Basal rates of soil respiration are correlated with photosynthesis in a mixed temperate forest

Many field studies have demonstrated that soil temperature explains most of the temporal variation in soil respiration (SR). However, there is increasing evidence to suggest that SR is also influenced by current, or recent, photosynthate. Accordingly, seasonal changes in SR nominally attributed to temperature may, in part, be due to seasonality in photosynthesis. Within a mixed coniferous–deciduous temperate forest, we measured SR and used the process model SECRETS to test whether seasonal changes in photosynthesis influence seasonal differences in SR. Measurements were made in six adjacent plots (from pure evergreen to pure deciduous) that exhibited a gradient in the seasonality of photosynthesis. Within all six plots, we found strong correlations between the basal rate of SR (BR; defined as the SR at 10°C) and modeled photosynthesis (i.e. gross primary productivity; GPP). Moreover, we observed larger seasonal changes in BR in those plots that exhibited larger seasonal changes in photosynthesis, as compared with plots with smaller changes in photosynthesis. This is relevant because estimates of the Q₁₀ of SR (Q₁₀ is the relative change in a process rate per temperature change of 10°C) typically assume a constant BR. Our results therefore support the hypothesis that differences in the apparent Q₁₀ of SR (apparent Q₁₀=Q₁₀ derived from field measurements of SR and temperature) among studies may, in large part, be related to seasonal differences in photosynthesis. We suggest that variation in stand structure and species composition and, thus, in the photosynthetic signatures, induce different seasonal changes in BR via differences in the belowground supply of labile carbon. If these seasonal changes in BR are not properly accounted for, fitted apparent Q₁₀ values may not express the temperature response of respiratory processes in the soil.     

Four years of litter input manipulation changes soil microbial characteristics in a temperate mixed forest

Aboveground litter not only is an important source of nutrients to soil microbes but also regulates the microclimate in topsoil. How the changes in aboveground litter quantity would affect the microbial biogeochemical cycles is still unclear. Here we conducted a litter input manipulation experiment in a temperate mixed forest to investigate how different amounts of litter input affect soil organic carbon (SOC) and soil respiration via their regulation on soil microbes. We found that although neither SOC stock nor soil CO₂ efflux was affected by litter manipulation, soil microbial characteristics had responded after four years of litter addition or removal treatments. Microbial biomass carbon (MBC) in the O horizon was higher in litter addition plots than in litter removal plots as a result of the changed availability of labile C under litter treatments. Both double litter and no litter treatments changed microbial compositions, which was probably due to the increased soil pH in no litter treatment and the increased labile C in double litter treatment. The null change in soil respiration could be attributed to the offset between the negative effect of decreased substrate and the positive effect of increased temperature on soil respiration in litter removal plots. Due to the important role of soil microbes in carbon cycling, the altered microbial properties under litter manipulation treatments suggested the inevitable changes in biogeochemical cycling in the long run and call for long-term studies on SOC dynamics in the future.

     

Denitrification studies in a temperate forest catena soil

Summary Formation of ammonium during the reduction of nitrate under moderate and strict anaerobic incubation of two topsoils of a temperate forest catena, an acid mull and an anmoor was studied. In mull, both conditions of incubation caused reduction of nitrate and release of ammonium. The accumulation of ammonium continued even when there was no nitrate left hence indicating the formation of ammonium apparently through desamination of organic matter. Whereas, in anmoor neither any such formation of ammonium nor any significant reduction of nitrate was observed in the case of moderate anaerobic incubation. But under strict anaerobic incubation, progressive disappearance of nitrate was encountered from the beginning up to 30 days and this was accompanied by an increasing accumulation of ammonium in this soil. Yet this accumulation stopped when there was no nitrate left. Thus, the formation of ammonium is caused by the reduction of nitrate in anmoor.     

