GROWTH AND DEVELOPMENT

Growth is defined as an irreversible or permanent increase in size and dry weight of an organism.
It is caused by cell division mainly mitosis. Cell division leads to increase in cell number, also
involves cell expansion and cell differentiation in cell differentiation the cells become
specialized to perform particular functions i.e. division of labor. They undergo changes both in
shape and form and join others of similar kind to form tissues or organs. Such cells do not divide
any more therefore growth is usually accompanied by an increase in complexity of an organism
called development.

What is development?

It is the process whereby cells after division differentiate and increase in complexity leading to
changes both in shape and form. The process of development is closely linked with growth in
that growth and development are commonly used to describe processes which are thought of as
growth. In multicellular organisms, growth is divided into 3 phases:

i) Cell division (hyperplasia) which is an increase in cell number as a result of cell division
ii) Cell expansion (hypertrophy) which is an irreversible increase in cell size as a result of
uptake of water or synthesis of living material
iii) Cell differentiation: this is when cells become specialized. This also includes development.

All stages of growth involve bio chemical activity. Protein synthesis is important since it is the
means by which the DNA message is expressed in terms of enzymes synthesized by the cell.
The enzymes control cell activities. The changes at the cell level bring about the changes of the
overall form and structure both of individual organs and the organism as a whole.

Growth may be positive or negative. Positive growth occurs when anabolism exceeds catabolism
whereas negative growth occurs when catabolism exceeds anabolism. E.g. during germination
of a seed and the production of seed structures, various parameters increase in magnitude such as
cell number, cell size, fresh mass, length, volume and complexity while others such as dry mass
may actually decrease.

Types of growth

1. A twelve year old boy after following the same diet for 1 year can gain 3kg in weight.
2. A tree which is well watered increases in height by 10cm and in trunk circumference or girth
by 1cm.

Examples of development

A fertilized ovum of a mammal gives rise to relatively undifferentiated cells of the embryo, some
of those cells differentiate to form nerve cells specialized for receiving and conducting impulses,
and some differentiate into erythrocytes for transporting O2.

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In all organisms, growth involves synthesis of new cellular substances from raw food materials, a
process called assimilation.

The accumulation of new cellular materials results in cellular enlargement. Cells cannot enlarge
indefinitely because enlargement leads to a decrease in S.A to V.R and this in turn lowers the
cells capacity for exchange of materials with its environment. This problem is solved by cell
division where by one cell gives rise to two daughter cells and each daughter cell takes in raw
materials, increase in size and divides once again. In multicellular organisms, cell division leads
to an increase in the number of cells and consequently growth of the individual. In unicellular
organisms, cell division leads to an increase in the number of cells in the population, i.e. it leads
to growth of the population. In multicellular organisms growth is always accompanied by
development.

MEASUREMENT OF GROWTH

Growth is estimated by measuring some parameters or variables over a period of time. The
parameter used depends on the organism whose growth is to be determined. The parameters
used may include:

 Length (height) this may however be misleading in case of a bush because growth in length
may stop but it continues growing sideways.
 number of leaves
 girth or circumference of a tree
 Fresh weight/mass i.e. mass of an organism under normal conditions. This is very easy to
measure and may not damage the organism. However, it is inaccurate due to temporary
fluctuations in water content and it is also not very easy to determine the fresh mass for large
organisms like big trees.
 Dry weight or mass, this involves removing all water by drying the organism before
weighing. However it is difficult to carry out and permanently destroys the organism
involved but gives an accurate measure of growth.
 Area e.g. leaf area
 volume

GROWTH PATTERNS

When any parameter of growth is measured against set intervals of time, a growth curve is
produced. For many populations organisms/organs the curve is S shaped and called “sigmoid
curve”

Sigmoid growth curve

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A sigmoid curve is divided into the following parts:

1. Large phase
This is the initial phase during which little growth occurs. The organisms are adapting to the
new environment and multiplying very slowly.
2. Acceleration phase
This is also referred to as rapid growth phase or phase of exponential growth. The organisms
are already adapted to the new generation.
3. Steady phase (4)
This is also referred to as constant growth phase. This phase, there is deficiency of nutrients
and accumulation of toxic waste products. The organisms begin to die resulting in an
equivalent between rate of death and the rate of multiplication.

