El Niño-Southern Oscillation forcing on carbon and water cycling in a Bornean tropical rainforest.

Carbon dioxide and water vapor exchanges between tropical forest canopies and the atmosphere through photosynthesis, respiration, and evapotranspiration (ET) influence carbon and water cycling at the regional and global scales. Their inter- and intra-annual variations are sensitive to seasonal rhythms and longer-timescale tropical climatic events. In the present study, we assessed the El Niño-Southern Oscillation (ENSO) influence on ET and on the net ecosystem exchange (NEE), using eddy-covariance flux observations in a Bornean rainforest over a 10-y period (2010-2019) that included several El Niño and La Niña events. From flux model inversions, we inferred ecophysiological properties, notably the canopy stomatal conductance and "big-leaf" maximum carboxylation rate (Vcmax25_BL). Mean ET values were similar between ENSO phases (El Niño, La Niña, and neutral conditions). Conversely, the mean net ecosystem productivity was highest during La Niña events and lowest during El Niño events. Combining Shapley additive explanation calculations for nine controlling factors with a machine-learning algorithm, we determined that the primary factors for ET and NEE in the La Niña and neutral phases were incoming shortwave solar radiation and Vcmax25_BL, respectively, but that canopy stomatal conductance was the most significant factor for both ET and NEE in the El Niño phase. A combined stomatal-photosynthesis model approach further indicated that Vcmax25_BL differences between ENSO phases were the most significant controlling factor for canopy photosynthesis, emphasizing the strong need to account for ENSO-induced ecophysiological factor variations in model projections of the long-term carbon balance in Southeast Asian tropical rainforests.

Image analysis procedure for the optical scanning of fine-root dynamics: errors depending on the observer and root-viewing window size.

Clarifying the dynamics of fine roots is critical to understanding carbon and nutrient cycling in forest ecosystems. An optical scanner can potentially be used in studying fine-root dynamics in forest ecosystems. The present study examined image analysis procedures suitable for an optical scanner having a large (210 mm × 297 mm) root-viewing window. We proposed a protocol for analyzing whole soil images obtained by an optical scanner that cover depths of 0-210 mm. We tested our protocol using six observers with different experience in studying roots. The observers obtained data from the manual digitization of sequential soil images recorded for a Bornean tropical forest according to the protocol. Additionally, the study examined the potential tradeoff between the soil image size and accuracy of estimates of fine-root dynamics in a simple exercise. The six observers learned the protocol and obtained similar temporal patterns of fine-root growth and biomass with error of 10-20% regardless of their experience. However, there were large errors in decomposition owing to the low visibility of decomposed fine roots. The simple exercise revealed that a smaller root-viewing window (smaller than 60% of the original window) produces patterns of fine-root dynamics that are different from those for the original window size. The study showed the high applicability of our image analysis approach for whole soil images taken by optical scanners in estimating the fine-root dynamics of forest ecosystems.

Vertical variations in wood CO2 efflux for live emergent trees in a Bornean tropical rainforest.

Difficult access to 40-m-tall emergent trees in tropical rainforests has resulted in a lack of data related to vertical variations in wood CO2 efflux, even though significant variations in wood CO2 efflux are an important source of errors when estimating whole-tree total wood CO2 efflux. This study aimed to clarify vertical variations in wood CO2 efflux for emergent trees and to document the impact of the variations on the whole-tree estimates of stem and branch CO2 efflux. First, we measured wood CO2 efflux and factors related to tree morphology and environment for seven live emergent trees of two dipterocarp species at four to seven heights of up to ∼ 40 m for each tree using ladders and a crane. No systematic tendencies in vertical variations were observed for all the trees. Wood CO2 efflux was not affected by stem and air temperature, stem diameter, stem height or stem growth. The ratios of wood CO2 efflux at the treetop to that at breast height were larger in emergent trees with relatively smaller diameters at breast height. Second, we compared whole-tree stem CO2 efflux estimates using vertical measurements with those based on solely breast height measurements. We found similar whole-tree stem CO2 efflux estimates regardless of the patterns of vertical variations in CO2 efflux because the surface area in the canopy, where wood CO2 efflux often differed from that at breast height, was very small compared with that at low stem heights, resulting in little effect of the vertical variations on the estimate. Additionally, whole-tree branch CO2 efflux estimates using measured wood CO2 efflux in the canopy were considerably different from those measured using only breast height measurements. Uncertainties in wood CO2 efflux in the canopy did not cause any bias in stem CO2 efflux scaling, but affected branch CO2 efflux.

