Vertical and horizontal light heterogeneity along gradients of secondary succession in cool- and warm-temperate forests.

Aims: Light availability varies drastically in forests, both vertically and horizontally. Vertical light heterogeneity (i.e., patterns of light attenuation from the forest canopy to the floor) may be related to light competition among trees, while horizontal light heterogeneity (i.e., variations in light intensity at a given height within forests) may be associated with light-niche partitioning among tree species. However, light heterogeneity in vertical and horizontal directions and their associations with forest structure are rarely studied to date. Here we report the first comprehensive study to compare the vertical and horizontal light heterogeneity in differently aged forests in two forest types. Location: Twelve forest stands of different ages in cool-temperate forests (consisting of deciduous broad-leaved trees) and five of different ages in warm-temperate forests (evergreen conifer and deciduous broad-leaved trees) in Japan. Methods: We measured vertical light profiles at 1-m intervals from the understorey (1 m above the ground) to the top canopy (12-22 m depending on stands) at 16 locations for each stand (20 m × 20 m). We also measured structural parameters (diameter at breast height, height, and crown dimensions) for all major trees in these stands. Results: Along the secondary successional gradients, the vertical and horizontal light heterogeneity changed in a systematic manner in both forests. The vertical light attenuation rate was steeper in early succession and more gradual in late succession, and the horizontal light heterogeneity was relatively small in early succession and more pronounced in late succession. The vertical light attenuation rate was different between the two forest types; the light intensity dropped more sharply from the canopy surface in the cool-temperate forests due to the crown being vertically shorter and denser (i.e., higher leaf density per unit volume). Conclusion: In early succession, a steeper light attenuation rate is likely related to the strong light competition among co-occurring trees and thus a self-thinning process. In late succession, the high spatial light heterogeneity in forests (i.e., larger horizontal light heterogeneity and gradual light attenuation rate) may allow more species to partition light, and thus may enhance species coexistence and diversity.

Root climbers may alter invertebrate communities on tree trunks: an indirect effect of lianas.

Root climbers are a group of lianas which typically climb the host tree using adhesive roots and do not compete with the host for light. They are relatively more abundant in high-latitude forests and were recently documented to have no harmful effect on their host trees. Although previous studies have examined the direct negative effects of lianas on their host trees, little is known about potential indirect effects via effects on the animal and plant communities. Here, we aimed to determine the effects of root climbers on the animal and plant community composition of their host trunk and whether these alterations indirectly affect host trees. To answer these questions, we compared the invertebrate community, herbaceous plant presence, moss cover and herbivory rates in more than 22 pairs of host and control trees in cool-temperate forests in Hokkaido, Japan. We selected Quercus crispula Blume and Hydrangea petiolaris Sieb. et Zucc. as the study species. Generalised linear mixed models (GLMMs) were used to determine the influence of lianas on tree trunk communities and host trees. Our results showed that the presence of root climbers increased the abundance of Araneae and herbaceous plants and reduced the abundance of Formicidae on host trees, implying a reduction of herbivory on hosts. These results imply that liana presence may reduce the herbivory rate on the host tree, potentially by altering the trunk animal and plant community structure. Our results have implications for the understanding of the effects of lianas on host trees and their function in high-latitude forests.

Genetic differentiation in the timing of budburst in Fagus crenata in relation to temperature and photoperiod.

Climate change is expected to influence plant productivity particularly through changes in the timing of budburst. Nonetheless, knowledge about the intraspecific variation of the timing of budburst and its relationship with climate is insufficient for most tree species. Based on the common garden experiments of Fagus crenata, we investigated the interrelationships between the day of budburst, cumulative degree-days (temperature sum), chilling duration, and photoperiod at the timing of budburst for the trees of different combinations of 11 sites of seed origin and seven experimental sites in Japan. We found that the relationship between the latitude of experimental sites and the timing of budburst differed for the trees of different latitudes of origins. The timing of budburst was earlier for the trees of more northern populations throughout the latitudes of experimental sites. Variation in the timing of budburst among the trees of different seed origins was smaller for more northern experimental sites. Such patterns were caused by directional changes in the relationships between temperature sum, chilling duration, and photoperiod among the trees of different origins: the asymptotes of the curvilinear relationship between chilling duration and temperature sum, chilling duration and photoperiod, and temperature sum and photoperiod, decreased for more northern populations. With the northward expansion of species distribution, the responses of budburst to climate probably changed genetically in such ways in this species. Our results suggest that intraspecific variations in the relationships between the timing of budburst and associated meteorological factors inevitably influence the overall pattern of the timing of budburst at the geographic scale, and the timing of budburst might deviate from predictions when intraspecific variations are not considered.

