亚热带常绿林不同冠层小枝叶面积-叶生物量关系研究

叶面积与叶生物量的关系对于理解植物叶片的碳收益和投资权衡策略具有重要意义。收益递减假说认为植物的叶面积与叶生物量成显著异速生长关系,其异速生长指数<1.0,但该假说是否适用于不同生活型（常绿与落叶）亚热带木本植物不同冠层高度（上下冠层）当年生小枝的叶片仍不清楚。以江西亚热带常绿阔叶林的69种常绿与落叶木本植物当年生小枝上的叶为研究对象,采用标准化主轴回归估计（standardized major axis estimation,SMA）方法检验不同冠层高度和生活型叶面积与叶生物量的异速生长关系。结果显示：（1）当年生小枝叶生物量在不同冠层高度和生活型的植物中无显著差异（P>0.05）,叶面积在常绿和落叶植物中有显著差异（P<0.05）,常绿和落叶植物的比叶重存在显著差异（P<0.05）,而落叶植物的比叶重在不同冠层高度存在显著差异（P<0.05）,同一冠层,常绿植物比叶重显著高于落叶植物（P<0.05）;（2）69种植物的叶面积与叶生物量异速生长指数具有物种特异性,60.9%的物种叶面积与叶生物量呈等速生长关系;（3）不同冠层和生活型植物的叶面积与叶生物量呈等速生长关系,但其异速生长常数在不同冠层高度与生活型间存在差异。这些结果表明冠层高度和生活型未改变叶面积-生物量之间的等速生长关系,不支持"收益递减"假说。     

Above- and below-ground biomass relationships across 1534 forested communities

BACKGROUND AND AIMS: Prior work has shown that above- and below-ground dry biomass across individual plants scale in a near isometric manner across phyletically and ecologically diverse species. Allometric theory predicts that a similar isometric scaling relationship should hold true across diverse forest-types, regardless of vegetational composition. METHODS: To test this hypothesis, two compendia for forest-level above- and below-ground dry biomass per hectare (M(A) and M(R), respectively) were examined to test the hypothesis that M(A) vs. M(R) scales isometrically and in the same manner as reported for data from individual plants. Model Type II regression protocols were used to compare the numerical values of M(A) vs. M(R) scaling exponents (i.e. slopes of log-log linear relationships) for the combined data sets (n =1534), each of the two data sets, and data sorted into a total of 17 data subsets for community- and biome-types as well as communities dominated by angiosperms or conifers. KEY RESULTS: Among the 20 regressions examined, 15 had scaling exponents that were indistinguishable from that reported for M(A) vs. M(R) across individual plants. The isometric hypothesis could not be strictly rejected on statistical grounds; four of these 15 exponents had broad 95% confidence intervals resulting from small sample sizes. Significant variation was observed in the y-intercepts of the 20 regression curves, because of absolute differences in M(A) or M(R). CONCLUSIONS: The allometries of forest- and individual plant-level M(A) vs. M(R) relationships share strikingly similar scaling exponents, but differ because of considerable variation in y-intercepts. These results support prior allometric theory and provide boundary conditions for the scaling of M(A) and M(R).     

On the allometry of biomass partitioning and light harvesting for plants with leafless stems

Prior explicit allometric models are extended to predict the scaling relationship between the ability of plants with leafless stems to harvest sunlight H and total standing plant biomass M(T) (which equals the sum of standing stem and root biomass, M(S) and M(R)). Provided that H scales in a directly proportional manner (isometrically) with respect to either stem surface area (i.e.H proportional, variant SA(S) ) or total stem biomass (i.e. H proportional, variant M(S)), the allometric model presented here predicts that SA(S) proportional, variant M(T)(3/4) or M(S) proportional, variant M(T)(3/4), respectively. These alternative predictions are tested empirically using data for standing stem and root biomass gathered for the large columnar cactus species Pachycereus pringlei. Statistical comparisons between observed and predicted scaling relationships indicate that SA(S) proportional, variant M(T)(3/4), whereas M(S) proportional, variant M(T)(3/4) is mathematically inconsistent with the observation that stem biomass scales nearly isometrically with respect to root biomass. The contention that the H of leafless stems scales isometrically with respect to stem surface area is thus reasonable both theoretically and empirically.     

