稳定同位素技术在植物水分利用研究中的应用

近20a稳定同位素技术在植物生态学研究中的应用得到了长足发展,使得对植物与水分关系也有了更深一步的了解。介绍稳定同位素性碳、氢、氧同位素在研究植物水分关系中的应用及进展,以期能为国内植物水分利用研究提供参考。由于植物根系从土壤中吸收水分时并不发生同位素分馏,对木质部水分同位素分析有助于对植物利用水分来源,生态系统中植物对水分的竞争和利用策略的研究,更好地了解生态系统结构与功能。稳定碳同位素作为植物水分利用效率的一个间接指标,在不同水分梯度环境中,及植物不同代谢产物与水分关系中有着广泛的应用。同位素在土壤-植被-大气连续体水分中的应用,有助于了解生态系统的水分平衡。随着稳定同位素方法的使用,植物与水分关系的研究将取得更大的进展。     

Water source niche overlap increases with site moisture availability in woody perennials

Classical niche partitioning theory posits increased competition for and partitioning of the most limiting resource among coexisting species. Coexisting plant species may vary in rooting depth, reflecting niche partitioning in water source use. Our goal was to assess the soil water partitioning of woody plant communities across northern Arizona along an elevational moisture gradient using stem and soil water isotopes from two sampling periods to estimate the use of different water sources. We hypothesized that niche overlap of water sources would be higher and monsoon precipitation uptake would be lower at sites with higher moisture availability. Pairwise niche overlap of coexisting species was calculated using mixing model estimates of proportional water use for three sources. Across the moisture gradient, niche overlap increased with site moisture index (precipitation/potential evapotranspiration) across seasons, and site moisture index explained 37% of the variation in niche overlap of intermediate and deeper sources of water. Desert trees utilized more winter source water than desert shrubs, suggesting the partitioning of water sources between functional groups. However, seasonal differences in surface water use were primarily found at intermediate levels of site moisture availability. Our findings support classical niche partitioning theory in that plants exhibit higher overlap of water sources when water is not a limiting resource.     

滇池流域水文生态系统服务权衡与协同时空异质性及其归因分析

城市化扩张影响流域水文生态系统服务供给,导致水文生态环境压力增加。明晰滇池流域水文生态系统服务权衡与协同关系及归因,对于滇池流域社会经济发展与生态可持续管理的双赢具有重要意义。基于InVEST模型估算了滇池流域2000、2010、2018年碳存储、氮输出、土壤保持和产水量4种水文生态系统服务,分析了水文生态系统服务时空格局变化,利用双变量Moran′s I指数分析了滇池流域水文生态系统服务间时空异质性,通过冗余分析和条件效应分析等方法识别水文生态系统服务归因。结果表明：(1)水文生态系统服务的高值分布与土地利用覆盖有密切关系。(2)水文生态系统服务空间变化主要体现在滇池周边人类活动干扰强烈的区域,表现为氮输出、土壤保持和产水量增加,碳存储减小。(3)水文生态系统服务权衡与协同关系变化明显。碳存储和土壤保持为协同;氮输出和产水量先呈协同后呈权衡;产水量、氮输出均与碳存储和土壤保持为权衡,前者空间变化不明显,后者协同范围扩大,权衡范围缩小。(4)水文生态系统服务主要受多个因素共同影响,主要包括：高程、耕地比例、林地比例、建设用地比例。研究结果可为研究区水文生态系统环境治理和土地利用优化配置...     

Stable isotopes of carbon and nitrogen in soil ecological studies

The development of stable isotope techniques is one of the main methodological advances in ecology of the last decades of the 20th century. Many biogeochemical processes are accompanied by changes in the ratio between stable isotopes of carbon and nitrogen (¹²C/¹³C and ¹⁴N/¹⁵N), which allows different ecosystem components and different ecosystems to be distinguished by their isotopic composition. Analysis of isotopic composition makes it possible to trace matter and energy flows through biological systems and to evaluate the rate of many ecological processes. The main concepts and methods of stable isotope ecology and patterns of stable isotope fractionation during organic matter decomposition are considered with special emphasis on the fractionation of isotopes in food chains and the use of stable isotope studies of trophic relationships between soil animals in the field.     

