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。     

代谢异速生长理论及其在微生物生态学领域的应用

新陈代谢是生物的基本生理过程,影响生物在不同环境中参与物质循环和能量转化的过程.代谢速率作为生物体重要的生命过程指标,几乎影响所有的生物活性速率,且在很多研究中均表现出异速生长现象.所谓代谢异速是指生物体代谢速率与其个体大小(或质量)之间存在的幂函数关系.代谢异速生长理论的提出,从机制模型角度解释了代谢异速关系这一普遍存在的生命现象.该理论利用分形几何学及流体动力学等原理,从生物能量学角度阐释了异速生长规律的机理,证实了3/4权度指数的存在;但同时有研究表明,权度指数因环境因素等影响处于2/3-1范围之间而非定值.随着研究工作的深入,代谢异速生长理论研究从起初的宏观动植物领域拓展到了微生物领域,在研究微生物的代谢异速生长理论时,可将微生物的可操作分类单元(Operational taxonomic unit,OTU)或具有特定功能的功能群视为一个微生物个体,基于其遗传多样性和功能多样性特征进行表征,以便于将微生物群落多样性与其生态功能性联系起来,使该理论在微生物生态学领域得到有效的补充和完善.尽管细菌具有独特的生物学特性,但与宏观生物系统中观测到的现象表现出明显的一致性.有研究表明,3个农田土壤细菌基于遗传多样性的OTU数的平均周转率分别为0.71、0.80和0.84,介于2/3与1之间,可能与生物代谢异速指数有一定关联,为微生物代谢异速指数的研究提出了一个参考解决方案.鉴于微生物个体特征和生物学特性,在分析代谢速率与个体大小关系中,从微生物单位个体的定义、个体大小表征到计量单位的统一,仍需更多的理论支持.分析了代谢异速生长理论在微生物与生态系统功能关系研究中的可能应用,延伸了该理论的应用范围,并对尚待加强的研究问题进行了评述和展望.     

四种一年生荒漠植物构件形态与生物量间的异速生长关系

1年生草本植物是古尔班通古特沙漠夏秋季节草本层片的主要组成部分。选择4种藜科1年生草本植物(刺沙蓬、沙蓬、角果藜和对节刺)作为研究对象,对比分析了各物种构件形态特征、生物量分配以及它们之间的异速生长关系。结果表明:沙蓬个体最大,其次为刺沙蓬,角果藜和对节刺个体最小;对节刺具有最大的根冠比(R/S),其次是刺沙蓬,沙蓬和角果藜R/S最小。4种植物构件形态与生物量间均呈显著正相关,表现出强烈的协同变化趋势。R/S与绝大部分指标间呈显著负相关,表明随个体增大地下生物量分配比例逐渐减小。4种植物各构件形态、地上及地下生物量间大部分呈指数1.0的异速生长关系,但各物种间的异速生长指数绝大部分无显著差异且具有共同的异速生长指数。研究表明:尽管物种不同、个体大小迥异,但4种1年生植物构件特征间多具有相同的生长速率和协同变化特征,体现了1年生植物这一生活型内的不同物种对干旱荒漠环境的趋同适应。     

植物种群自疏过程中构件生物量与密度的关系

不论是在对植物种群自疏规律还是在对能量守衡法则的研究中,个体大小(M)大多针对植物地上部分生物量,地下部分和构件生物量及其动态十分重要又多被忽视。以1年生植物荞麦为材料研究了自疏种群地下部分生物量、包括地下部分的个体总生物量以及各构件生物量与密度的关系。结果表明:平均地上生物量和个体总生物量与密度的异速关系指数(γabove-ground和γindividual)分别为-1.293和-1.253,与-4/3无显著性差异(P>0.05),为-4/3自疏法则提供了有力证据;平均根生物量-密度异速指数γroot(-1.128)与-1无显著性差异(P>0.05),与最终产量恒定法则一致;平均茎生物量-密度异速指数γstem(-1.263)接近-4/3(P>0.05),平均叶生物量-密度异速指数γleaf(-1.524)接近-3/2(P>0.05),分别符合-4/3自疏法则与-3/2自疏法则;而繁殖生物量与密度的异速关系指数γreproductive(-2.005)显著小于-3/2、-4/3或-1(P<0.001)。因此,不存在一个对植物不同构件普适的生物量-密度之间的关系。光合产物在地上和地下构件的生物量分配格局以及构件生物量与地上生物量之间特异的异速生长关系导致不同构件具有不同的自疏指数。无论对于地上生物量还是个体总生物量,荞麦种群能量均守衡,而对于地下生物量,荞麦种群能量不守衡。     

