Species traits and shoot-root biomass allocation in 20 dry-grassland species

Aims A plant has a limited amount of resources at any time and it allocates them to different structures. In spite of the large number of previous studies on allocation patterns within single species, knowledge of general patterns in species allocation is still very limited. This is because each study was done in different conditions using different methodology, making generalization difficult. We investigate intraspecific above- versus below-ground biomass allocation among individuals across a spectrum of dry-grassland plant species at two different developmental stages and ask whether allocation is age- and species specific, and whether differences among species can be explained by their life-history traits and phylogeny.Methods We collected data on above- and below-ground biomass of seedlings and adult plants of 20 species from a common garden experiment. We analysed data on shoot–root biomass allocation allometrically and studied the relationship between the allometric exponents (slopes on log–log scale), species life-history traits and phylogenetic distances.Important findings We found isometric as well as allometric patterns of biomass allocation in the studied species. Seedlings and adult individuals of more than half of the species differed in their above- versus below-ground biomass allometric exponents. Seedlings and adult individuals of the remaining species differed in their allometric coefficients (intercepts). Annual species generally allocated proportionally more to above- than below-ground biomass as seedlings than as adults, whereas perennial species showed the opposite pattern. Plant life-history traits, such as plant life span, age of first flowering, month in which the species begin flowering and specific leaf area were much more important in explaining differences in shoot–root allometry among species than were phylogenetic relationships. This suggests that allocation patterns vary greatly among closely related species but can be predicted based on species life-history traits.     

Above- and Belowground Biomass Allocation in Shrub Biomes across the Northeast Tibetan Plateau

Biomass partitioning has been explored across various biomes. However, the strategies of allocation in plants still remain contentious. This study investigated allocation patterns of above- and belowground biomass at the community level, using biomass survey from the Tibetan Plateau. We explored above- and belowground biomass by conducting three consecutive sampling campaigns across shrub biomes on the northeast Tibetan Plateau during 2011–2013. We then documented the above-ground biomass (AGB), below-ground biomass (BGB) and root: shoot ratio (R/S) and the relationships between R/S and environment factors using data from 201 plots surveyed from 67 sites. We further examined relationships between above-ground and below-ground biomass across various shrub types. Our results indicated that the median values of AGB, BGB, and R/S in Tibetan shrub were 1102.55, 874.91 g m^-2, and 0.85, respectively. R/S showed significant trend with mean annual precipitation (MAP), while decreased with mean annual temperature (MAT). Reduced major axis analysis indicated that the slope of the log-log relationship between above- and belowground biomass revealed a significant difference from 1.0 over space, supporting the optimal hypothesis. Interestingly, the slopes of the allometric relationship between log AGB and log BGB differed significantly between alpine and desert shrub. Our findings supported the optimal theory of above- and belowground biomass partitioning in Tibetan shrub, while the isometric hypothesis for alpine shrub at the community level.     

Patterns of above- and belowground biomass allocation in China’s grasslands: Evidence from individual-level observations

Above- and belowground biomass allocation not only influences growth of individual plants, but also influences vegetation structures and functions, and consequently impacts soil carbon input as well as terrestrial ecosystem carbon cycling. However, due to sampling difficulties, a considerable amount of uncertainty remains about the root: shoot ratio (R/S), a key parameter for models of terrestrial ecosystem carbon cycling. We investigated biomass allocation patterns across a broad spatial scale. We collected data on individual plant biomass and systematically sampled along a transect across the temperate grasslands in Inner Mongolia as well as in the alpine grasslands on the Tibetan Plateau. Our results indicated that the median of R/S for herbaceous species was 0.78 in China’s grasslands as a whole. R/S was significantly higher in temperate grasslands than in alpine grasslands (0.84 vs. 0.65). The slope of the allometric relationship between above- and belowground biomass was steeper for temperate grasslands than for alpine. Our results did not support the hypothesis that aboveground biomass scales isometrically with belowground biomass. The R/S in China’s grasslands was not significantly correlated with mean annual temperature (MAT) or mean annual precipitation (MAP). Moreover, comparisons of our results with previous findings indicated a large difference between R/S data from individual plants and communities. This might be mainly caused by the underestimation of R/S at the individual level as a result of an inevitable loss of fine roots and the overestimation of R/S in community-level surveys due to grazing and difficulties in identifying dead roots. Our findings suggest that root biomass in grasslands tended to have been overestimated in previous reports of R/S.     

