Sensitivity of a dynamic global vegetation model to climate and atmospheric CO2

The equilibrium carbon storage capacity of the terrestrial biosphere has been investigated by running the Lund–Potsdam–Jena Dynamic Global Vegetation Model to equilibrium for a range of CO₂ concentrations and idealized climate states. Local climate is defined by the combination of an observation-based climatology and perturbation patterns derived from a 4 × CO₂ warming simulations, which are linearly scaled to global mean temperature deviations, Δ T _glob. Global carbon storage remains close to its optimum for Δ T _glob in the range of ±3°C in simulations with constant atmospheric CO₂. The magnitude of the carbon loss to the atmosphere per unit change in global average surface temperature shows a pronounced nonlinear threshold behavior. About twice as much carbon is lost per degree warming for Δ T _glob above 3°C than for present climate. Tropical, temperate, and boreal trees spread poleward with global warming. Vegetation dynamics govern the distribution of soil carbon storage and turnover in the climate space. For cold climate conditions, the global average decomposition rate of litter and soil decreases with warming, despite local increases in turnover rates. This result is not compatible with the assumption, commonly made in global box models, that soil turnover increases exponentially with global average surface temperature, over a wide temperature range.     

Soil microbial responses to increased concentrations of atmospheric CO2

Terrestrial ecosystems respond to an increased concentration of atmospheric CO₂. While elevated atmospheric CO₂ has been shown to alter plant growth and productivity, it also affects ecosystem structure and function by changing below-ground processes. Knowledge of how soil microbiota respond to elevated atmospheric CO₂ is of paramount importance for understanding global carbon and nutrient cycling and for predicting changes at the ecosystem-level. An increase in the atmospheric CO₂ concentration not only alters the weight, length, and architecture of plant roots, but also affects the biotic and abiotic environment of the root system. Since the concentration of CO₂ in soil is already 10–50 times higher than that in the atmosphere, it is unlikely that increasing atmospheric CO₂ will directly influence the rhizosphere. Rather, it is more likely that elevated atmospheric CO₂ will affect the microbe–soil–plant root system indirectly by increasing root growth and rhizodeposition rates, and decreasing soil water deficit. Consequently, the increased amounts and altered composition of rhizosphere-released materials will have the potential to alter both population and community structure, and activity of soil- and rhizosphere-associated microorganisms. This occurrence could in turn affect plant health and productivity and plant community structure. This review covers current knowledge about the response of soil microbes to elevated concentrations of atmospheric CO₂.     

Arbuscular mycorrhizae respond to elevated atmospheric CO2 after long-term exposure: evidence from a CO2 spring in New Zealand supports the resource balance model

The resource balance model predicts that under elevated atmospheric CO₂, plants should preferentially allocate photosynthate to acquiring below-ground resources. Only short-term experiments are available to test this hypothesis, while long-term responses are really of interest in global change ecology. Arbuscular mycorrhizae represent one mode of below-ground nutrient acquisition available to the vast majority of plants. Percent root colonization by arbuscular mycorrhizal fungi (AMF), AMF soil hyphal length, and soil concentrations of the AMF protein glomalin increased linearly along a CO₂ gradient provided in a grassland by a CO₂ spring in Northland, New Zealand. These results are an important confirmation of numerous short-term studies, and present the first test of the resource balance model, applied to AMF, after long-term elevated CO₂ exposure.     

陆地生态系统地下碳输入与输出过程研究进展

生态系统地下碳输入与输出过程是陆地生态系统碳分配和转化的核心,并直接影响着全球碳循环。陆地生态系统凋落物、根系周转、根系分泌物、土壤有机碳、土壤微生物和土壤呼吸是地下碳输入与输出过程中的重要组成部分。由于这些组分非常复杂且其研究技术和方法受到限制,目前人们对陆地生态系统地下碳输入与输出过程尚缺乏全面的认识,故在陆地生态系统碳循环研究中存在诸多的不确定性。该文概述了凋落物、根系周转、根系分泌物、土壤有机碳、土壤微生物和土壤呼吸的研究方法,以及它们对气候变化的响应,探讨了陆地生态系统地下碳输入与输出过程中的研究难点,并对未来需要深入探究的一些领域进行了展望。     

Mycorrhizal fungi have a potential role in soil carbon storage under elevated CO2 and nitrogen deposition

