration Synthesis. This decline is both
However in some speciess there is increase due to their accelerated
in respiration degradation as well as
decreased Synthesis
4. Increased membrane permeability
5. Membranus subcelular inclusions are 5. Increase in contents of
Disrupted hydrolytic enzymes such as
Protease and nuclease.
6. Cells undergo reduction of their structure 6. Decrease in inorganic ions
and Various nutrients.
160

30.6 SIGNIFICANCE OF SENESCENCE: The main purpose of leaf senescence is to

recover the nutrients, specially nitrogen and carbon for the growth of younger leaves
and other developing organs on the plant. The senescence of leaves in deciduous
trees is also a mechanism of avoiding extreme environmental conditions such as
severe cold. Of late scientists are trying to develop ‘stay green varieties‘ which can
retard the process of leaf senescence and can maintain green leaves for a longer
period.
30.7 ABSCISSION AND ITS RELATION TO SENESCENCE:

The term abscission is used to describe ‘the processes involved in the
shedding of plant structure, characterized by the degradation of cell walls at
the point of weakening’. Cells surrounding the fracture line produce and secrete
cell wall degrading enzymes which hydrolyze the central region of the wall allowing
the cells to separate for fracture to occur. This fracture occurs in “Separation layer”
which is 1-3 cells wide. Plants do not have the ability to produce separation layer any
where. They are genetically limited to specific locations called “Abscission Zones”
which are 5-50 cells wide.
Cells of abscission zone are somatic and have more persistent meristematic
activity. Lignified structural elements like fibers and sclereids are absent in
abscission zone and are replaced by collenchymas. Lignified walls are extremely
resistant to enzymatic hydrolysis while collenchymas walls are readily degradable.
Abscisic acid was originally isolated as an “abscission causing factor”.
However it is evident that ABA stimulates abscission of organs in only few species
and that the primary hormone causing abscission is ethylene. On the other hand
ABA is clearly involved in senescence, and through promotion of senescence it might
indirectly increase ethylene formation and stimulate abscission. Senescence acts as
a signal for inducing abscission and senescence of plant parts usually precedes the
abscission.
But there are examples where leaf senescence occurs in the autumn and
abscission does not occur until the following spring. Thus, the linkage between
senescence and abscission can be broken since, both processes occur
independently.
*******
161

Lecture:31
POST HARVEST PHYSIOLOGY
SEED DORMANCY
Seeds harvested at physiological maturity are known to possess maximum

viability and vigor. The ability of these seeds to germinate depends on the internal
and external environment of the seeds. Any disturbance in external or internal
condition leads to failure of germination in seeds.
31.1 Definition: Dormancy is defined as physical or physiological condition of the

seed that prevents germination in the presence of otherwise favorable conditions
for germination.
Dormancy may occur with in the embryo (Ground nut) or in the seed coat
(Sunflower). The period of dormancy varies from a few days to several months
depending on the plant species
After-ripening: The period of rest after harvest that is necessary for germination
is sometimes referred to as after – ripening period and the changes that take
place during the rest are described as after – ripening.
Quiscence: is the phenomenon in which the seeds fail to germinate for want of a
particular environmental factor.
31.2 TYPES OF DORMANCY:
A. Primary dormancy: This is otherwise called as innate dormancy. These seeds

enter in to dormancy much before they are harvested i.e when they are still on their
mother plant itself.
e.g; 2-3 months in Virginia runners of Ground nut
Up to 40 days after the seed is formed in sunflower.
B. Secondary dormancy: This is called as induced dormancy. These seeds are

non dormant when they are harvested from the mother plant. How ever when they
are exposed to brief periods of un favorable environmental condition they show
dormancy.
e.g: Mustard seed exposed to high concentration of CO2
Wheat stored in high moisture content in air tight containers at 50 0 C
31.3 ADVANTAGES AND DISADVANTAGES OF DORMANCY
The survival of natural populations depends mainly on their ability to exploit

the favorable and avoid the unfavorable weather conditions to which they are
cyclically exposed in their natural habitats. The state of dormancy equips organisms
162

