生态化学计量学理论的应用、完善与扩展

生态化学计量学结合生物学、化学和物理学等基本原理, 研究碳、氮、磷等化学元素在各种生态过程中的平衡。由于生态化学计量学研究可以把生态实体的各个层次在元素水平上统一起来, 因此生态化学计量学已成为许多生态系统的新兴研究工具。目前, 生态化学计量学研究已深入到生态学的各个层次(分子、细胞、个体、种群、群落、生态系统)及区域等不同尺度。由于C、N、P 对有机体和生态系统的结构和功能的重要作用, C∶N∶P 化学计量学成为各种生态过程研究中的核心内容, 其基本理论(动态平衡理论、生长速率理论)围绕C∶N∶P 化学计量比而展开阐述。将生态化学计量学理论应用于全球格局下的生态系统研究时, 产生了许多崭新的成果(如植物营养全球格局等)。希望对生态化学计量学的概念、核心理论和全球格局下的应用以及该学说的完善与发展状况的简单介绍, 能有助于推动我国在此领域的相关研究。     

生态系统碳氮磷元素的生态化学计量学特征

生态系统元素平衡是当前全球变化生态学和生物地球化学循环的研究热点和焦点.在系统介绍生态化学计量学与碳氮磷元素循环研究进展的基础上,重点从土壤C:N:P化学计量比的分布特征、指示作用、对碳固定的影响,以及人类活动对C:N:P比的影响等方面探讨了C:N:P比在养分限制、生物地球化学循环、森林演替与退化等领域中的应用等问题,并展望了生态系统碳氮磷平衡的元素化学计量学未来研究的发展方向.通过对生态化学计量学理论和方法的研究,可以深入认识植物-凋落物-土壤相互作用的养分调控因素,对于揭示碳氮磷元素之间的相互作用及平衡制约关系,为减缓温室效应提供新思路和理论依据,具有重要的现实意义.     

生态化学计量学特征及其应用研究进展

生态化学计量学已成为生态学研究的热点问题。作为一门新兴学科,综观国内外最新研究进展,相关研究目前尚存在着许多不足。基于此,从全球与区域尺度、功能群尺度及个体水平3个方面阐述生态化学计量学特征,从空间、时间、生境和植物类型等生物与非生物因素综述生态化学计量学特征的驱动因素。并讨论生态化学计量学特征在限制性养分判断、生态系统稳定性、生长率与C:N:P关系中的应用。     

生态化学计量学研究进展

生态化学计量学结合生物学、化学和物理学等基本原理,研究能量和碳、氮、磷等化学元素在生态系统中,特别是各种生态系统过程(如竞争、捕食、寄生、共生等)参与者中的变化,以及它们之间的动态平衡,并分析这种平衡对生态系统的影响。目前,C∶N∶P化学计量学研究已深入到生态学的各个层次(细胞、个体、种群、群落、生态系统)及区域等不同尺度。近年来,由于认识到化学计量学研究可以把生态实体的各个层次在元素水平上统一起来,因此生态化学计量学已成为许多生态系统的新兴研究工具。其中,C∶N∶P化学计量学是各种生态过程研究中的核心内容。论述了生态化学计量学在物种、群落、生态系统等各层次的应用现状,并指出了C∶N∶P化学计量学研究的应用前景和发展趋势,以期引起同行的重视并推动该领域的进一步发展。     

