Integrated drought responses of black poplar: how important is phenotypic plasticity?

Climate change is expected to increase drought frequency and intensity which will threaten plant growth and survival. In such fluctuating environments, perennial plants respond with hydraulic and biomass adjustments, resulting in either tolerant or avoidant strategies. Plants' response to stress relies on their phenotypic plasticity. The goal of this study was to explore physiology of young Populus nigra in the context of a time‐limited and progressive water deficit in regard to their growth and stress response strategies. Fourteen French 1‐year‐old black poplar genotypes, geographically contrasted, were subjected to withholding water during 8 days until severe water stress. Water fluxes (i.e. leaf water potentials and stomatal conductance) were analyzed together with growth (i.e. radial and longitudinal branch growth, leaf senescence and leaf production). Phenotypic plasticity was calculated for each trait and response strategies to drought were deciphered for each genotype. Black poplar genotypes permanently were dealing with a continuum of adjusted water fluxes and growth between two extreme strategies, tolerance and avoidance. Branch growth, leaf number and leaf hydraulic potential traits had contrasted plasticities, allowing genotype characterization. The most tolerant genotype to water deficit, which maintained growth, had the lowest global phenotypic plasticity. Conversely, the most sensitive and avoidant genotype ceased growth until the season's end, had the highest plasticity level. All the remaining black poplar genotypes were close to avoidance with average levels of traits plasticity. These results underpinned the role of plasticity in black poplar response to drought and calls for its wider use into research on plants' responses to stress.     

Phenotypic plasticity and genetic differentiation of quantitative traits in genotypes of Leymus chinensis北大核心CSCD

Aims Adaptation mechanisms of plants to environment can be classified as genetic differentiation and phenotypic plasticity (environmental modification). The strategy and mechanism of plant adaptation is a hot topic in the field of evolutionary ecology. Leymus chinensis is one of constructive species in the Nei Mongol grassland. Particularly, Leymus chinensis is a rhizomatous and clonally reproductive grass, a genotype that can play an important role in the community. In this study, we aimed to (1) investigate the phenotypic plasticity of L. chinensis under different conditions, and (2) test the genetic differentiation and reaction norms (the relationship between the environment and the phenotype of an individual or a group of individuals) under four environmental conditions among different genotypes of L. chinensis. Methods Ten genotypes of L. chinensis were randomly selected. Under the control condition, we studied the effects of genotype, defoliation, drought and their interactions on 11 quantitative traits of growth (8 traits including photochemical efficiency of photosystem II, maximum net photosynthetic rate, transpiration rate, specific leaf area, relative growth rate, the number of tillers increased, aboveground and underground biomass growth), defense (total phenol concentration of leaf) and tolerance (non-structural carbohydrate content of root, root/shoot ratio) of L. chinensis. We studied the phenotypic plasticity, genetic differentiation and reaction norms mainly through tested the effect of environment and genotype on these traits. Important findings First, all 11 traits showed obvious phenotypic plasticity (i.e., significant effect of drought, defoliation and their interactions). The expression of 10 genotypes of L. chinensis was divergent under different environmental conditions. Interactions of genotype and environment significantly affected the maximum net photosynthetic rate, transpiration rate, specific leaf area, relative growth rate, total phenolic concentration of leaf, and total non-structural carbohydrate content of root. This indicated that the phenotypic plasticity of these five traits exhibited genetic differentiation. Second, the increase of number of tillers, belowground biomass and non-structural carbohydrate content of root did not show genetic differentiation under the same condition. The other eight traits showed significantly genetic differentiation, and the heritabilities (H2) of six traits related to growth were higher than 0.5. The leaf total phenol concentration and root/shoot ratio showed genetically differentiation only under the drought and defoliation condition, with the heritabilities being 0.145 and 0.201, respectively. These results explained why L. chinensis widely distributed in the Nei Mongol grassland, and provided genetic and environmental basis for related application and species conservation in this grassland ecosystem.     

