Rainfall manipulation experiments as simulated by terrestrial biosphere models: Where do we stand?

Changes in rainfall amounts and patterns have been observed and are expected to continue in the near future with potentially significant ecological and societal consequences. Modelling vegetation responses to changes in rainfall is thus crucial to project water and carbon cycles in the future. In this study, we present the results of a new model-data intercomparison project, where we tested the ability of 10 terrestrial biosphere models to reproduce the observed sensitivity of ecosystem productivity to rainfall changes at 10 sites across the globe, in nine of which, rainfall exclusion and/or irrigation experiments had been performed. The key results are as follows: (a) Inter-model variation is generally large and model agreement varies with timescales. In severely water-limited sites, models only agree on the interannual variability of evapotranspiration and to a smaller extent on gross primary productivity. In more mesic sites, model agreement for both water and carbon fluxes is typically higher on fine (daily-monthly) timescales and reduces on longer (seasonal-annual) scales. (b) Models on average overestimate the relationship between ecosystem productivity and mean rainfall amounts across sites (in space) and have a low capacity in reproducing the temporal (interannual) sensitivity of vegetation productivity to annual rainfall at a given site, even though observation uncertainty is comparable to inter-model variability. (c) Most models reproduced the sign of the observed patterns in productivity changes in rainfall manipulation experiments but had a low capacity in reproducing the observed magnitude of productivity changes. Models better reproduced the observed productivity responses due to rainfall exclusion than addition. (d) All models attribute ecosystem productivity changes to the intensity of vegetation stress and peak leaf area, whereas the impact of the change in growing season length is negligible. The relative contribution of the peak leaf area and vegetation stress intensity was highly variable among models.

Volatiles and Tannins in Pistacia lentiscus and Their Role in Browsing Behavior of Goats (Capra hircus).

Goat herding is an important tool in the ecologically sound management of Mediterranean shrublands and woodlands, although effective levels of woody biomass removal by the goats is neither guaranteed nor easy to predict. Preliminary observations indicated that one reason for this may be poor understanding of plant-herbivore interactions that operate intraspecifically at the local spatial scale. We asked, whether goats show intraspecific preferences among neighboring plants when foraging a small local population of Pistacia lentiscus, a dominant tall shrub. First, we characterized and quantified the profile of stored and emitted volatile organic compounds (VOCs) and the PEG-binding capacity of tannins (a proxy for protein binding capacity) in the foliage of P. lentiscus shrubs, sampled within an area of 0.9 ha. We then tested goat preference between pairs of these shrubs that differed in chemical composition. Almost all sampled P. lentiscus shrubs were allocated to one of two distinct VOC chemotypes: one dominated by germacrene D and limonene (designated chemotype L) and the other by germacrene D and α-pinene (chemotype P). In contrast, continuous moderate variability was found in the binding capacity of tannins in the foliage. Goats showed preference for shrubs of chemotype L over those of chemotype P, and their preference was negatively correlated with the binding capacity of tannins. Possible influences of VOCs on goat preference that may explain the observed patterns are discussed in the light of possible context-dependent interpretation of plant VOC signals by large mammalian herbivores.

Insights into the Drought and Heat Avoidance Mechanism in Summer-Dormant Mediterranean Tall Fescue.