Modelling below- and above-ground biomass for non-woody and woody plants

BACKGROUND AND AIMS: Intraspecific relationships between below- and above-ground biomass (MB and MA, respectively) have been studied extensively to evaluate environmental effects on growth and development at the level of the individual plant. However, no current theoretical model for this relationship exists for broad interspecific trends. The aims of this paper are to provide a model and to test its predictions using a recently assembled, large database (1406 data entries for 257 species). METHODS: An allometric model was derived to predict the relationship between MB and MA for non-woody and woody plants based on previously developed scaling relationships for leaf, stem and root standing biomass and annual growth rates. The predictions of this model were tested by comparing the numerical values of predicted scaling exponents (the slopes of log-log regression curves) with those observed for the database. KEY RESULTS AND CONCLUSIONS: For non-woody plants and the juveniles of woody species, the model predicts an isometric scaling relationship (i.e. MB proportional, variant MA). For woody plants, a complex scaling function is predicted. But, for a particular set of biologically reasonable conditions, the model predicts MB proportional, variant MA across woody plants. These predictions accord reasonably well with observed statistical trends when non-woody and woody plants are studied separately (n=1061 and 345 data entries, respectively). Although the reliability of regression formulas to estimate MB based on MA measurements increased with increasing plant size, estimates of MB can be as much as two orders of magnitude off, even when using regression formulas with r2 >0.90 and F >53,000.     

Effects of Grazing on Above- vs. Below-Ground Biomass Allocation of Alpine Grasslands on the Northern Tibetan Plateau

Biomass allocation is an essential concept for understanding above- vs. below-ground functions and for predicting the dynamics of community structure and ecosystem service under ongoing climate change. There is rare available knowledge of grazing effects on biomass allocation in multiple zonal alpine grassland types along climatic gradients across the Northern Tibetan Plateau. We collected the peak above- and below-ground biomass (AGB and BGB) values at 106 pairs of well-matched grazed vs. fenced sites during summers of 2010–2013, of which 33 pairs were subject to meadow, 52 to steppe and 21 to desert-steppe. The aboveground net primary productivity (ANPP) was represented by the peak AGB while the belowground net primary productivity (BNPP) was estimated from ANPP, the ratio of living vs. dead BGB, and the root turnover rate. Two-ways analyses of variance (ANOVA) and paired samples comparisons with t-test were applied to examine the effects of pasture managements (PMS, i.e., grazed vs. fenced) and zonal grassland types on both ANPP and BNPP. Allometric and isometric allocation hypotheses were also tested between logarithmically transformed ANPP and BNPP using standardized major axis (SMA) analyses across grazed, fenced and overall sites. In our study, a high community-dependency was observed to support the allometric biomass allocation hypothesis, in association with decreased ANPP and a decreasing-to-increasing BNPP proportions with increasing aridity across the Northern Tibetan Plateau. Grazing vs. fencing seemed to have a trivial effect on ANPP compared to the overwhelming influence of different zonal grassland types. Vegetation links above- and below-ground ecological functions through integrated meta-population adaptive strategies to the increasing severity of habitat conditions. Therefore, more detailed studies on functional diversity are essentially to achieve conservation and sustainability goals under ongoing climatic warming and intensifying human influences.     

Isometric scaling of above- and below-ground biomass at the individual and community levels in the understorey of a sub-tropical forest

Background and Aims Empirical studies and allometric partitioning (AP) theory indicate that plant above-ground biomass (M_A) scales, on average, one-to-one (isometrically) with below-ground biomass (M_R) at the level of individual trees and at the level of entire forest communities. However, the ability of the AP theory to predict the biomass allocation patterns of understorey plants has not been established because most previous empirical tests have focused on canopy tree species or very large shrubs.Methods In order to test the AP theory further, 1586 understorey sub-tropical forest plants from 30 sites in south-east China were harvested and examined. The numerical values of the scaling exponents and normalization constants (i.e. slopes and y-intercepts, respectively) of log–log linear M_A vs. M_R relationships were determined for all individual plants, for each site, across the entire data set, and for data sorted into a total of 19 sub-sets of forest types and successional stages. Similar comparisons of M_A/M_R were also made.Key Results The data revealed that the mean M_A/M_R of understorey plants was 2·44 and 1·57 across all 1586 plants and for all communities, respectively, and M_A scaled nearly isometrically with respect to M_R, with scaling exponents of 1·01 for all individual plants and 0·99 for all communities. The scaling exponents did not differ significantly among different forest types or successional stages, but the normalization constants did, and were positively correlated with M_A/M_R and negatively correlated with scaling exponents across all 1586 plants.Conclusions The results support the AP theory’s prediction that M_A scales nearly one-to-one with M_R (i.e. M_A ∝ M_R ^≈1·0) and that plant biomass partitioning for individual plants and at the community level share a strikingly similar pattern, at least for the understorey plants examined in this study. Furthermore, variation in environmental conditions appears to affect the numerical values of normalization constants, but not the scaling exponents of the M_A vs. M_R relationship. This feature of the results suggests that plant size is the primary driver of the M_A vs. M_R biomass allocation pattern for understorey plants in sub-tropical forests.     

