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Auxins and the fungal inhibitor fusicoccin. This fungal agent has very dramatic shortterm stimulatory
effects on growth of a wide array of plant tissues, and always causes a simultaneous drop in pH (Marre et
al., 1974).

A different approach to the rapit responses to auxin was proposed by evans an ray (1969), who
suggested that some growth-limiting protein (or other limiting substrate) may be used up in the rapid
growth reaction. Penny (1971) did some clever experiments whitch led her to conclude that there is
indeed such a growth-limiting protein; when she blocked its synthesis with cycloheximide, she estimated
that this pool has a half-life of 12 to 17 min; she asserts that once this pool is exhausted, new protein
synthesis is necessary before the rapid auxin response can be obtained again

It has been reported that cyclic AMP can markedly enhance auxin stimulations of growth (Kamisaka and
Masuda, 1970). And one might infer that this mediating factor for the actions of many animal hormones
may be involved likewise in auxin action. The effects of added cyclic AMP are-certainly
small(Hartung,1972), but, on the other hand, some plants are very rich in the enzymes which split the
cyclic phosphate ring (Lin and Varner, 1972), and this could make in vivo tests rather insensitive to added
cyclic AMP. Another candidate for a cofactor of hormone action is a stimulator of RNA polymerase, which
can be displaced off membrane preparations by auxin (Hardin et al , 1972). Such a cofactor could well
provide a link between a plasmalemma site of auxin attachment and the auxin regulation of nucleic acid-
directed protein synthesis.

AUXIN INHIBITIONS
Since inhibitory actions of auxin may be even more widespread than protive actions on growth. It is
unfortunate that so little information is avaiblelable on their mechanisms. Three suggestions have been
made to explain auxin inhibitions. Burstrom (1955) has concluded from work with wheat-root inhibitions
that auxin may catalyze the hardening of the cell wall, thus leading to a shorter duration of growth and a
shorter final cell length. The enhanced hardening action is deduced from data like those in fig. 4-31. A
second suggested expenetion of auxin inhibitions is that the hormone exerts a self-inhibitions by the
attachment of separate molekules each of the two presumed sites of auxin attachment (foster et
al.,1952).this type of action would be similar to the substrate in hibition of some enzymes, in which
there is more than one attachment site for a substrate. Each of the two molecules would serve to
prevent the completion of two- point attachment by the other molecule. A third suggested explenation
of auxin inhibition is thst ethylene is responsible for the inhibitory effect ( Burg and Burg, 1966). In this
case, the inhibition would be due to an action quite separate from the mechanism of growth stimulation.
The ethy ene explenation rests principally on the finding that inhibition occur at concentrations of auxin
which do stimulate ethylene formation, as shown in fig. 4-32. It may serve to explain auxin inhibitions of
peasteam growth and inhibitons of bud growth but apparently not inhibitions of coleooptile growth
since these tissues produce little ethylene and are fairly insensitive to it (burg and burg, 1968a).

It is appealing to think that the characteristic two-phase curve of auxin responses, like those in fig. 4-2
and 4-3, might reflect a single auxin action resultin in diverse growth responses; however, since many
tissues show very strong auxin inhibitions and little or no auxin promotive responses, e.g., in roots, the
idea of selfinhibition or inhibition by self competition reactions becomes less attractive and the idea of
inhibition by mechanisms quite separate from promotive actions seems a more reasonable alternative.
Thus the auxin inhibition of wheat roots may be through a stimulation of cell-well hardening; the
inhibition of pea steams may be through the stimulation of ethlene production;

[grafik]

Fig. 4-32 auxin concentrations which inhibit the grouth of pea-stem sections, as indicated by the
decrease in length achieved, are correlated with those concentrations which b-ing about ethylene
biosynthesis (Burg and Burg, 1966)

The inhibition of bud growth may be a result of ethylene production and the consequent inhibition of
cell division (Burg and Burg, 1968b); and the inhibition of abscission may be through a suppression of
senescence (Abeles et al.,1967).

SUMMARY
Three major features of auxins stand out strikingly: (1) the diversity of auxin effects, (2) the diversity of
the other chemical control which may be interwomen witth the auxin effects, and (3) the systemic
patterns of the auxin effects.

The diverse effects of auxins are readily apparent in the large ar-d expanding list of growth and
differntiation activities which are influenced by endogenous auxins (of; Table 18-1) in addition to cell-
elongation, epical dominance, and abscission processes, there are effects on flower initiation and
development, pollen-tube growth, fruit-set and fruit growth the formation of compression wood in
conhers tuber and bulb formation, and seed germination. Almost every dynamic part of plant growth
and development seems to be affected by auxin. In the individuall cell there are effects on the plasticity
and elasticity of the wall, on cytoplasmic viscosity. On protoplasmic streaming, on respiration rates, on
metabolic [athways, on changes in oxidative states, on the contents of nucleic acids, and on the activities
of many enzymes.

Allied with the diversity of physiological effects of auxins is a diversity of moleculas species which can
bring about the responses. The fact that IAA can share its biological effectiveness with such diverse
synthetic chemicals as naphthaleneacetic acid, 2,4-dichlorophenoxyaceticacid, 2,3,6-trichlorobenzoic
acid, and a host of others attests to the wide range of compounds which can act as auxins in plants. Most
auxins are able not only to stimulate growth of stems and coleoptiles but also to alter differentiation,
abacission, and other developmental effects. The systemic correlative effects are not shared by all
auxins, for the limited ability of many to move in the auxin-transport system restricts their partisipation
in normal correlation effects. They are readily swept along in the stream of phloem translocation, but
such movement does not conform to the polar qualities of the hormone-transport system.

The diversity of agents which can participate in growth regulation is a strikingly repetitive theme. Cell
enlargement may be directed by gibberellins, auxins, kinins, ethylene, and various inhibitors; thre are
strong growth-promoting effects of chelating agents, fatty acids, and even organic acids. When auxin was
first discovered, went (1928) said, Ohne Wuchsstoff, kein Wachstum. Now there is evidence that such a
concise, all-or-none dependence on auxin is not a general characteristic of cell enlargement (cf. Kefford
and Goldacre, 1961). Many cells do not appear to need auxin for growth. The auxin influerces on
physiological and developmental processes generally require other substances as cofactors, including the
factors for cell division, cell enlargement, xylem differentiation, root initiation, and apical dominance.
The chemicals regulating growth and development are manifold.

The systemastic patterns of auxin effects in the plant, representing thr correlation effects on growth and
differentiation, reveal auxin as a chemical messenger influencing many patterns of plant development. A
system of chemical mesengers is a principal ingredient for the creation of a multicellular organism out of
what would otherwise be only a multicellular colony, and auxin is the outstanding known participant in
such a control system

General REFERENCES
Audus, L. J. 1972. Plant Growth Substances. Leonard Hill Books, London. 533 pp.
Evanar, M. L. 1973. Rapid stimulation of plant cell elongation by hormonal and non-hormonal factors.
Bioscience, 23:711-718.
Thimann, K. V. 1972. The natural plant hormone. F. C. Steward, ed..Plant Physiology, a Treatise.
Academic Press, New York, pp. 3-332.