The possible roles of algae in restricting the increase in atmospheric CO2 and global temperature

ABSTRACT

Anthropogenic inputs are increasing the CO₂ content of the atmosphere, and the CO₂ and total inorganic C in the surface ocean and, to a lesser degree, the deep ocean. The greenhouse effect of the increased CO₂ (and, to a lesser extent, other greenhouse gases) is very probably the major cause of present global warming. The warming increases temperature of the atmosphere and the surface ocean to a greater extent than the deep ocean, with shoaling of the thermocline, decreasing nutrient flux to the surface ocean where there is greater mean photosynthetic photon flux density. These global changes influence algae in nature. However, it is clear that algae are important, via the biological pump, in decreasing the steady state atmospheric and ocean surface CO₂, and thus decreasing radiative forcing, a reduction enhanced by algal increases in albedo. As well as these natural processes there are possibilities that algae can, with human intervention, partly offset the increase in atmospheric CO₂. One possibility is to grow algae as sources of fuel for transport, in principle providing an energy source that is close to CO₂-neutral. The other possibility is to increase the role of algae in sequestering CO₂ as organic C over periods of hundreds or more years in the deep ocean and marine sediments and/or increasing albedo and decreasing radiative forcing of temperature. There are problems, currently unresolved, in the economically viable production of algal biofuels without carbon trading subsidies. Enhanced algal CO₂ sequestration also has costs, both in resource input (phosphorus (P) from high P content rocks, a limited resource with a competing use as an agricultural fertilizer) and adverse environmental effects. For example, ocean anoxic zones producing N₂O and increased algal production of short-lived halocarbons by algae that both, through breakdown, destroy O₃ and increase UV flux to the Earth’s surface.     

Significance of ocean carbonate budgets for the global carbon cycle

Changes in the trace gas composition of the atmosphere over glacial–interglacial cycles are linked to changes in the oceanic carbon cycle. This paper examines the role of biologically driven fluxes of organic and inorganic carbon in modifying the carbon dioxide chemistry of the oceans, and the corresponding implications for the partitioning of CO₂ between the atmosphere and ocean. Relevant details of the marine carbon system are presented together with an assessment of the significance of remineralization and dissolution processes. Recent estimates of the marine carbonate fluxes show significant uncertainties and inconsistencies which must be resolved in order to assess fully the role of the oceans' biota in the marine carbon system. Various types of ocean carbon cycle models have been developed in order to interpret the changes in past atmospheric carbon dioxide. Some take account of the role of the oceans' biota, focussing in the main on the cycling of organic matter. Relatively few have considered the role of the carbonate pump and the subtle interactions between organic and inorganic carbon cycling. The significance of carbonate formation and dissolution, and of the effects of global change on the marine carbonate system, for air–sea fluxes of CO₂ are discussed. Finally some recommendations for future research are made in order to improve our understanding of how spatial and temporal variation in marine carbonate fluxes, in conjunction with processes determining the oxidation and burial of organic matter in the oceans, affect levels of CO₂ in the atmosphere.     

Air-sea carbon dioxide equilibrium: Will it be possible to use seaweeds for carbon removal offsets?

To limit global warming below 2°C by 2100, we must drastically reduce greenhouse gas emissions and additionally remove ~100–900 Gt CO₂ from the atmosphere (carbon dioxide removal, CDR) to compensate for unavoidable emissions. Seaweeds (marine macroalgae) naturally grow in coastal regions worldwide where they are crucial for primary production and carbon cycling. They are being considered as a biological method for CDR and for use in carbon trading schemes as offsets. To use seaweeds in carbon trading schemes requires verification that seaweed photosynthesis that fixes CO₂ into organic carbon results in CDR, along with the safe and secure storage of the carbon removed from the atmosphere for more than 100 years (sequestration). There is much ongoing research into the magnitude of seaweed carbon storage pools (e.g., as living biomass and as particulate and dissolved organic carbon in sediments and the deep ocean), but these pools do not equate to CDR unless the amount of CO₂ removed from the atmosphere as a result of seaweed primary production can be quantified and verified. The draw-down of atmospheric CO₂ into seawater is via air-sea CO₂ equilibrium, which operates on time scales of weeks to years depending upon the ecosystem considered. Here, we explain why quantifying air-sea CO₂ equilibrium and linking this process to seaweed carbon storage pools is the critical step needed to verify CDR by discrete seaweed beds and nearshore and open ocean aquaculture systems prior to their use in carbon trading.     

