Atsushi Ishida

Leaf nitrogen distribution in relation to crown architecture in the tall canopy species, Fagus crenata

The theory of optimal leaf N distribution predicts that the C gain of plants is maximized when the N content per unit area (Narea) scales with light availability, but most previous studies have demonstrated that the N distribution is not proportional to light availability. In tall trees, the leaves are often clustered on twigs (leaf cluster) and not evenly distributed within the crowns. Thus, we hypothesized that the suboptimal N distribution is partly caused by the limited capacity to translocate N between leaf clusters, and consequently, the relationship between light and Narea differs for leaves in different clusters. We investigated the light availability and N content of all individual leaves within several leaf clusters on tall trees of a deciduous canopy species Fagus crenata in Japan. We observed that the within-cluster leaf N distribution patterns differed from the between-cluster patterns and the slopes of the relationships between light and Narea were lower within clusters than between clusters. According to the detailed analysis of the N distribution within leaf clusters, Narea was greater for current-year shoots with greater light availability or a larger total leaf area. The latter pattern was probably caused by the greater sink strength of the current-year shoots with a larger leaf area. These N distribution patterns suggest that leaf clusters are fairly independent with respect to their N use, and the productivity of real F. crenata crowns may be less than optimal.

Recovery of Physiological Traits in Saplings of Invasive Bischofia Tree Compared with Three Species Native to the Bonin Islands under Successive Drought and Irrigation Cycles.

Partial leaf shedding induced by hydraulic failure under prolonged drought can prevent excess water consumption, resulting in delayed recovery of carbon productivity following rainfall. To understand the manner of water use of invasive species in oceanic island forests under a fluctuating water regime, leaf shedding, multiple physiological traits, and the progress of embolism in the stem xylem under repeated drought-irrigation cycles were examined in the potted saplings of an invasive species, Bischofia javanica Blume, and three endemic native species, Schima mertensiana (Sieb. Et Zucc,) Koitz., Hibiscus glaber Matsum, and Distylium lepidotum Nakai, from the Bonin Islands, Japan. The progress of xylem embolism was observed by cryo-scanning electron microscopy. The samples exhibited different processes of water saving and drought tolerance based on the different combinations of partial leaf shedding involved in embolized conduits following repeated de-rehydration. Predawn leaf water potential largely decreased with each successive drought-irrigation cycle for all tree species, except for B. javanica. B. javanica shed leaves conspicuously under drought and showed responsive stomatal conductance to VPD, which contributed to recover leaf gas exchange in the remaining leaves, following a restored water supply. In contrast, native tree species did not completely recover photosynthetic rates during the repeated drought-irrigation cycles. H. glaber and D. lepidotum preserved water in vessels and adjusted leaf osmotic rates but did not actively shed leaves. S. mertensiana exhibited partial leaf shedding during the first cycle with an osmotic adjustment, but they showed less responsive stomatal conductance to VPD. Our data indicate that invasive B. javanica saplings can effectively use water supplied suddenly under drought conditions. We predict that fluctuating precipitation in the future may change tree distributions even in mesic or moist sites in the Bonin Islands.

The dynamics of carbon stored in xylem sapwood to drought-induced hydraulic stress in mature trees

Climate-induced forest die-off is widespread in multiple biomes, strongly affecting the species composition, function and primary production in forest ecosystems. Hydraulic failure and carbon starvation in xylem sapwood are major hypotheses to explain drought-induced tree mortality. Because it is difficult to obtain enough field observations on drought-induced mortality in adult trees, the current understanding of the physiological mechanisms for tree die-offs is still controversial. However, the simultaneous examination of water and carbon uses throughout dehydration and rehydration processes in adult trees will contribute to clarify the roles of hydraulic failure and carbon starvation in tree wilting. Here we show the processes of the percent loss of hydraulic conductivity (PLC) and the content of nonstructural carbohydrates (NSCs) of distal branches in woody plants with contrasting water use strategy. Starch was converted to soluble sugar during PLC progression under drought, and the hydraulic conductivity recovered following water supply. The conversion of NSCs is strongly associated with PLC variations during dehydration and rehydration processes, indicating that stored carbon contributes to tree survival under drought; further carbon starvation can advance hydraulic failure. We predict that even slow-progressing drought degrades forest ecosystems via carbon starvation, causing more frequent catastrophic forest die-offs than the present projection.

