Fluxes of terrestrial and aquatic carbon by emergent mosquitoes: a test of controls and implications for cross-ecosystem linkages

Adult aquatic insects are a common resource for many terrestrial predators, often considered to subsidize terrestrial food webs. However, larval aquatic insects themselves consume both aquatic primary producers and allochthonous terrestrial detritus, suggesting that adults could provide aquatic subsidy and/or recycled terrestrial energy to terrestrial consumers. Understanding the source of carbon (aquatic vs. terrestrial) driving aquatic insect emergence is important for predicting magnitude of emergence and effects on recipient food web dynamics; yet direct experimental tests of factors determining source are lacking. Here, we use Culex mosquitoes in experimental pools as an exemplar to test how variation in general factors common to aquatic systems (terrestrial plant inputs and light) may alter the source and amount of energy exported to terrestrial ecosystems in adult aquatic insects that rely on terrestrial resources as larvae. We found strong sequential effects of terrestrial plant inputs and light on aquatic insect oviposition, diet, and emergence of Culex mosquitoes. Ovipositing mosquitoes laid ~3 times more egg masses in high terrestrial input pools under low light conditions. This behavior increased adult emergence from pools under low light conditions; however, high input pools (which had the highest mosquito densities) showed low emergence rates due to density-dependent mortality. Mosquito diets consisted mainly of terrestrial resources (~70–90 %). As a result, the amount of aquatic carbon exported from pools by mosquitoes during the experiment was ~18 times higher from low versus high light pools, while exports of terrestrial carbon peaked from pools receiving intermediate levels of inputs (3–6 times higher) and low light (~6 times higher). Our results suggest that understanding the interplay among terrestrial plant inputs, light availability and biotic responses of aquatic insects may be key in predicting source and magnitude of emergence, and thus the strength and effects of aquatic–terrestrial linkages in freshwater systems.     

Dissolved Organic Carbon in Terrestrial Ecosystems: Synthesis and a Model

The movement of dissolved organic carbon (DOC) through soils is an important process for the transport of carbon within ecosystems and the formation of soil organic matter. In some cases, DOC fluxes may also contribute to the carbon balance of terrestrial ecosystems; in most ecosystems, they are an important source of energy, carbon, and nutrient transfers from terrestrial to aquatic ecosystems. Despite their importance for terrestrial and aquatic biogeochemistry, these fluxes are rarely represented in conceptual or numerical models of terrestrial biogeochemistry. In part, this is due to the lack of a comprehensive understanding of the suite of processes that control DOC dynamics in soils. In this article, we synthesize information on the geochemical and biological factors that control DOC fluxes through soils. We focus on conceptual issues and quantitative evaluations of key process rates to present a general numerical model of DOC dynamics. We then test the sensitivity of the model to variation in the controlling parameters to highlight both the significance of DOC fluxes to terrestrial carbon processes and the key uncertainties that require additional experiments and data. Simulation model results indicate the importance of representing both root carbon inputs and soluble carbon fluxes to predict the quantity and distribution of soil carbon in soil layers. For a test case in a temperate forest, DOC contributed 25% of the total soil profile carbon, whereas roots provided the remainder. The analysis also shows that physical factors—most notably, sorption dynamics and hydrology—play the dominant role in regulating DOC losses from terrestrial ecosystems but that interactions between hydrology and microbial–DOC relationships are important in regulating the fluxes of DOC in the litter and surface soil horizons. The model also indicates that DOC fluxes to deeper soil layers can support a large fraction (up to 30%) of microbial activity below 40 cm. Received 14 January 2000; accepted 6 September 2000     

大气二氧化碳浓度升高对植物根际微生物及菌根共生体的影响

综述了大气CO2浓度升高条件下,植物根际土壤环境、根际土壤微生物和植物菌根形成的变化趋势等方面的研究进展,CO2浓度升高,运转到根系的碳水化合物增加,根际环境、根际微生物活性、微生物群落结构以及菌根共生体的形成发生变化．提出在CO2浓度升高条件下,根际微生物和菌根真菌群落的变化对植物群落和陆地生态系统碳动态的调节是今后的研究趋向。     

Virus ecology of fluvial systems: a blank spot on the map?

