Noriyuki Osada

The worldwide leaf economics spectrum

Bringing together leaf trait data spanning 2,548 species and 175 sites we describe, for the first time at global scale, a universal spectrum of leaf economics consisting of key chemical, structural and physiological properties. The spectrum runs from quick to slow return on investments of nutrients and dry mass in leaves, and operates largely independently of growth form, plant functional type or biome. Categories along the spectrum would, in general, describe leaf economic variation at the global scale better than plant functional types, because functional types overlap substantially in their leaf traits. Overall, modulation of leaf traits and trait relationships by climate is surprisingly modest, although some striking and significant patterns can be seen. Reliable quantification of the leaf economics spectrum and its interaction with climate will prove valuable for modelling nutrient fluxes and vegetation boundaries under changing land-use and climate.

The mechanism of general flowering in Dipterocarpaceae in the Malay Peninsula

The mechanism of general flowering in Dipterocarpaceae in the Malay Peninsula is revealed through field survey and meteorological data analyses. The regions of general flowering coincide with those which experienced a low night-time temperature (LNT) c. 2 mo before flowering. This supports the hypo- thesis that low air temperature induces the development of floral buds of diptero- carps. LNT was found to be caused by radiative cooling during dry spells in winter when the northern subtropical ridge (STR) occasionally migrates southwards with a dry air mass into the equatorial region. LNT events usually occur in La Nifia episodes, not in El Nifio episodes as believed previously. This is because the south- ward migration of the STR is associated with the intensification of local meridional Hadley Circulation in the western Pacific, which is strengthened in a La Nifia episode. Results suggest that El Nifio-like climate change in increased atmospheric carbon dioxide concentrations may be critical for the tropical rain forest biome in south-east Asia.

BAAD: a biomass and allometry database for woody plants

Understanding how plants are constructed—i.e., how key size dimensions and the amount of mass invested in different tissues varies among individuals—is essential for modeling plant growth, carbon stocks, and energy fluxes in the terrestrial biosphere. Allocation patterns can differ through ontogeny, but also among coexisting species and among species adapted to different environments. While a variety of models dealing with biomass allocation exist, we lack a synthetic understanding of the underlying processes. This is partly due to the lack of suitable data sets for validating and parameterizing models. To that end, we present the Biomass And Allometry Database (BAAD) for woody plants. The BAAD contains 259 634 measurements collected in 176 different studies, from 21 084 individuals across 678 species. Most of these data come from existing publications. However, raw data were rarely made public at the time of publication. Thus, the BAAD contains data from different studies, transformed into standard units and variable names. The transformations were achieved using a common workflow for all raw data files. Other features that distinguish the BAAD are: (i) measurements were for individual plants rather than stand averages; (ii) individuals spanning a range of sizes were measured; (iii) plants from 0.01–100 m in height were included; and (iv) biomass was estimated directly, i.e., not indirectly via allometric equations (except in very large trees where biomass was estimated from detailed sub-sampling). We included both wild and artificially grown plants. The data set contains the following size metrics: total leaf area; area of stem cross-section including sapwood, heartwood, and bark; height of plant and crown base, crown area, and surface area; and the dry mass of leaf, stem, branches, sapwood, heartwood, bark, coarse roots, and fine root tissues. We also report other properties of individuals (age, leaf size, leaf mass per area, wood density, nitrogen content of leaves and wood), as well as information about the growing environment (location, light, experimental treatment, vegetation type) where available. It is our hope that making these data available will improve our ability to understand plant growth, ecosystem dynamics, and carbon cycling in the worlds vegetation.

Changes in crown development patterns and current-year shoot structure with light environment and tree height in Fagus crenata (Fagaceae)

The relative effects of light and tree height on the architecture of leader crowns (i.e., the leading section of the main trunk, 100 cm in length) and current-year shoots for a canopy species, Fagus crenata, occupying both the ridge top and the valley bottom in a cool-temperate forest in Japan were investigated. For leader crowns, the number of current-year shoots and leaves increased with increasing tree height, whereas the mean length of current-year shoots increased with increasing relative photon flux density (PFD). The leader crown area decreased, and the depth and leaf area index of leader crowns increased, with increasing relative PFD. The mass of current-year shoots increased with relative PFD. However, this total mass was allocated differently between stems and leaves depending on tree height, such that the relative allocation to stems increased with increasing tree height. Furthermore, stem structures within current-year shoots also changed with height, such that taller trees produced thicker and shorter stems of the same volume. In contrast, leaf structure and leaf biomass allocations changed with relative PFD. Specific leaf area decreased with increasing relative PFD. In addition, leaf number increased more rapidly with increasing individual leaf mass for trees exposed to greater relative PFD. Consequently, the total leaf area supported by a stem of a given diameter decreased with increasing tree height and relative PFD. Thus, the architecture of leader crowns and current-year shoots were related differently to light and tree height, which are considered important for efficient light capture and the growth of small and tall trees in different environments.

Community-level flowering phenology and fruit set : Comparative study of 25 woody species in a secondary forest in Japan

We investigated the flowering phenology, pollinator visitation, and fruit set of 25 animal-pollinated woody species in a warm temperate secondary forest in Japan. Various species flowered sequentially from February to October. The principal pollinators were bumblebees, honey-bees, flies and/or beetles and birds; bumblebees and flies/beetles pollinated most trees. The duration of flowering was shorter for species that bloomed in the middle of the season than it was for species that bloomed earlier or later in the season. The timing of flowering was more synchronous within species that had a shorter flowering duration; this was also detected when phylogenetically independent contrasts were calculated. This could be important for the effective pollination of species with a short flowering duration because such species bloom sequentially over a short period of less than 1 month around May. Fruit set was related not to pollinator type, sex expression, flowering sequence (in order of the date of peak flowering) or flowering duration, but to the relative abundance of the species in the forest. This correlation was detected for fly- and beetle-pollinated species but not for bumblebee-pollinated species. Thus, relatively rare plant species with opportunistic pollinators might experience limited fruit set because of insufficient pollinator services. Bagging experiments conducted on eight hermaphrodite species revealed that the fruit set of bagged flowers was nearly zero, lower than that of control flowers. These results indicate the importance of pollinators for successful reproduction and thus for the coexistence of plants in this secondary forest.

