We observed diurnal and seasonal patterns of leaf-scale gas exchange within the crown of a Dipterocarpus sublamellatus Foxw. tree growing in a lowland dipterocarp forest at Pasoh, Peninsular Malaysia. Observations were carried out nine times over 6 years, from September 2002 to December 2007. Observation periods included both wet and mild-dry periods, and natural and saturated photosynthetic photon flux density (PPFD) light conditions. In situ measurements of the diurnal change in net photosynthetic rate and in stomatal conductance were carried out on canopy leaves of a 40-m-tall D. sublamellatus tree, which was accessed from a canopy corridor. A diurnal change in electron transport rate was observed under saturated PPFD conditions. The maximum net assimilation rate was approximately 10 micromol m(-2) s(-1). There was a clear inhibition of the net assimilation rate coupled with stomatal closure after late morning and this inhibition occurred year-round. Although the electron transport rate decreased alongside this inhibition, it sometimes followed on. Numerical analysis showed that the main factor in the inhibition of the net assimilation rate was patchy bimodal stomatal closure, which occurred in both mild-dry and wet periods. The midday depression occurred year-round, though there are fluctuations in soil moisture during the mild-dry and wet periods. The magnitude of the inhibition was not related to soil water content but was related to vapor pressure deficit (VPD): that is, whether the days were sunny and hot or cloudy and cool. On cloudy, cool days in the wet period, the net photosynthesis was only moderately inhibited, but it still decreased in the afternoon and was coupled with patchy stomatal closure, even in quite moderate VPD, leaf temperature and PPFD conditions. Our results suggest that patchy stomatal closure signaled by the increase in VPD, in transpiration and by circadian rhythms, was the key factor in constraining midday leaf gas exchange of the D. sublamellatus canopy leaves.
Effects of heterogeneity in stomatal behavior on gas-exchange characteristics of leaves from four tree species growing in different climates, including temperate, tropical monsoon and tropical rain forest, were investigated by combining gas-exchange measurements and the pressure-infiltration method. Field observations indicated linear relationships between whole-leaf conductance and the ratio of infiltrated to non-infiltrated leaf area (open stomata area) in Dipterocarpus sublamellatus Foxw. and Neobalanocarpus heimii (King) Ashton in a tropical rain forest in Peninsular Malaysia, whereas the ratio of infiltrated to non-infiltrated area rapidly increased up to the whole-leaf conductance at which the entire leaf was infiltrated in Cinnamomum camphora Sieb. in a temperate evergreen forest in Japan and in Azadirachta indica Juss. in a tropical monsoon area in Thailand. These results strongly suggest small ranges in bell-shaped stomatal conductance distributions in C. camphora and A. indica and bimodal stomatal conductance distributions in D. sublamellatus and N. heimii. The values of normalized maximum carboxylation rate at 25 degrees C (V(cmax25)) derived from gas-exchange measurements were not constant, but decreased with decreasing whole-leaf conductance in D. sublamellatus and N. heimii. A gas-exchange model analysis revealed a linear relationship between whole-leaf conductance and the ratio of infiltrated to non-infiltrated leaf area for bimodal stomatal conductance distributions, whereas for bell-shaped distributions, the relationships were nonlinear. Midday depression of apparent V(cmax25) in these species was mainly caused by bimodal stomatal closure. The bimodal stomatal distribution model could also explain diurnal changes in photosynthetic assimilation and transpiration rates in these species.
To evaluate water use and the supporting water source of a tropical rainforest, a 4-year assessment of evapotranspiration (ET) was conducted in Pasoh Forest Reserve, a lowland dipterocarp forest in Peninsular Malaysia. The eddy covariance method and isotope signals of rain, plant, soil, and stream waters were used to determine forest water sources under different moisture conditions. Four sampling events were conducted to collect soil and plant twig samples in wet, moderate, dry, and very dry conditions for the identification of isotopic signals. Annual ET from 2012 to 2015 was quite stable with an average of 1,182 ± 26 mm, and a substantial daily ET was observed even during drought periods, although some decline was observed, corresponding with volumetric soil water content. During the wet period, water for ET was supplied from the surface soil layer between 0 and 0.5 m, whereas in the dry period, approximately 50% to 90% was supplied from the deeper soil layer below 0.5-m depth, originating from water precipitated several months previously at this forest. Isotope signatures demonstrated that the water sources of the plants, soil, and stream were all different. Water in plants was often different from soil water, probably because plant water came from a different source than water that was strongly bound to the soil particles. Plants showed no preference for soil depth with their size, whereas the existence of storage water in the xylem was suggested. The evapotranspiration at this forest is balanced and maintained using most of the available water sources except for a proportion of rapid response run-off.