Crop Science 1 Module1 3A

Introduction
Physiological processes are fundamental to the growth, development, and

productivity of crops. Understanding these processes is essential for farmers and
agronomists to optimize agricultural practices and improve crop yields. Physiological
processes allow farmers to implement targeted strategies, including proper irrigation,
nutrient application, and crop protection, to optimize crop production and enhance overall
agricultural sustainability.
This module presents the different physiological processes as occurs in plants, the
factors affecting the different processes and the plant characteristics that differentiate the
occurrence among the processes in plants. At the end of this unit the students should be
able to: Explain how the physiological processes affect crop production, concepts of
growth and development, phases of plant growth and development, plant movements and
crop adaptations, concepts related to plant growth. Students may also define
photosynthesis, respiration, transpiration and translocation and relate the physiological
processes occurring in plants in the formation/production of economic useful parts.
A. CONCEPTS OF GROWTH AND DEVELOPMENT
1. Growth
Growth is defined as an irreversible increase in the size of the organism.

Elongation of the roots and stem, leaf enlargement and increased in the size of flowers
and fruits are example of growth. Growth can be measured as increase in length, width,
area, volume, mass, or weight (either fresh or dry weight). Thus, it can be expressed
quantitatively. In crop science, dry weight is the more meaningful measure of growth.
The growth of plants involves both the production of new cells and their
subsequent enlargement. Growth serves not only to increase a plant’s size but also to
provide the plant with a limited means of movement and orientation for placing itself in a
more favorable position with regard to light, nutrients, reproduction, and dispersal.
There are two aspects of plant growth: primary and secondary. Primary growth
takes place in young organs resulting in an increase in length of shoots and roots.
Secondary growth follows primary growth in some plants and results in an increased girth
as layers of woody tissue are laid down. Monocots and herbaceous dicots typically exhibit
only primary growth. Woody dicots exhibit both primary and secondary growth (Fig. 1).
58 | C r o p S c i e n c e 1 ( P r i n c i p l e s o f C r o p P r o d u c t i o n )
Fig.1. Primary and secondary growth
The formation of new cells takes place in regions known as meristems. At the tip
or apex of each stem and root is the apical meristem, where cells are actively dividing
(Figure 2). Each apical meristem produces three other meristems (protoderm, ground
meristem, and procambium) called primary meristems. Tissues derived from the primary
meristems are called primary tissues and include epidermis, cortex, pith and primary
xylem and phloem (vascular tissues). The elongation of cells produced by the primary
meristem’s accounts for most of the increased length of stems and roots.
Fig. 2. Meristems and tissues
2. Development
Development or morphogenesis – is the series of changes by which an

individual plant or animal passes from a lower to a higher state of being or from embryonic
condition to maturity. It is a change in the complexity of the organism. For example, a
dividing cell resulting to many cells is development and a dividing tissue giving rise to
other tissues is also development. A change from a less complex to a more complex
being is the essence of development.
The orderly cycle of development that the whole plant undergoes involves complex
patterns of change in cells, tissues, and organs. The different phases or stages of
development are the following: (1) germination; (2) juvenility; (3) maturation; and (4)
senescence.
The cycle begins with seed germination and progresses through juvenility,
maturity, and flowering. Upon fruiting, the essential cycle of plant development is
completed. In perennials, the plant is ready to repeat the cycle after a rest period. In
annuals and biennials, fruiting is a signal to the organism to enter the final phases of plant
growth and development – senescence and death.
Table 1. Comparison of Plant and Animal Development
Plant Animal
Cell organelle and structure
Plants cells have chloroplasts for Animal cells do not have chloroplasts.
photosynthesis. Plant cells are Animal cells are round. Vacuoles in animal
rectangular. Vacuoles store water and cells store water, ions and waste.
maintain turgidity of the cell
Cell Movement
Plant cells are positionally fixed. Animal cell are motile.
Plant cells are trapped in rigid cell walls Animal tissues may be folded and moved
made of cellulose, which prevents against each other easily. At metazoan
movement of cells and tissues. Plants form gastrulation this way a triple layered
three basic tissue systems as well (dermal, system is built (ectoderm, mesoderm and
ground and vascular), yet without ectoderm). Some animal cells may even
gastrulation. move to other sites autonomously.
Rigidity of the body shape
Plant development is highly regulated The animal body plan is in most parts
by the environment clearly determined
As in most cases may not choose or The basic body plan of an animal during its
change its environment, it has to adapt to different life stages is mostly clearly
it. The body plan is variable and determined by its genes. If the
characterized by multiple times occurring, environment changes, they may react e.g.,
often iterative structures. Proportions and by moving to another place or changing
frequency of organs may vary their short- and long-term behavior.
Meiosis
Plants undergo no gametic meiosis, In animals’ gametes are formed directly
but a sporic meiosis through meiosis
In plants the meiosis produces spores and There is nothing that could be compared
not gametes. First the gametophyte is to the gametophyte in plants.
formed by mitotic divisions, which then
form the gametes
Plasticity
Plants show an enormous plasticity in Animal cells are determinate early in
their development development
If, for example, a shoot is nibbled by an When animal cells develop into tissues,
herbivore, axillary meristems often grow they are clean and, in most cases,
out to substitute for the lost part. This irreversibly determinate. While most
strategy resembles (limb) regeneration in tissues are regenerated from stem cells,
some animals. Whole plants can even be the regeneration of whole organs just
regenerated from single cells. occurs at some animal species like
Furthermore, the form of a plant (including Ambystoma mexicanum.
branching, height and relative portions of
vegetative and reproductive structures) is
strongly affected by environmental factors
such as light and temperature, resulting in
a great variety of morphologies from the
same genotyope. This amazing level of
plasticity helps plants compensate for their
lack of mobility
Morphogenesis
Plants go through a longer period of Animals develop to a distinct, complete
morphogenesis body shape
During their development plants do not During their life stages still, some
head for a distinct body plan. Many plants reorganization may take place, yet only in
just grow and develop on and on till they seldom cases new structures will develop.
die. Areas of actively dividing, Some animals develop stepwise into
undifferentiated cells, called meristems, different shapes.
allow for iterative growth and the formation
of more and more new organs and
structures during a plant’s life.
Cellular Basis of Growth & Development
1. Cell Division
Cell division and growth in plants are active in the following plant parts:
o the root tips,
o the stem tips,
o the cambium in dicotyledons, and
o the intercalary meristem in monocotyledons.
In a multicellular plant, cells undergo many changes as they develop from a
newly formed cell in a meristem to a functional mature cell. This differentiation
during or after enlargement forms cells highly specialized in size, shape, and
function.
Some cells complete all structural changes in a few days; others take much
longer. Some live for years; others die in a few hours depending on the plant.
2. Cell Enlargement
After mitotic division, the combined volume of the two new cells is about equal to
the volume of the parent cell.
The bulk of the increase comes in the vacuoles, which are small and
numerous in young cells, but larger and fewer in older ones.
Many cells increase as much as 500 times in volume as they age, with most of the
increase being due to water intake in the vacuoles.
Non-lignified plant tissues are supported by the pressure of cell contents against
the (primary) cell walls of their tissues. This turgor pressure is caused by the uptake of
water by the cytoplasm of the cells so that pressure is exerted at the plasma membrane
on the cell wall.
3. Cellular Differentiation
Differentiation is controlled by genotype, environment and its interaction. Once
the cell reached its final volume it becomes specialized.
Location of growth (meristem - apical, lateral, intercalary, cambium)
There are two aspects of plant growth: primary and secondary.
a) Primary growth takes place in young, herbaceous organs resulting in an increase

in length of shoots and roots.
b) Secondary growth follows primary growth in some plants and results in an increase
girth as layers of woody tissue are laid down.
Monocots and herbaceous dicots typically exhibit only primary growth.
Woody dicots exhibit both primary and secondary kind of growth.
Meristems and Growth

1. Primary Growth - growth in length that gives rise to primary (herbaceous) tissues
called the primary plant body.
Two Types
a) Apical meristem or apex - the growing points located at the tips of stems and
roots
b) Intercalary meristem - the growth region at the base of grass leaves which
causes leaves to elongate.
2. Secondary Growth - growth in width or diameter which gives rise to secondary
(woody or corky) tissues called the secondary plant body.
1. Lateral meristem - meristematic regions along the sides of stems and roots.
Two Types
a) vascular cambium or cambium - gives rise to secondary xylem (wood) on the
inside and phloem on the outside.
b) cork cambium or phellogen - gives rise to the periderm (bark).
Stem Anatomy
1. Monocot stems differ from dicot stems in that they lack secondary growth
2. Monocots has no vascular cambium nor cork cambium
3. Stems usually uniform in diameter
4. Scattered vascular bundles (not in a ring like dicot stems)
Types of Growth
1. Accretion. Growth of non-living organisms. Increases in size of a body of rock

or a landmass as a result of material accumulating on or around it. Ex. addition
of residues or deposits like stalagmites. (Salt springs)
2. Biological growth. Increases in size of living organism
2.1. Apoposition. Cell growth in which layers of material are deposited on

already existing ones. Ex. addition of secondary and tertiary walls to primary
walls.
2.2. Intussusception. Cell wall growth: the growth of the surface area of a cell
wall by the incorporation of particles into the wall.
Patterns of Growth
Plants are both axial and polar structures
1. An axial structure is symmetrical about a line or axis.
2. Polar structure. The opposite ends of the axis are different. Plants have many
axes. The main axis runs between the shoot and the root. Subsidiary axes
include branches and lateral roots.
The tip of a root is morphologically different from its base (Figure 14) and thus
shows polarity at the organ level. Within the root, polarity along files of cells can be seen
in the form of gradient of differentiation from apex to base.
Polarity can also exist at the level of individual cell. A mature root cap secrets
mucilage only across its outer wall (near the soil) and, thus forms different cell walls in
this same cell adjacent to the inside of root cap cells.
Polarity is established at the start of cell life. The first division of fertilized cell
(embedded in an ovule of a flower) determines the polarity of the plant (i.e., it determines
the axis and which end of it will give rise to the shoot or the root).
The growth of any one part of a plant always takes place in coordination with the
other parts. This is the correlative control growth. Important morphogenetic signals are
carried out by internal chemical messengers known as hormones for coordination and
regulation. It is clear that growth and development are controlled and influenced by three
general factors, environment (such as nutrition, light, temperature, water, etc), hormone,
and genetic factors.
B. PHASES OF PLANT GROWTH AND DEVELOPMENT
1. Phases of Growth
An idealized S-shaped (sigmoid) growth curve exhibited by numerous annual

plants and individual parts of both annual and perennial plants. The basic pattern of
growth in annuals is represented by an S-shaped curve, orSigmoid curve (Figure 3).
Fig. 3. S-shaped curve, or Sigmoid curve
Three phases of growth can be identified in the Sigmoid curve:
Phase I. Lag phase (Establishment and

