Oxygen deficiency in the rooting zone occurs with poor drainage after rain or irrigation, causing depressed growth and yield of dryland species, in contrast with native wetland vegetation that tolerates such conditions. This review examines how roots are injured by O2 deficiency and how metabolism changes during acclimation to low concentrations of O2. In the root apical meristem, cell survival is important for the future development; metabolic changes under anoxia help maintain cell survival by generating ATP anaerobically and minimizing the cytoplasmic acidosis associated with cell death. Behind the apex, where cells are fully expanded, ethylene-dependent death and lysis occurs under hypoxia to form continuous, gas-filled channels (aerenchyma) conveying O2 from the leaves. This selective sacrifice of cells may resemble programmed cell death and is distinct from cell death caused by anoxia. Evidence concerning alternative possible mechanisms of anoxia tolerance and avoidance is presented.
SUMMARY
Barley plants were grown for 21 days in sand culture, continuously irrigated with nutrient solution. The rooting depth was divided into three compartments, one above another, such that different zones of the root system could be supplied with very low or high concentrations of a single inorganic nutrient, all other nutrients being maintained at a high concentration.
Exposure of parts of the main seminal roots (axes) to high concentrations of phosphate caused a localized promotion of the initiation and subsequent extension of both first and second order laterals, compared with zones receiving very low concentrations of phosphate. This resulted in considerable modification to root form, but with only a small depression in shoot growth, compared with control plants receiving an ample supply to all parts of the root system. The extension of seminal axes was little affected by the concentration of phosphate to which they were exposed.
Similar responses to those described for phosphate occurred with variation in concentration of nitrate or ammonium, but with potassium a localized supply promoted the growth of laterals to approximately the same extent as controls throughout the entire root system.
The experiments show that adequate external concentrations of nitrogen and phosphorus are required by any part of the root system for optimal growth of laterals, but not axes. Possible mechanisms which compensate shoot growth when nutrients are supplied to only part of the root system, and agronomic implications, are discussed.
The roots and stem base of intact, 10 day old maize (Zea mays L. cv. LG11) plants, grown in nutrient solution, were continuously aerated either with ethylene (5 μl l(-1)) in air or with air alone. Ethylene treatment hastened the emergence of adventitious (nodal) roots from the base of the shoot, but slowed their subsequent extension. Ethylene also promoted the collapse of cells in the cortex of these roots, with lysigenous development of prominent air spaces (aerenchyma). Non-aeration of the nutrient solution caused endogenously produced ethylene to accumulate in the roots, and stimulated both the emergence of adventitious roots and the formation of cortical air spaces in them. With non-aeration the concentration of oxygen did not fall below 1% in the equilibrium gas phase (air=20.8%). Complete deoxygenation of the nutrient solution, produced by passing oxygen-free nitrogen gas, prevented both air space formation and the evolution of ethylene by root segments.These results suggest that adventitious rooting and cortical air space formation in nodal roots in Zea mays may be stimulated by enhanced concentrations of endogenous ethylene arising either from entrapment of the gas by unstirred water layers around the roots and/or by increased biosynthesis. These responses are considered conducive to survival in waterlogged soil.
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