Effects of tree size and organ age on variations in carbon,nitrogen,and phosphorus stoichiometry in Pinus koraiensis

Carbon(C),nitrogen(N),and phosphorus(P)are of fundamental importance for growth and nutrient dynamics within plant organs and deserve more attention at regional to global scales.However,our knowledge of how these nutrients vary with tree size,organ age,or root order at the individual level remains limited.We determined C,N,and P contents and their stoichiometric ratios(i.e.,nutrient traits)in needles,branches,and fine roots at different organ ages(0-3-year-old needles and branches)and root orders(1st-4th order roots)from 64 Pinus koraiensis of varying size(Diameter at breast height ranged from 0.3 to 100 cm)in northeast China.Soil factors were also measured.The results show that nutrient traits were regulated by tree size,organ age,or root order rather than soil factors.At a whole-plant level,nutrient traits decreased in needles and fine roots but increased in branches with tree size.At the organ level,age or root order had a negative effect on C,N,and P and a positive effect on stoichiometric ratios.Our results demonstrate that nutrient variations are closely related to organ-specific functions and ecophysiological processes at an individual level.It is suggested that the nutrient acquisition strategy by younger trees and organ fractions with higher nutrient content is for survival.Conversely,nutrient storage strategy in older trees and organ fractions are mainly for steady growth.Our results clarified the nutrient utilization strategies during tree and organ ontogeny and suggest that tree size and organ age or root order should be simultaneously considered to understand the complexities of nutrient variations.

Leaf nitrogen resorption is more important than litter nitrogen mineralization in mediating the diversity–productivity relationship along a nitrogen-limited temperate forest succession chronosequence

The resorption of nutrients by plants before litter fall and the mineralization of nutrients from plant litter by soil processes are both important pathways supporting primary productivity. While the positive relationship between plant biodiversity and primary productivity is widely accepted for natural ecosystems, the roles of nutrient resorption and mineralization in mediating that relationship remains largely unknown. Here, we quantified the relative importance of nitrogen(N) resorption and N mineralization in driving plant community N investment and the correlation between species diversity and community productivity along an N-limited successional chronosequence of the mixed broadleaved–Korean pine(Pinus koraiensis) forest in northeastern China. Leaf N resorption efficiency(NRE) at the community level increased significantly along the successional chronosequence,whereas litter N mineralization rate decreased significantly. Leaf NRE was more important than litter N mineralization rate in driving the diversity–productivity relationship. However, higher leaf NRE led to less N mineralization as succession progressed along the chronosequence. Our results highlight the importance of the N resorption pathway rather than the N mineralization pathway for forest N acquisition with community succession,and they provide mechanistic insights into the positive effects of biodiversity on ecosystem functioning. In future forest management practices, we recommend appropriate application of N fertilizer to mitigate the adverse effects of N-poor soil on seedling regeneration during late succession and thus maintain the sustainable development of temperate forest ecosystems.

Allometric models for leaf area and leaf mass predictions across different growing seasons of elm tree(Ulmus japonica)

Convenient and effective methods to determine seasonal changes in individual leaf area (LA) and leaf mass (LM) of plants are useful in research on plant physiology and forest ecology. However, practical methods for estimating LA and LM of elm (Ulmus japonica) leaves in different periods have rarely been reported. We collected sample elm leaves in June, July and September. Then, we developed allometric models relating LA, LM and leaf parameters, such as leaf length (L) and width (W) or the product of L and W (LW). Our objective was to find optimal allometric models for conveniently and effectively estimating LA and LM of elm leaves in different periods. LA and LM were significantly correlated with leaf parameters (P < 0.05), and allometric models with LW as an independent variable were best for estimating LA and LM in each period. A linear model was separately developed to predict LA of elm leaves in June, July and September, and it yielded high accuracies of 93, 96 and 96%, respectively. Similarly, a specific allometric model for predicting LM was developed separately in three periods, and the optimal model form in both June and July was a power model, but the linear model was optimal for September. The accuracies of the allometric models in predicting LM were 88, 83 and 84% for June, July and September, respectively. The error caused by ignoring seasonal variation of allometric models in predicting LA and LM in the three periods were 1-4 and 16-59%, respectively.

