Paper 603: Unit V: B.

Sc 3rd Year Botany Major (

6th Semester)

Chapter: Plant Growth and Development:

Plant Growth Regulators

Plant Growth Regulators are defined as small, simple chemicals produced

naturally by plants to regulate their growth and development.

➢ Characteristics
✓ Plant Growth Regulators can be of a diverse chemical composition
such as gases (ethylene), terpenes (gibberellic acid) or carotenoid
derivates (abscisic acid). They are also referred to as plant growth
substances, phytohormones or plant hormones. Based on their
action, they are broadly classified as follows:

✓ Plant Growth Promoters – They promote cell division, cell

enlargement, flowering, fruiting and seed formation. Examples are
auxins, gibberellins and cytokinins.
✓ Plant Growth Inhibitors – These chemicals inhibit growth and
promote dormancy and abscission in plants. An example is an
abscisic acid. Note: Ethylene can be a promoter or an inhibitor, but is
largely a Plant Growth Inhibitor.

Plant Growth and Development

• Growth and its Phases

• Vernalisation
• Photoperiodism
All plant growth regulators were discovered accidentally. Let’s take a
detailed look at each regulator and learn about it more closely:

Auxins:

Discovery: Auxins were the first growth hormone to be discovered. They

were discovered due to the observations of Charles Darwin and his son,
Francis Darwin. The Darwins observed that the coleoptile (protective
sheath) in canary grass grows and bends towards the source of light. This
phenomenon is ‘phototropism’. In addition, their experiments showed that
the coleoptile tip was the site responsible for the bending. Finally, this led to
the isolation of the first auxin by F. W. Went from the coleoptile tip of oat
seedlings.

(Source: Wikimedia Commons)

✓ Types
First isolated from human urine, auxin is a term applied to natural and
synthetic compounds that have growth regulating properties. Plants produce
natural auxins such as Indole-3-acetic acid (IAA) and Indole butyric acid
(IBA). Natural auxins are found in growing stems and roots from where
they migrate to their site of action. Naphthalene acetic acid (NAA) and 2, 4-
dichlorophenoxyacetic (2, 4-D) are examples of synthetic auxins.
 ✓ Effects

• Promote flowering in plants like pineapple.

• Help to initiate rooting in stem cuttings.
• Prevent dropping of fruits and leaves too early.
• Promote natural detachment (abscission) of older leaves and fruits.
• Control xylem differentiation and help in cell division.

✓ Applications

• Used for plant propagation.

• To induce parthenocarpy i.e. the production of fruit without prior
fertilization.
• 2, 4-D is widely used as a herbicide to kill dicotyledonous weeds.
• Used by gardeners to keep lawns weed-free.
Note: The growing apical bud in higher plants inhibits the growth of the
lateral buds. This phenomenon is ‘Apical Dominance‘. Removal of the
apical bud allows the lateral buds to grow. This technique is commonly
used in tea plantations and hedge-making.

(Source: Wikimedia Commons)

 ➢ Gibberellins

➢ Discovery
It is the component responsible for the ‘bakane’ disease of rice seedlings.
The disease is caused by the fungal pathogen Gibberella fujikuroi. E.
Kurosawa treated uninfected rice seedlings with sterile filtrates of the
fungus and reported the appearance of disease symptoms. Finally, the
active substance causing the disease was identified as gibberellic acid.

➢ Types
There exist more than 100 gibberellins obtained from a variety of
organisms from fungi to higher plants. They are all acidic and are
denoted as follows – GA1, GA2, GA3 etc. GA3 (Gibberellic acid) is the
most noteworthy since it was the first to be discovered and is the most
studied.

➢ Effects

• Increase the axis length in plants such as grape stalks.

• Delay senescence (i.e. ageing) in fruits. As a result, their market
period is extended.
• Help fruits like apples to elongate and improve their shape.
Applications

• The brewing industry uses GA3 to speed the malting process.

• Spraying gibberellins increase sugarcane yield by lengthening the
stem.
• Used to hasten the maturity period in young conifers and promote
early seed production.
• Help to promote bolting (i.e. sudden growth of a plant just before
flowering) in cabbages and beet.

➢ Cytokinins

• Discovery
F. Skoog and his co-workers observed a mass of cells called ‘callus’ in
tobacco plants. These cells proliferated only when the nutrient medium
contained auxins along with yeast extract or extracts of vascular tissue.
Skoog and Miller later identified the active substance responsible for
proliferation and called it kinetin.