Responses of gas-exchange rates and water relations to annual fluctuations of weather in three species of urban street trees

The frequency of extreme weather has been rising in recent years. A 3-year study of street trees was undertaken in Tokyo to determine whether: (i) street trees suffer from severe water stress in unusually hot summer; (ii) species respond differently to such climatic fluctuations; and (iii) street trees are also affected by nitrogen (N) deficiency, photoinhibition and aerosol pollution. During the study period (2010-12), midsummers of 2010 and 2012 were unusually hot (2.4-2.8u2005°C higher maximum temperature than the long-term mean) and dry (6-56% precipitation of the mean). In all species, street trees exhibited substantially decreased photosynthetic rate in the extremely hot summer in 2012 compared with the average summer in 2011. However, because of a more conservative stomatal regulation (stomatal closure at higher leaf water potential) in the hot summer, apparent symptoms of hydraulic failure were not observed in street trees even in 2012. Compared with Prunusu2005×u2005yedoensis and Zelkova serrata, Ginkgo biloba, a gymnosperm, was high in stomatal conductance and midday leaf water potential even under street conditions in the unusually hot summer, suggesting that the species had higher drought resistance than the other species and was less susceptible to urban street conditions. This lower susceptibility might be ascribed to the combination of higher soil-to-leaf hydraulic conductance and more conservative water use. Aside from meteorological conditions, N deficiency affected street trees significantly, whereas photoinhibition and aerosol pollution had little effect. The internal CO2 and δ(13)C suggested that both water and N limited the net photosynthetic rate of street trees simultaneously, but water was more limiting. From these results, we concluded that the potential risk of hydraulic failure caused by climatic extremes could be low in urban street trees in temperate regions. However, the size of the safety margin might be different between species.

Anthropogenic effects on a tropical forest according to the distance from human settlements.

The protection of tropical forests is one of the most urgent issues in conservation biology because of the rapid deforestation that has occurred over the last 50 years. Even in protected forests, the anthropogenic effects from newly expanding villages such as harvesting of medicinal plants, pasturing cattle and forest fires can induce environmental modifications, especially on the forest floor. We evaluated the anthropogenic effects of the daily activities of neighboring residents on natural forests in 12 plots extending from the village boundary into a natural forest in Thailand. The basal area per unit land area did not present a significant trend; however, the species diversity of woody plants decreased linearly towards the village boundary, which caused a loss of individual density because of severe declines in small saplings compared with adult trees and large saplings in proximity to the village. An analysis of tree-size categories indicates a lack of small samplings near the village boundary. The current forest appears to be well protected based on the adult tree canopy, but regeneration of the present-day forests is unlikely because of the loss of seedlings.

Physiological mechanisms of drought-induced tree die-off in relation to carbon, hydraulic and respiratory stress in a drought-tolerant woody plant

Drought-induced tree die-off related to climate change is occurring worldwide and affects the carbon stocks and biodiversity in forest ecosystems. Hydraulic failure and carbon starvation are two commonly proposed mechanisms for drought-induced tree die-off. Here, we show that inhibited branchlet respiration and soil-to-leaf hydraulic conductance, likely caused by cell damage, occur prior to hydraulic failure (xylem embolism) and carbon starvation (exhaustion of stored carbon in sapwood) in a drought-tolerant woody species, Rhaphiolepis wrightiana Maxim. The ratio of the total leaf area to the twig sap area was used as a health indicator after drought damage. Six adult trees with different levels of tree health and one dead adult tree were selected. Two individuals having the worst and second worst health among the six live trees died three months after our study was conducted. Soil-to-leaf hydraulic conductance and leaf gas exchange rates decreased linearly as tree health declined, whereas xylem cavitation and total non-structural carbon remained unchanged in the branchlets except in the dead and most unhealthy trees. Respiration rates and the number of living cells in the sapwood decreased linearly as tree health declined. This study is the first report on the importance of dehydration tolerance and respiration maintenance in living cells.