How is light interception efficiency related to shoot structure in tall canopy species?

Coexistence of multiple species is a fundamental aspect of plant and forest ecology. Although spatial arrangement of leaves within crowns is an important determinant of light interception and productivity, shoot structure varies considerably among coexisting canopy species. We investigated the relative importance of structural traits in determining the light availability of leaves (I) and light interception efficiency at the current-year shoot level (LIECS; the total light interception of leaves divided by shoot biomass) at the top of crowns of 11 canopy species in a cool-temperate forest in Japan. In accordance with Corner's rules, the total mass, stem mass, total mass of leaf laminae, individual leaf area, and stem cross-sectional area of current-year shoot were positively correlated with each other, and branching intensity (the number of current-year shoots per branch unit of 1-m length) was inversely correlated with these traits across species. In contrast, I was correlated not with these traits, but with leaf elevation angle (a L). Moreover, variation in LIECS across species was caused by variation in I (thus in a L). Thus, a L is a key parameter for the leaf light interception of canopy shoots in this cool-temperate forest. Differences in a L across species might be related to different physiological strategies that developed in the high light and water-limited environment of forest canopies. Small variation in the length of current-year shoots among species implies that variations in I and LIECS would be important for the coexistence of these canopy species.

Efficacy of generic allometric equations for estimating biomass: a test in Japanese natural forests.

Accurate estimation of tree and forest biomass is key to evaluating forest ecosystem functions and the global carbon cycle. Allometric equations that estimate tree biomass from a set of predictors, such as stem diameter and tree height, are commonly used. Most allometric equations are site specific, usually developed from a small number of trees harvested in a small area, and are either species specific or ignore interspecific differences in allometry. Due to lack of site-specific allometries, local equations are often applied to sites for which they were not originally developed (foreign sites), sometimes leading to large errors in biomass estimates. In this study, we developed generic allometric equations for aboveground biomass and component (stem, branch, leaf, and root) biomass using large, compiled data sets of 1203 harvested trees belonging to 102 species (60 deciduous angiosperm, 32 evergreen angiosperm, and 10 evergreen gymnosperm species) from 70 boreal, temperate, and subtropical natural forests in Japan. The best generic equations provided better biomass estimates than did local equations that were applied to foreign sites. The best generic equations included explanatory variables that represent interspecific differences in allometry in addition to stem diameter, reducing error by 4-12% compared to the generic equations that did not include the interspecific difference. Different explanatory variables were selected for different components. For aboveground and stem biomass, the best generic equations had species-specific wood specific gravity as an explanatory variable. For branch, leaf, and root biomass, the best equations had functional types (deciduous angiosperm, evergreen angiosperm, and evergreen gymnosperm) instead of functional traits (wood specific gravity or leaf mass per area), suggesting importance of other traits in addition to these traits, such as canopy and root architecture. Inclusion of tree height in addition to stem diameter improved the performance of the generic equation only for stem biomass and had no apparent effect on aboveground, branch, leaf, and root biomass at the site level. The development of a generic allometric equation taking account of interspecific differences is an effective approach for accurately estimating aboveground and component biomass in boreal, temperate, and subtropical natural forests.

Geographic variation in shoot traits and branching intensity in relation to leaf size in Fagus crenata: A common garden experiment.

PREMISE OF THE STUDY: Differences in leaf size are expected to be coordinated with various shoot traits and branching intensity because these relationships will influence light capture efficiency, water use, and biomechanics. Previous studies have mainly focused on interspecific patterns of these trait relationships, but not on intraspecific patterns at the geographic scale. We investigated intraspecific variation in shoot traits and branching intensity of Fagus crenata in Japan. METHODS: Allometric relationships between the traits of current-year shoots and branching intensity per branch unit of 1-m length on the main axis (BI) and its coordination with latitude were investigated using trees from 10 provenances in a common garden. KEY RESULTS: Individual trees originating from lower latitudes have smaller leaves with greater leaf mass per area and nitrogen content per area, greater Huber value (stem cross-sectional area per total leaf area [ATL]) of current-year shoots, and greater BI. Notably, the slope of the log-log relationship between BI and ATL was close to -1.0 across the trees from different source sites, implying that branching in this species occurs to control leaf area. CONCLUSIONS: Shoot traits and branching intensity were apparently coordinated with leaf size to control leaf area deployment in this species. Such patterns probably reflect differences in competition for hydraulic conductance among nearby shoots within crowns, as a consequence of different meteorological conditions across the source sites.

