Resourceful heterotrophs make the most of light in the coastal ocean

The carbon cycle in the coastal ocean is affected by how heterotrophic marine bacterioplankton obtain their energy. Although it was previously thought that these organisms relied on the organic carbon in seawater for all of their energy needs, several recent discoveries now suggest that pelagic bacteria can depart from a strictly heterotrophic lifestyle by obtaining energy through unconventional mechanisms that are linked to the penetration of sunlight into surface waters. These newly discovered mechanisms involve the harvesting of energy, either directly from light or indirectly from inorganic compounds that are formed when dissolved organic carbon absorbs light. In coastal systems, these mixed metabolic strategies have implications for how efficiently organic carbon is retained in the marine food web and how climatically important gases are exchanged between the ocean and the atmosphere.     

The production-to-respiration ratio and its implication in Lake Biwa,Japan

Production-to-respiration (P:R) ratio was estimated at an offshore site of Lake Biwa in order to examine whether the plankton and benthic community is subsidized with allochthonous organic carbon, and to clarify the role of this lake as potential source or sink of carbon dioxide. The respiration rate of protozoan and metazoan plankton was calculated from their biomass and empirical equations of oxygen consumption rates, and that of bacterioplankton was derived from their production rate and growth efficiency. In addition, the carbon mineralization rate in the lake sediments was estimated from the accumulation rate of organic carbon, which was determined using a ²¹⁰Pb dating technique. On an annual basis, the sum of respiration rates of heterotrophic plankton was comparable to net primary production rate measured by the ¹³C method. However, when the mineralization rate in the lake sediments was included, the areal P:R ratio was 0.89, suggesting that Lake Biwa is net heterotrophic at the offshore site with the community being subsidized with allochthonous organic carbon. Such a view was supported by the surface water pCO₂ that was on average higher than that of the atmosphere. However, the estimate of net CO₂ release rate was close to that of carbon burial rate in the sediments. The result suggests that the role of Lake Biwa in relation to atmospheric carbon is almost null at the offshore site, although the community is supported partially by organic carbon released from the surrounding areas.     

Net Ecosystem Production and Carbon Greenhouse Gas Fluxes in Three Prairie Wetlands

In central North America, prairie wetlands provide many important ecosystem services including attenuating floods, improving water quality, and supporting biodiversity. However, over half of these wetlands have been drained for agriculture. Relatively little is known about the functioning of these ecosystems either in their natural state or restored after drainage. We characterized net ecosystem production and carbon greenhouse gas (GHG) fluxes (carbon dioxide [CO₂] and methane) in the open-water zones of three prairie wetlands over two ice-free seasons. These wetlands included a natural site and sites restored 3 and 14 years prior to study (hereafter “recently restored” and “older restored”). We also assessed how two techniques for estimating metabolic status, the diel oxygen method (used to measure NEP) and net CO₂ fluxes, compared. The diel oxygen method suggested that the restored wetlands were net heterotrophic across years, whereas the natural wetland was net heterotrophic in 1 year and net autotrophic in the other. Similar conclusions arose from quantifying net CO₂ fluxes, although this technique proved to be relatively insensitive for understanding metabolic status at a daily temporal scale owing to the influence of geochemical processes on CO₂ concentrations. GHG efflux was greatest from the recently restored wetland, followed by the older restored and natural wetlands. Overall, GHG flux rates were high and variable compared with other inland aquatic ecosystems. Although restoration may progressively return wetland functioning to near-natural states, our results highlight the necessity of preventing wetland drainage in the first place to preserve ecosystem functions and services.     

Aquatic Microbiology for Ecosystem Scientists: New and Recycled Paradigms in Ecological Microbiology