• Types
Cytokinins were discovered as kinetin. Kinetin does not occur naturally
but scientists later discovered several natural (example – zeatin) and
synthetic cytokinins. Natural cytokinins exist in root apices and
developing shoot buds – areas where rapid cell division takes place.

• Effects

• Help in the formation of new leaves and chloroplast.

• Promote lateral shoot growth and adventitious shoot formation.
• Help overcome apical dominance.
• Promote nutrient mobilisation which in turn helps delay leaf
senescence.

➢ Abscisic Acid

• Discovery
Three independent researchers reported the purification and
characterization of three different inhibitors – Inhibitor B, Abscission II
and Dormin. Later, it was found that all three inhibitors were chemically
identical and were, therefore, together were given the name abscisic acid.
Abscisic acid mostly acts as an antagonist to Gibberellic acid.

• Effects

o Regulate abscission and dormancy.

o Inhibit plant growth, metabolism and seed germination.
o Stimulates closure of stomata in the epidermis.
o It increases the tolerance of plants to different kinds of stress and
is, therefore, called ‘stress hormone’.
o Important for seed development and maturation.
o It induces dormancy in seeds and helps them withstand
desiccation and other unfavourable growth factors.

• Ethylene

• Discovery
A group of cousins showed that a gaseous substance released from
ripe oranges hastens the ripening of unripe oranges. Consequently,
they found that the substance was ethylene – a simple gaseous Plant
Growth Regulator. Ripening fruits and tissues undergoing senescence
produce ethylene in large amounts.

• Effects

o Affects horizontal growth of seedlings and swelling of the axis in

dicot seedlings.
o Promotes abscission and senescence, especially of leaves and
flowers.
 o Enhances respiration rate during ripening of fruits. This
phenomenon is ‘respiratory climactic’.
o Increases root growth and root hair formation, therefore helping
plants to increase their absorption surface area.

• Application
Ethylene regulates many physiological processes and is, therefore,
widely used in agriculture. The most commonly used source of
ethylene is Ethephon. Plants can easily absorb and transport an
aqueous solution of ethephon and release ethylene slowly.

o Used to break seed and bud dormancy and initiate germination in

peanut seeds.

o To promote sprouting of potato tubers.

o Used to boost rapid petiole elongation in deep water rice plants.

o To initiate flowering and synchronising fruit-set in pineapples.

o To induce flowering in mango.

Ethephon hastens fruit ripening in apples and tomatoes and increases

yield by promoting female flowering in cucumbers. It also accelerates
abscission in cherry, walnut and cotton.

In summary, one or the other plant growth regulator influences every

phase of growth or development in plants. These roles could be
individualistic or synergistic; promoting or inhibiting. Additionally, more
 than one regulator can act on any given life event in a plant. Along with
genes and extrinsic factors, plant growth regulators play critical roles in
plant growth and development. Factors like temperature and light affect
plant growth events (vernalisation) via plant growth regulators.

Plants Growth: Characteristics, Development,

Phases and Factors
Characteristics of Plants Growth:
Growth is the manifestation of life. All organisms, the simplest as well as
the most intricate, are slowly changing the whole time they are alive. They
transform material into more of themselves.

From such ingredients as minerals, proteins, carbohydrates, fats, vitamins,

hormones etc., organisms form additional protoplasm. The formation of
protoplasm is called assimilation.

A large part of the food which a plant manufactures is used as a source of

energy. Food may be consumed soon after it is produced, or it may be
stored and used as a source of energy for the plant or its offspring weeks,
months, or even years later.

A healthy plant, however, manufactures more food than is necessary to

maintain the activities of its living substance, and the surplus may be built,
more or less permanently, into its tissues, producing new protoplasm and
new cell walls and thus promoting the growth of the plant body. Growth
represents the excess of constructive over destructive metabolism.

Growth involves an irreversible increase in size which is usually, but not

necessarily, accompanied by an increase in dry weight. The basic process
of growth is the production of new protoplasm, which is clearly evident in
the regions of active cell division.
The next stage in growth is increase in plant size, which is the result of
absorption of water and the consequent stretching of the tissues, a process
which in the strict sense is not growth at all, since it involves little or no
increase in the characteristic material of the plant itself.

