Journal of Advanced Laboratory Research in Biology

E-ISSN: 0976-7614
Volume 5, Issue 4, October 2014
PP 152-168
https://e-journal.sospublication.co.in

Review Article

A Review of Plant Growth Substances: Their Forms, Structures, Synthesis

and Functions
Ogunyale O.G.1, Fawibe O.O.1, Ajiboye A.A.2 and Agboola D.A.1*
1
Dept. of Biological Sciences, Federal University of Agriculture; P.M.B 2240, Abeokuta, Ogun State, Nigeria.
2
Department of Biological Sciences, P.M.B. 4494, Osun State University, Osogbo, Osun State, Nigeria.

Abstract: Plant growth substances are compounds, either natural or synthetic that modifies or controls through
physiological action, the growth and maturation of plants. If the compound is produced within the plant, it is called a
plant hormone or phytohormone. In general, it is accepted that there are five major classes of plant hormones. They
are Auxins (IAA), Cytokinins, Gibberellins, Ethylene and Abscisic Acid. However, there are still many plant growth
substances that cannot be grouped under these classes, though they also perform similar functions, inhibiting or
promoting plant growth. These substances include Brassinosteroids (Brassins), Salicylic Acid, Jasmonic Acid,
Fusicoccin, Batasins, Strigolactones, Growth stimulants (e.g. Hymexazol and Pyripropanol), Defoliants (e.g.
Calcium Cyanamide, Dimethipin). Researchers are still working on the biosynthetic pathways of some of these
substances. Plant growth substances are very useful in agriculture in both low and high concentrations. They affect
seed growth, time of flowering, the sex of flowers, senescence of leaves and fruits, leaf formation, stem growth, fruit
development and ripening, plant longevity, and even plant death. Some synthetic regulators are also used as
herbicides and pesticides. Therefore, attention should be paid to the production and synthesis of these substances so
that they affect plants in a way that would favour yield.

Keywords: Phytohormone, Auxins, Cytokinins, Gibberellins, Ethylene, Abscisic acid, Plant Growth Regulators.

CHAPTER ONE

1. Introduction The internodes in the terminal bud are very short so that
the developing leaves grow above the apical meristem
Growth can be defined as an increase in some that produced them and thus protect it. New meristems,
quantity over time. The quantity can be physical (e.g. the lateral buds, develop at the nodes, each just above
Growth in height), abstract (e.g. a system becoming the point where a leaf is attached. When the lateral buds
more complex, an organism becoming more mature). develop, they produce new stem tissue, and thus
Growth can also refer to the quantitative change that branches are formed.
accompanies development. Growth in plants can be Under special circumstances (such as changes in
defined as an irreversible increase in volume of an photoperiod), the apical meristem is converted into a
organism (Taiz and Zeiger, 2002). flower bud. This develops into a flower. The conversion
Growth in plants is obtainable majorly at of the apical meristem to a flower bud "uses up" the
meristems where rapid mitosis provides new cells. As meristem so that no further growth of the stem can
these cells differentiate, they provide new plant tissue. occur at that point. However, lateral buds behind the
In stems, mitosis in the apical meristem of the shoot flower can develop into branches.
apex (also called the terminal bud) produces cells that
enable the stem to grow longer and, periodically, cells 1.1 Measurement of Plant Growth
that will give rise to leaves. The point on the stem
where leaves develop is called a node. The region To capture enough data on the overall growth of
between a pair of adjacent nodes is called the Internode. plants, it is recommended that at least one final weight

*Corresponding author:
E-mail: [email protected].
Plant Growth Substances Agboola et al

measure is recorded, one measure of root health, and all useful for root vegetables such as beets, carrots and
of the observation measurements that pertain to the type potatoes that have a large root (Mulanax, 2005).
of plant being used.
1.1.3 Root Shoot Ratio
1.1.1 Weighing Plants: Fresh vs. Dry Weight
Roots allow a plant to absorb water and nutrients
Measuring Fresh Weight: While the fresh from the surrounding soil, and a healthy root system is
weight of plants can be technically measured without key to a healthy plant. The root: shoot ratio is one
harming them, the simple act of removing a plant from measure to help assess the overall growth of the plants.
its growing "medium" can cause trauma and affect the The control group of plants will provide a "normal"
ongoing growth rate and thus the experiment. root: shoot ratio for each of the plant types, any changes
Measuring the fresh weight of plants is tricky and from this normal level (either up or down) would be an
should probably be saved as a final measure of growth indication of a change in the overall health of the plant.
at the end of the experiment. Here is the process of It is important to combine the data from the root: shoot
measuring fresh weight; Plants are removed from the ratio with data from observations to get an accurate
soil and washed to remove any loose soil. The plants measurement. For example, an increase in root: shoot
are then blotted gently with soft paper towel to remove ratio could be an indication of a healthier plant,
any free surface moisture. They are weighed provided the increase came from greater root size and
immediately (plants have a high composition of water, NOT from a decrease in shoot weight. To measure the
so waiting to weigh them may lead to some drying and root: shoot ratio:
therefore produce inaccurate data). The plants are removed from soil and washed to
Measuring dry weight: Since plants have a high remove any loose soil. They are blotted to remove any
composition of water and the level of water in a plant free surface moisture. They are then dried in an oven
will depend on the amount of water in its environment set to low heat (100 degrees) overnight. The plants are
(which is very difficult to control), using dry weight as left to cool in a dry environment (a Ziploc bag will keep
a measure of plant growth tends to be more reliable. moisture out) - in a humid environment the tissue will
You can only capture this data once as a final measure take up water. Once the plants have cooled, they are
at the conclusion of the experiment (Wood and Roper, weighed on a scale.
2007). The root is separated from the top (cut at the soil
Plants are removed from the soil and washed. line), then both are weighed and recorded separately for
They are again blotted to remove any free surface each plant. (Dry weight for roots/dry weight for top of
moisture. The plants are then dried in an oven set to low plant = root/shoot ratio). The root/shoot ratio can be
heat (100 degrees) overnight. They are left to cool in a calculated for each treatment. Plants contain mostly
dry environment (a Ziploc bag will keep moisture out) - water, so make sure you have a scale that goes down to
in a humid environment the plant tissue will take up milligrams since a dry plant will not weight very much
water. Once the plants have cooled weigh them on a (MBG, 2003).
scale. Plants contain mostly water, so a scale that goes
down to milligrams should be used since a dry plant 1.2 Plant Growth Substances
will not weight very much.
A plant growth regulator is an organic compound,
1.1.2 Root Mass either natural or synthetic, that modifies or controls one
or more specific physiological processes within a plant.
Root mass is recommended as a final If the compound is produced within the plant it is called
measurement as the plant must be removed from its a plant hormone. A plant regulator is defined by the
growing medium in order to capture accurate data. Environmental Protection Agency as "any substance or
There are quite a few different methods for measuring mixture of substances intended, through physiological
root mass depending on the type and structure of the action, to accelerate or retard the rate of growth or
roots. maturation, or otherwise alter the behaviour of plants or
Grid intersects technique: The plant is their produce.
removed from the soil. When working with thin or light Additionally, plant regulators are characterized by
roots, the roots may be dyed using an acidic stain. The their low rates of application; high application rates of
roots are then laid on a grid pattern this is then followed the same compounds often are considered herbicidal
by counting the number of times the roots intersect the (Peggy, 1999). Simply put, plant growth regulators
grid. The roots are traced on paper, with each of the (also known as growth regulators or plant hormones)
tracings measured, and the root length is calculated are chemicals used to alter the growth of a plant or
from the tracings. The numbers of roots are then plant part. Hormones are substances naturally produced
counted and the diameter measured. This is especially by plants, substances that control normal plant
functions, such as root growth, fruit set and drop,