seedling growth)
This phase relates to the

germination of the seed and seedling
growth. Crop seeds that germinate below
ground are dependent on the stored
materials in the cotyledons or endosperm
until the seedlings emerge into the light
and photosynthesis starts. In theseearly
stages, the dry weight may decrease as
stored materials are respired or broken
down. No growth has occurred in this
period but the tissues have undergone
differentiation into roots, leaves and
stem, and that development has occurred
at the expense of growth. Once the
expanded or first true leaves develop
chlorophyll, photosynthetic increases in
dry matter exceeds respiratory losses
and growth proceeds rapidly.
Phase II. Log/Exponential Figure 4. A nearly ideal sigmoid growth curve (a) and
(Period of rapid growth, stem elongation bell-shaped growth rate curve (b) for maize.
and flowering)
This phase of growth is characterized by a rapid and often linear increase in dry
matterproduction. This is associated with vegetative growth in annuals and terminates
with theonset of flowering. In cereals, the early part of this phase is associated with the
production of tillers and leaves. As the final leaf or flag leaf expands, the rate of stem
elongation is reduced and the emergence of the previously formed inflorescence occurs.
During this phase of growth, the root system increases in size.
Phase III. Stationary phase (Ripening and Senescence)
This phase is marked by a reduction in growth rate until growth stops at crop
maturity. In cereals, the seeds begin to develop and the growth of stems and leaves stops
after flowering. During this period, re-translocation of assimilates stored in the leaves and
stems occur to partially sustain seed growth. At the end of the growth period,water is lost
from the aerial parts of the plants, photosynthesis stops, and the crop beginsto senesce.
This pattern of growth is exhibited not only by the whole plant but also by any partof the
plant.
2. Phases of Development
The orderly cycle of development that the whole plant undergoes involves complex
patterns of change in cells, tissues, and organs. The different phases or stages of
development are the following: (1) germination; (2) juvenility; (3) maturation; and (4)
senescence.
The cycle begins with seed germination and progresses through juvenility,
maturity, and flowering. Upon fruiting, the essential cycle of plant development is
completed. In perennials, the plant is ready to repeat the cycle after a rest period. In
annuals and biennials, fruiting is a signal to the organism to enter the final phases of plant
growth and development – senescence and death.
A. Germination Phase
Seed germination is defined as the emergence and development from the seed
embryo of those essential structures that indicate the ability of the seed to develop into a
normal plant under favorable conditions. The germination phase begins with imbibition of
water and ends when the seedling is self-sustaining.
The major events occurring in germination are: water imbibition, enzyme

activation; initiation of embryo growth; rupture of the seed coat and emergence of
the seedling; and finally, seedling establishment.
Fig. 3. The germination phase
Water Imbibition
Water is absorbed by the seed through natural openings and the seed coat. The
seed swells and the seed coat often ruptures, facilitating both water and gas uptake, and
emergence of the growing points later (Figure 4).
Fig. 4. Water imbibition
Enzyme Activation
Enzymes are synthesized or activated.

Complex reserve substances are enzymatically
converted to simple forms and translocated to
the growing points. Some substances undergo
respiratory breakdown and release energy;
others are used to synthesize compounds
needed for growth (Figure 5).
Enzyme activation refers to the process

by which an enzyme becomes catalytically
active, allowing it to facilitate specific chemical
reactions. Enzymes are biological catalysts that
accelerate chemical reactions in living
organisms by lowering the activation energy
required for the reaction to occur.
Fig. 5. Enzyme activation
Initiation of Embryo Growth
The synthesis of new

materials results in an
increase the size of the root-
shoot axis (epicotyl,
hypocotyl, radicle) (Figure
6). The initiation of embryo
growth is a critical step in the
development of a new plant.
It involves the activation of
processes that lead to the
formation and expansion of
embryonic tissues,
eventually resulting in the
germination of seeds. The
initiation of embryo growth is
a tightly regulated and
coordinated process that
ensures the successful
transition from a dormant
seed to an actively growing Fig. 6. Initiation of embryo growth
and metabolically active
seedling. Environmental factors such as temperature, water availability, and light play
crucial roles in influencing the timing and success of this process.
Seedling Establishment
This is the transition when the seedling produces part of its food requirement and
is partly dependent on reserve food. When the seedling is totally independent, that is, it
produces its entire food requirement, the germination phase is complete.
Three conditions must be fulfilled before germination can occur. First, the seed
must be viable, that is, the seed is alive and capable of germination. Second, internal
conditions of the seed must be favorable for germination, that is, physical and chemical
barriers to germination must have disappeared. Third, the seed must be subjected to
appropriate environmental conditions. The essential requirements are water, favorable
temperature, oxygen, and for some species (e.g., lettuce), light.
A seed, although viable, may not germinate. The inability of a viable seed to
germinate even when subjected to favorable environmental conditions is called seed
dormancy.
Seed dormancy can be caused by several reasons:
Causes of seed dormancy Methods of breaking dormancy
A. Impermeable seed coat: 1. Mechanical scarification by:
a. to water (Examples: legumes,) a. abrasion - break the seed coat so that water
b. thick seedcoat (Examples: mango, can be absorbed by the seed
ampalaya b. impaction
c. to oxygen (Example: grasses) c. combination of the two
2. Acid scarification by sulfuric acid treatment
3. Dissolve the seed coat by:

a. hot water
b. wax solvent - acetone, alcohol
4. Dry heat at 60-80°C for one-half to two

hours. Dry heat causes swelling of tissues
5. Extreme cold (-50°C to -150°C). Contraction

of tissues at very low temperatures breaks
the seed coat
6. Use of 0.2% KNO3 solution

B. Immature embryo. Some seeds are 1. Requires after-ripening. This is affected by
shed before the embryo is mature storage
C. Presence of inhibitor: 1. Overcome by washing in water
- seedling growth inhibitor in rice 2. Gibberellic acid treatment
- inhibitor in barley seed, tomato
D. Light sensitivity (Example: tobacco, 1. Exposure to light
lettuce)
Water is essential for enzyme activation and the breakdown, translocation, and
use of reserve storage material. For germination to proceed, the seed must be able to
absorb moisture until it reaches the critical moisture content, which differs with species.
For example, the critical moisture content of corn is 30.5%; rice, 32-35%; and peanut, 50-
55%.
The response to temperature depends on species, variety, growing region, and

duration of time from harvest. For example, the cardinal temperatures
(minimum/optimum/maximum) for the germination of rice seeds are 10-12oC/30- 37oC/40-
42oC; for soybean, 8oC/32oC/40oC; and for corn, 8-10oC/32-35oC/40-44oC. As a general
rule, temperate region seeds require lower temperatures than do tropical region seeds.
Respiration increases sharply during seed germination. Since respiration is

essentially an oxidative process, an adequate supply of oxygen must be available.
Atmospheric air contains almost 21% oxygen. At oxygen concentration substantially
below 20%, germination of most seeds is retarded. There are seeds, such as rice, that
can germinate in the absence of oxygen; anaerobic respiration enables them to
germinate.
Light is required by certain species (e.g., lettuce). If high temperature induces

dormancy, high light intensities will erase that effect.
Two types of germination occur. In epigeous or epigeal germination, the

hypocotyl elongates and raises the cotyledons above the ground. In hypogenous or
hypogeal germination, the lengthening of the epicotyl, or in the case of grasses, the
mesocotyl, does not raise the cotyledon above the ground, and only the epicotyl emerges
(Figure 7).
Fig. 7. Epigeal and hypogeal germination
Good germination is the foundation of a good crop. It is a requirement, although

it is not sufficient, for the attainment of the desired yield. Ideally, there must be high, fast,
and uniform seed germination.
B. Juvenile Phase
A seed is considered germinated when it has produced a plant that, under proper
environmental conditions, is potentially capable of continuous and uninterrupted growth.
From the time this stage is reached until the first flower primordium is initiated, the plant
is considered to be in the vegetative phase of growth. If during this vegetative phase the
plant cannot be made to flower, regardless of the environmental conditions imposed, it is
said to be juvenile.
The juvenile phase is characterized by the most rapid rate of plant growth. It is
devoted exclusively to rapid vegetative growth in which the plant achieves a size that will
be competitively stronger in the plant community. In this phase, the plant develops
characteristic morphological forms of leaves, stems, and roots. It is therefore important in
the formation of the leaf area that will support yield.
As the juvenile plant grows up, structural differences along its stem axis may reflect
the gradual change from juvenile to mature types of growth. One of the most readily
observed morphological expressions of juvenility is leaf form. Among the Angiosperms,
juvenile leaves may be simple and mature leaves compound (bean). On rare occasions,
the reverse is true. In the pea, the juvenile leaves are reduced to scales. Increasing
amounts of dissection or lobing of the leaves is often associated with increasing maturity.
This has been described for cotton as a changing extent of leaf lobing during the
development of the leaf. The cotton seedlings produce a simple entire leaf, and as the
plant becomes mature, the leaf form gradually becomes palmate. After the plant has
started to fruit, the leaves tend to return to the entire shape. Ideally, one might find an
entire series from juvenile to mature and senescent leaf forms arrayed from the base to
the tip of the cotton plant.
The end of the juvenile phase is indicated when the plant initiates the flower
primordium or responds to flower-inducing stimuli.
C. Maturity Phase
A plant is mature if it is potentially capable of reproduction, that is, flowering. The

mature plant, although capable, may not necessarily flower. The environment to which
the plant is exposed at the time of maturity determines if the plant will reach this ultimate
expression of the mature state.
Juvenility and maturity are relative terms. In many species, these growth phases
blend into each other, a part of the plant juvenile, a part mature.
Flowering
The first event, the most critical, in flowering is the transformation of the stem
primordia from the vegetative state to the reproductive state. This transformation is
irreversible, and the flower parts will continue to develop until anthesis (the time at which
the flower is fully open) even though environmental conditions change (Figure 8).
Fig. 8. Flowering
Two conditions are required for a plant to initiate the flower: (a) the juvenile phase
has been completed; and (b) the required environmental stimulus has been supplied. For
certain plants, only the first condition needs to be satisfied. Once they finish the juvenile
phase, they will initiate the flower. For others, both conditions have to be satisfied.
The two environmental stimuli required for the flowering of some plants are
daylength (or photoperiod) and low temperature.
1. Daylength. The response of plants to daylength is called photoperiodism. In spite

of the name, the length of the dark period is the critical factor in the photoperiodic
response. There are three categories of plants based on photoperiodism:
a. short-day plants – require a dark period exceeding the critical night length in order
to flower. Example, poinsettia, rice, winged bean
b. long-day plants – inhibited from flowering when the dark period is shorter than
the critical night length, and they can flower under continuous illumination.
Example, winter wheat.
c. day-neutral plants – can initiate the flower under any night length. Most plants
are day-neutral plants
Short-day plants may be obligate (or qualitative) short-day plants or facultative (or
quantitative) short-day plants. Qualitative short-day plants absolutely require the proper
night length for them to flower. If they are not subjected to the proper night length, theywill
not flower. Quantitative short-day plants will flower even when the proper night length is
not supplied, but the flowers are few. The same is true with long-day plants (Figure 9).
Fig. 9. Difference between short-day to long-day plants
2. Low temperature. Some plants require exposure to low temperature (around 0-

10oC), a process called vernalization, to initiate the flower. In temperate countries,
this happens naturally during winter.
Senescence
The deteriorative processes which naturally terminate the functional life of a cell,
tissue, organ, or organism is called senescence. It begins at full maturity and ends in
death. In some plants, this occurs as the gradual encroachment of deteriorative
processes and in other plants there may be fairly abrupt deterioration leading to death.
Chlorophyll degradation, photosynthetic deterioration of leaves, changes in amounts of
key metabolites (e.g., protein breakdown), and alterations in cell walls and membrane
permeability are senescence phenomena.
There are two general types of senescence. Partial senescence refers to the
deterioration and death of plant organs, such as leaves, stems, flowers, and fruits. Overall
or complete senescence refers to the deterioration and death of the entire plant, except
for the seeds. In annual grains, the entire plant dies by some systematic function. The
senescence of perennials appears as a gradual erosion of growth and viability. In
perennial herbs, the above-ground portion may die, but the root system and underground
stem remains viable. The death of the whole plant commonly occurs in annual and
biennial species upon completion of fruiting.
Senescence is initiated by flowering and accentuated by fruiting. Removal of the

flowers and fruits of several species defers or prevents senescence.
In crop production, it is sometimes desirable to delay senescence to increase yield.