The third and the last stage in growth involves the entry of plenty of
building materials, chiefly carbohydrates, into the expanded young tissues.
This results in an increase in the dry weight but no visible increase in
external size of the plant. Growth is, however, more than just an
increasing amount of the plant. Differential growth of plant parts results in
a characteristic shape. Each plant species has a distinctive form,
development by growth patterns.

Differentiation:
Differentiation can be recognized at cell level, tissue level, organ level, and
at the level of an organism. It becomes more obvious at the level of organ
and organism. For instance, if we consider flower as an organ of plant, is
bears sepals for photosynthesis and protection of inner floral parts;
beautiful, coloured petals to attract insects for cross-pollination ; stamens
for producing male gametes; and the carpels for bearing the ovules which
after fertilization produce seeds.

Considering an angiosperm as an organism, we observe that it possesses

the roots for absorption of water and minerals and fixation in the soil; the
trunk and stem branches bear leaves for photosynthesis, flowers and
fruits; the fruits for bearing the seeds which on germination form each a
new plant.

Development:
Development implies a whole sequence of qualitative structural changes
that a plant undergoes from the zygote stage to its death. The
developmental changes may be gradual or abrupt. Examples of certain
abrupt changes are germination, flowering and senescence (ageing leading
to death).
Slow developmental changes include formation and maturation of tissues,
formation of vegetative and floral buds and the formation of reproductive
organs. Unlike growth, development is a qualitative change. It cannot be
measured in quantitative terms, and is either described or illustrated with
the help of photographs or drawings. Development includes growth (cell
division, enlargement and differentiation), morphogenesis, maturation
and senescence.

The growth cycle of annual, monocarpic, flowering plants (angiosperms)

begins with the fertilized egg, the zygote. The zygote develops into an
embryo following cell divisions and differentiation (embryonal stage). The
embryo is enclosed within a seed where it undergoes a period of inactivity
(dormancy). The resting embryo resumes growth during the germination
of seed and develops into a seedling (seedling stage).

The seedling grows into a vegetative plant (vegetative phase). After some
period of vegetative growth, the plant undergoes maturation and enters
the reproductive phase. It develops flowers and fruits, the latter containing
the seeds. Finally senescence sets in (senescence stage) leading to the
death of the plant.

In unicellular organisms, growth consists of an increase in the size or

volume (enlargement) of the cell. This increase is due to the synthesis of
new protoplasm. Growth in unicellular organisms thus consists of single
phase or step. Growth leads to maturation (“adults”) or full grown
individuals. Cell division in unicellular organisms results in their
multiplication or reproduction.

In simple multicellular organisms like Spirogyra, growth involves two

phases or steps, cell division and enlargement. Cell division results in
increase in the number of cells in the filamentous alga. The newly formed
cells enlarge or increase in size. As a result, the filament of Spirogyra
grows. In, flowering plants, however, growth involves three phases cell
division, enlargement and differentiation.
Growth Regions in Animals and Plants:
Cell division and differentiation are important aspects of growth and
development in both animals and plants. In mammals, the growth is
diffuse and it is very difficult to specify the regions where the growth
occurs. In animals, the growth of the embryo is completed quite early,
although the mature size may be gained at specific periods.

In plants, the growth may be diffuse or localized. Diffuse growth occurs in

lower forms of life i.e., filamentous algae. Here each cell of the
multicellular plant body can divide and enlarge. The higher plants,
especially, the trees are built up in a modular fashion i.e. their
development is relatively open-ended and their structure never complete.

In such plants, the growth continues throughout with the new organs
forming, replacing the old ones. Here the growth is localized i.e. growth is
confined to certain specific regions, the growing points. Localized growth
occurs due to the activity of a group of cells called the meristems.
Depending upon the location of the meristems, the growth may be apical,
intercalary and lateral.

Phases of Plant Growth:

As a plant is made up of cells, its growth will be the sum total of the
growth of its cells.

The growth of cells involves three main phases:

(1) The phase of cell division (formative phase),

(2) Cell enlargement and cell differentiation.

(3) Cell Differentiation or Cell Maturation.

1. Phase of Cell Division (Formative Phase):

Cell division is the basic event for the growth of multicellular plants. All
cells in an organism result from the division of pre-existing cells. The type
of cell division that occurs during the growth of an organism is mitosis. It
is a quantitative as well as qualitative division that is generally completed in
two stages: the division of the nucleus (karyokinesis), followed by the
division of the cytoplasm (cytokinesis).

