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Coastal dunes are characterised by strong gradients of abiotic stress, typically increasing in severity from inland areas towards the shoreline. Thus, dune gradients represent unique opportunities to study intraspecific responses to environmental changes and to investigate which factors drive community change. This study aims to examine functional trait variation in two coexisting species in response to environmental changes along a dune gradient in NW Spain. Trait convergence was also investigated and compared between both ends of the gradient. We measured functional leaf traits related to plant efficiency in the use of light, water and nutrients, also possible stressors (salt content and pH) and availability of limiting resources (water and nutrients) in the soil. Most soil variables showed changes following a non-directional gradient. Differences in soil variables were site specific and depended on growth of the study species. Structural and functional traits depended on species and/or plant position on the gradient, except for effective quantum yield of PSII and leaf δ15N. The pattern of variation was mostly directional for reflectance indices related to leaf physiology. Multivariate analyses showed significant interspecific differences in the set of traits they exhibited along positions in the gradient. Species also differed in the combination of traits selected under given environmental conditions. Coexisting species display a specific set of traits that reflects different strategies to environmental stress. Our study highlights the overly simplistic nature of some previous studies that assume dune gradients are monotonically directional, without considering that these gradients may be differentially modified by species activity.

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Brassica oleracea cultivars include important vegetable and forage crops grown worldwide, whereas the wild counterpart occurs naturally on European sea cliffs. Domestication and selection processes have led to phenotypic and genetic divergence between domesticated plants and their wild ancestors that inhabit coastal areas and are exposed to saline conditions. Salinity is one of the most limiting factors for crop production. However, little is known about how salinity affects plants in relation to domestication of B. oleracea. The objective of this study was to determine the influence of domestication status (wild, landrace or cultivar) on the response of different B. oleracea crops to salinity, as measured by seed germination, plant growth, water content and mineral concentration parameters at the seedling stage. For this purpose, two independent pot experiments were conducted with six accessions of B. oleracea, including cabbage (group capitata) and kale (group acephala), in a growth chamber under controlled environmental conditions. In both taxonomic groups, differences in domestication status and salt stress significantly affected all major process such as germination, changes in dry matter, water relations and mineral uptake. In the acephala experiment, the domestication × salinity interaction significantly affected water content parameters and shoot Na+ allocation. At early stages of development, wild plants are more succulent than cultivated plants and have a higher capacity to maintain lower Na+ concentrations in their shoots in response to increasing levels of salinity. Different responses of domesticated and cultivated accessions in relation to these traits indicated a high level of natural variation in wild B. oleracea. Exclusion of Na+ from shoots and increasing succulence may enhance salt tolerance in B. oleracea exposed to extreme salinity in the long term. The wild germplasm can potentially be used to improve the salt tolerance of crops by the identification of useful genes and incorporation of these into salinity-sensitive cultivars.

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BACKGROUND AND AIMS: One of the most striking attributes of clonal plants is their capacity for physiological integration, which enables movement of essential resources between connected ramets. This study investigated the capacity of physiological integration to buffer differences in resource availability experienced by ramets of the clonal wild strawberry plant, Fragaria vesca. Specifically, a study was made of the responses of connected and severed offspring ramets growing in environments with different water availability conditions (well watered or water stressed) and nitrogen forms (nitrate or ammonium). METHODS: The experimental design consisted of three factors, 'integration' (connected, severed) 'water status' (well watered, water stressed) and 'nitrogen form' (nitrate, ammonium), applied in a pot experiment. The effects of physiological integration were studied by analysing photochemical efficiency, leaf spectral reflectance, photosynthesis and carbon and nitrogen isotope discrimination, the last of which has been neglected in previous studies. KEY RESULTS: Physiological integration buffered the stress caused by water deprivation. As a consequence, survival was improved in water-stressed offspring ramets that remained connected to their parent plants. The nitrogen isotope composition (δ(15)N) values in the connected water-stressed ramets were similar to those in ramets in the ammonium treatment; however, δ(15)N values in connected well-watered ramets were similar to those in the nitrate treatment. The results also demonstrated the benefit of integration for offspring ramets in terms of photochemical activity and photosynthesis. CONCLUSIONS: This is the first study in which carbon and nitrogen isotopic discrimination has been used to detect physiological integration in clonal plants. The results for nitrogen isotope composition represent the first evidence of preferential transport of a specific form of nitrogen to compensate for stressful conditions experienced by a member clone. Water consumption was lower in plants supplied with ammonium than in plants supplied with nitrate, and therefore preferential transport of ammonium from parents to water-stressed offspring could potentially optimize the water use of the whole clone.