荒漠草原不同雨量带土壤-植物-微生物C、N、P及其化学计量特征

降水作为关键性驱动因子深刻影响着荒漠草原生态系统养分循环过程。采用生态化学计量学方法,调查了荒漠草原不同雨量带土壤-植物-微生物C、N、P及其生态化学计量特征对降水格局的适应性规律。研究区不同雨量带土壤C、N、P随降水梯度的递减亦呈现递减趋势。平均土壤C∶N∶P比例为28.9∶2.7∶1,主要受到P元素控制。不同雨量带平均土壤MBC∶MBN∶MBP比例为108.6∶5.6∶1,表现出明显的C富集现象。不同雨量带平均植物C∶N∶P比例为117.4∶6.7∶1,表现为明显的C、N缺乏或P富集。降水为主的气候原因造成了研究区环境中P含量相对较高,并直接反映在了植物化学计量特征上。研究区土壤C和N之间具有极显著的正相关关系(P0.01),相关系数高达0.98。植物N和P之间具有显著的正相关关系(P0.05),相关系数为0.90。土壤N与植物C、P分别呈显著正相关和显著负相关(P0.05),相关系数分别为0.84和-0.82。降水在塑造荒漠草原生态格局以及驱动生态系统养分循环过程中发挥了关键性作用。     

N：P化学计量学在生态学研究中的应用

化学计量学很早就被应用于生态学研究中,但长期以来几乎被生态学家所忽视。近年来,由于认识到化学计量学研究可以把生态实体的各个层次在元素水平上统一起来,因此元素化学计量学成为近年来新兴的一个生态学研究领域。氮磷作为植物生长的必需矿质营养元素和生态系统常见的限制性元素,在植物体内存在功能上的联系,二者之间具有重要的相互作用。近年来由于人类活动的强烈影响,这两种元素的循环在速度和规模上都发生了前所未有的改变,导致一系列环境问题的出现,因此N：P化学计量学研究就显得极为重要。本文论述了N：P化学计量学在物种、群落、生态系统等各层次的应用现状,同时从分子生物学角度分析了应用N：P化学计量学的可行性,并指出了N：P化学计量学研究的应用前景和存在的缺陷。     

海洋浮游生物的生态化学计量学研究进展

生态化学计量学可以简单定义为从分子到生物圈的元素生物学,其跨越了环境和生命的各个层次,是构建从分子到生态系统统一化理论的新思路,是生态科学发展的必然趋势.海洋生物占地球生物圈总生物量的50%,是全球生物地球化学循环的重要组成部分,而浮游生物作为海洋生态系统物质循环和能量流动的重要环节,在海洋生态系统元素循环过程中起着关键作用.但是目前关于海洋浮游生物生态化学计量学的研究较零散和缺乏.因此,本文从限制元素影响海洋浮游生物的生态现象和机理、生化物质对营养限制的响应、营养限制的食物链传递与反馈4方面,对海洋浮游生物化学计量学研究进行综述,分析了该领域当前存在的问题,并对我国海洋浮游生物生态化学计量学研究的发展重点提出了展望.
     

中国典型生态脆弱区生态化学计量学研究进展

在人类活动和自然环境变化的相互作用下,生态脆弱区生态系统随之变迁,荒漠化、盐碱化、水土流失、植被生产力下降等是生态脆弱区面临的重要问题。生态化学计量学作为当前多学科交叉研究的热点领域,强调从生态系统能量与元素平衡角度,揭示元素生物地球化学循环和生态系统对环境变化的调控机制。为了促进对生态脆弱区碳（C）、氮（N）、磷（P）生态化学计量的深入理解,本文重点总结了近年来有关我国典型生态脆弱区植物、凋落物、土壤和土壤微生物量C、N、P生态化学计量及其对环境变化响应的研究成果,并展望未来研究方向,以期促进生态化学计量学的发展和生态脆弱区生态保护与恢复研究。研究表明,植物-凋落物-土壤-土壤微生物系统C、N、P化学计量具有较强相关性,并受土壤因子、气候因子、生物因子和人类活动的显著影响。在生态脆弱区,我国北方荒漠及荒漠化地区由于较高的N∶P比值易受P限制,而青藏高原脆弱区、西南岩溶石漠化地区和黄土高原脆弱区等生态脆弱区更易受N限制;随着植被恢复,养分限制逐渐由N限制向P限制转变。生态脆弱区相对较低的养分含量和C∶N∶P比值或许可在一定程度上解释植被生产力较低的原因,而具有较高N、P化学计量内稳性的植物在贫瘠环境中具有较强竞争力和更高稳定性。今后可加强多尺度、不同生态系统植物-凋落物-土壤-土壤微生物系统生态化学计量和长期、多因子交互控制实验的研究。     

