Involvement of The Hormones Ethylene and Abscisic Acid in Some Adaptive Responses of Plants to Submergence, Soil Waterlogging and Oxygen Shortage
Ethylene and abscisic acid are two of five substances or groups of substances with established regulatory (hormonal) roles in plant development. The lack of detailed discussion here about the three remaining groups of hormones (gibberellins, cytokinins and auxins) does not imply that they are unimportant for the growth of inundated plants. Instead, it reflects a lack of experimental attention, despite some early and promising work with these compounds in waterlogged tomato and sunflower (Phillips, 1964; Burrows and Carr, 1969; Reid, Crozier and Harvey, 1969; Railton and Reid, 1973). Evidence that implicates ethylene in plant responses to over-wet conditions is particularly strong and comes from research from several laboratories working with different species and growth processes over a period of approximately 15 years. Studies with abscisic acid (ABA) are less extensive and have concentrated on stomatal closure. Only with the recent adoption of reliable physicochemical methods for ABA analysis, and the use of ABA-deficient mutants has the part played by this hormone in closing the stomata of waterlogged plants been demonstrated convincingly. Hormones such as ethylene and abscisic acid, possess several characteristics which make them likely mediators of developmental change initiated by aeration stress. First, they can be active in extremely small concentrations (e.g. ethylene is active in promoting leaf epinasty at 0.01 volumes per million, Leather, Forrence and Abeles, 1972). This suggests that quite small changes in hormone titre, induced quickly by environmental factors, can be physiologically significant. Secondly, only certain tissues or cells within a plant or organ are highly responsive to a particular hormone at a given time. Thus hormone effects are not located indiscriminately about the plant but instead are selective for particular tissues and locations. Such spatial precision is a prerequisite for any explanation of how excess water comes to modify specific morphogenetic processes. Hormones are also known to act very quickly. For example, applications of ABA to a leaf may close stomata within 5 min (Cummins, Kende and Raschke, 1971). This rapidity of action is necessary if hormones are to explain several effects of inundation which can be very fast (e.g. less than 10 min, Musgrave and Walters, 1974). At least one of the responses to inundation can also be reversed if water levels fall. It is therefore apposite that the effects of some hormones are reversible upon withdrawing treatment. The overriding characteristic of hormones that links them with aeration stress is the close visual similarity between several morphological consequences of flooding and those documented in the hormone literature as reponses to application of these substances (see Jackson and Goss, 1978; Jackson and Drew, 1984; Reid and Bradford, 1984 for reviews). The extent to which hormone-mediated developmental phenomena aid survival by aeration-stressed plants is uncertain, although this seems likely on intuitive grounds. However, there is as yet little direct experimental evidence on this point.
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