浮游生物异速生长关系研究进展

生物个体的生物学特征(新陈代谢速率等)与个体大小(体积和生物量等)的依赖性关系称为异速生长关系, 这种关系经过尺度转换之后, 还影响着种群、群落和生态系统等。海洋环境较为复杂, 浮游生物的代谢速率不仅受到个体大小影响, 同时光照、温度和营养物质也是重要的影响因素, 因此3/4 规律只有在理想环境状况下才能实现。对于大多数浮游生物来讲, 其尺度因子并非单一的, 存在着多元尺度指数。由于单一和多元尺度指数都忽略了环境温度的影响, 生态学代谢理论在考虑个体大小的基础上, 加上温度这个参数后能够解释不同层次的生态学过程。在综述了异速生长关系发展历史的基础上, 着重探讨了该关系在浮游生物中的研究进展。     

密度对尖头叶藜生物量分配格局及异速生长的影响

植物器官指示植物不同的功能,而植物器官生物量分配比例的变化表征了植物对资源获取能力的调整。在植物生长发育过程中,植物各器官呈一种明显的异速生长规律。利用异速生长分析方法,通过模拟不同密度(16、44.4、100、400株/m~2)下尖头叶藜(Chenopodium acuminatum)的生长特性,研究密度对尖头叶藜器官生物量分配格局及异速生长的影响。结果表明,随密度增加,尖头叶藜地上和地下器官都存在不同程度的竞争:其中,根和主茎生物量分配增加,茎和地上生物量分配减少,而叶和繁殖生物量分配不随密度变化而变化。研究发现,尖头叶藜各器官间具有显著的异速生长关系:其中叶∶主茎、根∶地上部分、根∶茎、根∶主茎、繁殖器官∶地上部分及繁殖器官∶根生物量间的异速生长不随密度变化而变化,属于表观可塑性;而叶∶地上部分、叶∶根、叶∶茎、茎∶地上部分、主茎∶地上部分、繁殖器官∶茎、繁殖器官∶主茎生物量间具有极显著的异速生长关系,异速指数和个体大小显著受密度变化影响,属于真正可塑性,这表明密度能够影响尖头叶藜各器官的生长变化。尖头叶藜叶∶主茎、叶∶根及主茎∶地上部分生物量间的异速指数在D4-密度时与3/4差异不显著(P0.05),符合生态代谢理论,而在D1—D3密度时与3/4差异显著(P0.05),表明充分竞争的植株更符合代谢理论,而竞争不激烈的植株对资源的投入具有物种特异性。     

古尔班通古特沙漠中4种荒漠草本植物的生物量与化学计量特征

对于荒漠植物不同生长期生物量分配和化学计量比的研究有助于深入了解荒漠植物的功能结构,更好掌握环境对植物的生存影响。本研究选取古尔班通古特沙漠4种一年生荒漠草本植物沙蓬（Agriophyllum sqarrosum（L.）Moq.）、雾冰藜（Bassssia dasyphylla（Fisch.et Mey.）O.Kuntze）、角果藜（Ceratocarpus arenarius L.）和碱蓬（Suaeda glauca（Bge.）Bge.）对比研究了4种生物量分配与N、P化学计量学随植物生长的变化。结果显示：（1）4种荒漠植物生物量积累配过程中,根冠比随生长季的延长逐渐降低。地上、地上生物量相关生长关系表明,角果藜为等速→异速的变化过程,沙蓬从采样初期至末期的变化过程为异速→等速生长,而沙蓬和碱蓬的相关生长指数相反,分别为异速→等速→异速、等速→异速→等速的变化格局;（2）4种植物N、P含量随生长期的延长逐步降低。在整个生长季节4种植物的N与P含量的变化趋势均为相似。（3）植物N、P含量间达到正相关显著,除沙蓬和雾冰藜的N：P之外其余指标分别与植物的地上、地下生物量及总生物量间呈负相关显著,而根冠比、相关生长指数与化学计量特征间未达到显著水平,表明了二者较弱的相关性。研究表明,4种荒漠草本植物生物量与化学计量特征的相关性不大,说明化学计量比并不是影响植物生物量及生活策略的主要因素,而更多是受植物自身遗传特性的影响。同时也体现了荒漠草本植物在养分匮乏的条件下,形成了自身独特的生理生态特征,且具有相对稳定的适应特性。     