This lengthens the period of photosynthetic activity of the leaves. In cereals and annual
legumes, senescence can be delayed by foliar application of nitrogen at flowering.
C. PLANT MOVEMENTS/CROP ADAPTATION
Plant Movements
Plants exhibit various movements and adaptations in response to environmental

stimuli. These movements and adaptations are critical for their survival and reproduction.
Plant movements are essential adaptive responses that allow plants to interact
with their environment, optimize resource acquisition, and respond to various stimuli.
These movements, often slow and non-directional, are crucial for the survival and
reproductive success of plants. Here's a discussion on different types of plant movements
and their significance:
1. Tropisms
• Phototropism. The growth or movement of a plant towards or away from light.

This ensures that leaves and stems are positioned optimally for photosynthesis.
• Gravitropism. Growth in response to gravity. Roots show positive gravitropism by
growing downward, while shoots exhibit negative gravitropism by growing against
gravity.
2. Nastic Movements
• Thigmonasty: Non-directional movement in response to touch or mechanical

stimulation. Examples include the folding of leaves in the sensitive plant (Mimosa
pudica).
• Nyctinasty: Daily rhythmic movements in response to light or temperature
changes. For example, the opening and closing of certain flowers during the day
and night.
3. Circadian Rhythms
• Plants exhibit daily rhythms in physiological and biochemical processes,

influencing activities like stomatal opening and closure, nutrient uptake, and flower
opening. These rhythms are important for plant growth and adaptation to
environmental changes.
4. Heliotropism
• Movement or growth of plant parts in response to the direction of the sun.

Sunflowers are well-known for following the sun's movement throughout the day,
maximizing light absorption for photosynthesis.
5. Hydrotropism
• Growth or movement of plant roots towards or away from water. This helps roots
explore the soil for moisture, ensuring optimal water uptake.
6. Chemotropism
• Growth or movement of plant parts in response to chemical gradients. This is

particularly relevant in the growth of pollen tubes towards ovules during pollination.
7. Photonasty
• Movement in response to light intensity. Examples include the opening of flowers

in response to sunlight.
Significance and Adaptive Advantages
1. Resource Optimization
• Plant movements help optimize resource acquisition by ensuring exposure to
sunlight for photosynthesis, efficient nutrient uptake, and exploration of the soil for
water.
2. Defense Mechanisms
• Nastic movements, such as thigmonasty, can act as defense mechanisms against
herbivores by making the plant less palatable or accessible.
3. Reproductive Success
• Movements related to flower opening and pollination enhance the chances of
successful reproduction by facilitating interactions with pollinators and optimizing
fertilization.
4. Environmental Adaptation
• Circadian rhythms and responses to environmental factors like temperature, light,
and humidity enable plants to adapt to changing conditions and survive in diverse
habitats.
5. Survival Strategies
• Tropisms and other movements represent adaptive strategies that allow plants to
respond dynamically to their surroundings, enhancing their overall survival in a
competitive environment.
Understanding plant movements is crucial not only for basic botanical knowledge but
also for agricultural practices. Manipulating these movements can have applications in
optimizing crop growth, improving yield, and enhancing stress tolerance in various
environmental conditions.
Crop Adaptations
Crop adaptations refer to the evolutionary or agricultural modifications that enable

plants to thrive in specific environmental conditions or under certain cultivation practices.
These adaptations can involve a range of characteristics, from physiological and
morphological features to genetic traits that enhance the crop's resilience, productivity,
and overall performance
1. Drought Tolerance
• Deep root systems, reduced transpiration rates, and mechanisms for water
retention.
• Crop Examples: Drought-tolerant varieties of maize, sorghum, and millet have
been developed with enhanced water-use efficiency.
2. Heat and Cold Tolerance

• Heat-resistant enzymes, protective structures, and the ability to adjust metabolic
rates.
• Crop Examples: Varieties of wheat, rice, and fruits bred for heat or cold tolerance
to withstand extreme temperature fluctuations.
3. Salinity Tolerance
• Mechanisms to exclude or tolerate high salt concentrations, such as salt glands or
ion-exclusion.
• Crop Examples: Salt-tolerant varieties of rice, barley, and certain vegetables
designed for cultivation in saline soils.
4. Pest Resistance
• Production of toxic compounds, attractant-repellant strategies, and symbiotic
relationships with beneficial organisms.
• Crop Examples: Genetically modified (GM) crops with resistance to specific pests,
like Bt cotton expressing a toxin against certain insect pests.
5. Pathogen Resistance
• Production of secondary metabolites, disease resistance genes, and physical
barriers.
• Crop Examples: Breeding for disease-resistant varieties, such as wheat with
resistance to rust pathogens.
Significance and Future Directions
1. Sustainable Agriculture
• Crop adaptations contribute to sustainable agriculture by reducing the reliance on
chemical inputs and minimizing environmental impact.
2. Global Food Security
• Developing crops with diverse adaptations ensures food security by expanding
cultivation into diverse environments and climates.
3. Genetic Engineering
• Advances in biotechnology enable the introduction of specific adaptive traits
through genetic engineering, providing tools for precision breeding.
4. Climate Change Resilience

• Crop adaptations are crucial in addressing challenges posed by climate change,
helping agriculture withstand unpredictable weather patterns and extremes.
5. Biodiversity Conservation
• Preserving and utilizing crop adaptations contribute to the conservation of genetic
diversity, ensuring the availability of resources for future breeding efforts.
Understanding and harnessing crop adaptations are essential for sustainable and
resilient agriculture, ensuring that crops can withstand various environmental stresses
and meet the increasing demand for food in a changing world. Advances in breeding
techniques, genetic engineering, and precision agriculture continue to play key roles in
enhancing crop adaptations for the future.
D. OTHER CONCEPTS RELATED TO PLANT GROWTH
1. The Law of the Minimum
In his book, published in Germany

in 1840 and translated as Organic
Chemistry in its Applications to Agriculture
and Physiology, Justus Liebig formulated
his law of the minimum. The law states:
“The growth of a plant is dependent upon
the amount of “foodstuff” presented to it in
minimum quantities.” Applied to crop
production, it means yield is determined by
a minimum factor, i.e., a factor that is
insufficiently available. The factor might be
one or more of many things besides
mineral nutrients: water, damage by pests
(diseases, insects), competition from
weeds, CO2 concentration, or the plant’s
genes (Fig. 10).
The “Law of the Minimum” states

that growth is controlled by the scarcest Fig. 10. Law of Minimum
resource (limiting factor). This law is usually believed to be the result of Justus von
Liebig’s research (1840) but the agronomist and chemist Carl Sprengel published in 1828
an article that contained in essence the Law of the Minimum and this law can be called
the Sprengel–Liebig Law of the Minimum.
This law is also known as the “barrel” concept. A wooden barrel is composed of
vertical staves. If the staves are of different lengths, the shortest one determines the
capacity of the barrel, which states that if one of the essential plant nutrients is deficient,
plant growth will be poor even when all other essential nutrients are abundant.
It states that growth is controlled not by the total originality applied to plant growth,
where it was found that increasing the amount of plentiful nutrients did not increase plant
growth. Only by increasing the amount of the limiting nutrient (the most scarce) in relation
to “need”, was the growth of the plant improved.
The Law of the Minimum takes on added importance when fertilizer prices —
especially of nitrogen (N) and phosphate (P2O5) products — are high. This may tempt
some growers to reduce or even eliminate applications of micronutrient or secondary
nutrient fertilizers that provide balanced potassium (K), magnesium (Mg) and sulfur (S).
But von Liebig’s “Law” tells us clearly that if a soil is deficient in, say, Mg, yields will be
depressed regardless of how much N-P-K product you apply.
2. The Law of Optima and Limiting Factors
F.E. Blackman (1905) in

England proposed the term
limiting factor for that “foodstuff
presented … in minimum
quantities.” Example, if plant yield
is limited by insufficient amounts
of nitrogen, then more nitrogen is
applied. When nitrogen has been
applied, then perhaps
phosphorus becomes limiting
and needs to be applied. The
response is linear (Figure 2).
The “law of the minimum”

proved to be unsatisfactory, Fig. 2. Graph illustrating Blackman’s law of limiting factor
because experiments showed
that yield does not increase linearly with increased application (dosage) of the minimum
factor. The minimum law was therefore modified to the “optimum law” (Liebscher, 1895).
The minimum or limiting factor affects yield more the more the other factors approach
their optimum dosage.
Photosynthesis, for example, is a biological process that depends on multiple
factors. The general chemical reaction of photosynthesis is
6CO2+12H2O+energy=C6H12O6+6O2+6H2O. Based on this equation, CO2, H2O, and light
energy (sunlight) are the limiting factors of this reaction. If any of them become accessible
at a pace slower or lower than usual, the rate of photosynthesis is expected to become
slow based on the pace of the slowest factor.
For example, if CO2 concentration becomes scarce (e.g., due to closure

of stomatal openings in response to elevated temperatures in the environment), the rate
of photosynthesis becomes slow even if H2O and light energy levels are amply available.
The same result will occur if light energy becomes less available or less intense,
the rate of photosynthesis will be slower despite the abundance of CO2 and H2O. Light
becomes a limiting factor in photosynthesis when the plant is unable to collect light, for
instance, due to shade resulting from the dense population of plants.
3. The Law of Diminishing Returns
Finally,
Mitscherlich (1906)
formulated the law of
diminishing returns, also
known as the law of
decreasing productivity.
Yield depends on each
growth factor (now
appropriately called
production factor) with a
factor-specific production
coefficient. The increase
in yield per increase in
growth factor is
proportional to the
difference between the
actual yield and the
maximum yield. The
response is therefore
curvilinear, not linear as in
the law of the minimum or
limiting factor.
In this formulation, two important aspects are stressed: (1) Every production factor
potentially limits yield; and (2) an increase in each production factor increases the yield if
the optimum has not been reached or surpassed. Mitscherlich’s law can be formulated
for a certain growth factor (experimental factor), e.g., for a macroelement, as follows:
∆y/∆x = c (A-y)
Where A is the maximum yield (experimental factor optimal), y is the actual yield
(experimental factor suboptimal), A-y is the difference compared with the maximum yield,
x is the dosage of experimental factor and c is the production coefficient. The production
coefficient is an empirical parameter which can be determined comparatively, for example
at given conditions of soil and climatic factors, for macroelements.
On the other hand, Mitscherlich’s Law of Diminishing Returns states a decreasing

marginal productivity as levels of the limiting factor are raised. According to this law, plants
cannot grow indefinitely and there is a maximum of production. This law assumes that
returns from fertilization are proportional to the difference between maximum and current
productions, in such a way that the returns tend to zero as production approaches its
maximum value.
The Law of Diminishing Returns, states that if an increasing of a variable factor are
applied to a fixed quantity of other factors per unit of time, the increments in total output
will first increase but beyond some point, it begins to decline.
E. PLANT LIFE PROCESSES
Plant life processes encompass a variety of essential activities that occur within
plants to maintain life, sustain growth, and ensure reproductive success. These
processes are crucial for the survival of plants and contribute to the overall functioning of
ecosystems.
1. PHOTOSYNTHESIS
Photosynthesis is a biochemical process that serves as the connecting link