Invading Monotypic Stands of Phalaris arundinacea: A Test of Fire, Herbicide, and Woody and Herbaceous Native Plant Groups

Phalaris arundinacea L. is an aggressive species that can dominate wetlands by producing monotypic stands that suppress native vegetation. In this study invasion windows were created for native species in monotypic stands of P. arundinacea with either fire or herbicide. Three native species groups, herbaceous plants, herbaceous seeds, and woody shrubs, were planted into plots burned or treated with herbicide in the early spring. Fire did not create an effective invasion window for native species; there was no difference in P. arundinacea root and shoot biomass or cover between burned and control plots (p≥ 0.998). Herbicide treatment created an invasion window for native species by reducing P. arundinacea root and shoot biomass for two growing seasons, but that invasion window was fast closing by the end of the second growing season because P. arundinacea shoot biomass had nearly reached the shoot biomass levels in the control plots (p= 0.053). Transplant mortality, frost, and animal herbivory prevented the herbaceous species and woody seedlings from becoming fully established in the plots treated with herbicide during the first year of the experiment. Transplanted monocots had a greater survival than dicots. By the second growing season the herbaceous group had the greatest mean areal cover (5%), compared to the woody seedlings (3%) and seed group (0%). Long‐term monitoring of the plots will determine whether the herbaceous transplants will compete effectively with P. arundinacea and whether the woody species will survive, shade the P. arundinacea, and accelerate forest succession.     

Allometric scaling relationship between above‐ and below‐ground biomass within and across five woody seedlings

Allometric biomass allocation theory predicts that leaf biomass (M_L) scaled isometrically with stem (M_S) and root (M_R) biomass, and thus above‐ground biomass (leaf and stem) (M_A) and root (M_R) scaled nearly isometrically with below‐ground biomass (root) for tree seedlings across a wide diversity of taxa. Furthermore, prior studies also imply that scaling constant should vary with species. However, litter is known about whether such invariant isometric scaling exponents hold for intraspecific biomass allocation, and how variation in scaling constants influences the interspecific scaling relationship between above‐ and below‐ground biomass. Biomass data of seedlings from five evergreen species were examined to test scaling relationships among biomass components across and within species. Model Type II regression was used to compare the numerical values of scaling exponents and constants among leaf, stem, root, and above‐ to below‐ground biomass. The results indicated that M_L and M_S scaled in an isometric or a nearly isometric manner with M_R, as well as M_A to M_R for five woody species. Significant variation was observed in the Y‐intercepts of the biomass scaling curves, resulting in the divergence for intraspecific scaling and interspecific scaling relationships for M_L versus M_S and M_L versus M_R, but not for M_S versus M_R and M_A versus M_R. We conclude, therefore, that a nearly isometric scaling relationship of M_A versus M_R holds true within each of the studied woody species and across them irrespective the negative scaling relationship between leaf and stem.     

植物代谢速率与个体生物量关系研究进展

植物的各项生理生态功能（例如,呼吸、生长和繁殖）都与个体生物量成异速生长关系。West, Brown及Enquist基于分形网络结构理论所提出的WBE模型认为：植物的代谢（呼吸）速率正比于个体生物量的3/4次幂。然而,恒定的“3/4异速生长指数”与实测数据、植物生理生态学等研究之间存在矛盾,引发激烈的争论。论文分析了不同回归方法对代谢指数的影响,重点对植物代谢速率与个体生物量异速生长关系研究进展进行了综述,分析并得出了植物代谢指数在小个体时接近1.0,并随着生物量的增加而系统减小,且其密切依赖于氮含量的调控的结论。据此,提出了进一步深入研究植物代谢速率个体生物量关系需要解决的一些科学问题。     

Relationships among the Stem, Aboveground and Total Biomass across Chinese Forests

