Drought responses,phenotypic plasticity and survival of Mediterranean species in two different microclimatic sites

• Climate models predict a further drying of the Mediterranean summer. One way for plant species to persist during such climate changes is through acclimation. Here, we determine the extent to which trait plasticity in response to drought differs between species and between sites, and address the question whether there is a trade‐off between drought survival and phenotypic plasticity.
• Throughout the summer we measured physiological traits (photosynthesis – A_max, stomatal conductance – g_s, transpiration – E, leaf water potential – ψl) and structural traits (specific leaf area – SLA, leaf density – LD, leaf dry matter content – LDMC, leaf relative water content – LRWC) of leaves of eight woody species in two sites with slightly different microclimate (north‐ versus south‐facing slopes) in southern Spain. Plant recovery and survival was estimated after the summer drought period.
• We found high trait variability between species. In most variables, phenotypic plasticity was lower in the drier site. Phenotypic plasticity of SLA and LDMC correlated negatively with drought survival, which suggests a trade‐off between them. On the other hand, high phenotypic plasticity of SLA and LDMC was positively related to traits associated with rapid recovery and growth after the drought period.
• Although phenotypic plasticity is generally seen as favourable during stress conditions, here it seemed beneficial for favourable conditions. We propose that in environments with fluctuating drought periods there can be a trade‐off between drought survival and growth during favourable conditions. When climate become drier, species with high drought survival but low phenotypic plasticity might be selected for.

     

Geographical variation and the role of climate in leaf traits of a relict tree species across its distribution in China

• Intraspecific trait variation and trait–climate relationships are crucial for understanding a species’ response to climate change. However, these phenomena have rarely been studied for tree species. Euptelea pleiospermum is a relict tree species with a wide distribution in China that offers a novel opportunity to examine such relationships.
• Here, we measured 13 leaf traits of E. pleiospermum in 20 sites across its natural distribution in China. We investigated the extent of trait variation at local and regional scales, and developed geographic and climate models to explain trait variation at the regional scale.
• We documented intraspecific trait variation among leaf traits of E. pleiospermum at local and regional scales. Five traits exhibited relatively high trait variation: leaf area, leaf density and three leaf economic traits (leaf dry matter content, specific leaf area [SLA] and leaf phosphorus concentration). Significant trait–geography correlations were mediated by local climate. Most leaf trait variation could be explained (from 24% to 64%) by geographic or climate variables, except leaf width, leaf thickness, leaf dry matter content and leaf length–width ratio. Latitude and temperature were the strongest predictors of trait variation throughout the distribution of E. pleiospermum in China, and temperature explained more leaf trait variation than precipitation. In particular, we showed that leaves had longer petiole lengths, higher SLA and lower densities in northern E. pleiospermum populations. We suggest that northern E. pleiospermum populations are adapting to higher latitudinal environments via high growth rate (higher SLA) and low construction investment strategies (lower leaf densities), benefitting northern migration.
• Overall, we demonstrate that intraspecific trait variation reflects E. pleiospermum response to the local environment. We call for consideration of intraspecific trait variation to examine specific climate response questions. In addition, provenance experiments using widely distributed species are needed to separate trait variation resulting from genetic differentiation and plastic responses to environmental change.

     

Leaf shape variation and herbivore consumption and performance: a case study with Ipomoea hederacea and three generalists

The effect of leaf shape variation on plant-herbivore interactions has primarily been studied from the perspective of host seeking behavior. Yet for leaf shape to affect plant-herbivore coevolution, there must be reciprocal effects of leaf shape variation on herbivore consumption and performance. We investigated whether alternative leaf morphs affected the performance of three generalist insect herbivores by taking advantage of a genetic polymorphism and developmental plasticity in leaf shape in the Ivyleaf morning glory, Ipomoea hederacea. Across four experiments, we found variable support for an effect of leaf shape genotype on insects. For cabbage loopers (Trichoplusia ni) and corn earworms (Helicoverpa zea) we found opposing, non-significant trends: T. ni gained more biomass on lobed genotypes, while H. zea gained more biomass on heart-shaped genotypes. For army beetworms (Spodoptera exigua), the effects of leaf shape genotype differed depending on the age of the plants and photoperiod of growing conditions. Caterpillars feeding on tissue from older plants (95 days) grown under long day photoperiods had significantly greater consumption, dry biomass, and digestive efficiency on lobed genotypes. In contrast, there were no significant differences between heart-shaped and lobed genotypes for caterpillars feeding on tissue from younger plants (50 days) grown under short day photoperiods. For plants grown under short days, we found that S. exigua consumed significantly less leaf area when feeding on mature leaves than juvenile leaves, regardless of leaf shape genotype. Taken together, our results suggest that the effects of leaf shape variation on insect performance are likely to vary between insect species, growth conditions of the plant, and the developmental stage and age of leaves sampled. Handling editor: May Berenbaum.     