Nitrogen as a key regulator of flowering in Fagus crenata: understanding the physiological mechanism of masting by gene expression analysis.

The role of resource availability in determining the incidence of masting has been widely studied, but how floral transition and initiation are regulated by the resource level is unclear. We tested the hypothesis that floral transition is stimulated by high resource availabiltiy in Fagus crenata based on a new technique, the expression analyses of flowering genes. We isolated F. crenata orthologues of FLOWERING LOCUS T, LEAFY and APETALA1, and confirmed their functions using transgenic Arabidopsis thaliana. We monitored the gene expression levels for 5 years and detected a cycle of on and off years, which was correlated with fluctuations of the shoot-nitrogen concentration. Nitrogen fertilisation resulted in the significantly higher expression of flowering genes than the control, where all of the fertilised trees flowered, whereas the control did not. Our findings identified nitrogen as a key regulator of mast flowering, thereby providing new empirical evidence to support the resource budget model.

Brown bear digging decreases tree growth: Implication for ecological role of top predators in anthropogenic landscapes.

Large carnivores have recently increased in number and recolonized in human-dominated landscapes; however, their ecological roles in these landscapes have not been well studied. In the Shiretoko World Heritage (SWH) site, brown bears have recolonized a previously abandoned mosaic landscape of natural forests and conifer plantations after land abandonment. We previously reported that the bears had recently begun to dig for cicada nymphs in association with the creation of larch plantations. As a result, this digging activity decreased soil nutrients. To deepen the understanding of the novel ecological role of brown bears within human-modified landscapes, we examined the impacts of brown bear digging on the growth of larch trees. We found that brown bear digging decreased fine root biomass of larch, soil water, and nitrogen availability. Brown bear digging negatively affected needle nitrogen content, but not carbon isotope ratios, a water stress index of trees. Tree ring data suggest that digging negatively affected the radial growth of larches. The results imply that digging decreases tree growth due to limited soil nitrogen uptake. Our findings indicate that the ecological roles of large carnivores may differ between natural and anthropogenic landscapes.

Robustness of trait distribution metrics for community assembly studies under the uncertainties of assembly processes.

Numerous studies have revealed the existence of nonrandom trait distribution patterns as a sign of environmental filtering and/or biotic interactions in a community assembly process. A number of metrics with various algorithms have been used to detect these patterns without any clear guidelines. Although some studies have compared their statistical powers, the differences in performance among the metrics under the conditions close to actual studies are not clear. Therefore, the performances of five metrics of convergence and 16 metrics of divergence under alternative conditions were comparatively analyzed using a suite of simulated communities. We focused particularly on the robustness of the performances to conditions that are often uncertain and uncontrollable in actual studies; e.g., atypical trait distribution patterns stemming from the operation of multiple assembly mechanisms, a scaling of trait-function relationships, and a sufficiency of analyzed traits. Most tested metrics, for either convergence or divergence, had sufficient statistical power to distinguish nonrandom trait distribution patterns without uncertainty. However, the performances of the metrics were considerably influenced by both atypical trait distribution patterns and other uncertainties. Influences from these uncertainties varied among the metrics of different algorithms and their performances were often complementary. Therefore, under the uncertainties of an assembly process, the selection of appropriate metrics and the combined use of complementary metrics are critically important to reliably distinguish nonrandom patterns in a trait distribution. We provide a tentative list of recommended metrics for future studies.

Contribution of tree community structure to forest productivity across a thermal gradient in eastern Asia.

Despite their fundamental importance the links between forest productivity, diversity and climate remain contentious. We consider whether variation in productivity across climates reflects adjustment among tree species and individuals, or changes in tree community structure. We analysed data from 60 plots of humid old-growth forests spanning mean annual temperatures (MAT) from 2.0 to 26.6 °C. Comparing forests at equivalent aboveground biomass (160 Mg C ha-1), tropical forests ≥24 °C MAT averaged more than double the aboveground woody productivity of forests <12 °C (3.7 ± 0.3 versus 1.6 ± 0.1 Mg C ha-1 yr-1). Nonetheless, species with similar standing biomass and maximum stature had similar productivity across plots regardless of temperature. We find that differences in the relative contribution of smaller- and larger-biomass species explained 86% of the observed productivity differences. Species-rich tropical forests are more productive than other forests due to the high relative productivity of many short-stature, small-biomass species.