In all ecosystems, bacteria are the most numerous organisms and through them flows a large fraction of annual primary production. In the past decade we have learned a great deal about some of the factors that regulate bacteria and their activities, and how these activities, in turn, alter ecosystem-level processes. Here I review three areas in which recent progress has been made with particular reference to pelagic ecosystems: the problem of bacterial cell dormancy; the effect of solar radiation on organic matter lability; and, the maintenance of net heterotrophy. In a system in which grazing is the major source of mortality for bacteria, bacterial cell dormancy is something of a paradox. I argue that the degree to which bacteria are grazed by flagellates (highly selective grazers) versus other grazers (cladocerans, bivalves) may explain the degree and variation in the proportion of active cells which recent evidence shows to be large. Another factor affecting bacterial activity that has come to the fore in recent years is solar radiation. Irradiation, especially in the ultra-violet range has long been thought of as simply deleterious to some bacteria. A wealth of newer evidence shows that refractory dissolved organic compounds may be converted into microbially labile compounds by solar radiation in several wavebands. This interaction between irradiation and organic matter (photolysis) may explain, in part, how dissolved organic carbon (C) may be refractory in the dark environment of the soil but become labile in the illumunated surface waters of lakes or rivers. The newer evidence shows that aquatic ecosystems, at least oligotrophic ones, are significantly subsidized by terrestrially-produced organic matter. I review here multiple lines of evidence that suggest that freshwater ecosystems are predominantly systems which respire more organic C than they produce by photosynthesis, and are therefore net heterotrophic. While net heterotrophy is an interesting exception for terrestrial ecosystems, it appears to be commonplace for aquatic systems and represents an important linkage between terrestrial and aquatic ecosystems.     

Pelagic food webs: Responses to environmental processes and effects on the environment

Changes in both the environment and environmental research have led to the development of new protocols and approaches. These new approaches consider both the effects of changes in the global environment on living organisms (i.e. the responses of ecosystems to environmental processes) and the feedback responses of these organisms and ecosystems (i.e. the effects of living organisms on the environment). The present paper focuses on pelagic food webs in aquatic ecosystems. We examine three major effects of global environmental changes on aquatic organisms: (i) the release of pollutants and biological agents in lakes and nearshore marine waters; (ii) the loss of biodiversity and the collapse of commercially exploited resources that were heretofore renewable. We develop detailed examples of the effects of human activities on marine organisms (i.e. the effects of nutrient supply on the structure of pelagic food webs in marine systems. Finally, we examine (iii) the food-web-controlled exchanges of CO₂ between the atmosphere and the ocean, as a feedback effect of pelagic ecosystems on the global environment with respect to the ongoing climate change.     

Terrestrial Subsidies of Organic Carbon Support Net Ecosystem Production in Temporary Forest Ponds: Evidence from an Ecosystem Experiment

Recent research suggests that secondary production in aquatic systems can be driven by inputs of energy from terrestrial sources. Temporary forest ponds appear to be unproductive ecosystems that are reliant upon allochthonous inputs of energy to support secondary production, but the functioning of these systems has not been well quantified. To assess the metabolic state of this type of ecosystem as well as to quantify the importance of terrestrial subsidies of carbon to ecosystem function, we conducted an experiment in which we manipulated the amount of leaf litter in ponds. Litter was either removed or removed and replaced (that is, control) from the dry basins of ponds immediately after leaf abscission. Once the ponds filled, we monitored net ecosystem production (NEP) on a biweekly basis from 9 April to 27 May 2002. All ponds were consistently net heterotrophic; however, NEP was significantly less negative in removal ponds. Furthermore, removal ponds also had lower levels of respiration (R) and higher dissolved oxygen levels than control ponds. The removal of litter had no effect on gross primary production, indicating that the difference in NEP between treatments was driven by the change in R. Therefore, it appears that terrestrial inputs of organic carbon support heterotrophic respiration in these ponds, and that the endogenous production of carbon is insufficient to support secondary production.     

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.     

Role of lakes for organic carbon cycling in the boreal zone

We calculated the carbon loss (mineralization plus sedimentation) and net CO₂ escape to the atmosphere for 79 536 lakes and total running water in 21 major Scandinavian catchments (size range 437–48 263 km²). Between 30% and 80% of the total organic carbon that entered the freshwater ecosystems was lost in lakes. Mineralization in lakes and subsequent CO₂ emission to the atmosphere was by far the most important carbon loss process. The withdrawal capacity of lakes on the catchment scale was closely correlated to the mean residence time of surface water in the catchment, and to some extent to the annual mean temperature represented by latitude. This result implies that variation of the hydrology can be a more important determinant of CO₂ emission from lakes than temperature fluctuations. Mineralization of terrestrially derived organic carbon in lakes is an important regulator of organic carbon export to the sea and may affect the net exchange of CO₂ between the atmosphere and the boreal landscape.     

