Cool-temperate Deciduous Forest Research Articles

The Nitrogen Cycle of a Cool-Temperate Deciduous Broad-Leaved Forest

The nitrogen (N) cycle, a major biogeochemical cycle in forest ecosystems, notably affects ecosystem multifunctionality. However, the magnitude and role of organic N and the snow season remain uncertain in this cycle. We assessed the N flux and pool data of a temperate deciduous broad-leaved forest to clarify N cycle processes. The results showed that the most important component of the N pool was the soil N pool. The N demand of the site amounted to 139.4 kg N ha−1 year−1 and was divided into tree production (83.8%) and bamboo production (16.2%). We clarified that retranslocation (37.4%), mineralization at a soil depth of 0–5 cm (15.3%), litter leachate (4.6%), throughfall (2.3%), and canopy uptake (0.5%) provided 60.1% of the N demand. In terms of soil at 0–5 cm in depth, the net mineralization rate during the snow season contributed to 30% of the annual mineralization. We concluded that the study site was not N-saturated as a result of a positive N input–output flux budget. More than half of the total N was accounted for by dissolved organic N flowing through several pathways, indicating that organic N plays a vital role in the cycle. The mineralization rate in the soil layer during the snow season is an important link in the N cycle.

Long-term changes in vegetation and land use in mountainous areas with heavy snowfalls in northern Japan: an 80-year comparison of vegetation maps

Comparison of old and new vegetation maps is an effective way to detect vegetation dynamics. Recent developments in computer technology have made it possible to accurately compare old paper vegetation maps with current digitized vegetation maps to reveal long-term vegetation dynamics. Recently, a 1:50,000 scale vegetation map of the Hakkoda Mountains in northern Japan, located in the ecotone of cool temperate and subalpine forests in northern Japan under an East Asian monsoon climate, from 1930 was discovered. We compared the 1930s vegetation map with the most recent 2010 vegetation map to test the following hypotheses: 1) the occurrence of upward expansion of the upper limit of cool-temperate deciduous forests, and 2) whether designation as a national park in 1936 would have reduced forestry and land use, expanded beech forests, and cool-temperate deciduous forests. To compare vegetation changes, 67 types of vegetation legends for the 1930 and 2010 maps were unified to 21 based on plant species composition. Consequently, vegetation has changed substantially over the past 80 years. 1) In the subalpine zone above 1,000 m, the coniferous forest area decreased by half. In the cool temperate zone below 1,000 m, the area of beech forests increased 1.48 times, and some of them could be shifted upwards, replacing subalpine fir forests in the lower part of the subalpine zone. 2) In areas below 700 m, deciduous oak forests once used as thickets were almost halved. Instead, climax and beech forests expanded. However, we also found that even after the area was declared a national park, oak forests were cleared and converted to commercial forests such as cedar plantations, cattle ranches, and horse pastures in some areas. These results will be useful for future ecosystem and biodiversity research/conservation and will provide baseline information for climate change adaptation policies.

Contributions of the understory and midstory to total canopy solar-induced chlorophyll fluorescence in a ground-based study in conjunction with seasonal gross primary productivity in a cool-temperate deciduous broadleaf forest

Solar-induced chlorophyll fluorescence (SIF) can represent gross primary productivity (GPP) in many types of terrestrial vegetation. In principle, the chlorophyll-a fluorescence signal responds to the amount of light absorption and the fraction of energy distribution in photosystems. Therefore, it is mechanistically linked with CO2 assimilation. Recently, radiative transfer models have traced the processes of emission, interception and reabsorption within a canopy in heterogeneous tree stands. These processes influence the expected relationship between SIF and GPP. We inferred that the vertical profile of SIF could reveal those processes. In addition, the sum of SIF emission from different pathways within the canopy must be equivalent to the total SIF emission from all leaves. The purposes of this study were: (1) to clarify the seasonal and diurnal variations of ground-based SIF observed above the overstory, midstory and understory layers (strata); (2) to examine whether the sum of SIF observed from the three layers could represent total SIF emissions, and (3) to examine whether SIF above the understory could reveal the photosynthetic activity within the vertical profile of a forest. To estimate how much SIF was present after its emission from leaves within the canopy, we conducted spectral radiation observations in three vertical layers: above the understory, midstory and overstory (8, 14 and 18 m from the ground, respectively) to retrieve the red and far-red SIF in a deciduous broadleaf forest ecosystem in Japan from April to November 2020. We found that SIF above the overstory and the sum of SIF from the three layers increased sharply at the leaf-onset of the overstory and was correlated with each season's GPP. The sum of far-red SIF from the three layers was proportional to the total SIF estimated using the escape ratio approach for SIF above the overstory. In this study we demonstrated that SIF above the understory could detect the increased photosynthesis of the understory in spring, compared with the GPP and the vertical profile of the CO2 concentrations. In summary, SIF observed in three forest layers detected the seasonal change of GPP within the canopy. Thus, a multi-layer approach can be used to understand the relationship between SIF and photosynthesis within the canopy.