N.B the two phase i.e. accelerating and steady phases constitute the log phase during which
growth proceeds exponentially. During this phase, the rate of growth is proportional to the
amount of material or numbers of cells already present.

The nature of the curve during stationery phase or steady phase marks the period where overall
growth has ceased and the parameter under consideration remains constant. It may also vary
depending on the nature of the parameter, the species and internal factors.

In some cases the curve may continue to increase slightly until the organism dies. This indicates
positive growth which is typical of many invertebrates, fish and some reptiles. In some cases like
in cnidarians (coelenterates) the curve flattens out indicating no change in growth. In others,
growth curves trail off indicating a period of negative growth; this is typical of many mammals
including humans and is called negative growth. It is a sign of senescence with increasing age.

TYPES OF GROWTH CURVES

1) The absolute growth/ actual growth curve.

This is obtained by plotting data from any physical parameter such as dry mass against time.
This curve shows the overall growth pattern and the extent of growth.

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 Graph in B.S page 760

2) Absolute growth rate curve

This is obtained by plotting the change in the parameter against time. It can be obtained also by
calculating the slope of the absolute growth curve at various points and plotting these against
time. This curve shows how the rate of growth changes during tie of study. The peak of the
absolute growth rate curve marks the point of inflexion after which the rate of growth decreases
as the adult size is attained.

Graph in B.S page 760

3) Relative growth rate curve/specific rate growth

This is obtained by dividing the values obtained from each scope of absolute growth rate by the
aunt of growth at the beginning of time period and plotting the values against time. OR this is
measure of efficiency of growth.

Graph in BS page 760

Relative growth rate curve is also increase in growth over a period of time expressed as a
percentage of the growth that has already taken place hence relative growth rate curve may be
called percentage growth curves.

ISOMETRIC AND ALLOMETRIC GROWTHS

Isometric growth occurs when an organ grows at the same mean rate as the rest of the body. In
this situation, change in size of the organism is not accompanied by a change in shape or external
form of an organism, i.e. the proportion of the 2 structures remains the same. This type of
growth is typical of fish, insects e.g. locusts, except for the wings and genitalia, leaves of most
plants etc.

Allometric growth occurs when an organ grows at different rate from the rest of the body. This
produces a change in size of the organism which is accompanied by a change in shape of the
organism. This pattern of growth is characteristic of mammals. The relative proportions of
growth and development are shown in the figure below:

Allometric growth as shown by Human organs and tissues

Graph in BS page 762 fig. 22.9

Lymph tissue which produces white blood cells to fight infection grow rapidly early in life when
the risk of diseases in greater as immunity has not been yet acquired. By adult life, the mass of

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lymph tissue is less than half of what it was in early adolescence because of the immunity
acquired.

Reproductive organs grow very slowly in early life or remain small in size but develop rapidly
with an onset of sexual maturity (puberty) when they grow rapidly to adult size.

The human head and brain grow very early in life reaching about 90% of adult size at 6 years.
The human brain grows early in life to enable the organism or infants coordinate the changes
within the new environment. Growth of the rest of the body is fast during infancy slowing during
childhood and then growing faster after puberty. The result is that the human fetus and infants
have a head that is large relative to the rest of the body as compared to an adult. As a person
grows, the head grows slowly relative to the rest of the body resulting in an overall change in the
shape of the person.

LIMITED AND UNLILMITED GROWTH

LIMITED GROWTH (DEFINITE)

It is found in annual plants and majority of animals. In limited growth an organism grows to an
average maximum size which is characteristic of the species and no further growth. This is
followed by senescence (decline in old age and death). It is also observed in insects and birds.
The growth curve obtained is typically sigmoid except there may be an initial decrease in mass
during early stages of germination in plants. And once leaves start photosynthesizing, growth
proceeds in a sigmoid, and when the seeds and fruits are released at the end of the growing
period, the mass of the plant may decrease prior to death.

