Verification of our empirical understanding of the physiology and ecology of two contrasting plantation species using a trait database.

The effects of climate change on forest ecosystems take on increasing importance more than ever. Information on plant traits is a powerful predictor of ecosystem dynamics and functioning. We reviewed the major ecological traits, such as foliar gas exchange and nutrients, xylem morphology and drought tolerance, of Cryptomeria japonica and Chamaecyparis obtusa, which are major timber species in East Asia, especially in Japan, by using a recently developed functional trait database for both species (SugiHinokiDB). Empirically, C. obtusa has been planted under drier conditions, whereas C. japonica, which grows faster but thought to be less drought tolerant, has been planted under wetter conditions. Our analysis generally support the empirical knowledge: The maximum photosynthetic rate, stomatal conductance, foliar nutrient content and soil-to-foliage hydraulic conductance were higher in C. japonica than in C. obtusa. In contrast, the foliar turgor loss point and xylem pressure corresponding to 50% conductivity, which indicate drought tolerance, were lower in C. obtusa and are consistent with the drier habitat of C. obtusa. Ontogenetic shifts were also observed; as the age and height of the trees increased, foliar nutrient concentrations, foliar minimum midday water potential and specific leaf area decreased in C. japonica, suggesting that nutrient and water limitation occurs with the growth. In C. obtusa, the ontogenetic shits of these foliar traits were less pronounced. Among the Cupressaceae worldwide, the drought tolerance of C. obtusa, as well as C. japonica, was not as high. This may be related to the fact that the Japanese archipelago has historically not been subjected to strong dryness. The maximum photosynthetic rate showed intermediate values within the family, indicating that C. japonica and C. obtusa exhibit relatively high growth rates in the Cupressaceae family, and this is thought to be the reason why they have been selected as economically suitable timber species in Japanese forestry. This study clearly demonstrated that the plant trait database provides us a promising opportunity to verify out empirical knowledge of plantation management and helps us to understand effect of climate change on plantation forests by using trait-based modelling.

Estimating spatial variation in the effects of climate change on the net primary production of Japanese cedar plantations based on modeled carbon dynamics.

Spatiotemporal prediction of the response of planted forests to a changing climate is increasingly important for the sustainable management of forest ecosystems. In this study, we present a methodology for estimating spatially varying productivity in a planted forest and changes in productivity with a changing climate in Japan, with a focus on Japanese cedar (Cryptomeria japonica D. Don) as a representative tree species of this region. The process-based model Biome-BGC was parameterized using a plant trait database for Japanese cedar and a Bayesian optimization scheme. To compare productivity under historical (1996-2000) and future (2096-2100) climatic conditions, the climate scenarios of two representative concentration pathways (i.e., RCP2.6 and RCP8.5) were used in five global climate models (GCMs) with approximately 1-km resolution. The seasonality of modeled fluxes, namely gross primary production, ecosystem respiration, net ecosystem exchange, and soil respiration, improved after two steps of parameterization. The estimated net primary production (NPP) of stands aged 36-40 years under the historical climatic conditions of the five GCMs was 0.77 ± 0.10 kgC m-2 year-1 (mean ± standard deviation), in accordance with the geographical distribution of forest NPP estimated in previous studies. Under the RCP2.6 and RCP8.5 scenarios, the mean NPP of the five GCMs increased by 0.04 ± 0.07 and 0.14 ± 0.11 kgC m-2 year-1, respectively. The increases in annual NPP were small in the southwestern region because of the decreases in summer NPP and the small increases in winter NPP under the RCP2.6 and RCP8.5 scenarios, respectively. Under the RCP2.6 scenario, Japanese cedar was at risk in the southwestern region, in accordance with previous studies, and monitoring and silvicultural practices should be modified accordingly.

Subspecies divergence and pronounced phylogenetic incongruence in the East-Asia-endemic shrub Magnolia sieboldii.

BACKGROUND AND AIMS: The biogeographic patterns of the East-Asia-endemic shrub Magnolia sieboldii, in which the range of the subsp. sieboldii is interposed with the disjunct distribution of subsp. japonica, implies a complex evolutionary history, involving rapid speciation and hybridization. Here, we aim to reveal the evolutionary and phylogeographic histories of the species with a particular focus on the time of subspecies divergence, the hypothesis of secondary hybridization and the Pleistocene survival of each subspecies, using a combination of genetic analyses and ecological niche modelling. METHODS: Genetic variation, genetic structures and phylogenetic relationships were elucidated based on nuclear low-copy genes, chloroplast DNA, and nuclear simple sequence repeats (SSRs). A scenario selection analysis and divergence time estimation were performed using coalescent simulation in DIYABC and *BEAST. Ecological niche modelling and a test of niche differentiation were performed using Maxent and ENMTools. KEY RESULTS: All marker types showed deep, but pronouncedly incongruent, west-east genetic divergences, with the subspecies being delineated only by the nuclear low-copy genes. Phylogenetic tree topologies suggested that ancient hybridization and introgression were likely to have occurred; however, this scenario did not receive significant support in the DIYABC analysis. The subspecies differentiated their niches, but both showed a dependence on high humidity and were predicted to have persisted during the last glacial cycle by maintaining a stable latitudinal distribution via migration to lower altitudes. CONCLUSIONS: We found a deep genetic divergence and a pronounced phylogenetic incongruence among the two subspecies of M. sieboldii, which may have been driven by major paleogeographic and paleoclimatic events that have occurred since the Neogene in East Asia, including global cooling, climate oscillations and the formation of land bridges. Both subspecies were, however, considered to persist in situ in stable climatic conditions during the late Pleistocene.