Drone-Sensed and Sap Flux-Derived Leaf Phenology in a Cool Temperate Deciduous Forest: A Tree-Level Comparison of 17 Species

Understanding the relationship between leaf phenology and physiological properties has important implications for improving ecosystem models of biogeochemical cycling. However, previous studies have investigated such relationships only at the ecosystem level, limiting the biological interpretation and application of the observed relationships due to the complex vegetation structure of forest ecosystems. Additionally, studies focusing on transpiration are generally limited compared to those on photosynthesis. Thus, we investigated the relationship between stem sap flux density (SFD) and crown leaf phenology at the individual tree level using the heat dissipation method, unmanned aerial vehicle (UAV)-based observation, and ground-based visual observation across 17 species in a cool temperate forest in Japan, and assessed the potential of UAV-derived phenological metrics to track individual tree-level sap flow phenology. We computed five leaf phenological metrics (four from UAV imagery and one from ground observations) and evaluated the consistency of seasonality between the phenological metrics and SFD using Bayesian modelling. Although seasonal trajectories of the leaf phenological metrics differed markedly among the species, the daytime total SFD (SFDday) estimated by the phenological metrics was significantly correlated with the measured ones across the species, irrespective of the type of metric. Crown leaf cover derived from ground observations (CLCground) showed the highest ability to predict SFDday, suggesting that the seasonality of leaf amount rather than leaf color plays a predominant role in sap flow phenology in this ecosystem. Among the UAV metrics, Hue had a superior ability to predict SFDday compared with the other metrics because it showed seasonality similar to CLCground. However, all leaf phenological metrics showed earlier spring increases than did sap flow in more than half of the individuals. Our study revealed that UAV metrics could be used as predictors of sap flow phenology for deciduous species in cool, temperate forests. However, for a more accurate prediction, phenological metrics representing the spring development of sap flow must be explored.

Combining both spectral and textural indices for alleviating saturation problem in forest LAI estimation using Sentinel-2 data

Leaf area index (LAI) is an important structural parameter that controls energy, water, and gas exchanges of plant ecosystems. At present, remote sensing techniques, especially passive optical remote sensing, are the most widely used approaches to estimate LAI. However, the prevalent usage of spectral information alone often saturates, which is a major issue for retrieving LAI in particular for forests. In this study, our goal was to improve LAI estimation using Sentinel-2 data by combining both spectral and texture information to minimize the saturation effect in a typical cool-temperate deciduous forest in Japan. Two LAI field datasets obtained using digital hemispherical photographs, representing a range of LAI conditions, were used to investigate the performance of spectral and texture features or a combination of both. Texture information was derived using the grey-level co-occurrence matrix (GLCM) method. Commonly used vegetation indices and newly developed textural indices were computed for comparison. Spatial autocorrelation was determined prior to exploratory regression analyses to explore methods for estimating LAI using a combination of spectral- and texture-based features. Regression methods included generalized least squares (GLS), stepwise, principal components (PCR), and partial least squares (PLS). The results demonstrate that the texture information derived from Sentinel-2 imagery helps estimate LAI by minimizing the saturation problem, especially in the period of maximum LAI values. The combination of both features using the PLS model provided a higher accuracy estimation scheme for LAI (R 2: 0.66–0.75, RMSE: 0.23–0.37) and we foresee the wide application of the approach especially for a closed-canopy forest facing serious saturation problem when deriving LAI from reflected spectral information.

Diverse Heat Tolerance of the Yeast Symbionts of Platycerus Stag Beetles in Japan.