Phenotypic plasticity and genetic differentiation of quantitative traits in genotypes of Leymus chinensis

Aims Adaptation mechanisms of plants to environment can be classified as genetic differentiation and phenotypic plasticity (environmental modification). The strategy and mechanism of plant adaptation is a hot topic in the field of evolutionary ecology. Leymus chinensis is one of constructive species in the Nei Mongol grassland. Particularly, Leymus chinensis is a rhizomatous and clonally reproductive grass, a genotype that can play an important role in the community. In this study, we aimed to (1) investigate the phenotypic plasticity of L. chinensis under different conditions, and (2) test the genetic differentiation and reaction norms (the relationship between the environment and the phenotype of an individual or a group of individuals) under four environmental conditions among different genotypes of L. chinensis. Methods Ten genotypes of L. chinensis were randomly selected. Under the control condition, we studied the effects of genotype, defoliation, drought and their interactions on 11 quantitative traits of growth (8 traits including photochemical efficiency of photosystem II, maximum net photosynthetic rate, transpiration rate, specific leaf area, relative growth rate, the number of tillers increased, aboveground and underground biomass growth), defense (total phenol concentration of leaf) and tolerance (non-structural carbohydrate content of root, root/shoot ratio) of L. chinensis. We studied the phenotypic plasticity, genetic differentiation and reaction norms mainly through tested the effect of environment and genotype on these traits. Important findings First, all 11 traits showed obvious phenotypic plasticity (i.e., significant effect of drought, defoliation and their interactions). The expression of 10 genotypes of L. chinensis was divergent under different environmental conditions. Interactions of genotype and environment significantly affected the maximum net photosynthetic rate, transpiration rate, specific leaf area, relative growth rate, total phenolic concentration of leaf, and total non-structural carbohydrate content of root. This indicated that the phenotypic plasticity of these five traits exhibited genetic differentiation. Second, the increase of number of tillers, belowground biomass and non-structural carbohydrate content of root did not show genetic differentiation under the same condition. The other eight traits showed significantly genetic differentiation, and the heritabilities (H²) of six traits related to growth were higher than 0.5. The leaf total phenol concentration and root/shoot ratio showed genetically differentiation only under the drought and defoliation condition, with the heritabilities being 0.145 and 0.201, respectively. These results explained why L. chinensis widely distributed in the Nei Mongol grassland, and provided genetic and environmental basis for related application and species conservation in this grassland ecosystem.     

Capturing candidate drought tolerance traits in two native Andean potato clones by transcription profiling of field grown plants under water stress.

Candidate traits for drought tolerance were targeted by analyzing water stress responses in two moderately drought-tolerant native Andean potato clones, SA2563 and Sullu (Solanum tuberosum L. subsp, andigena (Juz, Bukasov) Hawkes) under field conditions. SA2563 exhibited increased root growth under drought, while Sullu retained a higher relative leaf water content. Gene expression profiling using the TIGR 10 K microarray revealed 1713 significantly differentially expressed genes, 186 of these genes were up-regulated in both clones. In addition to these commonly up-regulated genes, each clone induced a specific gene set in response to drought. Gene expression and metabolite analysis pinpointed candidate traits for drought tolerance present either in one or both of the clones under investigation. These traits included osmotic adjustment, changes in carbohydrate metabolism, membrane modifications, strengthening of cuticle and cell rescue mechanisms, such as detoxification of oxygen radicals and protein stabilization. Many of the up-regulated genes have been identified previously in laboratory studies on model plants using shock treatments, and the present study confirms the importance of these factors under field conditions.     

Trait means predict performance under water limitation better than plasticity for seedlings of Poaceae species on the eastern Tibetan Plateau

Water availability may be altered by changes in precipitation under global climate change in alpine areas. Trait means and plasticity are important for plants in response to a changing environment. In an examination of alpine plant responses to changed water availability, and for determination of how trait means and plasticity predict the performance (e.g., biomass) of these species, seeds of ten Poaceae species from the eastern Tibetan Plateau were sown and grown in a manipulated environment during a growing season in which rainfall was removed and other climate conditions remained unchanged. Growth and leaf traits of these species were measured. We found significant effects of moderate water stress on the seedling biomass of these species; however, the responses of these species to changed water condition were strongly dependent on species identity. For example, the biomass of some species significantly decreased under moderate drought, whereas that of others were either significantly increased or unaffected. This pattern was also observed for growth and leaf traits. Overall, the alpine Poaceae species showed low plasticity of traits in response to water availability relative to reports from other areas . Notably, the results show that trait means were better correlated with the productivity than with the plasticity of traits; thus, we argue that the trait means were better predictors of performance than plasticity for alpine Poaceae species. Poaceae species in alpine areas are important for forage production and for water catchment health worldwide, and these species may face water shortage because of current and future climate change. Understanding the response of alpine Poaceae species to water availability would facilitate our ability to predict the impacts of climate change on the alpine vegetation.     