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growth and other development processes. Plant growth plants would be mostly a mass of undifferentiated cells.
regulators are regulated as pesticides by the Florida So they are also called growth factors or growth
Department of Agriculture and Consumer Services hormones.
(FDACS) and must be registered with the FDACS for
lawful use in Florida like any pesticide lawfully used in 1.3 Forms of Plant Growth Substances
Florida.
Plant hormones (also known as phytohormones) According to the American Society for
are chemicals that regulate plant growth, which, in the Horticultural Science, plant growth regulators fall into
UK, are termed 'plant growth substances'. The term six major classes (Fishel, 2006). These classes include
'Phytohormone' was coined by Thimann in 1948. Plant Auxins, Gibberellins, Cytokinins, ethylene generators,
hormones are not nutrients, but chemicals that in small Growth inhibitors and Growth retardants. But in
amounts promote and influence the growth, general, it is accepted that there are five major classes
development, and differentiation of cells and tissues of plant hormones, some of which are made up of many
(Opik et al., 2005). Plant hormones are signal different chemicals that can vary in structure from one
molecules produced within the plant and occur in plant to the next. The chemicals are each grouped
extremely low concentrations. Hormones regulate together into one of these classes based on their
cellular processes in targeted cells locally and when structural similarities and on their effects on plant
moved to other locations of the plant. Hormones also physiology. Other plant hormones and growth
determine the formation of flowers, stems, leaves, the regulators are not easily grouped into these classes; they
shedding of leaves, and the development and ripening exist naturally or are synthesized by humans or other
of fruit (Salisbury and Ross, 1992). organisms, including chemicals that inhibit plant
Plants, unlike animals, lack glands that produce growth or interrupt the physiological processes within
and secrete hormones. Instead, each cell is capable of plants. A large number of related chemical compounds
producing hormones. Plant hormones shape the plant, are synthesized by humans. They are used to regulate
affecting seed growth, time of flowering, the sex of the growth of cultivated plants, weeds, and in vitro-
flowers, senescence of leaves, and fruits. They affect grown plants and plant cells; these man-made
which tissues grow upward and which grow downward, compounds are called Plant Growth Regulators or
leaf formation and stem growth, fruit development and PGRs for short. Early in the study of plant hormones,
ripening, plant longevity, and even plant death. "phytohormone" was the commonly used term, but its
Hormones are vital to plant growth, and, lacking them, use is less widely applied now.

Table 1. Plant Growth Hormones and Regulators.

Plant Growth Hormones Plant Growth Regulators

Auxins
Naphthalene acetic acid (NAA),
Auxins 2,4-Dichlorophenoxyacetic acid (2,4-D),
Indole-3-acetic acid (IAA) 2,methyl-4-chlorophenoxyacetic acid (MCPA),
Indoleacetonitrile (IAN) 2,4,5-Trichlorophenoxyacetic acid (2,4,5-T)
Indole ethanol (Iethanol) Phenylacetic acid (PAA)
Indole pyruvic acid (IPA) Indolepropionic acid (IPA)
Indolebutyric acid (IBA)
Phenoxyacetic acid (PAC)
Gibberellins
Gibberellic acids, 1 > 100
Cytokinins Cytokinins
Zeatin, ribosyl zeatin, 6-methylaminopurine. Kinetin, 2-Isopentenyl adenine (2, i-p), 6-Benzylaminopurine (6,BAP) or Benzyladenine
Abscisic Acid
Ethylene inhibitors such as aviglycine, ethylene releasers such as etacelasil, ethephon,
Ethylene
glyoxime.
Others
Others
Gametocides (fenridazon, maleic hydrazide), Morphactins (chlorfluren, chlorflurenol,
Fusicoccin, Brassins, Turgorins, Salicylic Acid,
dichlorflurenol, flurenol), Growth retardants (chlormequat, daminozide, flurprimidol,
Jasmonic Acid, Batasins, Plant Peptide
mefluidide, paclobutrazol, tetcydasis, uniconazole), Growth stimulators (brassinolide,
Hormones, Polyamines, Nitric Oxide,
brassinolide-ethyl, forchlorfenuron, hymexazol, prosuler, pyripropanol, triacontanol),
Strigolactones and Karrikins
Defoliants (calcium cyanamide, dimethipin, endothal, ethephon, merphos, metoxuron,
pentachlorophenol, thidiazuron, tribufos).