Testing the long-term stability of marine isoscapes in shelf seas using jellyfish tissues

Maps of the spatial distribution of stable isotope ratios across wide geographic areas (isoscapes) are increasingly used to study mechanisms of nutrient flux, movements of animals, and to improve trophic information derived from stable isotope analyses. Isoscapes are usually constructed from reference samples collected from known geographic positions, a time consuming and costly process. In this study, we test the temporal stability of isoscapes of carbon and nitrogen isotopes across the North Sea over a ten-year period. Using jellyfish tissues as reference organisms, we show that hydrodynamic and biogeochemical processes controlling the distribution of carbon and nitrogen isotope values, and thus that the underlying isoscapes, are temporally stable. Remarkably, broad geographic variations in stable carbon and nitrogen isotope distributions across the North Sea are consistent with previously published variations seen in archaeological cod bones from the ninth to seventeenth centuries, despite dramatic changes in land use in the past 1,000 years. Stable isotope-based studies of trophic interactions or movements in animals with lifespans of a year or more in shelf ecosystems can consequently be referenced to previously published isoscape models, and do not require construction of temporally explicit isotope baseline corrections. Scyphomedusan jellyfish are excellent reference organisms for ecological stable isotope analyses in pelagic ecosystems, due to their widespread distributions, well-defined life histories, and fast tissue growth.     

Multidimensional stable isotope analysis illuminates resource partitioning in a sub‐Antarctic island bird community

A central theme in community ecology is understanding how similar species co‐exist and how their interactions may evolve in the context of climate change. Most studies of resource partitioning among central place foragers, particularly birds, focus on the offspring‐rearing period, when they are accessible, but breeding success may be determined earlier and little is known about how such species partition resources at the onset of breeding. We used a non‐invasive approach to evaluate resource partitioning in co‐existing females at a sub‐Antarctic island during their pre‐laying periods. Three hypotheses were tested using carbon, nitrogen and oxygen stable isotope ratios measured in shells and membranes of hatched eggs as ecological tracers: 1) resource partitioning by geographic location and trophic level will exist among the 12 bird species and will be enhanced within taxonomic groups; 2) given the absence of strong oxygen gradients in the Southern Ocean we will not detect spatial structuring based on oxygen isotopes, but differences will exist between resident and oceanic species as the former may use meteoric water; 3) capital and income breeder strategies can be differentiated using stable isotopes of egg remains. Two and three dimensional isotopic data showed resource partitioning among species. As predicted, segregation was evident within the four main taxonomic groups: penguins, albatrosses, burrowing petrels and giant petrels. Unexpectedly, oxygen isotopes revealed widespread use of meteoric water among a suite of sub‐Antarctic birds. Stable isotopes allowed us to identify females of most species as income breeders at the onset of breeding, with the exception of the females of the two crested penguin exhibiting a mix of income and capital resources use. Multidimensional isotopic analyses revealed that resource partitioning exists at multiple stages of the annual cycle in ways likely to be important under global change, exhibiting wide potential for ecosystem analysis.     

Nitrogen and carbon isotope fractionation by two species of marine amphipods: Laboratory and field studies

The fractionation of the stable nitrogen and carbon isotopes between two species of marine amphipods and their food sources was investigated in both field and laboratory studies. The nitrogen isotope ratios showed a strong species effect. Amphithoe valida Smith grown in the laboratory averaged ?0.3‰ relative to several food sources. Parhyale hawaiensis (Dana) was +2.3‰ relative to its diets. Animals from the field showed similar differences from the macrophytic algae on which they were found. The carbon isotope composition of both species averaged ?0.7‰ relative to their diets. Laboratory animals fed fresh algae were slightly more negative than field animals or animals fed detrital algae.     

