Chapter 4: PLANT GROWTH SUBSTANCES

Auxin: The Growth Hormone Auxin is the first growth hormone to be discovered in plants and is one of an expanding lists of
chemical signaling agents that regulate plant development. It is from the Greek auxein, meaning “to increase” or “to grow.”
The most common naturally occurring form of auxin is indole-3-acetic acid (IAA). Several other auxins in higher plants were
discovered later, but IAA is by far the most abundant and physiologically relevant.
An early definition of auxins included all natural and synthetic chemical substances that stimulate elongation in coleoptiles and
stem sections. However, auxins affect many developmental processes besides cell elongation. Bending in response to light
(phototropism) or gravity (gravitropism) results from the lateral redistribution of auxin. In response to a directional light stimulus, the
auxin produced at the tip, instead of being transported basipetally, is transported laterally toward the shaded side.

Auxin regulates apical dominance. In most higher plants, the growing apical bud inhibits the growth of lateral (axillary buds) – a
phenomenon called apical dominance. Removal of the shoot apex (decapitation) usually results in the growth of one or more of the
lateral buds. Not long after the discovery of auxin, it was found that IAA could substitute for the apical bud in maintaining the
inhibition of lateral buds. Kenneth V. Thimann and Folke Skoog originally proposed that auxin from the shoot apex inhibits the
growth of the axillary bud directly – the so-called direct-inhibition model.

Auxin promotes the formation of lateral and adventitious roots. Although elongation of the primary root is inhibited by auxin
concentrations greater than 10-8 M, initiation of lateral (branch) roots and adventitious roots is stimulated by high auxin levels.

Auxin delays the onset of leaf abscission. The shedding of leaves, flowers, and fruits from the living plant is known as abscission.
These parts abscise in a region called the abscission zone, which is located near the base of the petiole of leaves. In most plants,
leaf abscission is preceded by the differentiation of a distinct layer of cells, the abscission layer, within the abscission zone.

Auxin levels are high in young leaves, progressively decrease in maturing leaves, and are relatively low in senescing leaves when
the abscission process begins. The role of auxin in leaf abscission can be readily demonstrated by excision of the blade from a
mature leaf, leaving the petiole intact on the stem.

These results suggest the following:

• Auxin transported from the blade normally prevents abscission
• Abscission is triggered during leaf senescence when auxin is no longer being produced.

Auxin transport regulates floral bud development. Polar auxin transport in the inflorescence meristem is required for normal floral
development. Apparently, the developing floral meristem depends on auxin being transported to it from subapical tissues.

Auxin promotes fruit development. Auxin is produced in pollen and in the endosperm and the embryo of developing seeds, and the
initial stimulus for fruit growth may result from pollination.

The distribution of IAA in the cell appears to be regulated largely by pH. The distribution of IAA and its metabolites has been
studied in tobacco cells. About one-third of the iAA is found in the chloroplast, and the remainder is located in the cytosol.

Auxin Transport

The main axes of shoots and roots, along with their branches, exhibit apex-base structural polarity, and this structural polarity has
its origin in the polarity of auxin transport. It was discovered that IAA moves mainly from the apical to the basal end in excised oat
coleoptile sections. This type of unidirectional transport is termed polar transport. Auxin is the only plant growth hormone known to
be transported polarly. Polar transport requires energy and is independent of gravity.

Gibberellins: Regulator of Plant Height Gibberellins (GAs)

are a family of compounds defined by their structure. Gibberellins are most often associated with the promotion of stem growth,
and the application of gibberellin to intact plants can induce large increases in plant height. Gibberellins induce dramatic internode
elongation in certain types of plants, such as dwarf and rosette species and grasses.

Effects of Gibberellin on Growth and Development Gibberellins stimulate stem growth in dwarf and rosette plants. Applied
gibberellin promotes internodal elongation in a wide range of species. However, the most dramatic stimulations are seen in dwarf
and rosette species. Accompanying this effect are a decrease in stem thickness, a decrease in leaf size, and a pale green color of
the leaves. Gibberellins promote fruit set. Applications of gibberellins can cause fruit set (the initiation of fruit growth following
pollination) and growth of some fruits, in cases where auxin may have no effect.
Gibberellins promote seed germination. Seed germination may require gibberellins for one of several possible steps: the activation
of vegetative growth of the embryo, the weakening of a growth-constraining endosperm layer surrounding the embryo, and the
mobilization of stored food reserves of the endosperm.

Cytokinins: Regulators of Cell Division

Mature plant cells generally do not divide in the intact plant, but they can be stimulated to divide by wounding, by infection with
certain bacteria, and by plant hormones, including cytokinins. Cytokinins are N6 -substituted aminopurines that will initiate cell
proliferation in many plant cells when they are cultured on a medium that also contains an auxin. The principal cytokinin of higher
plants – zeatin, or trans-6-(4-hydroxy-3- methylbut-2-enylamino) purine – is also present in plants as a riboside or ribotide and as
glycosides.

Cytokinin oxidases degrade cytokinin irreversibly and may play a role in regulation of the levels of this hormone. This hormone is
most abundant in the young, rapidly dividing cells of the shoot and root apical meristems.

