Plant phenology plays an important role in nutrient cycling and carbon balance in forest ecosystems, but its response to the interaction of global warming and precipitation reduction remains unclear. In this study, an experiment with factorial soil warming (ambient, ambient +5 °C) and precipitation exclusion (ambient, ambient -50 %) was conducted in a subtropical Chinese fir (Cunninghamia lanceolata) plantation. We investigated the effects of soil warming, precipitation exclusion, and their interactions on Chinese fir phenology involving tree height and fine root growth. In the meantime, the impact of tree height growth and related climatic factors on fine root production was also assessed. The results showed that: (1) more variable phenology responses were observed in fine root growth than in tree height growth to the climatic treatments; the duration of fine root growth and tree height growth was significantly reduced by the precipitation exclusion and warming treatment, respectively; phenology differences of fine root and tree height growth caused by the solo warming and precipitation exclusion treatment were further enhanced by the combined treatment; and despite the greater inter-annual phenology stability of tree height growth than that of fine root growth, both of them showed insignificant response to all the climate treatments; (2) asynchrony of phenology between tree height and fine root growth was significantly enlarged by solo warming and precipitation exclusion treatments, and further enlarged by the combined treatment; (3) fine root production was significantly and positively correlated with air, and soil temperature, and tree height growth as well, which was altered by warming and precipitation exclusion treatments. Our results demonstrated that climatic changes significantly and differently alter phenology of, and extend the phenology asynchrony between, above and below ground plant components, and also highlight the climate-sensitive and variable nature of root phenology. Overall, these phenology responses to climatic change may weaken the close link between fine root production and tree height growth, which may result in temporal mismatch between nutrient demand and supply in Chinese fir plantation.

Physiological responses of soil microorganisms to global warming are important for soil ecosystem function and the terrestrial carbon cycle. Here, we investigate the effects of weeks, years, and decades of soil warming across seasons and time on the microbial protein biosynthesis machineries (i.e. ribosomes), the most abundant cellular macromolecular complexes, using RNA:DNA and RNA:MBC (microbial biomass carbon) ratios as proxies for cellular ribosome contents. We compared warmed soils and non-warmed controls of 15 replicated subarctic grassland and forest soil temperature gradients subject to natural geothermal warming. RNA:DNA ratios tended to be lower in the warmed soils during summer and autumn, independent of warming duration (6 weeks, 8-14 years, and > 50 years), warming intensity (+3°C, +6°C, and +9°C), and ecosystem type. With increasing temperatures, RNA:MBC ratios were also decreasing. Additionally, seasonal RNA:DNA ratios of the consecutively sampled forest showed the same temperature-driven pattern. This suggests that subarctic soil microorganisms are depleted of ribosomes under warm conditions and the lack of consistent relationships with other physicochemical parameters besides temperature further suggests temperature as key driver. Furthermore, in incubation experiments, we measured significantly higher CO2 emission rates per unit of RNA from short- and long-term warmed soils compared to non-warmed controls. In conclusion, ribosome reduction may represent a widespread microbial physiological response to warming that offers a selective advantage at higher temperatures, as energy and matter can be reallocated from ribosome synthesis to other processes including substrate uptake and turnover. This way, ribosome reduction could have a substantial effect on soil carbon dynamics.

Oligotrophic mountain lakes act as sensitive indicators of landscape-scale changes in mountain regions due to their low nutrient concentration and remote, relatively undisturbed watersheds. Recent research shows that phosphorus (P) concentrations are increasing in mountain lakes around the world, creating more mesotrophic states and altering lake ecosystem structure and function. The relative importance of atmospheric deposition and climate-driven changes to local biogeochemistry in driving these shifts is not well established. In this study, we test whether increasing temperatures in watershed soils may be contributing to the observed increases in mountain lake P loading. Specifically, we test whether higher soil temperatures increase P mobilization from mountain soils by accelerating the rate of geochemical weathering and soil organic matter decomposition. We used paired soil incubation (lab) and soil transplant (field) experiments with mountain soils from around the western United States to test the effects of warming on rain-leachable P concentration, soil P mobilization, and soil respiration. Our results show that while higher temperature can increase soil P mobilization, low soil moisture can limit the effects of warming in some situations. Soils with lower bulk densities, higher pH, lower aluminum oxide contents, and lower ratios of carbon to nitrogen had much higher rain-leachable P concentration across all sites and experimental treatments. Together, these results suggest that mountain watersheds with high-P soils and relatively high soil moisture could have the largest increases in P mobilization with warming. Consequently, lakes and streams in such watersheds could become especially susceptible to soil-driven eutrophication as temperatures rise.

