Climate change due to greenhouse gas emissions is predicted to raise the mean global temperature by 1.0-3.5°C in the next 50-100 years. The direct and indirect effects of this potential increase in temperature on terrestrial ecosystems and ecosystem processes are likely to be complex and highly varied in time and space. The Global Change and Terrestrial Ecosystems core project of the International Geosphere-Biosphere Programme has recently launched a Network of Ecosystem Warming Studies, the goals of which are to integrate and foster research on ecosystem-level effects of rising temperature. In this paper, we use meta-analysis to synthesize data on the response of soil respiration, net N mineralization, and aboveground plant productivity to experimental ecosystem warming at 32 research sites representing four broadly defined biomes, including high (latitude or altitude) tundra, low tundra, grassland, and forest. Warming methods included electrical heat-resistance ground cables, greenhouses, vented and unvented field chambers, overhead infrared lamps, and passive night-time warming. Although results from individual sites showed considerable variation in response to warming, results from the meta-analysis showed that, across all sites and years, 2-9 years of experimental warming in the range 0.3-6.0°C significantly increased soil respiration rates by 20% (with a 95% confidence interval of 18-22%), net N mineralization rates by 46% (with a 95% confidence interval of 30-64%), and plant productivity by 19% (with a 95% confidence interval of 15-23%). The response of soil respiration to warming was generally larger in forested ecosystems compared to low tundra and grassland ecosystems, and the response of plant productivity was generally larger in low tundra ecosystems than in forest and grassland ecosystems. With the exception of aboveground plant productivity, which showed a greater positive response to warming in colder ecosystems, the magnitude of the response of these three processes to experimental warming was not generally significantly related to the geographic, climatic, or environmental variables evaluated in this analysis. This underscores the need to understand the relative importance of specific factors (such as temperature, moisture, site quality, vegetation type, successional status, land-use history, etc.) at different spatial and temporal scales, and suggests that we should be cautious in "scaling up" responses from the plot and site level to the landscape and biome level. Overall, ecosystem-warming experiments are shown to provide valuable insights on the response of terrestrial ecosystems to elevated temperature.
Stimulation of terrestrial productivity by rising CO~2~ concentration is projected to reduce the airborne fraction of anthropogenic CO~2~ emissions; coupled climate-carbon (C) cycle models, including those used in the IPCC Fourth Assessment Report (AR4), are sensitive to this negative feedback on atmospheric CO~2~^1^. The representation of the so-called CO~2~ fertilization effect in the 11 models used in AR4 and subsequent models^2,3^ was broadly consistent with experimental evidence from four free-air CO~2~ enrichment (FACE) experiments, which indicated that net primary productivity (NPP) of forests was increased by 23 +/- 2% in response to atmospheric CO~2~ enrichment to 550 ppm^4^. Substantial uncertainty remains, however, because of the expectation that feedbacks through the nitrogen (N) cycle will reduce the CO~2~ stimulation of NPP^5,6^; these feedbacks were not included in the AR4 models and heretofore have not been confirmed by experiments in forests^7^. Here, we provide new evidence from a FACE experiment in a deciduous Liquidambar styraciflua (sweetgum) forest stand in Tennessee, USA, that N limitation has significantly reduced the stimulation of NPP by elevated atmospheric CO~2~ concentration (eCO~2~). Isotopic evidence and N budget analysis support the premise that N availability in this forest ecosystem has been declining over time, and declining faster in eCO~2~. Model analyses and evidence from leaf- and stand-level observations provide mechanistic evidence that declining N availability constrained the tree response to eCO2. These results provide a strong rationale and process understanding for incorporating N limitation and N feedback effects in ecosystem and global models used in climate change assessments.
Direct quantification of terrestrial biosphere responses to global change is crucial for projections of future climate change in Earth system models. Here, we synthesized ecosystem carbon-cycling data from 1,119 experiments performed over the past four decades concerning changes in temperature, precipitation, CO 2 and nitrogen across major terrestrial vegetation types of the world. Most experiments manipulated single rather than multiple global change drivers in temperate ecosystems of the USA, Europe and China. The magnitudes of warming and elevated CO 2 treatments were consistent with the ranges of future projections, whereas those of precipitation changes and nitrogen inputs often exceeded the projected ranges. Increases in global change drivers consistently accelerated, but decreased precipitation slowed down carbon-cycle processes. Nonlinear (including synergistic and antagonistic) effects among global change drivers were rare. Belowground carbon allocation responded negatively to increased precipitation and nitrogen addition and positively to decreased precipitation and elevated CO 2. The sensitivities of carbon variables to multiple global change drivers depended on the background climate and ecosystem condition, suggesting that Earth system models should be evaluated using site-specific conditions for best uses of this large dataset. Together, this synthesis underscores an urgent need to explore the interactions among multiple global change drivers in underrepresented regions such as semi-arid ecosystems, forests in the tropics and subtropics, and Arctic tundra when forecasting future terrestrial carbon-climate feedback.
Increasing temperatures in northern high latitudes are causing permafrost to thaw 1 , making large amounts of previously frozen organic matter vulnerable to microbial decomposition 2 . Permafrost thaw also creates a fragmented landscape of drier and wetter soil conditions 3,4 that determine the amount and form (carbon dioxide (CO 2 ), or methane (CH 4 )) of carbon (C) released to the atmosphere. The rate and form of C release control the magnitude of the permafrost C feedback, so their relative contribution with a warming climate remains unclear 5,6 . We quantified the e ect of increasing temperature and changes from aerobic to anaerobic soil conditions using 25 soil incubation studies from the permafrost zone. Here we show, using two separate meta-analyses, that a 10 • C increase in incubation temperature increased C release by a factor of 2.0 (95% confidence interval (CI), 1.8 to 2.2). Under aerobic incubation conditions, soils released 3.4 (95% CI, 2.2 to 5.2) times more C than under anaerobic conditions. Even when accounting for the higher heat trapping capacity of CH 4 , soils released 2.3 (95% CI, 1.5 to 3.4) times more C under aerobic conditions. These results imply that permafrost ecosystems thawing under aerobic conditions and releasing CO 2 will strengthen the permafrost C feedback more than waterlogged systems releasing CO 2 and CH 4 for a given amount of C.High-latitude ecosystems store almost twice as much C in soils than what is contained in the atmosphere 7,8 . As the global climate warms, northern high latitudes are experiencing rapid increases in temperature 9 that have the potential to not only increase C emissions from previously frozen C in permafrost and the active layer 10 but also to indirectly affect the C cycle through changes in regional and local hydrology. Warmer temperatures increase thawing of icerich permafrost and the melting of ground ice, which causes the land surface to collapse into the space that was previously filled by ice resulting in thermokarst terrain 11 . Permafrost thawing can also gradually increase active layer thickness (seasonally thawed ground), causing poorly drained soil conditions in lowlands or drier conditions in uplands where natural drainage can increase 3 . On the other hand, permafrost thaw and collapse can cause soils to become waterlogged where anaerobic conditions prevail and C is released in the form of CO 2 and CH 4 . One major uncertainty in determining the climate forcing impact of permafrost ecosystems is understanding the relative magnitudes of the effects of shifting subsurface hydrology versus increasing temperatures on greenhouse gas release in permafrost ecosystems.In addition to soil temperature and moisture, the chemical composition (for example, carbon to nitrogen ratio) 12 , physical protection by soil minerals, microbial community dynamics, and other environmental controls, such as pH and nutrient availability, also impact the amount of C released to the atmosphere 13 . While temperature and soil moisture (that is, oxygen availability) a...
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