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 (2.4-2.8 °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 Prunus × yedoensis 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.

Responses to nitrogen pulses and growth under low nitrogen availability in invasive and native tree species with differing successional status.

Invasive species are frequently found in recently disturbed sites. To examine how these disturbance-dependent invasive species exploit resource pulses resulting from disturbance, twelve physiological and morphological traits, including age-dependent responsiveness in leaf traits to nitrogen pulse, were compared between Bischofia javanica, an invasive tree species in Ogasawara islands, and three native Ogasawara species, each having a different successional status. When exposed to a nitrogen pulse, invasive B. javanica showed higher increases in photosynthetic capacity, leaf area, epidermal cell number and cell size in leaves of broad age classes, and root nitrogen absorption ability than two native mid-/late or late-successional species, but showed no particular superiority to a native pioneer species in these responses. Under low nitrogen, however, it showed the largest relative growth rate among the four species, while the native pioneer showed the lowest growth. From these results, we concluded that the combination of moderately high responsiveness to resource pulses and the ability to maintain steady growth under resource limitations may give B. javanica a competitive advantage over a series of native species with different successional status from early to late-successional stages.

A paradox of leaf-trait convergence: why is leaf nitrogen concentration higher in species with higher photosynthetic capacity?

It is well known that leaf photosynthesis per unit dry mass (A(mass)) is positively correlated with nitrogen concentration (N(mass)) across naturally growing plants. In this article we show that this relationship is paradoxical because, if other traits are identical among species, plants with a higher A(mass) should have a lower N(mass), because of dilution by the assimilated carbon. To find a factor to overcome the dilution effect, we analyze the N(mass)-A(mass) relationship using simple mathematical models and literature data. We propose two equations derived from plant-growth models. Model prediction is compared with the data set of leaf trait spectrum obtained on a global scale. The model predicts that plants with a higher A(mass) should have a higher specific nitrogen absorption rate in roots (SAR), less biomass allocation to leaves, and/or greater nitrogen allocation to leaves. From the literature survey, SAR is suggested as the most likely factor. If SAR is the sole factor maintaining the positive relationship between N(mass) and A(mass), the variation in SAR is predicted to be much greater than that in A(mass); given that A(mass) varies 130-fold, SAR may vary more than 2000-fold. We predict that there is coordination between leaf and root activities among species on a global scale.

Correlation between relative growth rate and specific leaf area requires associations of specific leaf area with nitrogen absorption rate of roots.

Close correlations between specific leaf area (SLA) and relative growth rate (RGR) have been reported in many studies. However, theoretically, SLA by itself has small net positive effect on RGR because any increase in SLA inevitably causes a decrease in area-based leaf nitrogen concentration (LNCa), another RGR component. It was hypothesized that, for a correlation between SLA and RGR, SLA needs to be associated with specific nitrogen absorption rate of roots (SAR), which counteracts the negative effect of SLA on LNCa. Five trees and six herbs were grown under optimal conditions and relationships between SAR and RGR components were analyzed using a model based on balanced growth hypothesis. SLA varied 1.9-fold between species. Simulations predicted that, if SAR is not associated with SLA, this variation in SLA would cause a47% decrease in LNCa along the SLA gradient, leading to a marginal net positive effect on RGR. In reality, SAR was positively related to SLA, showing a 3.9-fold variation, which largely compensated for the negative effect of SLA on LNCa. Consequently, LNCa values were almost constant across species and a positive SLA-RGR relationship was achieved. These results highlight the importance of leaf-root interactions in understanding interspecific differences in RGR.

Applicability and limitations of optimal biomass allocation models: a test of two species from fertile and infertile habitats.

BACKGROUND AND AIMS: The practical applicability of optimal biomass allocation models is not clear. Plants may have constraints in the plasticity of their root : leaf ratio that prevent them from regulating their root : leaf ratio in the optimal manner predicted by the models. The aim of this study was to examine the applicability and limitations of optimal biomass allocation models and to test the assumption that regulation of the root : leaf ratio enables maximization of the relative growth rate (RGR). METHODS: Polygonum cuspidatum from an infertile habitat and Chenopodium album from a fertile habitat were grown under a range of nitrogen availabilities. The biomass allocation, leaf nitrogen concentration (LNC), RGR, net assimilation rate (NAR), and leaf area ratio (LAR) of each species were compared with optimal values determined using an optimal biomass allocation model. KEY RESULTS: The root : leaf ratio of C. album was smaller than the optimal ratio in the low-nitrogen treatment, while it was almost optimal in the high-nitrogen treatment. In contrast, the root : leaf ratio of P. cuspidatum was close to the optimum under both high- and low-nitrogen conditions. Owing to the optimal regulation of the root : leaf ratio, C. album in the high-nitrogen treatment and P. cuspidatum in both treatments had LNC and RGR (with its two components, NAR and LAR) close to their optima. However, in the low-nitrogen treatment, the suboptimal root : leaf ratio of C. album led to a smaller LNC than the optimum, which in turn resulted in a smaller NAR than the optimum and RGR than the theoretical maximum RGR. CONCLUSIONS: The applicability of optimal biomass allocation models is fairly high, although constraints in the plasticity of biomass allocation could prevent optimal regulation of the root : leaf ratio in some species. The assumption that regulation of the root : leaf ratio enables maximization of RGR was supported.