Climate change predictions suggest that arctic and subarctic ecosystems will be particularly affected by rising temperatures and extreme weather events, including severe heat waves. Temperature is one of the most important environmental factors controlling and regulating microbial decomposition in soils; therefore, it is critical to understand its impact on soil microorganisms and their feedback to climate warming. We conducted a warming experiment in a subarctic birch forest in North Sweden to test the effects of summer heat waves on the thermal trait distributions that define the temperature dependences for microbial growth and respiration. We also determined the microbial temperature dependences 10 and 12 months after the heat wave simulation had ended to investigate the persistence of the thermal trait shifts. As a result of warming, the bacterial growth temperature dependence shifted to become warm-adapted, with a similar trend for fungal growth. For respiration, there was no shift in the temperature dependence. The shifts in thermal traits were not accompanied by changes in α- or β-diversity of the microbial community. Warming increased the fungal-to-bacterial growth ratio by 33% and decreased the microbial carbon use efficiency by 35%, and both these effects were caused by the reduction in moisture the warming treatments caused, while there was no evidence that substrate depletion had altered microbial processes. The warm-shifted bacterial thermal traits were partially restored within one winter but only fully recovered to match ambient conditions after 1 year. To conclude, a summer heat wave in the Subarctic resulted in (i) shifts in microbial thermal trait distributions; (ii) lower microbial process rates caused by decreased moisture, not substrate depletion; and (iii) no detectable link between the microbial thermal trait shifts and community composition changes.

Plant litter is known to affect soil, community, and ecosystem properties. However, we know little about the capacity of litter to modulate grassland responses to climate change. Using a 7-yr litter removal experiment in a semiarid grassland, here we examined how litter removal interacts with a 2-yr drought to affect soil environments, plant community composition, and ecosystem function. Litter loss exacerbates the negative impacts of drought on grasslands. Litter removal increased soil temperature but reduced soil moisture and nitrogen mineralization, which substantially increased the negative impacts of drought on primary productivity and the abundance of perennial rhizomatous graminoids. Moreover, complete litter removal shifted plant community composition from grass-dominated to forb-dominated and reduced species and functional group asynchrony, resulting in lower ecosystem temporal stability. Our results suggest that ecological processes that lead to reduction in litter, such as burning, grazing, and haying, may render ecosystems more vulnerable and impair the capacity of grasslands to withstand drought events.

Climate warming advances the onset of tree growth in spring, but above- and belowground phenology are not always synchronized. These differences in growth responses may result from differences in root and bud dormancy dynamics, but root dormancy is largely unexplored. We measured dormancy in roots and leaf buds of Fagus sylvatica and Populus nigra by quantifying the warming sum required to initiate above- and belowground growth in October, January and February. We furthermore carried out seven experiments, manipulating only the soil and not air temperature before or during tree leaf-out to evaluate the potential of warmer roots to influence budburst timing using seedlings and adult trees of F. sylvatica and seedlings of Betula pendula. Root dormancy was virtually absent in comparison with the much deeper winter bud dormancy. Roots were able to start growing immediately as soils were warmed during the winter. Interestingly, higher soil temperature advanced budburst across all experiments, with soil temperature possibly accounting for c. 44% of the effect of air temperature in advancing aboveground spring phenology per growing degree hour. Therefore, differences in root and bud dormancy dynamics, together with their interaction, likely explain the nonsynchronized above- and belowground plant growth responses to climate warming.

Soils are the largest terrestrial carbon sink and the foundation of our food, fiber, and fuel systems. Healthy soils are carbon sinks, storing more carbon than they release. This reduces the amount of carbon dioxide released into the atmosphere and buffers against climate change. Soil microbes drive biogeochemical cycling and contribute to soil health through organic matter breakdown, plant growth promotion, and nutrient distribution. In this study, we determined how soil microbial growth traits respond to long-term soil warming. We found that bacterial isolates from warmed plots showed evidence of adaptation of optimum growth temperature. This suggests that increased microbial biomass and growth in a warming world could result in greater carbon storage. As temperatures increase, greater microbial activity may help reduce the soil carbon feedback loop. Our results provide insight on how atmospheric carbon cycling and soil health may respond in a warming world.