Interaction of ocean and biosphere in their transient responses to increasing atmospheric CO2

Increasing atmospheric CO₂ induces a net uptake of carbon in the ocean by a shift in chemical equilibrium in seawater, and in the terrestrial biosphere by a stimulated photosynthesis and productivity. The fractions absorbed in both biosphere and ocean decline with increasing dynamics of the release rate of CO₂ into the atmosphere. However, the relative portion of ocean absorption descends much faster with annual growth rate of CO₂ release than biospheric absorption does, due to a difference in dynamics. The equilibrium absorption capacity of the biosphere is estimated to be only one quarter of that of the ocean, but the current sink size of the biosphere is about half of that of the ocean.Apart from CO₂-stimulated carbon fixation, the biosphere releases CO₂ as a result of land use changes, in particular after deforestation. Both of these fluxes are of the order of 1–1.5 Pg of carbon per year. The CO₂-fertilization effect and regrowth together have turned the terrestrial biosphere as a whole from a source into a sink.     

Paradigm shifts in soil organic matter research affect interpretations of aquatic carbon cycling: transcending disciplinary and ecosystem boundaries

New conceptual models that highlight the importance of environmental, rather than molecular, controls on soil organic matter affect interpretations of organic matter (OM) persistence across terrestrial and aquatic boundaries. We propose that changing paradigms in our thinking about OM decomposition explain some of the uncertainties surrounding the fate of land-derived carbon (C) in marine environments. Terrestrial OM, which historically has been thought to be chemically recalcitrant to decay in soil and aquatic environments, dominates inputs to rivers yet is found in trace amounts in the ocean. We discuss three major transformations in our understanding of OM persistence that influence interpretations of the fate of aquatic OM: (1) a shift away from an emphasis on chemical recalcitrance as a primary predictor of turnover; (2) new interpretations of radiocarbon ages, which affect predictions of reactivity; and (3) the recognition that most OM leaving soils in dissolved form has been microbially processed. The first two explain rapid turnover for terrigenous OM in aquatic ecosystems once it leaves the soil matrix. The third suggests that the presence of terrestrial OM in aquatic ecosystems may be underestimated by the use of plant biomarkers. Whether these mechanisms occur in isolation of each other or in combination, they provide insight into the missing terrestrial C signature in the ocean. Spatially and temporally varying transformations of OM along land–water networks require that common terrestrial source indicators be interpreted within specific environmental contexts. We identify areas of research where collaborations between aquatic and terrestrial scientists will enhance quantification of C transfer from soils to inland water bodies, the ocean, and the atmosphere. Accurate estimates of OM processing are essential for improving predictions of the response of vulnerable C pools at the interface of soil and water to changes in climate and land use.     

Metabolic balance of streams draining urban and agricultural watersheds in central Japan

Empirical data that describe the metabolic balance of stream ecosystems in human-dominated watersheds are scarce. We measured ecosystem metabolism in 23 open-canopied lowland streams draining urban and agricultural areas in the Fuji River Basin, central Japan. Gross primary production (GPP) and community respiration (CR) were estimated using the diurnal dissolved oxygen (DO) change technique, with the reaeration coefficient (K ₂) determined from seven empirical depth-velocity equations. Because the predicted values of K ₂ showed variation among the depth-velocity equations, the estimates of stream metabolism also varied according to the equations. However, CR was almost always greater than GPP, resulting in negative net ecosystem production (NEP) and GPP/CR ratios below unity for most of the study reaches. Highly heterotrophic streams were found in intensively farmed watersheds, suggesting that organic matter loading from agricultural lands is likely to be a source of allochthonous carbon fueling excess respiration in the study streams. In contrast, streams draining more urbanized areas were less heterotrophic. The present results suggest that lowland streams in agriculturally developed watersheds are associated strongly with terrestrial ecosystems as a source of organic carbon. The resultant strong respiration might become the dominant process in ecosystem metabolism, as reported for headwater streams, large downstream rivers, and estuaries.     

Biogeochemistry of organic carbon, CO2, CH4, and trace elements in thermokarst water bodies in discontinuous permafrost zones of Western Siberia