Summer dormancy is an evolutionary response that some perennial cool-season grasses adopted as an avoidance strategy to escape summer drought and heat. It is correlated with superior survival after severe summer droughts in many perennial grass species originating from Mediterranean environments. Understanding the genetic mechanism and environmental determinants of summer dormancy is important for interpreting the evolutionary history of seasonal dormancy and for the development of genomic tools to improve the efficiency of genetic selection for this important trait. The objectives of this research are to assess morphological and biochemical attributes that seem to be specific for the characterization of summer dormancy in tall fescue, and to validate the hypothesis that genes underlying stem determinacy might be involved in the mechanism of summer dormancy. Our results suggest that vernalization is an important requirement in the onset of summer dormancy in tall fescue. Non-vernalized tall fescue plants do not exhibit summer dormancy as vernalized plants do and behave more like summer-active types. This is manifested by continuation of shoot growth and high root activity in water uptake during summer months. Therefore, summer dormancy in tall fescue should be tested only in plants that underwent vernalization and are not subjected to water deficit during summer months. Total phenolic concentration in tiller bases (antioxidants) does not seem to be related to vernalization. It is most likely an environmental response to protect meristems from oxidative stress. Sequence analysis of the TFL1 homolog CEN gene from tall fescue genotypes belonging to summer-dormant and summer-active tall fescue types showed a unique deletion of three nucleotides specific to the dormant genotypes. Higher tiller bud numbers in dormant plants that were not allowed to flower and complete the reproductive cycle, confirmed that stem determinacy is a major component in the mechanism of summer dormancy. The number of variables identified in these studies as potential players in summer dormancy in tall fescue including vernalization, TFL1/CEN, water status, and protection from oxidative stress are a further confirmation that summer dormancy is a quantitative trait controlled by several genes with varying effects and prone to genotype by environment interactions.

Seedling Emergence and Phenotypic Response of Common Bean Germplasm to Different Temperatures under Controlled Conditions and in Open Field.

Rapid and uniform seed germination and seedling emergence under diverse environmental conditions is a desirable characteristic for crops. Common bean genotypes (Phaseolus vulgaris L.) differ in their low temperature tolerance regarding growth and yield. Cultivars tolerant to low temperature during the germination and emergence stages and carriers of the grain quality standards demanded by consumers are needed for the success of the bean crop. The objectives of this study were (i) to screen the seedling emergence and the phenotypic response of bean germplasm under a range of temperatures in controlled chamber and field conditions to display stress-tolerant genotypes with good agronomic performances and yield potential, and (ii) to compare the emergence of bean seedlings under controlled environment and in open field conditions to assess the efficiency of genebanks standard germination tests for predicting the performance of the seeds in the field. Three trials were conducted with 28 dry bean genotypes in open field and in growth chamber under low, moderate, and warm temperature. Morpho-agronomic data were used to evaluate the phenotypic performance of the different genotypes. Cool temperatures resulted in a reduction of the rate of emergence in the bean genotypes, however, emergence and early growth of bean could be under different genetic control and these processes need further research to be suitably modeled. Nine groups arose from the Principal Component Analysis (PCA) representing variation in emergence time and proportion of emergence in the controlled chamber and in the open field indicating a trend to lower emergence in large and extra-large seeded genotypes. Screening of seedling emergence and phenotypic response of the bean germplasm under a range of temperatures in controlled growth chambers and under field conditions showed several genotypes, as landraces 272, 501, 593, and the cultivar Borlotto, with stress-tolerance at emergence, and high yield potential that could be valuable genetic material for breeding programs. Additionally, the potential genetic erosion in genebanks was assessed. Regarding bean commercial traits, under low temperature at sowing time seed reached larger size, and crop yield was higher compared to warmer temperatures at the sowing time. Therefore, early sowing of bean is strongly recommended.

Few multiyear precipitation-reduction experiments find a shift in the productivity-precipitation relationship.

