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To buffer against the high spatial and temporal heterogeneity of the riparian habitat, riparian tree species, such as black poplar (Populus nigra L.), may display a high level of genetic variation and phenotypic plasticity for functional traits. Using a multisite common garden experiment, we estimated the relative contribution of genetic and environmental effects on the phenotypic variation expressed for individual leaf area, leaf shape, leaf structure and leaf carbon isotope discrimination (Δ(13)C) in natural populations of black poplar. Twenty-four to 62 genotypes were sampled in nine metapopulations covering a latitudinal range from 48 °N to 42 °N in France and in Italy and grown in two common gardens at Orléans (ORL) and at Savigliano (SAV). In the two common gardens, substantial genetic variation was expressed for leaf traits within all metapopulations, but its expression was modulated by the environment, as attested by the genotype × environment (G × E) interaction variance being comparable to or even greater than genetic effects. For LA, G × E interactions were explained by both changes in genotype ranking between common gardens and increased variation in SAV, while these interactions were mainly attributed to changes in genotype ranking for Δ(13)C. The nine P. nigra metapopulations were highly differentiated for LA, as attested by the high coefficient of genetic differentiation (QST = 0.50 at ORL and 0.51 at SAV), and the pattern of metapopulation differentiation was highly conserved between the two common gardens. In contrast, they were moderately differentiated for Δ(13)C (QST = 0.24 at ORL and 0.25 at SAV) and the metapopulation clustering changed significantly between common gardens. Our results evidenced that the nine P. nigra metapopulations present substantial genetic variation and phenotypic plasticity for leaf traits, which both represent potentially significant determinants of populations' capacities to respond, on a short-term basis and over generations, to environmental variations., (© The Author 2015. Published by Oxford University Press. All rights reserved. For Permissions, please email: journals.permissions@oup.com.)

The ongoing global change could be an additional threat to the establishment and the long-term survival of Populus nigra L., an emblematic European riparian species. With the general aim of gaining insights into the adaptive potential of this species, we (i) quantified variations within and among three French P. nigra populations for key physiological attributes, i.e., water-use efficiency (assessed from bulk leaf carbon isotope discrimination, Δ(13)C), growth performance and related leaf traits, (ii) examined genotype and population by environment interactions, and (iii) explored the relationship between Δ(13)C and growth. Thirty genotypes were sampled in each of three naturally established populations and grown in two different sites, Orléans (ORL) and Guémené-Penfao (GMN). In ORL, two similar plots were established and different watering regimes were applied in order to test for the drought response. Significant variations were observed for all traits within and among populations irrespective of site and watering. Trait variation was larger within than among populations. The effect of drought was neither genotype- nor population-dependent, contrary to the effect of site. The population ranking was maintained in all sites and watering regimes for the two most complex traits: Δ(13)C and growth. Moreover, these two traits were unrelated, which indicates that (i) water-use efficiency and growth are largely uncoupled in this species, and (ii) the environmental factors driving genetic structuration for Δ(13)C and growth act independently. The large variations found within populations combined with the consistent differences among populations suggest a large adaptive potential for P. nigra.

Genetic variation of leaf proteome in drought response was investigated among eight Populus xeuramericana genotypes contrasting for their leaf carbon isotope discrimination (Delta), an estimate of intrinsic water-use efficiency. Plants were grown in open field on two similar plots. Drought was induced by an 86-day irrigation cessation on one plot, whereas a second plot remained regularly irrigated. Using 2-DE, 863 reproducible spots were detected; about 60% presented at least one significant effect i.e. treatment, genotype and/or genotype by treatment interaction effect. A significant genotype by treatment interaction was detected for 62 reliably identified proteins among which, about 65% consisted in chloroplast-associated proteins either involved in the Calvin cycle or in the electron-transport chains. The other proteins were involved in oxidative stress, amino acid or protein metabolisms. Correlations between protein abundance and Delta variations were found for 45 reliably identified proteins. The abundance of ribulose-1,5-bisphosphate carboxylase/oxygenase activase isoforms scaled negatively with Delta regardless of the treatment, suggesting that a large intrinsic water-use efficiency could be due to higher abundance of ribulose-1,5-bisphosphate carboxylase/oxygenase activase. Under control condition, abundance of enzymes involved in carbon fixation was also negatively correlated with Delta, whereas abundance of enzymes involved in photorespiration or respiration was positively correlated with Delta.

