Measurements of photosynthesis and respiration were made on leaves in summer in a Quercus rubra L. canopy at approximately hourly intervals throughout 5 days and nights. Leaves were selected in the upper canopy in fully sunlit conditions (upper) and in the lower canopy (lower). In addition, leaves in the upper canopy were shaded (upper shaded) to decrease photosynthesis rates. The data were used to test the hypothesis that total night-time respiration is dependent on total photosynthesis during the previous day and that the response is mediated through changes in storage in carbohydrate pools. Measurements were made on clear sunny days with similar solar irradiance and air temperature, except for the last day when temperature, especially at night, was lower than that for the previous days. Maximum rates of photosynthesis in the upper leaves (18.7 mumol m(-2) s(-1)) were approximately four times higher than those in the lower leaves (4.3 mumol m(-2) s(-1)) and maximum photosynthesis rates in the upper shaded leaves (8.0 mumol m(-2) s(-1)) were about half those in the upper leaves. There was a strong linear relationship between total night-time respiration and total photosynthesis during the previous day when rates of respiration were normalized to a fixed temperature of 20degreesC, removing the effects of temperature from this relationship. Measurements of specific leaf area, nitrogen and chlorophyll concentration and calculations of the maximum rate of carboxylation activity, V-cmax, were not significantly different between upper and upper shaded leaves 5 days after the shading treatment was started. There were small, but significant decreases in the rate of apparent maximum electron transport at saturating irradiance, J(max) (P>0.05), and light use efficiency, epsilon (P<0.05), for upper shaded leaves compared with those for upper leaves. This suggests that the duration of shading in the experiment was sufficient to initiate changes in the electron transport, but not the carboxylation processes of photosynthesis. Support for the hypothesis was provided from analysis of soluble sugar and starch concentrations in leaves. Respiration rates in the upper shaded leaves were lower than those expected from a relationship between respiration and soluble sugar concentration for fully exposed upper and lower leaves. However, there was no similar difference in starch concentrations. This suggests that shading for the duration of several days did not affect sugar concentrations but reduced starch concentrations in leaves, leading to lower rates of respiration at night. A model was used to quantify the significance of the findings on estimated canopy CO2 exchange for the full growing season. Introducing respiration as a function of total photosynthesis on the previous day resulted in a decrease in growing season night-time respiration by 23% compared with the value when respiration was held constant. This highlights the need for a process-based approach linking respiration to photosynthesis when modelling long-term carbon exchange in forest ecosystems.
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