Active processes of permafrost thaw in Western Siberia increase the number of soil subsidencies, thermokarst lakes and thaw ponds. In continuous permafrost zones, this process promotes soil carbon mobilisation to water reservoirs, as well as organic matter (OM) biodegradation, which produces a permanent flux of carbon dioxide (CO₂) to the atmosphere. At the same time, the biogeochemical evolution of aquatic ecosystems situated in the transition zone between continuous permafrost and permafrost-free terrain remains poorly known. In order to better understand the biogeochemical processes that occur in thaw ponds and lakes located in discontinuous permafrost zones, we studied ~30 small (1–100,000 m²) shallow (<1 m depth) lakes and ponds formed as a result of permafrost subsidence and thaw of the palsa bog located in the transition zone between the tundra and forest-tundra (central part of Western Siberia). There is a significant increase in dissolved CO₂ and methane (CH₄) concentration with decreasing water body surface area, with the largest supersaturation with respect to atmospheric CO₂ and CH₄ in small (<100 m²) permafrost depressions filled with thaw water. Dissolved organic carbon (DOC), conductivity, and metal concentrations also progressively increase from large lakes to thaw ponds and depressions. As such, small water bodies with surface areas of 1–100 m² that are not accounted for in the existing lake and pond databases may significantly contribute to CO₂ and CH₄ fluxes to the atmosphere, as well as to the stocks of dissolved trace elements and organic carbon. In situ lake water incubation experiments yielded negligible primary productivity but significant oxygen consumption linked to the mineralisation rate of dissolved OM by heterotrophic bacterioplankton, which produce a net CO₂ flux to the atmosphere of 5 ± 2.5 mol C m² year^?1. The most significant result of this study, which has long-term consequences on our prediction of aquatic ecosystem development in the course of permafrost degradation is CO₂, CH₄, and DOC concentrations increase with decreasing lake age and size. As a consequence, upon future permafrost thaw, the increase in the number of small water bodies, accompanied by the drainage of large thermokarst lakes to the hydrological network, will likely favour (i) the increase of DOC and colloidal metal stocks in surface aquatic systems, and (ii) the enhancement of CO₂ and CH₄ fluxes from the water surface to the atmosphere. According to a conservative estimation that considers that the total area occupied by water bodies in Western Siberia will not change, this increase in stocks and fluxes could be as high as a factor of ten.     

Floating Aquatic Macrophytes Can Substantially Offset Open Water CO<Subscript>2</Subscript> Emissions from Tropical Floodplain Lake Ecosystems

Tropical floodplain lake ecosystems are recognized as important sources of carbon (C) from the water to the atmosphere. They receive large amounts of organic matter and nutrients from the watershed, leading to intense net heterotrophy and carbon dioxide (CO₂) emission from open waters. However, the role of extensive stands of floating macrophytes colonizing floodplains areas is still neglected in assessments of net ecosystem exchange of CO₂ (NEE). We assessed rates of air-lake CO₂ flux using static chambers in both open waters and waters covered by the widespread floating aquatic macrophyte (water hyacinth; Eichornia sp.) in two tropical floodplain lakes in Pantanal, Brazil during different hydrological seasons. In both lakes, areas colonized by floating macrophytes were a net CO₂ sink during all seasons. In contrast, open waters emitted CO₂, with higher emissions during the rising and high water periods. Our results indicate that the lake NEE can be substantially overestimated (fivefold or more in the studied lakes) if the carbon fixation by macrophytes is not considered. The contribution of these plants can lead to neutral or negative NEE (that is, net uptake of CO₂) on a yearly basis. This highlights the importance of floating aquatic macrophytes for the C balance in shallow lakes and extensive floodplain areas.     

Small Players, Large Role: Microbial Influence on Biogeochemical Processes in Pelagic Aquatic Ecosystems

Although prokaryotes are small in size, they are a significant biomass component in aquatic planktonic ecosystems and play a major role in biogeochemical processes. A review of the recent literature shows that the relative importance of prokaryotes to material and energy fluxes is maximized in low-productivity (oligotrophic) ecosystems and decreases in high-productivity (eutrophic) ecosystems. We conclude that competition with eukaryotic autotrophs for dissolved nutrients and competition with phagotrophic heterotrophs and physical processes (sinking, photooxidation) for organic carbon (C) play important roles in determining the relative abundance and impact of prokaryotes in aquatic systems. Oligotrophic systems have low nutrient concentrations, with high proportions of dissolved nutrients in organic form, which favors prokaryotic heterotrophs over phytoplankton. Furthermore, a high proportion of the available organic C is dissolved rather than particulate, which favors prokaryotic heterotrophs over phagotrophic heterotrophs. In eutrophic systems, increased relative concentrations and loading of inorganic nutrients and increased relative concentrations of particulate organic C select for phytoplankton and phagotrophic heterotrophs over prokaryotic heterotrophs. Increased particle sinking fluxes and/or decreased excretion of organic carbon (EOC) may also decrease the relative importance of prokaryotic heterotrophs in eutrophic systems. In oligotrophic systems, interactions between autotrophs and heterotrophs are tightly coupled because the dominant heterotrophs are similar in size and growth rates, as well as having similar nutrient composition to the dominant autotrophs, small phytoplankton. In eutrophic systems, increased productivity passes through zooplankton that are larger and have slower growth rates than the autotrophs, leading to a greater potential for decoupled auto- and heterotrophic production and increased export production. Received 18 July 2000; Accepted 13 September 2001.     