Enhanced regional terrestrial carbon uptake over Korea revealed by atmospheric CO2 measurements from 1999 to 2017

Understanding changes in terrestrial carbon balance is important to improve our knowledge of the regional carbon cycle and climate change. However, evaluating regional changes in the terrestrial carbon balance is challenging due to the lack of surface flux measurements. This study reveals that the terrestrial carbon uptake over the Republic of Korea has been enhanced from 1999 to 2017 by analyzing long‐term atmospheric CO₂ concentration measurements at the Anmyeondo Station (36.53°N, 126.32°E) located in the western coast. The influence of terrestrial carbon flux on atmospheric CO₂ concentrations (ΔCO₂) is estimated from the difference of CO₂ concentrations that were influenced by the land sector (through easterly winds) and the Yellow Sea sector (through westerly winds). We find a significant trend in ΔCO₂ of ?4.75 ppm per decade (p < .05) during the vegetation growing season (May through October), suggesting that the regional terrestrial carbon uptake has increased relative to the surrounding ocean areas. Combined analysis with satellite measured normalized difference vegetation index and gross primary production shows that the enhanced carbon uptake is associated with significant nationwide increases in vegetation and its production. Process‐based terrestrial model and inverse model simulations estimate that regional terrestrial carbon uptake increases by up to 18.9 and 8.0 Tg C for the study period, accounting for 13.4% and 5.7% of the average annual domestic carbon emissions, respectively. Atmospheric chemical transport model simulations indicate that the enhanced terrestrial carbon sink is the primary reason for the observed ΔCO₂ trend rather than anthropogenic emissions and atmospheric circulation changes. Our results highlight the fact that atmospheric CO₂ measurements could open up the possibility of detecting regional changes in the terrestrial carbon cycle even where anthropogenic emissions are not negligible.     

Elevated CO2 enhances nitrogen fixation and growth in the marine cyanobacterium Trichodesmium

The increases in atmospheric pCO₂ over the last century are accompanied by higher concentrations of CO₂(aq) in the surface oceans. This acidification of the surface ocean is expected to influence aquatic primary productivity and may also affect cyanobacterial nitrogen (N)‐fixers (diazotrophs). No data is currently available showing the response of diazotrophs to enhanced oceanic CO₂(aq). We examined the influence of pCO₂ [preindustrial∼250 ppmv (low), ambient∼400, future∼900 ppmv (high)] on the photosynthesis, N fixation, and growth of Trichodesmium IMS101. Trichodesmium spp. is a bloom‐forming cyanobacterium contributing substantial inputs of ‘new N’ to the oligotrophic subtropical and tropical oceans. High pCO₂ enhanced N fixation, C : N ratios, filament length, and biomass of Trichodesmium in comparison with both ambient and low pCO₂ cultures. Photosynthesis and respiration did not change significantly between the treatments. We suggest that enhanced N fixation and growth in the high pCO₂ cultures occurs due to reallocation of energy and resources from carbon concentrating mechanisms (CCM) required under low and ambient pCO₂. Thus, in oceanic regions, where light and nutrients such as P and Fe are not limiting, we expect the projected concentrations of CO₂ to increase N fixation and growth of Trichodesmium. Other diazotrophs may be similarly affected, thereby enhancing inputs of new N and increasing primary productivity in the oceans.     