Spatial and temporal patterns of carbon flow in a temperate,large river food web

Using natural abundances of stable carbon (δ¹³C) and nitrogen (δ¹⁵N) isotopes, we quantified spatial and temporal patterns of carbon flow through the main channel food web in the lowland section of New Zealand’s longest river, the Waikato River. The study was undertaken with the objective of determining whether the Waikato River conforms to contemporary theoretical concepts regarding carbon flow in large river food webs. Potential organic carbon sources and invertebrate and fish consumers were sampled from three different hydrogeomorphic zones on six occasions, representing a range of seasonal and flow conditions. In line with the predictions of the riverine productivity model and riverine ecosystem synthesis, autochthonous algae and biofilms were the most important basal carbon source contributing to consumer biomass. These were often supported by C₃ aquatic macrophytes and allochthonous C₃ riparian plants. The relative importance of organic carbon sources differed between zones and appeared to change depending on season, presumably in response to water temperature and flow, particularly in the unconstrained zone of the lower river. We also demonstrate that to draw robust conclusions, consideration must be given to quantifying the isotopic signatures of organisms lower in the food web, as these can change significantly between sampling times and hydrogeomorphic zones.     

Estimating animal abundance in ground beef batches assayed with molecular markers

Estimating animal abundance in industrial scale batches of ground meat is important for mapping meat products through the manufacturing process and for effectively tracing the finished product during a food safety recall. The processing of ground beef involves a potentially large number of animals from diverse sources in a single product batch, which produces a high heterogeneity in capture probability. In order to estimate animal abundance through DNA profiling of ground beef constituents, two parameter-based statistical models were developed for incidence data. Simulations were applied to evaluate the maximum likelihood estimate (MLE) of a joint likelihood function from multiple surveys, showing superiority in the presence of high capture heterogeneity with small sample sizes, or comparable estimation in the presence of low capture heterogeneity with a large sample size when compared to other existing models. Our model employs the full information on the pattern of the capture-recapture frequencies from multiple samples. We applied the proposed models to estimate animal abundance in six manufacturing beef batches, genotyped using 30 single nucleotide polymorphism (SNP) markers, from a large scale beef grinding facility. Results show that between 411～1367 animals were present in six manufacturing beef batches. These estimates are informative as a reference for improving recall processes and tracing finished meat products back to source.     

Stable Isotope Turnover and Half-Life in Animal Tissues: A Literature Synthesis

Stable isotopes of carbon, nitrogen, and sulfur are used as ecological tracers for a variety of applications, such as studies of animal migrations, energy sources, and food web pathways. Yet uncertainty relating to the time period integrated by isotopic measurement of animal tissues can confound the interpretation of isotopic data. There have been a large number of experimental isotopic diet shift studies aimed at quantifying animal tissue isotopic turnover rate λ (%·day^-1, often expressed as isotopic half-life, ln(2)/λ, days). Yet no studies have evaluated or summarized the many individual half-life estimates in an effort to both seek broad-scale patterns and characterize the degree of variability. Here, we collect previously published half-life estimates, examine how half-life is related to body size, and test for tissue- and taxa-varying allometric relationships. Half-life generally increases with animal body mass, and is longer in muscle and blood compared to plasma and internal organs. Half-life was longest in ecotherms, followed by mammals, and finally birds. For ectotherms, different taxa-tissue combinations had similar allometric slopes that generally matched predictions of metabolic theory. Half-life for ectotherms can be approximated as: ln (half-life) = 0.22*ln (body mass) + group-specific intercept; n = 261, p<0.0001, r² = 0.63. For endothermic groups, relationships with body mass were weak and model slopes and intercepts were heterogeneous. While isotopic half-life can be approximated using simple allometric relationships for some taxa and tissue types, there is also a high degree of unexplained variation in our models. Our study highlights several strong and general patterns, though accurate prediction of isotopic half-life from readily available variables such as animal body mass remains elusive.     

Use of the doubly labeled water method for measurement of energy expenditure, total body water, water intake, and metabolizable energy intake in humans and small animals

The basis of the doubly labeled water method is measurement of the differential rates of disappearance of two isotopes of water (H2 18O and either 2H2O or 3H2O, administered at the start of the study) from body water. Published studies indicate that, in its current forms, this technique can be used to provide accurate and reasonably precise information on carbon dioxide production, total body water, and water intake in free-living humans and many small animals. Total energy expenditure can be calculated from carbon dioxide production with little loss of precision. Metabolizable energy intake can also be predicted, as the sum of total energy expenditure plus an estimate for the change in body energy stores during the measurement, but this prediction is unlikely to be accurate and precise unless the subject is in approximate energy balance.     

