It has been argued that scale is the central problem in ecology (Levin, 1992). Studies on carbon cycles and global climate change, the current major themes in modern ecology, require the interfacing of phenomena that occur on different scales of space, time, and ecological organization. For several decades, tremendous efforts have been made to reveal the general patterns of, and the mechanisms for the global carbon cycles. However, many uncertainties remain, particularly on local to regional scales. To reduce these uncertainties, regional collaborations across the board of nations are required.

Aims Tropical forest plays a key role in global C cycle; however, there are few studies on the C budget in the tropical rainforests in Asia. This study aims to (i) reveal the seasonal patterns of total soil respiration (R T), litter respiration (R L) and soil respiration without surface organic litter (R NL) in the primary and secondary Asian tropical mountain rainforests and (ii) quantify the effects of soil temperature, soil moisture and substrate availability on soil respiration.
Methods The seasonal dynamics of soil CO₂ efflux was measured by an automatic chamber system (Li-8100), within the primary and secondary tropical mountain rainforests located at the Jianfengling National Reserve in Hainan Island, China. The litter removal treatment was used to assess the contribution of litter to belowground CO₂ production.
Important findings The annual R T was higher in the primary forest (16.73±0.87 Mg C ha^-1) than in the secondary forest (15.10±0.26 Mg C ha^-1). The rates of R T, R NL and R L were all significantly higher in the hot and wet season (May–October) than those in the cool and dry season (November–April). Soil temperature at 5cm depth could explain 55–61% of the seasonal variation in R T, and the temperature sensitivity index (Q 10) ranked by R L (Q 10 = 3.39)> R T (2.17)> R NL (1.76) in the primary forest and by R L (4.31)> R T (1.86)> R NL (1.58) in the secondary forest. The contribution of R L to R T was 22–23%, while litter input and R T had 1 month time lag. In addition, the seasonal variation of R T was mainly determined by soil temperature and substrate availability. Our findings suggested that global warming and increased substrate availability are likely to cause considerable losses of soil C in the tropical forests.

Aims Clear-cutting is a common forest management practice, especially in subtropical China. However, the potential ecological consequences of clear-cutting remain unclear. In particular, the effect of clear-cutting on soil processes, such as the carbon cycle, has not been quantified in subtropical forests. Here, we investigated the response of soil respiration (Rs) to clear-cutting during a 12-month period in a subtropical forest in eastern China.
Methods We randomly selected four clear-cut (CC) plots and four corresponding undisturbed forest (UF) plots. Measurements of Rs were made at monthly time points and were combined with continuous climatic measurements in both CC and UF. Daily Rs was estimated by interpolating data with an exponential model dependent on soil temperature. Daily Rs was cumulated to annual Rs estimates.
Important findings In the first year after clear-cutting, annual estimates of Rs in CC (508±23g C m ?2 yr^-1) showed no significant difference to UF plots (480±12g C m ?2 yr^-1). During the summer, soil temperatures were usually higher, whereas the soil volumetric water content was lower in CC than in UF plots. The long-term effects of clear-cutting on Rs are not significant, although there might be effects during the first several months after clear-cutting. Compared with previous work, this pattern was more pronounced in our subtropical forest than in the temperate and boreal forests that have been studied by others. With aboveground residuals off-site after clear-cutting, our results indicate that the stimulation of increasing root debris, as well as environmental changes, will not lead to a significant increase in Rs. In addition, long-term Rs will not show a significant decrease from the termination of root respiration, and this observation might be because of the influence of fast-growing vegetation after clear-cutting in situ .

Aims Boreal forest is the largest and contains the most soil carbon among global terrestrial biomes. Soil respiration during the prolonged winter period may play an important role in the carbon cycles in boreal forests. This study aims to explore the characteristics of winter soil respiration in the boreal forest and to show how it is regulated by environmental factors, such as soil temperature, soil moisture and snowpack.
Methods Soil respiration in an old-growth larch forest (Larix gmelinii Ruppr.) in Northeast China was intensively measured during the winter soil-freezing process in 2011 using an automated soil CO₂ flux system. The effects of soil temperature, soil moisture and thin snowpack on soil respiration and its temperature sensitivity were investigated.
Important findings Total soil respiration and heterotrophic respiration both showed a declining trend during the observation period, and no significant difference was found between soil respiration and heterotrophic respiration until the snowpack exceeded 20cm. Soil respiration was exponentially correlated with soil temperature and its temperature sensitivity (Q 10 value) for the entire measurement duration was 10.5. Snow depth and soil moisture both showed positive effects on the temperature sensitivity of soil respiration. Based on the change in the Q 10 value, we proposed a 'freeze–thaw critical point' hypothesis, which states that the Q 10 value above freeze–thaw critical point is much higher than that below it (16.0 vs. 3.5), and this was probably regulated by the abrupt change in soil water availability during the soil-freezing process. Our findings suggest interactive effects of multiple environmental factors on winter soil respiration and recommend adopting the freeze–thaw critical point to model soil respiration in a changing winter climate.