Variations of soil enzyme activities in a temperate forest soil

Soil enzyme activities (dehydrogenase, urease, phosphatase, and arylsulfatase) in a temperate forest soil were determined in relation to landscape position and seasons. Overstory of the area is dominated by Quercus mongolica, Kalopanax pictus, Carpicus cordata, and Acer pseudo-sieboldianum. The activities were measured in three patches, namely a north-facing backslope, a ridge, and a south-facing backslope in autumn and spring over 2 years. In addition, spatially more detailed analysis for phosphatase was conducted before and after litterfall production in six patches. Dehydrogenase, urease, phosphatase, and arylsulfatase activities varied 1.8–18.5 μg INT-formazan g⁻¹ h⁻¹, 45.4–347.0 μg NH₄ ⁺ g⁻¹ h⁻¹, 0.9–4.5 mmol pNP g⁻¹ h⁻¹, and 0.7–2.6 mmol pNP g⁻¹ h⁻¹, respectively. In general, higher enzyme activities were found in the northern aspect than in the southern aspect. This variation appears to be related to differences in chemical properties (e.g., Fe, Al, and Mg) of soil as well as distribution of leaf litter. Two patterns were discernible in relation to seasonal variations. Dehydrogenase and urease exhibited a positive correlation with mean air temperature, suggesting that temperature would be a major controlling variable for those enzymes. In contrast, higher activities were detected in autumn for phosphatase and arylsulfatase activities, which appeared to be closely related to litter production and distribution. Overall results of this study indicate that soil enzyme activities in a forest floor are influenced by several variables such as temperature, nutrient availability, and input of leaf litter, which are closely related to landscape position.     

Temperature-independent diel variation in soil respiration observed from a temperate deciduous forest

The response of soil respiration (R_s) to temperature depends largely on the temporal and spatial scales of interest and how other environmental factors interact with this response. They are often represented by empirical exponential equations in many ecosystem analyses because of the difficulties in separating covarying environmental responses and in observing below ground processes. The objective of this study was to quantify a soil temperature‐independent component in R_s by examining the diel variation of an R_s time series measured in a temperate deciduous forest located at Oak Ridge, TN, USA between March and December 2003. By fitting 2 hourly, continuous automatic chamber measurements of CO₂ efflux at the soil surface to a Q₁₀ function to obtain the temperature‐dependent respiration (R_t) and plotting the diel cycles of R_t, R_s, and their difference (R_i), we found that an obvious temperature‐independent component exists in R_s during the growing season. The diel cycle of this component has a distinct day/night pattern and agrees well with diel variations in photosynthetically active radiation (PAR) and air temperature. Elevated canopy CO₂ concentration resulted in similar patterns in the diel cycle of the temperature‐independent component but with different daily average rates in different stages of growing season. We speculate that photosynthesis of the stand is one of the main contributors to this temperature‐independent respiration component although more experiments are needed to draw a firm conclusion. We also found that despite its relatively small magnitude compared with the temperature‐dependent component, the diel variation in the temperature‐independent component can lead to significantly different estimates of the temperature sensitivity of soil respiration in the study forest. As a result, the common practice of using fitted temperature‐dependent function from night‐time measurements to extrapolate soil respiration during the daytime may underestimate daytime soil respiration.     

High abundance of Crenarchaeota in a temperate acidic forest soil

The objective of the study was to elucidate the depth distribution and community composition of Archaea in a temperate acidic forest soil. Numbers of Archaea and Bacteria were measured in the upper 18 cm of the soil, and soil cores were sampled on two separate occasions using quantitative PCR targeting 16S rRNA genes. Maximum numbers of Archaea were 0.6-3.8 x 10(8) 16S rRNA genes per gram of dry soil. Numbers of Bacteria were generally higher, but Archaea always accounted for a high percentage of the total gene numbers (12-38%). The archaeal community structure was analysed by the construction of clone libraries and by terminal restriction length polymorphism (T-RFLP) using the same Archaea-specific primers. With the reverse primer labelled, T-RFLP analysis led to the detection of four T-RFs. Three had lengths of 83, 185 and 218 bp and corresponded to uncultured Crenarchaeota. One (447 bp) was assigned to Thermoplasmales. Labelling of the forward primer allowed further separation of the T-RF into Crenarchaeota Group I.1c and Group I.1b, and indicated that Crenarchaeota of the Group I.1c were the predominant 16S rRNA genotype (相似文献   

Effects of experimental drought on soil respiration and radiocarbon efflux from a temperate forest soil