The genus Platycerus (Coleoptera: Lucanidae) is a small stag beetle group, which is adapted to cool-temperate deciduous broad-leaved forests in East Asia. Ten Platycerus species in Japan form a monophyletic clade endemic to Japan and inhabit species-specific climatic zones. They are reported to have co-evolutionary associations with their yeast symbionts of the genus Sheffersomyces based on host cytochrome oxidase subunit I (COI) and yeast intergenic spacer (IGS) phylogenies. Here we examined the heat tolerances of the yeast colonies isolated from the mycangia of 37 females belonging ten Japanese Platycerus species. The upper limits of growth and survival temperatures of each colony were decided by cultivating it at ten temperature levels between 17.5 and 40°C. Although both temperatures varied during 25.0–31.25°C, the maximum survival temperatures (MSTs) were a little higher than the maximum growth temperatures (MGTs) in 16 colonies. Pearson’s correlations between these temperatures and environmental factors (elevation and 19 bioclimatic variables from Worldclim database) of host beetle collection sites were calculated. These temperatures were significantly correlated with elevation negatively, the maximum temperature of the warmest month (Bio5) positively, and some precipitative variables, especially in the warm season (Bio12, 13, 16, 18) negatively. Sympatric Platycerus kawadai and Platycerus albisomni share the same lineage of yeast symbionts that exhibit the same heat tolerance, but the elevational lower range limit of P. kawadai is higher than that of P. albisomni. Based on the field survey in their sympatric site, the maximum temperature of host wood of P. kawadai larvae is higher about 2–3°C than that of P. albisomni larvae in the summer, which may restrict the elevational range of P. kawadai to higher area. In conclusion, it is suggested that the heat tolerance of yeast symbionts restricts the habitat range of their host Platycerus species or/and that the environmental condition that host Platycerus species prefers affect the heat tolerance of its yeast symbionts.

Exploring the instability of the relationship between maximum potential electron transport rate and maximum carboxylation rate in cool-temperate deciduous forests

A clear understanding of the relationship between two photosynthetic capacity parameters, maximum carboxylation rate (Vcmax) and maximum electron transport rate (Jmax), is expected to reveal plant photosynthetic physiology strategies and is critical in determining the carbon and water cycle in the terrestrial biosphere. Increasing evidence in recent suggests the relationship is unstable in spatiotemporal contexts, casting a doubt on deriving Jmax solely from a single linear function of Vcmax as in most gas exchange models do. In this study, we have examined the instability of the relationship in terms of seasonal and interannual courses (from 2006 to 2014) as well as at different altitudes (550, 900, and 1500 m a.s.l.) and in different leaf types (sunlit and shaded) of the only dominant species of Fagus crenata Blume in typical cool temperate forests in Japan based on a long-term field dataset. Our results clearly indicated that there is not only remarkable seasonal variability in the two parameters themselves but also large variations in their ratios. The slope parameter of the best fitted single linear function for estimating Jmax from Vcmax varied also in different seasons and different years and different leaf types, as well as at different sites. Further exploration of the slope parameter with internal leaf traits and external climatic drivers revealed that the leaf chlorophyll content had a significant correlation with the slope parameter (R2 = 0.72, RMSE = 0.08, RPD = 2.00) but the impacts of climatic drivers were rather versatile. Our findings should hence advance understanding of the dynamics of photosynthetic capacity and the instability of the Jmax–Vcmax relationship, which should be considered explicitly in the future in order to substantially improve model predictions in terrestrial carbon cycle models.

Including leaf trait information helps empirical estimation of jmax from vcmax in cool-temperate deciduous forests

Understanding the uncertainty in the parameterization of the two photosynthetic capacity parameters, leaf maximum carboxylation rate (Vcmax), and maximum electron transport rate (Jmax), is crucial for modeling and predicting carbon fluxes in terrestrial ecosystems. In gas exchange models, to date, Jmax is typically estimated from Vcmax based on a linear regression. However, recent studies have revealed that this relationship varies, dependent upon species, leaf groups, and time, so it is doubtful that the regression applies universally. Furthermore, far less is known regarding how other leaf traits affect the regression. In this study we analyzed the two key photosynthetic parameters and popularly measurable leaf traits, leaf chlorophyll concentration and leaf mass per area (LMA), of cool-temperate forest stands in Japan, aiming to construct a simple regression applicable to temperate deciduous forests, at least. The analysis was based on a long-term field dataset covering years of data for both sunlit and shaded leaves at different altitudes. Results showed that the best-fitted slope of the regression differed markedly from those previously reported, which were typically acquired from sunlit leaves. LMA had a significant effect on the regression, producing the lowest root mean square errors and the highest ratio of performance to deviation values (RPD = 2.017). Although more data are needed to validate in other ecosystems, our approach at least provides a promising way to substantially improve photosynthesis model predictions, by introducing leaf traits into the popular empirical regression of Jmax against Vcmax, and ultimately to better understand the functioning of the photosynthetic machinery.