Well-defined productivity-precipitation relationships of ecosystems are needed as benchmarks for the validation of land models used for future projections. The productivity-precipitation relationship may be studied in two ways: the spatial approach relates differences in productivity to those in precipitation among sites along a precipitation gradient (the spatial fit, with a steeper slope); the temporal approach relates interannual productivity changes to variation in precipitation within sites (the temporal fits, with flatter slopes). Precipitation-reduction experiments in natural ecosystems represent a complement to the fits, because they can reduce precipitation below the natural range and are thus well suited to study potential effects of climate drying. Here, we analyse the effects of dry treatments in eleven multiyear precipitation-manipulation experiments, focusing on changes in the temporal fit. We expected that structural changes in the dry treatments would occur in some experiments, thereby reducing the intercept of the temporal fit and displacing the productivity-precipitation relationship downward the spatial fit. The majority of experiments (72%) showed that dry treatments did not alter the temporal fit. This implies that current temporal fits are to be preferred over the spatial fit to benchmark land-model projections of productivity under future climate within the precipitation ranges covered by the experiments. Moreover, in two experiments, the intercept of the temporal fit unexpectedly increased due to mechanisms that reduced either water loss or nutrient loss. The expected decrease of the intercept was observed in only one experiment, and only when distinguishing between the late and the early phases of the experiment. This implies that we currently do not know at which precipitation-reduction level or at which experimental duration structural changes will start to alter ecosystem productivity. Our study highlights the need for experiments with multiple, including more extreme, dry treatments, to identify the precipitation boundaries within which the current temporal fits remain valid.

Ant Abundance along a Productivity Gradient: Addressing Two Conflicting Hypotheses.

The number of individuals within a population or community and their body size can be associated with changes in resource supply. While these relationships may provide a key to better understand the role of abiotic vs. biotic constraints in animal communities, little is known about the way size and abundance of organisms change along resource gradients. Here, we studied this interplay in ants, addressing two hypotheses with opposite predictions regarding variation in population densities along resource gradients- the 'productivity hypothesis' and the 'productivity-based thinning hypothesis'. The hypotheses were tested in two functional groups of ground-dwelling ants that are directly primary consumers feeding on seeds: specialized seed-eaters and generalist species. We examined variations in colony density and foraging activity (a size measurement of the forager caste) in six ant assemblages along a steep productivity gradient in a semi-arid region, where precipitation and plant biomass vary 6-fold over a distance of 250km. An increase in the density or foraging activity of ant colonies along productivity gradients is also likely to affect competitive interactions among colonies, and consequently clinal changes in competition intensity were also examined. Ant foraging activity increased with productivity for both functional groups. However, colony density revealed opposing patterns: it increased with productivity for the specialized seed-eaters, but decreased for the generalist species. Competition intensity, evaluated by spatial partitioning of species at food baits and distribution of colonies, was uncorrelated with productivity in the specialized seed-eaters, but decreased with increasing productivity in the generalists. Our results provide support for two contrasting hypotheses regarding the effect of resource availability on the abundance of colonial organisms- the 'productivity hypothesis' for specialized seed-eaters and the 'productivity-based thinning hypothesis' for generalist species. These results also stress the importance of considering the role of functional groups in studies of community structure.

Countervailing effects on pine and oak leaf litter decomposition in human-altered Mediterranean ecosystems.

Species affect the dynamics of litter decay through the intrinsic properties of their litter, but also by influencing the environmental conditions imposed by their canopy, roots, and litter layers. We examined how human-induced changes in the relative abundances of two dominant Mediterranean trees-Pinus halepensis and Quercus calliprinos-impact leaf litter decomposition. A reciprocal transplant experiment tested decomposition of pine, oak, and mixed leaf litter in oak woodland and pine forest ecosystems with different relative abundances of pine and oak. Using likelihood methods, we tested the importance and magnitude of the environmental effects of local species abundance, litter layer composition, and soil properties on litter mass loss. Oak litter decomposition was slower than pine, and had an antagonistic effect on mixed litter decay. These results differ from other reported pine-oak associations, and are probably associated with a higher content of tannins and phenols in oak compared to pine litter in our study sites. The environmental effects of the two species were opposite to their litter decomposition dynamics. An increased proportion of pine in the oak woodlands and a higher content of pine needles in the litter layer of pine forests reduced decay rates. The presence of more oak and broadleaf litter in the litter layer accelerated decomposition in pine forests. Our results highlight the importance of considering multidimensional species effects mediated by both chemical and physical properties, and imply that man-made changes in the composition and configuration of plant communities may result in complex unpredicted consequences to ecosystem biogeochemistry.

