Yoshiko Kosugi

Impact of leaf physiological characteristics on seasonal variation in CO2, latent and sensible heat exchanges over a tree plantation

This study investigated the impact of leaf physiological characteristics on CO 2, sensible heat, and latent heat exchanges over a plant community. Measurements were carried out over plantation trees using an eddy correlation system, under well-watered soil conditions for several days spanning the year, and under soil drought conditions during autumn only. The following leaf physiological characteristics were investigated: the maximum rate of carboxylation (VcMAX25: [Planta 149 (1980) 78–90]), the relationship between stomatal conductance (gs) and net assimilation rate (An) adjusted by relative humidity (hs), and CO2 concentration at the leaf surface (cs) [An analysis of stomatal conductance. Ph.D. Thesis. Stanford University, CA, 89 pp.]. As the leaf physiological parameters, VcMAX and the relationship between stomatal conductance (gs) and Anhs/cs of four evergreen trees were estimated from gas exchange measurements at the single leaf scale. Numerical simulations were carried out using a multi-layer model consisting of a second-order closure model for atmospheric diffusion coupled with a radiation transfer model, a Farquhar type photosynthesis model, and Ball’s stomatal conductance model. The results suggest that the value of VcMAX25 changes every day, and that appropriate values should be used in simulations. The relationship between stomatal conductance (gs) and Ahs/cs changed under soil drought conditions, and this change should also be allowed for in simulations. These results show that changes in leaf physiological characteristics contribute considerably to seasonal variation in CO 2, latent heat, and sensible heat exchanges over plantation trees.

CO 2 exchange in a temperate Japanese cypress forest compared with that in a cool-temperate deciduous broad-leaved forest

To examine the characteristics of carbon exchange in coniferous forests, we analysed the seasonal and diurnal patterns of CO2 exchange, as measured using the eddy covariance method, in a Japanese cypress forest in the Kiryu Experimental Watershed (KEW) in central Japan. The net CO2 exchange data during periods of low-friction velocity conditions and during periods of missing data were interpolated. The daily CO2 uptake was observed throughout the year, with maximum values occurring in early summer. Periods of low carbon uptake were seen in late summer owing to high respiratory CO2 efflux. The diurnal and seasonal patterns of daytime CO2 exchange at KEW were compared with those in a cool-temperate deciduous forest of the Tomakomai Experimental Forest (TOEF) in Japan. The environmental differences between evergreen and deciduous forests affected the seasonal patterns of carbon uptake. Although there were great differences in the mean monthly air temperatures between the sites, the mean monthly daytime carbon uptake was almost equal at both sites during the peak growing period. The carbon-uptake values at the same PAR level were greater before noon than after noon, especially at TOEF, suggesting the stomatal regulation of carbon uptake.

Patterns of root respiration rates and morphological traits in 13 tree species in a tropical forest

The root systems of forest trees are composed of different diameters and heterogeneous physiological traits. However, the pattern of root respiration rates from finer and coarser roots across various tropical species remains unknown. To clarify how respiration is related to the morphological traits of roots, we evaluated specific root respiration and its relationships to mean root diameter (D) of various diameter and root tissue density (RTD; root mass per unit root volume; gcm(-3)) and specific root length (SRL; root length per unit root mass; mg(-1)) of the fine roots among and within 14 trees of 13 species from a primary tropical rainforest in the Pasoh Forest Reserve in Peninsular Malaysia. Coarse root (2-269mm) respiration rates increased with decreasing D, resulting in significant relationships between root respiration and diameter across species. A model based on a radial gradient of respiration rates of coarse roots simulated the exponential decrease in respiration with diameter. The respiration rate of fine roots (<2mm) was much higher and more variable than those of larger diameter roots. For fine roots, the mean respiration rates for each species increased with decreasing D. The respiration rates of fine roots declined markedly with increasing RTD and increased with increasing SRL, which explained a significant portion of the variation in the respiration among the 14 trees from 13 species examined. Our results indicate that coarse root respiration in tree species follows a basic relationship with D across species and that most of the variation in fine root respiration among species is explained by D, RTD and SRL. We found that the relationship between root respiration and morphological traits provides a quantitative basis for separating fine roots from coarse roots and that the pattern holds across different species.