ELEVATED ATMOSPHERIC CARBON DIOXIDE INCREASES ORGANIC CARBON FIXATION BY EMILIANIA HUXLEYI (HAPTOPHYTA), UNDER NUTRIENT‐LIMITED HIGH‐LIGHT CONDITIONS1

Phytoplankton play a key role in determining the partitioning of CO₂ between the atmosphere and the ocean on seasonal, interannual, and millennial time scales. The magnitude of biological draw‐down of atmospheric CO₂ and C storage in the oceans is affected by concurrent changes in other environmental factors, like nutrient supply. Furthermore, variations in carbon‐to‐nitrogen (C:N) and carbon‐to‐phosphorus (C:P) assimilation ratios modify the oceanic CO₂ storage capacity. Here we show that increased atmospheric CO₂ concentration enhances CO₂ fixation into organic matter by a noncalcifying strain of Emiliania huxleyi (Lohmann) Hay & Mohler only under certain conditions, namely high light and nutrient limitation. Enhanced organic matter production was accompanied by marked deviations of the C:N:P ratio from the canonical stoichiometry of marine particulate matter of 106:16:1 (C:N:P) known as the Redfield ratio. Increased cell organic carbon content, C:N, and C:P were observed at high light when growth was either nitrogen or phosphorus limited. Elevated CO₂ led to further increases in the particulate C:N and C:P ratios. Enhanced CO₂ uptake by phytoplankton such as E. huxleyi, in response to elevated atmospheric CO₂, could increase carbon storage in the nitrogen‐limited regions of the oceans and thus act as a negative feedback on rising atmospheric CO₂ levels.     

The role of phytoplankton photosynthesis in global biogeochemical cycles

Phytoplankton biomass in the world's oceans amounts to only 1–2% of the total global plant carbon, yet these organisms fix between 30 and 50 billion metric tons of carbon annually, which is about 40% of the total. On geological time scales there is profound evidence of the importance of phytoplankton photosynthesis in biogeochemical cycles. It is generally assumed that present phytoplankton productivity is in a quasi steady-state (on the time scale of decades). However, in a global context, the stability of oceanic photosynthetic processes is dependent on the physical circulation of the upper ocean and is therefore strongly influenced by the atmosphere. The net flux of atmospheric radiation is critical to determining the depth of the upper mixed layer and the vertical fluxes of nutrients. These latter two parameters are keys to determining the intensity, and spatial and temporal distributions of phytoplankton blooms. Atmospheric radiation budgets are not in steady-state. Driven largely by anthropogenic activities in the 20th century, increased levels of IR- absorbing gases such as CO₂, CH₄ and CFC's and NO_x will potentially increase atmospheric temperatures on a global scale. The atmospheric radiation budget can affect phytoplankton photosynthesis directly and indirectly. Increased temperature differences between the continents and oceans have been implicated in higher wind stresses at the ocean margins. Increased wind speeds can lead to higher nutrient fluxes. Throughout most of the central oceans, nitrate concentrations are sub-micromolar and there is strong evidence that the quantum efficiency of Photosystem II is impaired by nutrient stress. Higher nutrient fluxes would lead to both an increase in phytoplankton biomass and higher biomass-specific rates of carbon fixation. However, in the center of the ocean gyres, increased radiative heating could reduce the vertical flux of nutrients to the euphotic zone, and hence lead to a reduction in phytoplankton carbon fixation. Increased desertification in terrestrial ecosystems can lead to increased aeolean loadings of essential micronutrients, such as iron. An increased flux of aeolean micronutrients could fertilize nutrient-replete areas of the open ocean with limiting trace elements, thereby stimulating photosynthetic rates. The factors which limit phytoplankton biomass and photosynthesis are discussed and examined with regard to potential changes in the Earth climate system which can lead the oceans away from steady-state. While it is difficult to confidently deduce changes in either phytoplankton biomass or photosynthetic rates on decadal time scales, time-series analysis of ocean transparency data suggest long-term trends have occurred in the North Pacific Ocean in the 20th century. However, calculations of net carbon uptake by the oceans resulting from phytoplankton photosynthesis suggest that without a supply of nutrients external to the ocean, carbon fixation in the open ocean is not presently a significant sink for excess atmospheric CO₂.The submitted paper has been authored under Contract No. DE-AC02-76H00016 with the US Department of Energy. Accordingly, the US Government retains a non-exclusive, royalty-free license to publish or reproduce the published form of this contribution, or allow others to do so, for US Government purposes.     