between solar energy and life on earth. It is a constructive (anabolic) process by which
food of the plant is manufactured from simple inorganic materials through the agency of
chlorophyll; the energy needed is derived from sunlight. In this process, solar energy is
captured and converted into chemical energy stored in the form of carbohydrates. It is the
process where carbon dioxide (CO2) is fixed by the plants. The products of
photosynthesis are sources of energy and building materials for plants and animals. They
are also sources of industrial materials. Approximately 90% of the human population is
engaged directly or indirectly in the production, processing, and sale of the products and
by-products of photosynthesis.
Photosynthesis is a complex process and manufacture of sugars and its

precursors by green plants in the presence of light and chlorophyll. The summary
chemical equation is as follows:
6CO2 + 12H2O ---→ C6H12O6 + 6O2 + 6H2O
Fig. 14. Biosynthesis of Photosynthesis
Chloroplast
Are usually lens-shaped bounded by a

double membrane
The inner membrane invaginates parallel
to the surface and become organized into
specialized cytoplasmic body consisting
of a stack thylakoid called granum’s
which are embedded in a proteinaceous
matrix called the stroma.
It is the photosynthetically active
organelle of a plant cell.
It transforms solar radiation into chemical
forms of energy.
Inside the chloroplast are two adjacent
membranes fused along their
peripheries, forming disk-shaped
compartments called thylakoids.
Chlorophyll
Chlorophyll is a green
photosynthetic pigment
found in plants, algae, and
cyanobacteria.
Chlorophyll absorbs mostly
in the blue and to lesser
extent red portions of the
electromagnetic spectrum,
hence its intense green
color.
Green substance in
producers that traps light
energy from the sun, which is then used to combine carbon dioxide and water into
sugars in the process of photosynthesis Chlorophyll is vital for photosynthesis,
which helps plants get energy from light.
Chlorophyll molecules are specifically arranged in and around pigment protein
complexes called photosystems, which are embedded in the thylakoid membranes
of chloroplasts.
In addition to giving plants their green color, chlorophyll is vital for photosynthesis
as it helps to channel the energy of sunlight into chemical energy.
With photosynthesis, chlorophyll absorbs energy and then transforms water and
carbon dioxide into oxygen and carbohydrates.
The process of photosynthesis converts solar energy into a usable form for plants,
and the animals that eat them, forming the foundation of some food chains.
The process can be understood in terms of three partial steps as follows:
1. Diffusion of carbon dioxide to the chloroplast. Carbon dioxide in the air is

transported by turbulence or diffusion to the leaf stomata through which it diffuses
to the chloroplast.
2. Light reactions or photochemical reactions. In these reactions, light is

essential, while temperature is unimportant. They take place in the thylakoid
membranes of the chloroplast. Light energy is used to split water producing
molecular oxygen (O2), reduced nicotinamide adenine dinucleotide phosphate
(NADPH) and adenosine triphosphate (ATP). The light reactions can be divided
into three components:
a. Photolysis of water. Water is split into hydrogen ions (H+), electrons (e-) and
molecular oxygen (O2).
2H2O + light 4H+ + 4e- + O2
b. Reduction of NADP+ to NADPH. The electrons from water are transferred

through the photosynthetic electron transfer system to NADP+ to form NADPH.
NADP+ + H+ + 2e- + light NADPH
c. Photophosphorylation. Light and inorganic phosphate (Pi) are used to

convert adenosine diphosphate (ADP) to ATP.
ADP + Pi + light ATP
Based on the Adaptation of Crop Production
A. Classification of plants on the basis of temperature adaptation

Cool season or temperate crops. Those plants such as wheat, cauliflower, carrot
and sugar beet prefer a monthly temperature between 15ºC and 18ºC for growth
and development.
Warm season crops or tropical plants. Those plants adapted to tropical belt such
as corn, sugarcane, rice and sorghum require warm temperature of between 18ºC
and 27ºC during the growing season.
B. According to growing season

Dry season crops. Those that are suited under dry conditions provided irrigation is
supplied to them.
Wet season crops. Crops that can tolerate excess soil moisture during their
growing period.
Off-season crops. Crops that are grown out of their normal growing season usually
under enclosed protective structures (green house) during unfavorable weather
conditions.
C. Response to light
Sciophytes (shade loving plants). Plants that prefer shady conditions, e.g.,
some orchids
Heliophytes (sun loving plants). Plant that like sunlight or a plant can survive
and grow under direct sunlight or that grow best in direct sunlight, example corn,
peanut, field beans, and rice.
Facultative sciophytes (sun loving plants that can tolerate shade). Plants that
grown best under direct sunlight but can tolerate shady condition.
D. According to light requirement
The fixation or reduction of CO2 into carbohydrates can occur via three pathways:
1. Carbon-fixing (Dark reactions or biochemical reactions). In these reactions,

light is not required, but temperature plays an important role. The products of the
light reactions (NADPH and ATP) are used to fix carbon dioxide. The CO2 fixation
can occur by one of three pathways: (a) the Calvin-Benson or C3 pathway; (b) the
Hatch and Slack or C4 pathway; and (c) the Crassulacean Acid Metabolism or
CAM pathway.
In the C3 (C3 (Calvin-Benson Cycle/Reductive Pentose) pathway,

ribulose bisphosphate (RUBP) is the CO2 acceptor. 3-phosphoglyceric acid (PGA)
is the first stable product. The name of the pathway came from the fact that the
first stable product (PGA) has three carbon atoms. The Calvin-Benson cycle also
occurs in C4 plants and CAM plants as the last stage of the CO2-fixation process.
Plants that require one-third full
sunlight requirement (15ºC -
25ºC).
Fixation and reduction of one
molecule of CO2 requires three
molecules of ATP and 2 NADPH
(coming from light reaction)
Occurs in the mesophyll cell
chloroplast
CO2 acceptor is RUBP
RUBP carboxylase enzyme is
needed
The first product is 3-PGA
Example: rice, most vegetable
species, peas, beet and wheat.
2. C4 or Hatch Slack Pathway. In the C4 pathway, phosphoenol pyruvic acid (PEP)

is the initial CO2 acceptor and the first stable product is oxaloacetic acid (OAA),
which is converted to either malic acid or aspartic acid. OAA has four carbon
atoms, hence the name of the pathway. It is also called the Hatch and Slack
pathway to honor the two scientists who elucidated the pathway. The Calvin-
Benson-cycle part of the pathway occurs in the bundle-sheath cells, while the initial
fixation of CO2 occurs in the mesophyll cells. C4 photosynthesis is a series of
biochemical and anatomical modifications that concentrate carbon dioxide (CO2)
around the carboxylating enzyme Rubisco. This increases photosynthetic
efficiency in conditions that promote high rates of photorespiration.
C4 pathway requires the presence of two types of photosynthetic cells, i.e.,

mesophyll cells and bundle sheath cells.
The bundle sheath cells are arranged in a wreath like manner.
This kind of arrangement of cells is called Kranz anatomy (Kranz: wreath).
In Kranz anatomy, the mesophyll and bundle sheath cells are connected by
plasmodesmata or cytoplasmic bridges.
The C4 plants contain dimorphic chloroplasts. The chloroplasts in mesophyll
cells are granal, whereas in bundle sheath cells they are agranal.
The granal chloroplasts contain thylakoids which are stacked to form grana, as
formed in C3 plants. However, in agranal chloroplasts of bundle sheath cells
grana are absent and thylakoids are present only as stroma lamellae.
The presence of two types of cells (granal and agranal) allows occurrence of
light and carbon (dark) reactions separately in each type.
Here, release of O2 takes place in one type, while fixation of CO2 catalyzed by
Rubisco enzyme occurs in another type of cells.
In the C4 pathway, phosphoenol pyruvic acid (PEP) is the initial CO2
acceptor and the first stable product is oxaloacetic acid (OAA), which is
converted to either malic acid or aspartic acid. OAA has four carbon atoms,
hence the nameof the pathway. It is also called the Hatch and Slack pathway to
honor the two scientists who elucidated the pathway. The Calvin-Benson-cycle
part of thepathway occurs in the bundle-sheath cells, while the initial fixation of
CO2 occurs in the mesophyll cells.
3. CAM (Crassulacean Acid Metabolism) plants (35-45ºC 1/10th of full sunlight –
mesophilic plants highly adapted to arid environments. Plant whose stomates are
close during the day and open during the night. Example: Most cacti, pineapple,
agave, most orchids.
In the CAM pathway, CO2 is fixed at night because the stomata of CAM
plants are closed during the day and open at night as a way of conserving moisture.
It has many reactions similar to those of the C4 pathway, but the Calvin- Benson
cycle is separated from the rest of the photosynthetic pathway in time – it occurs
during the day while the initial fixation of CO2 occurs during the night. Based on
the photosynthetic pathway followed, plants are classified as C3 plants, C4 plants,
and CAM plants.
CAM – short for “Crassulacean Acid Metabolism” – is a method of carbon
fixation evolved by some plants in dry circumstances.
In most plants, the stomata – which are like tiny mouths that take in oxygen
all along the surfaces of their leaves – open during the day to take in
CO2 and release O2.
Plants must take in CO2 because they use it as a source for carbon atoms
to build sugars, proteins, nucleotides, and the other building blocks of life.
They must also release waste O2, which is the bi product that is left over
after the carbon atom from has CO2 been incorporated into a sugar.
Most plants open their stomata during the day because that is when energy
is received from the Sun. The energy from the Sun is harvested by the
chloroplasts and used to make ATP and NADPH. These short-term energy
storage molecules are then used to power the fixation of carbon into sugar.
In plants living in very dry environments, however, dangerous amounts of
water can be lost if the stomata are open during the hot, dry days. During
the night, which tends to be much cooler in dry environments, far less water
is lost by opening the stomata.
In order to meet their needs to combine the Sun’s energy with CO2 from the
air, CAM plants take in CO2 at night and store it in the form of a four-carbon
acid called “malate.” Then the malate is released during the day, where it
can be combined with the ATP and NADPH created by the Sun’s energy.
This allows the plants to conserve their water by closing their stomata during
the hot daytimes.
The name “Crassulacean Acid Metabolism” comes from the Crassula plant,
which was the first place that CAM metabolism was discovered and studied.
Steps of CAM Photosynthesis
1. CAM photosynthesis begins at night, when the plant’s stomata open and CO2 gas
is able to diffuse into the cytoplasm of CAM mesophyll cells.
In the cytoplasm of those cells, the CO2 molecules encounter hydroxyl ions, OH−,
which they combine with to become HCO3 the enzyme phosphoenolpyruvate
carboxylase (PEP carboxylase).
CO2 + OH− → HCO3
2. The PEP carboxylase enzyme catalyzes the following reaction to add the CO2 to
a molecule called phosphoenolpyruvate (PEP).
PEP + HCO3− → OXALOACETATE
3. Oxaloacetate then receives an electron from NADH and becomes a molecule of

malate. This reaction is catalyzed by the enzyme Malate Dehydrogenase (MDH).
That reaction looks like:
OXALOACETATE + NADPH + MDH → MALATE + NADP+
Interestingly, malate dehydrogenase catalyzes a reversible reaction, meaning that

it can either add electrons to oxaloacetate, or take electrons away from molecules
of malate.
4. Malate is now stored in vacuoles within the plant cells, until the sun rises and
photosynthesis begins. When that happens, malate enters the Calvin Cycle, just
like 3-phosphoglycerate would in a plant using a 3-carbon, or “C3” pathway for
carbon fixation.
CAM cycle
In this pathway, stomata open at night, which allows CO2 to diffuse into the leaf to
be combined with PEP and form malate. This acid is then stored in large central vacuoles
until daytime. During the day, malate is released from the vacuoles and decarboxylated.
Table 1 shows a comparison of the three types of plants.
Characteristics C3 plants C4 plants CAM plants
Typically, temperate species,