Midday depression of leaf CO2 exchange within the crown of Dipterocarpus sublamellatus in a lowland dipterocarp forest in Peninsular Malaysia

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.

Vertical variation in leaf gas exchange parameters for a Southeast Asian tropical rainforest in Peninsular Malaysia

Vertical variation in leaf gas exchange characteristics of trees grown in a lowland dipterocarp forest in Peninsular Malaysia was investigated. Maximum net photosynthetic rate, stomatal conductance, and electron transport rate of leaves at the upper canopy, lower canopy, and forest floor were studied in situ with saturated condition photosynthetic photon flux density. The dark respiration rate of leaves at the various heights was also studied. Relationships among gas exchange characteristics, and also with nitrogen content per unit leaf area and leaf dry matter per area were clearly detected, forming general equations representing the vertical profile of several important parameters related to gas exchange. Numerical analysis revealed that the vertical distribution of gas exchange parameters was well determined showing both larger carbon gain for the whole canopy and at the same time positive carbon gain for the leaves of the lowest layer. For correct estimation of gas exchange at both leaf and canopy scales using multi-layer models, it is essential to consider the vertical distribution of gas exchange parameters with proper scaling coefficients.

Vertical profiles and storage fluxes of CO2, heat and water in a tropical rainforest at Pasoh, Peninsular Malaysia

Ambient CO2 concentration, air temperature and relative humidity were measured intermittently for a 3-year period from the floor to the canopy top of a tropical rainforest in Pasoh, Peninsular Malaysia. Mean diurnal CO2 storage flux (Sc; μmol m-2 s-1) and sensible and latent heat storage fluxes (Qa and Qw; W m-2) ranged from -12.7 to 3.2 μmol m-2 s-1, -15 to 27 W m-2 and -10 to 20 W m-2, respectively. Small differences in diurnal changes were observed in Sc and Qa between the driest and wettest periods. Compared with the ranges of mean diurnal CO2 eddy flux (-14.7 to 4.9 μmol m-2 s-1), sensible eddy flux (-12 to 169 W m-2) and latent eddy flux (0 to 250 W m-2), the contribution of CO2 storage flux was especially large. Comparison with summertime data from a temperate Japanese cypress forest suggested a higher contribution of Sc in the tropical rainforest, probably mainly due to the difference in nighttime friction velocity at the sites. On the other hand, differences in Qa and Qw were smaller than the difference in Sc, probably because of the smaller nighttime sinks/sources of heat and water vapour.

Determination of the gas exchange phenology in an evergreen coniferous forest from 7 years of eddy covariance flux data using an extended big-leaf analysis

We defined gas exchange phenology as the seasonality of the gas exchange characteristics of a forest canopy, and investigated how the gas exchange phenology could be directly detected from an eddy covariance (EC) dataset and its influence on the canopy fluxes within an evergreen Japanese cypress forest. For the detection of gas exchange phenology, we derived three bulk parameters of the extended big-leaf model (Kosugi et al. 2005) inversely from EC flux data over a 7-year period: surface conductance (gc), maximum rate of carboxylation of the “big leaf” (VCMAX), and intercellular CO2 concentration of the “big leaf” (CI). The relationship between gc and the vapor pressure deficit declined in winter and spring. The relationship between the daily ecosystem respiration and air temperature was greater in the spring than in the other seasons. The temperature dependence curve of VCMAX decreased substantially in the winter and was different from that of an evergreen broadleaved forest. A decrease in CI was occasionally coupled with the decrease in canopy gross primary production during April and August, indicating that stomatal closure was responsible for a decline in canopy photosynthesis. Gas exchange phenology should be quantified when understanding the determining factors of the seasonality of canopy fluxes at evergreen coniferous forests.