Effect of eutrophication on air–sea CO2 fluxes in the coastal Southern North Sea: a model study of the past 50 years

The RIVERSTRAHLER model, an idealized biogeochemical model of the river system, has been coupled to MIRO‐CO₂, a complex biogeochemical model describing diatom and Phaeocystis blooms and carbon and nutrient cycles in the marine domain, to assess the dual role of changing nutrient loads and increasing atmospheric CO₂ as drivers of air–sea CO₂ exchanges in the Southern North Sea with a focus on the Belgian coastal zone (BCZ). The whole area, submitted to the influence of two main rivers (Seine and Scheldt), is characterized by variable diatom and Phaeocystis colonies blooms which impact on the trophic status and air–sea CO₂ fluxes of the coastal ecosystem. For this application, the MIRO‐CO₂ model is implemented in a 0D multibox frame covering the eutrophied Eastern English Channel and Southern North Sea and receiving loads from the rivers Seine and Scheldt. Model simulations are performed for the period between 1951 and 1998 using real forcing fields for sea surface temperature, wind speed and atmospheric CO₂ and RIVERSTRAHLER simulations for river carbon and nutrient loads. Model results suggest that the BCZ shifted from a source of CO₂ before 1970 (low eutrophication) towards a sink during the 1970–1990 period when anthropogenic DIN and P loads increased, stimulating C fixation by autotrophs. In agreement, a shift from net annual heterotrophy towards autotrophy in BCZ is simulated from 1980. The period after 1990 is characterized by a progressive decrease of P loads concomitant with a decrease of primary production and of the CO₂ sink in the BCZ. At the end of the simulation period, the BCZ ecosystem is again net heterotroph and acts as a source of CO₂ to the atmosphere. R‐MIRO‐CO₂ scenarios testing the relative impact of temperature, wind speed, atmospheric CO₂ and river loads variability on the simulated air–sea CO₂ fluxes suggest that the trend in air–sea CO₂ fluxes simulated between 1951 and 1998 in the BCZ was mainly controlled by the magnitude and the ratio of inorganic nutrient river loads. Quantitative nutrient changes control the level of primary production while qualitative changes modulate the relative contribution of diatoms and Phaeocystis to this flux and hence the sequestration of atmospheric CO₂.     

Nitrogen source and pCO2 synergistically affect carbon allocation,growth and morphology of the coccolithophore Emiliania huxleyi: potential implications of ocean acidification for the carbon cycle

Coccolithophores are unicellular phytoplankton that produce calcium carbonate coccoliths as an exoskeleton. Emiliania huxleyi, the most abundant coccolithophore in the world's ocean, plays a major role in the global carbon cycle by regulating the exchange of CO₂ across the ocean‐atmosphere interface through photosynthesis and calcium carbonate precipitation. As CO₂ concentration is rising in the atmosphere, the ocean is acidifying and ammonium (NH₄⁺) concentration of future ocean water is expected to rise. The latter is attributed to increasing anthropogenic nitrogen (N) deposition, increasing rates of cyanobacterial N₂ fixation due to warmer and more stratified oceans, and decreased rates of nitrification due to ocean acidification. Thus, future global climate change will cause oceanic phytoplankton to experience changes in multiple environmental parameters including CO₂, pH, temperature and nitrogen source. This study reports on the combined effect of elevated pCO₂ and increased NH₄⁺ to nitrate (NO₃^?) ratio (NH₄⁺/NO₃^?) on E. huxleyi, maintained in continuous cultures for more than 200 generations under two pCO₂ levels and two different N sources. Herein, we show that NH₄⁺ assimilation under N‐replete conditions depresses calcification at both low and high pCO₂, alters coccolith morphology, and increases primary production. We observed that N source and pCO₂ synergistically drive growth rates, cell size, and the ratio of inorganic to organic carbon. These responses to N source suggest that, compared to increasing CO₂ alone, a greater disruption of the organic carbon pump could be expected in response to the combined effect of increased NH₄⁺/NO₃^? ratio and CO₂ level in the future acidified ocean. Additional experiments conducted under lower nutrient conditions are needed prior to extrapolating our findings to the global oceans. Nonetheless, our results emphasize the need to assess combined effects of multiple environmental parameters on phytoplankton biology to develop accurate predictions of phytoplankton responses to ocean acidification.     