Climate change scenarios of herbaceous production along an aridity gradient: vulnerability increases with aridity.

Climate change is expected to reduce annual precipitation by 20% and increase its standard deviation by 20% in the eastern Mediterranean. We have examined how these changes may affect herbaceous aboveground net primary production (ANPP) and its inter-annual coefficient of variation (CV) in natural rangelands along a desert-Mediterranean precipitation gradient, at five sites representing arid, semi-arid, and Mediterranean-type ecosystems, respectively, all showing positive linear relationships between herbaceous ANPP and annual precipitation. Scenarios of reduced annual precipitation and increased inter-annual precipitation variability were defined by manipulating mean annual precipitation (MAP) and its standard deviation. We simulated precipitation and calculated ANPP using current ANPP-precipitation relationships. Our model predicts that reduced precipitation will strongly reduce ANPP in arid and semi-arid sites. Moreover, the effect of reduced precipitation on the CV of ANPP along the entire gradient may be modified by changes in inter-annual variability in MAP. Reduced precipitation combined with increased precipitation variability was the scenario most relevant to the wet end of the gradient, due to the increased likelihood for both dry and rainy years. In contrast, the scenario most relevant to the arid end of the gradient combined reduced precipitation with decreased precipitation variability, due to the strong effect on mean ANPP. All scenarios increased variability of ANPP along the entire gradient. However, the higher sensitivity of vegetation at arid and semi-arid sites (i.e., lower forage production) to future changes in the precipitation regime emphasizes the need to adapt grazing management in these ecosystems to secure their long-term viability as sustainable rangelands.

Testing the limits of resistance: a 19-year study of Mediterranean grassland response to grazing regimes.

A synthesis of a long-term (19 years) study assessing the effects of cattle grazing on the structure and composition of a Mediterranean grassland in north-eastern Israel is presented, with new insights into the response of the vegetation to grazing management and rainfall. We hypothesized that the plant community studied would be resistant to high grazing intensities and rainfall variability considering the combined long history of land-use and unpredictable climatic conditions where this community evolved. Treatments included manipulations of stocking densities (moderate, heavy, and very heavy) and of grazing regimes (continuous vs. seasonal), in a factorial design. The effect of interannual rainfall variation on the expression of grazing impacts on the plant community was minor. The main effects of grazing on relative cover of plant functional groups were related to early vs. late seasonal grazing. Species diversity and equitability were remarkably stable across all grazing treatments. A reduction in tall grass cover at higher stocking densities was correlated with increased cover of less palatable groups such as annual and perennial thistles, as well as shorter and prostrate groups such as short annual grasses. This long-term study shows that interannual fluctuations in plant functional group composition could be partly accounted for by grazing pressure and timing, but not by the measured rainfall variables. Grazing affected the dominance of tall annual grasses. However, the persistence of tall grasses and more palatable species over time, despite large differences in grazing pressure and timing, supports the idea that Mediterranean grasslands are highly resistant to prolonged grazing. Indeed, even under the most extreme grazing conditions applied, there were no signs of deterioration or collapse of the ecosystem. This high resistance to grazing intensity and interannual fluctuation in climatic conditions should favor the persistence of the plant community under forecasted increasing unpredictability due to climate change.

Middle-Eastern plant communities tolerate 9 years of drought in a multi-site climate manipulation experiment.

For evaluating climate change impacts on biodiversity, extensive experiments are urgently needed to complement popular non-mechanistic models which map future ecosystem properties onto their current climatic niche. Here, we experimentally test the main prediction of these models by means of a novel multi-site approach. We implement rainfall manipulations--irrigation and drought--to dryland plant communities situated along a steep climatic gradient in a global biodiversity hotspot containing many wild progenitors of crops. Despite the large extent of our study, spanning nine plant generations and many species, very few differences between treatments were observed in the vegetation response variables: biomass, species composition, species richness and density. The lack of a clear drought effect challenges studies classifying dryland ecosystems as most vulnerable to global change. We attribute this resistance to the tremendous temporal and spatial heterogeneity under which the plants have evolved, concluding that this should be accounted for when predicting future biodiversity change.