The ecology of viruses has been studied only in a limited number of rivers and streams. In light of a recent re‐appraisal of the global fluvial surface area, issues such as abundance and production, host mortality and the influence of suspended particles and biofilms are addressed. Viral life cycles, potential impacts of viruses on water biochemistry and carbon flow, and viral diversity are considered. Variability in trophic levels along with the heterogeneous nature and hydrological dynamics of fluvial environments suggest a prevailingly physical control of virus‐related processes under lotic conditions and more biological control under lentic conditions. Viral lysis likely contributes to a pool of rapidly cycling carbon in environments typically characterized by high proportions of recalcitrant terrestrial carbon. On average, 33.6% (equalling 0.605 Pg C year^?1) of the globally respired carbon from fluvial systems may pass through a viral loop. Virus distribution and the proportion of organic material in horizontal transport versus processes in retention zones remain to be determined in detail. The need for up‐scaling the contribution of virus‐related processes in fluvial systems is of global relevance. Further, the role of climate change and the effect of anthropogenic alterations of fluvial systems on viruses require attention. The identification of these considerable knowledge gaps should foster future research efforts.     

陆地生态系统碳平衡主要研究方法评述

陆地生态系统碳平衡是全球变化科学中的核心问题之一,目前也是生态科学中的前沿与热点问题,而陆地生态系统的复杂性与不确定性决定了对陆地生态系统碳平衡估测的复杂性和不确定性。为研究这一复杂性问题,已发展了许多研究方法。可分为“自下而上”与“自上而下”两种,各种方法都有其自身的优势和劣势。相关方向也已经有了大量的研究报道,但是,不同的研究由于在方法、时间与空间尺度等存在的差异,使得许多研究结果和预测很难被有效的整合或适用于大范围甚至全球水平。综述了陆地碳平衡的主要研究方法,分析和比较了各方法的特点,指出在研究中对不同方法的结果进行分析和比较,以及采用综合方法的必要性。     

The importance of rapid, disturbance-induced losses in carbon management and sequestration

Management of terrestrial carbon fluxes is being proposed as a means of increasing the amount of carbon sequestered in the terrestrial biosphere. This approach is generally viewed only as an interim strategy for the coming decades while other longer‐term strategies are developed and implemented — the most important being the direct reduction of carbon emissions. We are concerned that the potential for rapid, disturbance‐induced losses may be much greater than is currently appreciated, especially by the decision‐making community. Here we wish to: (1) highlight the complex and threshold‐like nature of disturbances — such as fire and drought, as well as the erosion associated with each — that could lead to carbon losses; (2) note the global extent of ecosystems that are at risk of such disturbance‐induced carbon losses; and (3) call for increased consideration of and research on the mechanisms by which large, rapid disturbance‐induced losses of terrestrial carbon could occur. Our lack of ability as a scientific community to predict such ecosystem dynamics is precluding the effective consideration of these processes into strategies and policies related to carbon management and sequestration. Consequently, scientists need to do more to improve quantification of these potential losses and to integrate them into sound, sustainable policy options.     

Stoichiometric flexibility as a regulator of carbon and nutrient cycling in terrestrial ecosystems under change

Ecosystems across the biosphere are subject to rapid changes in elemental balance and climatic regimes. A major force structuring ecological responses to these perturbations lies in the stoichiometric flexibility of systems - the ability to adjust their elemental balance whilst maintaining function. The potential for stoichiometric flexibility underscores the utility of the application of a framework highlighting the constraints and consequences of elemental mass balance and energy cycling in biological systems to address global change phenomena. Improvement in the modeling of ecological responses to disturbance requires the consideration of the stoichiometric flexibility of systems within and across relevant scales. Although a multitude of global change studies over various spatial and temporal scales exist, the explicit consideration of the role played by stoichiometric flexibility in linking micro-scale to macro-scale biogeochemical processes in terrestrial ecosystems remains relatively unexplored. Focusing on terrestrial systems under change, we discuss the mechanisms by which stoichiometric flexibility might be expressed and connected from organisms to ecosystems. We suggest that the transition from the expression of stoichiometric flexibility within individuals to the community and ecosystem scales is a key mechanism regulating the extent to which environmental perturbation may alter ecosystem carbon and nutrient cycling dynamics.     