Effective distance of volatile cues for plant-plant communication in beech.

In response to volatiles emitted from a plant infested by herbivorous arthropods, neighboring undamaged conspecific plants become better defended against herbivores; this is referred to as plant‒plant communication. Although plant‒plant communication occurs in a wide range of plant species, most studies have focused on herbaceous plants. Here, we investigated plant‒plant communication in beech trees in two experimental plantations in 2018 and one plantation in 2019. Approximately 20% of the leaves of a beech tree were clipped in half in the spring seasons of 2018 and 2019 (clipped tree). The damage levels to leaves in the surrounding undamaged beech trees were evaluated 90 days after the clipping (assay trees). In both years, the damage levels decreased with a reduction in the distance from the clipped tree. In 2019, we also recorded the damage levels of trees that were not exposed to volatiles (nonexposed trees) as control trees and found that those that were located <5 m away from clipped trees had significantly less leaf damage than nonexposed trees. By using a gas chromatograph-mass spectrometer, ten and eight volatile compounds were detected in the headspaces of clipped and unclipped leaves, respectively. Among them, the amount of (Z)-3-hexenyl acetate in clipped leaves was significantly higher than that in nonclipped leaves. Our result suggests that green leaf volatiles such as (Z)-3-hexenol and (Z)-3-hexenyl acetate and other volatile organic compounds emitted from clipped trees induced defenses in the neighboring trees within the 5 m radius. The effective distances of plant‒plant communication in trees were discussed from the viewpoint of the arthropod community structure in forest ecosystems.

Diversification of terpenoid emissions proposes a geographic structure based on climate and pathogen composition in Japanese cedar.

Biogenic volatile organic compounds emitted from plants are important constituents of atmospheric chemistry and play a major role in the resistance of plants against various environmental stresses. However, little is known about how abiotic and biotic environments on a geographic scale relate to diversifications of the emission. Here, we present variations of terpenes stored in and emitted from leaves of a single species in a common garden, using genetically differentiated local populations of Japanese cedar, the most dominant and widely distributed tree species in Japan. Furthermore, we determined the composition of fungal communities in 50 locations, based on the presence or absence of 158 fungal species inhabiting the cedar. The results showed that terpenoids, especially those that are emitted, were highly diversified and geographically structured among the 12 populations. The total amount of stored terpenes was negatively affected by warm and less-snow climates. On the other hand, variations in some emitted terpenoid species among the populations were correlated to antagonistic fungal species inhabiting the Japanese cedar. We propose that the diversification of composition and amount of stored and emitted terpenoids in the tree species is not only structured by climate, but also antagonistic fungal communities through biological interactions.

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.

Five-year nitrogen addition affects fine root exudation and its correlation with root respiration in a dominant species, Quercus crispula, of a cool temperate forest, Japan.

In forest ecosystems, fine root respiration directly contributes to belowground carbon (C) cycling. Exudation from fine roots indirectly affects C cycling via enhanced microbial decomposition of soil organic matter. Although these root-derived C fluxes are essential components of belowground C cycling, how nitrogen (N) addition affects these fluxes and their correlations remains unclear. In this study, fine root exudation, respiration and chemical/morphological traits were measured in a dominant canopy species, Quercus crispula Blume, found in a cool temperate forest, the Tomakomai Experimental Forest, Hokkaido University, which has undergone 5-year N addition. Soil-dissolved organic carbon (DOC) was also measured in both bulk and rhizosphere soils to evaluate the impact of fine root exudation on soil C cycling. Compared with a control plot with no N treatment, fine roots in the N addition plot exhibited larger diameters and higher N concentrations, but lower specific root lengths and areas. On a root-weight basis, respiration was not different between plots, but exudation was slightly higher under N addition. On a root-area basis, exudation was significantly higher in the N addition plot. Additionally, differences in DOC between rhizosphere and bulk soils were two times higher in the N addition plot than the control plot. Although fine root respiration was positively correlated with exudation in both the control and N addition plots, the ratio of exudation C to respiration C decreased after 5-year N addition. Nitrogen addition also affected absolute C allocation to fine root exudation and changed the C allocation strategy between exudation and respiration fluxes. These findings will help enhance predictions of belowground C allocation and C cycling under N-rich conditions in the future.