Effects of six years of simulated N deposition on gross soil N transformation rates in an old-growth temperate forest

Elevated atmospheric nitrogen(N) deposition has been detected in many regions of China, but its effects on soil N transformation in temperate forest ecosystems are not well known. We therefore simulated N deposition with four levels of N addition rate(N0, N30, N60, and N120) for6 years in an old-growth temperate forest in Xiaoxing’an Mountains in Northeastern China. We measured gross N transformation rates in the laboratory usingN tracing technology to explore the effects of N deposition on soil gross N transformations taking advantage of N deposition soils. No significant differences in gross soil N transformation rates were observed after 6 years of N deposition with various levels of N addition rate. For all N deposition soils, the gross NH~+ immobilization rates were consistently lower than the gross N mineralization rates,leading to net N mineralization. Nitrate(NO~-) was primarily produced via oxidation of NH~+(i.e., autotrophic nitrification), whereas oxidation of organic N(i.e., heterotrophic nitrification) was negligible. Differences between the quantity of ammonia-oxidizing bacteria and ammonia-oxidizing archaea were not significant for any treatment, which likely explains the lack of a significant effect on gross nitrification rates. Gross nitrification rates were much higher than the total NO~- consumption rates,resulting in a build-up of NO~-, which highlights the high risk of N losses via NO~- leaching or gaseous N emissions from soils. This response is opposite that of typical N-limited temperate forests suffering from N deposition,suggesting that the investigated old-growth temperate forest ecosystem is likely to approach N saturation.

Effects of ontogenetic stage and leaf age on leaf functional traits and the relationships between traits in Pinus koraiensis

Investigating the effects of ontogenetic stage and leaf age on leaf traits is important for understanding the utilization and distribution of resources in the process of plant growth.However,few studies have been conducted to show how traits and trait-trait relationships change across a range of ontogenetic stage and leaf age for evergreen coniferous species.We divided 67 Pinus koraiensis Sieb.et Zucc.of various sizes(0.3-100 cm diameter at breast height,DBH)into four ontogenetic stages,i.e.,young trees,middle-aged trees,mature trees and over-mature trees,and measured the leaf mass per area(LMA),leaf dry matter content(LDMC),and mass-based leaf nitrogen content(N)and phosphorus content(P)of each leaf age group for each sampled tree.One-way analysis of variance(ANOVA)was used to describe the variation in leaf traits by ontogenetic stage and leaf age.The standardized major axis method was used to explore the effects of ontogenetic stage and leaf age on trait-trait relationships.We found that LMA and LDMC increased significantly and N and P decreased significantly with increases in the ontogenetic stage and leaf age.Most trait-trait relationships were consistent with the leaf economic spectrum(LES)at a global scale.Among them,leaf N content and LDMC showed a significant negative correlation,leaf N and P contents showed a significant positive correlation,and the absolute value of the slopes of the trait-trait relationships showed a gradually increasing trend with an increasing ontogenetic stage.LMA and LDMC showed a significant positive correlation,and the slopes of the trait-trait relationships showed a gradually decreasing trend with leaf age.Additionally,there were no significant relationships between leaf N content and LMA in most groups,which is contrary to the expectation of the LES.Overall,in the early ontogenetic stages and leaf ages,the leaf traits tend to be related to a"low investment-quick returns"resource strategy.In contrast,in the late ontogenetic stages and leaf ages,they tend to be related to a"high investment-slow returns"resource strategy.Our results reflect the optimal allocation of resources in Pinus koraiensis according to its functional needs during tree and leaf ontogeny.

Stand development patterns of forest cover types in the natural forests of northern Baekdudaegan in South Korea

The purpose of this study was to classify current forest cover types,and to investigate stand development patterns for natural forests in six areas in northern Baekdudaegan,South Korea.Twenty-eight independent forest communities were aggregated into eight forest cover types by species composition in the overstory of each forest community.The forest cover types were of mixed mesophytic,‘‘others’ ’ deciduous,Quercus mongolica dominant,Q.mongolica pure,Pinus densiflora–Q.mongolica,P.densiflora,Betula ermanii,and Q.mongolica–P.koraiensis.The ecological information was organized by importance value and species diversity for each forest type.Based on the correlation between species diversity index and the abundance of Q.mongolica plus P.densiflora for corresponding forest cover types,we compared the developmental process and approximate successional pathway between each cover type.The P.densiflora forest cover type changes into the P.densiflora–Q.mongolica cover type,followed by the Q.mongolica dominant cover type through continuous invasion of the oak trees.Furthermore,the Q.mongolica pure cover type would spread toward the Q.mongolica dominant cover type with a mixture of various deciduous tree species.The Q.mongolica dominant cover type progresses through the other deciduous cover types to the mixed mesophytic cover type with diversified composition and structure.On the mid to lower slopes,with loamy soils and good moisture conditions,various deciduous forest types should progress,by ecological succession,toward the mixed mesophytic cover type without any further disturbance.