Overwintering evergreen oaks reverse typical relationships between leaf traits in a species spectrum

The leaf economics spectrum has given us a fundamental understanding of the species variations in leaf variables. Across plant species, tight correlations among leaf mass per area (LMA), mass-based nitrogen (Nm) and photosynthetic rate (Am) and leaf lifespan have been well known as trade-offs in leaf carbon economy. However, the regional or biome-level correlations may not be necessary to correspond with the global-scale analysis. Here, we show that almost all leaf variables in overwintering evergreen oaks in Japan were relatively well included within the evergreen-broadleaved trees in worldwide temperate forests, but Nm was more consistent with that in deciduous broadleaved trees. Contrary to the universal correlations, the correlation between Am and Nm among the evergreen oaks was negative and the correlation between Am and LMA disappeared. The unique performance was due to specific nitrogen allocation within leaves, i.e. the evergreen oaks with later leaf maturation had lower Nm but higher nitrogen allocation to photosynthetic enzymes within leaves, to enhance carbon gain against the delayed leaf maturation and the shortened photosynthetic period due to cold winters. Our data demonstrate that correlations between leaf variables in a local scale are occasionally different from averaged global-scale datasets, because of the constraints in each biome.

Microhabitat locality allows multi-species coexistence in terrestrial plant communities

Most terrestrial plant communities exhibit relatively high species diversity and many competitive species are ubiquitous. Many theoretical studies have been carried out to investigate the coexistence of a few competitive species and in most cases they suggest competitive exclusion. Theoretical studies have revealed that coexistence of even three or four species can be extremely difficult. It has been suggested that the coexistence of many species has been achieved by the fine differences in suitable microhabitats for each species, attributing to niche-separation. So far there is no explicit demonstration of such a coexistence in mathematical and simulation studies. Here we built a simple lattice Lotka-Volterra model of competition by incorporating the minute differences of suitable microhabitats for many species. By applying the site variations in species-specific settlement rates of a seedling, we achieved the coexistence of more than 10 species. This result indicates that competition between many species is avoided by the spatial variations in species-specific microhabitats. Our results demonstrate that coexistence of many species becomes possible by the minute differences in microhabitats. This mechanism should be applicable to many vegetation types, such as temperate forests and grasslands.

Effective use of high CO₂ efflux at the soil surface in a tropical understory plant.

Many terrestrial plants are C3 plants that evolved in the Mesozoic Era when atmospheric CO2 concentrations ([CO2]) were high. Given current conditions, C3 plants can no longer benefit from high ambient [CO2]. Kaempferia marginata Carey is a unique understory ginger plant in the tropical dry forests of Thailand. The plant has two large flat leaves that spread on the soil surface. We found a large difference in [CO2] between the partly closed space between the soil surface and the leaves (638u2005µmol mol−1) and the atmosphere at 20u2005cm above ground level (412u2005µmol mol−1). This finding indicates that the plants capture CO2 efflux from the soil. Almost all of the stomata are located on the abaxial leaf surface. When ambient air [CO2] was experimentally increased from 400 to 600 μmol mol−1, net photosynthetic rates increased by 45 to 48% under near light-saturated conditions. No significant increase was observed under low light conditions. These data demonstrate that the unique leaf structure enhances carbon gain by trapping soil CO2 efflux at stomatal sites under relatively high light conditions, suggesting that ambient air [CO2] can serve as an important selective agent for terrestrial C3 plants.