Role of first-order consumers in ecosystem carbon flow

Herbivores and detritus consumers (i.e. microbial decomposers and invertebrate and vertebrate detritivores) are pivotal components of trophic food webs and thus play a paramount role in the trophic transference and turnover of producer‐fixed carbon. Hence, elucidating patterns in carbon flux through these first‐order consumers is important to understand the nature and controls of carbon flow in ecosystems. Here, using the largest literature compilation to date, I show that, in contrast with the current belief, aquatic herbivores accumulate on average three times as much biomass as do terrestrial herbivores for a given level of primary production and, as a consequence, turn over the ingested carbon only slightly faster than do terrestrial herbivores. Conversely, aquatic detritus consumers generally accumulate a much lower biomass (i.e. over ten times lower) than their terrestrial counterparts for a given level of primary production and, thus, they turn over the ingested carbon much more quickly (i.e. over ten times faster). Because the detrital pathway generally dominates the trophic flow of carbon in both aquatic and terrestrial ecosystems, carbon also tends to flow through the total compartment of first order‐consumers (both herbivores and detritus consumers) at a much faster rate in aquatic than in terrestrial ecosystems. Thus, aquatic systems, because of faster carbon recycling rates through their basal and first‐order levels of the food chain, appear to have a lower capacity than do terrestrial systems for retaining carbon under natural or anthropogenic increases in photosynthetic fixation.     

Leaf litter decomposition in an ephemeral karst lake (Chaney Lake,Kentucky, U.S.A.)

Litter decomposition in temporary aquatic environments has not been experimentally studied as much as it has in perennial systems. However, litter is likely a critical resource for organisms inhabiting ephemeral aquatic habitats. In this study, we used litterbags under different conditions of submergence and water physical and chemical properties/characteristics to study mass and nutrient losses of terrestrial materials in an ephemeral karst lake in south-central Kentucky (USA). In the first experiment, which was designed to compare decomposition rates in submerged and dry sites, total mass and carbon declined more rapidly in the litter at fully submerged sites than in dry sites. In the second experiment, which was designed to compare decomposition rates in two different submerged environments, total mass and carbon showed similar decomposition trends between the two submerged areas with different seasonal temperature patterns. Nitrogen patterns were variable but in general nitrogen levels increased in the litter in both experiments over a period of several months. These results are similar to those found in some perennially inundated systems and indicate that litter decomposition dynamics in this temporary lake can be greatly affected by lake hydrology. Year-to-year variations in hydrology may thus have strong impacts on nutrient and energy release within this system, which may affect the organisms within this karst lake and in other areas of the karst ecosystem that are ecologically connected to it.     

陆地碳循环的动态非平衡假说

生态系统维持物质与能量的动态平衡是地球系统孕育与维持生物多样性的重要基础。自工业革命以来, 人类活动导致陆地生态系统的碳循环转变为动态非平衡,进而使陆地生态系统的结构与功能出现许多难以预测的变化动态。本文阐释了陆地生态系统碳循环的动态非平衡假说。该假说构建于陆地碳循环内部过程的四点基本特征和五类外部驱动因素。基于这些内部特征与外部因素, 本文归纳了陆地生态系统碳循环动态非平衡在不同时间与空间尺度的表达现象, 并从观测、实验与模型的角度讨论了其检测方法。陆地生态系统碳循环的动态非平衡假说不仅有助于我们理解复杂的陆地碳循环现象, 也为预测未来陆地碳汇动态提供了新的理论框架。     

Extracellular enzymes in terrestrial, freshwater, and marine environments: perspectives on system variability and common research needs

Extracellular enzymes produced by heterotrophic microbial communities are major drivers of carbon and nutrient cycling in terrestrial, freshwater, and marine environments. Although carbon and nutrient cycles are coupled on global scales, studies of extracellular enzymes associated with terrestrial, freshwater, and marine microbial communities are not often compared across ecosystems. In part, this disconnect arises because the environmental parameters that control enzyme activities in terrestrial and freshwater systems, such as temperature, pH, and moisture content, have little explanatory power for patterns of enzyme activities in marine systems. Instead, factors such as the functional diversity of microbial communities may explain varying patterns of enzyme activities observed in the ocean to date. In any case, many studies across systems focus on similar issues that highlight the commonalities of microbial community organization. Examples include the effective lifetime of enzymes released into the environment; the extent to which microbial communities coordinate enzyme expression to decompose complex organic substrates; and the influence of microbial community composition on enzyme activities and kinetics. Here we review the often-disparate research foci in terrestrial, freshwater, and marine environments. We consider the extent to which environmental factors may regulate extracellular enzyme activities within each ecosystem, and highlight commonalities and current methodological challenges to identify research questions that may aid in integrating cross-system perspectives in the future.     