Transpiration characteristics of a rubber plantation in central Cambodia.

The rapid and widespread expansion of rubber plantations in Southeast Asia necessitates a greater understanding of tree physiology and the impacts of water consumption on local hydrology. Sap flow measurements were used to study the intra- and inter-annual variations in transpiration rate (Et) in a rubber stand in the low-elevation plain of central Cambodia. Mean stand sap flux density (JS) indicates that rubber trees actively transpire in the rainy season, but become inactive in the dry season. A sharp, brief drop in JS occurred simultaneously with leaf shedding in the middle of the dry season in January. Although the annual maxima of JS were approximately the same in the two study years, the maximum daily stand Et of ∼2.0 mm day(-1) in 2010 increased to ∼2.4 mm day(-1) in 2011. Canopy-level stomatal response was well explained by changes in solar radiation, vapor pressure deficit, soil moisture availability, leaf area, and stem diameter. Rubber trees had a relatively small potential to transpire at the beginning of the study period, compared with average diffuse-porous species. After 2 years of growth in stem diameter, transpiration potential was comparable to other species. The sensitivity of canopy conductance (gc) to atmospheric drought indicates isohydric behavior of rubber trees. Modeling also predicted a relatively small sensitivity of gc to the soil moisture deficit and a rapid decrease in gc under extreme drought conditions. However, annual observations suggest the possibility of a change in leaf characteristics with tree maturity and/or initiation of latex tapping. The estimated annual stand Et was 469 mm year(-1) in 2010, increasing to 658 mm year(-1) in 2011. Diagnostic analysis using the derived gc model showed that inter-annual change in stand Et in the rapidly growing young rubber stand was determined mainly by tree growth rate, not by differences in air and soil variables in the surrounding environment. Future research should focus on the potentially broad applicability of the relationship between Et and tree size as well as environmental factors at stands different in terms of clonal type and age.

Patchy stomatal behavior during midday depression of leaf CO2 exchange in tropical trees.

We investigated effects of heterogeneous stomatal behavior on diurnal patterns of leaf gas exchange in 10 tree species. Observations were made in middle and upper canopy layers of potted tropical rainforest trees in a nursery at the Forest Research Institute Malaysia. Measurements were taken from 29 January to 3 February 2010. We measured in situ diurnal changes in net photosynthetic rate and stomatal conductance in three leaves of each species under natural light. In both top-canopy and sub-canopy species, midday depression of net assimilation rate occurred in late morning. Numerical analysis showed that patchy bimodal stomatal behavior occurred only during midday depression, suggesting that the distribution pattern of stomatal apertures (either uniform or non-uniform stomatal behavior) varies flexibly within single days. Direct observation of stomatal aperture using Suzuki's Universal Micro-Printing (SUMP) method demonstrated midday patchy stomatal closure that fits a bimodal pattern in Shorea leprosula Miq., Shorea macrantha Brandis. and Dipterocarpus tempehes V.Sl. Inhibition of net assimilation rate and stomatal conductance appears to be a response to changes in vapor pressure deficit (VPD). Variable stomatal closure with increasing VPD is a mechanism used by a range of species to prevent excess water loss from leaves through evapotranspiration (viz., inhibition of midday leaf gas exchange). Bimodal stomatal closure may occur among adjacent stomata within a single patch, rather than among patches on a single leaf. Our results suggest the occurrence of patches at several scales within single leaves. Further analysis should consider variable spatial scales in heterogeneous stomatal behavior between and within patches and within single leaves.