Phenotypic plasticity closely linked to climate at origin and resulting in increased mortality under warming and frost stress in a common grass

Phenotypic plasticity is important for species responses to global change and species coexistence. Phenotypic plasticity differs among species and traits and changes across environments. Here, we investigated phenotypic plasticity of the widespread grass Arrhenatherum elatius in response to winter warming and frost stress by comparing phenotypic plasticity of 11 geographically and environmentally distinct populations of this species to phenotypic plasticity of populations of different species originating from a single environment. The variation in phenotypic plasticity was similar for populations of a single species from different locations compared to populations of functionally and taxonomically diverse species from one environment for the studied traits (leaf biomass production and root integrity after frost) across three indices of phenotypic plasticity (RDPI, PIN, slope of reaction norm). Phenotypic plasticity was not associated with neutral genetic diversity but closely linked to the climate of the populations’ origin. Populations originating from warmer and more variable climates showed higher phenotypic plasticity. This indicates that phenotypic plasticity can itself be considered as a trait subject to local adaptation to climate. Finally, our data emphasize that high phenotypic plasticity is not per se positive for adaptation to climate change, as differences in stress responses are resulting in high phenotypic plasticity as expressed by common plasticity indices, which is likely to be related to increased mortality under stress in more plastic populations.     

Characterizing the contribution of plasticity and genetic differentiation to community‐level trait responses to environmental change

The match between functional trait variation in communities and environmental gradients is maintained by three processes: phenotypic plasticity and genetic differentiation (intraspecific processes), and species turnover (interspecific). Recently, evidence has emerged suggesting that intraspecific variation might have a potentially large role in driving functional community composition and response to environmental change. However, empirical evidence quantifying the respective importance of phenotypic plasticity and genetic differentiation relative to species turnover is still lacking. We performed a reciprocal transplant experiment using a common herbaceous plant species (Oxalis montana) among low‐, mid‐, and high‐elevation sites to first quantify the contributions of plasticity and genetic differentiation in driving intraspecific variation in three traits: height, specific leaf area, and leaf area. We next compared the contributions of these intraspecific drivers of community trait–environment matching to that of species turnover, which had been previously assessed along the same elevational gradient. Plasticity was the dominant driver of intraspecific trait variation across elevation in all traits, with only a small contribution of genetic differentiation among populations. Local adaptation was not detected to a major extent along the gradient. Fitness components were greatest in O. montana plants with trait values closest to the local community‐weighted means, thus supporting the common assumption that community‐weighted mean trait values represent selective optima. Our results suggest that community‐level trait responses to ongoing climate change should be mostly mediated by species turnover, even at the small spatial scale of our study, with an especially small contribution of evolutionary adaptation within species.     

Effects of fire,topography and year-to-year climatic variation on species composition in tallgrass prairie

Native unploughed tallgrass prairie from Konza Prairie, Kansas, USA is described with respect to plant species compositional changes over a five year period in response to fire and topography. The principal gradient of variation in the vegetation is related to time since burning. Species show an individualistic response in terms of relative abundance to this gradient. Both the percentage of and cover of C₄ species and all grasses decrease as the prairie remains unburnt. Forb and woody plant species numbers and abundance increase along this gradient. A secondary gradient of variation reflects topography (i.e. upland versus lowland soils). Upland soils support a higher species richness and diversity. Upland and lowland plant assemblages are distinct except on annually burnt prairie. The interaction between burning regime, topography and year-to-year climatic variation affects the relative abundance of the plant species differentially. The most dominant species overall, Andropogon gerardii, was affected only by year-to-year variation (i.e. climate). Its position at the top of the species abundance hierarchy was unaffected by burning regime or soil type. The other dominant species showed a suite of varying responses to these factors.Deceased May, 1986.     