N:P化学计量学在生态学研究中的应用

化学计量学很早就被应用于生态学研究中,但长期以来几乎被生态学家所忽视.近年来,由于认识到化学计量学研究可以把生态实体的各个层次在元素水平上统一起来,因此元素化学计量学成为近年来新兴的一个生态学研究领域.氮磷作为植物生长的必需矿质营养元素和生态系统常见的限制性元素,在植物体内存在功能上的联系,二者之间具有重要的相互作用.近年来由于人类活动的强烈影响,这两种元素的循环在速度和规模上都发生了前所未有的改变,导致一系列环境问题的出现,因此N:P化学计量学研究就显得极为重要.本文论述了N:P化学计量学在物种、群落、生态系统等各层次的应用现状,同时从分子生物学角度分析了应用N:P化学计量学的可行性,并指出了N:P化学计量学研究的应用前景和存在的缺陷.     

西南喀斯特典型石漠化生态系统土壤养分生态化学计量特征及其影响因素

喀斯特石漠化生态系统土壤养分元素生态化学计量特征及其对环境变异的生态响应是喀斯特退化森林生态系统恢复重建必需明确的关键科学问题。为探明喀斯特石漠化土壤C、N、P、K养分元素生态化学计量特征,探讨其对环境因子的响应,对西南喀斯特3个典型石漠化调查点(贵州毕节鸭池、清镇红枫湖和关岭-贞丰花江) 90个样方土壤及环境因子调查取样,研究了其土壤有机碳(C)、全氮(N)、全磷(P)及全钾(K)的化学计量特征及其影响因素。结果表明:西南喀斯特典型石漠化生态系统土壤C、N、P、K平均含量分别为45.61、2.54、0.79 g/kg和3.33 g/kg,计量比C∶N、C∶P、C∶K、N∶P、N∶K、P∶K平均值分别为19.56、65.07、23.65、3.45、1.32和0.39。4个土壤养分元素中,K元素表现明显高于其他元素的波动性。土壤养分含量及化学计量比在不同调查点、石漠化等级及植被覆盖率环境均有显著差异。无石漠化环境土壤养分C、N、P含量显著大于潜在、轻度、中度和强度石漠化,而强度石漠化环境土壤养分K含量却显著高于其他等级石漠化。土壤养分含量之间及其与化学计量比之间多具有显著的非线性相关关系。降水、温度、岩石裸露率和土地覆被是西南喀斯特石漠化生态系统土壤养分及其化学计量比最主要的影响因素。研究结果对丰富土壤生态化学计量学科学理论和我国西南喀斯特石漠化退化植被科学恢复具有重要意义。     

生态化学计量学在水生态系统中的研究与应用

生态化学计量学主要是研究碳、氮、磷等元素在各种生态过程中平衡的一门科学,其核心问题是揭示生物体元素组成的差异对生态功能的影响。由于生态化学计量学研究可以把生态实体的各个层次在元素水平上统一起来,因此生态化学计量学已成为许多生态系统的新兴研究工具。目前,生态化学计量学的研究与应用已深入到生态学的各个层次(分子、细胞、个体、种群、群落、生态系统及区域等不同尺度)。该文围绕生态化学计量学的两个重要组成理论,并结合笔者近年来的研究,归纳总结了生态化学计量学在水生态系统中的研究与应用及未来研究重点,希望有助于推动我国生态化学计量学在水生态系统中的应用研究。     

SEED: A framework for integrating ecological stoichiometry and eco-evolutionary dynamics