Stable isotope views on ecosystem function: challenging or challenged?

Stable isotopes and their potential for detecting various and complex ecosystem processes are attracting an increasing number of scientists. Progress is challenging, particularly under global change scenarios, but some established views have been challenged. The IX meeting of the Spanish Association of Terrestrial Ecology (AAET, Úbeda, 18–22 October 2009) hosted a symposium on the ecology of stable isotopes where the linear mixing model approach of partitioning sinks and sources of carbon and water fluxes within an ecosystem was challenged, and new applications of stable isotopes for the study of plant interactions were evaluated. Discussion was also centred on the need for networks that monitor ecological processes using stable isotopes and key ideas for fostering future research with isotopes.     

Ovariectomy in grasshoppers increases somatic storage, but proportional allocation of ingested nutrients to somatic tissues is unchanged

Reduced reproduction increases storage and extends lifespan in several animal species. The disposable soma hypothesis suggests this life extension occurs by shifting allocation of ingested nutrients from reproduction to the soma. A great deal of circumstantial evidence supports this hypothesis, but no direct tracking of nutrients has been performed in animals that are long-lived because of direct reduction in reproduction. Here, we use the stable isotopes to track carbon and nitrogen from ingestion to somatic organs in long-lived, ovariectomized grasshoppers. Three estimates of somatic storage (viz., quantity of hemolymph storage proteins, amount of femur muscle carbohydrates, and size of the fat body) all doubled upon ovariectomy. In stark contrast, ovariectomy did not increase the proportion of these tissues that were made from recently ingested foods. In other words, the physiology underlying relative allocation to these somatic tissues was not affected by ovariectomy. Thus, at the level of whole tissue storage, these results are consistent with a trade-off between reproduction and longevity. In contrast, our stable isotope data are inconsistent with the prediction that enhanced storage in ovariectomized females results from a physiological shift in allocation of ingested nutrients.     

不同植物叶片水分利用效率对光和CO2的响应与模拟

植物叶片水分利用效率的高低取决于气孔控制的光合作用和蒸腾作用两个相互耦合的过程,模拟水分利用效率对环境变化的响应特征和机制是理解生态系统碳循环和水循环及其耦合关系的基础。研究通过人工控制光强和CO₂浓度,对叶片水分利用效率进行了研究。提出了植物水分利用效率在光强和CO₂浓度共同作用下的估算模型。数据分析表明,该模型在包括C3和C4植物、草本和木本植物在内的9种植物上能很好地模拟水分利用效率对光强和CO₂浓度共同作用的响应。该模型可以用于估算CO₂浓度升高条件下光合速率的提高和蒸腾速率的降低对水分利用效率提高的贡献量。CO₂浓度变化条件下,水分利用效率在不同植物之间有巨大差异,研究区域尺度植物的水分利用效率时至少需要将植物区分为C4植物和C3植物,其中C3植物区分为草本和木本植物3种生态功能型才能较为准确地估算植物的整体水分利用效率。应用本研究提出的水分利用效率估算模型和植物水分利用效率生态功能型分类标准,可以为建立以植物的水分利用效率为基本参数的陆地生态系统水循环模型和陆地生态系统生产力模型提供重要依据。     

基于稳定同位素的湿地食物源判定和食物网构建研究进展

湿地生物营养动力学是湿地生态系统结构和功能评价研究的基础.碳、氮稳定同位素作为识别营养关系的方法,已在湿地生态系统食物来源、组成和食物链传递研究中得到广泛运用.本文系统综述了稳定同位素食物贡献度计算模型和营养级确定的基本方法和理论;讨论了动物营养分馏值和基线的选择依据;概括了湿地生态系统典型食物源及其稳定同位素变化特征;总结了草食、杂食和肉食等不同营养级动物的食物来源.指出了稳定同位素在湿地食物源溯源和食物网研究中的不足;基于国内外研究现状和发展趋势及需求,展望了未来同位素技术在湿地食物网生态学研究中的运用前景和研究重点,提出需要加强稳定同位素营养分馏和基线的影响因素、样品处理和保存方式研究以及胃含物、分子标记物和多元素同位素结合分析.     