Plasticity of functional traits varies clinally along a rainfall gradient in Eucalyptus tricarpa

Widespread species often occur across a range of climatic conditions, through a combination of local genetic adaptations and phenotypic plasticity. Species with greater phenotypic plasticity are likely to be better positioned to cope with rapid anthropogenic climate changes, while those displaying strong local adaptations might benefit from translocations to assist the movement of adaptive genes as the climate changes. Eucalyptus tricarpa occurs across a climatic gradient in south‐eastern Australia, a region of increasing aridity, and we hypothesized that this species would display local adaptation to climate. We measured morphological and physiological traits reflecting climate responses in nine provenances from sites of 460 to 1040 mm annual rainfall, in their natural habitat and in common gardens near each end of the gradient. Local adaptation was evident in functional traits and differential growth rates in the common gardens. Some traits displayed complex combinations of plasticity and genetic divergence among provenances, including clinal variation in plasticity itself. Provenances from drier locations were more plastic in leaf thickness, whereas leaf size was more plastic in provenances from higher rainfall locations. Leaf density and stomatal physiology (as indicated by δ¹³C and δ¹⁸O) were highly and uniformly plastic. In addition to variation in mean trait values, genetic variation in trait plasticity may play a role in climate adaptation.     

桂西南喀斯特地区优势木本经济植物叶功能性状变异及其适应策略

植物叶功能性状可直接或间接地反映植物对环境变化的响应与适应策略.该文采用野外气体交换测量和实验室分析相结合的方法,以桂西南典型喀斯特灌丛常见种龙须藤和黄荆为对照,对该区域5种优势木本经济植物的叶功能性状指标及其关系进行研究.结果表明:(1)11个叶功能性状指标种内均存在不同程度的变异,除胞间CO2浓度和水分利用效率外,...     

Genetic variation in circadian regulation of nocturnal stomatal conductance enhances carbon assimilation and growth

Circadian resonance, whereby a plant's endogenous rhythms are tuned to match environmental cues, has been repeatedly shown to be adaptive, although the underlying mechanisms remain elusive. Concomitantly, the adaptive value of nocturnal transpiration in C₃ plants remains unknown because it occurs without carbon assimilation. These seemingly unrelated processes are interconnected because circadian regulation drives temporal patterns in nocturnal stomatal conductance, with maximum values occurring immediately before dawn for many species. We grew individuals of six Eucalyptus camaldulensis genotypes in naturally lit glasshouses and measured sunset, predawn and midday leaf gas exchange and whole‐plant biomass production. We tested whether sunrise anticipation by the circadian clock and subsequent increases in genotype predawn stomatal conductance led to rapid stomatal opening upon illumination, ultimately affecting genotype differences in carbon assimilation and growth. We observed faster stomatal responses to light inputs at sunrise in genotypes with higher predawn stomatal conductance. Moreover, early morning and midday stomatal conductance and carbon assimilation, leaf area and total plant biomass were all positively correlated with predawn stomatal conductance across genotypes. Our results lead to the novel hypothesis that genotypic variation in the circadian‐regulated capacity to anticipate sunrise could be an important factor underlying intraspecific variation in tree growth.     

Effects of endophytic fungi on the phenotypic plasticity of Lolium perenne (Poaceae)

The effects of high vs. low levels of endophytic fungi on the phenotypic plasticity of cloned genotypes were examined in perennial ryegrass (Lolium perenne L.). The objectives were to determine whether endophytic fungi influence plastic responses of host genotypes to variable soil nutrients and whether or not endophyte infection and host genotype interact to determine the extent of this plasticity. Twelve infected genotypes were cloned into ramets: half the ramets were treated with the systemic fungicide Benomyl to reduce or eliminate the endophyte, while the other half were untreated. Ramets of each genotype were subjected to high, medium, or low levels of nutrients in the greenhouse for 11 wk. Tiller number, leaf area, and leaf mass were determined after 11 and 25 wk. The fungicide significantly reduced the level of endophyte infection. Responses to nutrient conditions in relation to fungicide treatment were genotype specific: for some genotypes, high levels of endophytic fungi appeared to reduce plasticity, while for other genotypes the endophyte had no effect. The potential for microscopic symbionts to affect phenotypic plasticity in genetically variable populations has not often been recognized. However, the clandestine effects of symbionts on the plasticity of host genotypes could impact microevolutionary processes occurring within plant populations that occupy heterogeneous environments.     