Characterising the extent and sources of intraspecific variation and their ecological consequences is a central challenge in the study of eco-evolutionary dynamics. Ecological stoichiometry, which uses elemental variation of organisms and their environment to understand ecosystem patterns and processes, can be a powerful framework for characterising eco-evolutionary dynamics. However, the current emphasis on the relative content of elements in the body (i.e. organismal stoichiometry) has constrained its application. Intraspecific variation in the rates at which elements are acquired, assimilated, allocated or lost is often greater than the variation in organismal stoichiometry. There is much to gain from studying these traits together as components of an ‘elemental phenotype’. Furthermore, each of these traits can have distinct ecological effects that are underappreciated in the current literature. We propose a conceptual framework that explores how microevolutionary change in the elemental phenotype occurs, how its components interact with each other and with other traits, and how its changes can affect a wide range of ecological processes. We demonstrate how the framework can be used to generate novel hypotheses and outline pathways for future research that enhance our ability to explain, analyse and predict eco-evolutionary dynamics.     

Spatial stoichiometry: cross‐ecosystem material flows and their impact on recipient ecosystems and organisms

Cross‐ecosystem material flows, in the form of inorganic nutrients, detritus and organisms, spatially connect ecosystems and impact food web dynamics. To date research on material flows has focused on the impact of the quantity of these flows and largely ignored their elemental composition, or quality. However, the ratios of elements like carbon, nitrogen and phosphorus can influence the impact material flows have on food web interactions through stoichiometric mismatches between resources and consumers. The type and movement of materials likely vary in their ability to change stoichiometric constraints within the recipient ecosystem and materials may undergo changes in their own stoichiometry during transport. In this literature review we evaluate the importance of cross‐ecosystem material flows within the framework of ecological stoichiometry. We explore how movement in space and time impacts the stoichiometry of material flow, as these transformations are essential to consider when assessing the ability of these flows to impact food web productivity and ecosystem functioning. Our review suggests that stoichiometry of cross‐ecosystem material flows are highly dynamic and undergo changes during transport across the landscape or from human influence. These material flows can impact recipient organisms if they change stoichiometry of the abiotic medium, or provide resources that have a different stoichiometry to in situ resources. They might also alter consumer excretion rates, in turn altering the availability of nutrients in the recipient ecosystem. These alterations in stoichiometric constraints of recipient organisms can have cascading trophic effects and shape food web dynamics. We highlight significant gaps in the literature and suggest new avenues for research that explore how cross‐ecosystem material flows impact recipient ecosystems when considering differences in stoichiometric quality, their movement through the landscape and across ecosystem boundaries, and the nutritional constraints of the recipient organisms.     

Recent advances in ecological stoichiometry: insights for population and community ecology

Conventional theories of population and community dynamics are based on a single currency such as number of individuals, biomass, carbon or energy. However, organisms are constructed of multiple elements and often require them (in particular carbon, phosphorus and nitrogen) in different ratios than provided by their resources; this mismatch may constrain the net transfer of energy and elements through trophic levels. Ecological stoichiometry, the study of the balance of elements in ecological processes, offers a framework for exploring ecological effects of such constraints. We review recent theoretical and empirical studies that have considered how stoichiometry may affect population and community dynamics. These studies show that stoichiometric constraints can affect several properties of populations (e.g. stability, oscillations, consumer extinction) and communities (e.g. coexistence of competitors, competitive interactions between different guilds). We highlight gaps in general knowledge and focus on areas of population and community ecology where incorporation of stoichiometric constraints may be particularly fruitful, such as studies of demographic bottlenecks, spatial processes, and multi-species interactions. Finally, we suggest promising directions for new research by recommending potential study systems (terrestrial insects, detritivory-based webs, soil communities) to improve our understanding of populations and communities. Our conclusion is that a better integration of stoichiometric principles and other theoretical approaches in ecology may allow for a richer understanding of both population and community structure and dynamics.     