Genetic and Environmental Controls on the Establishment of the Invasive Grass, Phalaris arundinacea

Whether an exotic species becomes integrated into a community or aggressively takes it over depends upon many interacting factors. Using contextual analyses, we combined genetic data about an invasive plant with information about the neighboring species, the community, and the environment to determine what factors enable a genotype or species to invade. We transplanted 50 individuals of each of three clones of the invasive grass Phalaris arundinacea, reed canary grass, into 150 random locations within a Vermont pasture. For each individual, we recorded clonal identity, neighbor identity, community indices (species richness and species diversity), and an environmental variable (soil moisture). The response variables were survivorship, above-ground biomass, below-ground biomass and the ratio of above- to below-ground biomass. Clonal identity affected both survivorship and below-ground biomass. The fastest tillering clone had poor survivorship but survivors produced a large amount of below-ground biomass, making this clone more likely to successfully overwinter. Neighbor species affected above- and below-ground biomass. Reed canary grass produced more above- and below-ground biomass when Anthoxanthum odoratum, a common pasture grass species, was abundant. Community attributes also influenced growth. Although we expected diverse plots to repel the invasion, plants in the more diverse plots had higher amounts of below-ground biomass. Finally, environmental effects also influenced growth. Reed canary grass produced more above-ground biomass in wetter plots, confirming that it does well under wet conditions.     

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。     

Evidence for "diminishing returns" from the scaling of stem diameter and specific leaf area

Research indicates that increases in total leaf area (A(T)) may fail to keep pace with increases in total leaf mass (M(L)) across plants differing in size (e.g., as measured by stem diameter, D). This "diminishing returns" hypothesis predicts that the scaling exponent for A(T) vs. M(L) will be less than one and that the exponent for specific leaf mass (i.e., A(T) / M(L)) vs. D will be negative. These predictions were examined using data from 46 plants ranging between 0.125 cm ≤ D ≤ 0.485 m across 25 woody dicot species. Standardized major axis slopes were used to quantify scaling exponents and random effects models were used to quantify species and size effects on the numerical values of exponents. The exponents for A(T) vs. M(L) and A(T) / M(L) vs. D differed among species and different species groupings. In general, the exponent for A(T) vs. M(L) was less than one and the exponent for A(T) / M(L) vs. D was negative, as predicted. However, random effects models indicated that species effects overshadowed size effects, although size effects were statistically significant. The diminishing returns hypothesis therefore receives statistical support, i.e., although the numerical values of exponents are "species-dependent," they are less than unity, as predicted by theory.     

Reexamination of a canonical model for plant organ biomass partitioning

A previously proposed "canonical" model for the scaling relations among leaf, stem, and root biomass (M(L), M(S), and M(R), respectively) asserts that the proportional relations M(L) ∝ M(S)(3/4) ∝ M(R)(3/4) and M(S) ∝ M(R) hold across seed plant species. This model is scrutinized by determining whether the scaling relations between M(L), M(S), and M(R) vs. basal stem diameter D(S) and between M(L), M(S), and M(R) vs. plant height h are logically consistent with previously predicted scaling exponents. For example, if M(L) is observed to scale as the 2-power of D(S) and the model asserts that M(L) ∝ M(S)(3/4), then M(S) must scale as the 8/3-power of D(S) if the model is valid. Using a large data base for species with self-supporting stems, statistical support was found for most such comparisons between predicted and observed scaling relationships. However, this judgement is predicated on (1) the assertion that the scaling exponents for M(R) with respect to D(S) (or h) are numerically "deflated" due to a systematic underestimate of fine and small root biomass and (2) the stringent protocol used to calculate the 95% confidence intervals of scaling exponents, which favors rejection of the model. In light of these features, the "canonical" model is logically consistent with the new scaling relations reported here. Therefore, the model is judged valid within the context of this evaluation.     