磷影响陆地生态系统碳循环过程及模型表达方法

全球气候变暖已大大改变了陆地植物碳吸收能力, 提高了全球植被净初级生产力。随着气候变暖的加剧, 磷对植物生长的限制作用逐渐显现且不断增强, 磷影响陆地生态系统碳循环的机理和模型研究已成为研究热点。该文系统分析了磷影响陆地生态系统碳循环的相关机理以及模型对相关过程的定量化表达方法。综合对比分析了国际上的Carnegie- Ames-Stanford Approach-CNP (CASA-CNP)、Community Land Model-CNP (CLM-CNP)和Jena Scheme for Biosphere-Atmosphere Coupling in Hamburg-CNP (JSBACH-CNP)等碳、氮、磷耦合模型中磷影响植物光合作用与同化物分配过程、植物对磷的吸收过程、土壤中磷的转化过程以及生态系统磷输入与输出等过程的相关数学表达方法, 指出了模型算法的局限与不确定性以及未来模型发展与改进的方向。同时综合对比分析了CASA-CNP、CLM-CNP、JSBACH-CNP模型的基本特征, 总结了磷循环模型的建模方法, 为未来开展磷影响陆地生态系统碳循环的模型模拟研究提供了借鉴方法与参考思路。     

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.     

Plumbing the Global Carbon Cycle: Integrating Inland Waters into the Terrestrial Carbon Budget

ABSTRACT Because freshwater covers such a small fraction of the Earth’s surface area, inland freshwater ecosystems (particularly lakes, rivers, and reservoirs) have rarely been considered as potentially important quantitative components of the carbon cycle at either global or regional scales. By taking published estimates of gas exchange, sediment accumulation, and carbon transport for a variety of aquatic systems, we have constructed a budget for the role of inland water ecosystems in the global carbon cycle. Our analysis conservatively estimates that inland waters annually receive, from a combination of background and anthropogenically altered sources, on the order of 1.9 Pg C y⁻¹ from the terrestrial landscape, of which about 0.2 is buried in aquatic sediments, at least 0.8 (possibly much more) is returned to the atmosphere as gas exchange while the remaining 0.9 Pg y⁻¹ is delivered to the oceans, roughly equally as inorganic and organic carbon. Thus, roughly twice as much C enters inland aquatic systems from land as is exported from land to the sea. Over prolonged time net carbon fluxes in aquatic systems tend to be greater per unit area than in much of the surrounding land. Although their area is small, these freshwater aquatic systems can affect regional C balances. Further, the inclusion of inland, freshwater ecosystems provides useful insight about the storage, oxidation and transport of terrestrial C, and may warrant a revision of how the modern net C sink on land is described.     

Carbon storage in terrestrial ecosystems: do browsing and grazing herbivores matter?

Large mammalian herbivores manifest a strong top‐down control on ecosystems that can transform entire landscapes, but their impacts have not been reviewed in the context of terrestrial carbon storage. Here, we evaluate the effects of plant biomass consumption by large mammalian herbivores (>10 kg adult biomass), and the responses of ecosystems to these herbivores, on carbon stocks in temperate and tropical regions, and the Arctic. We calculate the difference in carbon stocks resulting from herbivore exclusion using the results of 108 studies from 52 vegetation types. Our estimates suggest that herbivores can reduce terrestrial above‐ and below‐ground carbon stocks across vegetation types but reductions in carbon stocks may approach zero given sufficient periods of time for systems to respond to herbivory (i.e. decades). We estimate that if all large herbivores were removed from the vegetation types sampled in our review, increases in terrestrial carbon stocks would be up to three orders of magnitude less than many of the natural and human‐influenced sources of carbon emissions. However, we lack estimates for the effects of herbivores on below‐ground biomass and soil carbon levels in many regions, including those with high herbivore densities, and upwards revisions of our estimates may be necessary. Our results provide a starting point for a discussion on the magnitude of the effects of herbivory on the global carbon cycle, particularly given that large herbivores are common in many ecosystems. We suggest that herbivore removal might represent an important strategy towards increasing terrestrial carbon stocks at local and regional scales within specific vegetation types, since humans influence populations of most large mammals.     

森林生态系统碳循环对全球氮沉降的响应

森林土壤和植被储存着全球陆地生态系统大约46%的碳,在全球碳平衡中起着非常重要的作用。过去几十年来,森林生态系统的碳循环和碳吸存受到了全球氮沉降的深刻影响,因为氮沉降改变了陆地生态系统的生产力和生物量积累。以欧洲和北美温带森林区域开展的研究为基础,综述了氮沉降对植物光合作用、土壤呼吸、土壤DOM及林木生长的影响特征和机理,探讨了森林生态系统碳动态对氮沉降响应的不确定性因素。热带森林C、N循环与大部分温带森林不同,人为输入的氮对热带生态系统过程的影响也可能不同,因此指出了在热带地区开展碳氮循环耦合研究的必要性和紧迫性。     