Soil moisture affects microbial decay of SOM and rhizosphere respiration (RR) in temperate forest soils, but isolating the response of soil respiration (SR) to summer drought and subsequent wetting is difficult because moisture changes are often confounded with temperature variation. We distinguished between temperature and moisture effects by simulation of prolonged soil droughts in a mixed deciduous forest at the Harvard Forest, Massachusetts. Roofs constructed over triplicate 5 × 5 m² plots excluded throughfall water during the summers of 2001 (168 mm) and 2002 (344 mm), while adjacent control plots received ambient throughfall and the same natural temperature regime. In 2003, throughfall was not excluded to assess the response of SR under natural weather conditions after two prolonged summer droughts. Throughfall exclusion significantly decreased mean SR rate by 53 mg C m^?2 h^?1 over 84 days in 2001, and by 68 mg C m^?2 h^?1 over 126 days in 2002, representing 10–30% of annual SR in this forest and 35–75% of annual net ecosystem exchange (NEE) of C. The differences in SR were best explained by differences in gravimetric water content in the Oi horizon (r²=0.69) and the Oe/Oa horizon (r²=0.60). Volumetric water content of the A horizon was not significantly affected by throughfall exclusion. The radiocarbon signature of soil CO₂ efflux and of CO₂ respired during incubations of O horizon, A horizon and living roots allowed partitioning of SR into contributions from young C substrate (including RR) and from decomposition of older SOM. RR (root respiration and microbial respiration of young substrates in the rhizosphere) made up 43–71% of the total C respired in the control plots and 41–80% in the exclusion plots, and tended to increase with drought. An exception to this trend was an interesting increase in CO₂ efflux of radiocarbon‐rich substrates during a period of abundant growth of mushrooms. Our results suggest that prolonged summer droughts decrease primarily heterotrophic respiration in the O horizon, which could cause increases in the storage of soil organic carbon in this forest. However, the C stored during two summers of simulated drought was only partly released as increased respiration during the following summer of natural throughfall. We do not know if this soil C sink during drought is transient or long lasting. In any case, differential decomposition of the O horizon caused by interannual variation of precipitation probably contributes significantly to observed interannual variation of NEE in temperate forests.     

Microbial consumption of atmospheric isoprene in a temperate forest soil

Isoprene (2-methyl-1,3 butadiene) is a low-molecular-weight hydrocarbon emitted in large quantities to the atmosphere by vegetation and plays a large role in regulating atmospheric chemistry. Until now, the atmosphere has been considered the only significant sink for isoprene. However, in this study we performed both in situ and in vitro experiments with soil from a temperate forest near Ithaca, N.Y., that indicate that the soil provides a sink for atmospheric isoprene and that the consumption of isoprene is carried out by microorganisms. Consumption occurred rapidly in field chambers (672.60 +/- 30.12 to 2,718.36 +/- 86.40 pmol gdw day) (gdw is grams [dry weight] of soil; values are means +/- standard deviations). Subsequent laboratory experiments confirmed that isoprene loss was due to biological processes: consumption was stopped by autoclaving the soil; consumption rates increased with repeated exposure to isoprene; and consumption showed a temperature response consistent with biological activity (with an optimum temperature of 30 degrees C). Isoprene consumption was diminished under low oxygen conditions (120 +/- 7.44 versus 528.36 +/- 7.68 pmol gdw day under ambient O(2) concentrations) and showed a strong relationship with soil moisture. Isoprene-degrading microorganisms were isolated from the site, and abundance was calculated as 5.8 x 10 +/- 3.2 x 10 cells gdw. Our results indicate that soil may provide a significant biological sink for atmospheric isoprene.     

Variation in soil respiration under the tree canopy in a temperate mixed forest,central China,under different soil water conditions

The forest canopy cover can directly and indirectly affect soil conditions and hence soil carbon emission through soil respiration. Little is known, however, on the effects of canopy cover on soil respiration under the canopy of different tree species and soil water conditions. We have examined the variation in soil respiration at different soil water conditions (dry <10 %, wet >20 %, v/v) under different tree canopy covers in comparison with the canopy interspace in a temperate coniferous (Pinus armandii Franch) and broadleaved (Quercus aliena var. acuteserrata) mixed forest in central China. The results show that soil respiration measured under tree canopy cover varied with canopy size and soil water content. Soil respiration under small-sized canopies of P. armandii (PS) was higher than that under large-sized (PL) canopies, but the difference was only significant under the dry soil condition. However, soil respiration under large-sized canopies of Q. aliena (QL) was significantly greater than that under small-sized (QS) canopies under both dry and wet soil conditions. The difference in soil respiration between differently sized canopies of Q. aliena (33.5–35.8 %) was significantly greater than that between differently sized canopies of P. armandii (2.4–8.1 %). Differences in soil respiration between inter-plant gaps and under QS canopies in both the dry and wet soil conditions were significant. Significant increases in soil respiration (9.7–32.2 %) during the transition from dry to wet conditions were found regardless of canopy size, but the increase of soil respiration was significantly lower under P. armandii canopies (9.7–17.7 %) than under Q. aliena canopies (25.9–31.5 %). Our findings that the canopy cover of different tree species influences soil respiration under different soil moisture conditions could provide useful information for parameterizing and/or calibrating carbon flux models, especially for spatially explicit carbon models.     