Forest biomass plays a key role in the global carbon cycle. In the present study, a general allometric model was derived to predict the relationships among the stem biomass Ms, aboveground biomass MA and total biomass MT, based on previously developed scaling relationships for leaf, stem and root standing biomass. The model predicted complex scaling exponents for MT and/or MA with respect to Ms. Because annual biomass accumulation in the stem, root and branch far exceeded the annual increase in standing leaf biomass, we can predict that MT ∝MA ∝ Ms as a simple result of the model. Although slight variations existed in different phyletic affiliations （i.e. conifers versus angiosperms）, empirical results using Model Type Ⅱ （reduced major axis） regression supported the model＇s predictions. The predictive formulas among stem, aboveground and total biomass were obtained using Model Type I （ordinary least squares） regression to estimate forest biomass. Given the low mean percentage prediction errors for aboveground （and total biomass） based on the stem biomass, the results provided a reasonable method to estimate the biomass of forests at the individual level, which was insensitive to the variation in local environmental conditions （e.g. precipitation, temperature, etc.）.     

The role of belowground competition and plastic biomass allocation in altering plant mass–density relationships

Metabolic scaling theory (MST) predicts a ‘universal scaling law’ for plant mass–density relationships, but empirical observations are more variable. Possible explanations of this variability include plasticity in biomass allocation between the above‐ and belowground compartment and different modes of competition, which can be asymmetric or symmetric. Although complex interactions of these factors are likely to occur, so far the majority of modelling and empirical studies has focussed on mono‐factorial explanations. We here present a generic individual‐based model, which allows exploring the plant mass–density relationship in realistic settings by representing plasticity of biomass allocation and different modes of competition in the above‐ and belowground compartment. Plants grew according to an ontogenetic growth model derived from MST. To evaluate the behavior of the simulated plants related to the allocation patterns and to validate model predictions, we conducted greenhouse experiments with tree seedlings. The model reproduced empirical patterns both at the individual and population level. Without belowground resource limitation, aboveground processes dominated and the slopes of mass–density relationships followed the predictions of MST. In contrast, resource limitation led to an increased allocation of biomass to belowground parts of the plants. The subsequent dominance of symmetric belowground competition caused significantly shallower slopes of the mass–density relationship, even though the growth of individual plants followed MST. We conclude that changes in biomass allocation induced by belowground resource limitation explain the deviations from the mass–density relationship predicted by MST. Taking into account the plasticity of biomass allocation and its linkage to the above‐ and belowground competition is critical for fully representing plant communities, in particular for correctly predicting their response of carbon storage and sequestration to changing environmental conditions.     

Drought effect on plant biomass allocation: A meta‐analysis

Drought is one of the abiotic stresses controlling plant function and ecological stability. In the context of climate change, drought is predicted to occur more frequently in the future. Despite numerous attempts to clarify the overall effects of drought stress on the growth and physiological processes of plants, a comprehensive evaluation on the impacts of drought stress on biomass allocation, especially on reproductive tissues, remains elusive. We conducted a meta‐analysis by synthesizing 164 published studies to elucidate patterns of plant biomass allocation in relation to drought stress. Results showed that drought significantly increased the fraction of root mass but decreased that of stem, leaf, and reproductive mass. Roots of herbaceous plants were more sensitive to drought than woody plants that reduced reproductive allocation more sharply than the former. Relative to herbaceous plants, drought had a more negative impact on leaf mass fraction of woody plants. Among the herbaceous plants, roots of annuals responded to drought stress more strongly than perennial herbs, but their reproductive allocation was less sensitive to drought than the perennial herbs. In addition, cultivated and wild plants seemed to respond to drought stress in a similar way. Drought stress did not change the scaling exponents of the allometric relationship between different plant tissues. These findings suggest that the allometric partitioning theory, rather than the optimal partitioning theory, better explains the drought‐induced changes in biomass allocation strategies.     

On the vegetative biomass partitioning of seed plant leaves, stems, and roots

A central goal of comparative life-history theory is to derive the general rules governing growth, metabolic allocation, and biomass partitioning. Here, we use allometric theory to predict the relationships among annual leaf, stem, and root growth rates (GL, GS, and GR, respectively) across a broad spectrum of seed plant species. Our model predicts isometric scaling relationships among all three organ growth rates: GL is proportional to GS is proportional to GR. It also provides a conceptual basis for understanding the differences in the absolute amounts of biomass allocated to construct the three organ types. Analyses of a large compendium of biomass production rates across diverse seed plant species provide strong statistical support for the predictions of the theory and indicate that reproductive investments may scale isometrically with respect to vegetative organ growth rates. The general rules governing biomass allocation as indexed by the scaling exponents for organ growth rates are remarkably indifferent to plant size and taxonomic affiliation. However, the allometric "constants" for these relationships differ numerically as a function of phenotypic features and local environmental conditions. Nonetheless, at the level of both inter- and intraspecific comparisons, the same proportional biomass allocation pattern holds across extant seed plant species.     