Aims Root and heterotrophic respiration may respond differently to environmental variability, but little evidence is available from large-scale observations. Here we aimed to examine variations of root and heterotrophic respiration across broad geographic, climatic, soil and biotic gradients.
Methods We conducted a synthesis of 59 field measurements on root and heterotrophic respiration across China's forests.
Important findings Root and heterotrophic respiration varied differently with forest types, of which evergreen broadleaf forest was significantly different from those in other forest types on heterotrophic respiration but without statistically significant differences on root respiration. The results also indicated that root and heterotrophic respiration exhibited similar trends along gradients of precipitation, soil organic carbon and satellite-indicated vegetation growth. However, they exhibited different relationships with temperature: root respiration exhibited bimodal patterns along the temperature gradient, while heterotrophic respiration increased monotonically with temperature. Moreover, they showed different relationships with MOD17 GPP, with increasing trend observed for root respiration whereas insignificant change for heterotrophic respiration. In addition, root and heterotrophic respiration exhibited different changes along the age sequence, with insignificant change for root respiration and decreasing trend for heterotrophic respiration. Overall, these results suggest that root and heterotrophic respiration may respond differently to environmental variability. Our findings could advance our understanding on the different environmental controls of root and heterotrophic respiration and also improve our ability to predict soil CO₂ flux under a changing environment.

Aims Understanding carbon (C) and nitrogen (N) dynamics and their dependence on the stand density of an even-aged, mature forest provides knowledge that is important for forest management. This study investigated the differences in ecosystem total C and N storage and flux between a low-density stand (LD) and a high-density stand (HD) and examined the effects of stand density on aboveground net primary productivity (ANPP), total belowground C allocation (TBCA) and net ecosystem production (NEP) in a naturally regenerated, 65- to 75-year-old Pinus densiflora S. et Z. forest.
Methods LD (450 trees ha^-1) and HD (842 trees ha^-1) were established in an even-aged, mature P. densiflora forest in September 2006. The forest had been naturally regenerated following harvesting, and the stand density was naturally maintained without any artificial management such as thinning. The diameter at breast height (DBH ≥ 5.0cm) of all live stems within the stands was measured yearly from 2007 to 2011. To compare C and N storage and fluxes in LD and HD, C and N pools in aboveground and belowground biomass, the forest floor, coarse woody debris (CWD) and soil; soil CO₂ efflux (R S); autotrophic respiration (R A); litter production; and soil N availability were measured. Further, ANPP, TBCA and NEP were estimated from plot-based measurement data.
Important findings Ecosystem C (Mg C ha^-1) and N (Mg N ha^-1) storage was, respectively, 173.0±7.3 (mean ± SE) and 4.69±0.30 for LD and 162±11.8 and 4.08±0.18 for HD. There were no significant differences in C and N storage in the ecosystem components, except for soils, between the two stands. In contrast, there were significant differences in aboveground ANPP and TBCA between the two stands (P < 0.05). Litterfall, biomass increment and R S were major C flux components with values of, respectively, 3.89, 3.74 and 9.07 Mg C ha^-1 year^-1 in LD and 3.15, 2.94 and 7.06 Mg C ha^-1 year^-1 in HD. Biometric-based NEP (Mg C ha^-1 year^-1) was 4.18 in LD and 5.50 in HD. Although the even-aged, mature P. densiflora forest had similar C and N allocation patterns, it showed different C and N dynamics depending on stand density. The results of the current study will be useful for elucidating the effects of stand density on C and N storage and fluxes, which are important issues in managing natural mature forest ecosystems.