Genets of dwarf bamboo do not die after one flowering event: evidence from genetic structure and flowering pattern.

Dwarf bamboos in the genus Sasa are believed to be long-lived, synchronously flowering, and monocarpic plants. However, the monocarpy of dwarf bamboo has not been confirmed, because whether all ramets within one genet flower at the same time cannot be determined without differentiating the genetic structure among ramets. This study aims to evaluate the reproductive traits of Sasa pubiculmis by verifying the monocarpy and physiological integration between flowering ramets and non-flowering ramets during a 4-year flowering period. One genotypically identified genet, which covered an area of approximately 3 ha, had both flowering and non-flowering patches of ramets during the 4-year flowering period (2004-2007). A fraction of the flowering genet remained non-flowering during the 4 years of observation, and did not die after mass flowering. Flowering ramets were physically connected to non-flowering ramets via rhizomes, and assimilated (13)C was allocated from non-flowering ramets to flowering ramets. Consequently, we clarified that this dwarf bamboo potentially has polycarpic reproductive traits rather than monocarpic, and a genet can keep rhizomes and non-flowering patches alive to sustain the organism after mass flowering.

Intraspecific differences in spring leaf phenology in relation to tree size in temperate deciduous trees.

Spring leaf phenology strongly influences plant productivity in temperate deciduous forests. Many studies have detected earlier budburst and leaf maturation in smaller trees within species, and have discussed the adaptive significance of increasing carbon gain before canopy closure in small trees. However, some previous studies have found the opposite pattern, and the physiological and environmental bases for this discrepancy are incompletely understood. We investigated the spring leaf phenology of 11 deciduous species in a cool-temperate forest in Japan for 2 years with different amounts of snowfall, and also gathered data on the day of budburst from multiple studies to assess whether and how the timing of budburst is related to tree size, phylogeny, temperature and annual snowfall of study sites. We found that differences in the timing of budburst and leaf maturation between saplings (<1 m height) and short trees (1-10 m height) are larger than those between short trees and tall trees (>10 m height), resulting in non-linear relationships between timing and height in most species. Cumulative degree-days to the day of budburst were smaller in saplings than in tall trees, probably because saplings are selected to outcompete the other individuals/species to become established. Moreover, phylogenetic relatedness did not explain the difference in spring leaf phenology between saplings and tall trees in the literature survey. In addition, our literature survey showed that budburst occurred earlier in saplings of most species in sites with less snowfall, whereas budburst occurred earlier in tall trees at sites with heavy snowfall. These results suggest that the opposite patterns found in some studies may be due to (i) differences in the target size, as saplings show larger phenological discrepancies than short and tall trees, and (ii) the microclimate experienced by the tree, as sites with heavy snow show delayed sapling phenology.

Changes in petiole hydraulic properties and leaf water flow in birch and oak saplings in a CO2-enriched atmosphere.

Water relations in woody species are intimately related to xylem hydraulic properties. High CO(2) concentrations ([CO(2)]) generally decrease transpiration and stomatal conductance (g(s)), but there is little information about the effect of atmospheric [CO(2)] on xylem hydraulic properties. To determine the relationship between water flow and hydraulic structure at high [CO(2)], we investigated responses of sun and shade leaves of 4-year-old saplings of diffuse-porous Betula maximowicziana Regel and ring-porous Quercus mongolica Fisch. ex Ledeb. ssp. crispula (Blume) Menitsky grown on fertile brown forest soil or infertile volcanic ash soil and exposed to 500 micromol CO(2) mol(-1) for 3 years. Regardless of species and soil type, elevated [CO(2)] consistently decreased water flow (i.e., g(s) and leaf-specific hydraulic conductivity) and total vessel area of the petiole in sun leaves; however, it had no effect on these parameters in shade leaves, perhaps because g(s) of shade leaves was already low. Changes in water flow at elevated [CO(2)] were associated with changes in petiole hydraulic properties.

Seasonal changes of δD and δ18O in tree-ring cellulose of Quercus crispula suggest a change in post-photosynthetic processes during earlywood growth.