Temporal and intraclonal variation of flowering and pseudovivipary in Poa bulbosa.

BACKGROUND AND AIMS: Versatility in the reproductive development of pseudoviviparous grasses in response to growth conditions is an intriguing reproduction strategy. To better understand this strategy, this study examined variation in flowering and pseudovivipary among populations, co-occurring clones within populations, and among tillers in individual clones of Poa bulbosa, a summer-dormant geophytic grass that reproduces sexually by seed, and asexually by basal tiller bulbs and bulbils formed in proliferated panicles. METHODS: Clones were collected from 17 populations across a rainfall gradient. Patterns of reproduction were monitored for 11 years in a common garden experiment and related to interannual differences in climatic conditions. Intraclonal variation in flowering and pseudovivipary was studied in a phytotron, under daylengths marginal for flowering induction. KEY RESULTS: Clones showed large temporal variability in their reproductive behaviour. They flowered in some years but not in others, produced normal or proliferated panicles in different years, or became dormant without flowering. Proliferating clones did not show a distinct time sequence of flowering and proliferation across years. Populations differed in incidence of flowering and proliferation. The proportion of flowering clones increased with decreasing rainfall at the site of population origin, but no consistent relationship was found between flowering and precipitation in the common garden experiment across years. In contrast, flowering decreased at higher temperatures during early growth stages after bulb sprouting. Pulses of soil fertilization greatly increased the proportion of flowering clones and panicle production. High intraclonal tiller heterogeneity was observed, as shown by the divergent developmental fates of daughter plants arising from bulbs from the same parent clone and grown under similar conditions. Panicle proliferation was enhanced by non-inductive 8 h short days, while marginally inductive 12 h days promoted normal panicles. CONCLUSIONS: Interannual variation in flowering and proliferation in P. bulbosa clones was attributed to differences in the onset of the rainy season, resulting in different daylength and temperature conditions during the early stages of growth, during which induction of flowering and dormancy occurs.

An integrative analysis of the dynamics of landscape- and local-scale colonization of Mediterranean woodlands by Pinus halepensis.

Afforestation efforts have resulted in extensive plantations of either native or non-native conifers, which in many regions has led to the spread of those conifers into surrounding natural vegetation. This process of species colonization can trigger profound changes in both community dynamics and ecosystem processes. Our study disentangled the complexity of a process of colonization in a heterogeneous landscape into a simple set of rules. We analyzed the factors that control the colonization of natural woodland ecosystems by Pinus halepensis dispersing from plantations in the Mediterranean region of Israel. We developed maximum-likelihood models to explain the densities of P. halepensis colonizing natural woodlands. Our models unravel how P. halepensis colonization is controlled by factors that determine colonization pressure by dispersing seeds and by factors that control resistance to colonization of the natural ecosystems. Our models show that the combination of different seed arrival processes from local, landscape, and regional scales determine pine establishment potential, but the relative importance of each component varied according to seed source distribution. Habitat resistance, determined by abiotic and biotic conditions, was as important as propagule input in determining the density of pine colonization. Thus, despite the fact that pine propagules disperse throughout the landscape, habitat heterogeneity within the natural ecosystems generates significant variation in the actual densities of colonized pine. Our approach provides quantitative measures of how processes at different spatial scales affect the distribution and densities of colonizing species, and a basis for projection of expected distributions. Variation in colonization rates, due to landscape-scale heterogeneity in both colonization pressure and resistance to colonization, can be expected to produce a diversity of new ecosystems. This work provides a template for understanding species colonization processes, especially in light of anthropogenic impacts, and predicting future transformation of natural ecosystems by species invasion.

