When the detached first leaves of green or etiolated oat (Avena sativa cv. Victory) seedlings senesce in the dark, their oxygen consumption shows a large increase, beginning after 24 hours and reaching a peak of up to 2.5 times the initial rate by the 3rd day. This effect takes place while the chlorophyll of green leaves, or the carotenoid of etiolated leaves, is steadily decreasing. Kinetin, at a concentration which inhibits the decrease in pigment, completely prevents the respiratory rise; instead, the oxygen consumption drifts downwards. Lower kinetin concentrations have a proportional effect, 50% reduction of respiration being given by about 0.1 mg/l. About one-fifth of the respiratory rise may be attributed to the free amino acids which are liberated during senescence; several amino acids are shown to cause increases of almost 50% in the oxygen consumption when supplied at the concentrations of total amino acid present during senescence. A smaller part of the rise may also be due to soluble sugars liberated during senescence, largely coming from the hydrolysis of a presumptive fructosan. The remainder, and the largest part, of the increase is ascribed to a natural uncoupling of respiration from phosphorylation. This is deduced from the fact that dinitrophenol causes a similar large rise in the oxygen consumption of the fresh leaves or of leaf segments kept green with kinetin, but causes only a very small rise when the oxygen consumption is near its peak in senescent controls. The respiration of these leaves is resistant to cyanide, and 10 mM KCN even increases it by some 30%; in contrast, etiolated leaves of the same age, which undergo a similar rise in oxygen consumption over the same time period, show normal sensitivity to cyanide. The respiratory quotient during senescence goes down as low as 0.7, both with and without kinetin, though it is somewhat increased by supplying sugars or amino acids; glucose or alanine at 0.3 M bring it up to 1.0 and 0.87, respectively. N6-Benzylaminopurine and A-2-isopentenylaminopurine act similarly to kinetin in repressing the respiratory rise, the former being five times as active as kinetin, while the latter has only 1 % of the activity of kinetin. Zeatin also powerfully prevents senescence. Because the repression of the respiratory rise is shown by each cytokinin at the concentration at which it inhibits senescence, the action is ascribed in both cases to the maintenance of a tight coupling between respiration and phosphorylation. It is pointed out that such an effect would explain many features of cytokinin action.I This work was supported in part by Grants 35238X and 11867 from the National Science Foundation to K. V. Thimann.'This paper is dedicated to the memory of Solon A. Gordon because of his continued interest in plant metabolism and his many contributions to it, and in recognition of a long standing friendship.A change in the methodology of the senescence experiments is described and compared with the method previously used, and the influence of temperat...
A comparison has been made of the progress of senescence in the first leaf of 7-day-old oat plants (Avena sativa cv. Victory) in darkness and in white light. Light delays the senescence, and intensities not over 100 to 200 ft-c (1000-2000 lux) suffice for the maximum effect. In such intensities, chlorophyll loss and amino acid liberation stiOl go on in detached leaves at one-third to one-half the rate observed in darkness; however, when the leaves are attached to the plant, the loss of chlorophyH in 5 days is barely detectable. (17); in the last instance, a long photoperiod was found more effective than a short one. In rice, red light was effective in delaying senescence and far red had the opposite effect (16). In the presence of EDTA, however, white light was reported to promote bleaching (11), and although this effect has twice been confirmed (9, 23), its relationship to normal senescence in light is not clear. Recently, Takegami (20) has found light to prevent the decrease in total RNA which occurs in senescing tobacco leaves. The effect differed in the different fractions, the fall in 16S and 23S components being somewhat inhibited, whereas the 18S and 25S fractions were actually increased by white light.
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