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Honckenya peploides is a subdioecious dune plant that reproduces both sexually and by clonal growth. In northwest Spain this species was found to exhibit an extreme spatial segregation of the sexes, and our objective was to investigate genetic variation in unisexual clumps. Genetic variation was studied in six unisexual clumps of H. peploides, three of them exclusively composed of males and three exclusively female. In total, 193 samples were analysed using isozyme analysis and 80 samples were analysed using two AFLP primer combinations. Both techniques revealed considerably high genetic diversity (average proportion of distinguishable genotypes: 0.22 for isozymes and 0.36 for AFLP; average Simpson's D: 0.65 for isozymes and 0.68 for AFLP). Our results show that, in spite of clonal growth, each unisexual clump consists of different genotypes. Genetic diversity within clumps is similar for both sexual morphs. Reasons for unisexuality of the clumps are discussed.

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The gender of dimorphic plant species is often affected by ecophysiological variables. Differences have been interpreted as a response of the sexes to meet specific resource demands associated with reproduction. This study investigated whether sex-specific variations in ecophysiological traits in response to water availability determine the performance of each sex in different habitats, and therefore promote extreme spatial segregation of the sexes in the subdioecious plant, Honckenya peploides. Twenty-seven plants of each sex were individually potted in dune sand and assigned randomly to one of three water treatments. Well-watered plants were watered daily to field capacity, whereas plants in the moderate and high-water stress treatments received 40% and 20%, respectively, of the water given to well-watered plants. Photochemical efficiency, leaf spectral properties and components of relative growth rate (leaf area ratio and net assimilation rate) were measured. Photochemical efficiencies integrated over time were higher in male than in female plants. Water deficit decreased maximum quantum yield in female plants more rapidly than in male plants, but female plants (unlike male plants) had recovered to initial values by the end of the experiment. Maximum quantum yield in male plants was more affected by water stress than in female plants, indicating that male plants were more susceptible to photoinhibition. The two sexes did not differ in growth rate, but male plants invested a higher proportion of their biomass in leaves, had a higher leaf area per unit biomass and lower net assimilation rate relative to female plants. Female plants had a higher water content and succulence than male plants. Differences in stomatal density between the sexes depended on water availability. The results suggest that the two sexes of H. peploides have different strategies for coping with water stress. The study also provides evidence of sex differences in allocation traits. We conclude that between-sex differences in ecophysiological and allocation traits may contribute to explain habitat-related between-sex differences in performance and, therefore, the spatial segregation of the sexes.

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BACKGROUND AND AIMS: In clonal plants, internode connections allow translocation of photosynthates, water, nutrients and other substances among ramets. Clonal plants form large systems that are likely to experience small-scale spatial heterogeneity. Physiological and morphological responses of Fragaria vesca to small-scale heterogeneity in soil quality were investigated, together with how such heterogeneity influences the placement of ramets. As a result of their own activities plants may modify the suitability of their habitats over time. However, most experiments on habitat selection by clonal plants have not generally considered time as an important variable. In the present study, how the foraging behaviour of clonal plants may change over time was also investigated. METHODS: In a complex of environments with different heterogeneity, plant performance was determined in terms of biomass, ramet production and photosynthetic activity. To identify habitat selection, the number of ramets produced and patch where they rooted were monitored. KEY RESULTS: Parent ramets in heterogeneous environments showed significantly higher maximum and effective quantum yields of photosystem II than parents in homogeneous environments. Parents in heterogeneous environments also showed significantly higher investment in photosynthetic biomass and stolon/total biomass, produced longer stolons, and had higher mean leaf size than parents in homogeneous environments. Total biomass and number of offspring ramets were similar in both environments. However, plants in homogeneous environments showed random allocation of offspring ramets to surrounding patches, whereas plants in heterogeneous environments showed preferential allocation of offspring to higher-quality patches. CONCLUSIONS: The results suggest that F. vesca employs physiological and morphological strategies to enable efficient resource foraging in heterogeneous environments and demonstrate the benefits of physiological integration in terms of photosynthetic efficiency. The findings indicate that short-term responses cannot be directly extrapolated to the longer term principally because preferential colonization of high-quality patches means that these patches eventually show reduced quality. This highlights the importance of considering the time factor in experiments examining responses of clonal plants to heterogeneity.