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CHAPTER TWO

2. Auxins and Cytokinins

2.1 Auxins

Auxins were the first class of growth regulators

discovered. Indole-3-acetic acid (IAA) is the main
auxin in higher plants and has profound effects on plant
growth and development. Both plants and some plant
pathogens can produce IAA to modulate plant growth
(Zhao, 2010). The in vitro bioassay in which auxin- Fig. 4. Chemical Structure of 2,4,5-trichlorophenoxyacetic acid
containing agar blocks stimulated the growth of oat (C₈H₅Cl₃O₃).
coleoptile segments led to the identification of indole-3-
acetic acid (IAA) as the main naturally occurring auxin
in plants. Applications of IAA or synthetic auxins to
plants cause profound changes in plant growth and
development (Bonner, 1952).

Fig. 5. Chemical Structure of Naphthoxyacetic Acid (C₁₂H₁₀O₃).

Fig. 1. Chemical Structure of Indole-3-Acetic Acid (C₁₀H₉NO₂).

Apart from IAA, there are other natural auxins

found in plants, they are indole ethanol (Iethanol),
indoleacetonitrile (IAN) and indole pyruvic acid (IPA).
Synthetic auxins which are also known as man-made
auxins include 2,4-dichlorophenoxyacetic acid (2,4-D), Fig. 6. Chemical Structure of Naphthalene Acetic Acid (C₁₂H₁₀O₂).
naphthalene acetic acid (NAA), indole butyric acid
(IBA) etc. 2.1.1 Discovery of Auxin

The coleoptiles of grasses (like that of oat, the

Avena - coleoptile) are popular test objects of plant
physiology. A growing coleoptile that is illuminated
unilaterally does grow towards the light source. This
behaviour is common among plants and is known as
phototropism. Charles Darwin (assisted by his son
Francis Darwin) attributed in his work "The Power of
Movement in Plants" (1880) a decisive function in the
recognition of a light stimulus to the coleoptiles tip, and
Fig. 2. Chemical Structure of Indole Butyric Acid (C₁₂H₁₃NO₂). he observed that the actual bending occurs in a zone
below the tip. He concluded that a transmission of
impulse had to take place in the tissue. Studies of plant
anatomical nature revealed that the growth towards
light is caused by an elongation of the cells at the side
that is shielded from the light. The phototropic reaction
does not happen if the coleoptiles tip is removed,
though it can be induced again by the replacement of
the tip. This indicates the existence of a substance that
is spread from tip to bottom (basipetal direction) and
that causes the elongation.
Fig. 3. Chemical Structure of 2,4-dichlorophenoxyacetic acid
(C₈H₆Cl₂O₃).

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The Danish botanist P. Boysen-Jensen routes have been proposed, but none of the proposed
interrupted the assumed substance flow by inserting a pathways in plants have been fully determined. In
mica sheet into the shielded side, thus separating the addition, several pathways are known to exist in the
coleoptiles tip from the tissue below (1913). The water- tryptophan-dependent pathway, including the
impermeable sheet interrupted the phototropic reaction. tryptamine (TAM), indole-3-pyruvic acid (IPA) and
Consequently occurs no transport of the effector around indole-3-acetaldoxime (IAOx) pathways (Soeno et al.,
the small tile. The phototropic reaction remained intact 2010). Plants also make IAM, but the biosynthesis
when the mica sheet was inserted into the illuminated routes of IAM in plants are not defined (Sugawara et
side or along the coleoptiles vertical axis. al., 2009). IAM can be converted to IAA by
In the late twenties was the material nature of the Arabidopsis AMIDASE1 (Lehmann et al., 2010). In
effector finally proved by the Dutch plant physiologist some plant pathogenic bacteria, IAA is synthesized
F. Went. He assumed that a substance that flows from from Trp by the Trp monooxygenase iaaM and the
tip to bottom should also flow through a small cube of hydrolase iaaH. The iaaM converts Trp to indole-3-
agar. In order to test his assumption did he place cut acetamide (IAM) that is subsequently hydrolyzed to
coleoptile tips with the cutting side on top of small IAA by iaaH (Comai and Kosuge, 1982).
cubes of agar. Sometime later did he remove the tips Trp can also be converted into indole-3-
and placed the agar cubes that he believed to contain acetaldoxime (IAOx) by the P450 enzymes CYP79B2
the effector onto the decapitated coleoptiles. He wrote and -B3 (Zhao et al., 2002). The routes from IAOx to
about the carrying out of the decisive experiment: IAA are not understood, although both IAM and indole-
"When I removed the tip after an hour and 3-acetonitrile have been suggested as intermediates
placed the agar cube on one side of the seedling, (Sugawara et al., 2009). IAOx is probably not the main
nothing happened at first. But in the course of the night, auxin biosynthesis intermediate, because
the stump started to curve away from the agar block. It (i) A complete elimination of IAOx production in
had acquired the capacity of the stem tip to grow! At Arabidopsis only leads to subtle growth defects
3:00 A.M. on April 17, 1926" (Salisbury and Ross, (Zhao et al., 2002);
1978). Went called the effector auxin (or growth- (ii) The enzymes CYP79B2 and -B3 seemed only to
regulating substance). Its chemical name is indole-3- exist in a small group of plants, and
acetic acid (IAA). The formula shows that it is a (iii) There is no detectable IAOx in rice and maize
tryptophane derivative. (Sugawara et al., 2009).
Auxin biosynthesis in plants is extremely
2.1.2 Biosynthesis of Auxin complex. Multiple pathways likely contribute to de
novo auxin production. IAA can also be released from
Auxin is an essential hormone, but its IAA conjugates by hydrolytic cleavage of IAA-amino
biosynthetic routes in plants have not been fully acids, IAA-sugar, and IAA-methyl ester (Li et al.,
defined. Auxin is an essential regulator for various 2008). Furthermore, although plants share
plant developmental processes. Indole-3-acetic acid evolutionarily conserved core mechanisms for auxin
(IAA), the main auxin in plants, can be synthesized biosynthesis, different plant species may also have
from tryptophan (Trp) –dependent and -independent unique strategies and modifications to optimize their
pathways (Zhao, 2010). Several auxin biosynthesis IAA biosynthesis.

Fig. 7. Biosynthesis of Auxin.