Area-ratio Fraunhofer line depth (aFLD) method approach to estimate solar-induced chlorophyll fluorescence in low spectral resolution spectra in a cool-temperate deciduous broadleaf forest.

Solar-induced chlorophyll fluorescence (SIF) emissions were estimated by the "area-ratio Fraunhofer line depth (aFLD) method", a new retrieval methodology in spectra from a low spectral resolution (SR) spectroradiometer (MS-700: full width half maximum (FWHM) of 10nm and spectral sampling interval of 3.3nm), assisted with a scaling to reference SIF detected from high SR spectrum. The sparse pixels of a spectrum of low SR misses detecting the minimum of the O2A absorption band around at 760nm, which makes the SIF detection by conventional FLD methods lose accuracy considerably. To overcome this, the aFLD method uses the definite integral of spectra over a wide interval between 750 and 780nm. The integration of the spectrum is insusceptible to the change in shape of the depression curve, leading to higher accuracy of the aFLD method. Daily SIF, calculated by the aFLD method using the spectra obtained with MS-700, was scaled to reference daily SIF calculated by the spectral fitting method using the spectra obtained from August to December 2019 with an ultrafine SR spectroradiometer (QE Pro, FWHM = 0.24nm). As a result, SIF calculated from MS-700 spectra by aFLD method was strongly correlated with the reference SIF from QE Pro spectra (r2 = 0.81) and was successfully scaled. Then, the scaled 11-year SIF from MS-700 at a deciduous broadleaf forest showed the correlation with GPP at multiple time steps: daily, monthly, and yearly, consistently during 2008-2018. The comparison of aFLD-derived SIF with the global Orbiting Carbon Observatory-2 (OCO-2) SIF data set (GOSIF) showed high correlation on monthly values during 2008-2017 (r2 = 0.85). The combining approach of the aFLD method with a scaling to reference SIF successfully detected long-term canopy SIF emissions, which has great potential to provide essential information on ecosystem-level photosynthesis.

Crown Structure Explains the Discrepancy in Leaf Phenology Metrics Derived from Ground- and UAV-Based Observations in a Japanese Cool Temperate Deciduous Forest

Unmanned aerial vehicles (UAV) provide a new platform for monitoring crown-level leaf phenology due to the ability to cover a vast area while offering branch-level image resolution. However, below-crown vegetation, e.g., understory vegetation, subcanopy trees, and the branches of neighboring trees, along with the multi-layered structure of the target crown may significantly reduce the accuracy of UAV-based estimates of crown leaf phenology. To test this hypothesis, we compared UAV-derived crown leaf phenology results against those based on ground observations at the individual tree scale for 19 deciduous broad-leaved species (55 individuals in total) characterized by different crown structures. The mean crown-level green chromatic coordinate derived from UAV images poorly explained inter- and intra-species variations in spring leaf phenology, most probably due to the consistently early leaf emergence in the below-crown vegetation. The start dates for leaf expansion and end dates for leaf falling could be estimated with an accuracy of <1-week when the influence of below-crown vegetation was removed from the UAV images through visual interpretation. However, a large discrepancy between the phenological metrics derived from UAV images and ground observations was still found for the end date of leaf expansion (EOE) and start date of leaf falling (SOF). Bayesian modeling revealed that the discrepancy for EOE increased as crown length and volume increased. The crown structure was not found to contribute to the discrepancy in SOF value. Our study provides evidence that crown structure is a pivotal factor to consider when using UAV photography to reliably estimate crown leaf phenology at the individual tree-scale.