In this review, we discuss the potential for mycorrhizal fungi to act as a source or sink for carbon (C) under elevated CO₂ and nitrogen deposition. Mycorrhizal tissue has been estimated to comprise a significant fraction of soil organic matter and below-ground biomass in a range of systems. The current body of literature indicates that in many systems exposed to elevated CO₂, mycorrhizal fungi might sequester increased amounts of C in living, dead and residual hyphal biomass in the soil. Through this process, the fungi might serve as a negative feedback on the rise in atmospheric CO₂ levels caused by fossil fuel burning and deforestation. By contrast, a few preliminary studies suggest that N deposition might increase turnover rates of fungal tissue and negate CO₂ effects on hyphal biomass. If these latter responses are consistent among ecosystems, C storage in hyphae might decline in habitats surrounding agricultural and urban areas. When N additions occur without CO₂ enrichment, effects on mycorrhizal growth are inconsistent. We note that analyses of hyphal decomposition under elevated CO₂ and N additions are extremely sparse but are critical in our understanding of the impact of global change on the cycling of mycorrhizal C. Finally, shifts in the community composition of arbuscular and ectomycorrhizal fungi with increasing CO₂ or N availability are frequently documented. Since mycorrhizal groups vary in growth rate and tissue quality, these changes in species assemblages could produce unforeseeable impacts on the productivity, survivorship, or decomposition of mycorrhizal biomass.     

Elevated CO2 and moisture effects on soil carbon storage and cycling in temperate grasslands

In grassland ecosystems, most of the carbon (C) occurs below-ground. Understanding changes in soil fluxes induced by elevated atmospheric CO₂ is critical for balancing the global C budget and for managing grassland ecosystems sustainably. In this review, we use the results of short-term (1–2 years) studies of below-ground processes in grassland communities under elevated CO₂ to assess future prospects for longer-term increases in soil C storage.
Results are broadly consistent with those from other plant communities and include: increases in below-ground net primary productivity and an increase in soil C cycling rate, changes in soil faunal community, and generally no increase in soil C storage. Based on other experimental data, future C storage could be favoured in soils of moderate nutrient status, moderate-to-high clay content, and low (or moderateIy high) soil moisture status. Some support for these suggestions is provided by preliminary results from direct measurements of soil C concentrations near a New Zealand natural CO₂-venting spring, and by simulations of future changes in grassland soils under the combined effects of CO₂ fertilization and regional climate change.
Early detection of any increase in soil C storage appears unlikely in complex grassland communities because of (a) the difficulty of separating an elevated CO₂ effect from the effects of soil factors including moisture status, (b) the high spatial variability of soil C and (c) the effects of global warming. Several research imperatives are identified for reducing the uncertainties in the effects of elevated atmospheric CO₂ on soil C.     

Latitudinal differentiated water table control of carbon dioxide, methane and nitrous oxide fluxes from hydromorphic soils: feedbacks to climate change

The possibility of carbon (C) being locked away from the atmosphere for millennia is given in hydromorphic soils. However, the water-table-dependent feedback from soil organic matter (SOM) decomposition to the climate system is less clear. At least three greenhouse gases are produced: carbon dioxide (CO₂), methane (CH₄) and nitrous oxide (N₂O). These gases show emission peaks at different water table positions and have different global warming potentials (GWP), for example a factor of 23 for CH₄ and 296 for N₂O as compared with the equivalent mass of CO₂ on a 100-year time horizon. This review of available annual data on all three gases revealed that the radiative forcing effect of SOM decomposition is principally dictated by CO₂ despite its low GWP. Anaerobic SOM decomposition generally has a lower potential feedback to the climatic system than aerobic SOM decomposition. Concrete values are constrained by a lack of data from tropical and subarctic regions. Furthermore, data on N₂O and on plant effects are generally rare. However, there is a clear latitudinal differentiation for the GWP of soils under anaerobic conditions compared with aerobic conditions when looking at CO₂ and CH₄: in the tropical and temperate regions, the anaerobic GWP showed a range of 25–60% of the aerobic value, but values varied between 80% and 110% in the boreal zone. Hence, particularly in the vulnerable boreal zone, the feedback from ecosystems to climate change will highly depend on plant responses to changing water tables at elevated temperatures.     