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2.1.3 Auxin Transport Bending toward a light source (phototropism),

downward root growth in response to gravity
The distribution of auxin within plant tissues is (geotropism), flower formation, fruit set and growth.
subject to clear and recognizable rules that indicate a NAA and IBA is often the active ingredient in products
transport simultaneously active and polar. This means that stimulate root formation on plant cuttings. Auxins
that IAA-specific carriers have to exist beside the may be used to enhance fruit production or harvesting,
receptor (or receptors). At least six reasons speak for but these effects are species specific. Auxin, along with
this: Transport occurs always directed: it is polarized. cytokinin, is used in for culturing plant tissues for mass
The transport velocity is higher than expected from propagation. Auxins increase the plasma current, the
simple diffusion. Transport can occur against a plasticity of the cell wall, and they cause a proton efflux
concentration gradient. Transport is energy-consuming out of the cell. Experiments show that auxins increase
and is drastically reduced in the absence of oxygen. The the rate of transcription, control the activity of certain
transport system is substrate-specific. It transports enzymes, and have an influence on the ion pumps
certain auxin molecules like IAA or naphthylacetic acid within the membrane.
faster than 2,4-dichlorphenoxy acetic acid, for example.
The transport system can be blocked by specific 2.2 Cytokinins
inhibitors.
Experiments with plasma membrane vesicles Cytokinins are compounds with a structure
showed that the accumulation of auxin is dependent on resembling adenine which promotes cell division and
pH and electron potential. The transport of auxins have other similar functions to kinetin. Kinetin was the
through the membrane is directed. Auxin specifically first cytokinin discovered and so named because of the
binds tonoplasts, and it influences the release of compounds ability to promote cytokinesis (cell
calcium ions from the vacuole in vitro. Different division). Though it is a natural compound, It is not
carriers for import and export have been detected: a made in plants and is therefore usually considered a
proton carrier (S) that causes symport and an auxin- "synthetic" cytokinin (meaning that the hormone is
anion-carrier (AC) that is an active antiport-carrier. synthesized somewhere other than in a plant). The most
Moreover, was it demonstrated that these auxin carriers common form of naturally occurring cytokinin in plants
are distributed asymmetrically within the membrane. today is called zeatin which was isolated from corn
(Zea mays).
2.1.4 Auxin Functions and Physiological Effects Cytokinins have been found in almost all higher
plants as well as mosses, fungi, bacteria, and also in
Auxins physiological effects in plants are highly tRNA of many prokaryotes and eukaryotes. Today there
related to its functions. These effects include Auxins are more than 200 natural and synthetic cytokinins
are compounds that positively influence cell combined. Cytokinin concentrations are highest in
enlargement, bud formation and root initiation. They meristematic regions and areas of continuous growth
also promote the production of other hormones and in potential such as roots, young leaves, developing fruits
conjunction with cytokinins, they control the growth of and seeds (Salisbury and Ross, 1992).
stems, roots, and fruits, and convert stems into flowers
(Osborne, 2005). They affect cell elongation by altering
cell wall plasticity. They stimulate cambium, a subtype
of meristem cells, to divide and in stems cause
secondary xylem to differentiate. Auxins act to inhibit
the growth of buds lower down the stems (apical
dominance), and also to promote lateral and
adventitious root development and growth. Leaf
abscission is initiated by the growing point of a plant
ceasing to produce auxins. Auxins in seeds regulate Fig. 8. Chemical Structure of Cytokinins (C₁₀H₉N₅₀).
specific protein synthesis, (Walz et al., 2002) as they
develop within the flower after pollination, causing the 2.2.1 Discovery of Cytokinins
flower to develop a fruit to contain the developing
seeds. Auxins are toxic to plants in large Unlike other hormones, cytokinins are found in
concentrations; they are most toxic to dicots and less so both plants and animals. Cytokinins or CKs are a group
to monocots. Because of this property, synthetic auxin of chemicals that influence cell division and shoot
herbicides, including 2,4-D and 2,4,5-T have been formation. They were called kinins in the past when the
developed and used for weed control. Auxins, first cytokinins were isolated from yeast cells. It has
especially 1-Naphthaleneacetic acid (NAA) and Indole- been tried for a long time to cultivate, plant tissue on
3-butyric acid (IBA), are also commonly applied to artificial nutrient medium. The first approaches go back
stimulate root growth when taking cuttings of plants. to the Austrian plant anatomist G. Haberlandt (1854-

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Plant Growth Substances Agboola et al

1945, professor at Graz, later at Berlin). At first, posed

the composition of a suitable nutrient medium large
problem. The addition of coconut milk causes a drastic
increase in the growth of plant embryos and tissue
cultures (Overbeek, 1941). Coconut milk is an
endosperm product that has under natural conditions,
too, a growth-stimulating effect on the developing
coconut embryo. The question which components cause
the growth stimulation arose immediately.
In contrast to auxin, growth is not by cell
elongation but by cell division. Aged or autoclaved
Fig. 10. Chemical Structure Of 6-Benzylaminopurine (C12H11N5).
DNA preparations have the same effect, while fresh
DNA preparations display no effect at all (Miller and
Skoog, 1955). In the end was the adenine derivative 6- 2.2.2 Biosynthesis and Metabolism of Cytokinins
furfurylaminopurine (kinetin) identified as the effective
Cytokinin is generally found in higher
substance.
concentrations in meristematic regions and growing
Kinetin is physiologically extraordinary active,
tissues. They are believed to be synthesized in the roots
although it could not be isolated from any plant cell.
and translocated via the xylem to shoot. Cytokinin
Instead was a wide range of similar compounds found.
The first that was isolated from a natural source (unripe biosynthesis happens through the biochemical
modification of adenine. The process by which they are
corn seed) was zeatin (Letham et al., 1964).
synthesized is as follows: A product of the mevalonate
pathway called isopentyl pyrophosphate is isomerized.
This isomer can then react with adenosine
monophosphate with the aid of an enzyme called
isopentenyl AMP synthase. The result is isopentenyl
adenosine-5'-phosphate (isopentenyl AMP). This
product can then be converted to isopentenyl adenosine
by removal of the phosphate by a phosphatase and
further converted to isopentenyl adenine by removal of
Fig. 9. Chemical Structure of Kinetin (C₁₀H₉N₅₀).
the ribose group. Isopentenyl adenine can be converted
to the three major forms of naturally occurring
cytokinins. Other pathways or slight alterations of this
one probably lead to the other forms. Degradation of
cytokinins occurs largely due to the enzyme cytokinin
oxidase. This enzyme removes the side chain and
releases adenine. Derivatives can also be made, but the
pathways are more complex and poorly understood
(Salisbury and Ross, 1992).