Here we tested whether some leaf traits could be used as predictors for productivity in a range of Populus deltoides x P. nigra clones. These traits were assessed in 3-yr-old rooted cuttings from 29 clones growing in an open field trial, in a five randomized complete block design, under optimal irrigation. Variables were assigned to four groups describing productivity (above-ground biomass, total leaf area), leaf growth (total number of leaves increment rate), leaf structure (area of the largest leaf, specific leaf area, carbon and nitrogen contents), and carbon isotope discrimination in the leaves (Delta). High-yielding clones displayed larger total leaf area and individual leaf area, while no correlation could be detected between productivity and either leaf structure or Delta. By contrast, Delta was negatively correlated with number of leaves increment rate and leaf N content. Our study shows that there is a potential to improve water-use efficiency in poplar without necessarily reducing the overall productivity.

To test if some leaf parameters are predictors of productivity in a range of Populus deltoides (Bartr.) Marsh. x P. nigra L. clones, we assessed leaf traits and productivity in 2-month-old rooted cuttings from 31 clones growing in 4-l pots in a greenhouse, under conditions of controlled temperature and optimal irrigation. We evaluated four groups of variables describing (1) productivity (total biomass), (2) leaf growth (total leaf number increment and total leaf area increment rate), (3) leaf structure (specific leaf area and nitrogen and carbon contents) and (4) carbon isotope discrimination (delta), which is negatively correlated with time-integrated water-use efficiency. High-yielding clones did not necessarily display high leaf growth rates, but they displayed a larger total leaf area, lower specific leaf area and lower leaf nitrogen concentration than clones with low productivity. Total leaf area was mainly controlled by maximal individual leaf area and total leaf area increment rate (r = 0.51 and 0.56, respectively). Carbon isotope discrimination did not correlate with total biomass, but it was associated with total number of leaves and total leaf area increment rate (r = 0.39 and 0.45, respectively). Therefore, leaf area and specific leaf area were better indicators of productivity than leaf growth traits. The observed independence of delta from biomass production provides opportunities for selecting poplar clones combining high productivity and high water-use efficiency.

Responses to successive drought and re-watering cycles (1-3 cycles) were compared in greenhouse-grown cuttings of Populus x canadensis (Moench) clones, 'Luisa_ Avanzo' and 'Dorskamp.' Total leaf number increment rate, duration of leaf expansion, total and individual leaf area expansion rates and stomatal conductance were recorded periodically during the experiment. Soil water content (SWC) and predawn leaf water potential (Psi(WP)) were measured four times during each drought cycle. In parallel, relative leaf water content (RWC) and specific leaf area (SLA) were estimated on leaves collected from the top to bottom of each cutting. Under well-watered conditions, 'Luisa_Avanzo' and 'Dorskamp' differed in their patterns of leaf area expansion. Although duration of leaf expansion was similar between clones, 'Luisa_ Avanzo' exhibited higher total leaf number increment rates and individual leaf area increases than 'Dorskamp.' As a result, 'Luisa_Avanzo' cuttings reached larger individual and total leaf areas than 'Dorskamp.' 'Dorskamp' leaves had lower SLA than 'Luisa_Avanzo' leaves. In response to successive drought cycles, both clones underwent decreases in total leaf number increment rates and in total leaf area expansion rates, but both whole-plant and individual leaf areas were drastically reduced only in 'Luisa_Avanzo.' 'Dorskamp' maintained a constant leaf area as a result of an increase in the duration of leaf expansion during drought and a significant stimulation of individual leaf area expansion rate and total leaf number increment rate in response to re-watering. Drought caused a greater decrease in SLA in 'Luisa_Avanzo' than in 'Dorskamp.' Expanded leaves of 'Dorskamp' were constitutively dense or thick, or both, whereas leaves of 'Luisa_Avanzo' became dense or thick, or both, only in response to drought. In both clones, re-watering caused partial recovery of SLA to control values. Our data confirm previous field and greenhouse observations that 'Dorskamp' is more tolerant to successive drought and re-watering cycles than 'Luisa_Avanzo.' We conclude that this difference is mainly attributable to the plasticity of 'Dorskamp' after re-watering, rather than to clonal differences in drought responses.