Characteristics of Plant Growth Regulators
The plant growth regulators (PGRs) are small, simple molecules of diverse chemical composition. They could be indole compounds (indole-3-acetic acid, IAA); adenine derivatives (N6-furfurylamino purine, kinetin), derivatives of carotenoids (abscisic acid, ABA); terpenes (gibberellic acid, GA3) or gases (ethylene, C2H4). Plant growth regulators are variously described as plant growth substances, plant hormones or phytohormones in literature.
The plant growth regulators can be broadly divided into two groups based on their functions in a living plant body. One group of PGRs are involved in growth promoting activities, such as cell division, cell enlargement, pattern formation, tropic growth, flowering, fruiting and seed formation. These are also called plant growth promoters, e.g., auxins, gibberellins and cytokinins. The PGRs of the other group play an important role in plant responses to wounds and stresses of biotic and abiotic origin. They are also involved in various growth inhibiting activities such as dormancy and abscission. The PGR abscisic acid belongs to this group. The gaseous PGR, ethylene, could fit either of the groups, but it is largely an inhibitor of growth activities.
The Discovery of Plant Growth Regulators
Interestingly, the discovery of each of the five major groups of plant growth regulators have been accidental. All this started with the observation of Charles Darwin and his son Francis Darwin when they observed that the coleoptiles of canary grass responded to unilateral illumination by growing towards the light source (phototropism). After a series of experiments, it was concluded that the tip of coleoptile was the site of transmittable influence that caused the bending of the entire coleoptile (Figure 10). Auxin was isolated by F.W. Went from tips of coleoptiles of oat seedlings.
The ‘bakane’ (foolish seedling) a disease of rice seedlings, was caused by a fungal pathogen Gibberalla fujikuroi. E. Kurosawa reported the appearance of symptoms of the disease in uninfected rice seedlings when they were treated with sterile filtrates of the fungus. The active substances were later identified as gibberellic acid.
F. Skoog and his co-workers observed that from the intermodal segments of tobacco stems the callus (a mass of undifferentiated cells) proliferated only if, in addition to auxins the nutrients medium was supplemented with one of the following: extracts of vascular tissues, yeast extract, coconut milk or DNA. Skoog and Miller, later identified and crystallised the cytokinesis promoting active substance that they termed kinetin.
During mid-1960s, three independent researches reported the purification and chemical characterisation of three different kinds of inhibitors: inhibitor-B, abscission II and dormin. Later all the three were proved to be chemically identical. It was named abscisic acid (ABA).
Cousins confirmed the release of a volatile substance from ripened oranges that hastened the ripening of stored unripened bananas. Later this volatile substance was identified as ethylene, a gaseous PGR.
Physiological Effects of Plant Growth Regulators
Auxins (from Greek ‘auxein’ : to grow) was first isolated from human urine. The term ‘auxin’ is applied to the indole-3-acetic acid (IAA), and to other natural and synthetic compounds having certain growth regulating
properties. They are generally produced by the growing apices of the stems and roots, from where they migrate to the regions of their action. Auxins like IAA and indole butyric acid (IBA) have been isolated from plants.
NAA (naphthalene acetic acid) and 2, 4-D (2, 4-dichlorophenoxyacetic) are synthetic auxins. All these auxins have been used extensively in agricultural and horticultural practices.
Uses of Auxins
Auxins help to initiate rooting in stem cuttings, an application widely used for plant propagation. Auxins promote flowering e.g. in pineapples. They help to prevent fruit and leaf drop at early stages but promote the abscission of older mature leaves and fruits.
In most higher plants, the growing apical bud inhibits the growth of the lateral (axillary) buds, a phenomenon called apical dominance. Removal of shoot tips (decapitation) usually results in the growth of lateral buds (Figure 11). It is widely applied in tea plantations, hedge-making. Can you explain why?
Auxins also induce parthenocarpy, e.g., in tomatoes. They are widely used as herbicides. 2, 4-D, widely used to kill dicotyledonous weeds, does not affect mature monocotyledonous plants. It is used to prepare weed-free lawns by gardeners. Auxin also controls xylem differentiation and helps in cell division.