7种木本植物的分支指数与代谢指数

West、Brown和Enquist提出的植物分形网络模型(简称WBE模型)认为: 植物的分支指数(1/a, 1/b)决定植物的代谢指数, 当分支指数1/a、1/b分别为理论值2.0、3.0时, 代谢速率与个体大小的3/4次幂成正比, 但是恒定的3/4代谢指数并不能全面地反映植物的代谢情况。基于分支指数的协同变化, Price、Enquist和Savage对WBE模型进行扩展, 提出植物分支参数协同变化模型(简称PES模型)。该文借助于PES模型分析了7种木本植物的分支指数和代谢指数。结果表明: 物种间叶面积与叶生物量呈异速生长关系, 基于叶面积得到的分支指数1/a和代谢指数θ在物种间无显著差异, 基于叶生物量得到的分支指数1/a、1/b和代谢指数θ在物种间均存在显著差异, 但基于叶面积和叶生物量分别拟合出的整体分支指数1/a、1/b和代谢指数θ与理论值均无显著差异, 且用叶面积作为代谢速率的替代指标比用叶生物量分析得出的代谢指数与理论值更接近。今后研究应当关注植物叶面积与叶生物量的异速生长关系对植物代谢速率及相关功能特性的影响。     

Branching and metabolic exponents in seven woody plants

West、Brown和Enquist提出的植物分形网络模型(简称WBE模型)认为: 植物的分支指数(1/a, 1/b)决定植物的代谢指数, 当分支指数1/a、1/b分别为理论值2.0、3.0时, 代谢速率与个体大小的3/4次幂成正比, 但是恒定的3/4代谢指数并不能全面地反映植物的代谢情况。基于分支指数的协同变化, Price、Enquist和Savage对WBE模型进行扩展, 提出植物分支参数协同变化模型(简称PES模型)。该文借助于PES模型分析了7种木本植物的分支指数和代谢指数。结果表明: 物种间叶面积与叶生物量呈异速生长关系, 基于叶面积得到的分支指数1/a和代谢指数θ在物种间无显著差异, 基于叶生物量得到的分支指数1/a、1/b和代谢指数θ在物种间均存在显著差异, 但基于叶面积和叶生物量分别拟合出的整体分支指数1/a、1/b和代谢指数θ与理论值均无显著差异, 且用叶面积作为代谢速率的替代指标比用叶生物量分析得出的代谢指数与理论值更接近。今后研究应当关注植物叶面积与叶生物量的异速生长关系对植物代谢速率及相关功能特性的影响。     

黑河中游荒漠草地地上和地下生物量的分配格局

草地生态系统中地上和地下生物量的分配方式对于研究生态系统碳储量和碳循环有着重要的意义。为了解黑河中游荒漠草地的地上和地下生物量分配格局, 从群落和个体两个水平对黑河中游的地上和地下生物量进行了调查。结果表明: 群落水平上地上生物量介于3.2-559.2 g·m^-2之间, 地下生物量介于3.3-188.2 g·m^-2之间, 个体水平上地上生物量介于6.1-489.0 g·株^-1之间, 地下生物量介于2.4-244.2 g·株^-1之间, 群落水平上的根冠比(R/S)为0.10-2.49, 个体水平上为0.07-1.55, 地下生物量均小于地上生物量, 群落水平上R/S值大于个体水平。群落和个体水平地上和地下生物量的拟合斜率分别为1.1001和0.9913, 与1没有显著差异, 说明地上与地下生物量呈等速生长关系。群落和个体水平土壤表层0-20 cm和0-30 cm的根系生物量分别占全部根系生物量的89.81%、96.95%和81.42%、93.62%, 表明地下生物量主要集中在0-20 cm和0-30 cm土壤表层。     