to escape the detrimental effects of adverse natural environments, thereby

enhancing their chances of survival.
Plants with a long history of domestication generally show much less seed
dormancy than wild or recently domesticated species. Thus crop domestication has
promoted rapid seed germinability. This may often result in premature germination
of seeds within ears/pods (vivipary) when the crops are exposed to a wet weather
favorable for germination just before harvest. In such cases, a pre-harvest rain leads
to deterioration in the quality of crop produce, e.g., in wheat, it reduces seed quality
and vigor, milling and baking quality, and even grain yield. Therefore, a certain
degree of seed dormancy is often deliberately selected for in order to prevent pre-
harvest sprouting in cereals. A brief period of dormancy provides adequate time to
farmers to harvest, thresh and store the seeds, thereby avoiding considerable
losses.
Disadvantage: The seeds with dormancy cannot be used immediately after

harvest for seed purpose.
31.3 CAUSES FOR DORMANCY:
31.3.1 Seed coat factors
(a) Seed coat impermeable to water: Common in seeds of Leguminaceae,

Malvaceae, convolvulaceae and chenopodiaceae. Seed coat with thick
waxy cuticle, lignin and suberin barriers makes seeds impermeable to
water.
(b) Seed coat impermeable to oxygen: Any disturbance to the entry of
oxygen and exit of CO2 decreases respiration and there by remains in
dormant condition.
(c) Mechanically resistant seed coat: Hard seed coats of nuts make it
difficult for embryo to germinate and break the seed coat. High salt
concentration in water also aids to cause mechanical resistance to
germination.
31.3.2 Embryo Factors
(a) Immature/redimentary embryo : found in Ranunculus plantago . These

embryos require after ripening though their growth is completed
morphologically. This is also called as physiological dormancy.
31.3.3 Inhibitory Factors
The inhibitors ( e.g : coumarin, caffeic acid , ferulic acid and ABA) may
present in the embryo, endosperm or seed coat or pericarp.These inhibitors
deactivate enzymes like Amylase, protease and phytase. This will limit the
supply of simple substances like sugars, fatty acids and P necessary for
163

germination. If the ratio of these inhibitors is higher than the that of

endogenous hormones especially GA the seeds remain dormant till a
balance between them is reached in favor of growth promoters.
31.4 REMEDIAL MESURES / METHODS TO BREAK DORMANCY:
A number of methods of overcoming dormancy have been developed; these

methods are either aim at breaking/softening of seed coats or at promoting seed
germination through stimulation of embryo. The various treatments for overcoming
dormancy may be divided into the following three groups: (I) seed coat treatments,
(II) Embryo treatments and (III miscellaneous approaches.
31.4.1 Seed Coat Treatments:
These treatments are either physical or chemical in nature, and aim at making
hard seed coats permeable to water and / or gases by either cracking or softening
them; the process is usually referred to as scarification.
a) Mechanical scarification:
- Rotating the seeds in machines having drums with abrasive surfaces.
- rubbing the seed against abrasive surfaces manually . Eg. Coriander
b) Chemical scarification: This is achieved by the use of sulphuric acid, Hcl,
NaOH, alcohol, acetone, oxidizing agents etc.
-Treating in 3% nitric acid solution for 6-8 hrs can relieve the seed
dormancy in Rice.
Among the several methods available the most suitable method to break
seed dormancy at farmers level is nitric acid treatment. - Soaking the seed in o.1 N
nitric acid i.e., 6.3 ml per lit. of water for 12 to 24 hours effectively breaks the seed
dormancy in less or moderately dormant varieties. However, the varieties like MTU-
1001 ( VIJETHA) which is having 8 weeks and above dormancy duration should be
treated with higher nitric acid concentration i.e., 10ml per lit. of water. The seeds
can be utilized for sowing immediately after the treatment or they can be dried
thoroughly and can be utilized later for sowing.
Scarifications must be done with caution and care; otherwise it may damage
the seeds. It may be done in one of the following ways.
 Scarification may be achieved by rubbing the seeds on a sand paper manually

or by using a mechanical scarifier; care should be taken to avoid damage to
the embryonic axis. This treatment is effective in species like ‘subabool’,
green gram etc.
164