Landscape-scale density-dependent recruitment of oaks in planted forests: more is not always better.

Plant colonization studies usually address density-dependent processes in the narrow sense of recruitment constraints due to negative density-dependent seed and seedling mortality. However, complex density-dependent effects may be involved in additional stages of the recruitment process. We hypothesized that seed arrival and seedling establishment are influenced by density dependence acting at small scales at the site of colonization, and at larger scales as a function of the colonizing species' landscape abundance. These hypotheses were tested in a study of colonization of pine forests by oaks in a heterogeneous Mediterranean landscape. Maximum-likelihood models show that density effects switch from positive to negative along the range of landscape-scale oak seed source abundance. Contrary to expectations, high seed source densities limited oak recruitment, suggesting a landscape-scale Janzen-Connell effect. We propose a range of mechanisms that generate positive or negative density dependence during colonization, resulting in nonlinear density-dependent feedbacks that can generate unexpected colonization patterns.

Ecotypic variation of summer dormancy relaxation associated with rainfall gradient in the geophytic grass Poa bulbosa.

BACKGROUND AND AIMS: Summer dormancy is an adaptive trait in geophytes inhabiting regions with a Mediterranean climate, allowing their survival through the hot and dry summers. Summer dormancy in Poa bulbosa is induced by increasing day-length and temperature and decreasing water availability during spring. Populations from arid habitats became dormant earlier than those from mesic habitats. Relaxation of dormancy was promoted by the hot, dry summer conditions. Here we test the hypothesis that dormancy relaxation is also delayed in ecotypes of P. bulbosa inhabiting arid regions, as a cautious strategy related to the greater unpredictability of autumn rains associated with decreasing precipitation. METHODS: Ecotypes collected across a precipitation gradient (100-1200 mm year(-1)) in the Mediterranean climate region were grown under similar conditions in a net-house in Israel. Differences among ecotypes in dormancy induction and dormancy relaxation were determined by measuring time to dormancy onset in spring, and time to sprouting after the first effective rain in autumn. Seasonal and ecotype variation in dormancy relaxation were assessed by measuring time to sprouting initiation, rate of sprouting and maximal sprouting of resting dry bulbs sampled in the net-house during late spring, and mid- and late summer, and planted in a wet substrate at temperatures promoting (10 degrees C) or limiting (20 degrees C) sprouting. KEY RESULTS: Earlier dormancy in the spring and delayed sprouting in autumn were correlated with decreasing mean annual rainfall at the site of ecotype origin. Seasonal and ecotype differences in dormancy relaxation were expressed in bulbs planted at 20 degrees C. During the summer, time to sprouting decreased while rate of sprouting and maximal sprouting increased, indicating dormancy relaxation. Ecotypes from more arid sites across the rainfall gradient showed delayed onset of sprouting and lower maximal sprouting, but did not differ in rate of sprouting. Planting at 10 degrees C promoted sprouting and cancelled differences among ecotypes in dormancy relaxation. CONCLUSIONS: Both the induction and the relaxation of summer dormancy in P. bulbosa are correlated with mean annual precipitation at the site of population origin. Ecotypes from arid habitats have earlier dormancy induction and delayed dormancy relaxation, compared with those from mesic habitats.

Leaf traits capture the effects of land use changes and climate on litter decomposability of grasslands across Europe.