Interactions between increasing CO2 concentration and temperature on plant growth

The global environment is changing with increasing temperature and atmospheric carbon dioxide concentration, [CO₂]. Because these two factors are concomitant, and the global [CO₂] rise will affect all biomes across the full global range of temperatures, it is essential to review the theory and observations on effects of temperature and [CO₂] interactions on plant carbon balance, growth, development, biomass accumulation and yield. Although there are sound theoretical reasons for expecting a larger stimulation of net CO₂ assimilation rates by increased [CO₂] at higher temperatures, this does not necessarily mean that the pattern of biomass and yield responses to increasing [CO₂] and temperature is determined by this response. This paper reviews the interactions between the effects of [CO₂] and temperature on plants. There is little unequivocal evidence for large differences in response to [CO₂] at different temperatures, as studies are confounded by the different responses of species adapted and acclimated to different temperatures, and the interspecific differences in growth form and development pattern. We conclude by stressing the importance of initiation and expansion of meristems and organs and the balance between assimilate supply and sink activity in determining the growth response to increasing [CO₂] and temperature.     

Global patterns of root turnover for terrestrial ecosystems

Root turnover is a critical component of ecosystem nutrient dynamics and carbon sequestration and is also an important sink for plant primary productivity. We tested global controls on root turnover across climatic gradients and for plant functional groups by using a database of 190 published studies. Root turnover rates increased exponentially with mean annual temperature for fine roots of grasslands ( r ² = 0.48) and forests ( r ² = 0.17) and for total root biomass in shrublands ( r ² = 0.55). On the basis of the best-fit exponential model, the Q ₁₀ for root turnover was 1.4 for forest small diameter roots (5 mm or less), 1.6 for grassland fine roots, and 1.9 for shrublands. Surprisingly, after accounting for temperature, there was no such global relationship between precipitation and root turnover. The slowest average turnover rates were observed for entire tree root systems (10% annually), followed by 34% for shrubland total roots, 53% for grassland fine roots, 55% for wetland fine roots, and 56% for forest fine roots. Root turnover decreased from tropical to high-latitude systems for all plant functional groups. To test whether global relationships can be used to predict interannual variability in root turnover, we evaluated 14 yr of published root turnover data from a shortgrass steppe site in northeastern Colorado, USA. At this site there was no correlation between interannual variability in mean annual temperature and root turnover. Rather, turnover was positively correlated with the ratio of growing season precipitation and maximum monthly temperature ( r ² = 0.61). We conclude that there are global patterns in rates of root turnover between plant groups and across climatic gradients but that these patterns cannot always be used for the successful prediction of the relationship of root turnover to climate change at a particular site.     

Turnover of Grassland Roots in Mountain Ecosystems Revealed by Their Radiocarbon Signature: Role of Temperature and Management

Root turnover is an important carbon flux component in grassland ecosystems because it replenishes substantial parts of carbon lost from soil via heterotrophic respiration and leaching. Among the various methods to estimate root turnover, the root’s radiocarbon signature has rarely been applied to grassland soils previously, although the value of this approach is known from studies in forest soils. In this paper, we utilize the root’s radiocarbon signatures, at 25 plots, in mountain grasslands of the montane to alpine zone of Europe. We place the results in context of a global data base on root turnover and discuss driving factors. Root turnover rates were similar to those of a subsample of the global data, comprising a similar temperature range, but measured with different approaches, indicating that the radiocarbon method gives reliable, plausible and comparable results. Root turnover rates (0.06–1.0 y^-1) scaled significantly and exponentially with mean annual temperatures. Root turnover rates indicated no trend with soil depth. The temperature sensitivity was significantly higher in mountain grassland, compared to the global data set, suggesting additional factors influencing root turnover. Information on management intensity from the 25 plots reveals that root turnover may be accelerated under intensive and moderate management compared to low intensity or semi-natural conditions. Because management intensity, in the studied ecosystems, co-varied with temperature, estimates on root turnover, based on mean annual temperature alone, may be biased. A greater recognition of management as a driver for root dynamics is warranted when effects of climatic change on belowground carbon dynamics are studied in mountain grasslands.     