Fine root mass in relation to soil N supply in a cool temperate forest

Soil inorganic nitrogen supply and fine root mass in the top layers of mineral soil (0–5 and 5–10cm) were investigated at upper and lower sites of a cool temperate forest where Fagus crenata and Quercus crispula dominate. At both sites, soil inorganic nitrogen supply was greatest in the 0–5cm layer. The predominant forms of soil inorganic nitrogen supply were NH₄⁺-N at the upper site and NO₃^–-N at the lower site. Fine roots were concentrated in the 0–5cm layer at the upper site, but not at the lower site. The form of supplied soil inorganic nitrogen supply can be important in determining the vertical distribution of fine roots.     

Nitrogen cycling and nitrogen saturation in temperate forest ecosystems

The last decade has seen a dramatic shift in the focus of nitrogen cycling research in forest ecosystems. Concerns over nitrogen deficiencies and effects of removal in harvest have given way to concerns over excess nitrogen availability and the potential for forest decline and surface water pollution. Driving this paradigm shift is the increase in atmospheric deposition of nitrogen to forests due to industrial and agricultural activity. At the core of the new paradigm is the concept of 'nitrogen saturation' of forest ecosystems. The purpose of this review is to synthesize recent advances in research relating to nitrogen deposition effects on temperate zone forest ecosystems, and the further effects of nitrogen saturation on environmental quality.     

Contribution of litter layer to soil greenhouse gas emissions in a temperate beech forest

Background and aims

The litter layer is a major source of CO₂, and it also influences soil-atmosphere exchange of N₂O and CH₄. So far, it is not clear how much of soil greenhouse gas (GHG) emission derives from the litter layer itself or is litter-induced. The present study investigates how the litter layer controls soil GHG fluxes and microbial decomposer communities in a temperate beech forest.

Methods

We removed the litter layer in an Austrian beech forest and studied responses of soil CO₂, CH₄ and N₂O fluxes and the microbial community via phospholipid fatty acids (PLFA). Soil GHG fluxes were determined with static chambers on 22 occasions from July 2012 to February 2013, and soil samples collected at 8 sampling events.

Results

Litter removal reduced CO₂ emissions by 30 % and increased temperature sensitivity (Q₁₀) of CO₂ fluxes. Diffusion of CH₄ into soil was facilitated by litter removal and CH₄ uptake increased by 16 %. This effect was strongest in autumn and winter when soil moisture was high. Soils without litter turned from net N₂O sources to slight N₂O sinks because N₂O emissions peaked after rain events in summer and autumn, which was not the case in litter-removal plots. Microbial composition was only transiently affected by litter removal but strongly influenced by seasonality.

Conclusions

Litter layers must be considered in calculating forest GHG budgets, and their influence on temperature sensitivity of soil GHG fluxes taken into account for future climate scenarios.

     

Soil water content and temperature as independent or confounded factors controlling soil respiration in a temperate mixed hardwood forest

