MODULE 5

Plant Growth, Development and Reproduction

Overview
This module presents the difference between plant growth to
development, the concepts related to plant growth and the types of plant
reproduction.

Time Allotment (2 Weeks)

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

1. Understand the concept of growth and development, and their

significance to plant growth and development
2. Discuss concepts of growth and relate to crop production.
3. Explain the significance of the physiological processes and how it
affects crop productivity
4. Differentiate sexual and asexual propagation and discuss the
advantages and disadvantages of each method

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

1. What is the difference between growth to development?

2. Can banana plant be propagated using seeds? Why or why not?

Discussion

This section begins with the discussion of the difference of growth to development. It
then moves on the presentation of concepts related to plant growth. The last part of this
section provides the difference between sexual to asexual propagation.
 Growth is defined as an irreversible increase in the size of the organism. 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).

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
meristems accounts for most of the increased length of stems and roots.

Fig. 2. Meristems and tissues

The basic pattern of growth in annuals is represented by an S-shaped curve, or

Sigmoid curve (Figure 3).

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 these
early 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 (Period of rapid growth, stem elongation and flowering)
This phase of growth is characterized by a rapid and often linear increase in dry matter
production. This is associated with vegetative growth in annuals and terminates with the
onset 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 begins
to senesce.
This pattern of growth is exhibited not only by the whole plant but also by any part
of the plant.

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).

Fig. 5. Enzyme activation

Initiation of Embryo Growth
The synthesis of new materials result in an increase the size of the root-shoot
axis (epicotyl, hypocotyl, radicle) (Figure 6).

Fig. 6. Initiation of embryo growth

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:

- to water (Examples: legumes,)
- thick seedcoat (Examples: mango,
ampalaya
- to oxygen (Example: grasses)
B. Immature embryo. Some seeds are
shed before the embryo is mature
a. abrasion - break the seed coat so
that water can be absorbed by
the seed
b. impaction
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
1. Requires after-ripening. This is
effected by storage

C. Presence of inhibitor:
- seedling growth inhibitor in rice 1. Overcome by washing in water
- inhibitor in barley seed, tomato 1. Gibberellic acid treatment

D. Light sensitivity 1. Exposure to light

Example: tobacco, 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 hypogeous 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, they
will 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.

CONCEPTS RELATED TO PLANT GROWTH

A. 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.

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.

B. 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.

The “law of the minimum” proved to be unsatisfactory, 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.
C. 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.

MODE OF PLANT REPRODUCTION

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

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 reproduction involve the use of the following:

1. Underground stems – Modified stems 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 – These modifications include runner (e.g. strawberry),
stolon, sucker (e.g. chrysantemum)

3. Bulbils – 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
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 eggcell 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.

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.
c.) Pistillate flower – contains pistil but not stamen.
d.) Monoecious species – staminate and pistillate flowers occur on the same plant
Ex. maize, castor bean, coconut.
e) 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

a.1 Microsporogenesis – production of microspores

a.2 Megasporogenesis – production of megaspores

B. Gametogenesis – the production of male and female gametes in the microspores and
megaspores.

b.1 Microgametogenesis – refers to the production of male gamete or sperm.

b.2 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:

1. Cleistogamy – flowers do not open at all.

2. Chasmogamy – flowers open, but only after pollination has taken place.
3. Stigmas are closely surrounded by anthers.
4. Flowers open but the stamens and the stigma are hidden by other floral organs.
5. Stigmas become receptive and elongate through the staminal columns.

Mechanisms Promoting Cross Pollination

1. Dicliny or unisexuality – is a condition in which the flowers are either staminate
(male) or pistillate (female).

1a. Monoecy – Staminate and pistillate flowers occur in the same plant, either in
the same inflorescence or in separate inflorescences.

1b. Dioecy – The male and female flowers are present on different plants
 2. Dichogamy – stamens and pistils of hermaphrodite flowers may mature at different
times

2a. Protogyny – pistils mature before stamens

2b. Protandry – stamens mature before pistils

3. Stigmas are covered with a waxy film.

4. Combination of two or more of the above mechanisms.
5. Self-incompatibility – refers to the failure of a flower to fertilize the same flower or
other flowers on the same plant.
6. Male sterility – refers to the absence of functional pollen grains in otherwise
hermaphrodite flowers.

Summary
 Growth refers to the irreversible increase in the size of the organism (increase in
length, width, area, volume, mass, or weight) while development refers to 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.
 There are two types of germination: epigeal germination and hypogeal
germination.
 There are three concepts related to plant growth: Law of the Minimum, Law of
Optima and Limiting Factors, and Law of Diminishing Returns.
 Asexual propagation uses vegetative parts for reproduction while sexual
propagation uses seeds.

Post-Assessment
Online Quiz. The ten items online will be posted on November 12, 2021 (Friday) at
1:30pm. The allotted time is only 20minutes. The google form will be closed after 20
minutes. Special online quiz will be assigned to those who fail to take the online quiz in
time.

Reference
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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 Book
Co., U.S.A.
Mohr, H and P. Schopfer. 1995. Plant Physiology. Springer-Verlag Berlin Heidelberg,
Germany.
Salisbury, F.B. and C.W. Ross. 1992. Plant Physiology. (4th ed.) Belmont, CA: Wadsworth
Publishing Co.