 The seed coat may be pierced by a needle or a small incision may be made in
it at the abaxial end of the seed, e.g., in bitter gourd (Memordica charantia).
 In some cases, the seed coat may be completely removed by breaking e.g., in
rubber (Hevea Spp). In this technique, each seed has to be handled
individually, which makes the treatment slow, time taking and tedious.
 The seed may be soaked in a concentrated or dilute solution of sulphuric
acid for 1 to 60 minutes, followed by through washing with tap water to
remove all traces of acid e.g., in cotton (Gossypium spp)
 In some species e.g. lentil, Bengal gram etc. soaking the seeds in hot water
(800C) for 1-5 minutes effectively softens their seed coats. But seeds of some
species may be highly sensitive to this treatment e.g. a treatment of more
than 1 minute reduces the germinability of Bengal gram seeds
Mechanical scarification, especially, manual scarification, is the most

commonly used technique and is relatively safer but, quite often tedious.
31.4.2 Embryo treatments
When dormancy is due to factors located within the embryo such treatments
have to be applied that are capable of inducing the embryo to resume growth. Some
of the common treatments are briefly described below
A. Stratification : stratification is the incubation of seeds at a suitable low

temperature (usually,0-5oC) over a moist substratum before transferring them
to a temperature optimum for germination; it is a common embryo treatment
designed to overcome dormancy. It is commonly used in crops like cherry
(Prunus cerasus), mustard (Brassica campestris)species of family Rosaceae
(2-6 months at 5-10oC)etc.
B. High temperature treatment: In some plants incubation at 40-500C for few
hours to 1-5 days may be effective in overcoming dormancy. Care should be
taken that the moisture content of seeds should be less than 15% For
example rice (Oryza sativa ) seeds having less than 15% moisture are
incubated at 40-500C for 4-5 days for overcoming dormancy.
C. Chemical / Hormonal treatments: The growth regulators most commonly

used for this purpose are GA, (100ppm is the most commonly used
concentration) and kinetin (concentration range, usually 10-15ppm). Benzyl
adenine at 2 ppm and ethrel at 250ppm are effective in breaking seed
dormancy of sunflower. Ethrel 75ppm and GA at 60 – 75 ppm are effective in
controlling seed dormancy in groundnut.
Other widely used chemicals are potassium nitrate (0.2%) and
thiourea (0.5 to 3%) Potassium nitrate breaks the dormancy of seeds
requiring light and allows them to germinate in dark e.g. in case of oats, barley
165

tomato, etc. Thiourea breaks the dormancy of seeds requiring light and / or
chilling e.g.in lettuce, Gladiolus etc.
31.4.3 Misscellaneous Treatments
Exposing the seeds of many species to red or white Light leads to a

termination of dormancy For example lettuce seeds exposed to red light at
660nm or to white light are induced to germinate. Generally seeds are
placed initially in red light, and they are subsequently transferred to dark or
white light for germination.
31.5 OPTIMUM SEED STORAGE CONDITIONS
Seeds that can be stored in a state of low moisture content are called
“Orthodox” seeds ( Rice, Mung, sorghum, cotton etc.)The seeds that can be stored
in a state of high moisture content are called as recalcitrant seeds (Cacao,
Rubber,Tea etc.). Their viability depends on storage conditions and follows some
general rules: (Harrington thumb rules) (ISTA rules) (International Seed Testing
Association).
a) For each 1% decrease in seed moisture content, the storage life of the seeds is
doubled.
b) For each 100F (5.60C) decrease in seed storage temperature, the storage life of
seed is doubled.
c) The arithmetic sum of storage temperature in degrees F and the percent relative
humidity should not exceed 100, with no more than half the sum contributed by
the temperature.
These “rules of thumb” clearly indicate that temperature and moisture content
of the seed are major factors in determining the viability of seed.
31.6 FACTORS AFFECTING SEED VIABILITY DURING STORAGE:
31.6.1. Seed Moisture:

a) > 30% : Non dormant seeds may germinate
b) 18-30% : Rapid deterioration by microorganisms
c) 8-20% : Seeds respire rapidly and in poor ventilation, the generated
heat will kill them.
d) <8-9% : There is little or no insect activity
e) <4-5% : Immune from attack by insects and storage fungi, but they may
deteriorate faster than those maintained at a slightly higher
moisture content.
166