Fig. 10. Chemical Structure of Zeatin (C₁₀H₁₃N₅₀).

Fig. 11. Synthesis of Cytokinins (Source: Glover et al., 2008).

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Plant Growth Substances Agboola et al

2.2.3 Functions and Physiological Effects of increase cytokinins, too, the general rate of RNA and
Cytokinins protein synthesis. They reduce senescence and
stimulate the dark-germination of light-dependent
They also help delay senescence or the aging of seeds. In addition were several more selective effects
tissues, are responsible for mediating auxin transport observed. Cytokinins induce isocitrate lyase and
throughout the plant and affect the internodal length protease activities in cut pumpkin cotyledons; they
and leaf growth. They have a highly synergistic effect induce thiamine synthesis in growing callus cultures of
in concert with auxins, and the ratios of these two tobacco, thereby removing the thiamine need of the
groups of plant hormones affect most major growth calli. Cytokinins stimulate auxin synthesis in tissue
periods during a plant's lifetime. Cytokinins counter the cultures of tobacco; they also increase the carboxy
apical dominance induced by auxins; they in dismutase and the NADP-glyceraldehydes phosphate
conjunction with ethylene promote abscission of leaves, dehydrogenase activities in etiolated rice seedlings.
flower parts, and fruits (Sipes and Einset, 1983). Tissue Cytokinin stimulates the development of buds as well
cultures like that of tobacco or maple (Acer as the germination of seeds, and the accumulation of
pseudoplatanus) develop only after the addition of nitrate reductases in several embryos.
cytokinin. Besides, the rate of DNA replication,

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CHAPTER THREE

3. Gibberellin, Ethylene and Abscisic Acid hormone in the disease bakanae are present. Inactive
conjugates might be stored or translocated via the
3.1 Gibberellins phloem and xylem before their release (activation) at
the proper time and in the proper tissue (Sponsel,
Gibberellins, or GAs, include a large range of 1995).
chemicals that are produced naturally within plants and
by fungi. They were first discovered when Japanese 3.1.2 Functions and Physiological Effects of
researchers, including Eiichi Kurosawa, noticed a Gibberellins
chemical produced by a fungus called Gibberella
fujikuroi that produced abnormal growth in rice plants Gibberellins are important in seed germination,
(Grennan, 2006). affecting enzyme production that mobilizes food
Unlike the classification of auxins which are production used for growth of new cells. This is done
classified on the basis of function, gibberellins are by modulating chromosomal transcription. In grain
classified on the basis of structure as well as function. (rice, wheat, corn, etc.) seeds, a layer of cells called the
All gibberellins are derived from the ent-gibberellane aleurone layer wraps around the endosperm tissue.
skeleton. The structure of this skeleton derivative along Absorption of water by the seed causes production of
with the structure of a few of the active gibberellins is GA. The GA is transported to the aleurone layer, which
shown above. The gibberellins are named GA1....GAn responds by producing enzymes that break down stored
in order of discovery. Gibberellic acid, which was the food reserves within the endosperm, which are utilized
first gibberellin to be structurally characterised, is GA3. by the growing seedling. Gas produces bolting of
There are currently 136 GAs identified from plants, rosette-forming plants, increasing internodal length.
fungi and bacteria. GAs are widespread and so far They promote flowering, cellular division, and in seed
ubiquitous in flowering (angiosperms) and non- growth after germination. Gibberellins also reverse the
flowering (gymnosperms) plants as well as ferns. inhibition of shoot growth and dormancy induced by
Abscisic Acid (Tsai et al., 1997). Gibberellins also
stimulate stem elongation by stimulating cell division
and elongation. They stimulate bolting/flowering in
response to long days, break seed dormancy in some
plants which require stratification or light to induce
germination. Stimulates enzyme production (a-amylase)
in germinating cereal grains for mobilization of seed
reserves. Gibberellins induce maleness in dioecious
flowers (sex expression) and they cause parthenocarpic
Fig. 12. Chemical Structure of Gibberellic Acid (C₁₉H₂₂O₆). (seedless) fruit development. They also delay
senescence in leaves and citrus fruits.
3.1.1 Gibberellin Biosynthesis and Metabolism
3.2 Ethylene
Gibberellins are diterpenes synthesized from
acetyl CoA via the mevalonic acid pathway. They all Ethylene is unique in that it is found only in the
have either 19 or 20 carbon units grouped into either gaseous form and it is often found in high
four or five ring systems. The fifth ring is a lactone ring concentrations within cells at the end of a plant's life.
as shown in the structures above attached to ring A. Ethylene is a gas that forms through the Yang Cycle
Gibberellins are believed to be synthesized in young from the breakdown of methionine, which is in all cells.
tissues of the shoot and also the developing seed. It is Ethylene has a very limited solubility in water and does
uncertain whether young root tissues also produce not accumulate within the cell, but diffuses out of the
gibberellins. There is also some evidence that leaves cell and escapes out of the plant. Its effectiveness as a
may be the source of some biosynthesis (Sponsel, plant hormone is dependent on its rate of production
1995). versus its rate of escaping into the atmosphere.
Certain commercial chemicals which are used to Ethylene is produced at a faster rate in rapidly growing
stunt growth do so in part because they block the and dividing cells, especially in darkness. New growth
synthesis of gibberellins. Some of these chemicals are and newly germinated seedlings produce more ethylene
Phosphon D, Amo-1618, Cycocel (CCC), ancymidol, than can escape the plant, which leads to elevated
and paclobutrazol. During active growth, the plant will amounts of ethylene (Wang et al., 2007).
metabolize most gibberellins by hydroxylation to
inactive conjugates quickly with the exception of GA3.
GA3 is degraded much slower, which helps to explain
why the symptoms initially associated with the
Fig. 13. Chemical Structure of Ethylene (C₂H₄).