e.g., spinach, wheat, potato, Typically, tropical or
tobacco, sugar beet, soy semi-tropical species, Typically, arid zone
1. Plant species bean, sunflower, etc., and e.g., corn, sorghum, species, e.g., cacti,
following the tropical species, e.g., rice, sugarcane,amaranth, Agave, bromeliadssuch
pathway taro, sweet potato, cassava, plants adapted to high as pineapple,most
mungbean, light intensity and orchids, other
etc. Most plants areC3 plants. temperature. succulents.
Mesophyll withdistinct
Mesophyll with nodistinct Mesophyll with large
2. Leaf anatomy bundlesheath (Kranz
bundle sheath vacuoles.
anatomy).
3. Energy
requirement 1:3:2 1:5:2 1:6.5:2
(CO2: ATP: NADPH)
4. Optimum
temperature for
photosynthesis 15-25 25-40 35-45
(oC)
5. CO2 fixation
pathways One Two separated inspace Two separated intime
6. Light saturationfor Unsaturated even atfull
photosynthesis ¼ to ½ of fullsunlight 1/10 of full sunlight
sunlight
7. CO2 compensation
point (µl/l) 30-70 0-10 0-5
8. Maximum values
of net photosynthesis 15-30 35-45 1-5
(µmol CO2/m2/s)
9. Transpiration
ratio (H2O lost/CO2 450-950 250-350 18-125
fixed)
10. High (1/3 of total
Photorespiration Very low or absent Very low
photosynthesis)
The triosphosphate produced in the Calvin-Benson cycle (in C3, C4 and CAM
pathways) is the starting point for a wide range of biosynthetic pathways. The two major
end products of photosynthesis are sucrose and starch. However, their syntheses are not
parts of the Calvin-Benson cycle. Sucrose is the main form of carbohydrates translocated
in the plant. It is, therefore, an important raw material for the synthesis in different parts
of the plant of many other organic molecules that are necessary for growth, maintenance,
and yield formation.
Factors Affecting Photosynthesis
Photosynthesis and consequently, crop productivity are influenced by a host of

variable factors. Included in these are both the internal and external or environmental
factors. The internal factors include nutritional status of the plant, stomatal aperture and
leaf characteristics such as age, morphology, leaf area index, leaf angle and orientation.
On the other hand, the external factors include light or irradiance, temperature, carbon
dioxide concentration, moisture and wind.
1. Internal Factors
▪ Nutritional status of the plant. A deficiency of any of the essential element will
cause a decrease in the photosynthetic rate of the leaves. For example, chlorophyll
contains both nitrogen and magnesium; if they are limited in supply, chlorophyll
may not be formed. Precursor molecules for chlorophyll synthesis include iron, and
if it is not present, chlorophyll cannot be synthesized. In this case, the deficiency
of an element influenced the photosynthetic apparatus of the plant.
In rice (O. sativa), nitrogen content per leaf area is closely related with
photosynthetic rate. Nitrogen is one of the most important nutrients and it is often
deficient in rice-cultivated areas. When nitrogen is limiting, both the leaf area index
(LAI) and the rate of photosynthesis per unit leaf surface is reduced. The
combination of these two factors results in a lowered rate of photosynthesis.
▪ Stomatal aperture. When water is abundant and nutrition is adequate, the rate of
photosynthesis is usually controlled by the stomatal aperture or opening and the
number of stomata per unit of surface area. Since carbon dioxide (CO2) diffuses
or moves into green tissues through the stomata, the stomatal number and
apertures must be great enough for passage of adequate levels of CO2. In rice
plants, stomata are found on both the upper and lower surfaces of the leaf.
Stomatal aperture is reduced by water stress, insufficient light, and high carbon
dioxide concentration.
▪ Leaf age. The photosynthetic rate of young leaves is usually low but increases as
the leaf approaches full expansion. After reaching maturity, subsequent
photosynthetic activity varies widely with both species and environment.
Old, senescent leaves eventually become yellow and are unable to

photosynthesize because of chlorophyll breakdown and loss of functional
chloroplasts.
2. External Factors
▪ Light (Irradiance). Under natural conditions, the process of photosynthesis is

“driven” by the visible portion of the spectrum or radiant energy between 400 and
700 nanometer (nm) wavelength.
▪ Photosynthetic rate is partially a function of the level of irradiance. In total
darkness, no photosynthesis occurs and as irradiance increases, photosynthesis
increases until the light compensation point is reached, where photosynthetic
fixation of CO2 exactly equals respiratory release of CO2, i.e. there is no net
movement of CO2 into or out of the leaf (net photosynthesis = 0). As irradiance
further increases, net photosynthesis (gross photosynthetic CO2 fixation minus
respiration) increases rapidly until the light saturation level is reached, at which
point the photosynthetic rate levels off.
▪ The following are units or expressions of:
a. photosynthesis
▪ mg CO2 /dm 2 leaf area/ hr-
▪ mg CO2 /cm2 leaf area/sec
▪ u mol CO2 / m2 leaf area/sec
b. irradiance
▪ watt / m2
▪ cal /cm 2 / min
Light intensity varies with the season, for instance in Los Baños, Laguna the light
intensity during the wet and dry seasons are:
a. wet season, 300 cal / cm2 / day

b. dry season, 500-600 cal / cm 2 / day
This means that there is a doubling of the available light during summer and
explains why light is often a limiting factor to high yields during the wet season.
▪ Temperature. The light dependent reaction of photosynthesis is little affected by

temperature but the dark reaction is very temperature dependent.
▪ Within the temperature range of 20-35 0C, temperature has little effect on
photosynthesis. However, it does have a definite effect on respiration.
Enzymatically controlled reactions such as respiration occur in a temperature
range greater than 0oC and less than 50oC. Respiration increases by a factor of
2 or 3 for every increase in temperature of 10oC. This means that the high tropical
temperature has no effect on photosynthesis but will result in a higher rate of
respiration. This is one reason why yields are low in the tropics.
▪ Carbon dioxide (CO2) concentration. Carbon dioxide is a gaseous component

of air. Dry air contains 78% nitrogen (N2); 21% oxygen (O2); 0.93% argon (Ar);
0.038% (380ppm) CO2 and traces of other gases. Although carbon dioxide is at a
low concentration, 85% to 92% of a plant’s dry weight is derived from CO2 uptake
in photosynthesis.
▪ The average CO2 concentration of normal ambient (surrounding) air is about
0.038% or about 380 ppm. Diurnal fluctuation of CO2 concentration occurs over
the rice fields, increasing at night when only respiration is being carried on and
decreasing during the day when the rice plants are carrying out photosynthesis.
▪ Photosynthetic rates are enhanced by higher CO2 concentration, unless stomata

are closed by drought or water stress. The response of photosynthesis to CO2 is
similar to its response to irradiance.
▪ There is a very strong interaction between CO2 level and light level. At low
irradiance, low levels of CO2 will saturate the photosynthetic mechanisms but as
light level increases, progressively higher concentrations of CO2 are required to
“use up” the NADPH and ATP produced in the light reactions. Another aspect of
CO2 levels is that within the canopy of a crop, CO2 levels are often well below the
general atmospheric level. This depletion depends on several factors including
canopy density, wind speed and irradiance.
▪ Moisture. Water is a substrate for photosynthesis, but only about 0.1% of the total
water is used by the plant for photosynthesis. Transpiration accounts for 99% of
the water used by plants. Approximately 1% is used to hydrate the plant, maintain
turgor pressure and make growth possible.
▪ The supply of water or moisture affects stomatal aperture or opening. Stomata

open because the guard cells take up water and swell. When water is deficient,
stomata close, hence the rate of photosynthesis is reduced.
▪ Wind. Some preliminary data need to be examined in order to understand the role
of wind in photosynthesis.
▪ The maximum increase in dry weight per square meter per day observed in rice in
Los Baños, Laguna is about 30 g. If their mineral content accounts for 10% of the
dry matter, the net increase in organic matter would be 27 g / m2 / day. To obtain
27 g of organic matter (dry weight /m2), it is necessary for the plant to assimilate
39.6 g of CO2 / m2.
▪ The replacement of air above the rice crop with fresh air is important to the
photosynthetic efficiency of the crop. If the air were not replaced, it would quickly
become depleted of CO2. Air movement is accomplished primarily through wind
and turbulence. The movement of air through turbulence has been computed 104
times faster than normal diffusion would be. Normally, moderate winds and
turbulence are important to the production of high yields.
2. RESPIRATION
Two of the most important processes

carried out by green plants are
photosynthesis and respiration.
Photosynthesis, which takes place in the
chloroplasts, is constructive and reductive,
while respiration is degradative (destructive)
and oxidative.
All humanity and animal life is

dependent on the ability of plants to utilize
solar energy and to store it to provide
material for respiration.
Every organism must extract energy

from the organic fuel molecules that it either
manufactures or captures from the
environment. These fuel molecules are
transported to all the cells of a complex
organism, where they can be broken down
to provide the energy for cellular work.
Every plant cell then must extract energy from its long-term storage molecules –
the organic molecules it manufactures by photosynthesis. Within each plant cell, glucose
and other fuel molecules are broken down by the process of cellular respiration, a series
of chemical reactions that break apart molecules and transfer the energy stored in their
bonds to adenosine triphosphate (ATP) for use to carry out work in the cell.
Cells use different catabolic pathways to extract energy from the fuel molecules
they manufacture or ingest: aerobic respiration and anaerobic respiration. The type of
environment a cell inhabits may determine which catabolic pathway it uses to break down
fuel molecules. Cells that live in an environment where oxygen is plentiful use an aerobic
pathway, one that requires oxygen, whereas cells that inhabit waterlogged soils or
polluted water where oxygen is absent must use anaerobic pathways that do not require
oxygen.
A. The Process of Aerobic Respiration
Plant cells extract energy from fuel molecules (e.g., glucose, fatty acids and other
organic compounds) by using aerobic respiration. This process involves a long sequence
of 30 or more chemical reactions, each regulated by a specific enzyme. During aerobic
respiration, energy is released as fuel molecules are catabolized to CO2 and water. One
of the most common pathways of aerobic respiration involves breakdown of glucose.
Glucose is generally regarded as the starting point for the respiratory metabolism of
carbohydrates.
The overall process of respiration can be expressed by the equation below:
C6H12O6 + 6O2 6CO2 + 6H2O + Energy

glucose oxygen carbon dioxide water
Chemically, respiration is oxidation. In oxidation, either oxygen is added to the

material being oxidized or hydrogen is removed from it. A substrate, or more specifically
a respiratory substrate, is any organic plant constituent oxidized partially (to more
oxidized compounds) or completely (to carbon dioxide and water) in respiratory
metabolism.
The chemical reactions of aerobic respiration are grouped into four stages, namely:
▪ Glycolysis
▪ Formation of acetyl coenzyme A
▪ The citric acid cycle also known as Krebs cycle
▪ Electron transport and chemiosmosis
Glycolysis occurs in the cytoplasm; all other parts of aerobic respiration occur in
the mitochondrion.
B. Importance of Aerobic Respiration
1. It supplies the
energy needed for
the growth and
maintenance of the
plant.
2. It provides carbon
skeletons needed
for the synthesis of
a large number of
other essential
plant products.
These products
include amino
acids for proteins,
nucleotides for
nucleic acids, and carbon precursors for porphyrin pigments (such as chlorophyll
and cytochromes) and for fats, sterols, carotenoids, flavonoid pigments such as
anthocyanins, and certain other aromatic compounds such as lignin.
C. Factors Affecting the Rate of Aerobic Respiration
1. Temperature. Temperature affects respiration rates. Within the range of 0oC-

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

other factors being constant and not limiting, the rate of respiration decreases as
oxygen concentration decreases. Lowering the oxygen concentration by
increasing the concentration of carbon dioxide or nitrogen is useful in storing
certain fruit and vegetable crops. This modified storage atmosphere slows down
rapid respiration.
3. Soil condition. Compacted or water-logged soils are generally poorly aerated.