The glacial oceans

The accumulation of carbonate and carbonaceous matter increased during the glacial age due to intensified sink water formation in the northernmost Atlantic Ocean and the intensified upwellings in the Indo-Pacific Oceans. As a consequence of that there was a decrease of the ΣCO₂ concentration in the ocean as well as of the CO₂ content in the atmosphere.As a whole the accumulation area of calcareous matter was enlarged by ca. 20% during glacial times. The decreased accumulation area in the North Atlantic was well compensated by a large increase in the Indo-Pacific oceans.The northernmost part of the Atlantic Ocean was quasi-stagnant during the most intense deglacial times, thus causing a reduced exchange between surface water and deep water which, in its turn, led to a decrease of the accumulation of carbon (as CaCO₃ and organic matter). The concentration of ΣCO₂ rose in the seawater and the pCO₂ increased as well.     

Coral reefs modify their seawater carbon chemistry – implications for impacts of ocean acidification

Reviews suggest that that the biogeochemical threshold for sustained coral reef growth will be reached during this century due to ocean acidification caused by increased uptake of atmospheric CO₂. Projections of ocean acidification, however, are based on air‐sea fluxes in the open ocean, and not for shallow‐water systems such as coral reefs. Like the open ocean, reef waters are subject to the chemical forcing of increasing atmospheric pCO₂. However, for reefs with long water residence times, we illustrate that benthic carbon fluxes can drive spatial variation in pH, pCO₂ and aragonite saturation state (Ω_a) that can mask the effects of ocean acidification in some downstream habitats. We use a carbon flux model for photosynthesis, respiration, calcification and dissolution coupled with Lagrangian transport to examine how key groups of calcifiers (zooxanthellate corals) and primary producers (macroalgae) on coral reefs contribute to changes in the seawater carbonate system as a function of water residence time. Analyses based on flume data showed that the carbon fluxes of corals and macroalgae drive Ω_ain opposing directions. Areas dominated by corals elevate pCO₂ and reduce Ω_a, thereby compounding ocean acidification effects in downstream habitats, whereas algal beds draw CO₂ down and elevate Ω_a, potentially offsetting ocean acidification impacts at the local scale. Simulations for two CO₂ scenarios (600 and 900 ppm CO₂) suggested that a potential shift from coral to algal abundance under ocean acidification can lead to improved conditions for calcification in downstream habitats, depending on reef size, water residence time and circulation patterns. Although the carbon fluxes of benthic reef communities cannot significantly counter changes in carbon chemistry at the scale of oceans, they provide a significant mechanism of buffering ocean acidification impacts at the scale of habitat to reef.     

Reduced water repellency of a grassland soil under elevated atmospheric CO2

Water repellency is a widespread characteristic of soils that can modify soil moisture content and distribution and is implicated in important processes such as aggregation and carbon sequestration. Repellency arises as a consequence of organic matter inputs; as elevated atmospheric CO₂ is known to modify such inputs, we tested the repellency of a grassland soil after 5 years of exposure to elevated CO₂ in a free air carbon dioxide enrichment experiment. Using a water droplet penetration time test, we found a significant reduction in repellency at elevated CO₂ in samples at field moisture content. As many of the processes potentially influenced by repellency have been shown to be modified at elevated CO₂ (e.g. soil aggregation, C sequestration, recruitment from seed), we suggest that further exploration of this phenomenon could enhance our understanding of CO₂ effects on ecosystem function. The mechanism responsible for the change in repellency has not been identified.     