31.6.2 Temperature:
Cold storage of seeds at 0-50C is generally desirable if they are dried to safe
moisture limits and sealed in moisture proof containers.
31.6.3 Relative humidity
The activities of seed storage fungi are more influenced by the RH of the inter
seed atmosphere than by the moisture content of seeds. Oil seeds may differ in their
moisture content from starchy seeds even though both are at same atmospheric RH.
At 45% RH starchy seed have 11% moisture and oil seeds have 4-6% moisture.
31.6.4 Cultivar and harvest variability

Different cultivars and harvests of a particular species may show different
viability characteristics under the same storage conditions.
31.6.5 Pre and post harvest conditions

Environmental variation during seed development usually has little effect on
the viability of seeds, unless the ripening process is interrupted by premature
harvesting. Weathering of matured seeds in field particularly in excess moisture or
freezing temperatures, results in inferior storage potential. Mechanical damage
during harvesting severely reduces the viability of some seeds.
31.6.6 Oxygen pressure during storage:

If seeds are not maintained in hermetic storage at low moisture contents, even
under constant temperature and moisture, the gaseous environment may change as
a result of respiratory activity of the seeds and associated micro flora.
31.6.7 Fluctuating storage conditions:

Onion and Dandelion seeds stored under conditions of alternating high and
low RH lose viability proportionately to the length of time that the seeds are
subjected to the high RH.
*******
167

Lecture:32
POST HARVEST PHYSIOLOGY

FRUIT RIPENING
32.1 Definition: Ripening may be defined as those nonreversible, diverse, physical,

chemical and qualitative changes that render the fruit attractive for consumption at
the transition phase following maturation.
32.2 Metabolic changes during fruit ripening :

During the post harvest handling and storage period, and attempt is
generally made to maximize control over texutural changes to prevent, synchronize
or accelerate the process.
32.2.1 Seed maturation Quality attributes

32.2.2 Changes in pigmentation
a) Degradation of chlorophyll
b) Unmasking of existing pigments Color
c) Synthesis of carotenoids
d) Synthesis of anthocyanins
fruit ripening is usually associated with changes in the color of the fruit which
is due to changes in the pigment composition of chlorophyll, carotenoids and
anthocyanins. In the ripening fruit, there is a fast disappearance of chlorophyll
accompanied by accumulation of red and yellow carotenoid pigments in the
chloroplast. As the tomato fruit matures, the predominant carotenoid that is
synthesized is carotene. Some fruits like grapes, pomegranate produces
anthocyanins when mature. In tomato another pigment accumulates during ripening
is Lycopene.
32.2.3 Softening
a) Changes in pectin composition
b) Possible alterations in other cell Texture
wall components
c) Hydrolysis of storage materials
Softening may be a detrimental quality in some fruits like cucumber, squashes
which are consumed in unripe state. In others, it is an essential component in the
development of optimum quality. With the progress of ripening the fruit softens. The
softening is due to enzymatic hydrolysis of polysaccharides. The cell wall is made up
of cellulose, hemi cellulose, calcium pectate, polyuronides, and glycol protein. The
important cell wall hydrolyzing enzymes like pectin methyl esterase (PME), poly
galacturenase (PG) and cellulose increases during ripening and the dissolution of
middle lamella is observed. This is accompanied by increase in the enzyme PME.
168

Another enzyme poly galacturonase (PG) which is not present in green tomato
appears during ripening. This enzyme hydrolyses oligo galacturonoids.
32.2.4 Change in carbohydrate composition.

a) Starch conversion to sugar
b) Sugar inter conversions Flavor
32.2.5 Production of aromatic volatiles
32.2.6 Changes in organic acids
During ripening, starch hydrolysis occurs, and sugar accumulates. For

example starch content of banana decreases from the initial 21% to about 1% in
ripened fruit. This is accompanied by accumulation of sugars mainly sucrose to the
extent of up to 20% by fresh weight. The taste of the fruit depends upon the sugar-
acid ratio and also on the absolute level of sugar and acid contents. The PH of all the
fruits is in the acidic range.
32.2.7. Fruit abscission occurs.