A Spatial Relationship between Canopy and Understory Leaf Area Index in an Old-Growth Cool-Temperate Deciduous Forest

Quantification of leaf area index (LAI) is essential for understanding forest productivity and the atmosphere–vegetation interface, where the majority of gas and energy exchange occurs. LAI is one of the most difficult plant variables to adequately quantify, owing to large spatial and temporal variability, and few studies have examined the horizontal and vertical distribution of LAI in forest ecosystems. In this study, we demonstrated the LAI distribution in each layer from the understory to canopy using multiple-point measurements (121 points) and examined the relationships among layers in a cool-temperate deciduous forest. LAI at each point, and the spatial distribution of LAI in each layer, varied within the forest. The spatial distribution of LAI in the upper layer was more heterogeneous than that of LAI at the scale of the entire forest. Significant negative correlations were observed between the upper- and lower-layer LAI. Our results indicate that the understory compensates for gaps in LAI in the upper layer; thus, the LAI of the entire forest tends to remain spatially homogeneous even in a mature forest ecosystem.

Carbon and Nutrient Inputs by Litterfall in Evergreen and Deciduous Forests in Korea

Knowledge about carbon and nutrient fluxes by litterfall is important for understanding nutrient cycling in geologically unique ecosystems. However, the determination of forest litterfall production patterns is difficult due to many biophysical factors influencing the process. In this study, we (1) quantified the litterfall production and carbon and nutrient fluxes in warm-temperate evergreen forest stands in Jeju Gotjawal and (2) compared these values to those of a typical cool-temperate deciduous forest stand by forest types and climate differences. Litterfall from evergreen broadleaved forests at Cheongsu (CS) and Seonheul (SHb), a mixed forest at Seonheul (SHm) in Jeju Gotjawal, and a deciduous broadleaved forest at Chungnam National University Forest (CNU) was collected for a full two years using litter traps. Samples were sorted into leaves, twigs, barks, seeds, and unidentified materials, and then weighed and measured for C, N, P, K, Ca, and Mg fluxes by litterfall. Results showed that the mean annual litterfall (846.3 g m−2, average of CS, SHb, and SHm) at Jeju Gotjawal was similar to that of CNU (885.5 g m−2), but varied by site in Jeju Gotjawal: CS (933.1 g m−2) was significantly higher than the average of SHb and SHm (802.9 g m−2). Seasonal patterns of litterfall production differed by forest types; evergreen broadleaved forests showed a bimodal peak in fall and spring while deciduous broadleaved forests showed a unimodal peak in fall. Jeju Gotjawal had significantly higher total macronutrient concentrations and contents (except for K) than CNU and they also varied by site in Jeju Gotjawal: CS had higher N, P, Ca, and Mg contents than SHb and SHm. We conclude that litterfall production and nutrient fluxes differed by forest stand as influenced by forest types and climate. Further, our findings are important for understanding carbon and nutrient dynamics in the geologically unique ecosystem of Jeju Gotjawal and other areas with similar characteristics.

Nitrogen Deposition and Responses of Forest Structure to Nitrogen Deposition in a Cool-Temperate Deciduous Forest

Few studies have reported the estimation of nitrogen (N) deposition, including dissolved organic N (DON) fluxes, through water flows and the contribution of snowfall in Asia. In this study, the concentrations and fluxes of DON and dissolved inorganic N (DIN) in bulk precipitation (BP), the throughfall (TF) of trees and understory dwarf bamboo, and stemflow (SF) were evaluated in a cool-temperate forest over three years to clarify N fluxes via precipitation and responses of trees and understory canopies to N deposition. The input of N to the study site in BP was 11.1 ± 1.71 kg N ha−1 year−1, with a significant contribution from DON (78%). Snowfall fluxes contributed up to 46% of the total N input, with variations related to the amount of snowfall (2.08–5.52 kg N ha−1 year−1). The forest canopy enriched DON (2.11 ± 0.42 kg N ha−1 year−1) but consumed NO3 + NO2–N (−0.73 ± 0.19 kg N ha−1 year−1). In contrast, through the understory bamboo canopy, DON (−1.02 ± 0.55 kg N ha−1 year−1) decreased while DIN (0.35 ± 0.44 kg N ha−1 year−1) increased. This study indicates that DON and snowfall should not be neglected when evaluating total N deposition into forest ecosystems, especially in remote regions. The canopy processes related to the dissolved N in the presence of understory plants might have significant implications for the internal N cycle in forest ecosystems.