During mitosis, the cell passes through prophase, metaphase, anaphase

and telophase, resulting in equal distribution of the genetical material and
the cytoplasm in each of the two daughter cells thus formed. Further, the
daughter cells are genetically similar to the parent cell. As a result of this
process, cells having the same genetic constitution get multiplied.

In higher plants, cell divisions continuously occur in the meristematic

regions, such as apical meristem. As a result, an increase in the number of
cells takes place in the meristematic region. Some of the daughter cells
retain the meristematic activity, while others enter the next phase of
growth— the phase of cell enlargement.

2. Phase of Cell Enlargement:

The cell enlargement plays an important role in contributing to the size of
the tissue and organs. The enlargement occurs by synthesizing
protoplasm, absorbing water (hydration), developing vacuoles and adding
new cell wall material to the stretched, thin elastic walls to make them
slightly thicker and permanent. Cell enlargement may be linear or in all
directions.

3. Phase of Cell Differentiation or Cell Maturation:

During the last phase, the enlarged cells eventually acquire a specific size
and form according to their location and role following biochemical,
physiological and morphological changes, i.e., the cells undergo
specialization or transformation. As a result, various kinds of cells get
differentiated. These differentiated cells form different kinds of simple
and complex tissues which perform different functions.

Experiment to Study Phases of Growth:

Germinate a few seeds of Pea or Bean in moist saw dust. Pick up a couple
of seedlings with straight radicle of 2-3 cm length. Wash the seedlings.
Blot the surface water. Mark the radicles from tip to base with 10-15
points at intervals of 2 mm with the help of water proof or India ink. As
soon as the ink dries up, place the seedlings on moist blotting paper in a
petri dish. Allow the seedling to grow for 1-2 days. Measure the intervals
between the marks.

Growth Rates:
The increased growth per unit time is termed as growth rate. Thus, rate of
growth can be expressed mathematically. An organism, or a part of the
organism can produce more cells in a variety of ways. The growth rate
shows an increase that may be arithmetic or geometrical (Figure 2.2).

In arithmetic growth, following mitotic cell division, only one daughter cell
continues to divide while the other differentiates and matures. The
simplest expression of arithmetic growth is exemplified by a root
elongating at a constant rate. Look at (Fig. 2.3). On plotting the length of
the organ against time, a linear curve is obtained.
Mathematically, it is expressed as:
Lt = L0 + rt
Lt = length at time ’t’
L0 = length at time ‘zero’
r = growth rate/elongation per unit time.

Let us now see what happens in geometrical growth. In most systems, the
initial growth is slow (lag phase), and it increases rapidly thereafter – at an
exponential rate. Here, both the progeny cells following mitotic division
retain the ability to divide and continue to do so (Fig. 2.4). Geometrical
growth can be expressed by “Grand Period of Growth” (Fig. 2.5).
Quantitative comparisons between the growth of living system can also be
made in two ways:
(i) Measurement and the comparison of total growth per unit time is
called the absolute growth rate,

(ii) The growth of the given system per unit time expressed on a common
basis, e.g., per unit initial parameter is called the relative growth rate.

In Figure 2.6 two leaves, A and B, are drawn that are of different sizes but
shows absolute increase in area in the given time to give leaves, A, and B’.
However, one of them shows much higher relative growth rate. Which
one and why?

The Grand Period of Growth:

The vegetative growth of most plants in general shows three phases,
starting slowly, becoming gradually faster and finally slowing again. These
three phases, which are together known as “grand period of growth”,
cover the whole of the vegetative history of an annual plant. In a perennial
plant such a grand period of growth is repeated annually with periods of
dormancy between the repetitions.

In order to explain the grand period of growth, a graph may be drawn

between the duration of growth and increase in the dry weight of the
plant. It is graphically represented by a ‘S’-shaped curve (a sigmoid curve)
(Fig. 2.5). These variations in growth occur due to several external and
internal factors.

The sigmoid curve shows following three distinct phases:

(1) The lag phase or initial phase:
It represents initial stages of growth. The rate of growth is naturally slow
during this phase.

(2) Log phase or exponential phase:

It is the period of maximum and rapid growth. Physiological activities of
cells are at their maximum.