Ontogenetic changes in leaf phenology of a canopy species, Elateriospermum tapos (Euphorbiaceae), in a Malaysian rain forest

Leaf phenology was studied in individuals of a canopy species, Elater- iospermum tapos (Euphorbiaceae), at various ontogenetic stages, in a Malaysian rain forest. The timing of leaf emergence was not synchronized among sapling indi- viduals, and was not correlated with any meteorological factors of the preceding month. The timing of leaf fall in saplings was positively correlated with net radi- ation, and maximum and minimum temperature, but negatively correlated with relative humidity the preceding month, although these correlations were weak. The leaf production rate was larger under higher light, but the leaf fall rate was not related to the light regime of the saplings. Thus, leaf production was enhanced by the light availability for each individual, while leaf fall may have been enhanced by drought stress. Non-synchronous leaf production appears to be important for sapling growth, allowing saplings to occupy better-lit space quickly. On the other hand, tall trees showed a clear synchronous leaf-fall pattern, with an annual cycle, and no meteorological factors were correlated with the timing of leaf fall. Mature trees of this species produced flowers simultaneously with new leaves, after shed- ding their leaves. This suggests that the need to synchronize flowering might be the primary determinant of leaf production phenology in mature individuals.

Effects of atmospheric CO2 concentration, irradiance, and soil nitrogen availability on leaf photosynthetic traits of Polygonum sachalinense around natural CO2 springs in northern Japan

Long-term exposure to elevated CO2 concentration will affect the traits of wild plants in association with other environmental factors. We investigated multiple effects of atmospheric CO2 concentration, irradiance, and soil N availability on the leaf photosynthetic traits of a herbaceous species, Polygonum sachalinense, growing around natural CO2 springs in northern Japan. Atmospheric CO2 concentration and its interaction with irradiance and soil N availability affected several leaf traits. Leaf mass per unit area increased and N per mass decreased with increasing CO2 and irradiance. Leaf N per area increased with increasing soil N availability at higher CO2 concentrations. The photosynthetic rate under growth CO2 conditions increased with increasing irradiance and CO2, and with increasing soil N at higher CO2 concentrations. The maximal velocity of ribulose 1,5-bisphosphate carboxylation (Vcmax) was affected by the interaction of CO2 and soil N, suggesting that down-regulation of photosynthesis at elevated CO2 was more evident at lower soil N availability. The ratio of the maximum rate of electron transport to Vcmax (Jmax/Vcmax) increased with increasing CO2, suggesting that the plants used N efficiently for photosynthesis at high CO2 concentrations by changes in N partitioning. To what extent elevated CO2 influenced plant traits depended on other environmental factors. As wild plants are subject to a wide range of light and nutrient availability, our results highlight the importance of these environmental factors when the effects of elevated CO2 on plants are evaluated.

Fruit dispersal of two dipterocarp species in a Malaysian rain forest

Seed dispersal is the predominant mobile stage for sessile plants, and critically affects the distribution patterns of the species (Nathan & Muller-Landau 2000). Thus, seed dispersal patterns are important in understanding the population dynamics of the species. Tropical rain forests of Southeast Asia are characterized by the dominance of the family Dipterocarpaceae. Most dipterocarp species reach the emergent layer of the forest, about 40-60 m in height (Whitmore 1984). The fruits have two to five long wings (elongated sepals) and are dispersed by wind. The fruits germinate without dormancy (Whitmore 1984). Therefore, the distance of primary dispersal by wind is critical for the distribution of most dipterocarp species (Suzuki & Ashton 1996). Wind speed is much higher in the emergent layer than in the canopy and understorey layers in a tropical rain forest (Aoki et al. 1978), and winddispersed tree species are especially abundant in the former (Suzuki & Ashton 1996). This suggests that wind speed and height strongly influence the fruit dispersal of wind-dispersed species. In addition, differences in fruit size

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.

Influence of between-year variation in the density of Rhus trichocarpa fruits on the removal of fruit by birds

The fruiting phenology and fruit removal patterns of Rhus trichocarpa Miq. (Anacardiaceae) were investigated in a warm-temperate secondary forest in Japan. Mature fruits of this species are dispersed by birds. Effects of fruit display size and canopy openness on fruit removal were investigated in years with different fruit densities (i.e., masting and non-masting years). Moreover, effects of increased canopy openness during winter on fruit removal were also investigated. Seasonal patterns of fruit removal were quite different between masting and non- masting years. In the non-mast year, fruits were removed by birds soon after maturation in the summer. In contrast, in the mast year, fruits were removed gradually by birds from summer to winter. Moreover, the rate of fruit removal was greater for trees with a larger display size in the non-mast year, whereas that was greater for trees with greater canopy openness in the mast year. Canopy openness increased in winter, and fruit removal in winter was enhanced in trees with a more open canopy only in the non-mast year. These observations strongly suggest that avian dispersers became satiated in the mast year, whereas fruit removal was enhanced in the non-mast year. In the mast year, although many fruits were not dispersed until winter, they were neither depredated nor rotten, and the long period of fruit removal by birds may have enhanced fruit dispersal. The large abundance of seedlings and saplings of this species in Japanese secondary forests suggests that this fruiting strategy is beneficial.