Variation in soil temperature can account for most of the seasonal and diel variation in soil CO₂ efflux, but the temperature effect is not always consistent, and other factors such as soil water content are known to influence soil respiration. The objectives of this research were to study the spatial and temporal variation in soil respiration in a temperate forested landscape and to evaluate temperature and soil water functions as predictors of soil respiration. Soil CO₂ fluxes were measured with chambers throughout an annual cycle in six study areas at the Harvard Forest in central Massachusetts that include soil drainage classes from well drained to very poorly drained. The mean annual estimate of soil CO₂ efflux was 7.2 Mg ha^–1, but ranged from 5.3 in the swamp site to 8.5 in a well-drained site, indicating that landscape heterogeneity is related to soil drainage class. An exponential function relating CO₂ fluxes to soil temperature accounted for 80% of the seasonal variation in fluxes across all sites (Q₁₀ = 3.9), but the Q₁₀ ranged from 3.4 to 5.6 for the individual study sites. A significant drought in 1995 caused rapid declines in soil respiration rates in August and September in five of the six sites (a swamp site was the exception). This decline in CO₂ fluxes correlated exponentially with decreasing soil matric potential, indicating a mechanistic effect of drought stress. At moderate to high water contents, however, soil water content was negatively correlated with soil temperature, which precluded distinguishing between the effects of these two confounded factors on CO₂ flux. Occurrence of high Q₁₀ values and variation in Q₁₀ values among sites may be related to: (i) confounding effects of high soil water content; (ii) seasonal and diel patterns in root respiration and turnover of fine roots that are linked to above ground phenology and metabolism; and (iii) variation in the depth where CO₂ is produced. The Q₁₀ function can yield reasonably good predictions of annual fluxes of CO₂, but it is a simplification that masks responses of root and microbial processes to variation in temperature and water content throughout the soil.     

Increased nitrate availability in the soil of a mixed mature temperate forest subjected to elevated CO2 concentration (canopy FACE)

In a mature temperate forest in Hofstetten, Switzerland, deciduous tree canopies were subjected to a free‐air CO₂ enrichment (FACE) for a period of 8 years. The effect of this treatment on the availability of nitrogen (N) in the soil was assessed along three transects across the experimental area, one under Fagus sylvatica, one under Quercus robur and Q. petraea and one under Carpinus betulus. Nitrate, ammonium and dissolved organic N (DON) were analysed in soil solution obtained with suction cups. Nitrate and ammonium were also captured in buried ion‐exchange resin bags. These parameters were related to the local intensity of the FACE treatment as measured from the ¹³C depletion of dissolved inorganic carbon in the soil solution. Over the 8 years of experiment, the CO₂ enrichment reduced DON concentrations, did not affect ammonium, but induced higher nitrate concentrations, both in soil solution and resin bags. In the nitrate captured in the resin bags, the natural abundance of the isotope ¹⁵N increased strongly. This indicates that the CO₂ enrichment accelerated net nitrification, probably as an effect of the higher soil moisture resulting from the reduced transpiration of the CO₂‐enriched trees. It is also possible that N mineralization was enhanced by root exudates (priming effect) or that the uptake of inorganic N by these trees decreased slightly as the result of a reduced N demand for fine‐root growth. In this mature deciduous forest, we did not observe any progressive N limitation due to elevated atmospheric CO₂ concentrations; on the contrary, we observed an enhanced N availability over the 8 years of our measurements. This may, together with the global warming projected, exacerbate problems related to N saturation and nitrate leaching, although it is uncertain how long the observed trends will last in the future.     

Nitrogen oxide gas emissions from temperate forest soils receiving long-term nitrogen inputs

From spring 2000 through fall 2001, we measured nitric oxide (NO) and nitrous oxide (N₂O) fluxes in two temperate forest sites in Massachusetts, USA that have been treated since 1988 with different levels of nitrogen (N) to simulate elevated rates of atmospheric N deposition. Plots within a pine stand that were treated with either 50 or 150 kg N ha^?1 yr^?1 above background displayed consistently elevated NO fluxes (100–200 µg N m^?2 h^?1) compared to control plots, while only the higher N treatment plot within a mixed hardwood stand displayed similarly elevated NO fluxes. Annual NO emissions estimated from monthly sampling accounted for 3.0–3.7% of N inputs to the high‐N plots and 8.3% of inputs to the Pine low‐N plot. Nitrous oxide fluxes in the N‐treated plots were generally < 10% of NO fluxes. Net nitrification rates (NRs) and NO production rates measured in the laboratory displayed patterns that were consistent with field NO fluxes. Total N oxide gas flux was positively correlated with contemporaneous measurements of NR and concentration. Acetylene inhibited both nitrification and NO production, indicating that autotrophic nitrification was responsible for the elevated NO production. Soil pH was negatively correlated with N deposition rate. Low levels (3–11 µg N kg^?1) of nitrite () were detected in mineral soils from both sites. Kinetic models describing NO production as a function of the protonated form of (nitrous acid [HNO₂]) adequately described the mineral soil data. The results indicate that atmospheric deposition may generate losses of gaseous NO from forest soils by promoting nitrification, and that the response may vary significantly between forest types under similar climatic regimes. The lowering of pH resulting from nitrification and/or directly from deposition may also play a role by promoting reactions involving HNO₂.     