Aims This study aimed to develop radial growth models and to predict the potential spatial distribution of Pinus densiflora (Japanese red pine) and Quercus spp. (Oaks) in South Korea, considering topographic and climatic factors.
Methods We used a dataset of diameter at breast height and radial growth estimates of individual trees, topographic and climatic factors in systematic sample plots distributed over the whole of South Korea. On the basis that radial growth is attributed primarily to tree age, we developed a radial growth model employing tree age as an explanatory variable. We estimated standard growth (SG), defined as radial growth of the tree at age 30, to eliminate the influence of tree age on radial growth. In addition, SG estimates including the Topographic Wetness Index, temperature and precipitation were calculated by the Generalized Additive Model.
Important findings As a result of variogram analysis of SG, we found spatial autocorrelation between SG, topographic and climatic factors. Incremental temperature had negative impacts on radial growth of P. densiflora and positive impacts on that of Quercus spp. Precipitation was associated with positive effects on both tree species. Based on the model, we found that radial growth of P. densiflora would be more vulnerable than that of Quercus spp. to climatic factors. Through simulation with the radial growth model, it was predicted that P. densiflora stands would be gradually replaced with Quercus spp. stands in eastern coastal and southern regions of South Korea in the future. The models developed in this study will be helpful for understanding the impact of climatic factors on tree growth and for predicting changes in distribution of P. densiflora and Quercus spp. due to climate change in South Korea.

Aims Understanding of the ecophysiological dynamics of forest canopy photosynthesis and its spatial and temporal scaling is crucial for revealing ecological response to climate change. Combined observations and analyses of plant ecophysiology and optical remote sensing would enable us to achieve these studies. In order to examine the utility of spectral vegetation indices (VIs) for assessing ecosystem-level photosynthesis, we investigated the relationships between canopy-scale photosynthetic productivity and canopy spectral reflectance over seasons for 5 years in a cool, temperate deciduous broadleaf forest at 'Takayama' super site in central Japan.
Methods Daily photosynthetic capacity was assessed by in situ canopy leaf area index (LAI), (LAI × V cmax [single-leaf photosynthetic capacity]), and the daily maximum rate of gross primary production (GPP max) was estimated by an ecosystem carbon cycle model. We examined five VIs: normalized difference vegetation index (NDVI), enhanced vegetation index (EVI), green–red vegetation index (GRVI), chlorophyll index (CI) and canopy chlorophyll index (CCI), which were obtained by the in situ measurements of canopy spectral reflectance.
Important findings Our in situ observation of leaf and canopy characteristics, which were analyzed by an ecosystem carbon cycling model, revealed that their phenological changes are responsible for seasonal and interannual variations in canopy photosynthesis. Significant correlations were found between the five VIs and canopy photosynthetic capacity over the seasons and years; four of the VIs showed hysteresis-type relationships and only CCI showed rather linear relationship. Among the VIs examined, we applied EVI–GPP max relationship to EVI data obtained by Moderate Resolution Imaging Spectroradiometer to estimate the temporal and spatial variation in GPP max over central Japan. Our findings would improve the accuracy of satellite-based estimate of forest photosynthetic productivity in fine spatial and temporal resolutions, which are necessary for detecting any response of terrestrial ecosystem to meteorological fluctuations.