稳定性同位素技术与Keeling曲线法在陆地生态系统碳/水交换研究中的应用

稳定性同位素技术和Keeling曲线法是现代生态学研究的重要手段和方法之一。稳定性同位素能够整合生态系统复杂的生物学、生态学和生物地球化学过程在时间和空间尺度上对环境变化的响应。Keeling曲线法是以生物过程前后物质平衡理论为基础,将CO₂或H₂O的同位素组成(δD、δ¹³C或δ¹⁸O)与其对应浓度测量结合起来,将生态系统净碳通量区分为光合固定和呼吸释放通量,或将整个生态系统水分蒸散区分为植物蒸腾和土壤蒸发。在全球尺度上,稳定性同位素技术、Keeling曲线法与全球尺度陆地生态系统模型相结合,还可区分陆地生态系统和海洋生态系统对全球碳通量的贡献以及不同植被类型(C₃或C₄)在全球CO₂同化量中所占的比例。然而,生态系统的异质性使得稳定性同位素技术和Keeling曲线法从冠层尺度外推到生态系统、区域或全球尺度时存在有一定程度的不确定性。此外,取样时间、地点的选取也会影响最终的研究结果。尽管如此,随着分析手段的不断精确和研究方法的日趋完善,稳定性同位素技术和Keeling曲线法与其它测量方法(如微气象法)的有机结合将成为未来陆地生态系统碳/水交换研究的重要手段和方法之一。     

Divergent phenological response to hydroclimate variability in forested mountain watersheds

Mountain watersheds are primary sources of freshwater, carbon sequestration, and other ecosystem services. There is significant interest in the effects of climate change and variability on these processes over short to long time scales. Much of the impact of hydroclimate variability in forest ecosystems is manifested in vegetation dynamics in space and time. In steep terrain, leaf phenology responds to topoclimate in complex ways, and can produce specific and measurable shifts in landscape forest patterns. The onset of spring is usually delayed at a specific rate with increasing elevation (often called Hopkins' Law; Hopkins, 1918), reflecting the dominant controls of temperature on greenup timing. Contrary with greenup, leaf senescence shows inconsistent trends along elevation gradients. Here, we present mechanisms and an explanation for this variability and its significance for ecosystem patterns and services in response to climate. We use moderate‐resolution imaging spectro‐radiometer (MODIS) Normalized Difference Vegetation Index (NDVI) data to derive landscape‐induced phenological patterns over topoclimate gradients in a humid temperate broadleaf forest in southern Appalachians. These phenological patterns are validated with different sets of field observations. Our data demonstrate that divergent behavior of leaf senescence with elevation is closely related to late growing season hydroclimate variability in temperature and water balance patterns. Specifically, a drier late growing season is associated with earlier leaf senescence at low elevation than at middle elevation. The effect of drought stress on vegetation senescence timing also leads to tighter coupling between growing season length and ecosystem water use estimated from observed precipitation and runoff generation. This study indicates increased late growing season drought may be leading to divergent ecosystem response between high and low elevation forests. Landscape‐induced phenological patterns are easily observed over wide areas and may be used as a unique diagnostic for sources of ecosystem vulnerability and sensitivity to hydroclimate change.     

The confounding effects of source isotopic heterogeneity on consumer–diet and tissue–tissue stable isotope relationships

Stable isotope analysis of consumer tissues document patterns of resource use because data are linearly related to isotope compositions of their source(s) (i.e., food, water, etc.). Deviations in parameters estimated for these relationships can arise from variations in consumer tissue-diet spacing (Δ(TS)) and the level of isotopic heterogeneity in the source(s). We present a set of simple hypotheses that distinguish between the effects of Δ(TS) and source isotope heterogeneity. The latter may arise via mixed diets, during tissue turnover, or by isotopic routing of dietary components. We apply these concepts to stable carbon and nitrogen isotope relationships between gut contents and body tissues of large mammal herbivores from mixed C(3)/C(4) South African savannas and test predictions based on the compound- and/or time-specific data archived within each material. Predicted effects of source isotope heterogeneity are readily detected in carbon isotope relationships between materials representing different time periods or comprising bulk versus protein-only diet components. Differences in Δ(TS) of carbon isotopes across mammal herbivore species with very different feeding niches (and diet isotope compositions) are likely to be small or non-existent in these habitats. Variations in Δ(TS) estimated for nitrogen isotopes are much greater, leading to inconsistencies that cannot be explained by diet or trophic level effects alone. The effects of source heterogeneity on isotopic relationships generate numerical artefacts that have been misinterpreted as variations in Δ(TS). We caution against generalized application of hypotheses based on assumptions of source isotopic homogeneity, even for single diets commonly used in laboratory studies. More careful consideration of how heterogeneity affects consumer-diet relationships is needed for many field and laboratory systems.     

