Permafrost soils in boreal and Arctic ecosystems store almost twice as much carbon as is currently present in the atmosphere. Permafrost thaw and the microbial decomposition of previously frozen organic carbon is considered one of the most likely positive climate feedbacks from terrestrial ecosystems to the atmosphere in a warmer world. The rate of carbon release from permafrost soils is highly uncertain, but it is crucial for predicting the strength and timing of this carbon-cycle feedback effect, and thus how important permafrost thaw will be for climate change this century and beyond. Sustained transfers of carbon to the atmosphere that could cause a significant positive feedback to climate change must come from old carbon, which forms the bulk of the permafrost carbon pool that accumulated over thousands of years. Here we measure net ecosystem carbon exchange and the radiocarbon age of ecosystem respiration in a tundra landscape undergoing permafrost thaw to determine the influence of old carbon loss on ecosystem carbon balance. We find that areas that thawed over the past 15 years had 40 per cent more annual losses of old carbon than minimally thawed areas, but had overall net ecosystem carbon uptake as increased plant growth offset these losses. In contrast, areas that thawed decades earlier lost even more old carbon, a 78 per cent increase over minimally thawed areas; this old carbon loss contributed to overall net ecosystem carbon release despite increased plant growth. Our data document significant losses of soil carbon with permafrost thaw that, over decadal timescales, overwhelms increased plant carbon uptake at rates that could make permafrost a large biospheric carbon source in a warmer world.

The carbon (C) storage capacity of northern latitude ecosystems may diminish as warming air temperatures increase permafrost thaw and stimulate decomposition of previously frozen soil organic C. However, warming may also enhance plant growth so that photosynthetic carbon dioxide (CO 2 ) uptake may, in part, offset respiratory losses. To determine the effects of air and soil warming on CO 2 exchange in tundra, we established an ecosystem warming experiment -the Carbon in Permafrost Experimental Heating Research (CiPEHR) project -in the northern foothills of the Alaska Range in Interior Alaska. We used snow fences coupled with spring snow removal to increase deep soil temperatures and thaw depth (winter warming) and open-top chambers to increase growing season air temperatures (summer warming). Winter warming increased soil temperature (integrated 5-40 cm depth) by 1.5 1C, which resulted in a 10% increase in growing season thaw depth. Surprisingly, the additional 2 kg of thawed soil C m À2 in the winter warming plots did not result in significant changes in cumulative growing season respiration, which may have been inhibited by soil saturation at the base of the active layer. In contrast to the limited effects on growing-season C dynamics, winter warming caused drastic changes in winter respiration and altered the annual C balance of this ecosystem by doubling the net loss of CO 2 to the atmosphere. While most changes to the abiotic environment at CiPEHR were driven by winter warming, summer warming effects on plant and soil processes resulted in 20% increases in both gross primary productivity and growing season ecosystem respiration and significantly altered the age and sources of CO 2 respired from this ecosystem. These results demonstrate the vulnerability of organic C stored in near surface permafrost to increasing temperatures and the strong potential for warming tundra to serve as a positive feedback to global climate change.

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As permafrost degrades, the amount of organic soil carbon (C) that thaws during the growing season will increase, but decomposition may be limited by saturated soil conditions common in high-latitude ecosystems. However, in some areas, soil drying is expected to accompany permafrost thaw as a result of increased water drainage, which may enhance C release to the atmosphere. We examined the effects of ecosystem warming, permafrost thaw, and soil moisture changes on C balance in an upland tundra ecosystem. This study was conducted at a water table drawdown experiment, established in 2011 and located within the Carbon in Permafrost Experimental Heating Research project, an ecosystem warming and permafrost thawing experiment in Alaska. Warming and drying increased cumulative growing season ecosystem respiration by~20% over 3 years of this experiment. Warming caused an almost twofold increase in decomposition of a common substrate in surface soil (0-10 cm) across all years, and drying caused a twofold increase in decomposition (0-20 cm) relative to control after 3 years of drying. Decomposition of older C increased in the dried and in the combined warmed + dried plots based on soil pore space 14 CO 2 . Although upland tundra systems have been considered CH 4 sinks, warming and ground thaw significantly increased CH 4 emission rates. Water table depth was positively correlated with monthly respiration and negatively correlated with CH 4 emission rates. These results demonstrate that warming and drying may increase loss of old permafrost C from tundra ecosystems, but the form and magnitude of C released to the atmosphere will be driven by changes in soil moisture.

Abstract. A large pool of organic carbon (C) has been accumulating in the Arctic for thousands of years because cold and waterlogged conditions have protected soil organic material from microbial decomposition. As the climate warms this vast and frozen C pool is at risk of being thawed, decomposed, and released to the atmosphere as greenhouse gasses. At the same time, some C losses may be offset by warming-mediated increases in plant productivity. Plant and microbial responses to warming ultimately determine net C exchange from ecosystems, but the timing and magnitude of these responses remain uncertain. Here we show that experimental warming and permafrost (ground that remains below 08C for two or more consecutive years) degradation led to a two-fold increase in net ecosystem C uptake during the growing season. However, warming also enhanced winter respiration, which entirely offset growing-season C gains. Winter C losses may be even higher in response to actual climate warming than to our experimental manipulations, and, in that scenario, could be expected to more than double overall net C losses from tundra to the atmosphere. Our results highlight the importance of winter processes in determining whether tundra acts as a C source or sink, and demonstrate the potential magnitude of C release from the permafrost zone that might be expected in a warmer climate.