上海辰山植物园不同生活型木本植物枝叶大小关系的比较

植物枝叶生长普遍存在显著的正相关关系，决定了植物构型塑造和生物量分配。以上海辰山植物园149种木本植物为对象，通过测定顶枝上70 cm长枝条的直径、叶面积及其生物量，比较相同生境条件下不同生活型木本植物的枝叶大小关系。结果表明，枝条截面积与总叶面积间呈异速生长关系（a=1.148 6，CI=1.000 6~1.302 3），枝条干重与叶干重间呈等速生长关系（a=1.054 2，CI=0.921 3~1.205 6），不同生活型均具有相同的斜率系数a。不同生活型的异速生长常量b（y轴截距）存在显著差异，相同枝条截面积下落叶乔木比常绿乔木和常绿灌木具有更大的叶面积，相同枝条干重下常绿乔木和落叶乔木比常绿灌木具有更大的叶干重。这可能与不同生活型木本植物水分竞争效率和叶建成成本差异有关。     

Scaling relationship between tree respiration rates and biomass

The WBE theory proposed by West, Brown and Enquist predicts that larger plant respiration rate, R, scales to the three-quarters power of body size, M. However, studies on the R versus M relationship for larger plants (i.e. trees larger than saplings) have not been reported. Published respiration rates of field-grown trees (saplings and larger trees) were examined to test this relationship. Our results showed that for larger trees, aboveground respiration rates R_A scaled as the 0.82-power of aboveground biomass M_A, and that total respiration rates R_T scaled as the 0.85-power of total biomass M_T, both of which significantly deviated from the three-quarters scaling law predicted by the WBE theory, and which agreed with 0.81–0.84-power scaling of biomass to respiration across the full range of measured tree sizes for an independent dataset reported by Reich et al. (Reich et al. 2006 Nature 439, 457–461). By contrast, R scaled nearly isometrically with M in saplings. We contend that the scaling exponent of plant metabolism is close to unity for saplings and decreases (but is significantly larger than three-quarters) as trees grow, implying that there is no universal metabolic scaling in plants.     

Estimation of biomass allocation and carbon density of Rhododendron simsii shrubland in the subtropical mountainous areas of China

Aims As an important potential carbon sink, shrubland ecosystem plays a vital role in global carbon balance and climate regulation. Our objectives were to derive appropriate regression models for shrub biomass estimation, and to reveal the biomass allocation pattern and carbon density in Rhododendron simsii shrubland.
Methods We conducted investigations in 27 plots, and developed biomass regression models for shrub species to estimate shrub biomass. The biomass of herb and litterfall were obtained through harvesting. Plant samples were collected from each plot to measure carbon content in different organs.
Important findings The results showed that the power and linear models were the most appropriate equation forms. The D and D²H (where D was the basal diameter (cm) and H was the shrub height (m)) were good predictors for organ biomass and total biomass of shrubs. All of the biomass models reached extremely significant level, and could be used to estimate shrub biomass with high accuracy. It was more difficult to predict leaf and annual branch biomass than stem biomass, because leaf and annual branch were susceptible to herbivores and inter-plant competition. The mean biomass of the shrub layer was 20.78 Mg·hm^-2, in which Rhododendron simsii and Symplocos paniculata biomass accounted for 93.63%. Influenced by both environment and species characteristics, the biomass of the shrub layer organs was in the order of stem > root > leaf > annual branch. The root:shoot ratio of the shrub layer was 0.32, which was less than other shrubs in subtropical regions. The relative higher aboveground biomass allocation reflected the adaptation of plants to the warm and humid environment for more photosynthesis. The mean total community biomass was 26.26 Mg·hm^-2, in which shrub layer, herb layer and litter layer accounted for 79.14%, 7.62% and 13.25%, respectively. Litter biomass was relatively high, which suggested that this community had high nutrient return. There were significant correlations among aboveground biomass, belowground biomass and total biomass of shrub layer and herb layer. The mean biomass carbon density of the community was 11.70 Mg·hm^-2 and the carbon content ratio was 44.55%. The carbon density was usually obtained using the conversion coefficient of 0.5 in previous studies, which could overestimate carbon density by 12.22%.     