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3.2.1 Biosynthesis of Ethylene

Fig. 14. Biosynthesis of Cytokinins (Source: McKeon et al., 1995).
In higher plants, ethylene is produced from L- 3.2.2 Ethylene Distribution and Transport
methionine. Methionine is activated by ATP to form S-
adenosylmethionine through the catalytic activity of S- Probably each plant tissue produces or takes up
adenosylmethionine synthetase. Starting from S- ethylene at some point during development and
adenosylmethionine two specific steps result in the responds to it in an appropriate fashion. Uptake is
formation of ethylene. The first step produces the non- usually not a problem because ethylene can diffuse
protein amino acid, 1-aminocyclopropane-1-carboxylic freely through membranes. The distribution of the gas
acid (ACC). It is catalyzed by ACC synthase with within the plant occurs through intercellular spaces and
pyridoxal phosphate acting as a co-factor. Formation of when dissolved in the symplast from cell to cell. Long-
ACC is the rate-limiting step in ethylene biosynthesis. distance distribution is achieved by releasing ACC into
ACC synthase is encoded by a medium-size multigene the vascular tissue where it is moved to the site of
family. Various signals, which influence ethylene action. There, is converted to ethylene. Tomatoes, for
synthesis, result in increased expression of single instance, produce ACC in water-logged roots. ACC is
members of the ACC synthase gene family. Production moved via the vascular bundles to the leaves where it is
of ethylene from ACC is catalyzed by ACC oxidase. converted to ethylene, which then induces epinastic
This reaction is oxygen-dependent. At anaerobic lowering of the petioles (Sauter, 2012).
conditions, ethylene formation is completely
suppressed. Fe2+ is a co-factor and ascorbate 3.2.3 Functions and Physiological Effects of
cosubstrate; CO2 was shown to activate ACC oxidase. Ethylene
ACC oxidases are encoded by small gene families in
plants. Leaf expansion: Ethylene affects cell growth
The concentration of ethylene in a plant tissue is and cell shape; when a growing shoot hits an obstacle
dependent on the rate of biosynthesis and on the while underground, ethylene production greatly
diffusion of the gas. Ethylene is neither actively increases, preventing cell elongation and causing the
transported nor degraded. Induction of ethylene stem to swell. The resulting thicker stem can exert more
synthesis by signals such as auxin or wounding usually pressure against the object impeding its path to the
occurs through activation of ACC synthase through surface. If the shoot does not reach the surface and the
increased gene expression. ACC oxidase activity, on ethylene stimulus becomes prolonged, it affects the
the other hand, is constitutively present in most stem's natural geotropic response, which is to grow
vegetative plant tissues. In some cases, further upright, allowing it to grow around an object. Studies
induction of ACC oxidase by ethylene is observed seem to indicate that ethylene affects the stem diameter
(Sauter, 2012). In maturing fruits, ethylene synthesis is and height: When stems of trees are subjected to wind,
autocatalytically enhanced, i.e., ethylene induces its causing lateral stress, greater ethylene production
own biosynthesis. The self-enhancing synthesis and occurs, resulting in thicker, more sturdy tree trunks and
diffusion of the gaseous hormone throughout the fruit branches. Ethylene affects fruit-ripening: Normally,
accelerate ripening and contribute to a synchronized when the seeds are mature, ethylene production
increases and builds-up within the fruit, resulting in a
climacteric event just before seed dispersal (Wang et
al., 2007).

3.3 Abscisic Acid

Abscisic acid also called ABA, was discovered and

researched under two different names before its
chemical properties were fully known, it was called
dormin and abscisin II. Once it was determined that the
two latter compounds are the same, it was named
abscisic acid. The name "abscisic acid" was given
because it was found in high concentrations in newly
abscissed or freshly fallen leaves.
This class of PGR is composed of one chemical
compound normally produced in the leaves of plants,
originating from chloroplasts, especially when plants
are under stress. In general, it acts as an inhibitory
chemical compound that affects bud growth, and seed
ripening process. and bud dormancy. It mediates changes within the
apical meristem, causing bud dormancy and the

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alteration of the last set of leaves into protective bud stomata (Yan et al., 2007). ABA exists in all parts of
covers. Since it was founded in freshly abscissed the plant and its concentration within any tissue seems
leaves, it was thought to play a role in the processes of to mediate its effects and function as a hormone; its
natural leaf drop, but further research has disproven degradation, or more properly catabolism, within the
this. Plant species from temperate parts of the world, it plant affects metabolic reactions and cellular growth
plays a role in leaf and seed dormancy by inhibiting and production of other hormones (Kermode, 2005).
growth, but, as it is dissipated from seeds or buds, Plants start life as a seed with high ABA levels. Just
growth begins. In other plants, as ABA levels decrease, before the seed germinates, ABA levels decrease;
growth then commences as gibberellin levels increase. during germination and early growth of the seedling,
Without ABA, buds and seeds would start to grow ABA levels decrease even more. As plants begin to
during warm periods in winter and be killed when it produce shoots with fully functional leaves, ABA levels
froze again. Since ABA dissipates slowly from the begin to increase, slowing down cellular growth in
tissues and its effects take time to be offset by other more "mature" areas of the plant. Stress from water or
plant hormones, there is a delay in physiological predation affects ABA production and catabolism rates,
pathways that provide some protection from premature mediating another cascade of effects that trigger
growth. It accumulates within seeds during fruit specific responses from targeted cells. Scientists are
maturation, preventing seed germination within the still piecing together the complex interactions and
fruit, or seed germination before winter. Abscisic acid's effects of this and other phytohormones.
effects are degraded within plant tissues during cold
temperatures or by its removal by water washing in out
of the tissues, releasing the seeds and buds from
dormancy (Feurtado et al., 2004).
In plants under water stress, ABA plays a role in
closing the stomata. Soon after plants are water-stressed
and the roots are deficient in water, a signal moves up
to the leaves, causing the formation of ABA precursors
there, which then move to the roots. The roots then
release ABA, which is translocated to the foliage
through the vascular system (Ren et al., 2007). It then
modulates the potassium and sodium uptake within the
Fig. 15. Chemical Structure of Abscisic acid (C₁₅H₂₀O₄).
guard cells, which then lose turgidity, closing the