32.2.8 Changes in respiration rate (respiration rate increases).
32.2.9 Increased permeability of tissue.
32.2.10. Quantitative and qualitative changes in protein occur.
32.2.11 Development of surface waxes occurs.
32.3 FACTORS INFLUENCING FRUIT RIPENING
A)Temperature affects the rate of synthesis of specific pigments and their final
concentration in the fruit. The optimum and maximum temperature for synthesis of a
specific pigment varies between species. For eg. Lycopene synthesis in tomato is
inhibited above 300C whereas in watermelon synthesis is not prevented until the fruit
temperature rises above 370C.
B)Oxygen is essential for carotenoid synthesis and increasing the oxygen

concentration enhances the synthesis of this pigments.
32.4 Climacteric and non- climacteric Fruits: Fruits of different species vary in
their ability to ripe, when detached from the parent plant. Many fruits must be
harvested only when fully ripe eg. Grape, cherry, lemon etc. they are not capable of
ripening after detachment and are called non climacteric fruits. Other fruits like
apple, banana, tomato can be harvested unripe but after fully mature stage. They
can undergo normal ripening even though detached from the parent plant and are
called climacteric fruits. These fruits often, undergo hydrolytic conversions in
storage materials and synthesize the pigments and flavors associated with a ripe
fruit.
169

32.5 HORMONAL REGULATION OF RIPENING

32.5.1 Ripening Induction
Application of ethephon promotes degreening and early ripening in grape,
tomato, coffee, peach, pear, plum and citrus. Smoking is commercially employed to
hasten degreening and ripening of banana and mango. Calcium carbide release
acetylene, on hydrolysis of which hasten ripening process. ABA (1 ppm) Thirourea
(20%), CCC (4000ppm) (2 chloro ethyl – trimethyl ammonium chloride), Etherel (200-
300 ppm) sprays one week before harvest hastens ripening.
32.5.2 Delay of Ripening

The self life of fruits like apple, banana and other can be improved by storing
the fruit in low oxygen tension (2-3%) or by absorbing ethylene with a suitable
absorbent like alumina or silica gel impregnated with potassium permanganate.
Another commercial practice is to preserve the fruits in cold storage. Maleic
Hydrazide, GA(10-6M), IAA (10-6M), Kinetin (10-5M), sprays one to two weeks before
harvesting and post harvest dip of cycocel, Alar, GA (150 ppm), vit K3 (menadione
Sodium bisulfite), KMnO4, CaCl2, waxol delays ripening.
32.5.3 Sugarcane Ripening

Glyphosine, glyphosate and ethephon hasten ripening of sugarcane. Other
compounds like chlormequat, mefluidide, Polaris and ripenthol also used for
sugarcane ripening.
32.5.4 USE OF HORMONES TO IMPROVE SHELFLIFE OF CUT FLOWERS
The use of preservative solutions to promote the quality and longevity of cut
flowers is known for many years. Flower preservatives are composed mainly of
sugars and germicides and some times include mineral solutes, organic acids, salts,
antioxidants and ethylene inhibitors. Use of hormones in preservation solution is
relatively limited. However cytokinins are widely used.
Sl
. Type of Optimum conc. And
Crop plants Nature of action
N compound purpose
o.
1 Cytokinins Carnations, rose,  100 ppm for pulsing  Ethylene

(kinetin, BA, iris, tulip, gerbera,  10-100 ppm for bud production is
IPA, PBA) anthurium opening & holding reduced.
solution  Burst in ethylene
production is
delayed.
 Improved water
uptake and
maintained
turgidity.
170

