Tomo’omi Kumagai

Spatial variations in xylem sap flux density in evergreen oak trees with radial-porous wood: Comparisons with anatomical observations

To estimate whole-tree water use when employing sap flow measurements, integration of the sap flux density (Fd) over the sapwood area is needed. Accordingly, it is necessary to obtain information on the characteristics of stem water transportation such as spatial variations in Fd and the active xylem area in the stem cross-section. Although evergreen oak trees with radial-porous wood represent a major component of secondary forests in western Japan, detailed information on their stem water transportation characteristics remains unclear. In the present study, we used the heat dissipation method (Granier method) to conduct measurements of azimuthal and radial variations in the Fd of Quercus glauca Thunb. ex Murray, a representative evergreen broad-leaved tree in western Japan. Further, by analyzing the anatomy of the xylem structure, we examined why Fd varies spatially in the stem cross-section. By using a dye solution injected into a radial hole bored into the tree trunk, we confirmed that the entire stem is hydroactive. We also compared the spatial variations in Fd and water conductivity per xylem area (Ks) which were estimated by using the observed vessel diameters and their density over the stem cross-section and Hagen–Poiseuille’s law. Azimuthal and radial variations in Fd reached about 60 and 50% of the maximum values, respectively, and could be explained by spatial variation in Ks. As a result, we obtained statistical parameters describing the spatial variation in Fd in Q. glauca and determined that whole-tree water use estimated from measurements in one direction had at most ±20% potential errors for studied trees.

Effects of sample size on sap flux-based stand-scale transpiration estimates

In this study, we aimed to assess how sample sizes affect confidence of stand-scale transpiration (E) estimates calculated from sap flux (F(d)) and sapwood area (A(S_tree)) measurements of individual trees. In a Japanese cypress plantation, we measured F(d) and A(S_tree) in all trees (n = 58) within a 20 x 20 m study plot, which was divided into four 10 x 10 subplots. We calculated E from stand A(S_tree) (A(S_stand)) and mean stand F(d) (J(S)) values. Using Monte Carlo analyses, we examined the potential errors associated with sample sizes in E, A(S_stand) and J(S) using the original A(S_tree) and F(d) data sets. Consequently, we defined the optimal sample sizes of 10 and 15 for A(S_stand) and J(S) estimates, respectively, in the 20 x 20 m plot. Sample sizes larger than the optimal sample sizes did not decrease potential errors. The optimal sample sizes for J(S) changed according to plot size (e.g., 10 x 10 and 10 x 20 m), whereas the optimal sample sizes for A(S_stand) did not. As well, the optimal sample sizes for J(S) did not change in different vapor pressure deficit conditions. In terms of E estimates, these results suggest that the tree-to-tree variations in F(d) vary among different plots, and that plot size to capture tree-to-tree variations in F(d) is an important factor. The sample sizes determined in this study will be helpful for planning the balanced sampling designs to extrapolate stand-scale estimates to catchment-scale estimates.

Modeling CO2 exchange over a Bornean tropical rain forest using measured vertical and horizontal variations in leaf‐level physiological parameters and leaf area densities

Southeast Asian tropical rain forests are among the worlds most important biomes in terms of global carbon cycling; nevertheless, the impact of environmental factors on the ecosystem CO 2 flux remains poorly understood. One-dimensional multilayer biosphere-atmosphere models such as soil-vegetation-atmosphere transfer (SVAT) models are promising tools for understanding how interactions between environmental factors and leaf-level physiological parameters might impact canopy-level CO 2 exchange. To examine application of the SVAT model in tropical rain forests, which is expected to be difficult partly because of the complex canopy structure and large number of tree species, we measured vertical and horizontal variations in leaf-level physiological parameters and leaf area densities together with eddy covariance measurements using a canopy crane in a tropical rain forest in Sarawak, Malaysia. Despite differences in species and canopy positions, leaf nitrogen per unit area (N a ) within the canopy could be one-dimensionally described as a linear function of height. N a also clearly explained the other leaf-level physiological parameters across species and canopy positions. Even though the leaf area density profile likely varies in this tropical forest, the SVAT model satisfactorily reproduced the eddy covariance measurements. Furthermore, the CO 2 flux calculated on the assumption that N a measured in the upper canopy was distributed evenly throughout was almost the same as that taking the vertical gradient into consideration. These findings suggest that when reproducing the CO 2 flux in tropical rain forests using the SVAT model, the leaf area density profile obtained from the leaf area index (LAI) measured at one point and leaf-level physiological properties measured across species in the upper canopy are sufficient.