Geographic variation and plasticity to water and nutrients in Pelargonium australe

Here, patterns of phenotypic plasticity and trait integration of leaf characteristics in six geographically discrete populations of the perennial herb Pelargonium australe were compared. It was hypothesized that populations would show local adaptation in trait means, but similar patterns of plasticity and trait integration. Further, it was questioned whether phenotypic plasticity was positively correlated with environmental heterogeneity and whether plasticity for water-use traits in particular was adaptive. Seedlings were grown in a glasshouse at six combinations of water and nutrient availability. Leaf anatomical, morphological and gas exchange traits were measured. High amounts of plasticity in leaf traits were found in response to changes in growth conditions and there was evidence of local adaptation among the populations. While there were significant correlations between plasticity and environmental heterogeneity, not all were positive. Notably, patterns of plasticity and trait integration varied significantly among populations. Despite that variation, some of the observed plasticity was adaptive: fitness was correlated with conservative water use when water was limiting. Pelargonium arrived in Australia approximately 5 million yr ago. It is concluded here that high amounts of plasticity, in some cases adaptive, and weak integration among traits may be key to the spread and success of this species.     

Influence of plant genotype and environment on oviposition preference and offspring survival in a gallmaking herbivore

Summary Plant resistance to insect herbivores may derive from traits influencing herbivore preference, traits influencing the suitability of the plant as a host, or both. However, the plant traits influencing host-plant selection by ovipositing insect herbivores may not completely overlap those traits that affect larval survival, and distinct traits may exhibit different levels of genetic vs. environmental control. Therefore, resource supply to the host plant could affect oviposition preference and larval performance differently in different plant genotypes. To test this hypothesis, the effects of resistance level, plant genotype, and resource supply to the host plant on oviposition preference and larval performance of a gallmaking herbivore, and on various plant traits that could influence these, were examined. Replicates of four genotypes of Solidago altissima, grown under low, medium, or high levels of nutrient supply in full sun or with medium levels of nutrients in shade, were exposed to mass-released Eurosta solidaginis. The number of plants ovipunctured was significantly affected by plant genotype and the interaction between genotype and nutrient supply to the host plant: one susceptible and one resistant genotype were more preferred, and preference tended to increase with nutrient supply in the more-preferred genotypes. The growth rate of ovipunctured plants during the oviposition period was significantly greater than that of unpunctured plants. Bud diameter (which was strongly correlated with plant growth rate), leaf area, and leaf water content were significant determinants of the percentage of plants ovipunctured, explaining 74% of the variance. The number of surviving larvae was significantly affected by plant genotype, but no effect of nutrient or light supply to the host plant was detected. The ratio of bud diameter to bud length was positively related to the percentage of ovipunctured plants that formed galls, suggesting that the accurate placement of eggs near the apical meristem by ovipositing females may be easier in short, thick buds. No significant correlation was observed between oviposition preference and larval survival at the population level. These results suggest that the plant traits affecting oviposition preference may exhibit different magnitudes of phenotypic plasticity than those affecting larval survival, and that the degree of phenotypic plasticity in plant traits affecting oviposition preference may differ among genotypes within a species.     

Environmental Heterogeneity and Population Differentiation in Plasticity to Drought in <Emphasis Type="Italic">Convolvulus Chilensis</Emphasis> (Convolvulaceae)

Plant populations may show differentiation in phenotypic plasticity, and theory predicts that greater levels of environmental heterogeneity should select for higher magnitudes of phenotypic plasticity. We evaluated phenotypic responses to reduced soil moisture in plants of Convolvulus chilensis grown in a greenhouse from seeds collected in three natural populations that differ in environmental heterogeneity (precipitation regime). Among several morphological and ecophysiological traits evaluated, only four traits showed differentiation among populations in plasticity to soil moisture: leaf area, leaf shape, leaf area ratio (LAR), and foliar trichome density. In all of these traits plasticity to drought was greatest in plants from the population with the highest interannual variation in precipitation. We further tested the adaptive nature of these plastic responses by evaluating the relationship between phenotypic traits and total biomass, as a proxy for plant fitness, in the low water environment. Foliar trichome density appears to be the only trait that shows adaptive patterns of plasticity to drought. Plants from populations showing plasticity had higher trichome density when growing in soils with reduced moisture, and foliar trichome density was positively associated with total biomass. Co-ordinating editor: F. Stuefer     