Land use and climate changes induce shifts in plant functional diversity and community structure, thereby modifying ecosystem processes. This is particularly true for litter decomposition, an essential process in the biogeochemical cycles of carbon and nutrients. In this study, we asked whether changes in functional traits of living leaves in response to changes in land use and climate were related to rates of litter potential decomposition, hereafter denoted litter decomposability, across a range of 10 contrasting sites. To disentangle the different control factors on litter decomposition, we conducted a microcosm experiment to determine the decomposability under standard conditions of litters collected in herbaceous communities from Europe and Israel. We tested how environmental factors (disturbance and climate) affected functional traits of living leaves and how these traits then modified litter quality and subsequent litter decomposability. Litter decomposability appeared proximately linked to initial litter quality, with particularly clear negative correlations with lignin-dependent indices (litter lignin concentr tion, lignin:nitrogen ratio, and fiber component). Litter quality was directly related to community-weighted mean traits. Lignin-dependent indices of litter quality were positively correlated with community-weighted mean leaf dry matter content (LDMC), and negatively correlated with community-weighted mean leaf nitrogen concentration (LNC). Consequently, litter decomposability was correlated negatively with community-weighted mean LDMC, and positively with community-weighted mean LNC. Environmental factors (disturbance and climate) influenced community-weighted mean traits. Plant communities experiencing less frequent or less intense disturbance exhibited higher community-weighted mean LDMC, and therefore higher litter lignin content and slower litter decomposability. LDMC therefore appears as a powerful marker of both changes in land use and of the pace of nutrient cycling across 10 contrasting sites.

Regulation of summer dormancy by water deficit and ABA in Poa bulbosa ecotypes.

BACKGROUND AND AIMS: Survival of many herbaceous species in Mediterranean habitats during the dry, hot summer depends on the induction of summer dormancy by changes in environmental conditions during the transition between the winter (growth) season to the summer (resting) season, i.e. longer days, increasing temperature and drought. In Poa bulbosa, a perennial geophytic grass, summer dormancy is induced by long days, and the induction is enhanced by high temperature. Here the induction of summer dormancy in a Mediterranean perennial grass by water deficit under non-inductive photoperiodic conditions is reported for the first time. METHODS: Plants grown under 22/16 degrees C and non-inductive short-day (9 h, SD) were subjected to water deficit (WD), applied as cycles of reduced irrigation, or sprayed with ABA solutions. They were compared with plants in which dormancy was induced by transfer from SD to inductive long-day (16 h, LD). Responses of two contrasting ecotypes, from arid and mesic habitats were compared. Dormancy relaxation in bulbs from these ecotypes and treatments was studied by comparing sprouting capacity in a wet substrate at 10 degrees C of freshly harvested bulbs to that of dry-stored bulbs at 40 degrees C. Endogenous ABA in the bulbs was determined by monoclonal immunoassay analysis. KEY RESULTS: Dormancy was induced by WD and by ABA application in plants growing under non-inductive SD. Dormancy induction by WD was associated with increased levels of ABA. Bulbs were initially deeply dormant and their sprouting capacity was very low, as in plants in which dormancy was induced by LD. Dormancy was released after 2 months dry storage at 40 degrees C in all treatments. ABA levels were not affected by dormancy relaxation. CONCLUSIONS: Summer dormancy in P. bulbosa can be induced by two alternative and probably additive pathways: (1) photoperiodic induction by long-days, and (2) water deficit. Increased levels of endogenous ABA are involved in both pathways.

Assessing the effects of land-use change on plant traits, communities and ecosystem functioning in grasslands: a standardized methodology and lessons from an application to 11 European sites.

BACKGROUND AND AIMS: A standardized methodology to assess the impacts of land-use changes on vegetation and ecosystem functioning is presented. It assumes that species traits are central to these impacts, and is designed to be applicable in different historical, climatic contexts and local settings. Preliminary results are presented to show its applicability. METHODS: Eleven sites, representative of various types of land-use changes occurring in marginal agro-ecosystems across Europe and Israel, were selected. Climatic data were obtained at the site level; soil data, disturbance and nutrition indices were described at the plot level within sites. Sixteen traits describing plant stature, leaf characteristics and reproductive phase were recorded on the most abundant species of each treatment. These data were combined with species abundance to calculate trait values weighed by the abundance of species in the communities. The ecosystem properties selected were components of above-ground net primary productivity and decomposition of litter. KEY RESULTS: The wide variety of land-use systems that characterize marginal landscapes across Europe was reflected by the different disturbance indices, and were also reflected in soil and/or nutrient availability gradients. The trait toolkit allowed us to describe adequately the functional response of vegetation to land-use changes, but we suggest that some traits (vegetative plant height, stem dry matter content) should be omitted in studies involving mainly herbaceous species. Using the example of the relationship between leaf dry matter content and above-ground dead material, we demonstrate how the data collected may be used to analyse direct effects of climate and land use on ecosystem properties vs. indirect effects via changes in plant traits. CONCLUSIONS: This work shows the applicability of a set of protocols that can be widely applied to assess the impacts of global change drivers on species, communities and ecosystems.