The Biological Roles of Cytokinins

Cytokinins participate in the regulation of many plant processes, including cell division, morphogenesis of shoots and roots,
chloroplast maturation, cell enlargement, and senescence. In addition to cell division, the ratio of auxin to cytokinin determines the
differentiation of cultured plant tissues into either roots or buds: High ratios promote roots; low ratios, buds.

Ethylene: The Gaseous Hormone

Ethylene is formed in most organs of higher plants. Senescing tissues and ripening fruits produce more ethylene than do young or
mature tissues. The biosynthesis of this hormone is triggered by various developmental processes, by auxins, and by
environmental stresses. Ethylene regulates fruit ripening and other processes associated with leaf and flower senescence, leaf and
fruit abscission, root hair development, seedling growth, and hook opening.

Ethylene promotes the ripening of some fruits. Ethylene has long been recognized as the hormone that accelerates the ripening of
edible fruits. Exposure of such fruits to ethylene hastens the processes associated with ripening, and a dramatic increase in
ethylene production accompanies the initiation of ripening.

Ethylene breaks seed and bud dormancy in some species. Seeds that fail to germinate under normal conditions are said to be
dormant. Ethylene has ability to break dormancy and initiate germination in certain seeds. Ethylene promotes the elongation
growth of submerged aquatic species. Although usually thought of as an inhibitor of stem elongation, ethylene is able to promote
stem and petiole elongation in various submerged or partially submerged aquatic plants. Growth is stimulated in the submerged
plants because ethylene builds up in the tissues. In the absence of O2, ethylene synthesis is diminished, but the loss of ethylene
by diffusion is retarded under water.

. Abscisic Acid: A Seed Maturation and Antistress Signal

Abscisic acid (ABA) plays major roles in seed and bud dormancy, as well as responses to water stress. ABA is a 15-carbon
terpenoid compound derived from the terminal portion of carotenoids. ABA is synthesized in almost all cells that contain plastids
and is transported via both the xylem and the phloem. The level of ABA fluctuates dramatically in response to developmental and
environmental changes.

ABA is required for the development of desiccation tolerance in the developing embryo, the synthesis of storage proteins, and the
acquisition of dormancy. Seed dormancy and germination are controlled by the ratio of ABA to GA, and ABA-deficient embryos
may exhibit precocious germination and vivipary. ABA is also antagonized by ethylene and brassinosteroid promotion of
germination.

During water stress, the ABA level of the leaf can increase 50-fold. In addition to closing stomata, ABA increases the hydraulic
conductivity of the root and increases the root:shoot ratio at low water potentials. ABA and an alkalinization of the xylem sap are
thought to be two chemical signals that the root sends to the shoot as the soil dries.

PHYTOCHROMES AND PHOTOPERIODISM

The term photomorphogenesis refers to the dramatic effects of light on plant development cellular metabolism. Red light exerts the
strongest influence, and the effects of red light are often reversible by far-red light. Among the different pigments that can promote
photomorphogenic1 responses in plants, the most important are those that absorb red and blue light. Phytochrome is a blue
protein pigment with a molecular mass of about 125 kDa (kilodaltons) that absorbs red and far-red light most strongly, but that also
absorbs blue light.
The Photochemical and Biochemical Properties of Phytochrome Phytochrome exists in two forms: a red light-absorbing form (Pr)
and a far-red light-absorbing form (Pfr). Phytochrome can Interconvert between Pr and Pfr Forms In dark-grown or etiolated plants,
phytochrome is present in a red light-absorbing form, referred to as Pr because it is synthesized in this form. Pr, which to the
human eye is blue, is converted by red light to a far-red light-absorbing form called Pfr, which is blue-green. Pfr, in turn, can be
converted back to Pr by far-red light. Known as photoreversibility, this conversion/reconversion property is the most distinctive
property of phytochrome, and it may be expressed in abbreviated form as follows: ‘

When Pr molecules are exposed to red light, most of them absorb it and are converted to Pfr, but some of the Pfr also absorbs the
red light and is converted back to Pr because both Pr and Pfr absorb red light. In addition to absorbing red light, both forms of
phytochrome absorb light in the blue region of spectrum. Therefore, phytochrome effects can be elicited also by blue light, which
can convert Pr to Pfr and vice versa.

Localization of Phytochrome in Tissues and Cells

In etiolated seedlings the highest phytochrome levels are usually found in meristematic regions or in regions that were recently
meristematic. However, differences in expression patterns between monocots and dicots and between Type I and Type II
phytochromes are apparent when sensitive methods are used.

Phytochrome responses can be distinguished by the amount of light required to induce them. The amount of light is referred to as
the fluence, which is defined as the number of photons impinging on a unit surface area. In addition to the fluence, some
phytochrome responses are sensitive to the irradiance, or fluence rate, of light.

Ecological Functions: Shade Avoidance Phytochrome plays important ecological roles for plants growing in the environment.
Phytochrome Enables Plants to Adapt to Changing Light Conditions The presence of a red/far-red reversible pigment in all green
plants. From algae to dicots, suggests that these wavelengths of light provide information that helps plants adjust to their
environment.