Animal board invited review: OneARK: Strengthening the links between animal production science and animal ecology

Wild and farmed animals are key elements of natural and managed ecosystems that deliver functions such as pollination, pest control and nutrient cycling within the broader roles they play in contributing to biodiversity and to every category of ecosystem services. They are subjected to global changes with a profound impact on the natural range and viability of animal species, the emergence and spatial distribution of pathogens, land use, ecosystem services and farming sustainability. We urgently need to improve our understanding of how animal populations can respond adaptively and therefore sustainably to these new selective pressures. In this context, we explored the common points between animal production science and animal ecology to identify promising avenues of synergy between communities through the transfer of concepts and/or methodologies, focusing on seven concepts that link both disciplines. Animal adaptability, animal diversity (both within and between species), selection, animal management, animal monitoring, agroecology and viability risks were identified as key concepts that should serve the cross-fertilization of both fields to improve ecosystem resilience and farming sustainability. The need for breaking down interdisciplinary barriers is illustrated by two representative examples: i) the circulation and reassortment of pathogens between wild and domestic animals and ii) the role of animals in nutrient cycles, i.e. recycling nitrogen, phosphorus and carbon through, for example, contribution to soil fertility and carbon sequestration. Our synthesis identifies the need for knowledge integration techniques supported by programmes and policy tools that reverse the fragmentation of animal research toward a unification into a single Animal Research Kinship, OneARK, which sets new objectives for future science policy. At the interface of animal ecology and animal production science, our article promotes an effective application of the agroecology concept to animals and the use of functional diversity to increase resilience in both wild and farmed systems. It also promotes the use of novel monitoring technologies to quantify animal welfare and factors affecting fitness. These measures are needed to evaluate viability risk, predict and potentially increase animal adaptability and improve the management of wild and farmed systems, thereby responding to an increasing demand of society for the development of a sustainable management of systems.     

Multi‐decadal and ontogenetic trophic shifts inferred from stable isotope ratios of pinniped teeth

Identifying and characterizing top predators’ use of trophic resources provides important information about animal ecology and their response to changing conditions. Information from sources such as stable isotopes can be used to infer changes in resource use as direct observations in the wild are difficult to obtain, particularly in the marine environment. Stable carbon and nitrogen isotope values were recovered from the canine teeth of grey seals collected from haul outs in the central North Sea in the 1970/1980s (n = 44) and 2000s (n = 25), spanning a period of marked ecosystem changes in the region. Extracting material deposited during juvenile and adult life‐stages, we reconstructed a multi‐decadal record of δ¹⁵N and δ¹³C variation. Using established correlations between stable isotope ratios and sea bottom temperature we created a proxy for baseline isotopic variability to account for this source of temporal change. We found 1) a significant long‐term decline in juvenile grey seal δ¹⁵N values, suggesting trophic position has decreased over time; 2) a decline in adult δ¹⁵N values and contraction in stable isotopic niche space after the North Sea regime shift, signifying both a decline in trophic position and change in foraging habits over the 20th century; and 3) evidence for dietary segregation between juvenile and adult animals, showing juvenile individuals feeding at a lower trophic position and in more nearshore areas than adults. Our results demonstrate the efficacy of mining archived biological samples to address ecological questions and imply important ontogenetic and long‐term shifts in the feeding ecology of a top predator. Long‐term changes in grey seal trophic dynamics may be partly in response to well documented ecosystem changes in the North Sea. Such indirect monitoring of marine predators may have utility when set in the context of ecosystem assessments where paucity of long‐term monitoring data is prevalent.