Leaf photosynthetic and post-photosynthetic processes modulate the isotope ratios of tree-ring cellulose. Post-photosynthetic processes, such as the remobilization of stored starch in early spring, are important to understanding the mechanisms of xylem formation in tree stems; however, untangling the isotope ratio signals of photosynthetic and post-photosynthetic processes imprinted on tree rings is difficult. Portions of carbon-bound hydrogen and oxygen atoms are exchanged with medium water during post-photosynthetic processes. We investigated the δD and δ18O values of tree-ring cellulose using Quercus crispula Blume trees in two different habitats to evaluate seasonal changes in the exchange rate (f-value) of hydrogen or oxygen with medium water, and examined the associations of the post-photosynthetic processes. Theoretically, if the f-value is constant, δD and δ18O would be positively correlated due to meteorological factors, while variation in the f-value will create a discrepancy and weak correlation between δD and δ18O due to the exchange of carbon-bound hydrogen and oxygen with medium water. The values of δD decreased drastically from earlywood to latewood, while those of δ18O increased to a peak and then decreased toward the latewood. The estimated seasonal f-value was high at the beginning of earlywood and decreased toward the latewood. The post-photosynthetic processes associated with changes in the f-value were the remobilization of stored starch and triose cycling during cellulose synthesis because of the shortage of photo-assimilates in early spring. Although we did not evaluate relevant physiological parameters, the seasonal pattern of δD and δ18O in tree-ring cellulose of Q. crispula was clear, suggesting that the dual isotope (δD and δ18O) approach can be used to reveal the resource allocation mechanisms underlying seasonal xylem formation.

Seasonal changes in the temperature response of photosynthesis in canopy leaves of Quercus crispula in a cool-temperate forest.

Understanding seasonal changes in photosynthetic characteristics of canopy leaves is indispensable for modeling the carbon balance in forests. We studied seasonal changes in gas exchange characteristics that are related to the temperature dependence of photosynthesis in canopy leaves of Quercus crispula Blume, one of the most abundant species in cool-temperate forests in Japan. Photosynthetic rate and ribulose-1,5-bisphosphate (RuBP) carboxylation capacity (V(cmax)) at 20 degrees C increased from June to August and then decreased in September. The activation energy of V(cmax), a measure of the temperature dependence of V(cmax), was highest in summer, indicating that V(cmax) was most sensitive to leaf temperature at this time. The activation energy of V(cmax) was significantly correlated with growth temperature. Other parameters related to the temperature dependence of photosynthesis, such as intercellular CO(2) partial pressure and temperature dependence of RuBP regeneration capacity, showed no clear seasonal trend. It was suggested that leaf senescence affected the balance between carboxylation and regeneration of RuBP. The model simulation showed that photosynthetic rate and its optimal temperature were highest in summer.

The effect of interspecific variation in photosynthetic plasticity on 4-year growth rate and 8-year survival of understorey tree seedlings in response to gap formations in a cool-temperate deciduous forest.

Gap formation increases the light intensity in the forest understorey. The growth responses of seedlings to the increase in light availability show interspecific variation, which is considered to promote biodiversity in forests. At the leaf level, some species increase their photosynthetic capacity in response to gap formation, whereas others do not. Here we address the question of whether the interspecific difference in the photosynthetic response results in the interspecific variation in the growth response. If so, the interspecific difference in photosynthetic response would also contribute to species coexistence in forests. We also address the further relevant question of why some species do not increase their photosynthetic capacity. We assumed that some cost of photosynthetic plasticity may constrain acquisition of the plasticity in some species, and hypothesized that species with larger photosynthetic plasticity exhibit better growth after gap formation and lower survivorship in the shade understorey of a cool-temperate deciduous forest. We created gaps by felling canopy trees and studied the relationship between the photosynthetic response and the subsequent growth rate of seedlings. Naturally growing seedlings of six deciduous woody species were used and their mortality was examined for 8 years. The light-saturated rate of photosynthesis (Pmax) and the relative growth rate (RGR) of the seedlings of all study species increased at gap plots. The extent of these increases varied among the species. The stimulation of RGR over 4 years after gap formation was strongly correlated with change in photosynthetic capacity of newly expanded leaves. The increase in RGR and Pmax correlated with the 8-year mortality at control plots. These results suggest a trade-off between photosynthetic plasticity and the understorey shade tolerance. Gap-demanding species may acquire photosynthetic plasticity, sacrificing shade tolerances, whereas gap-independent species may acquire shade tolerances, sacrificing photosynthetic plasticity. This strategic difference among species would contribute to species coexistence in cool-temperate deciduous forests.