Here, both the progeny cells following mitotic cell division retain the
ability to divide and continue to do so. However, with limited nutrient
supply, the growth slows down leading to a stationary phase_

The exponential growth can be expressed as

W1 = W0ert
W1 = final size (weight, height, number etc.)
W0 = initial size at the beginning of the period
r = growth rate

t = time of growth

e = base of natural logarithms

Here, r is the relative growth rate and is also the measure of the ability of
the plant to produce new plant material, referred to as efficiency index.
Hence, the final size of W1 depends on the initial size, W0.
(3) Adult phase or stationary phase:
This phase is characterized by a decreasing growth rate. The plant reaches
maturity, hence the physiological activity of cells also slows down and
plant begins to senesce.

Factors Affecting Plant Growth:

(I) External Factors:
Regardless of the habitat in which a plant is growing, it is continuously
subjected to the variability’s of a complex set of environmental factors.
Environmental factors play an important role in the growth and
development of any plant. Important among these environmental factors
are temperature, light, oxygen, water and nutrients.

(1) Temperature:
Temperature is one of the most important environmental factors that
effect the growth of any plant. However, the minimum, optimum and
maximum limits of temperature for growth vary from species to species.
For instance, the winter cereals make some growth at temperatures of 34°
to 40°F, whereas in that temperature range pumpkins and melons do not
grow ,it all.

As the temperature increases above the minimum, growth is accelerated

until a certain optimum temperature is attained, above which it becomes
slower and ultimately completely retarded. The optimum temperature
greatly varies with the species of plant; it also varies with the age of the
plant. The optimum temperatures for the growth of tropical plants are
higher than the temperate ones.

Arctic and alpine species may grow at the freezing point or even at a
temperature slightly below the freezing point. Their optimum temperature
is usually no higher than 10°C. The optimum temperature for most of the
tropical species varies from 30° to 35°C, and for temperate species it
usually varies from 25° to 30°C.

Effect of duration for which a plant is exposed to a particular temperature

also varies with the species. For instance, a plant may make considerable
growth if exposed to a temperature of 86°F for a short duration—the same
temperature has deleterious effects on growth if maintained for a longer
duration.

Soil temperature also greatly influences the growth of roots and shoots.
Under natural conditions, temperature is a cyclic environmental factor.
Normally the temperatures of day and night greatly vary and with only few
exceptions plants grow better when night temperatures are lower than day
temperatures. Sometimes, the term thermo-periodicity is used to
designate the effects of an alternation of temperature between the day and
night upon the growth and other reactions of the plants.

(2) Light:
Light is another important factor that variously effects the growth and
development of all plants. Light intensity, quality of light and duration of
light affect growth through several ways. It greatly influences several
important physiological processes like chlorophyll synthesis, stomata
movements, photosynthesis, formation of anthocyanin, temperature of
aerial organs, absorption of minerals, permeability, rate of transpiration,
streaming of protoplasm etc.

(i) Intensity of light:

The intensity of light greatly influences plant growth. Variations in the
intensity of sunlight are always invariably associated with changes in the
quality of light, and under natural conditions, variations in light intensity
have more significant effects upon growth pattern of plants than changes
in the quality of light. Most crops and ornamental plants, for instance,
wheat, corn, peas, tobacco make vigorous and stocky growth and flower
profusely with full sun. Such plants are called “sun plant”.

When grown with intermediate light intensities, sun plants become taller
and have larger, thinner leaves, but fewer flowers. They make very poor
growth in low light intensity. Shirley (1929, 1935), however, observed in a
number of plant species that the absolute weight, percentage of dry matter
in the tops, thickness and rigidity of the stem and leaf thickness all
increase with increase in the light intensity up to full sun light, provided no
other factor is limiting. Low light intensity results in poor flower
development and consequently very poor fruit setting.

(ii) Quality of light:

Different wave lengths of sun light have significant effects upon the growth
of plants. Most of the experiments conducted in this direction indicate
that overall development of a plant and increase in its dry weight take
place most effectively in the full spectrum of visible light. Plants grown in
blue and violet light tend to be dwarf, those in red light, tall and spindly.
The ultraviolet and infra red radiations of sunlight do not promote
growth.