Stand biomass of Pinus sylvestris var. mongolica plantations benefits from high density monocultures in the boreal zone

Pinus sylvestris var.mongolica(P.sylvestris)plantations are extensively established in the boreal zone.Increasing stand biomass of these plantations can effectively enhance carbon stock,which is crucial for mitigating climate change.However,the current understanding of optimizing plantation strategies to maximize stand biomass is primarily derived from experiments in tropical and subtropical zones,which is difficult to extend to the boreal due to substantial climatic differences.Based on a comprehensive dataset from 1,076 sample plots of P.sylvestris plantations in the boreal zone of China,we evaluated the effects of tree species richness and stand density on tree height,diameter at breast height(DBH),and stand biomass to investigate the optimal plantation strategy.Furthermore,we examined how these effects changed with stand age and investigated their relative importance.We found that monocultures at a high stand density of 2,000–2,500​ha^(−1) were the optimal plantation strategy to maximize stand biomass(107.5​Mg·ha^(−1)),and this held true at almost all stand ages.Unfortunately,this strategy resulted in low species richness and small individual trees(10.6​m height and 9.8​cm DBH),thus presenting a trade-off.In addition,as stand age increased,the effect of tree species richness on stand biomass shifted from positive to negative,but the effect of stand density was always positive.Overall,stand age had the greatest effect on stand biomass,followed by stand density and then tree species richness.Our findings reveal a distinct plantation strategy for optimizing stand biomass of P.sylvestris plantations in the boreal zone.More importantly,this study highlights that(1)maximizing stand biomass in the boreal zone may compromise tree species richness;(2)net effects of tree species richness on stand biomass are not always positive,as negative selection effects offset positive complementary effects.

N limitation increases along a temperate forest succession:evidences from leaf stoichiometry and nutrient resorption

Forest productivity and carbon(C) sequestration largely depend on soil N and P availability.To date,however,the temporal variation of nutrient limitation along forest succession is still under debate.Leaf stoichiometry and nutrient resorption are important indicators for predicting nutrient limitation of plant growth.Here,we measured nitrogen(N)and phosphorus(P)concentrations in green leaves and leaf liter for all woody species at four stages of temperate forest succession,and analyzed how abiotic and biotic factors affect leaf stoichiometry and nutrient resorption along forest succession.At the individual scale,leaf N and P concentrations had a significant increase at the end of the succession,while no change in leaf N:P ratio was detected.Nitrogen resorption efficiency(NRE)increased significantly with succession,but P resorption efficiency(PRE)first increased and then decreased.Significant increases in NRE:PRE ratios only occurred at the end of the succession.Moreover,plant N cycling was less responsive to soil nutrient than P cycling.At the community scale,we found that leaf N and P concentrations first decreased and then increased along forest succession,which were mainly affected by Shannon-Wiener index and species richness.Leaf N:P ratio significantly varied with succession and was mainly determined by community-weighted mean diameter at breast height(DBH).NRE increased and was significantly influenced by species richness and DBH,while PRE was relatively stable along forest succession.Thus,the NRE:PRE ratios significantly increased,indicating that N limitation is exacerbated with the temperate forest succession.These results might reflect the intense interspecific competition for limiting resource in a higher biodiversity community.In conclusion,our findings highlight the importance of biotic factors in driving forest ecosystem nutrient cycling and provide valuable information for sustainable fertilizer management practices in China's temperate and boreal forests.

Functional traits influence plant survival depending on environmental contexts and life stages in an old-growth temperate forest

Aims Functional traits are usually used to predict plant demographic rates without considering environmental contexts.However,previous studies have consistently found that traits have low explanatory power for plant demographic rates.We hypothesized that accounting for environmental contexts instead of focusing on traits alone could improve our understanding of how traits influence plant demographic rates.Methods We used generalized linear mixed-effect models to analyse the effects of functional traits(related to leaf,stem,seed and whole plant),environmental gradients(soil nutrients,water and elevation)and their interactions on the survival dynamics of 14133 saplings and 3289 adults in a 9-ha old-growth temperate forest plot.Important Findings We found that environmental variables,neighbour crowding and traits alone(i.e.main effects)influenced plant survival.However,the effects of the latter two variables varied between saplings and adults.The trait–environment interactions influenced plant survival,such that resource conservative traits increased plant survival under harsh conditions but decreased survival under mild conditions.The elevational gradient was the most important environmental factor driving these effects in our plot.Our results support the hypothesis that functional traits influence plant survival depending on environmental contexts in local communities.These results also imply that one species with limited trait variation cannot occupy all environments,which can promote species diversity.

