The effects of ontogeny and environmental oscillations on plant responses to oxygen deprivation

Abstract

The effects of ontogeny and environmental oscillations on plant responses to oxygen deprivation were investigated for a wide spectrum of species, ranging from agricultural to forestry crops, and from temperate to tropical plants. The extent to which hypoxia- or anoxia-tolerance were affected by ontogeny and environmental oscillations, was assessed mainly through changes in survival and growth and physiological parameters such as respiratory activity, ethanol production and carbohydrate depletion. Anoxia-tolerance of germinating seeds of chickpea (Cicer arietlnum L.) was found to vary, even within the earlier stages of germination, according to the length of the aerobic imbibition period previous to the anoxic shock. The notable fact was that 6 minutes of seed aerobic imbibition prior to anoxic treatment was sufficient to significantly increase post-anoxic survival after 4 days anoxia, compared to seeds not allowed to previously imbibe aerobically. These survival results were mirrored by the significant increase in the oxygen uptake by the embryos of seeds which were allowed to imbibe aerobically for 2 hours prior to anoxia, compared to the embryos of anaerobically imbibed seeds. Germination stage also affected the response of barley (Hordeum vulgare L.) to washing under anoxia, a treatment used to investigate membrane stability and other factors associated with anoxic injury. Temperature, frequency of washing and nutrients present in the washing solution also affect the post-anoxic responses of barley. Daily washing of seedlings under anoxia was often detrimental to post-anoxic survival. One washing only, at the end of the anoxic period, often enhanced survival. This positive effect seemed to be more linked to protection against plasmamembrane leakage due to calcium ions present in the washing solution than to removal of anaerobically-produced potentially toxic volatiles. Two Brazilian tree species were also studied. The seeds of Parkia pendula (Willd.) Benth. ex. Walp., a species typically present in unflooded areas in the Amazon, were still able to germinate after seven months submergence. P.pendula one-month old seedlings, however, did not survive longer than one month flooding, which can be a considerable disadvantage in the ca. six-month long flooding period of the Amazon floodplains. However, adult trees can still be found, although rarely, in flooded areas. The possible strategies involved in an eventual establishment of P.pendula individuals in flooded areas of the Brazilian Amazon are discussed. A contrast is drawn between the responses to flooding of this species and flood-tolerant Parkia discolor. Enterolobium contortisiliquum (Veil.) Morong, is a tree species which is present in both the dry soils of the cerrados (neotropical savannas) in Central Brazil and in the flood-prone Gallery Forests. This species showed considerable tolerance to flooding and drought, as reflected by the various morphological and metabolic adaptations observed in response to these stresses. The role of the xylopodium, a rigid wood tuber, in such tolerance to flood and drought stresses is discussed. Water-stressed roots of some crop species studied presented higher ethanol levels than control plants. Additionally, these same drought-treated roots showed a considerable amount of shrinkage compared to control roots, as measured by root diameter. It is argued that drought causes root shrinkage, which possibly reduces ability of such roots to capture oxygen and results in hypoxia in the tissues and consequent increase in ethanol production. This hypothesis of drought-induced hypoxia is compared with several recent findings in the literature, and is discussed as a possible factor which allows drought, under specific circumstances, to acclimatise plants to a subsequent flooding. From preliminary experiments with alternation of flooding and drought in Eucalyptus species, it appeared that a previous stress affects a plant's response to a subsequent stress. This effect was not always negative, and in E.regnans a five-week drought allowed a subsequent 3-fold increase in flooding survival. This experiment, however, needs to be repeated in order to confirm these results. A common cause for anaerobic injury seems to be unlikely for the diverse plants studied, and anoxia survival often seemed to be related to a combination of morphological and metabolic adaptations. A critical reflection on the risks of labelling plants as tolerant or sensitive to oxygen deprivation is provided, as well as a discussion on the perspectives of applied research which may further the development of ecophysiological theory.