Opposite effects of daylength and temperature on flowering and summer dormancy of Poa bulbosa.

BACKGROUND AND AIMS: The timing of flowering and summer dormancy induction plays a central role in the adaptation of Mediterranean geophytes to changes in the length of the growth season along rainfall gradients. Our aim was to analyse the role of the variation in the responses of flowering and summer dormancy to vernalization, daylength and growth temperature for the adaptation of Poa bulbosa, a perennial geophytic grass, to increasing aridity. METHODS: Flowering and dormancy were studied under controlled daylengths [9 h short day (SD) vs. 16 h long day (LD)] and temperatures (16/10, 22/16 and 28/22 degrees C day/night) in four ecotypes originating in arid, semi-arid and mesic habitats (110, 276 and 810 mm rain year(-1), respectively) and differing in flowering capacity under natural conditions: arid-flowering, semi-arid-flowering, semi-arid-non-flowering and mesic-non-flowering. KEY RESULTS: Flowering and dormancy were affected in opposite ways by daylength and growth temperature. Flowering occurred almost exclusively under SD. In contrast, plants became dormant much earlier under LD than under SD. In both daylengths, high temperature and pre-chilling (6 weeks at 5 degrees C) enhanced dormancy imposition, but inhibited or postponed flowering, respectively. Induction of flowering and dormancy in the different ecotypes showed differential responsiveness to daylength and temperature. Arid and semi-arid ecotypes had a higher proportion of flowering plants and flowering tillers as well as more panicles per plant than mesic ecotypes. 'Flowering' ecotypes entered dormancy earlier than 'non-flowering' ecotypes, while the more arid the site of ecotype origin, the earlier the ecotype entered dormancy. CONCLUSIONS: Variation in the flowering capacity of ecotypes differing in drought tolerance was interpreted as the result of balanced opposite effects of daylength and temperature on the flowering and dormancy processes.

Variation in onset of summer dormancy and flowering capacity along an aridity gradient in Poa bulbosa L., a geophytic perennial grass.

Variation in the onset of summer dormancy and flowering capacity of 16 populations of Poa bulbosa, collected along a steep north-south aridity gradient in Israel (810-110 mm rain year(-1)), was studied under controlled conditions in a phytotron (16 h daylength, 22/16 degrees C day/night) and under natural conditions in a garden experiment in a net-house. Plant age at the onset of dormancy varied markedly amongst populations (7-16 weeks under controlled conditions) and was positively correlated with mean annual precipitation at the site of origin of the population, i.e. dormancy was earlier as aridity increased. Flowering capacity in the different populations was negatively correlated with rainfall in the original habitat and, consequently, also with the age at onset of dormancy, i.e. the lower the mean annual precipitation, the earlier the onset of dormancy and the higher the proportion of flowering plants and panicles per plant. Differences in xeromorphic leaf traits were also observed among populations from locations differing in aridity. Plants from the more arid sites (110-310 mm year(-1)) generally had greyish and curved leaves, whereas plants from more humid sites (500-810 mm year(-1)) tended to have green and straight leaves. Thus, plants with curved and/or greyish leaves generally had a higher flowering capacity and entered dormancy earlier than plants with straight and/or green leaves. The significance of the association among these traits for the adaptation of P. bulbosa to increasing aridity is discussed.