Pattern and control of biomass allocation across global forest ecosystems

The underground part of a tree is an important carbon sink in forest ecosystems. Understanding biomass allocation between the below‐ and aboveground parts (root:shoot ratios) is necessary for estimation of the underground biomass and carbon pool. Nevertheless, large‐scale biomass allocation patterns and their control mechanisms are not well identified. In this study, a large database of global forests at the community level was compiled to investigate the root:shoot ratios and their responses to environmental factors. The results indicated that both the aboveground biomass (AGB) and belowground biomass (BGB) of the forests in China (medians 73.0 Mg/ha and 17.0 Mg/ha, respectively) were lower than those worldwide (medians 120.3 Mg/ha and 27.7 Mg/ha, respectively). The root:shoot ratios of the forests in China (median = 0.23), however, were not significantly different from other forests worldwide (median = 0.24). In general, the allocation of biomass between the belowground and aboveground parts was determined mainly by the inherent allometry of the plant but also by environmental factors. In this study, most correlations between root:shoot ratios and environmental factors (development parameter, climate, altitude, and soil) were weak but significant (p < .01). The allometric model agreed with the trends observed in this study and effectively estimated BGB based on AGB across the entire database.     

Bud composition, branching patterns and leaf phenology in cerrado woody species

BACKGROUND AND AIMS: Plants have complex mechanisms of aerial biomass exposition, which depend on bud composition, the period of the year in which shoot extension occurs, branching pattern, foliage persistence, herbivory and environmental conditions. METHODS: The influence of water availability and temperature on shoot growth, the bud composition, the leaf phenology, and the relationship between partial leaf fall and branching were evaluated over 3 years in Cerrado woody species Bauhinia rufa (BR), Leandra lacunosa (LL) and Miconia albicans (MA). KEY RESULTS: Deciduous BR preformed organs in buds and leaves flush synchronously at the transition from the dry to the wet season. The expansion time of leaves is <1 month. Main shoots (first-order axis, A1 shoots) extended over 30 d and they did not branch. BR budding and foliage unfolds were brought about independently of inter-annual rainfall variations. By contrast, in LL and MA evergreen species, the shoot extension rate and the neoformation of aerial organs depended on rainfall. Leaf emergence was continuous for 2-6 months and lamina expansion took place over 1-4 months. The leaf life span was 5-20 months and the main A1 shoot extension happened over 122-177 d. Both evergreen species allocated biomass to shoots, leaves or flowers continuously during the year, branching in the middle of the wet season to form second-order (A2 shoots) and third-order (A3 shoots) axis in LL and A2 shoots in MA. Partial shed of A1 shoot leaves would facilitate a higher branching intensity A2 shoot production in LL than in MA. MA presented a longer leaf life span, produced a lower percentage of A2 shoots but had a higher meristem persistence on A1 and A2 shoots than LL. CONCLUSIONS: It was possible to identify different patterns of aerial growth in Cerrado woody species defined by shoot-linked traits such as branching pattern, bud composition, meristem persistence and leaf phenology. These related traits must be considered over and above leaf deciduousness for searching functional guilds in a Cerrado woody community. For the first time a relationship between bud composition, shoot growth and leaf production pattern is found in savanna woody plants.     

Power and metabolic scope of bird flight: a phylogenetic analysis of biomechanical predictions

For flying animals aerodynamic theory predicts that mechanical power required to fly scales as P proportional, variant m (7/6) in a series of isometric birds, and that the flight metabolic scope (P/BMR; BMR is basal metabolic rate) scales as P (scope) proportional, variant m (5/12). I tested these predictions by using phylogenetic independent contrasts from a set of 20 bird species, where flight metabolic rate was measured during laboratory conditions (mainly in wind tunnels). The body mass scaling exponent for P was 0.90, significantly lower than the predicted 7/6. This is partially due to the fact that real birds show an allometric scaling of wing span, which reduces flight cost. P (scope) was estimated using direct measurements of BMR in combination with allometric equations. The body mass scaling of P (scope) ranged between 0.31 and 0.51 for three data sets, respectively, and none differed significantly from the prediction of 5/12. Body mass scaling exponents of P (scope) differed significantly from 0 in all cases, and so P (scope) showed a positive body mass scaling in birds in accordance with the prediction.