Fine-scale genetic structure of red deer (Cervus elaphus) in a French temperate forest

Despite the classic population genetic view of a population as a network of sub-populations consisting of randomly mating individuals, the mating system and dispersal patterns of social animals affect the distribution of genetic variation on a local scale. The spatially open, forest-dwelling red deer (Cervus elaphus) population at the Petite Pierre National Reserve in north-eastern France is culled annually, with the management aim of maximising the number of adult males in the population, and is a typical example of an exploited red deer population from continental Europe. Through a change in management policy, the number of adult males in the population has increased over time, leading to a reduction in variance of male reproductive success (Bonenfant et al., 2002). In this study, we investigate the fine-scale genetic structure of the population using 14 microsatellite loci and attempt to find evidence for a change in this genetic structure over time. DNA was extracted from bone powder obtained by drilling into antlers and mandibular condyles. DNA was successfully extracted from up to 30-year-old samples, but it was necessary to genotype samples in duplicate to obtain reliable genetic profiles. Our results point towards a pattern of fine-scale spatial structure amongst female red deer in the study area, but not amongst males, as would be expected for a typical mammalian system with male-biased dispersal and female philopatry. In addition, our results hint at a decrease in spatial genetic structure amongst females over time, which might be related to a change in management policy, but small sample size limited the robustness of this conclusion.     

Shifts in root dynamics along a hillslope in a mixed,mesic temperate forest

Plant and Soil - Complex topography, a common feature of forested areas, generates unique environmental gradients that may shape root dynamics in unexpected ways. Nevertheless, belowground studies...     

Complementarity in the use of nitrogen forms in a temperate broad-leaved mixed forest

Background: The complementary use of different forms of soil nitrogen (N) might lead to a higher productivity of mixed forests than monocultures, but convincing evidence for temperate mixed forests is scarce.

Aims: We searched for species differences in N uptake rates and the preference for NH₄⁺, NO₃^? or glycine among five temperate broad?leaved tree species (Acer pseudoplatanus, Carpinus betulus, Fagus sylvatica, Fraxinus excelsior, Tilia cordata) in a mature mixed stand.

Methods: ¹⁵N tracer was added to the soil and its accumulation in fine root biomass was analysed after 10 min, 1 h and 1 d.

Results: The estimated root uptake rates of the species were in the range of 5–46 µg N g^?1 root h^?1 for NH₄⁺, 6–86 µg N g^?1 h^?1 for NO₃^? and 4–29 µg N g^?1 h^?1 for glycine during the first hour after tracer application. Carpinus, Tilia and Acer tended to prefer NH₄⁺ over NO₃^?, while Fraxinus showed equal preference for both N forms and Fagus seemed to prefer NO₃^?.

Conclusions: The five co-existing tree species differed in uptake rates and partly in their N form preference, but complementarity in the use of different N forms seems to be of minor importance in this forest because tree species appear to be rather flexible in their N form use.     

Population dynamics of coniferous and broad-leaved trees in a Japanese temperate mixed forest

The growth and survival of coniferous and broad-leaved trees were followed over a 5-yr period in a temperate old-growth mixed forest in Japan, and dynamic features of the forest were studied in relation to the life history of the dominants, the coniferous Abies homolepis and the broad-leaved Fagus crenata. During this period, the gap formation rate was 31m² ha^?1yr^?1, the mortality of trees > 2m high was 1.7%/yr, and the rate of loss in basal area 1.4%/yr. These values were much higher than the recruitment, 0.3%/yr, and the total growth of surviving and new trees, 0.6%/yr, owing to the inhibition of regeneration by understorey dwarf bamboo (Sasa borealis). A transition matrix model based on DBH size classes predicts that the basal area of the forest will decrease by 14% in 50 yr, but that the DBH distribution of trees > 10 cm diameter will change little. Equilibrium DBH distributions assuming recruitment being equal to mortality, were quite different between broad-leaved and coniferous trees, reflecting different survivorship curves of the two dominants. The composition and structure of the forest may change depending on the pattern and frequency of disturbances, or episodic events, notably the synchronous death of Sasa borealis.