Mechanisms driving plant functional trait variation in a tropical forest

Plant functional trait variation in tropical forests results from taxonomic differences in phylogeny and associated genetic differences, as well as, phenotypic plastic responses to the environment. Accounting for the underlying mechanisms driving plant functional trait variation is important for understanding the potential rate of change of ecosystems since trait acclimation via phenotypic plasticity is very fast compared to shifts in community composition and genetic adaptation. We here applied a statistical technique to decompose the relative roles of phenotypic plasticity, genetic adaptation, and phylogenetic constraints. We examined typically obtained plant functional traits, such as wood density, plant height, specific leaf area, leaf area, leaf thickness, leaf dry mass content, leaf nitrogen content, and leaf phosphorus content. We assumed that genetic differences in plant functional traits between species and genotypes increase with environmental heterogeneity and geographic distance, whereas trait variation due to plastic acclimation to the local environment is independent of spatial distance between sampling sites. Results suggest that most of the observed trait variation could not be explained by the measured environmental variables, thus indicating a limited potential to predict individual plant traits from commonly assessed parameters. However, we found a difference in the response of plant functional traits, such that leaf traits varied in response to canopy‐light regime and nutrient availability, whereas wood traits were related to topoedaphic factors and water availability. Our analysis furthermore revealed differences in the functional response of coexisting neotropical tree species, which suggests that endemic species with conservative ecological strategies might be especially prone to competitive exclusion under projected climate change.     

SHORT INTERNODES-like genes regulate shoot growth and xylem proliferation in Populus

? Genes controlling plant growth and form are of considerable interest, because they affect survival and productivity traits, and are largely unknown or poorly characterized. The SHORT INTERNODES(SHI) gene is one of a 10-member SHI-RELATED SEQUENCE (SRS) gene family in Arabidopsis that includes important developmental regulators. ? Using comparative sequence analysis of the SRS gene families in poplar and Arabidopsis, we identified two poplar proteins that are most similar to SHI and its closely related gene STYLISH1 (STY1). The two poplar genes are very similar in sequence and expression and are therefore probably paralogs with redundant functions. ? RNAi suppression of the two Populus genes enhanced shoot and root growth, whereas the overexpression of Arabidopsis SHI in poplar reduced internode and petiole length. The suppression of the two genes increased fiber length and the proportion of xylem tissue, mainly through increased xylem cell proliferation. The transgenic modifications were also associated with significant changes in the concentrations of gibberellins and cytokinin. ? We conclude that Populus SHI-RELATED SEQUENCE (SRS) genes play an important role in the regulation of vegetative growth, including wood formation, and thus could be useful tools for the modification of biomass productivity, wood quality or plant form.     

The genetics and genomics of the drought response in Populus

The genetic nature of tree adaptation to drought stress was examined by utilizing variation in the drought response of a full-sib second generation (F(2)) mapping population from a cross between Populus trichocarpa (93-968) and P. deltoides Bart (ILL-129) and known to be highly divergent for a vast range of phenotypic traits. We combined phenotyping, quantitative trait loci (QTL) analysis and microarray experiments to demonstrate that 'genetical genomics' can be used to provide information on adaptation at the species level. The grandparents and F(2) population were subjected to soil drying, and contrasting responses to drought across genotypes, including leaf coloration, expansion and abscission, were observed, and QTL for these traits mapped. A subset of extreme genotypes exhibiting extreme sensitivity and insensitivity to drought on the basis of leaf abscission were defined, and microarray experiments conducted on these genotypes and the grandparent species. The extreme genotype groups induced a different set of genes: 215 and 125 genes differed in their expression response between groups in control and drought, respectively, suggesting species adaptation at the gene expression level. Co-location of differentially expressed genes with drought-specific and drought-responsive QTLs was examined, and these may represent candidate genes contributing to the variation in drought response.     