Will ocean acidification affect marine microbes?

The pH of the surface ocean is changing as a result of increases in atmospheric carbon dioxide (CO₂), and there are concerns about potential impacts of lower pH and associated alterations in seawater carbonate chemistry on the biogeochemical processes in the ocean. However, it is important to place these changes within the context of pH in the present-day ocean, which is not constant; it varies systematically with season, depth and along productivity gradients. Yet this natural variability in pH has rarely been considered in assessments of the effect of ocean acidification on marine microbes. Surface pH can change as a consequence of microbial utilization and production of carbon dioxide, and to a lesser extent other microbially mediated processes such as nitrification. Useful comparisons can be made with microbes in other aquatic environments that readily accommodate very large and rapid pH change. For example, in many freshwater lakes, pH changes that are orders of magnitude greater than those projected for the twenty second century oceans can occur over periods of hours. Marine and freshwater assemblages have always experienced variable pH conditions. Therefore, an appropriate null hypothesis may be, until evidence is obtained to the contrary, that major biogeochemical processes in the oceans other than calcification will not be fundamentally different under future higher CO₂/lower pH conditions.     

Emiliania huxleyi increases calcification but not expression of calcification-related genes in long-term exposure to elevated temperature and pCO2

Increased atmospheric pCO₂ is expected to render future oceans warmer and more acidic than they are at present. Calcifying organisms such as coccolithophores that fix and export carbon into the deep sea provide feedbacks to increasing atmospheric pCO₂. Acclimation experiments suggest negative effects of warming and acidification on coccolithophore calcification, but the ability of these organisms to adapt to future environmental conditions is not well understood. Here, we tested the combined effect of pCO₂ and temperature on the coccolithophore Emiliania huxleyi over more than 700 generations. Cells increased inorganic carbon content and calcification rate under warm and acidified conditions compared with ambient conditions, whereas organic carbon content and primary production did not show any change. In contrast to findings from short-term experiments, our results suggest that long-term acclimation or adaptation could change, or even reverse, negative calcification responses in E. huxleyi and its feedback to the global carbon cycle. Genome-wide profiles of gene expression using RNA-seq revealed that genes thought to be essential for calcification are not those that are most strongly differentially expressed under long-term exposure to future ocean conditions. Rather, differentially expressed genes observed here represent new targets to study responses to ocean acidification and warming.     

造林再造林、森林采伐、气候变化、CO2浓度升高、火灾和虫害对森林固碳能力的影响

森林生态系统具有吸收大气CO_2、缓解气候变化的作用。造林再造林作为京都议定书认可的大气CO_2减排途径,是提高森林固碳能力的低成本、有效策略。森林生态系统固碳能力还受森林采伐、气候变化、大气CO_2浓度升高、火灾以及虫害等自然因素和人为因素的强烈影响。综述了全球和区域造林再造林的固碳能力,以及目前较受重视的一些因素(森林采伐、气候变化、大气CO_2浓度升高、火灾以及虫害)对森林生态系统固碳能力的影响。结果表明,全球造林再造林固碳能力为148—2400TgC/a;采伐造成的全球森林碳损失最大为900 TgC/a,其次是火灾为300 TgC/a,虫害造成森林碳释放最小在2—107 TgC/a之间。建议在今后的研究中,应关注固碳措施和多种环境因素对森林生态系统固碳能力,尤其是对森林土壤固碳能力的影响,严格控制森林采伐和火灾发生,以及减少或避免造林再造林活动引起的碳泄漏。     