Growth curve of an annual plant

Diagram in BS page 762 fig. 22.10

UNLIMITED GROWTH

In perennial plants, the growth pattern is an annual series of sigmoid curves. They grow
continuously throughout their lives i.e. their curve never flattens out. They have an unlimited or
indefinite growth i.e. they have a continuous growth curve. It is also found in fungi, algae,
animals particularly invertebrates, fish and reptiles.

Diagram in BS page 762 fig. 22.11

INTERMITTENT GROWTH (Discontinuous growth)

Crustaceans and other arthropods such as insects have a characteristic growth pattern. As their
exoskeleton is incapable of expansion, they have to moult periodically during growth. Before a
new skeleton has fully hardened it is capable of some expansion. During this, time insect may

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take up water in order to expand the exoskeleton as much as possible. This gives room for
further growth and the organism changes abruptly to a new size after every moult.

A plot of length of such an organism against time gives a characteristic star-like curve. The steep
portions correspond to the time of moulting. The flat portions correspond to the stages between
moults also known as instars.

Growth curve of short horned grass hopper

Graph in BS page 763 fig. 22.12

GROWTH AND DEVELOPMENT IN PLANTS

GERMINATION

According to ISTA (International seed Testing Association) 1966; Seed germination is the
emergency and development from the embryo of those essential structures which indicate ability
to develop into a normal plant under favorable conditions within the soil.

Germination takes place in seeds when provided with water, O 2 and optimum temperature. It
begins with rapid uptake of water through the micropyle which is accompanied by considerable
swelling and an increase in mass of the seed. Swelling of the inner tissues causes rapture of the
soft seed coats. The absorbed water activates enzymes, thus leads to hydrolysis of food materials
e.g. starch and protein which are stored in the cotyledons or endosperm. The soluble food
materials are trapped to the growing points of the embryo where they are used for reproduction
to provide energy and synthesis of new cellular materials. The radical is the first to emerge, it
grows down wards between soil particles, and root hairs develop a short distance from the root
cap and start absorbing water and mineral salts.

Types of germination

Germination is of 2 types;

i) Epigeal germination

This is where the cotyledons appear above the ground due to rapid elongation of the hypocotyls.
E.g. beans, tomato, cotton, mangoes, etc. The cotyledons on exposure to sun light turn green and
become photosynthetic. This is because they contain proto-chlorophyll which is transformed
into chlorophyll. This is when they assume the function of making food.

ii) Hypogeal germination

The cotyledons remain underground due to the epicotyls growing faster than hypocotyls. Seeds
showing hypogeal germination are endospermic. Examples include maize and G nuts.

Environmental conditions necessary for germination

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Seed germination is only possible under conditions that are suitable for germination, they
include: water, light, soil structure, Temperature, O2 and Micro organisms

PHYSIOLOGY OF GERMINATION

A typical seed stores carbohydrates, lipids and proteins either in its endosperm or cotyledon of
the embryo. Lipids are usually stored in form of oils mainly by legumes like ground nuts. In
grasses including cereals, starch is the major food reserve. Legumes also store proteins. In
addition seeds store mineral salts e.g. phosphorus and vitamins. Food reserves in seeds are
insoluble in water and cannot be trapped in the seedling. They must be broken down into
relatively simple soluble substances which dissolve in water to be moved to the growing apices
of the plumule and radicle.

i) Carbohydrates:

They are hydrolyzed to starch and into soluble disaccharide sugar maltose catalyzed by enzyme
amylase. Maltose is for the hydrolyzed to glucose by maltase. `

Maltose is converted to sucrose for translocation to the growing apices of the embryo. Sucrose is
used for synthesis of cellulose, hemicelluloses and pectin compounds which are the main
components of the cell wall. Sucrose is also respired to provide energy for growth.