Microbial carbon limitation: The need for integrating microorganisms into our understanding of ecosystem carbon cycling

Numerous studies have demonstrated that fertilization with nutrients such as nitrogen, phosphorus, and potassium increases plant productivity in both natural and managed ecosystems, demonstrating that primary productivity is nutrient limited in most terrestrial ecosystems. In contrast, it has been demonstrated that heterotrophic microbial communities in soil are primarily limited by organic carbon or energy. While this concept of contrasting limitations, that is, microbial carbon and plant nutrient limitation, is based on strong evidence that we review in this paper, it is often ignored in discussions of ecosystem response to global environment changes. The plant‐centric perspective has equated plant nutrient limitations with those of whole ecosystems, thereby ignoring the important role of the heterotrophs responsible for soil decomposition in driving ecosystem carbon storage. To truly integrate carbon and nutrient cycles in ecosystem science, we must account for the fact that while plant productivity may be nutrient limited, the secondary productivity by heterotrophic communities is inherently carbon limited. Ecosystem carbon cycling integrates the independent physiological responses of its individual components, as well as tightly coupled exchanges between autotrophs and heterotrophs. To the extent that the interacting autotrophic and heterotrophic processes are controlled by organisms that are limited by nutrient versus carbon accessibility, respectively, we propose that ecosystems by definition cannot be ‘limited’ by nutrients or carbon alone. Here, we outline how models aimed at predicting non‐steady state ecosystem responses over time can benefit from dissecting ecosystems into the organismal components and their inherent limitations to better represent plant–microbe interactions in coupled carbon and nutrient models.     

Warming,eutrophication, and predator loss amplify subsidies between aquatic and terrestrial ecosystems

The exchange of organisms and energy among ecosystems has major impacts on food web structure and dynamics, yet little is known about how climate warming combines with other pervasive anthropogenic perturbations to affect such exchanges. We used an outdoor freshwater mesocosm experiment to investigate the interactive effects of warming, eutrophication, and changes in top predators on the flux of biomass between aquatic and terrestrial ecosystems. We demonstrated that predatory fish decoupled aquatic and terrestrial ecosystems by reducing the emergence of aquatic organisms and suppressing the decomposition of terrestrial plant detritus. In contrast, warming and nutrients enhanced cross‐ecosystem exchanges by increasing emergence and decomposition, and these effects were strongest in the absence of predators. Furthermore, we found that warming advanced while predators delayed the phenology of insect emergence. Our results demonstrate that anthropogenic perturbations may extend well beyond ecosystem boundaries by influencing cross‐ecosystem subsidies. We find that these changes are sufficient to substantially impact recipient communities and potentially alter the carbon balance between aquatic and terrestrial ecosystems and the atmosphere.     

Biodiversity and ecosystem productivity: implications for carbon storage

Recent experiments have found that Net Primary Productivity (NPP) can often be a positive saturating function of plant species and functional diversity. These findings raised the possibility that more diverse ecosystems might store more carbon as a result of increased photosynthetic inputs. However, carbon inputs will not only remain in plant biomass, but will be translocated to the soil via root exudation, fine root turnover, and litter fall. Thus, we must consider not just plant productivity (NPP), but also net productivity of the whole ecosystem (NEP), which itself measures net carbon storage. We currently know little about how plant diversity could influence soil processes that return carbon back to the atmosphere, such as heterotrophic respiration and decomposition of organic matter. Nevertheless, it is clear that any effects on such processes could make NPP a poor predictor of whole-ecosystem productivity, and potentially the ability of the ecosystem to store carbon. We examine the range of mechanisms by which plant diversity could influence net ecosystem productivity, incorporating processes involved with carbon uptake (productivity), loss (autotrophic and heterotrophic respiration), and residence time within the system (decomposition rate). Understanding the relationship between plant diversity and ecosystem carbon dynamics must be made a research priority if we wish to provide information relevant to global carbon policy decisions. This goal is entirely feasible if we utilize some basic methods for measuring the major fluxes of carbon into and out of the ecosystem.     