The impact of elevated CO2 and global climate change on arbuscular mycorrhizas: a mycocentric approach

Arbuscular mycorrhizal (AM) symbioses are a potentially important link in the chain of response of ecosystems to elevated atmospheric [CO₂]. By promoting plant phosphorus uptake and acting as a sink for plant carbon, they can alleviate photosynthetic down-regulation. Because hyphal turnover is likely to be fast, especially in warmer soils, they can also act as a rapid pathway for the return of carbon to the atmosphere. However, most experiments on AM responses to [CO₂] have failed to take into account the difference in growth of mycorrhizal and non- mycorrhizal plants; those that have done so suggest that AM colonization of roots is little altered by [CO₂], although this issue remains to be resolved. Very little is known about the effects of other factors of global environmental change on mycorrhizas. These issues need urgent attention. It is also necessary to understand the potential for the various AM fungal taxa to respond differentially to environmental changes, including carbon supply and soil temperature and moisture, especially because of the differential abilities of plant and fungal species to migrate in response to changing environments. Indeed, there is a need for a new approach to the study of mycorrhizal associations, which has been too plant-centred. It is essential to regard the fungus as an organism itself, and to understand its biology both as an entity and as part of a symbiosis.     

Elevated CO2 and conifer roots: effects on growth, life span and turnover

Elevated CO₂ increases root growth and fine (diam. 2 mm) root growth across a range of species and experimental conditions. However, there is no clear evidence that elevated CO₂ changes the proportion of C allocated to root biomass, measured as either the root:shoot ratio or the fine root:needle ratio. Elevated CO₂ tends to increase mycorrhizal infection, colonization and the amount of extramatrical hyphae, supporting their key role in aiding the plant to more intensively exploit soil resources, providing a route for increased C sequestration. Only two studies have determined the effects of elevated CO₂ on conifer fine-root life span, and there is no clear trend. Elevated CO₂ increases the absolute fine-root turnover rates; however, the standing crop root biomass is also greater, and the effect of elevated CO₂ on relative turnover rates (turnover:biomass) ranges from an increase to a decrease. At the ecosystem level these changes could lead to increased C storage in roots. Increased fine-root production coupled with increased absolute turnover rates could also lead to increases in soil organic C as greater amounts of fine roots die and decompose. Although CO₂ can stimulate fine-root growth, it is not known if this stimulation persists over time. Modeling studies suggest that a doubling of the atmospheric CO₂ concentration initially increases biomass, but this stimulation declines with the response to elevated CO₂ because increases in assimilation are not matched by increases in nutrient supply.     

Evolutionary responses of stomatal density to global CO2 change

Stomatal density is known to respond to CO₂ levels during leaf development. Current interest in the increasing concentration of atmospheric CO₂ has stimulated much experimentation on the responses of plants to relatively short-term exposure in artificially high CO₂ levels. Attempts to extrapolate from short-term to long-term responses raise fundamental questions concerning evolutionary change in response to rising global CO₂ levels. We consider the improved water use efficiency observed under elevated CO₂ levels to be the main driving force of natural selection affecting the genotypic component controlling stomatal density. Whether a response is merely phenotypic or becomes incorporated into the genotype depends on two factors: (i) the time scale of exposure and (ii) the generation time of a species. Measurements of stomatal density on fossil leaves of Salix herbacea through a glacial cycle covering the last 140000 years have shown a decrease in stomatal density in response to the rising CO₂ levels of this period. This accords with the shorter-term observations on leaves of trees seen in herbarium specimens where the stomatal density has decreased in response to the rising CO₂ levels of the last 200 years. The results indicate that natural selection over the 140000-year period may have favoured a similar response to that shown by trees phenotypically over the last 200 years. Since there is now some evidence for the genetic control of stomatal density, the role of natural selection affecting it must be considered when translating responses from short-term experiments to predict how stomatal density will be affected by long-term climatic and atmospheric change.     

Species-specific responses to atmospheric carbon dioxide and tropospheric ozone mediate changes in soil carbon

We repeatedly sampled the surface mineral soil (0–20 cm depth) in three northern temperate forest communities over an 11-year experimental fumigation to understand the effects of elevated carbon dioxide (CO₂) and/or elevated phyto-toxic ozone (O₃) on soil carbon (C). After 11 years, there was no significant main effect of CO₂ or O₃ on soil C. However, within the community containing only aspen ( Populus tremuloides Michx.), elevated CO₂ caused a significant decrease in soil C content. Together with the observations of increased litter inputs, this result strongly suggests accelerated decomposition under elevated CO_2. In addition, an initial reduction in the formation of new (fumigation-derived) soil C by O₃ under elevated CO₂ proved to be only a temporary effect, mirroring trends in fine root biomass. Our results contradict predictions of increased soil C under elevated CO₂ and decreased soil C under elevated O₃ and should be considered in models simulating the effects of Earth's altered atmosphere.     