Formation of disjunct plant distributions in Northeast Asia: a case study of Betula davurica using a species distribution model

Repeated climate change during glacial and interglacial periods of the Quaternary led to mass migrations that resulted in disjunct distributions for many species. However, few studies have examined the processes that form disjunct distributions in Northeast Asia (NEA). In this study, we examined the disjunct distribution of Betula davurica Pall. in the Japanese archipelago. This species is a dominant canopy tree found in cool-temperate deciduous broad-leaved forests of continental NEA. We hypothesized that Quaternary climate change caused the present disjunct distribution pattern of this species. To test this hypothesis, we adopted a species distribution model and examined a series of potential habitats in the Last Glacial Maximum (LGM), the mid-Holocene, and the present. We generated models in MaxEnt with B. davurica presence as the response variable and six bioclimatic variables as predictor variables. During the LGM, projected potential habitats were distributed around the Korean Peninsula, East China, and the Japanese archipelago, excluding Hokkaido. In the mid-Holocene, habitats retreated both from East China and western Japan, remained unchanged in the Korean Peninsula and central Honshu mountains, and expanded to northern China, the Russian Far East, as well as northern Japan (Hokkaido). Thus, post-LGM global warming led to an expansion of B. davurica distribution to northern parts of continental NEA, along with a retreat in the Japanese archipelago. This shift in populations formed the present disjunct distribution.

High soil microbial activity in the winter season enhances nitrogen cycling in a cool-temperate deciduous forest

The soil microbial community plays essential roles in bioavailable nitrogen (N) cycling, and microbial dynamics during the plant dormant season could affect N cycling in the subsequent growing season. Despite frequent observations of seasonal shifts in microbial community composition in forests, the biogeochemical consequences for bioavailable N cycling have not been well characterized. Here we examine the relationship between microbial community dynamics and bioavailable N dynamics throughout a year in a cool-temperate deciduous forest, with a focus on the dormant season. Specifically, we analyzed temporal changes in abundances of N-cycling microbial populations and N concentrations and transformations in soils. We also assessed temporal changes in bacterial community composition by deep sequencing of 16S rRNA genes. Even though the bacterial community composition was stable throughout the year, we found a drastic increase in abundances of total bacterial and fungal populations in the mid-winter, followed by a subsequent decrease during the snowmelt period in the early spring. We also found a temporary increase in abundances of NH3-oxidizing bacterial population in the mid-winter and of denitrifying bacterial and fungal populations during the snowmelt period. Correspondingly, soil NH4+, NO3−, and dissolved organic N concentrations sequentially peaked in the winter and early spring. These results suggest that successive growth of litter degraders, ammonifiers, nitrifiers, and denitrifiers in the dormant season drives the subsequent bioavailable N transformations. High microbial N metabolic activities during the dormant season could also support plant growth during the growing season and increase the efficiency of annual N cycling in the forest via the seasonal partitioning of N between plants and microbial communities.

Long-term characteristics of soil respiration in a Korean cool-temperate deciduous forest in a monsoon climate

ABSTRACTAnalysis of relationship between soil respiration and environmental factors has become essential for understanding changes in ecosystem carbon cycles under global warming. However, rough predictions have been made that soil respiration will increase with increasing temperature, but long-term data to support this theory were scarce. We measured soil respiration and environmental factors continuously using an automatic open-closed chamber system in a Korean cool-temperate forest from 2004 to 2016 to ascertain the reliability of this prediction and to more accurately predict changes in carbon cycle. Average air and soil temperatures were 11.0°C and 10.2°C. The increase in temperature was greater in winter (the inactive period for soil respiration) than in summer (the active period). Additionally, precipitation decreased sharply because of patter changes in 2012, and through 2016, it was approximately 69% of the previous period. Effect of precipitation on soil respiration was expected to be larger than temperature because the change in precipitation appeared in summer. Soil respiration exhibited a significant decline in 2012 because of precipitation. From 2004 to 2011, it averaged 344.4 mgCO2 m−2 h−1 and from 2012 to 2016 the average was 205.3 mgCO2 m−2 h−1. This phenomenon hasn’t been detected in short-term studies, suggesting that the prediction of previous studies is inaccurate. Additionally, to predict future ecosystem carbon cycle changes in a cool-temperate monsoon climate, changes in precipitation pattern should be regarded as equally important to temperature, and the prediction cannot be based solely on temperature. Therefore, long-term and continuous measurements are needed with consideration of the effects of both precipitation and temperature.Abbreviations: Rs: soil respiration; Ts: soil temperature; Ta: air temperature; AOCC: automatic open/closed chamber