Non-phototrophic CO 2 fixation by soil microorganisms

Although soils are generally known to be a net source of CO₂ due to microbial respiration, CO₂ fixation may also be an important process. The non-phototrophic fixation of CO₂ was investigated in a tracer experiment with ¹⁴CO₂ in order to obtain information about the extent and the mechanisms of this process. Soils were incubated for up to 91 days in the dark. In three independent incubation experiments, a significant transfer of radioactivity from ¹⁴CO₂ to soil organic matter was observed. The process was related to microbial activity and could be enhanced by the addition of readily available substrates such as acetate. CO₂ fixation exhibited biphasic kinetics and was linearly related to respiration during the first phase of incubation (about 20–40 days). The fixation amounted to 3–5% of the net respiration. After this phase, the CO₂ fixation decreased to 1–2% of the respiration. The amount of carbon fixed by an agricultural soil corresponded to 0.05% of the organic carbon present in the soil at the beginning of the experiment, and virtually all of the fixed CO₂ was converted to organic compounds. Many autotrophic and heterotrophic biochemical processes result in the fixation of CO₂. However, the enhancement of the fixation by addition of readily available substrates and the linear correlation with respiration suggested that the process is mainly driven by aerobic heterotrophic microorganisms. We conclude that heterotrophic CO₂ fixation represents a significant factor of microbial activity in soils.     

水生生态系统的碳循环及对大气CO2的汇

水生生态系统,特别是海洋无疑是大气CO2的一个巨大的汇。海洋对大气CO2的汇以及大气圈和海洋之间碳的变换量在很大程度上取于混合层碳酸盐化学、水中溶解碳的平流传输、CO2通过空气－－海水界面的扩散、海洋生物生产和所产生的有面碳化合物的沉隆等,现在已建立和发展了多种海洋碳子模型以对CO2的汇进行估测。根据国内外研究资料,综述了水生生态系统碳循环过程及“生物泵”作用机制等方面的研究进展;介绍了两大类主要的海洋碳子模型：厢式模型和普通环流模型,采用这些模型对海洋碳汇的估算约为1.2-2.4GtC/a;分析了湖泊、河流等对大气CO2汇的特点及向海洋的转移,并对影响水体生态系统碳循环和大气CO2汇的因素进行了讨论。     

A coupled atmosphere–ecosystem model of the early Archean Earth

A coupled photochemical‐ecosystem model has been developed to simulate the early Archean biosphere. The model incorporates kinetic and nutrient limitations on biological productivity, along with constraints imposed by metabolic thermodynamics. We have used this model to predict the biogenic CH₄ flux and net primary productivity (NPP) of the marine biosphere prior to the advent of oxygenic photosynthesis. Organisms considered include chemotrophic and organotrophic methanogens, H₂‐, H₂S‐, and Fe‐using anoxygenic phototrophs, S‐reducing bacteria, CO‐using acetogens, and fermentative bacteria. CH₄ production and NPP in our model are limited by the downward flux of H₂, CO, S₈, and H₂S through the atmosphere–ocean interface and by the upwelling rate of Fe²⁺ from the deep oceans. For reasonable estimates of the supply rates of these compounds, we find that the biogenic CH₄ flux should have ranged from approximately ¹/₃ to 2.5 times the modern CH₄ flux. In the anoxic Archean atmosphere, this would have produced CH₄ concentrations of 100 ppmv to as much as 35 000 ppmv (3.5%), depending on the rate at which hydrogen escaped to space. Recent calculations indicating that hydrogen escape was slow favour the higher CH₄ concentrations. Calculated NPP is lower than in the modern oceans by a factor of at least 40. In our model, H₂‐based metabolism is moderately more productive than Fe²⁺‐based metabolism, with S‐based metabolism being considerably less productive. Internal recycling of sulphur within the surface ocean could conceivably raise rates of sulphur metabolism by a factor of 10 higher than the values predicted by our model. Although explicit climate calculations have not been performed here, our results are consistent with the idea that the Archean climate was warm, and possibly very hot. Some or most of our ecosystem scenarios are consistent with the carbon isotope record, depending on how that record is interpreted. If the conventional view is correct and organic carbon burial accounted for approximately 20% of total carbon burial during the Archean, then only two of our phototroph‐based model ecosystems are plausible. However, if a recent alternative analysis is correct and only approximately 0–10% of total buried carbon was organic, then essentially all of our anaerobic ecosystems are plausible. A better understanding of both the geochemical and the biological records is needed to better constrain our models.