Impact of elevated carbon dioxide on the rhizosphere communities of Carex arenaria and Festuca rubra

The increase in atmospheric carbon dioxide (CO₂) levels is predicted to stimulate plant carbon (C) fixation, potentially influencing the size, structure and function of micro- and mesofaunal communities inhabiting the rhizosphere. To assess the effects of increased atmospheric CO₂ on bacterial, fungal and nematode communities in the rhizosphere, Carex arenaria (a nonmycorrhizal plant species) and Festuca rubra (a mycorrhizal plant species) were grown in three dune soils under controlled soil temperature and moisture conditions, while subjecting the aboveground compartment to defined atmospheric conditions differing in CO₂ concentrations (350 and 700 μL L⁻¹). Real-time polymerase chain reaction (PCR) and PCR-denaturing gradient gel electrophoresis methods were used to examine effects on the size and structure of rhizosphere communities. Multivariate analysis of community profiles showed that bacteria were most affected by elevated CO₂, and fungi and nematodes to a lesser extent. The influence of elevated CO₂ was plant dependent, with the mycorrhizal plant ( F. rubra ) exerting a greater influence on bacterial and fungal communities. Biomarker data indicated that arbuscular mycorrhizal fungi (AMF) may play an important role in the observed soil community responses. Effects of elevated CO₂ were also soil dependent, with greater influence observed in the more organic-rich soils, which also supported higher levels of AMF colonization. These results indicate that responses of soil-borne communities to elevated CO₂ are different for bacteria, fungi and nematodes and dependent on the plant type and soil nutrient availability.     

Soil CO2 efflux in a boreal pine forest under atmospheric CO2 enrichment and air warming

The response of forest soil CO₂ efflux to the elevation of two climatic factors, the atmospheric concentration of CO₂ (↑CO₂ of 700 μmol mol⁻¹) and air temperature (↑ T with average annual increase of 5°C), and their combination (↑CO₂+↑ T ) was investigated in a 4-year, full-factorial field experiment consisting of closed chambers built around 20-year-old Scots pines ( Pinus sylvestris L.) in the boreal zone of Finland. Mean soil CO₂ efflux in May–October increased with elevated CO₂ by 23–37%, with elevated temperature by 27–43%, and with the combined treatment by 35–59%. Temperature elevation was a significant factor in the combined 4-year efflux data, whereas the effect of elevated CO₂ was not as evident. Elevated temperature had the most pronounced impact early and late in the season, while the influence of elevated CO₂ alone was especially notable late in the season. Needle area was found to be a significant predictor of soil CO₂ efflux, particularly in August, a month of high root growth, thus supporting the assumption of a close link between whole-tree physiology and soil CO₂ emissions. The decrease in the temperature sensitivity of soil CO₂ efflux observed in the elevated temperature treatments in the second year nevertheless suggests the existence of soil response mechanisms that may be independent of the assimilating component of the forest ecosystem. In conclusion, elevated atmospheric CO₂ and air temperature consistently increased forest soil CO₂ efflux over the 4-year period, their combined effect being additive, with no apparent interaction.     

Experimentally increased soil temperature causes release of nitrogen at a boreal forest catchment in southern Norway

Boreal forest ecosystems are sensitive to global warming, caused by increasing emissions of CO₂ and other greenhouse gases. Assessment of the biological response to future climate change is based mainly on large-scale models. Whole-ecosystem experiments provide one of the few available tools by which ecosystem response can be measured and with which global models can be evaluated. Boreal ecosystem response to global change may be manifest by alterations in nitrogen (N) dynamics, as N is often the growth limiting nutrient. The CLIMEX (Climate Change Experiment) project entails catchment-scale manipulations of CO₂ (to 560 ppmv) and temperature (by + 3 to + 5 °C) to whole forest ecosystems in southern Norway. Soil temperature is increased at 400-m² EGIL catchment by means of electric cables placed on the soil surface. Soil warming at EGIL catchment caused an increase in nitrate and ammonium concentrations in runoff in the first year of treatment. We hypothesize that higher temperature increased N release by mineralization. Whether these responses are only transient will be shown by additional years' treatment.     