Modeling water transportation and storage in sapwood — model development and validation

I have described a single tree water flux model that combines the stomatal conductance model based on plant physiological knowledge with the concept of fluid mechanics used to describe water transportation and storage in sapwood. The model was parameterized and validated using field data of the water budget of Cryptomeria japonica [J. Jpn. For. Soc. 78 (1996) 66] determined by weighing a suspended cut tree. In their experiment, water transportation and storage in the sapwood and leaves and the sapwood potential were examined under ordinary conditions and extreme conditions where water uptake was stopped by sealing the cut stem surface. Numerical experiments show the information to reproduce the observed results: low intensity of stomatal response to decreasing leaf water potential and lower hydraulic conductance of the crown sapwood than the conductance of the trunk sapwood. The model reproduced the observed results under both ordinary and extreme conditions using only information of observed physical size and hydraulic properties, except for the information on low values of hydraulic conductance of the crown sapwood.

Strategies of a Bornean tropical rainforest water use as a function of rainfall regime: isohydric or anisohydric?

Although Bornean tropical rainforests are among the moistest biomes in the world, they sporadically experience periods of water stress. The observations indicate that these ecosystems tend to have little regulation of water use, despite episodes of relatively severe drought. This water-use behaviour is often referred to as anisohydric behaviour, as opposed to isohydric plants that regulate stomatal movement to prevent hydraulic failure. Although it is generally thought that anisohydric behaviour is an adaptation to more drought-prone habitats, we show that anisohydric plants may also be more favoured than isohydric plants under very moist environments where there is little risk of hydraulic failure. To explore this subject, we examined the advantages of isohydric and anisohydric species as a function of the hydroclimatic environment using a stochastic model of soil moisture and carbon assimilation dynamics parameterized by field observations. The results showed that under very moist conditions, anisohydric species tend to have higher productivity than isohydric plants, despite the fact that the two plant types show almost the same drought-induced mortality. As precipitation decreases, the mortality of anisohydric plants drastically increases whereas that of isohydric plants remains relatively constant and low; in these conditions, isohydric plants surpass anisohydric plants in their productivity.

Vertical Profiles of Environmental Factors within Tropical Rainforest, Lambir Hills National Park, Sarawak, Malaysia

Environmental factors, such as global solar radiation, wind speed, air temperature, humidity, and CO2 concentration, were measured above and within the canopy of a tropical rainforest in Lambir Hills National Park, Sarawak, Malaysia. Few data concerning the environment of this forest have been reported. Intensive observations were carried out in 1998, 1999, and 2000 with the following results: (1) The fraction of global solar radiation reaching the upper layer of the canopy varied with global solar radiation above the canopy. Even though the global solar radiation above the canopy fluctuated, the fraction of that reaching the lower canopy and the ground was constantly approximately 5%. (2) The fraction of wind speed reaching each layer of the canopy increased with wind speed above the canopy. Little wind was usually present at the lower canopy. (3) The daytime air temperature at the canopy top was higher than that near the ground. The maximum difference between the air temperature at the canopy top and that at the ground was about 5°C, and the diurnal temperature ranges at the canopy top and those at the ground were about 8°C and about 5°C, respectively. The highest daytime water vapor pressure occurred within the canopy and particularly near the ground. Vertical gradients of water vapor pressure during the day were steep, probably because of high transpiration. (4) In the 1998 observation the minimum and the maximum CO2 concentrations were 360 ppm in the day and 450 ppm at night, while in the 2000 observation the minimum and the maximum CO2 concentrations were 350 ppm in the day and 540 ppm at night. The higher CO2 concentration in the daytime and the lower concentration at night observed during the 1998 observation period were probably due to reduced photosynthesis and soil respiration caused by exceptional dry conditions during the observation period.