Summary:

Starch amylase maltose maltase glucose respiration, cellulose, hemicellulose,

pectin compounds

ii) proteins

They are hydrolyzed to peptides and amino acids by enzyme peptidase. Some amino acids are
moved in solution to the embryo. At the growing points of the plumule and radicle, amino acids
are used to synthesize structural and enzyme proteins.

Proteins peptidase polypeptides peptidase amino acids form structure in growing apices
and enzyme proteins

iii) lipids

The fats and oils are first hydrolyzed to fatty acids and glycerol by lipase enzyme. The fatty
acids may be oxidized to release energy, converted to sucrose for transport or used for membrane
synthesis. Glycerol is also converted to transportable sugars.

A well-studied example of germination of a polysaccharide-rich seed is that of barley grain.

Relative changes in dry mass of endosperm and embryo during germination of barley

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Question: the table below shows the changes observed in dry weight in mg of a barley seedling,
its embryo and endosperm during the first ten days after onset of germination.

Time/days Embryo Endosperm Whole seedling

0 2 41 45
2 2 39 43
4 7 32 41
6 15 21 38
8 22 11 35
10 35 6 43

a) Suggest how the experiment was carried out

b) Using a suitable scale and on the same set of axes plot graphs of dry weight of embryo,
endosperm and whole seedling against time.
c) Describe and account for the changes in weight shown by:
i) The embryo
ii) The endosperm
iii) Whole seedling during the period of the experiment.
d) Explain how you would expect the weight of the whole seedling to change if the experiment
was carried out in the dark.

Question:

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A study was conducted on the germination and early growth of sorghum. The grains were soaked
in cotton wool in a green-house and at two – day intervals, samples were taken and separated
into two components, of endosperm and embryo (seedling), which were then oven dried and
weighed. Figure 1 shows the variation of total dry mass, dry mass of endosperm and embryo.

Use the information to answer the questions that follow.

(a) Explain the variation with time of

(i) Dry mass of endosperm
(ii) Dry mass of embryo
(iii) Total dry mass of sorghum seedling.
(b) Explain why the following were done.
(i) Oven – drying the seeds
(ii) Separating the seed components
(iii) Sowing seeds in a greenhouse
(c) From the information given, name the method used to measure growth and give its
limitations.
(03 marks)

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 (d) (i) Name two internal factors in the seeds that would affect the results above.
(02
marks)
(ii) Suggest precautions that could have been taken to ensure reliable results.
(02
marks)
(e) What conclusion can be drawn from the graph after 8 days?
(01
marks)
(f) Explain what would happen if the experiment continued for another 10 days.
(04
marks)

RESPIRATION IN GERMINATING SEEDS

Respiratory rates in both storage tissues (food reserves) and embryo (growth centre) are high
owing to the intense metabolic activity in both regions. Substrates for respiration may differ in
each region and may also change during germination. This is revealed by changes in the
respiratory quotient.

Question: when castor oil seeds were analyzed for lipid and sugar content during germination in
darkness, the results obtained are shown in the figure below.

Changes in lipid and sugar content of castor oil seeds during germination in dark

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The R.Q of the seedlings was measured at day 5 and the embryo was found to have an R.Q of 1.0
while the remaining cotyledons had an RQ of 0.4 – 0.5

a) Suggest an explanation for these results.

b) What would you expect the RQ of the whole seedling to be on day 11? Explain briefly
c) Why was the experiment carried out in the dark?

Solutions:

a) Caster seeds have food reserves in form of lipids. By day 4 the lipids begin reducing while
the mass of sugars by day 4 increase. This is because during germination the lipids are
hydrolyzed to fatty acids and glycerol and the glycerol is converted into sugars, and then
transported to the embryo where it’s oxidized to yield energy among other uses.
At day 5 when the sugar is oxidized the R.Q is 1.0. In the cotyledons the RQ is between 0.4-
0.5 indicating that the lipids are the ones oxidized to yield energy.
b) At day 11, the R.Q of the whole seedling will be slightly less than 1.0 this is because at this
particular time all the lipid food reserves have been converted into sugars whose oxidation
gives an RQ of 1.0 and a small unit of lipids present when oxidized gives an RQ of between
0.4 – 0.5. A combination of both gives an R.Q slightly less than 1.0.