This condition reduces respiration in the roots resulting in poor plant growth.
Mineral nutrient deficiencies affect the respiratory enzymes, indirectly causing a
reduction in respiration.
4. Light. Plants that grow in low light intensities exhibit lower respiration rates. Low
light reduces photosynthesis, thus decreasing the amount of carbohydrates
available for respiration.
Table 2. A comparison of photosynthesis and aerobic respiration.
Items Photosynthesis Aerobic respiration

Raw materials CO2 , H2O C6H12O6, O2
End products C6H12O6, O2 CO2 , H2O
Plant cells that have
Plant cells that containchlorophyll All plant cells
these process
Parts of cellinvolved Chloroplast Cytoplasm, mitochondrion
Light energy → NADPH/ATP Energy stored in fuel
Pathway of energy → energy stored in carbohydrate molecules → NADH/ATP
molecules → energy for work in cell
3. TRANSPIRATION
Transpiration is the loss of water in the form of water vapor from aerial parts of
plants. It is basically an evaporative process. It involves two stages: (1) the evaporation
of water from cell surfaces
(dependent on energy, the
latent heat of vaporization,
which is equal to 539 cal/gram),
(2) the diffusion of water vapor
out of the plant, mainly through
the leaves. The driving force of
transpiration is the vapor
pressure gradient between the
leaf interior and the
atmosphere.
There are three types of

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

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

mobilization of soil nutrients toward the roots. It aids in the translocation of mineral
nutrients absorbed by the roots. It cools the plant, thereby maintaining a favorable plant
temperature for growth and development.
The factors that affect transpiration are:
1. solar radiation – it is the main source of energy for the evaporation of water;
visible radiation (light) can directly or indirectly induce the opening of stomata.
That’s why transpiration is high during the day, except in CAM plants.
2. temperature – an increase in temperature increases the capacity of the air to
absorb water vapor.
3. relative humidity – as relative humidity becomes lower, transpiration
increases.
4. wind – a gentle breeze increases the rate of transpiration compared to still air.
5. soil moisture availability – when there is lack of water in the soil, transpiration
is reduced.
6. carbon dioxide concentration – an increase in carbon dioxide concentration
induces partial closure of stomata.
Thus, transpiration is reduced plant adaptations such as modification of leaves into

scales or spines, degree of cuticular deposition, sunken stomata, closing of stomata
during the day (in CAM plants), presence of hairs on the leaf, solar tracking in which the
leaf is oriented such that its lamina is almost parallel to the rays of the sun thereby
reducing its heat load, etc.
4. TRANSLOCATION
Translocation and
partitioning of assimilates
refer to the movement and
distribution of organic
substances produced during
photosynthesis within a plant.
Assimilates are organic
compounds such as sugars,
amino acids, and lipids that
are produced in the leaves
through photosynthesis.
These assimilates are
essential for plant growth and
development.
Translocation involves
the movement of assimilates
from the site of production,
typically the leaves, to other
parts of the plant such as
roots, stems, flowers, and
fruits. This movement
primarily occurs through the
phloem, a specialized vascular tissue responsible for transporting sugars and other
organic compounds. The process of translocation relies on a pressure gradient created
by the active loading of sugars into the phloem cells in source tissues (usually leaves)
and their unloading in sink tissues (other parts of the plant).
Partitioning refers to the distribution or allocation of assimilates among different
plant organs or tissues according to their respective demands. Plants allocate assimilates
to support various physiological processes such as growth, respiration, storage, and
reproduction. For example, during periods of active growth, assimilates may be
partitioned towards developing leaves, stems, or roots. In contrast, during reproductive
stages, assimilates may be allocated towards the development of flowers and fruits.
The translocation and partitioning of assimilates are highly regulated processes

influenced by various internal and external factors including environmental conditions
(such as light intensity, temperature, and water availability), hormonal signals, and
developmental cues. Understanding these processes is crucial for optimizing crop yield
and quality in agriculture and for elucidating plant responses to changing environmental
conditions.
To maintain its metabolic activity, the plant needs to circulate to the various plant
parts a wide variety of compounds. In multicellular plants, the water and inorganic
substances absorbed by the roots have to be transported to the leaves. The sugars and
other organic compounds synthesized there have to be distributed to all parts of the plant,
where they are used for growth and maintenance, or stored. These parts include the parts
that we harvest.
To meet these needs, higher plants have developed two translocation systems,
the xylem and the phloem. The xylem provides the pathway by which the water and
mineral nutrients absorbed by the roots are distributed to all parts of the plant, including
the highest leaves. The phloem, on the other hand, provides the pathway by which the
sugars and other organic compounds produced in the leaves and some inorganic ions
move to the different parts of the plant.
Sugars, which are synthesized during photosynthesis, move throughout the plant,
principally through the phloem tissue. The movement can be downward, from leaves to
roots, but lateral and upward movement from leaves to buds or to fruits or to storage
organs also occurs. Translocation takes place in the long sieve elements connected end
to end to form sieve tubes.
The rate of translocation of sugars in the phloem is rapid, in some instances more
than a thousand times faster than simple diffusion of sugar through water. The rate of
translocation has been measured in many plants and average values of 1 to 6 g/cm2/hr
have been found in developing fruits and tubers.
Much of the carbohydrate translocated within plants is sucrose. This disaccharide

is formed by the linkage of glucose and fructose accompanied by the removal of a
molecule of water.
Movement of materials in living plants has been observed to occur in different

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

diseases, or mechanical breakage (as in lodging).
5. MINERAL NUTRITION
Mineral nutrition in plants refers to the acquisition, uptake, transport, and utilization
of essential mineral nutrients required for their growth, development, and overall
physiological functioning. These mineral nutrients are obtained primarily from the soil
through the plant's root system.
There are two main categories of mineral nutrients required by plants:

macronutrients and micronutrients.
a. Macronutrients:
Macronutrients are essential elements required by plants in relatively large

quantities for their growth, development, and overall functioning. These nutrients play
critical roles in various metabolic processes and are necessary for the synthesis of
important organic molecules.
The main macronutrients required by plants are:
1. Nitrogen (N)
Functions: Essential component of amino acids, proteins, nucleic acids,
chlorophyll, and various hormones. Plays a crucial role in photosynthesis, growth,
and development.
Deficiency Symptoms: Stunted growth, chlorosis (yellowing) of leaves, reduced
leaf size, and delayed flowering.
2. Phosphorus (P)
Functions: Involved in energy transfer processes (ATP), nucleic acid synthesis
(DNA, RNA), membrane structure, and root development. Important for flowering
and fruiting.
Deficiency Symptoms: Stunted growth, purplish or reddish discoloration of leaves,
poor root development, and delayed maturity.
3. Potassium (K)
Functions: Facilitates osmoregulation, regulates stomatal opening and closing,
activates enzymes involved in photosynthesis and respiration, and enhances
drought and disease resistance.
Deficiency Symptoms: Yellowing and necrosis of leaf margins, weak stems, poor
fruit quality, and susceptibility to stress.
4. Calcium (Ca)
Functions: Structural component of cell walls, essential for cell division, membrane
stability, and signal transduction. Regulates enzyme activity and cell permeability.
Deficiency Symptoms: Stunted growth, distorted or necrotic leaf margins, blossom-
end rot in fruits, and impaired root development.
5. Magnesium (Mg)
Functions: Central component of chlorophyll molecules, essential for
photosynthesis. Activates enzymes involved in ATP metabolism and protein
synthesis.
Deficiency Symptoms: Interveinal chlorosis (yellowing between leaf veins), leaf
curling, reduced growth, and poor fruit quality.
6. Sulfur (S)
Functions: Integral component of amino acids (cysteine and methionine), vitamins,
coenzymes, and proteins. Essential for the synthesis of sulfur-containing
compounds.
Deficiency Symptoms: Chlorosis of younger leaves, stunted growth, delayed
maturity, and reduced seed and fruit production.
b. Micronutrients (Trace Elements):
Micronutrients, also known as trace elements or minor elements, are

essential nutrients required by plants in small quantities for their growth,
development, and overall health. While they are needed in smaller amounts
compared to macronutrients, micronutrients play crucial roles in various
physiological processes and are indispensable for plant metabolism.
The main micronutrients required by plants are:
1. Iron (Fe)
Functions: Essential for chlorophyll synthesis, electron transport in photosynthesis
and respiration, and enzyme activation (e.g., cytochromes).
Deficiency Symptoms: Interveinal chlorosis in young leaves, reduced growth, and
poor chlorophyll formation.
2. Manganese (Mn)
Functions: Involved in photosynthesis, enzyme activation (e.g., superoxide
dismutase), and the synthesis of chlorophyll and other pigments.
Deficiency Symptoms: Interveinal chlorosis with brown or black spots, stunted
growth, and reduced fruiting.
3. Zinc (Zn)
Functions: Essential for enzyme activation (e.g., carbonic anhydrase), protein
synthesis, and auxin metabolism. Plays a role in chlorophyll formation and root
development.
Deficiency Symptoms: Stunted growth, shortened internodes, interveinal chlorosis
in younger leaves, and reduced fruit set.
4. Copper (Cu)
Functions: Required for photosynthesis, electron transport (cytochrome c
oxidase), lignin synthesis, and enzyme activation (e.g., superoxide dismutase).
Deficiency Symptoms: Dieback of shoot tips, wilting, distorted leaves (known as
"twisted tip" symptom), and reduced fertility.
5. Molybdenum (Mo)
Functions: Necessary for nitrogen metabolism, particularly in the conversion of
nitrate to ammonia (nitrogen fixation) and the synthesis of amino acids (e.g.,
methionine).
Deficiency Symptoms: Interveinal chlorosis with rolled or cupped leaves, reduced
growth, and impaired flowering.
6. Boron (B)
Functions: Essential for cell wall synthesis, carbohydrate metabolism, pollen
germination, and the movement of sugars within the plant.
Deficiency Symptoms: Brittle or corky tissues, distorted growth (e.g., "witches'
broom"), hollow stems or roots, and poor fruit development.
7. Chlorine (Cl)
Functions: Involved in photosynthesis (as a cofactor for OEC, oxygen-evolving
complex), osmoregulation, and stomatal function.
Deficiency Symptoms: Reduced growth, wilting, and leaf tip burn (necrosis).
Plants absorb these mineral nutrients in the form of ions dissolved in soil water
through their root system. The uptake process is facilitated by various mechanisms
including active transport, facilitated diffusion, and ion exchange. Once absorbed, the
minerals are transported via the vascular system (xylem and phloem) to different parts of
the plant where they are utilized for various metabolic processes.
Deficiencies or imbalances in mineral nutrition can lead to stunted growth,

chlorosis, necrosis, reduced yield, and overall poor plant health. Therefore, understanding
the requirements and interactions of mineral nutrients is essential for optimizing plant
growth and productivity in agriculture and horticulture. Soil testing, fertilization, and
nutrient management practices are commonly employed to address nutrient deficiencies
and maintain soil fertility.
Organic Matter
• Organic matter represents the remains of plants and animals at various stages of
decomposition.
• Its advantages are:
• Improves drainage, aeration, nutrient and water holding capacity of the soil
• Binds soil particles together into different sizes and forms (structure)
• Provides nutrients. It is especially a very good source of some
microelements.
• Soils high in organic matter are usually dark-colored and ideal for vegetable
production.
• When a soil is low in organic matter, it becomes hard and forms crust during
summer months.
Soil Reaction (pH)

• Soil reaction refers to the degree of acidity or alkalinity and is measured in terms
of pH.
• At pH 7, the soil is neither acidic nor alkalinic (neutral).
• At pH 7, the soil is acidic in reaction
• At pH greater than 7, the soil is alkaline in reaction
• The lower the pH, the more acidic is the soil
• The higher the pH, the more alkaline is the soil
Table 6. Soil classes according to pH.