羊草种群生物量分配动态模拟

基于4种密度羊草种群的温室模拟试验,分析了羊草种群地上生物量与地下生物量的动态变化．结果表明：羊草种群的地上生物量与地下生物量随生长期呈增加的趋势,且随密度的增加而增加,但不同密度间羊草地上生物量与地下生物量的相对生长速率没有显著差异;不同密度羊草种群的根冠比随生长进程而增加,但差异不显著,表明环境因子是影响羊草种群根冠比变化的主导因素;单株羊草及羊草种群的地上生物量和地下生物量间存在显著的幂函数关系,但幂函数指数与系数随羊草密度的变化而变化,表明资源竞争引起的微环境差异导致了羊草种群根冠比的动态变化,可为定量研究光合产物分配提供参考．     

Canonical rules for plant organ biomass partitioning and annual allocation

Here we review a general allometric model for the allometric relationships among standing leaf, stem, and root biomass (M(L), M(S), and M(R), respectively) and the exponents for the relationships among annual leaf, stem, and root biomass production or "growth rates" (G(L), G(S), and G(R), respectively). This model predicts that M(L) ∝ M(S)(3/4) ∝ M(R)(3/4) such that M(S) ∝ M(R) and that G(L) ∝ G(S) ∝ G(R). A large synoptic data set for standing plant organ biomass and organ biomass production spanning ten orders of magnitude in total plant body mass supports these predictions. Although the numerical values for the allometric "constants" governing these scaling relationships differ between angiosperms and conifers, across all species, standing leaf, stem, and root biomass, respectively, comprise 8%, 67%, and 25% of total plant biomass, whereas annual leaf, stem, and root biomass growth represent 30%, 57%, and 13% of total plant growth. Importantly, our analyses of large data sets confirm the existence of scaling exponents predicted by theory. These scaling "rules" emerge from simple biophysical mechanisms that hold across a remarkably broad spectrum of ecologically and phyletically divergent herbaceous and tree-sized monocot, dicot, and conifer species. As such, they are likely to extend into evolutionary history when tracheophytes with the stereotypical "leaf," "stem," and "root" body plan first appeared.     

Large‐scale pattern of biomass partitioning across China's grasslands

Aim To investigate large‐scale patterns of above‐ground and below‐ground biomass partitioning in grassland ecosystems and to test the isometric theory at the community level. Location Northern China, in diverse grassland types spanning temperate grasslands in arid and semi‐arid regions to alpine grasslands on the Tibetan Plateau. Methods We investigated above‐ground and below‐ground biomass in China's grasslands by conducting five consecutive sampling campaigns across the northern part of the country during 2001–05. We then documented the root : shoot ratio (R/S) and its relationship with climatic factors for China's grasslands. We further explored relationships between above‐ground and below‐ground biomass across different grassland types. Results Our results indicated that the overall R/S of China's grasslands was larger than the global average (6.3 vs. 3.7). The R/S for China's grasslands did not show any significant trend with either mean annual temperature or mean annual precipitation. Above‐ground biomass was nearly proportional to below‐ground biomass with a scaling exponent (the slope of log–log linear relationship between above‐ground and below‐ground biomass) of 1.02 across various grassland types. The slope did not differ significantly between temperate and alpine grasslands or between steppe and meadow. Main conclusions Our findings support the isometric theory of above‐ground and below‐ground biomass partitioning, and suggest that above‐ground biomass scales isometrically with below‐ground biomass at the community level.     