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。     

水分和种植密度对干热河谷车桑子生长性状及种内相互关系的影响

水分是干热河谷植物生长过程中最主要的限制因子,种植密度增加也会引起植物生长的资源限制,两者交互作用下植物生长性状及种内关系的变化特征还不清楚。以干热河谷优势植物——车桑子为研究对象,根据元谋干热河谷年均生长季降雨量设置3种水分梯度：高水分、中水分和低水分,同时在各水分梯度下设置4个种植密度：1、2、4、9株/盆,探究水分、种植密度及其交互作用对车桑子生长性状、生物量分配及种内相互作用的影响。结果表明：（1）低水分条件下,车桑子生长和水分生理受到抑制,但车桑子在较低的叶水势下依然能够保持较高的相对含水量;（2）干旱胁迫显著降低了车桑子总生物量和单株生物量,显著增加了枯叶生物量比例,低水分和中水分条件下,增加种植密度对总生物无显著影响;而高水分条件下,增加种植密度显著提高了车桑子总生物量;（3）低水分显著增加了茎、叶生物量的异速生长指数,将更多生物量分配到叶,而种植密度增加显著降低了茎、叶生物量的异速生长指数,增加了茎的生物量分配;（4）通过相对邻体效应的计算,各处理条件下,车桑子种内关系均表现为竞争作用,并且,这种竞争作用的强度随水分的减少和密度的增加而增加。在高密度条件下（9棵/盆）,增加水分不会减轻种内竞争作用。综上,水分和种植密度均会对车桑子个体的生理生长产生影响,在植被恢复过程中,应考虑水分和种植密度对车桑子个体产生的资源限制作用。     

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.     

WBE 模型及其在生态学中的应用：研究概述

介绍了WBE模型,综述了该模型在生态学中的应用进展。WBE模型,以及以该模型为基础的MTE模型,假设生物体为自相似分形网络结构,提出代谢速率和个体大小之间存在3/4指数关系,分别预测了从个体到生物圈多个尺度上的生物属性之间的异速生长关系,而且部分得到了验证。WBE模型的应用涵盖了个体组织生物量、年生长率,种群密度和生态系统单位面积产量、能量流动率等多个方面;即使在生物圈大尺度上,WBE模型也可用来预测试验中无法直接测量的特征变量的属性,如全球碳储量的估算等。至今,关于WBE和MTE模型仍然存在各种褒贬争论,讨论焦点主要集中于模型建立的前提假设以及权度指数的预测。今后的研究工作应规范试验技术和方法,考虑物种多样性和环境等因素的影响,提出符合各类生物的模型结构体系,使其具有更广泛的应用性和预测性。     

Lack of Evidence for 3/4 Scaling of Metabolism in Terrestrial Plants

Scaling, as the translation of information across spatial, temporal, and organizational scales, is essential to predictions and understanding in all sciences and has become a central issue in ecology. A large body of theoretical and empirical evidence concerning allometric scaling in terrestrial individual plants and plant communities has been constructed around the fractal volume-filling theory of West, Brown, and Enquist （the WBE model）. One of the most thought-provoking findings has been that the metabolic rates of plants, like those of animals, scale with their size as a 3/4 power law. The earliest, single most-important study cited in support of the application of the WBE model to terrestrial plants claims that whole-plant resource use in terrestrial plants scales as the 3/4 power of total mass, as predicted by the WBE model. However, in the present study we show that empirical data actually do not support such a claim. More recent studies cited as evidence for 3/4 scaling also suffer from several statistical and data-related problems. Using a forest biomass dataset including 1 266 plots of 17 main forest types across China, we explored the scaling exponents between tree productivity and tree mass and found no universal value across forest stands. We conclude that there is not sufficient evidence to support the existence of a single constant scaling exponent for the metabolism-biomass relationship for terrestrial plants.     