Group pH
Extremely acidic Below 4.5
Very extremely acidic 4.5 – 5.0
Strongly acidic 5.1 – 5.5
Medium acidic 5.6 – 6.0
Slightly acidic 6.1 – 6.5
Neutral 6.6 – 7.3
Slightly alkaline 7.4 – 7.8
Moderately alkaline 7.9 – 8.4
Strongly alkaline 8.5 – 9.0
Very strongly alkaline 9.0 +
• Vegetables usually grow well in slightly acid or slightly alkalinic soil.

• Below this pH range, calcium may become less available.
• Above this pH range, iron and manganese may become less available
Table 7. Optimum pH range for different vegetables
Crop Optimum range
Potato, garlic 5.0 – 6.5
Corn 5.3 – 7.3
Peanut 5.5 – 6.0
Squash, watermelon 5.5 – 6.5
Snap bean 5.5 – 6.7
Soybean, sweet potato, yam bean 5.5 – 7.0
Chinese cabbage, eggplant, lettuce 6.0 – 6.5
Cabbage 6.0 – 7.0
Radish, spinach, cassava 6.0 – 7.5
Asparagus 6.0 – 8.0
Presence of Salts
• Salt may be present in soils in quantities which vegetable crops may tolerate
• Saline soils are those soils with high soluble salts of ammonium, calcium,
magnesium and sodium
• Usually the salt is sodium chloride or sodium chloride
Table 8. Relative salt tolerance of vegetable crops

Relatively non- Moderately salt- Relatively salt- Highly salt-tolerant
tolerant tolerant tolerant
EC range in micromhes/cm at 25ºC
200 – 400 400 – 600 600 – 800 800 – 1200
Lima bean Tomato Green beet asparagus
Green bean Broccoli Kale
Celery Cabbage Spinach
Pepper Okra
Lettuce
Sweet corn
Onion
Pea
Watermelon
Cantaloupe
Squash
• Salt damage to plants which is expressed as low yield is due to excessive uptake
of substances, such as chloride in toxic quantities.
• Increased salts in the soil water also reduce the amount of water available to
plants, so crops grown in saline soils need more frequent irrigation than the same
crops grown in normal soils.
Fertilizer and Green Manuring
Fertilizer
• Materials applied to the soil to promote greater plant growth or better crop quality.
Food/Nutrients
• Materials given to plants for growth, building up of new plants parts or to give
strength in performing work.
How are food nutrients lost from the soil?

1. Leaching – nutrients is being carried by the water
2. Seepage – sideward loss of water in the soil
3. Percolation – downward movement/loss of water in the soil
4. Yearly removal of material through the crop
5. Mechanical erosion
6. Loss of nitrogen through denitrification
7. Volatilization
Forms of Commercial Fertilizer

1. Single element – carries only one element that is present in the source of fertilizer
2. Double element – fertilizer with two elements
3. Complete fertilizer – fertilizer contains the three most important elements namely,
N, P. and K
Kinds of Fertilizer
1. Organic Fertilizer
2. Inorganic Fertilizer
Table 9. Differences of Organic and Inorganic Fertilizers:

Organic Fertilizer Inorganic Fertilizer
Advantages
Economical Easy carriage
Longer effectivity Faster effectivity
Neutralizing Readily available
Improve soil texture
Inhibit the multiplication of harmful
microorganisms
Disadvantages
Laborious Expensive
Takes time to take effect Dangerous to the plant (sometimes)
Needs a larger space Makes the soil acidic
Need available composting materials Easily lost from the soil
Voluminous
Fertilizer requirements
• Soil analysis
• Plant tissue analysis
• Field fertilizer trials
Time of Application
• Basal – fertilizer is applied during or before planting
• Top dressing – fertilizer is broadcasted over head
• Side dressing – fertilizer is applied along the furrow’s side
Methods of Fertilizer Application

• Foliar spray
• Broadcast
• Drill
• Furrow method
Green manuring
• Is the growing of legumes and turning them under during the blooming stage.
Legumes
• Pod producing plants having root nodules
Benefits or advantages of Green Manuring

1. Addition of humus or vegetable matter to the soil will increase its capacity for
retaining moisture
a. Humus – a complex amorphous, colloidal substances that is resistant to
further decomposition. It is the end in organic matter decomposition
b. Importance:
i. Soil fertility
ii. Soil quality
2. Prevention of surface run – off or wash (erosion) on steeps lands
3. Improvement of the roots, condition of the soil brought by the action of the roots of
green manure crops
4. Protection of the soil and roots of crops from excessive heat of the sun
5. Suppression or control weeds
6. Renders more available materials for plant nutrition
7. Fixation of atmospheric nitrogen in the soil by the action of symbiotic
microorganisms (e.g., Rhizobium in the root nodules of legumes) that are present
in leguminous plants
Fertilizer Computation
• It is not enough that a farmer applied fertilizer to his crops. The amount he applies
may increase his profits or cause losses for him depending on how and how much
fertilizer he applies.
• Applying what you think is the right amount of fertilizer might still be too little to
produce the desired effect that would be profitable.
• Applying too much might be more than necessary, thus wasting money which
would otherwise have been profit.
• In essence, not applying the right amount of commercial fertilizer could be caused
by failure to calculate the quantity of commercial fertilizer in its equivalent
recommended dosage.
• Computing fertilizer is not really that difficult. One has to know how to follow a given
formula and perform the operations of multiplication, division, addition and
subtraction.
A. Working formula:
Rn
Qcf = ---------------------------- (A)
G
Where:
• Qcf – amount of commercial fertilizer expressed in grams, kilogram or bags of the
chosen fertilizer materials such as ammonium sulfate, ammophos or complete
fertilizer
• R – amount of recommended nutrients (N or P2O5 or K2O) expressed in kilograms

per hectare. To facilitate computation, the amount of nutrient recommended (Rn)
to be computed should correspond to the area (A) where fertilizer should be
applied.
• G – as mentioned earlier, this refers to the percent of either N, P2O5 contained in

the fertilizer material converted to a decimal number by dividing by 100. In
computing for the amount of commercial fertilizer materials needed or the amount
of nutrient elements present in a given amount of commercial fertilizer, the
corresponding analysis for any of the three elements in question should be used.
Example:
If the recommended amount of nutrient is 60 kg N/ha (Rn) and the area is 2

hectares (A), the total amount of nutrient becomes 120 kg N, (Rn x A = 60 kg N/ha x 2
hectares). Likewise, if the recommended amount if 60 + 30 + 0 kg/ha and the area is 1.5
hectares, the recommendation will be 90 + 40 + 0 (60 kg N/ha x 1.5 hectares) = 90 kg N;
30 kg P2O5/ha x 1.5 hectares = 45 kg P2O5
Formula:
RR
Fertilizer requirement = ------------------------- x 100 x area
A
Where:
RR = recommended rate
Ai = Active ingredient
100 = constant
A. Based on Ai:
1. Given: RR = 90 – 60 – 30 Fertilizer sources = 46 – 0 – 0

= 0 – 21 – 0
= 0 – 0 – 21
Solution:
30 60 90
a. --------------- x 100 b. -------------- x 100 c. -------------- x 100
21 21 46
= 142.85 kg of 0 – 0 – 21 = 285.71 kg of 0 – 21 – 0 = 195.65 kg of 46-0-0
2. Given: RR = 90 – 30 – 30
Fertilizer sources = 46 – 0 – 0
= 14 – 14 – 14
Solution:
30
a. --------------- x 100 = 214.28 kg of 14 – 14 – 14
14
90 – 30 – 30
b. 30 – 30 – 30
-------------------------
60 – 0 – 0
60
c. --------------- x 100 = 130.43 kg of 46 – 0 – 0
46
3. Given: RR = 90 – 60 – 30
Sources = 46 – 0 – 0
= 16 – 0 – 0
= 14 – 14 – 14
Solution:
30
a. --------------- x 100 = 214.28 kg of 14 – 14 – 14
14
90 – 60 – 30
b. 30 – 30 – 30
-------------------------
60 – 30 – 0
30
c. --------------- x 100 = 150 kg of 16 – 20 – 0
20
d. 150 x 0.16 = 24% N
60 – 30 – 30 36
e. 24 – 30 – 30 f. --------------- x 100
------------------------- 46
36 – 0 – 0
= 78.26 kg of 46 – 0 – 0
F. PLANT REPRODUCTION
Plant reproduction is the biological process by which plants produce offspring or

new individuals. Unlike animals, which often rely on mating and sexual reproduction
involving the fusion of gametes (sperm and egg), plants exhibit a diverse range of
reproductive strategies, including both sexual and asexual methods.
The mode of reproduction determines the genetic constitution of crop plants, that
is, whether the plants are normally homozygous or heterozygous. This, in turn,
determines the breeding methods applicable to a crop species. A knowledge of the mode
of reproduction of crop plants is also important for making artificial hybrids and provides
a basis for understanding the mechanism of heredity in plants.
The various modes of reproduction found in crop plants may be broadly grouped
into two types: (1) asexual and (2) sexual
1. Asexual Reproduction
This mode of reproduction does not involve fusion of male and female gametes. It
is prevalent in all plants that are devoid of seed set, have long reproduction cycle, have
interozygosity etc. New plants may develop from vegetative parts of the plant. (Vegetative
reproduction)
a. Vegetative Reproduction
Vegetative Propagation: Many plants can reproduce asexually through vegetative

structures such as rhizomes, stolons, bulbs, tubers, runners, or plantlets. In this process,
new individuals develop from specialized vegetative organs without the involvement of
seeds or fertilization.
Vegetative reproduction involves the use of the following:
1. Underground stems. A modified and specialized plant structures that grow below
the soil surface and serve various functions, including storage of nutrients, water,
and energy reserves, vegetative propagation, and survival during adverse
conditions like rhizome (e.g, ginger, banana, turmeric), tuber (e.g., potato), bulb
(e.g., onion, garlic) corm (e.g., Colocasia, yam), are used for multiplication.
2. Sub-aerial stems. Plant structures that grow partially above ground and partially
below ground, often serving as transitional structures between aboveground aerial
parts and underground structures These modifications include runner (e.g.,
strawberry), stolon, sucker (e.g., chrysanthemum)
3. Bulbils. Specialized structures produced by some plants as a means of