Net primary and ecosystem production and carbon stocks of terrestrial ecosystems and their responses to climate change

Evaluating the role of terrestrial ecosystems in the global carbon cycle requires a detailed understanding of carbon exchange between vegetation, soil, and the atmosphere. Global climatic change may modify the net carbon balance of terrestrial ecosystems, causing feedbacks on atmospheric CO₂ and climate. We describe a model for investigating terrestrial carbon exchange and its response to climatic variation based on the processes of plant photosynthesis, carbon allocation, litter production, and soil organic carbon decomposition. The model is used to produce geographical patterns of net primary production (NPP), carbon stocks in vegetation and soils, and the seasonal variations in net ecosystem production (NEP) under both contemporary and future climates. For contemporary climate, the estimated global NPP is 57.0 Gt C y^–1, carbon stocks in vegetation and soils are 640 Gt C and 1358 Gt C, respectively, and NEP varies from –0.5 Gt C in October to 1.6 Gt C in July. For a doubled atmospheric CO₂ concentration and the corresponding climate, we predict that global NPP will rise to 69.6 Gt C y^–1, carbon stocks in vegetation and soils will increase by, respectively, 133 Gt C and 160 Gt C, and the seasonal amplitude of NEP will increase by 76%. A doubling of atmospheric CO₂ without climate change may enhance NPP by 25% and result in a substantial increase in carbon stocks in vegetation and soils. Climate change without CO₂ elevation will reduce the global NPP and soil carbon stocks, but leads to an increase in vegetation carbon because of a forest extension and NPP enhancement in the north. By combining the effects of CO₂ doubling, climate change, and the consequent redistribution of vegetation, we predict a strong enhancement in NPP and carbon stocks of terrestrial ecosystems. This study simulates the possible variation in the carbon exchange at equilibrium state. We anticipate to investigate the dynamic responses in the carbon exchange to atmospheric CO₂ elevation and climate change in the past and future.     

森林生态系统细根周转规律及影响因素

根系周转是陆地生态系统碳循环的关键过程, 对研究土壤碳库变化及全球气候变化均具有重要意义。然而由于根系周转率的测量计算方法较多, 不同方法得出的结果差异较大, 且目前对全球区域尺度上森林生态系统根系周转的研究还不够充分, 使得全球森林生态系统根系周转变化规律仍不清楚。该研究通过收集文献数据并统一周转率计算方法, 对全球5种森林类型的细根周转空间格局进行整合, 同时结合土壤理化性质和气候数据, 得出影响森林生态系统细根周转的因子。结果表明, 不同森林类型细根周转率存在显著差异, 且随着纬度的升高逐渐降低; 森林生态系统细根周转率与年平均温度和年平均降水量呈正相关; 森林生态系统细根周转率与土壤有机碳含量呈正相关但与土壤pH值呈负相关。该研究为揭示森林生态系统细根周转规律及机制提供了科学依据。     

Water availability controls microbial temperature responses in frozen soil CO2 production

Soil processes in high-latitude regions during winter are important contributors to global carbon circulation, but our understanding of the mechanisms controlling these processes is poor and observed temperature response coefficients of CO₂ production in frozen soils deviate markedly from thermodynamically predicted responses (sometimes by several orders of magnitude). We investigated the temperature response of CO₂ production in 23 unfrozen and frozen surface soil samples from various types of boreal forests and peatland ecosystems and also measured changes in water content in them after freezing. We demonstrate that deviations in temperature responses at subzero temperatures primarily emanates from water deficiency caused by freezing of the soil water, and that the amount of unfrozen water is mainly determined by the quality of the soil organic matter, which is linked to the vegetation cover. Factoring out the contribution of water limitation to the CO₂ temperature responses yields response coefficients that agree well with expectations based on thermodynamic theory concerning biochemical temperature responses. This partitioning between a pure temperature response and the effect of water availability on the response of soil CO₂ production at low temperatures is crucial for a thorough understanding of low-temperature soil processes and for accurate predictions of C-balances in northern terrestrial ecosystems.