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Sex ratio and sexual dimorphism in physiology and growth were studied in the dioecious tree Ilex aquifolium at two localities in northern Spain. Genet sex ratio was significantly male biased in one locality but not in the other. However, ramet and flowering ramet sex ratios were male biased at both study sites. Males had significantly thicker main trunks than females in one locality and produced more ramets in the other. Growth rate, estimated from mean width of annual rings, did not differ between localities, but males produced wider rings than females at both sites. Mean annual growth rates over the last 10, 20, and 30 yr were significantly higher for males. Measurements of chlorophyll fluorescence indicated that the efficiency of photosynthesis of leaves on nonfruiting branches of females was higher than for leaves on branches of male plants under low-light conditions, though not under saturating-light conditions. Efficiency of photosynthesis was significantly lower on fruiting branches of female plants than on nonfruiting branches. We discuss whether the observed between-sex differences are attributable to the higher cost of reproduction in females and/or to pollen competition.

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Plants in nature live in populations of variable density, a characteristic which may influence individual plant responses to the environment. We investigated how the responses of Sinapis alba plants to different wind speeds and CO2 concentrations could be modified by plant density. In our wind-density experiment the expectation that mechanical and physiological effects of wind will be ameliorated by growing in high density, as a result of positive plant interactions, was realised. Although individual plants were smaller at higher densities, the effect of increasing windspeed was much less than at lower plant densities. A similar reduced sensitivity of individual plant growth under high densities was also observed under CO2 enrichment. When measured as a population or stand response, there was no effect of density on the CO2 responses, with all stands showing very similar increases in total biomass with CO2 enrichment. In the wind speed experiment, total biomass per stand increased significantly with density, although there was no effect of density on the wind speed response. Specific leaf area decreased with increasing wind speed and this response was significantly affected by the density at which the plants grew.

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We examined how independent and interactive effects of CO2 concentrations, water supply and wind speed affect growth rates, biomass partitioning, water use efficiency, diffusive conductance and stomatal density of plants. To test the prediction that wind stress will be ameliorated by increased CO2 and/or by unrestricted water supply we grew Sinapis alba L. plants in controlled chambers under combinations of two levels of CO2 (350 ppmv, 700 ppmv), two water regimes and two wind speeds (0.3 ms-1, 3.7 ms-1). We harvested at ten different dates over a period of 60 days. A growth analysis was carried out to evaluate treatment effects on plant responses. Plants grown both in increased CO2 and in low wind conditions had significantly greater stem length, leaf area and dry weights of plant parts. Water supply significantly affected stem diameter, root weight and leaf area. CO2 enrichment significantly increased the rate of biomass accumulation and the relative ratio of biomass increase to leaf area expansion. High wind speed significantly reduced plant growth rates and the rate of leaf area expansion was reduced more than the rate of biomass accumulation. Regression analysis showed significant CO2 effects on the proportion of leaf and stem dry weight to total dry weight. A marked plant-age effect was dependent on water supply, wind speed and CO2 concentration. A reduced water supply significantly decreased the stomatal conductance, and water use efficiency significantly increased with a limited water supply, low wind and increased CO2. We found significant CO2 x wind effects for water diffusion resistance, adaxial number of stomata and water use efficiencies and significant wind x water effect for water use efficiency. In conclusion, wind stress was ameliorated by growing in unrestricted water but not by growing in increased CO2.