Carbon and nitrogen dynamics during forest stand development: a global synthesis

Our knowledge of carbon (C) and nitrogen (N) dynamics during stand development is not only essential for evaluating the role of secondary forests in the global terrestrial C cycle, but also crucial for understanding long-term C-N interactions in terrestrial ecosystems. However, a comprehensive understanding of forest C and N dynamics over age sequence remains elusive due to the diverse results obtained across individual studies. Here, we synthesized the results of more than 100 studies to examine C and N dynamics during forest stand development. Our results showed that C accumulated in aboveground vegetation, litter and forest floor pools, while the mineral soil C pool did not exhibit significant changes in most studies. The rate of C changes declined with stand age and approached equilibrium during the later stage of stand development. The rate of N changes exhibited linear increases with that of C changes, indicating that N also accrued in various ecosystem components except mineral soil. These results demonstrate that substantial increases in C pools over age sequence are accompanied by N accretion in forest ecosystems. The concurrent C and N dynamics suggest that forest ecosystems may have an intrinsic ability to preclude progressive N limitation during stand development.     

Environmental dynamics of dissolved black carbon in wetlands

Wetlands are ecosystems commonly characterized by elevated levels of dissolved organic carbon (DOC), and although they cover a surface area less than 2 % worldwide, they are an important carbon source representing an estimated 15 % of global annual DOC flux to the oceans. Because of their unique hydrological characteristics, fire can be an important ecological driver in pulsed wetland systems. Consequently, wetlands may be important sources not only of DOC but also of products derived from biomass burning, such as dissolved black carbon (DBC). However, the biogeochemistry of DBC in wetlands has not been studied in detail. The objective of this study is to determine the environmental dynamics of DBC in different fire-impacted wetlands. An intensive, 2-year spatial and temporal dynamics study of DBC in a coastal wetland, the Everglades (Florida) system, as well as one-time sampling surveys for the other two inland wetlands, Okavango Delta (Botswana) and the Pantanal (Brazil), were reported. Our data reveal that DBC dynamics are strongly coupled with the DOC dynamics regardless of location, season or recent fire history. The statistically significant linear regression between DOC and DBC was applied to estimate DBC fluxes to the coastal zone through two main riverine DOC export routes in the Everglades ecosystem. The presence of significant amounts of DBC in these three fire-impacted ecosystems suggests that sub-tropical wetlands could represent an important continental-ocean carrier of combustion products from biomass burning. The discrimination of DBC molecular structure (i.e. aromaticity) between coastal and terrestrial samples, and between samples collected in wet and dry season, suggests that spatially-significant variation in DBC source strength and/or degree of degradation may also influence DBC dynamics.     

New insights into enzymes in the environment

This article introduces the thematic special issue “Enzymes in the Environment: Incorporating Enzymes and Microbial Physiology into Biogeochemical Models”. This collection of review articles, original studies, and models stemmed from a workshop organized by the Enzymes in the Environment Research Coordination Network, which was held at Colorado State University in Fort Collins, CO, USA in May 2012. The meeting, themed “Incorporating Enzymes and Microbial Physiology into Biogeochemical Models” featured talks from both modelers and experimentalists with experience ranging across terrestrial, aquatic, and marine ecosystems. This special issue represents a series of integrative discussions that occurred among experimentalists and modelers at the meeting. This issue highlights that a better understanding of carbon cycling pathways and subsequent carbon availability dynamics among ecosystems may be achieved by simultaneously integrating aspects of terrestrial and aquatic carbon cycling and microbial physiology into biogeochemical cycling models.     

火干扰与生态系统的碳循环

火干扰是陆地生态系统碳循环的重要影响因子。它改变着整个系统的碳循环过程与碳分布格局。正确评估火干扰在碳循环过程中的作用,对推进全球碳循环研究有着重要的意义。从4个方面系统的回顾了火干扰对碳循环的影响过程及其研究方法:(1)火烧过程中含碳痕量气体排放的估算;(2)火烧迹地恢复过程中净第一性生产力(NPP)与土壤呼吸的变化;(3)火干扰对生态系统碳源/汇的影响;(4)模型方法在火干扰与生态系统碳循环研究中的应用。目前火灾碳排量的估算方法业已成熟,但进行更精确的估算必须基于对受干扰生态系统的性质以及火势的时空变异性质的准确理解;相比之下,对于间接的、更为重要的影响,即对火烧迹地恢复过程中碳循环变化的研究则显不足。由于数据缺乏,现有研究大多限于对碳循环某一方面的观测与定量描述,缺乏全面的机理性分析。对此,实地观测、模型模拟与遥感观测的跨尺度集成将成为未来火干扰研究的一个主要方向。