Overall growth of plant in green light is much less than in either blue-
violet or orange red portions of the spectrum. This effect of green light is
partly due to lower efficiency of photosynthesis in the green light.
Different wave lengths of sun light do not have uniform effects on
different organs of a plant. For instance, orange-red light generally results
in poor development of stems and hypocotyls.

Greatest elongation of stems and hypocotyls in most of the plants takes

place in blue-violet portion of the spectrum, less in the green and still less
in the orange-red and least in the complete spectrum of visible light. On
the other hand maximum expansion of leaf blades occurs in the full
spectrum of visible light and least in the green.

(iii) Duration of light:

Duration, intensity and quality of light have marked influence on the rate
of photosynthesis and hence the rate of growth. During winters, when the
days are short, plants grow slowly; as the days get longer toward spring,
growth is accelerated.

Duration of light not only affects photosynthesis but also greatly influences
dormancy and flowering in plants. The short days of autumn bring about
retardation of growth in many plants, a phenomenon not related to
photosynthesis. A number of trees respond to the short days of autumn
by ceasing to grow and becoming dormant.

The length of day has a marked influence on flowering. Plants, according

to their requirement of light for flowering, are classified as-long-day plants,
short-day plants and day-neutral plants. The long-day plants in general
flower when the days are longer than 13 or 14 hour (depending upon the
species), while the short-day plants produce flowers when the days are
shorter than 13 or 14 hours. Flowering in the day-neutral plants is not
affected by the length of the day. They can very well flower under both
short and long day conditions.

(3) Oxygen supply:

With the exception of only those plants which are native to marshy and
boggy terrains, the growth of all terrestrial plants is greatly retarded in
poorly aerated soils. Usually the shoots of plants receive an ample supply
of oxygen, but the roots may or may not get sufficient oxygen to grow and
function normally. Plants in flooded fields or in water logged pots do not
thrive due to marked deficiencies in soil aeration. The retarded growth of
plants in poorly aerated soils is chiefly due to reduced absorption of
minerals and water.

(4) Water:
Water is one of the most essential requirements for growth of a plant.
With an inadequate water supply, growth is poor and yields low. Plants
grow well when ample but not excessive moisture is available. For most of
the plants a soil-water content in the capacity to just above the wilting
percentage is most favorable tor good growth. With a decrease in the soil-
water content, marked effects on growth do not appear until the
permanent wilting percentage is reached. At the permanent wilting
percentage all growth ceases. If the soil is continuously above field
capacity, as it may be in poorly drained fields, plants grow slowly because
roots are deprived of oxygen.

Plants vary in their response to moisture deficiency. For instance,

radishes, spinach and peppers wilt and cease to grow when soil-water
percentage is low. Cucurbits and tomatoes in the field stop growing and
their lower leaves respond by changing from a light green colour to a
darker green or bluish colour. The leaves of corn and many grasses curl
when the water supply is inadequate.

Deficient soil-water supply may affect the growth of a plant more at certain
stages in its development than others. Vegetative growth in many plants is
checked but the development of reproductive organs is not affected under
deficient soil-water supply.

(5) Soil Nutrients: The quantity and nature of soil nutrients have marked
influence on the growth and development of plants. For luxuriant growth
of any crop, field should be adequately rich in nutrients (both micro and
macronutrients). Furthermore, these mineral nutrients do not effect the
growth as such, but only when present in the form of ions, or as
constituents of molecules.
II. Internal Factors:
(1) Growth Regulators: Several classes of growth regulators are known.
While some growth regulators are growth promoting (e.g., auxins,
gibberellins, cytokinins, florigen etc.), others are growth inhibitors (e.g.,
abscisic acid, ethylene, chlorocholine chloride). Many of them are
synthesized by the plants, while few of them are synthetic.
(2) C/N Ratio: The ratio of carbohydrates and nitrogenous compounds
governs the pattern of growth. Presence of more carbohydrates compared
to nitrogenous compounds favours good vegetative growth, flowering and
fruiting. On the contrary, presence of more nitrogenous compounds
compared to carbohydrates results in poor vegetative growth, flowering
and fruiting.
(3) Genotype and Genetic Factor: All metabolic activities, growth and
development are under the control of genetic complement (genotype),
nuclear, as well as extra nuclear, of the cell. Expression of appropriate
genes in an appropriate sequence is controlled both by genes and the
environment. The genes, located in chromosomes, transcribe information
to m-RNA which translates it into structural and enzymic proteins.