The response of subarctic grassland\'s below-ground to soil warming is key to understanding this ecosystem\'s adaptation to future climate. Functionally different below-ground plant organs can respond differently to changes in soil temperature (Ts). We aimed to understand the below-ground adaptation mechanisms by analysing the dynamics and chemistry of fine roots and rhizomes in relation to plant community composition and soil chemistry, along with the duration and magnitude of soil warming.
We investigated the effects of the duration [medium-term warming (MTW; 11 years) and long-term warming (LTW; > 60 years)] and magnitude (0-8.4 °C) of soil warming on below-ground plant biomass (BPB), fine root biomass (FRB) and rhizome biomass (RHB) in geothermally warmed subarctic grasslands. We evaluated the changes in BPB, FRB and RHB and the corresponding carbon (C) and nitrogen (N) pools in the context of ambient, Ts < +2 °C and Ts > +2 °C scenarios.
BPB decreased exponentially in response to an increase in Ts under MTW, whereas FRB declined under both MTW and LTW. The proportion of rhizomes increased and the C-N ratio in rhizomes decreased under LTW. The C and N pools in BPB in highly warmed plots under MTW were 50 % less than in the ambient plots, whereas under LTW, C and N pools in warmed plots were similar to those in non-warmed plots. Approximately 78 % of the variation in FRB, RHB, and C and N concentration and pools in fine roots and rhizomes was explained by the duration and magnitude of soil warming, soil chemistry, plant community functional composition, and above-ground biomass. Plant\'s below-ground biomass, chemistry and pools were related to a shift in the grassland\'s plant community composition - the abundance of ferns increased and BPB decreased towards higher Ts under MTW, while the recovery of below-ground C and N pools under LTW was related to a higher plant diversity.
Our results indicate that plant community-level adaptation of below ground to soil warming occurs over long periods. We provide insight into the potential adaptation phases of subarctic grasslands.

Climate warming has been suggested to impact high latitude grasslands severely, potentially causing considerable carbon (C) losses from soil. Warming can also stimulate nitrogen (N) turnover, but it is largely unclear whether and how altered N availability impacts belowground C dynamics. Even less is known about the individual and interactive effects of warming and N availability on the fate of recently photosynthesized C in soil. On a 10-year geothermal warming gradient in Iceland, we studied the effects of soil warming and N addition on CO2 fluxes and the fate of recently photosynthesized C through CO2 flux measurements and a 13 CO2 pulse-labeling experiment. Under warming, ecosystem respiration exceeded maximum gross primary productivity, causing increased net CO2 emissions. N addition treatments revealed that, surprisingly, the plants in the warmed soil were N limited, which constrained primary productivity and decreased recently assimilated C in shoots and roots. In soil, microbes were increasingly C limited under warming and increased microbial uptake of recent C. Soil respiration was increased by warming and was fueled by increased belowground inputs and turnover of recently photosynthesized C. Our findings suggest that a decade of warming seemed to have induced a N limitation in plants and a C limitation by soil microbes. This caused a decrease in net ecosystem CO2 uptake and accelerated the respiratory release of photosynthesized C, which decreased the C sequestration potential of the grassland. Our study highlights the importance of belowground C allocation and C-N interactions in the C dynamics of subarctic ecosystems in a warmer world.

Soil dissolved organic carbon (DOC) is the most active part in forest soil carbon pool, the responses of which to climate warming has profound effects on forest carbon cycling. Based on a manipulative soil warming experiment in subtropical evergreen broad-leaved forests, we collected soil solutions in situ and used ultraviolet-visible, infrared and three-dimensional fluorescence spectroscopy analyses to explore the effects of soil warming (+4 ℃, 1 year) on soil DOC quantity and quality along the soil profile. The results showed that soil DOC flux remained constant along the soil profile. Soil DOC mainly included two humic-like fractions and one microbial metabolite. Warming significantly decreased soil DOC flux and the abundance of aromatic and hydrophobic components, and increased the amount of low molecular weight carbohydrates. Furthermore, soil warming increased the relative proportion of humic-like fractions in the surface soil layer (0-10 cm) and microbial metabolite in the deep soil layer (30-40 cm), indicating that warming might accelerate microbial turnover in the deep layer. Overall, soil warming not only decreased soil DOC content, but also simplified the composition of soil DOC in subtropical evergreen broad-leaved forests.
土壤可溶性有机碳(DOC)是森林土壤碳库中最活跃的部分,其对气候变暖的响应将深刻影响森林生态系统碳循环。本研究基于亚热带常绿阔叶林野外增温实验平台,原位收集土壤溶液,结合紫外可见光谱、红外光谱和三维荧光光谱分析,探究土壤剖面DOC通量及组成对增温(+4 ℃,1年)的响应。结果表明:土壤DOC通量在土层之间没有显著差异。土壤DOC主要由2个类腐殖酸组分和1个微生物代谢产物组分组成。与对照相比,增温显著降低了土壤DOC通量,降低了土壤溶液芳香化指数和疏水组分比例,增加了小分子碳水化合物的含量。此外,增温还提高了表层土壤(0～10 cm)类腐殖酸组分的相对贡献,增加了深层土壤(30～40 cm)微生物代谢产物组分的相对贡献,这在一定程度上表明变暖加速了深层土壤微生物周转。总体来说,土壤增温降低了亚热带常绿阔叶林土壤DOC数量,使DOC结构趋于简单。.