全球变化背景下陆地植物N/P生态化学计量学研究进展

生态化学计量学理论最早应用在水生生态系统的研究中,但最近20年来在陆地生态系统中也开展了大量的相关研究,特别是关于全球变化背景下陆地植物N/P生态化学计量学方面的研究得到很大的发展,极大地丰富和提升了我们对陆地植物包括生态系统生态过程的认识。就全球变化背景下陆地植物生态化学计量学的研究现状进行了综述,同时以中国科学院华南植物园90周庆为契机,总结我们关于南亚热带森林植物生态化学计量学的研究工作,进而分析当前存在的一些问题并提出今后研究的发展重点,以期促进和推动我园和我国生态化学计量学相关领域的研究。     

Think ratio! A stoichiometric view on biodiversity–ecosystem functioning research

Ecological stoichiometry (ES) has become one of the most pervasive theoretical frameworks in environmental sciences and biology in the last two decades. ES allows predicting processes on all organizational levels from subcellular structures to ecosystems by relating the elemental composition and demand of organisms to the relative availability of resources. However, ES has been rarely used to understand and predict the relationship between biodiversity and ecosystem functioning (BEF), although ES would be ideally suited as it makes predictions on both population processes underlying biodiversity as well as on matter transformations underlying ecosystem processes. Here, we propose to link the two fields of research on ES and BEF relationships and highlight a number of potential avenues for further research. First, we cast a stoichiometric view on drivers of biodiversity change. Second, we address the stoichiometric underpinning of biodiversity–productivity relationships. Third, we discuss potential interactions between stoichiometry and diversity in a food web context.     

Interactions between temperature and nutrients across levels of ecological organization

Temperature and nutrient availability play key roles in controlling the pathways and rates at which energy and materials move through ecosystems. These factors have also changed dramatically on Earth over the past century as human activities have intensified. Although significant effort has been devoted to understanding the role of temperature and nutrients in isolation, less is known about how these two factors interact to influence ecological processes. Recent advances in ecological stoichiometry and metabolic ecology provide a useful framework for making progress in this area, but conceptual synthesis and review are needed to help catalyze additional research. Here, we examine known and potential interactions between temperature and nutrients from a variety of physiological, community, and ecosystem perspectives. We first review patterns at the level of the individual, focusing on four traits – growth, respiration, body size, and elemental content – that should theoretically govern how temperature and nutrients interact to influence higher levels of biological organization. We next explore the interactive effects of temperature and nutrients on populations, communities, and food webs by synthesizing information related to community size spectra, biomass distributions, and elemental composition. We use metabolic theory to make predictions about how population‐level secondary production should respond to interactions between temperature and resource supply, setting up qualitative predictions about the flows of energy and materials through metazoan food webs. Last, we examine how temperature–nutrient interactions influence processes at the whole‐ecosystem level, focusing on apparent vs. intrinsic activation energies of ecosystem processes, how to represent temperature–nutrient interactions in ecosystem models, and patterns with respect to nutrient uptake and organic matter decomposition. We conclude that a better understanding of interactions between temperature and nutrients will be critical for developing realistic predictions about ecological responses to multiple, simultaneous drivers of global change, including climate warming and elevated nutrient supply.     

Using ecological stoichiometry to understand and predict infectious diseases

A key characteristic of host–parasite interactions is the theft of host nutrients by the parasite, yet we lack a general framework for understanding and predicting the interplay of host and parasite nutrition that applies across biological levels of organization. The elemental nutrients (C, N, P, Fe, etc.), and ecological stoichiometry provide a framework for understanding host–parasite interactions and their relation to ecosystem functioning. Here we use the ecological stoichiometry framework to develop hypotheses and predictions regarding the relationship between elemental nutrients and host–parasite interactions. We predict that a suite of host and parasite traits, stoichiometric homeostasis, host diet stoichiometry, and biogeochemical cycling are related to disease dynamics, host immunity and resistance, and bacterial growth form determination. We show that ecological stoichiometry is capable of expanding our understanding of host–parasite interactions, and complementing other approaches such as population and community ecology, and molecular biology, for studying infectious diseases.