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。     

氮添加对中国陆地植被地上-地下生物量分配的影响

大气氮沉降水平持续升高导致的外源氮输入增加,强烈影响了陆地生态系统的碳循环。目前,已有大量报道证实了氮沉降升高对全球陆地植被固碳的积极影响。虽然之前大部分研究将这一结果归因于光合作用增强导致的地上生物量增加,但最近的研究发现长期氮添加对植物地下根系的影响也同样重要。归纳整理了181篇公开发表的我国野外模拟氮沉降试验结果,采用整合分析(Meta-analysis)方法,定量评估了氮添加对我国陆地植被地上-地下生物量分配的影响特征和不同生态系统类型及施氮方式之间的影响差异。通过分析地上-地下生物量分配对氮添加的响应差异来探究植被碳增益对长期大气氮沉降增加的潜在响应机制。结果表明,氮添加显著增强了我国陆地植被的光合作用及碳固存,且植物碳增益在不同生态系统类型及施氮制度间有所差异。植物叶片的氮含量显著增加,使得叶片碳氮比及凋落物碳氮比显著降低,但并未显著影响细根的碳氮比。氮添加总体上显著提高了植物的净光合速率,但降低了光合利用效率。地上生物量,凋落物产量和根生物量平均分别显著增加了38%,17%和18%,总体上植物地上部分对氮添加的响应程度比地下部分更高。然而,不同生态系统类型的地上-地下生物...     

古尔班通古特沙漠南缘3种生活型草本植物生物量分配及相关生长关系

为了解古尔班通古特沙漠不同生活型草本植物的生物量分配特征,通过取样调查当地34种草本植物的生物量研究表明：（1）采集到的每个物种样本的总生物量大多小于4 g,其中多数个体（大于80%）的地上与地下生物量分别在2和0.5 g以内,根冠比主要集中在0.25以内;（2）类短命植物向地下部分分配较多生物量,根冠比较大,集中分布在1.67左右;短命和一年生长营养期草本向地下部分配的生物量远小于类短命植物,两者的根冠比较小,主要集中在0.15左右;（3）短命植物地下与地上部生物量间为等速生长关系,而类短命植物和一年生长营养期草本植物地下与地上生物量间为异速生长关系,其中类短命植物随个体增大向地下部分配生物量的比例增多,而一年生长营养期草本则相反。综上所述,短命、类短命和一年生长营养期草本植物地下与地上生物量间具有不同的分配特征,与各自独特的生活史特征相一致。     

甘肃南部7种高寒杜鹃生物量分配的异速生长关系

生物量分配模式影响着植物个体生长和繁殖到整个群落的质量和能量流动等所有层次的功能, 揭示高寒灌丛的生物量分配模式不仅可以掌握植物的生活史策略, 而且对理解灌丛碳汇不确定性具有重要意义。该研究以甘肃南部高山-亚高山区的常绿灌丛——杜鹃(Rhododendron spp.)灌丛的7个典型种为对象, 采用全株收获法研究了不同物种个体水平上各器官生物量的分配比例和异速生长关系。结果表明: 7种高寒杜鹃根、茎、叶生物量的分配平均比例为35.57%、45.61%和18.83%, 各器官生物量分配比例的物种差异显著; 7种高寒杜鹃的叶与茎、叶与根、茎与根以及地上生物量与地下生物量之间既有异速生长关系, 也有等速生长关系, 异速生长指数不完全支持生态代谢理论和小个体等速生长理论的参考值; 各器官异速生长关系的物种差异显著。结合最优分配理论和异速生长理论能更好地解释陇南山地7种高寒杜鹃生物量的变异及适应机制。     

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.     

Biomass allocation and leaf stoichiometric characteristics in four desert herbaceous plants during different growth periods in the Gurbantünggüt Desert,China

荒漠草本植物是荒漠生态系统物种多样性的主体, 对其生物量分配及叶片化学计量特征随植物生长的变化规律的研究有助于深入了解荒漠草本植物生存策略和功能特征。该文选择古尔班通古特沙漠4种优势草本(2种短命植物, 2种一年生长营养期植物)为研究对象, 通过野外原位多时段取样, 对比研究了四者生物量分配、叶片N-P化学计量学随植物生长的变化特征, 以及二者之间的关系。结果表明, 在生物量累积过程中, 4种植物根冠比逐渐减小, 地上与地下生物量间的相关生长关系也发生变化, 其中琉苞菊(Hyalea pulchella)和角果藜(Ceratocarpus arenarius)的相关生长指数先增加后减小, 并趋于稳定, 而尖喙牻牛儿苗(Erodium oxyrrhynchum)和沙蓬(Agriophyllum squarrosum)的相关生长指数由小到大并趋于等速生长。琉苞菊叶片N、P含量呈逐渐增长趋势, 而另外3种植物呈下降趋势, 表明所研究的荒漠植物在生长过程中, 叶片N-P化学计量发生改变, 叶片化学计量特征与生物量指标的相关性较弱。