Lack of Evidence for 3/4 Scaling of Metabolism in Terrestrial Plants

Scaling, as the translation of information across spatial, temporal, and organizational scales, is essential to predictions and understanding in all sciences and has become a central issue in ecology. A large body of theoretical and empirical evidence concerning allometric scaling in terrestrial individual plants and plant communities has been constructed around the fractal volume-filling theory of West, Brown, and Enquist (the WBE model). One of the most thought-provoking findings has been that the metabolic rates of plants, like those of animals, scale with their size as a 3/4 power law. The earliest, single most-important study cited in support of the application of the WBE model to terrestrial plants claims that whole-plant resource use in terrestrial plants scales as the 3/4 power of total mass, as predicted by the WBE model.However, in the present study we show that empirical data actually do not support such a claim. More recent studies cited as evidence for 3/4 scaling also suffer from several statistical and data-related problems. Using a forest biomass dataset including 1 266 plots of 17 main forest types across China, we explored the scaling exponents between tree productivity and tree mass and found no universal value across forest stands. We conclude that there is not sufficient evidence to support the existence of a single constant scaling exponent for the metabolism-biomass relationship for terrestrial plants.     

Sizing up allometric scaling theory

Metabolic rate, heart rate, lifespan, and many other physiological properties vary with body mass in systematic and interrelated ways. Present empirical data suggest that these scaling relationships take the form of power laws with exponents that are simple multiples of one quarter. A compelling explanation of this observation was put forward a decade ago by West, Brown, and Enquist (WBE). Their framework elucidates the link between metabolic rate and body mass by focusing on the dynamics and structure of resource distribution networks-the cardiovascular system in the case of mammals. Within this framework the WBE model is based on eight assumptions from which it derives the well-known observed scaling exponent of 3/4. In this paper we clarify that this result only holds in the limit of infinite network size (body mass) and that the actual exponent predicted by the model depends on the sizes of the organisms being studied. Failure to clarify and to explore the nature of this approximation has led to debates about the WBE model that were at cross purposes. We compute analytical expressions for the finite-size corrections to the 3/4 exponent, resulting in a spectrum of scaling exponents as a function of absolute network size. When accounting for these corrections over a size range spanning the eight orders of magnitude observed in mammals, the WBE model predicts a scaling exponent of 0.81, seemingly at odds with data. We then proceed to study the sensitivity of the scaling exponent with respect to variations in several assumptions that underlie the WBE model, always in the context of finite-size corrections. Here too, the trends we derive from the model seem at odds with trends detectable in empirical data. Our work illustrates the utility of the WBE framework in reasoning about allometric scaling, while at the same time suggesting that the current canonical model may need amendments to bring its predictions fully in line with available datasets.     

Computer simulations support a core prediction of a contentious plant model

? Premise of the study: An overarching but vigorously debated plant model proposed by the West, Brown, Enquist (WBE) theory predicts the scaling relationships for numerous botanical phenomena. However, few studies have evaluated this model's basic assumptions, one of which is that natural selection has resulted in hierarchal networks that minimize the energy required to distribute nutrients internally and have thus produced highly efficient organisms. ? Methods: If these core assumptions are correct, an "idealized" plant complying with all of the scaling relationships emerging from the WBE plant model should rapidly outcompete other plants, even those that differ slightly from it. To test this reasoning, a computer model was used to simulate competition between an idealized WBE plant, a generic "average" angiosperm (GA), and one of seven variants of the idealized WBE plant, each being similar to the GA in one of the GA's scaling parameters. ? Key results: Replicate simulations show that the idealized WBE plant rapidly outcompetes all other plants under light-shade and open-field conditions. However, changing only one of the WBE's scaling parameters results in death or in the coexistence of WBE and GA plants. ? Conclusions: These simulations support a core assumption of the WBE plant model and suggest why this idealized plant has not evolved.     

Plant physiology in theory and practice: An analysis of the WBE model for vascular plants

The theoretical model of West, Brown and Enquist (hereafter WBE) proposed the fractal geometry of the transport system as the origin of the allometric scaling laws observed in nature. The WBE model has either been criticized for some restrictive and biologically unrealistic constraints or its reliability debated on the evidence of empirical tests. In this work, we revised the structure of the WBE model for vascular plants, highlighting some critical assumptions and simplifications and discuss them with regard to empirical evidence from plant anatomy and physiology. We conclude that the WBE model had the distinct merit of shedding light on some important features such as conduit tapering. Nonetheless, it is over-simplistic and a revised model would be desirable with an ontogenetic perspective that takes some important phenomena into account, such as the transformation of the inner sapwood into heartwood and the effect of hydraulic constraints in limiting the growth in height.