propagation. These structures resemble bulbs in appearance and function, but
they are smaller and typically arise from the leaf axils or stems of the plant rather
than from the basal plate like true bulbs. Bulbils are modified flowers that develop
into plants directly without formation of seeds (e.g., Agaves).
4. Normal stem. It is commonly used for the propagation of many crop species.
Normal stem is used as a propagule by cuttings (e.g., sugarcane), layering (e.g.,
lemon, lichi), grafts (e.g., mango, apple), buddings.
b. Apomixis
Apomixis is a form of asexual reproduction in which seeds are produced without

fertilization. The ovule develops into a seed containing an embryo formed from maternal
tissues without genetic recombination.
In apomixis, seeds are formed but the embryos develop without fertilization. When
sexual reproduction also occurs, the apoximis is termed as facultative. But when sexual
reproduction is absent, it is referred to as obligate.
Apomixis is classified into:
1. Adventive Embryo – In here, embryos develop directly from vegetative cells of

the ovule, such as nucellus, integument, and chalaza.
2. Apospory – Some vegetative cells of the ovule develop into unreduced embryo
sacs after meiosis. The embryo may develop from egg cell or some other cell of
such an embryo sac.
3. Diplospory – Embryo sac is produced from the megaspore which may be haploid
or, more generally, diploid.
3a. Parthenogenesis – In parthenogenesis, the embryo develops from an egg
cell.
3b. Apogamy – In apogamy, synergids or antipodal cells develop into an embryo.
c. Fragmentation
Some plants can regenerate from fragments of their parent plant. For example,
pieces of stem, leaves, or roots can develop into new individuals under suitable
conditions.
Fragmentation is a method of asexual reproduction in which a parent organism

breaks into fragments, each of which can grow into a new individual. This process is
common in various organisms, including plants, fungi, and some animals, and it enables
rapid propagation and colonization. Fragmentation typically occurs in organisms with
modular body plans, where each module has the potential to develop into a complete
organism.
In plants, fragmentation can occur through various mechanisms:
1. Natural Breakage. Some plants naturally produce brittle stems or branches that
can break off easily due to environmental factors such as wind, water currents, or
animal activity. These broken fragments can then root and grow into new plants.
For example, pieces of stems or branches of willow trees can root and give rise to
new individuals when carried downstream.
2. Human-induced Fragmentation. Human activities such as pruning, cultivation,
or gardening can inadvertently cause fragmentation in plants. For instance, when
a gardener divides a clump of iris plants or separates runners from a strawberry
plant, each fragment has the potential to develop into a new plant.
3. Accidental Damage. Accidental damage to plants, such as physical trauma or
mechanical injury, can result in the fragmentation of plant tissues. For example, if
a branch of a succulent plant breaks off, it may produce roots and form a new plant
when placed in suitable growing conditions.
4. Rhizomatous Plants. Some plants produce rhizomes, underground stems that
can give rise to new shoots and roots. When these rhizomes are damaged or
disturbed, they may fragment into smaller pieces, each capable of generating a
new plant. Examples include bamboo and some grasses.
d. Agamospermy
Agamospermy is a form of asexual reproduction in plants in which seeds are

produced without the process of fertilization. In agamospermy, the embryo develops from
unfertilized egg cells or other cells of the ovule without the fusion of male and female
gametes. This process results in the formation of offspring that are genetically identical
to the parent plant, as there is no genetic recombination.
Agamospermy provides several advantages to plants:
1. It allows plants to reproduce rapidly without the need for pollination or the presence
of compatible mates, particularly in environments where pollinators are scarce or
unreliable.
2. It ensures genetic uniformity among offspring, which can be advantageous for
maintaining desirable traits or adaptations in a population.
3. It provides a mechanism for certain plant species to colonize new habitats or
expand their range quickly, as agamospermous seeds can germinate and
establish themselves under a wide range of environmental conditions.
However, agamospermy may also have disadvantages:
1. Lack of genetic diversity can limit the ability of plants to adapt to changing
environmental conditions or resist diseases and pests.
2. Agamospermous plants may face challenges in evolutionary processes such as
natural selection and genetic recombination, which are important for long-term
species survival and diversification.
2. Sexual Reproduction
Sexual reproduction involves fusion of male and female gametes to form a zygote,
which develops into an embryo to form seed. In crop plants, male and female gametes
are produced in specialized structures known as flowers.
Flower distribution on the plant
a. Perfect or hermaphrodite flower – contains both stamens and pistil. b.)

Staminate flower – contains stamens but not pistil.
b. Pistillate flower – contains pistil but not stamen.
c. Monoecious species – staminate and pistillate flowers occur on the same
plant Ex. maize, castor bean, coconut.
d. Dioecious species – staminate and pistillate flowers occur on different
plants. e.g., papaya, date palm, hemp.
Gamete Formation and Fertilization
a. Sporogenesis – production of microspores and megaspores
Microsporogenesis – production of microspores

Megasporogenesis – production of megaspores
b. Gametogenesis – the production of male and female gametes in the microspores and
megaspores.
Microgametogenesis – refers to the production of male gamete or sperm.

Megagametogenesis – the development of embryo sac from a megaspore.
c. Fertilization – The fusion of the sperm with the egg cell producing a diploid zygote
Anthesis – the first opening of a flower.
Modes of Pollination
Pollination refers to the transfer of pollen grains from anthers to stigmas.
a. Self-pollination (autogamy) –is the transfer of pollen from an anther to a stigma

within the same flower or to stigma of another flower on the same plant
b. Cross-pollination (allogamy) – is the transfer of pollen to the stigma in a flower on
a different plant
c. Often Cross-pollination – refers to out crossing of 15-30 percent in self-pollinated
crops.
Floral Mechanisms Promoting Self-Pollination:
a. Cleistogamy – flowers do not open at all.

b. Chasmogamy – flowers open, but only after pollination has taken place.
c. Stigmas are closely surrounded by anthers.
d. Flowers open but the stamens and the stigma are hidden by other floral organs.
e. Stigmas become receptive and elongate through the staminal columns.
Mechanisms Promoting Cross Pollination
a. Dicliny or unisexuality – is a condition in which the flowers are either staminate

(male) or pistillate (female).
Monoecy – Staminate and pistillate flowers occur in the same plant, either in the
same inflorescence or in separate inflorescences.
Dioecy – The male and female flowers are present on different plants
b. Dichogamy – stamens and pistils of hermaphrodite flowers may mature at

differenttimes
Protogyny – pistils mature before stamens

Protandry – stamens mature before pistils
c. Stigmas are covered with a waxy film.

d. Combination of two or more of the above mechanisms.
e. Self-incompatibility – refers to the failure of a flower to fertilize the same flower or
other flowers on the same plant.
f. Male sterility – refers to the absence of functional pollen grains in
otherwisehermaphrodite flowers.
References:
Competition. (2019). Retrieved from Marietta.edu website:

http://w3.marietta.edu/~biol/biomes/competition.htm
Copeland, L.O. 1970. Principles of Seed Science and Technology. Burgess Publishing Co.,U.S.A.
Galston, A.W., P.J. Davies and R.L. Satter. 1980. The Life of a Green Plant (3rt ed.), p. 249.
Engelwood Cliffs, NJ: Prentice Hall.
Galvez, C.T. 2004. Lecture Delivered During the Review for Licensure Examination for
Agriculture. Central Luzon State University. Science City of Muñoz, Nueva Ecija.
Harper, F. 1983. Principles of Arable Crop Production. Granada, New York.
Hopson, J. and N. Wessels 1990. Essentials of Biology. McGraw-Hill, Inc., U.S.A.Janick, J. 1981.
Plant Science. W.H. Freeman and Co., U.S.A.
Leopold, A. and P. Kriedmann. 1975. Plant Growth and Development. McGraw-Hill BookCo.,
U.S.A.
Mohr, H and P. Schopfer. 1995. Plant Physiology. Springer-Verlag Berlin Heidelberg,Germany.

Principle of limiting factors – Oxford Reference. (2017, June 16). Retrieved from
Oxfordreference.com website:
https://www.oxfordreference.com/view/10.1093/oi/authority.20110803100346211
Rose, W.C. (1931). Feeding Experiments. Journal of Biological Chemistry 94: 155–65.
Salisbury, F.B. and C.W. Ross. 1992. Plant Physiology. (4th ed.) Belmont, CA: Wadsworth
Publishing Co.
Shelford, V. E. (July 1, 1931). “Some Concepts of Bioecology”. Ecology. 12 (3): 455–467.

doi:10.2307/1928991.
Turchin, P. (2001). “Does Population Ecology Have General Laws?”. Oikos. 94 (1): 17-26.
doi:10.1034/j.1600-0706.2001.11310. x.

Crop Science 1 Module1 3A

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Introduction

Physiological processes are fundamental to the growth, development, and

A. CONCEPTS OF GROWTH AND DEVELOPMENT

Growth is defined as an irreversible increase in the size of the organism.

Fig. 2. Meristems and tissues

Development or morphogenesis – is the series of changes by which an

Table 1. Comparison of Plant and Animal Development

Cellular Basis of Growth & Development

Location of growth (meristem - apical, lateral, intercalary, cambium)

There are two aspects of plant growth: primary and secondary.

a) Primary growth takes place in young, herbaceous organs resulting in an increase

Meristems and Growth

1. Accretion. Growth of non-living organisms. Increases in size of a body of rock

2. Biological growth. Increases in size of living organism

2.1. Apoposition. Cell growth in which layers of material are deposited on

Plants are both axial and polar structures

1. An axial structure is symmetrical about a line or axis.

B. PHASES OF PLANT GROWTH AND DEVELOPMENT

An idealized S-shaped (sigmoid) growth curve exhibited by numerous annual

Fig. 3. S-shaped curve, or Sigmoid curve

Phase I. Lag phase (Establishment and

This phase relates to the

Phase III. Stationary phase (Ripening and Senescence)

The major events occurring in germination are: water imbibition, enzyme

Fig. 3. The germination phase

Fig. 4. Water imbibition

Enzymes are synthesized or activated.

Enzyme activation refers to the process

The synthesis of new

Seed dormancy can be caused by several reasons:

2. Acid scarification by sulfuric acid treatment

3. Dissolve the seed coat by:

4. Dry heat at 60-80°C for one-half to two

5. Extreme cold (-50°C to -150°C). Contraction

6. Use of 0.2% KNO3 solution

The response to temperature depends on species, variety, growing region, and

Respiration increases sharply during seed germination. Since respiration is

Light is required by certain species (e.g., lettuce). If high temperature induces

Two types of germination occur. In epigeous or epigeal germination, the

Fig. 7. Epigeal and hypogeal germination

Good germination is the foundation of a good crop. It is a requirement, although

A plant is mature if it is potentially capable of reproduction, that is, flowering. The

1. Daylength. The response of plants to daylength is called photoperiodism. In spite

2. Low temperature. Some plants require exposure to low temperature (around 0-

Senescence is initiated by flowering and accentuated by fruiting. Removal of the

In crop production, it is sometimes desirable to delay senescence to increase yield.

Plants exhibit various movements and adaptations in response to environmental

• Phototropism. The growth or movement of a plant towards or away from light.

• Thigmonasty: Non-directional movement in response to touch or mechanical

• Plants exhibit daily rhythms in physiological and biochemical processes,

• Movement or growth of plant parts in response to the direction of the sun.

• Growth or movement of plant parts in response to chemical gradients. This is

• Movement in response to light intensity. Examples include the opening of flowers

Significance and Adaptive Advantages

Crop adaptations refer to the evolutionary or agricultural modifications that enable

2. Heat and Cold Tolerance

Significance and Future Directions

4. Climate Change Resilience

D. OTHER CONCEPTS RELATED TO PLANT GROWTH

1. The Law of the Minimum