Azimuthal variations of sap flux density within Japanese cypress xylem trunks and their effects on tree transpiration estimates

Sap flow techniques are practical tools for estimating tree transpiration. Though many previous studies using sap flow techniques did not consider azimuthal variations of sap flux density (Fd) on xylem trunk to estimate tree transpiration, a few studies reported that ignoring the azimuthal variations in Fd could cause large errors in tree transpiration estimates for some tree species. Therefore, examining azimuthal variations in Fd for major plantation tree species is critical for estimating tree transpiration. Using the thermal dissipation method, we examined azimuthal variations in Fd in six trees of Japanese cypress Chamaecyparis obtusa (Sieb. et Zucc.) Endl., which is one of the most common plantation tree species in Japan. We recorded considerable variations among Fd at four different azimuthal directions. The Fd value for one aspect was more than 100% larger than those for the other aspects. We calculated differences between tree transpiration estimates based on Fd for one to three azimuthal directions and those based on Fd for four aspects. The differences relative to tree transpiration estimates based on Fd for four aspects were typically 30, 20, and 10% in accordance with the Fd for one, two, and three measurement aspects, respectively. This finding indicates that ignoring azimuthal variations could cause large errors in tree transpiration estimates for Japanese cypress.

Differences in transpiration characteristics of Japanese beech trees, Fagus crenata, in Japan

Japanese beech (Fagus crenata Blume) is widely distributed across the Japan archipelago. This species requires morphological and physiological plasticity to cope with the diverse environmental conditions across its geographical range. In this study, we monitored transpiration (E) to examine plasticity mechanisms as an example of geographical variation in whole-tree water use. We determined E by measuring the sap flux of Japanese beech trees in three stands: Kuromatsunai (KR), Kawatabi (KW) and Shiiba (SH), which were located in different areas in Japan. We conducted biometric measurements to characterize leaf and crown morphology and evaluated geographical variations in E characteristics, such as canopy aerodynamic conductance, canopy stomatal conductance (G(S)) and decoupling coefficient (Omega). Leaf morphology and crown shape showed clear geographical clines. Individual leaf areas decreased in the order KR > KW > SH. The crown shape in the KR and KW stands was cylindrical but planar in the SH stand. We evaluated the effects of leaf and crown morphology on E characteristics. The Omega values showed that, while E in the KW and SH stands was highly sensitive to G(S) and atmospheric evaporative demand, E in the KR stand was sensitive to radiative energy. To maximize carbon gain without further water loss, trees maintain a high G(S) in a moist habitat. For example, the KR trees may decrease E by reducing their absorbed radiation energy by adjusting the individual leaf size and crown structure. Our results indicate that the geographical variation in the water use pattern of Japanese beech is determined by the interaction between its physiological and morphological status.

Evapotranspiration of rubber (Hevea brasiliensis) cultivated at two plantation sites in Southeast Asia

To investigate the effects of expanding rubber (Hevea brasiliensis) cultivation on water cycling in Mainland Southeast Asia (MSEA), evapotranspiration (ET) was measured within rubber plantations at Bueng Kan, Thailand, and Kampong Cham, Cambodia. After energy closure adjustment, mean annual rubber ET was 1211 and 1459 mm yr(-1) at the Thailand and Cambodia sites, respectively, higher than that of other tree-dominated land covers in the region, including tropical seasonal forest (812-1140 mm yr(-1)), and savanna (538-1060 mm yr(-1)). The mean proportion of net radiation used for ET by rubber (0.725) is similar to that of tropical rainforest (0.729) and much higher than that of tropical seasonal forest (0.595) and savanna (0.548). Plant area index (varies with leaf area changes), explains 88.2% and 73.1% of the variance in the ratio of latent energy flux (energy equivalent of ET) to potential latent energy flux (LE/LEpot) for midday rain-free periods at the Thailand and Cambodia sites, respectively. High annual rubber ET results from high late dry season water use, associated with rapid refoliation by this brevideciduous species, facilitated by tapping of deep soil water, and by very high wet season ET, a characteristic of deciduous trees. Spatially, mean annual rubber ET increases strongly with increasing net radiation (R-n) across the three available rubber plantation observation sites, unlike nonrubber tropical ecosystems, which reduce canopy conductance at high R-n sites. High water use by rubber raises concerns about potential effects of continued expansion of tree plantations on water and food security in MSEA.

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.