Aims (i) To explore variations in nutrient resorption of woody plants and their relationship with nutrient limitation and (ii) to identify the factors that control these variations in forests of eastern China.
Methods We measured nitrogen (N) and phosphorus (P) concentrations in both green and senesced leaves of 172 woody species at 10 forest sites across eastern China. We compared the nutrient resorption proficiency (NuRP) and efficiency (NuRE) of N and P in plant leaves for different functional groups; we further investigated the latitudinal and altitudinal variations in NuRP and NuRE and the impacts of climate, soil and plant types on leaf nutrient resorptions.
Important findings On average, the leaf N resorption proficiency (NRP) and P resorption proficiency (PRP) of woody plants in eastern China were 11.1mg g ? 1 and 0.65 mg g ? 1, respectively; and the corresponding N resorption efficiency (NRE) and P resorption efficiency (PRE) were 49.1% and 51.0%, respectively. Angiosperms have higher NRP (are less proficient) values and lower NRE and PRE values than gymnosperms, but there are no significant differences in NRP, PRP and PRE values between species with different leaf habits (evergreen vs. deciduous angiosperms). Trees have higher NRE and PRE than shrubs. Significant geographical patterns of plant nutrient resorption exist in forests of eastern China. In general, NRP and PRE decrease and PRP and NRE increase with increasing latitude/altitude for all woody species and for the different plant groups. Plant functional groups show more controls than environmental factors (climate and soil) on the N resorption traits (NRP and NRE), while site-related variables present more controls than plant types on PRP and PRE. NRP increases and PRP and NRE decrease significantly with increasing temperature and precipitation for the overall plants and for most groups, except that significant PRE–climate relationship holds for only evergreen angiosperms. Leaf nutrient resorption did not show consistent responses in relation to soil total N and P stoichiometry, probably because the resorption process is regulated by the relative costs of drawing nutrients from soil versus from senescing leaves. These results support our hypothesis that plants growing in P-limited habitats (low latitudes/altitudes or areas with high precipitation/temperature) should have lower PRP and higher PRE, compared with their counterparts in relatively N-limited places (high latitudes/altitudes or areas with low precipitation/temperature). Our findings can improve the understanding of variations in N and P resorption and their responses to global change, and thus facilitate to incorporate these nutrient resorption processes into future biogeochemical models.

Aims Vast grasslands on the Tibetan Plateau are almost all under livestock grazing. It is unclear, however, what is the role that the grazing will play in carbon cycle of the grassland under future climate warming. We found in our previous study that experimental warming can shift the optimum temperature of saturated photosynthetic rate into higher temperature in alpine plants. In this study, we proposed and tested the hypothesis that livestock grazing would alter the warming effect on photosynthetic and respiration through changing physical environments of grassland plants.
Methods Experimental warming was carried by using an infrared heating system to increase the air temperature by 1.2 and 1.7°C during the day and night, respectively. The warming and ambient temperature treatments were crossed over to the two grazing treatments, grazing and un-grazed treatments, respectively. To assess the effects of grazing and warming, we examined photosynthesis, dark respiration, maximum rates of the photosynthetic electron transport (J max), RuBP carboxylation (V cmax) and temperature sensitivity of respiration Q 10 in Gentiana straminea, an alpine species widely distributed on the Tibetan grassland. Leaf morphological and chemical properties were also examined to understand the physiological responses.
Important findings 1) Light-saturated photosynthetic rate (A max) of G. straminea showed similar temperature optimum at around 16°C in plants from all experimental conditions. Experimental warming increased A max at all measuring temperatures from 10 to 25°C, but the positive effect of the warming occurred only in plants grown under the un-grazed conditions. Under the same measuring temperature, A max was significantly higher in plants from the grazed than the un-grazed condition. 2) There was significant crossing effect of warming and grazing on the temperature sensitivity (Q 10) of leaf dark respiration. Under the un-grazed condition, plants from the warming treatment showed lower respiration rate but similar Q 10 in comparison with plants from the ambient temperature treatment. However, under the grazed condition Q 10 was significantly lower in plants from the warming than the ambient treatment. 3) The results indicate that livestock grazing can alter the warming effects on leaf photosynthesis and temperature sensitivity of leaf dark respiration through changing physical environment of the grassland plants. The study suggests for the first time that grazing effects should be taken into account in predicting global warming effects on photosynthesis and respiration of plants in those grasslands with livestock grazing.

Anthropogenic nitrogen (N) emissions to atmosphere have increased dramatically in China since 1980s, and this increase has aroused great concerns on its ecological impacts on terrestrial ecosystems. Previous studies have showed that terrestrial ecosystems in China are acting as a large carbon (C) sink, but its potential in the future remains largely uncertain. So far little work on the impacts of the N deposition on C sequestration in China's terrestrial ecosystems has been assessed at a national scale. Aiming to assess and predict how ecological processes especially the C cycling respond to the increasing N deposition in China's forests, recently researchers from Peking University and their partners have established a manipulation experimental network on the ecological effects of the N deposition: Nutrient Enrichment Experiments in China's Forests Project (NEECF). The NEECF comprises 10 experiments at 7 sites located from north to south China, covering major zonal forest vegetation in eastern China from boreal forest in Greater Khingan Mountains to tropical forests in Hainan Island. This paper introduces the framework of the NEECF project and its potential policy implications.