Gibberellins are another kind of promoter plant growth regulator. There are more than 100 gibberellins reported from widely different organisms such as fungi and higher plants. They are denoted as GA1, GA2, GA3 and so on. However, Gibberellic acid (GA3) was one of the first gibberellins to be discovered and remains the most intensively studied form. All GAs are acidic. They produce a wide range of physiological responses in the plants. Their ability to cause an increase in length of axis is used to increase the length of grapes stalks. Gibberellins, cause fruits like apple to elongate and improve its shape. They also delay senescence. Thus, the fruits can be left on the tree longer so as to extend the market period. GA3 is used to speed up the malting process in brewing industry.
Sugarcane stores carbohydrate as sugar in their stems. Spraying sugarcane crop with gibberellins increases the length of the stem, thus increasing the yield by as much as 20 tonnes per acre. Spraying juvenile conifers with GAs hastens the maturity period, thus leading to early seed production. Gibberellins also promotes bolting (internode elongation just prior to flowering) in beet, cabbages and many plants with rosette habit.
Cytokinins have specific effects on cytokinesis, and were discovered as kinetin (a modified form of adenine, a purine) from the autoclaved herring sperm DNA. Kinetin does not occur naturally in plants. Search for natural substances with cytokinin-like activities led to the isolation of zeatin from corn-kernels and coconut milk. Since the discovery of zeatin, several naturally occurring cytokinins, and some synthetic compounds with cell division promoting activity, have been identified. Natural cytokinins are synthesised in regions where rapid cell division occurs, for example, root apices, developing shoot buds, young fruits etc. It helps to produce new leaves, chloroplasts in leaves, lateral shoot growth and adventitious shoot formation. Cytokinins help overcome the apical dominance. They promote nutrient mobilisation which helps in the delay of leaf senescence.
Ethylene is a simple gaseous plant growth regulator. It is synthesised in large amounts by tissues undergoing senescence and ripening fruits. Influences of ethylene on plants include horizontal growth of seedlings, swelling of the axis and apical hook formation in dicot seedlings. Ethylene promotes senescence and abscission of plant organs especially of leaves and flowers. Ethylene is highly effective in fruit ripening. It enhances the respiration rate during ripening of the fruits. This rise in rate of respiration is called respiratory climactic.
Ethylene breaks seed and bud dormancy, initiates germination in peanut seeds, sprouting of potato tubers. Ethylene promotes rapid internode/petiole elongation in deep water rice plants. It helps leaves/ upper parts of the shoot to remain above water. Ethylene also promotes root growth and root hair formation, thus helping the plants to increase their absorption surface.
Ethylene is used to initiate flowering and for synchronising fruit-set in pineapples. It also induces flowering in mango. Since ethylene regulates so many physiological processes, it is one of the most widely used PGR in agriculture. The most widely used compound as source of ethylene is ethephon. Ethephon in an aqueous solution is readily absorbed and transported within the plant and releases ethylene slowly. Ethephon hastens fruit ripening in tomatoes and apples and accelerates abscission in flowers and fruits (thinning of cotton, cherry, walnut). It promotes female flowers in cucumbers thereby increasing the yield.
As mentioned earlier, abscisic acid (ABA) was discovered for its role in regulating abscission and dormancy. But like other plant growth regulators, it also has other wide ranging effects on plant growth and development. It acts as a general plant growth inhibitor and an inhibitor of plant metabolism. ABA inhibits seed germination. ABA stimulates the closure of stomata in the epidermis and increases the tolerance of plants to various kinds of stresses. Therefore, it is also called the stress hormone. ABA plays an important role in seed development, maturation and dormancy. By inducing dormancy, ABA helps seeds to withstand desiccation and other factors unfavourable for growth. In most situations, ABA acts as an antagonist to GAs.
We may summarise that for any and every phase of growth, differentiation and development of plants, one or the other PGR has some role to play. Such roles could be complimentary or antagonistic. These could be individualistic or synergistic.
Similarly, there are a number of events in the life of a plant where more than one PGR interact to affect that event, e.g., dormancy in seeds/ buds, abscission, senescence, apical dominance, etc.
Remember, the role of plant growth regulator is of only one kind of intrinsic control. Along with genomic control and extrinsic factors, they play an important role in plant growth and development. Many of the extrinsic factors such as temperature and light, control plant growth and development via PGR. Some of such events could be: vernalisation, flowering, dormancy, seed germination, plant movements, etc.