N.B conversion of lipids to sugar leads to increase in dry mass of seedlings between day 6 and
day 7 beyond this point lipid reserves are being exhausted and the rate of use of the sugars starts
to exceed its rate of production. This results in a decrease in the net mass of sugars and the total
mass of the seedling. The sugar is used in respiration and anaerobic reactions.

SEED DORMANCY

This is a state in which a seed though viable will not germinate under conditions considered to be
adequate for germination. Dormant seeds usually have low metabolic activity, they have low
water content and ‘Zero’ growth rate i.e. the embryos do not grow in any way. There are 3 types
of seed dormancy:

i) Innate (primary dormancy):

In this type of dormancy seeds after dispersal cannot germinate immediately and majority of
seeds fall under this category. Such seeds only germinate after a period of ‘after ripening’ or
storage. This period leads to changes which are needed to improve germination.
ii) Induced dormancy (secondary dormancy):
The seeds achieve dormancy because of one factor missing.
iii) Enforced dormancy:
Is the one which is thrust onto the seeds by storage e.g. stores, refrigerators. Etc.

Causes of seed dormancy

They are divided into 3 categories

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 i) Environmental factors
ii) Anatomical structure of the seeds
iii) Internal physiological state of the seed and timing mechanism
i) Environment factors
 Lack of adequate water supply
 Lack of adequate oxygen
 Requirement for light in case of positively photoblastic seeds
 Requirement for dark in case of negatively photoblastic seeds.
 Requirement for suitable temperature range i.e. at high temperature, water is not
available and at very low temperature, water is frozen.
ii) Anatomical structure of the seeds
 Hard seed coats mechanically resisting emergency of seedling parts.
 Hard seed coats impermeable to water and air.

iii) Internal physiological state of the seeds timing mechanism.

 Embryological immaturity
 Embryological dormancy
 Presence of germination inhibitors like abscisc acid.
 Absence of germination promoters such as gibberellic acid, cytokines, etc.
 General after ripening requirement.

Breaking of seed dormancy

1) Under natural conditions fire exposes seeds to radiation and causes seeds to germinate
2) Clearing vegetation cover. This works for positively photoblastic seeds.
3) By attaining high and low temperature.
Some seeds in order to break dormancy need to be hydrated at low temperature (between 4-
100C). This is called stratification (cold storage). Stratification encourages embryo
development, synthesis of germination promoters e.g. Gibberellic acids and decrease of
germination inhibitors like ABA. It also encourages improvement of coat permeability.
4) Hard seeds coats are broken by:
 Break down by microorganisms in soil for example bacterial and fungi.
 Digestive actions of enzymes of mammals and birds. This works well for seeds of red
pepper, passion fruits, etc.
 Exposure to alternating high and low temperature
 Treatment of seeds using appropriate chemicals e.g. concentrated sulphuric acid and
alcohol.
 Clipping or breaking off pieces of seed coats.
5) Dormancy due to chemical inhibitors is broken by stratification and treatment of seeds using
germination stimulators like gibberellic acid and cytokines.
6) Dormancy as a result of embryo immaturity and embryo dormancy is broken by allowing
after ripening period and stratification.

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 Importance of after ripening

 During after ripening, there is completion of embryological development i.e. when the
embryo is immature.
 Physical and chemical changes may take place within the seed leading to improved
germination.
 There may be germination inhibitors in the seeds and these may disappear during after
ripening period.
 Germination stimulators may be synthesized during after ripening period leading to improved
germination like gibberellic acid.
 Chemical composition of storage/reserve materials will change leading to improved or
delayed germination e.g. Abscisc acid (ABA)

The requirement for this period varies from one species to another so in some varieties for
example barley is allowed more time for shading from their parent plants.