New data-driven estimation of terrestrial CO2 fluxes in Asia using a standardized database of eddy covariance measurements, remote sensing data, and support vector regression

The lack of a standardized database of eddy covariance observations has been an obstacle for data-driven estimation of terrestrial CO2 fluxes in Asia. In this study, we developed such a standardized database using 54 sites from various databases by applying consistent postprocessing for data-driven estimation of gross primary productivity (GPP) and net ecosystem CO2 exchange (NEE). Data-driven estimation was conducted by using a machine learning algorithm: support vector regression (SVR), with remote sensing data for 2000 to 2015 period. Site-level evaluation of the estimated CO2 fluxes shows that although performance varies in different vegetation and climate classifications, GPP and NEE at 8days are reproduced (e.g., r2=0.73 and 0.42 for 8day GPP and NEE). Evaluation of spatially estimated GPP with Global Ozone Monitoring Experiment 2 sensor-based Sun-induced chlorophyll fluorescence shows that monthly GPP variations at subcontinental scale were reproduced by SVR (r2=1.00, 0.94, 0.91, and 0.89 for Siberia, East Asia, South Asia, and Southeast Asia, respectively). Evaluation of spatially estimated NEE with net atmosphere-land CO2 fluxes of Greenhouse Gases Observing Satellite (GOSAT) Level 4A product shows that monthly variations of these data were consistent in Siberia and East Asia; meanwhile, inconsistency was found in South Asia and Southeast Asia. Furthermore, differences in the land CO2 fluxes from SVR-NEE and GOSAT Level 4A were partially explained by accounting for the differences in the definition of land CO2 fluxes. These data-driven estimates can provide a new opportunity to assess CO2 fluxes in Asia and evaluate and constrain terrestrial ecosystem models. (Less)

Amplitude and seasonality of storage fluxes for CO2, heat and water vapour in a temperate Japanese cypress forest

Storage fluxes of CO2, air temperature and relative humidity were measured in a temperate Japanese cypress forest for 3 yr. The 3-yr means of diurnal storage fluxes of CO2 (Sc), latent heat (Qa) and sensible heat (Qw) ranged from -1.3 to 1.2 μmol m-2 s-1, -9 to 12 W m-2 and -2 to 5 W m-2, respectively. The mean diurnal eddy fluxes of CO2 (Fc), sensible heat (H) and latent heat (λE) ranged from -13.1 to 2.6 μmol m-2 s-1, -37 to 187 and 7 to 146 W m-2, respectively. The variation and amplitude of diurnal changes in Sc, Qa and Qw were generally similar to those reported for other forests. However, the amplitude of diurnal changes in Sc was much smaller than that of tropical rain forests. The amplitudes of diurnal changes in Sc and Qw in summer were larger than those in winter, although the amplitude of Qa exhibited no clear seasonal variation. This was likely because of the small seasonal change in the source (sensible heat exchange) in comparison with CO2 and latent heat exchange.

Observation of the scale of patchy stomatal behavior in leaves of Quercus crispula using an Imaging-PAM chlorophyll fluorometer

Patchy stomatal closure occurs in plants with heterobaric leaves, in which vertical extensions of bundle sheath cells delimit the mesophyll and restrict the diffusion of CO(2). The scale of patchy stomatal behavior was investigated in this study. The distribution of PSII quantum yield (Φ(II)) obtained from chlorophyll fluorescence images was used to evaluate the scale of stomatal patchiness and its relationship with leaf photosynthesis in the sun leaves of 2-year-old saplings of Quercus crispula Blume. Fluorescent patches were observed only during the day with low stomatal conductance. Comparison of numerical simulation of leaf gas exchange and chlorophyll fluorescence images showed that heterogeneous distribution of electron transport rate through PSII (J) was observed following stomatal closure with a bimodal manner under both natural and saturated photosynthetic photon flux densities. Thus, fluorescence patterns can be interpreted in terms of patchy stomatal closure. The mapping of J from chlorophyll fluorescence images showed that the scale of stomatal patchiness was approximately 2.5-fold larger than that of anatomical patches (lamina areas bounded by bundle sheath extensions within lamina). Our results suggest the spatial scale of stomatal patches in Q. crispula leaves.