2 Auxin (2-4D) Carnation  500 ppm for holding  Inhibits ethylene

solution production
 Retards petal
senescence
3 GA3 Carnation  200 ppm holding  Delays
gladiolus solution, senescence
 20-35 ppm for bud
opening
4 ABA  1 ppm in continouos  Delays wilting by
holding solution stomatal closure
 10 ppm for 1 day
5 Growth Snap dragon  Snapdragon 10-50  Delays
retardants B- carnation roses ppm senescence
Nine  Carnation & roses –
500 ppm for pulsing
& 125 ppm for
holding solu
Chlormequat Gladiolus, tulip &  20-50 ppm for bud  Delays
carnation opening senescence
 10 ppm for holding
solution
6 Inhibitors Snap dragon,  0.5-1% for 30 min  Reduces
lupine for pulsing respiration of
Eg. MH  250-500 ppm for flowers
holding solution  Slows down
metabolism,
ageing &
development
*******
171

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Chief Editor : Dr.V.UMAMAHESH

Assistant Professor (Plant Physiology)

Agricultural College, Naira.

Co-Editors: Dr.K.Bala Krishna Reddy, Professor (Plant Physiology)

Dr.P.V. Rao, Professor (Plant Physiology)

Dr.A.Siva Sankar, Professor (Plant Physiology)

Dr.D.Vishnu Vardhan Reddy, Professor(Plant physiology)

Dr.K.L.Narasimha Rao, Professor (Plant Physiology)

Dr.V.Raja Rajeswari, Professor (Plant Physiology)

Dr. K. Ashoka Rani, Professor (Plant Physiology)

Dr. G. Rama Rao, Assoc.Professor(Plant Physiology)

Smt. V.Jayalalitha, Asst.professor (Plant physiology)

Dr.B.Srikanth, Asst.professor (Plant physiology)

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ACHARYA N.G. RANGA AGRICULTURAL UNIVERSITY

Detailed Lecture Outlines

1. Course No : PPHY 161

2. Course title : CROP PHYSIOLOGY

3. Credit hours : 3 (2+1)

4. General objective : To impart knowledge to the students on different plant

By the end of the course, the student will be able to

i) Study the growth and development of plants

iii) Understand the physiology of seeds and fruit ripening

By the end of the practical exercises, the students will be able to

i) Understand various plant metabolic processes, at different stages of plant

A) Theory Lecture Outlines

1. Introduction – definition of crop physiology – importance in agriculture and

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germination – morphological, physiological and biochemical changes during

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22. Photoperiodism and flowering – importance of photoperiodism – classification

1) Bidwell, R.G.S 1995, Plant Physiology, Macmillan Publishers Co., New

3) Frank, B. Salisbury and Cleon, W. Ross, 2005. Plant Physiology, CBS

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4) Gardener, P., Brent Pearce, R. and Roger, L. Mitchell, 1985. Physiology

5) Hopkins WG & Huner NPA. 2004. Introduction to Plant Physiology. John

6) Lincoln Taiz and Eduardo Zeiger 2006, Plant Physiology, Sinauer

7) Ray, G. Noggel and George, J. Fritz 1991. Introductory Plant Physiology.

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Internal processes & Plant growth & Development

Crop: is a group of plants grown as a community in a specific locality and, for a

1.1 Crop Physiology- Definition :

1.2 A BRIEF HISTORY OF CROP PHYSIOLOGY:

1771- Joseph priestly : Plants could regenerate oxygen in the atmosphere

1779- Ingenhouz : Studied the role of light in Photosyntheis

1865- Julius Sachs :published ‘Experimentelle Pflanzenphysiologi’ . By this time

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1963: Hesketh and Moss showed that photosynthesis by leaves of Maize,

Later on several research works were carried out to understand the

1.3 IMPORTANCE OF CROP PHYSIOLOGY IN AGRICULTURE AND HORTICULTURE:

Many aspects of Agriculture and Horticulture can benefit from more

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seedling growth, crop establishment, vegetative development, flowering, fruit and

 Seed is the most important input in agriculture. Germination of seed and

 Total amount of dry matter produced less the photosynthates used in

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 Nutriophysiology is yet another important area to under stand crop physiology.

Examples:Zinc deficiency leads to Khaira disease in Rice. This can be