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CHAPTER FOUR

4. Other Plant Growth Substances inhibit growth of certain plant parts and to strongly
promote leaf senescence (Salisbury and Ross, 1992).
4.1 Lunularic Acid The major function of JA and its various metabolites is
regulating plant responses to abiotic and biotic stresses
In liverworts, lunularic acid is present in gemmae. as well as plant growth and development (Delker et al.,
Lunularic acid prevents germination of these gemmae 2006). Regulated plant growth and development
until they fall from the mother plant’s thallus and the processes include growth inhibition, senescence, tendril
acid is leached out (Milborrow, 1984). Furthermore, the coiling, flower development and leaf abscission.
growth of the entire thallus seems to be partly
controlled lunularic acid in response to day length.
During short days the concentration of the inhibitor is
low and thalli grow rapidly, during long days the
reverse is true. Lunularic acid is present in many
species of lower plants, but not in algae (Gorham,
1977).

Fig. 17. Chemical Structure of Jasmonic Acid (C₁₂H₁₈O₃).

JA is also responsible for tuber formation in

potatoes, yams, and onions. It has an important role in
response to wounding of plants and systemic acquired
resistance. When plants are attacked by insects, they
respond by releasing JA, which activates the expression
Fig. 16. Structure of lunularic acid. of protease inhibitors, among many other anti-herbivore
defense compounds. These protease inhibitors prevent
4.2 Batasins the proteolytic activity of the insects' digestive
proteases, thereby stopping them from acquiring the
These are compounds found in yam plants needed nitrogen in the protein for their own growth
(Dioscorea batatas) that seem to cause dormancy in (Zavala et al., 2004).
bulbils (vegetative reproductive structures) that arise For example, research on Madagascar periwinkle
from the swelling of the aerial lateral buds. Batasins are (Catharanthus roseus), which makes some anti-cancer
concentrated in the skin of the bulbils and are absent alkaloids, identified a transcription factor that responds
from the core. A long duration of exposure to cold to JA by activating the expression of several genes
temperature (stratification or pre-chilling) that breaks encoding alkaloid biosynthetic genes (Van der Fits and
dormancy causes batasins to disappear, whereas their Memelink, 2000). This chemical may have a role in
amounts increase during the initial development of pest control, according to an October 2008 BBC News
dormant bulbils (Hasegawa and Hashimoto, 1975). report. JA affects several plant processes, such as pollen
However, it is not known whether batasins are development, root growth, fruit ripening and
accumulated in or transported into the bud cells whose senescence and it is also produced in response to the
failure to grow actually causes dormancy. presence of insect pests and disease-causing organisms.
JA triggers the production of enzymes that confer an
4.3 Jasmonic Acid increased resistance against herbivorous (plant eating)
insects. For example, caterpillar infested tomato plants
Jasmonic acid and its methyl ester (methyl release volatile jasmonic acid into the air that attracts
jasmonate) occur in several plant species and in the oil natural caterpillar enemies such as parasitic wasps. The
of jasmine (Parthier, 1990). Jasmonates have been wasps lay their eggs on the caterpillars’ bodies, when
found in over 150 families and 260 species including the eggs hatch, the wasp larvae consume the host
fungi, mosses and ferns, so they are perhaps ubiquitous insects. In one study, plants treated with jasmonic acid
in plants. JA is a lipid derived plant hormone, which is increased parasitism on caterpillar pests two folds over
structurally similar to prostaglandins in animals control fields that were not treated with it.
(Solomon et al., 2011). Biologists have made progress in identifying key
Jasmonic acid (JA) is derived from the fatty acid aspects of the jasmonic acid signalling pathway.
linolenic acid. It is a member of the jasmonate class of Interestingly the jasmonic acid pathway includes
plant hormones. It is biosynthesized from linolenic acid ubiquitin as in the auxin signalling pathway. The
by the octadecanoid pathway. JA has been shown to

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Plant Growth Substances Agboola et al

ubiquitin tags in the JA signalling pathway cause The brassinosteroids (BRs) are involved in several
degradation of transcription receptors, thereby altering aspects of growth and development. Arabidopsis
gene expression (Solomon et al., 2011). mutants that cannot synthesize BRs are dwarf plants
with small curly leaves and reduced fertility. This effect
4.4 Brassinosteroids can be reversed by researchers by applying BR
exogenously to the plant. Studies of these mutants
Brassinosteroids (also called brassins) are plant suggest that BRs are involved in multiple
steroid hormones, although steroid hormones in plants developmental processes such as cell division, vascular
were not well characterised until the mid 1990s. They development and seed germination.
have distinctive growth promoting activity in some Many steps of the BR signalling pathway have
plants, especially in stems. These substances were first been characterised. BRs bind to a receptor kinase in the
isolated from bee-collected pollen grains of rape plant plasma membrane (unlike steroid hormones in
(Brassica napus), mustard (Salisbury and Ross, 1992). animals which enter the cell and bind to receptors in the
The structure of one brassin known as brassinolide is cytoplasm or nucleus). When BR binds to its receptor,
shown below. the receptor phosphorylates molecules in the cytosol,
which in turn triggers a signalling cascade that
ultimately alters gene expression in the cell. Part of the
signal transduction process involves attaching ubiquitin
to nuclear proteins targeted for degradation in a
proteasome. As a result, the expression of BR target
genes is adjusted (Solomon et al., 2011).

Fig. 18. Chemical Structure of Brassinolide (C₂₈H₄₈O₆).

Fig. 19. Biosynthesis of Brassinosteroids, Abscisic Acid and Gibberellins (Source: Gianpiero, 2009).