Ocean acidification increases fatty acids levels of larval fish

Rising levels of anthropogenic carbon dioxide in the atmosphere are acidifying the oceans and producing diverse and important effects on marine ecosystems, including the production of fatty acids (FAs) by primary producers and their transfer through food webs. FAs, particularly essential FAs, are necessary for normal structure and function in animals and influence composition and trophic structure of marine food webs. To test the effect of ocean acidification (OA) on the FA composition of fish, we conducted a replicated experiment in which larvae of the marine fish red drum (Sciaenops ocellatus) were reared under a climate change scenario of elevated CO₂ levels (2100 µatm) and under current control levels (400 µatm). We found significantly higher whole-body levels of FAs, including nine of the 11 essential FAs, and altered relative proportions of FAs in the larvae reared under higher levels of CO₂. Consequences of this effect of OA could include alterations in performance and survival of fish larvae and transfer of FAs through food webs.     

Estimating soil carbon sequestration under elevated CO2 by combining carbon isotope labelling with soil carbon cycle modelling

Elevated CO₂ concentrations generally stimulate grassland productivity, but herbaceous plants have only a limited capacity to sequester extra carbon (C) in biomass. However, increased primary productivity under elevated CO₂ could result in increased transfer of C into soils where it could be stored for prolonged periods and exercise a negative feedback on the rise in atmospheric CO₂. Measuring soil C sequestration directly is notoriously difficult for a number of methodological reasons. Here, we present a method that combines C isotope labelling with soil C cycle modelling to partition net soil sequestration into changes in new C fixed over the experimental duration (C_new) and pre‐experimental C (C_old). This partitioning is advantageous because the C_new accumulates whereas C_old is lost in the course of time (ΔC_new>0 whereas ΔC_old<0). We applied this method to calcareous grassland exposed to 600 μL CO₂ L^?1 for 6 years. The CO₂ used for atmospheric enrichment was depleted in ¹³C relative to the background atmosphere, and this distinct isotopic signature was used to quantify net soil C_new fluxes under elevated CO₂. Using ¹³C/¹²C mass balance and inverse modelling, the Rothamsted model ‘RothC’ predicted gross soil C_new inputs under elevated CO₂ and the decomposition of C_old. The modelled soil C pools and fluxes were in good agreement with experimental data. C isotope data indicated a net sequestration of ≈90 g C_new m^?2 yr^?1 in elevated CO₂. Accounting for C_old‐losses, this figure was reduced to ≈30 g C m^?2 yr^?1 at elevated CO₂; the elevated CO₂‐effect on net C sequestration was in the range of≈10 g C m^?2 yr^?1. A sensitivity and error analysis suggests that the modelled data are relatively robust. However, elevated CO₂‐specific mechanisms may necessitate a separate parameterization at ambient and elevated CO₂; these include increased soil moisture due to reduced leaf conductance, soil disaggregation as a consequence of increased soil moisture, and priming effects. These effects could accelerate decomposition of C_old in elevated CO₂ so that the CO₂ enrichment effect may be zero or even negative. Overall, our findings suggest that the C sequestration potential of this grassland under elevated CO₂ is rather limited.     

Feedback of carbon and nitrogen cycles enhances carbon sequestration in the terrestrial biosphere

