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Three terms used routinely to describe various aspects of the changes that a plant
undergoes during its life cycle are growth, differentiation, and development.
3.1.2. Growth
Growth is the quantitative term, related to changes in size and mass. It can be assessed
by a variety of quantitative measures. Growth of cells in culture is sometimes measured as in
increase in cell number or the fresh weight of packed cells. For higher plants, however fresh
weight is not always a reliable measure. Although most plant tissues are approximately 80%
water, water content is highly variable and fresh weight will fluctuate widely with changes in
the water status of the plant. Dry weight, a measure of the amount of protoplasm or dry
matter, is used more often than fresh weight, but even dry weight can be misleading as a
measure of growth.
3.1.3. Differentiation
Differentiation is a qualitative term, referring to the differences other than size that arise
among cells, tissues and organs. Differentiation occurs when a dividing cell gives rise to two
daughter cells destined to assume different anatomical characteristics and functions. In the
earliest stage of development, for example, division of the zygote gives rise to cells that will
become the root or shoot of the plant. Unspecialized parenchyma cells differentiate into
xylem vessels or phloem sieve tubes, each with a distinct of morphology and specialized
function. Differentiation does not lend itself easily to quantitative interpretation but must
normally be described as a series of qualitative changes. Finally, although growth and
differentiation are normally concurrent events, examples abound of growth without
differentiation and differentiation without growth.
Differentiation is a two-way street. Even though plant cells may appear to be highly
differentiated or specialized, they may often be stimulated to revert to a more embryonic
form; that is cells dedifferentiate. It is a though the cells have been genetically
reprogrammed, allowing them to reverse the process and them to differentiate along new and
different paths.
The ability of differentiated cells to regenerate new plants demonstrates that all living
plants cells retain complete a genetic program, even though not all of the information is
actively used by the cell at any given time. This concept is known as totipotency. All that is
required to change the pattern of differentiation is the right input to select the appropriate
genetic information at the right time.
Table 3.1. The influence of plant hormone groups on different categories of development. An x
indicates a demonstrated effect of that hormone group on one or more aspects of that developmental
category. The absence of an x does not mean that the hormone is ineffective, only that an effect has not
been reported in literature. (Reprinted from G. Hopkins: Introduction to Plant Physiology. John
Wiley et Sons, New York, 1995.)
There are currently five recognized groups of plant hormones: auxins, gibberellins,
cytokinins, abscisic acid, and ethylene. In addition to the five principal hormones, two other
groups sometimes appear to be active in regulating plant growth, the brassinosteroids and
polyamines. Each of these groups will be briefly introduced at this time, followed by a
general discussion of its physiological role. The influence of plant hormone groups on
different categories of development we can see in Table 3.1.
Fig. 3.4. (1)The control of apical dominance by auxin. Experimental material: Field bean
(Vicia faba). After removal of the apical terminal bud (a) side buds grow in the lower leaf
axils (b). If the cut apical surflace is covered by an agar block containing IAA, lateral buds
remain inhibited (c). A control block (without IAA) has no effect (d). IAA is therefore able to
replace the end bud with respect to apical dominance. (from Mohr et Schopfer 1995, after
Bonner et Galston 1952)
Fig. 3.4. (2) Apical dominance in broadbean
(Vicia faba) (A) Control plants. (B) Removal
of the stem apex, a source of auxin, promotes
axillary bud growth. (C) Dominance can be
restored by applying auxin (in lanolin paste)
to the cut stem surface. (Reprinted from G.
Hopkins: Introduction to Plant Physiology.
John Wiley et Sons, New York, 1995.)
Fig. 3.5. Auxin (IAA) and cytokinin (kinetin) as factors limiting mitotic activity and formation
of organs in a tissue culture. Experimental material: Explant from stem pith of a tobacco
plant (Nicotiana tabacum). Relatively high concentrations of IAA and kinetin lead to the
formation of a callus after a few weeks. Development can be redirected to root or shoot
formation by either increasing or decreasing the ratio of IAA/kinetin. (from Mohr et Schopfer
1995, after Ray 1963)
Apical dominance is the inhibition of the development of some or all of the lateral buds
by the terminal (apical) bud of shoot. Removal of the terminal bud releases some of the lateral
buds from inhibition. This implies that a substance produced at the apex, most probably auxin,
is responsible for the inhibition (see Figure 3.4.). Cytokinins have been shown to promote the
growth of lateral buds.
Many effects of auxins are brought about by the combined action of auxin with other
growth substances. For example, the stimulation of cambial activity, the induction of
parthenocarpy, and the enhancement of internode elongation are all more effectively
promoted by a combination of auxin and gibberellin than by either substance alone. Similarly
appropriate concentrations of auxin and cytokinin are needed in culture media to promote cell
division in tissue explants. Depending on the relative concentration of each, root meristems
(high auxin: low cytokinin) or shoot meristems (low auxin: high cytokinin) may be initiated
(see Figure 3.5.).
Fig. 3.6. (A) Dwarfism in a GA deficiency mutant and its reversal by GA application.
Experimental material: Dwarf-5 mutant (D-5/d-5) and wild type of maize (Zea mays). The
mutant received in total 250 μg GA3 applied to the apex from the seedling stage at intervals
of 2 to 5d. From left to right: Wild type, untreated; wild type, treated; mutant, untreated;
mutant, treated. Analysis shows that stunted growthis based on the failure of one gene, which
controls the cyclisation of copalylpyrophosphate to ent-kaurene. The phenotypic defect can
therefore be reversed by application of ent-kaurene. (after Phinney and West 1960, from
Mohr et Schopfer 1995)
Fig. 3.6. (B) The effect of gibberellic acid on dwarf
pea. Left: Control, showing reduced internode
elongation. Right: Gibberellin treated. Enhanced
internode elongation following a foliar-drench
with 5 x 10-4 M gibberellin. (after Hopkins 1995)
Fig. 3.8. The effects of cytokinins on growth and development of cereal plants. (After Kamínek
et al. 2003, reprinted from Macháčková et Romanov (eds.): Phytohormones in Plant
Biotechnology and Agriculture. Kluwer Academic Publishers, Dordrecht, 2003).
Generally the most obvious effects of cytokinins include delay of senescence mature
leaves. Senescence is characterized by the breakdown of protein, nucleic acids and other
macromolecules, a loss of chlorophyll, and the accumulation of soluble nitrogen products
such as amino acids. Cytokinins also induce of flowering in certain species, and cut across of
dormancy in axillary buds and some seeds. Endogenous cytokinins are found in the greatest
concentrations in embryos and developing fruits, e.g. in the 'milk' of the coconut. The
application of cytokinins stimulates also release of axillary buds from apical dominance,
thus antagonizing the effect of auxins.