Climate change impacts on vegetation are mediated by soil processes that regulate rhizosphere water balance, nutrient dynamics, and ground-level temperatures. For ecosystems characterized by high fine-scale substrate heterogeneity such as grasslands on poorly developed soils, effects of climate change on plant communities may depend on substrate properties that vary at the scale of individuals (om 2 ), leading to fine-scale shifts in community structure that may go undetected at larger scales. Here, we show in a long-running climate experiment in species-rich limestone grassland in Buxton, England (UK), that the resistance of the community to 15-year manipulations of temperature and rainfall at the plot scale (9 m 2 ) belies considerable community reorganization at the microsite (100 cm 2 ) scale. In individual models of the abundance of the 25 most common species with respect to climate treatment and microsite soil depth, 13 species exhibited significant soil depth affinities, and nine of these have shifted their position along the depth gradient in response to one or more climate treatments. Estimates of species turnover across the depth gradient reviewed in relation to measurements of water potential, nitrogen supply, pH, and community biomass suggest that communities of shallow microsites are responding directly to microenvironmental changes induced by climate manipulation, while those of the deepest microsites are shifting in response to changes in competitive interference from more nutrient-demanding species. Moreover, for several species in summer drought and winter heated treatments, climate response in deep microsites was opposite that of shallow microsites, suggesting microsite variation is contributing to community stability at the whole-plot level. Our study thus demonstrates a strong link between community dynamics and substrate properties, and suggests ecosystems typified by fine-scale substrate heterogeneity may possess a natural buffering capacity in the face of climate change.

Increased summer drought will exacerbate the regeneration of many tree species at their lower latitudinal and altitudinal distribution limits. In vulnerable habitats, introduction of more drought-tolerant provenances or species is currently considered to accelerate tree species migration and facilitate forest persistence. Trade-offs between drought adaptation and growth plasticity might, however, limit the effectiveness of assisted migration, especially if introductions focus on provenances or species from different climatic regions. We tested in a common garden experiment the performance of Pinus sylvestris seedlings from the continental Central Alps under increased temperatures and extended spring and/or summer drought, and compared seedling emergence, survival and biomass allocation to that of P. sylvestris and closely related Pinus nigra from a Mediterranean seed source. Soil heating had only minor effects on seedling performance but high spring precipitation doubled the number of continental P. sylvestris seedlings present after the summer drought. At the same time, twice as many seedlings of the Mediterranean than the continental P. sylvestris provenance were present, which was due to both higher emergence and lower mortality under dry conditions. Both P. sylvestris provenances allocated similar amounts of biomass to roots when grown under low summer precipitation. Mediterranean seedlings, however, revealed lower phenotypic plasticity than continental seedlings under high precipitation, which might limit their competitive ability in continental Alpine forests in non-drought years. By contrast, high variability in the response of individual seedlings to summer drought indicates the potential of continental P. sylvestris provenances to adapt to changing environmental conditions.

The future trajectory of forest ecosystems under climate change is heavily debated. Previous studies on the impacts of climate change on forest ecosystems have focused mainly on direct effects of altered climatic conditions, whereas interactions with disturbance events have been largely neglected. The aim of this study is to explore interactions of drought with fire disturbance and to assess their effects on tree species shifts in the European Central Alps. Tree recruitment after a stand replacing wildfire in the Rhone valley, Switzerland, was measured along an altitudinal temperature moisture gradient. Recruitment was more successful in pioneer species (Betula pendula, Populus tremula and Salix appendiculata) than in pre-fire stand forming (PFSF) species (Larix decidua, Picea abies and Pinus sylvestris). Seedling and sapling density was not related to fire intensity, but it correlated with the distance to the forest edge in PFSF species. The window of opportunity for seedling establishment was short (1-2 years), and moisture deficit was the main limiting factor for tree recruitment at lower altitudes. We suggest that prolonged drought periods, as projected under continued global warming, will further aggravate tree recruitment success after fire disturbance at low altitudes of the Central Alps and may eventually lead to a shift from PFSF species to either more drought-tolerant species or to forest-free vegetation.

Soil nitrogen mineralisation (Nmin), the conversion of organic into inorganic N, is important for productivity and nutrient cycling. The balance between mineralisation and immobilisation (net Nmin) varies with soil properties and climate. However, because most global-scale assessments of net Nmin are laboratory-based, its regulation under field-conditions and implications for real-world soil functioning remain uncertain. Here, we explore the drivers of realised (field) and potential (laboratory) soil net Nmin across 30 grasslands worldwide. We find that realised Nmin is largely explained by temperature of the wettest quarter, microbial biomass, clay content and bulk density. Potential Nmin only weakly correlates with realised Nmin, but contributes to explain realised net Nmin when combined with soil and climatic variables. We provide novel insights of global realised soil net Nmin and show that potential soil net Nmin data available in the literature could be parameterised with soil and climate data to better predict realised Nmin.

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Microclimatic effects (light, temperature) are often neglected in phenological studies and little is known about the impact of resource availability (nutrient and water) on tree's phenological cycles.• Here we experimentally studied spring and autumn phenology in four temperate trees in response to changes in bud albedo (white-vs. black-painted buds), light conditions (nonshaded vs. ~70% shaded), water availability (irrigated, control and reduced precipitation) and nutrients (low vs. high availability). • We found that higher bud albedo or shade delayed budburst (up to +12 days), indicating that temperature is sensed locally within each bud. Leaf senescence was delayed by high nutrient availability (up to +7 days) and shade conditions (up to +39 days) in all species, except oak. Autumn phenological responses to summer droughts depended on species, with a delay for cherry (+ 7 days) and an advance for beech (-7 days). • The strong phenological effects of bud albedo and light exposure reveal an important role of microclimatic variation on phenology. In addition to the temperature and photoperiod effects, our results suggest a tight interplay between source and sink processes in regulating the end of the seasonal vegetation cycle, which can be largely influenced by resource availability (light, water and nutrients).

Insulin-like growth factor binding protein-3 (IGFBP-3), the product of a tumor suppressor target gene, can modulate cell proliferation and apoptosis by IGF-I-dependent and IGF-Iindependent mechanisms. IGFBP-3 controls the bioavailability of IGFs in the extracellular environment and is known to be subject to degradation by various extracellular proteases. Although nuclear localization and functions of IGFBP-3 have been described in the past, we show as the novel features of this study that the abundance of nuclear IGFBP-3 is directly regulated by ubiquitin/proteasome-dependent proteolysis. We show that IGFBP-3 degradation depends on an active ubiquitin-E1 ligase, specific 26S proteasome inhibitors can efficiently stabilize nuclear IGFBP-3, and the metabolic halflife of nuclear IGFBP-3 is strongly reduced relative to cytoplasmic IGFBP-3. Nuclear IGFBP-3 is highly polyubiquitinated at multiple lysine residues in its conserved COOHterminal domain and stabilized through mutation of two COOH-terminal lysine residues. Moreover, we show that IGFBP-3, if ectopically expressed in the nucleus, can induce apoptotic cell death. These results suggest that ubiquitin/proteasomemediated proteolysis of IGFBP-3 may contribute to downregulation of apoptosis. (Cancer Res 2006; 66(6): 3024-33)

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