Importance of seed dormancy

i) It allows seeds to be dispersed from their parent plants before germinating thus avoiding
overcrowding and competition around the parent plants.
ii) It ensures that germination occurs when conditions are favorable for growth. E.g. in many
temperate plants, dormancy is broken through exposure of seeds to winter cold. This
ensures that seeds are ready for germination in early spring so that they can grow through
the rest of spring and summer when temperature is suitable for growth.
iii) It allows the embryo to develop to maturity.

Longevity of seeds

This is the time the seeds last before they lose ability to germinate following shading from the
parent plants. The wild species seeds last longer than the cultivated relatives. E.g. ipomeas spp
seeds can germinate after a period of about 40 years, the Indian lotus spp can germinate after a
period of about 1000 years.

Measurement of plant growth

1. Size:

This includes length, area or volume of a single cell, tissue or organ or organism. This parameter
is difficult to measure when the plant becomes so complex. Measurement of size is applicable to
early stages of development.

2. Weight

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This is the commonly used parameter. This is normally associated with increased size of a plant.
We can increase fresh weight or dry weight of a plant.

i) Fresh weight:

This is the weight of a plant including water. However, this method is misleading because the
water content of a plant fluctuates with time of the day. E.g. in the morning fresh weight is high
because of low transpiration rate and fresh weight is low in the afternoon and evening. This
method is also destructive to the plant as it involves uprooting the plants so it is not applicable to
larger rooted plants. Generally this method is inaccurate.

ii) Dry weight:

This is the weight of a plant excluding water. It is a method widely used. It is more accurate than
fresh weight method. However this method has short comings like;

 It is destructive to the plant as it requires drying.

 During the process of weighing and drying there is possibility of loss of material which
gives inaccurate results.
 During the drying process some volatile materials may be lost.
 Plants accumulate inorganic salts which are incorporated into the organic structure of the
plant body and during the drying the process, they contribute to the dry weight giving
false results.
 Seedlings normally lose weight because they use stored food for embryo respiration.

This method is not applicable to large plants.

3. Length:

Applied to simple plant organs such as the stem, increase in length of leaves, length of branches
and length of roots. This method is restricted to early stages of plant growth.

4. Area:

This is applied to leaf area and simple leaves. This method is appropriate to plants with complex
morphology.

5. Quantitative changes in protoplasm

This method gives the fundamental measure of growth in plants however the method has short
comings and these include:

 The method is very difficult to manipulate

 Normally the total protein content of the tissue cannot be estimated because protein forms
most of the protoplasm. The protein content may be estimated by determining the nitrogen

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 content of the cell. Nitrogen forms a very small percentage of the total weight of proteins. A
tissue is burnt and nitrogen content turns into nitrogen gas which forms an acid.

6. Complexity of the plant

It is measured by increase in number of various organs e.g. leaves, flowers, fruits, etc. this can be
taken as a measure of growth but the method becomes difficult in case of very large plants.

7. Differentiation

This is when you determine the change in number of types of cells as the plant grows. This
method requires the use of microscope. Normally a cell divides, it expands and then
differentiates. However the method is destructive to plants.

TYPES OF GROWTH IN PLANTS

In plants with exception of the young embryo, growth is confined to certain regions known as
meristems. A meristem is a group of cells with the ability to divide by mitosis to form daughter
cells which grow to form the plant body. Meristems are of 3 types which include:

Apical meristems:

These are found in the shoot and root apex. I.e. shoot apical meristem and root apical meristem.
They are responsible for primary growth leading to increase in length.

Lateral meristems:

These are found in older parts of the plant and are parallel with long axis e.g. the cork cambium
and vascular cambium. Lateral meristems are responsible for increase in girth.

Intercalary meristems:

These are found at the nodes of many monocots. Intercalary meristems allow growth of length to
occur in areas other than the tips so are useful in case the tips are damaged and are also
responsible for increase in length.

PRIMARY GROWTH

Primary growth is the first form of growth to occur in a plant. It is the only form of growth found
in monocots and herbaceous plants. Meristematic cells are packed tightly with no air spaces
between them. These cells are known as initials. When they divide, one daughter cell remains in
the meristem while the other increases in size and differentiate to become part of the plant body.