The efforts to explain the ‘missing sink’ for anthropogenic carbon dioxide (CO₂) have included in recent years the role of nitrogen as an important constraint for biospheric carbon fluxes. We used the Nitrogen Carbon Interaction Model (NCIM) to investigate patterns of carbon and nitrogen storage in different compartments of the terrestrial biosphere as a consequence of a rising atmospheric CO₂ concentration, in combination with varying levels of nitrogen availability. This model has separate but closely coupled carbon and nitrogen cycles with a focus on soil processes and soil–plant interactions, including an active compartment of soil microorganisms decomposing litter residues and competing with plants for available nitrogen. Biological nitrogen fixation is represented as a function of vegetation nitrogen demand. The model was validated against several global datasets of soil and vegetation carbon and nitrogen pools. Five model experiments were carried out for the modeling periods 1860–2002 and 2002–2100. In these experiments we varied the nitrogen availability using different combinations of biological nitrogen fixation, denitrification, leaching of soluble nitrogen compounds with constant or rising atmospheric CO₂ concentrations. Oversupply with nitrogen, in an experiment with nitrogen fixation, but no nitrogen losses, together with constant atmospheric CO₂, led to some carbon sequestration in organismic pools, which was nearly compensated by losses of C from soil organic carbon pools. Rising atmospheric CO₂ always led to carbon sequestration in the biosphere. Considering an open nitrogen cycle including dynamic nitrogen fixation, and nitrogen losses from denitrification and leaching, the carbon sequestration in the biosphere is of a magnitude comparable to current observation based estimates of the ‘missing sink.’ A fertilization feedback between the carbon and nitrogen cycles occurred in this experiment, which was much stronger than the sum of separate influences of high nitrogen supply and rising atmospheric CO₂. The demand‐driven biological nitrogen fixation was mainly responsible for this result. For the modeling period 2002–2100, NCIM predicts continued carbon sequestration in the low range of previously published estimates, combined with a plausible rate of CO₂‐driven biological nitrogen fixation and substantial redistribution of nitrogen from soil to plant pools.     

Metal limitation of cyanobacterial N2 fixation and implications for the Precambrian nitrogen cycle

Nitrogen fixation is a critical part of the global nitrogen cycle, replacing biologically available reduced nitrogen lost by denitrification. The redox‐sensitive trace metals Fe and Mo are key components of the primary nitrogenase enzyme used by cyanobacteria (and other prokaryotes) to fix atmospheric N₂ into bioessential compounds. Progressive oxygenation of the Earth's atmosphere has forced changes in the redox state of the oceans through geologic time, from anoxic Fe‐enriched waters in the Archean to partially sulfidic deep waters by the mid‐Proterozoic. This development of ocean redox chemistry during the Precambrian led to fluctuations in Fe and Mo availability that could have significantly impacted the ability of prokaryotes to fix nitrogen. It has been suggested that metal limitation of nitrogen fixation and nitrate assimilation, along with increased rates of denitrification, could have resulted in globally reduced rates of primary production and nitrogen‐starved oceans through much of the Proterozoic. To test the first part of this hypothesis, we grew N₂‐fixing cyanobacteria in cultures with metal concentrations reflecting an anoxic Archean ocean (high Fe, low Mo), a sulfidic Proterozoic ocean (low Fe, moderate Mo), and an oxic Phanerozoic ocean (low Fe, high Mo). We measured low rates of cellular N₂ fixation under [Fe] and [Mo] estimated for the Archean ocean. With decreased [Fe] and higher [Mo] representing sulfidic Proterozoic conditions, N₂ fixation, growth, and biomass C:N were similar to those observed with metal concentrations of the fully oxygenated oceans that likely developed in the Phanerozoic. Our results raise the possibility that an initial rise in atmospheric oxygen could actually have enhanced nitrogen fixation rates to near modern marine levels, providing that phosphate was available and rising O₂ levels did not markedly inhibit nitrogenase activity.     

Soil respiration and the global carbon cycle

Soil respiration is the primary path by which CO₂fixed by land plants returns to the atmosphere. Estimated at approximately 75 × 10¹⁵gC/yr, this large natural flux is likely to increase due changes in the Earth's condition. The objective of this paper is to provide a brief scientific review for policymakers who are concerned that changes in soil respiration may contribute to the rise in CO₂in Earth's atmosphere. Rising concentrations of CO₂in the atmosphere will increase the flux of CO₂from soils, while simultaneously leaving a greater store of carbon in the soil. Traditional tillage cultivation and rising temperature increase the flux of CO₂from soils without increasing the stock of soil organic matter. Increasing deposition of nitrogen from the atmosphere may lead to the sequestration of carbon in vegetation and soils. The response of the land biosphere to simultaneous changes in all of these factors is unknown, but a large increase in the soil carbon pool seems unlikely to moderate the rise in atmospheric CO₂during the next century.