Soil depth distribution and the patterns of alpha- and beta-diversity of families of soil Collembola in cool-temperate deciduous natural forests and larch plantations of northern Japan

Soil depth distribution and the patterns of alpha- and beta-diversity of families of soil Collembola in cool-temperate deciduous natural forests and larch plantations of northern Japan

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.

Phylogeography of postglacial range expansion in Juglans mandshurica (Juglandaceae) reveals no evidence of bottleneck, loss of genetic diversity, or isolation by distance in the leading-edge populations

The past studies of postglacial recolonization patterns in high latitude regions have revealed a significant role of dispersal capacity in shaping the genetic diversity and population structure of temperate trees. However, most of these studies have focused on species with long-distance dispersal followed by exponential population growth and were therefore unable to reveal the patterns in the case of a gradual expansion. Here we studied the impacts of postglacial range expansions on the distribution of genetic diversity in the Manchurian walnut (Juglans mandshurica), a common tree of East Asian cool-temperate deciduous forests that apparently lacks long-distance seed dispersal ability. The genetic diversity and structure of 19 natural walnut populations in Northeast China and the Korean Peninsula were examined using 17 nuclear simple sequence repeat (SSR) loci. Potential habitats under current and past climatic conditions were predicted using the ecological niche modelling (ENM) method. Bayesian clustering analysis revealed three groups, which were inferred to have diverged through multiple glacial-interglacial cycles in multiple refugia during the Quaternary Period. ENM estimated a southward range shift at the LGM, but high suitability scores still occurred in the western parts of the Changbai Mountains (Northeast China), the Korean peninsula and the exposed seafloor of the Yellow Sea. In contrast to most other cool-temperate trees co-occurring in the same region, the Manchurian walnut did not show any evidence of a population bottleneck, loss of genetic diversity or isolation by distance during the postglacial expansion. Our study clearly indicates that current northern populations originated from one glacial lineage and recolonization via a gradually advancing front due to the lack of a long-distance seed dispersal mechanism led to no latitudinal decrease in genetic diversity.

Responses of Soil, Heterotrophic, and Autotrophic Respiration to Experimental Open-Field Soil Warming in a Cool-Temperate Deciduous Forest

How global warming will affect soil respiration (R S) and its source components is poorly understood despite its importance for accurate prediction of global carbon (C) cycles. We examined the responses of R S, heterotrophic respiration (R H), autotrophic respiration (R A), nitrogen (N) availability, and fine-root biomass to increased temperature in an open-field soil warming experiment. The experiment was conducted in a cool-temperate deciduous forest ecosystem in northern Japan. As this forest is subjected to strong temporal variation in temperature, on scales ranging from daily to seasonal, we also investigated the temporal variation in the effects of soil warming on R S, R H, and R A. Soil temperature was continuously elevated by about 4.0°C from 2007 to 2014 using heating wires buried in the soil, and we measured soil respiratory processes in all four seasons from 2012 to 2014. Soil warming increased annual R S by 32–45%, but the magnitude of the increase was different between the components: R H and R A were also stimulated, and increased by 39–41 and 17–18%, respectively. Soil N availability during the growing season and fine-root biomass were not remarkably affected by the warming treatment. We found that the warming effects varied seasonally. R H increased significantly throughout the year, but the warming effect showed remarkable seasonal differences, with the maximum stimulation in the spring. This suggests that warmer spring temperature will produce a greater increase in CO2 release than warmer summer temperatures. In addition, we found that soil warming reduced the temperature sensitivity (Q 10) of R S. Although the Q 10 of both R H and R A tended to be reduced, the decrease in the Q 10 of R S was caused mainly by a decrease in the response of R A to warming. These long-term results indicate that a balance between the rapid and large response of soil microbes and the acclimation of plant roots both play important roles in determining the response of R S to soil warming, and must be carefully considered to predict the responses of soil C dynamics under future temperature conditions.