Morphological and Physiological Acclimation Responses to Contrasting Light and Water Regimes in <Emphasis Type="Italic">Primula sieboldii</Emphasis>

The effects of the availability of light (high, medium and low) and soil water (wet and dry) on morphological and physiological traits responsible for whole plant carbon gain and ramet biomass accumulation were examined in a splitter-type clonal herbaceous species Primula sieboldii, a spring plant inhabiting broad range of light environments including open grassland and oak forest understory. Growth experiments were conducted for three genets originated from natural microhabitats differing in light and soil water availability. Ramets of a genet from high light and wet microhabitat, which were grown in low light (relative photon flux density: R-PPFD of 5%) showed 41% less light-saturated photosynthetic rate, 50% less dark respiration rate and earlier defoliation than the ramets in high light (R-PPFD of 61%). The estimation of daily photosynthesis revealed that the light acclimation response in leaf gas exchange contributes to efficient carbon gain of whole plants, irrespective of experimental light conditions. Water stress increased root weight ratio, decreased ramet leaf area, petiole length and photosynthetic capacity. These morphological effects of water stress were larger in high and medium light regimes than in low light regime. The consequence of the above responses was recognized in the relative growth rate of the ramets. The relative growth rate of the ramets in high light with wet regime was four-fold of that in low light plus wet regime, and was 1.5-fold of that in high light plus dry regime. However, even in low light and/or dry regimes, ramets kept positive relative growth rates and produced gemma successfully. We could not detect significant variation in growth responses among genets. The high photosynthetic plasticity revealed in the present study should enable Primula sieboldii to inhabit in a broad range of light and soil water availability.     

Leaf shape linked to photosynthetic rates and temperature optima in South African <Emphasis Type="Italic">Pelargonium</Emphasis> species

The thermal response of gas exchange varies among plant species and with growth conditions. Plants from hot dry climates generally reach maximal photosynthetic rates at higher temperatures than species from temperate climates. Likewise, species in these environments are predicted to have small leaves with more-dissected shapes. We compared eight species of Pelargonium (Geraniaceae) selected as phylogenetically independent contrasts on leaf shape to determine whether: (1) the species showed plasticity in thermal response of gas exchange when grown under different water and temperature regimes, (2) there were differences among more- and less-dissected leafed species in trait means or plasticity, and (3) whether climatic variables were correlated with the responses. We found that a higher growth temperature led to higher optimal photosynthetic temperatures, at a cost to photosynthetic capacity. Optimal temperatures for photosynthesis were greater than the highest growth temperature regime. Stomatal conductance responded to growth water regime but not growth temperature, whereas transpiration increased and water use efficiency (WUE) decreased at the higher growth temperature. Strikingly, species with more-dissected leaves had higher rates of carbon gain and water loss for a given growth condition than those with less-dissected leaves. Species from lower latitudes and lower rainfall tended to have higher photosynthetic maxima and conductance, but leaf dissection did not correlate with climatic variables. Our results suggest that the combination of dissected leaves, higher photosynthetic rates, and relatively low WUE may have evolved as a strategy to optimize water delivery and carbon gain during short-lived periods of high soil moisture. Higher thermal optima, in conjunction with leaf dissection, may reflect selection pressure to protect photosynthetic machinery against excessive leaf temperatures when stomata close in response to water stress.     

不同坡向川西亚高山林木竞争与叶片表型可塑性的关系研究

表型可塑性是植物生长响应外界环境变化的重要表现形式,体现了植物个体在环境胁迫下的适合度。但是关于植物表型可塑性的驱动机制仍然存在很多争议。为了探讨植物表型可塑性的影响因素,以四川省阿坝藏族羌族自治州位于同一海拔梯度但坡向相反的天然次生林为研究对象,分析了不同坡向竞争强度与10种树木叶片功能性状表型可塑性的关系的差异。研究发现：(1)研究样地中阴坡水分和养分资源优于阳坡;(2)阴坡上林木平均种内和种间竞争强度高于阳坡,阴坡上林木种内竞争强度随着树木个体大小的增加而显著性减少,阳坡上林木种内竞争强度却随着个体大小的增加而增加;(3)阴坡上叶片表型可塑性高于阳坡,表型可塑性随着个体大小的增加而增加,在阳坡上却随着个体大小增加而降低。这些结果表明阴坡上水分等资源环境条件优于阳坡,林木生长受到环境资源限制较少。在林木生长过程中,较高的竞争强度引起的资源重叠加剧,尤其是种内竞争强度的变化,从而导致了阴坡上较高的叶片表型可塑性。因为较高的竞争强度,随着林木个体大小的增加,树木需要更高的可塑性赢得竞争优势来获取更多的资源支持生长。但是在阳坡上,资源相对缺乏,环境资源对树木生长的限制降低了叶片表型的可塑...