PRIMARY GROWTH OF THE SHOOT

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The structure of a typical shoot apical meristem is shown to have 3 main regions i.e. region of
cell division, cell expansion and the region of cell differentiation.

A longitudinal section of the shoot apex shows that cells of the surface layer are regularly
arranged to form the outer zone of cells known as tunica or protoderm while the inner cells are
irregularly arranged to form the corpus.

The tunica/protoderm gives rise to the epidermis while the corpus includes the procambium
which gives rise to vascular tissues and the pericycle as well as the ground meristem which
produces parenchyma cells and the pith.

In the zone of expansion the daughter cells produced by the initials increase in size by osmotic
uptake of water into the cytoplasm and then into the vacuoles. Increase in the length of stems and
roots are brought about by elongation of cells during this stage. The small vacuoles increase in
size eventually fusing to form a single large vacuole. While this is happening, water flows into
the vacuoles by osmosis pushing the stretchable cell wall outwards.

Expansion phase of growth of a plant cell

The final shape acquired by the cell depends on the thickening of its cellulose walls and this is
the basis of differentiation.

If cellulose is laid uniformly, spherical cell parenchyma results, when cellulose is laid in columns
on the inside of the original walls collenchyma cells result, sclerenchyma cells are formed by
deposition of thick layers of lignin leading to death of cells.

The proto-cambium forms a series of longitudinally running strands whose cells first
differentiate into proto-xylem to the inside and proto-phloem and companion cells to the outside.

The proto-xylem and proto-phloem elements are later replaced by xylem and phloem in the zone
of differentiation. The mature xylem is characterized by deposition of lignin to the inside and this
gives extra rigidity compared with the proto-xylem and it indicates stoppage of elongation.

Development of the shoot also indicates the growth of leaves and lateral buds. The leaves arise
from swellings which form at regular intervals on both sides of the shoot apex. These swellings
are known as leaf primordial.

The leaf primordial then elongates rapidly increasing area to form leaf blades. When the leaves
start growing they develop axillary bud primordial in the axils which have the potential to grow
into lateral shoots and flowers.

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Note: the angle between the main stem and the leaf is called the axil.

At the axil, the axillary or lateral buds develop. The lateral buds have meristematic cells at their
tips whose growth leads to formation of side branches. The point of origin of a leaf is called the
node and the part between the two nodes is called the internode.

(Diagrams in FA page 426)

PRIMARY GROWTH OF THE ROOT

At the tip of the root apical meristem is a quiescent center; a group of initials (meristematic cells)
form which all other cells are thought to come from. The quiescent center has got much slower
mitosis than the cells of the apical meristem surrounding it. The root apex is covered by the root
cap to the outside. The outer most zones are known as protoderm which produces cells which
differentiate into root epidermis and root cap.

Inside the protoderm is the ground meristem which differentiates into the root cortex. Just
beneath the root apex is a single procambium. Some roots have an additional meristematic layer
known as calyptrogen which gives rise to the cells of the root cap. In the zone of differentiation,
strands of sieve tube elements and companion cells appear near the outside of the procambium.
Shortly a similar number of strands of pro-xylem cells alternating with the primary phloem
strands differentiate.

The meta-xylem cells differentiate last. The outermost procambium cells retain the ability to
divide and become the pericycle which produces the lateral roots. The ground meristem
differentiates into parenchyma cells of the root cortex. The inner most layer of the root cortex
differentiates into the endodermis.

The endodermal cells secrete a suberized casparian strip on their radial walls.

The endodermis controls the passage of absorbed water and mineral salts from the cortex into the
root vascular tissue. The root epidermis derived from the protoderm is different from that of the
shoot system in having no cuticle as this would prevent absorption of water and mineral salts.
Just behind the root apex, some epidermal cells develop tubular extensions known as root hairs
which increase the root surface area through which absorption from the soil occurs.

(Diagrams in FA page)

SECONDARY GROWTH

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