Plant size,environmental factors and functional traits jointly shape the stem radius growth rate in an evergreen coniferous species across ontogenetic stages

Aims Plant size,environmental conditions and functional traits are important for plant growth;however,it is less clear which combination of these factors is the most effective for predicting tree growth across ontogenetic stages.Methods We selected 65 individuals of an evergreen coniferous species,Pinus koraiensis,with diameters at breast height(DBH)from 0.3 to 100 cm in Northeast China.For each individual,we measured the stem radius growth rate(SRGR,µm/year)for the current year,environmental factors(light,soil nutrient and soil water)and functional traits(leaf,branch and root traits).Important Findings SRGR increased with DBH when the DBH was lower than 58 cm,whereas it decreased with DBH when the DBH was larger than 58 cm.Structural equation modeling analysis suggested that,when the DBH was 0–15 cm,plant size had a direct negative influence on SRGR and an indirect positive influence on SRGR due to the light intensity above the plant.Plant size had direct positive and negative effects when the DBH was 16–58 cm and 59–100 cm,respectively.When the DBH was larger than 15 cm,soil parameters were more important than light intensity for SRGR.The functional traits selected for use in the best model were changed from the specific leaf area and wood density to the root nitrogen concentration with increasing tree size.In summary,plant size,environmental factors and functional traits jointly shaped tree growth,and their relative influence varied with size,suggesting that the resources limiting tree growth may change from light to soil nutrient with increasing tree size.

Long-term(42 years)effect of thinning on soil CO_(2)emission in a mixed broadleaved-Korean pine(Pinus koraiensis)forest in Northeast China

Thinning is an important forest management practice that has great potential to influence regional soil carbon storage and dynamics.The present study measured soil respiration(RS,the efflux of CO2 emitted)and its two components(heterotrophic(RH)and autotrophic(RA)respiration)from soil 42 years after thinning in comparison to un-thinning(control).Autotrophic respiration was significantly greater in the thinning plot,approximately 44%higher compared to the control,while both RSand RHwere slightly,but not significantly,higher in the thinning plot.Higher fine root biomass might have contributed to the higher RAin the thinning plot.Both RSand RHshowed clear soil temperature-dependent seasonal patterns,whereas RAwas less responsive to changes in temperature,especially within one specific season.The annual and season-specific temperature sensitivities of RSand RHwere lower in the thinning plot,specifically during the mid-growing season.Furthermore,variations in the season-specific temperature sensitivity of RSand RHwere less intense in the thinning plot.We conclude that forest thinning can reduce the temperature sensitivity of RSand RHduring the mid-growing season and increase soil CO2 emission in the long term.

Factors that affect the assembly of ground-dwelling beetles at small scales in primary mixed broadleaved-Korean pine forests in north-east China

Disentangling the relative roles of environmental and spatial processes in community assembly is a major task of community ecology.It is necessary to uncover this question at multiple spatial scales;however,the relative importance of spatial and environmental processes on ground-dwelling beetle assembly at a small scale is still unclear.Based on two permanent plots(each 300 m)located in primary mixed broadleaved-Korean pine forests,the topographic,soil,and plant factors were collected,and the spatial variables(MEMs,distancebased Moran’s eigenvector maps)were calculated.A redundancy analysis(RDA)was used to evaluate the influence of topographic,soil,and plant variables on ground-dwelling beetle compositions.A variation partitioning analysis was used to quantify the relative contributions of environmental and spatial processes on the assembly of ground-dwelling beetles.The results of the RDA reported that the soil,plant,and topographic variables affected Staphylinidae and Silphidae beetle compositions in both plots.According to the results of variation partitioning,pure soil and plant variables were important for the assembly of Silphidae beetles in the LS plot.The contributions of pure topographic,soil,and plant variables were significantly lower than those of pure spatial variables.The contributions of pure spatial variables were significant for the assembly of Staphylinidae and Silphidae beetles in both plots.In addition,the relative importance of environmental and spatial processes was not significantly changed after including more environmental variables and the unexplained variations.Finally,this study suggests that both spatial and environmental variables are important for the assembly of ground-dwelling beetle communities,while pure spatial variables are more important than pure environmental variables at a small scale(300 m).