4.5 Salicylic Acid into the air. When nearby healthy plants receive the
airborne chemical signal, they begin synthesizing anti-
Salicylic acid, a phenolic extract from willow bark, viral proteins that enhance their resistance to the virus
was long used as an analgesic. It is now prepared (Solomon et al., 2011).
commercially and is the active ingredient of aspirin.
Salicylic acid is known to activate defense genes
for resistance against insect pests and pathogen
invaders, known as the hypersensitive response.
When a plant is under attack, the concentration of
SA increases and it spreads systemically throughout the
plant. Plants also use methyl salicylate, which is
volatile to signal nearby plants. Tobacco plants infected Fig. 20. Chemical structure of salicylic acid (C₇H₆O₃).
with Tobacco Mosaic Virus release methyl salicylate

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Plant Growth Substances Agboola et al

4.5.1 Pathways of SA biosynthesis in plants phytotoxin that stimulates both rapid proton extrusion
and transient growth in the stem and coleoptile sections
Isotope feeding experiments suggest that plants (Taiz and Zeiger, 2002). Research with both cucumber
synthesize SA from cinnamate produced by cotyledons and maize coleoptiles verified that
phenylalanine ammonia lyase (PAL). Genetic studies fusicoccin is a potent growth promoter and can acidify
have indicated that the bulk of SA is produced from cell walls enough to promote their growth. It has the
isochorismate. Given the importance of both PAL and remarkable abilities to activate a plasma membrane
ICS in SA accumulation demonstrated from ATPase that transports hydrogen ion from the cytosol
experiments using genetic mutants, gene silencing and into the wall to lower the wall pH, to enhance wall
chemical inhibition, it is possible that the PAL and loosening and to promote cell growth.
isochorismate synthase (ICS) pathways are integrated Even though fusicoccin can increase growth of
through a metabolic or regulatory grid in SA coleoptiles and cotyledons as it promotes hydrogen ion
biosynthesis. The recently identified PBS3 and EPS1 efflux, auxins cannot promote hydrogen ion efflux
are important for pathogen-induced SA production and enough to promote maize-coleoptile growth, nor can
may encode enzymes catalyzing related, and possibly cytokinins promote hydrogen ion efflux enough to
sequential, reactions in the synthesis of an important enhance cotyledon growth. This implies that, in effect,
precursor or regulatory molecule for SA biosynthesis auxins and other hormones must cause wall loosening
(Zhixiang et al., 2009). and cell expansion in some (maybe most) species by
some mechanism (Salisbury and Ross, 1992).

4.7 Tugorins

These sets of compounds were discovered by

Schildknecht (1983, 1984) using leaves of the plant
Mimosa and Acacia karroo. He isolated and identified
compounds that activated pulvini in these leaves which
are not sensitive to touch but exhibits nyctinasty.
A Mimosa leaf is placed in a solution with a
suspected active substance which is then transported in
the transpiration stream to the pulvini, where their
membranes respond and cause the leaf pinnules to fold
up if the substance is active. Two Periodic Leaf
Movement Factors (PLMFs) from Acacia proved to be
the β-glucosides of Gallic acid, with the glucoside
bonding at its para-hydroxyl group. Several other
compounds with similar structures were also identified
from other plants. The most active were the β-D-
glucoside-6-sulphate (called PLMF1) and the β-D-
glucoside-3,6-sulphate (PLMF 2) of Gallic acid.
Extracts from Mimosa contained PLMF 1 and PLMF 7,
another Npound called PLMF 3 which is derived from
protocatechuic acid were isolated from Oxalis stricta
(this species react to touch or shaking as much as
Mimosa does). Glucose-6-sulphate was found to be part
of the molecule. Schildknecht then suggested that these
compounds form a new class of plant hormones, which
he called Tugorins because they act on turgor of
pulvinus cells, and they are active at low concentrations
like other phytohormones (Salisbury and Ross, 1992).

Fig. 21. Biosynthetic Pathway of Salicylic Acid. 4.8 Strigolactones

4.6 Fusicoccin Strigolactones are plant hormones that stimulate
the branching and growth of symbiotic arbuscular
Fusicoccin is a diterpene glucoside that was mycorrhizal fungi, increasing the probability of contact
identified by plant pathologists in the 1960s as the and establishment of a symbiotic association between
principal toxin responsible for disease symptoms the plant and fungus (López-Ráez et al., 2009).
caused by the fungus Fusicoccum amygdali on peach, Strigolactones inhibit plant shoot branching (Gomez-
almond, and prune trees (Marre, 1979). It is a fungal

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Plant Growth Substances Agboola et al

Roldan et al., 2008). They also trigger germination of Arabidopsis thaliana respond sensitively and
parasitic plant seeds (for example Striga from which specifically to karrikins in smoke. These exciting
they gained their name) (Akiyama and Hayashi, 2006). discoveries might be explained if karrikins are
Strigolactones are carotenoid-derived and contain a produced in the environment by processes other than
labile ether bond that is easily hydrolyzed in the fire, such as by chemical or microbial degradation of
rhizosphere meaning that there is a large concentration vegetation in response to disturbance of the soil or
gradient between areas near the root and those further removal of the plant canopy. Another hypothesis is that
away. Recently, strigolactone biosynthesis were found plants contain endogenous karrikins that function
to be DWARF27 dependent (Lin et al., 2009). naturally in the control of seed germination and that
species from fire-prone habitats have evolved to
respond also to exogenous karrikins. A variant on this
hypothesis is that karrikins mimic endogenous plant
hormones such as terpenoids that control seed
germination. The evidence for these hypotheses is
discussed, but whatever the explanation karrikins are
now firmly established as an important family of
naturally occurring plant growth regulators.

Fig. 22: Chemical Structure of Strigolactones (C₁₇H₁₄O₅).

4.9 Karrikins

Karrikins are a chemically defined family of plant

growth regulators discovered in smoke from burning
plant material. Karrikins are potent in breaking
dormancy of seeds of many species adapted to
environments that regularly experience fire and smoke.
The recent discovery that karrikins trigger seed
germination and control seedling growth in taxa that
would rarely experience fire indicates that their Fig. 23. Chemical Structure of Karrikins.
significance could extend far beyond fire ecology. This
is exemplified by new studies showing that the seeds of

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Plant Growth Substances Agboola et al

CHAPTER FIVE

4. Conclusion flipper.diff.org/app/items/info/2127, retrieved

June 4th, 2012.
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