Genome‐wide association mapping for phenotypic plasticity in rice

Phenotypic plasticity of plants in response to environmental changes is important for adapting to changing climate. Less attention has been paid to exploring the advantages of phenotypic plasticity in resource‐rich environments to enhance the productivity of agricultural crops. Here, we examined genetic variation for phenotypic plasticity in indica rice (Oryza sativa L.) across two diverse panels: (1) a Phenomics of Rice Adaptation and Yield (PRAY) population comprising 301 accessions; and (2) a Multi‐parent Advanced Generation Inter‐Cross (MAGIC) indica population comprising 151 accessions. Altered planting density was used as a proxy for elevated atmospheric CO₂ response. Low planting density significantly increased panicle weight per plant compared with normal density, and the magnitude of the increase ranged from 1.10 to 2.78 times among accessions for the PRAY population and from 1.05 to 2.45 times for the MAGIC population. Genome‐wide‐association studies validate three E nvironmental R esponsiveness (ER) candidate alleles (qER1–3) that were associated with relative response of panicle weight to low density. Two of these alleles were tested in 13 genotypes to clarify their biomass responses during vegetative growth under elevated CO₂ in Japan. Our study provides evidence for polymorphisms that control rice phenotypic plasticity in environments that are rich in resources such as light and CO₂.     

Leaf morphological plasticity and stomatal conductance in three Populus alba L. genotypes subjected to salt stress

The effect of salt stress on leaf morphology and functionality was studied in three Populus alba genotypes differing in tolerance to salinity: 6K3 (sensitive), 2AS11 (moderately tolerant), and 14P11 (tolerant). Plants were subjected to an intense and progressive salt stress from 50 to 250 mM NaCl by 50 mM steps at 10-day intervals. The micromorphological results highlighted phenotypic variation among the three genotypes already in control plants, with the genotype 14P11 having significantly smaller epidermal cells and higher stomatal density. Salt-treated plants modulated differently the expansion of stomata compared with epidermal cells. Regression analysis showed significant correlations between decrease of stomatal area and stomatal conductance (g_s) in genotypes 14P11 and 6K3. So, the common reduction of stomatal area could be an early mechanism to save water in this species. However, only genotype 14P11 showed further significant decrease of this trait under the highest salinity level, combined with a significant reduction in leaf length. In addition, this genotype showed the lowest leaf abscission rate at the end of salt stress period. The genotype 6K3 was severely affected by leaf necrosis and showed the highest leaf abscission rate in salt stress conditions. In the moderately tolerant genotype 2AS11, an intermediate plastic behaviour in both leaf morphology and physiology was observed during the experiment. The phenotypic variation among the three genotypes in terms of micromorphology and stomatal conductance is discussed in relation to plant functionality in salt stress conditions. Overall results suggest that leaf morphological habit contributes to salt tolerance in P. alba.     

桂西南岩溶区八种适生植物光合性状的变异与关联

为探究岩溶植物的光合生理适应机制，采用Li-6400XT便携式光合作用测量系统，对广西平果市岩溶区8种适生植物的叶片净光合速率(P_n)、气孔导度(G_s)、胞间CO₂浓度(C_i)、蒸腾速率(T_r)、水分利用效率(WUE)和气孔限制值(L_s)等光合特征参数进行了测定分析。结果表明：(1)6个光合特征参数在种内和种间均存在不同程度的变异，并且种内变异均大于种间变异。(2)G_s和T_r的变化主要来源于种间变异(46.72%～49.76%),而P_n、C_i、WUE和L_s变化主要来源于种内变异(48.66%～64.50%)。在生活型水平上，P_n、G_s和T_r的种内变异表现为常绿植物小于落叶植物，而C_i、WUE和L_s则相反。(3)各参数的种间变异均表现为落叶植...     

Intraspecific variation in seedling growth responses of a relict tree species Euptelea pleiospermum to precipitation manipulation along an elevation gradient

Improving the accuracy of predictions regarding how plants respond to climate change is crucial to protecting biodiversity. However, little is known about the effects of seed source and elevation on the response of mountain plant species to reductions in precipitation. Here, we collected seeds of a tree species (Euptelea pleiospermum) from three seed sources and carried out a two-growing-season reciprocal transplant experiment with precipitation manipulation at three sites along an elevation gradient in the Shennongjia Mountains, central China. Variations in whole-plant traits, leaf traits, and root traits were investigated. We found that most plant traits of E. pleiospermum seedlings were affected by reductions in precipitation, and responses varied among different elevations and seed sources. Whole-plant traits, root biomass, and leaf traits related to photosynthesis capacity decreased under reduced precipitation treatments at mid and high elevation sites. Thus, climate change induced drought will likely have a negative influence on seedling growth at mid and high elevation regions. In addition, a home-site advantage in whole-plant traits and root traits was observed. However, the responses of leaf traits in most cases were not affected by seed source because of higher phenotypic plasticity. Our results suggested that both local adaptation and phenotypic plasticity were important in seedling growth responses to reduced precipitation. We also highlight the importance of taking intraspecific variation into account when studying the response of plants to changes in climate.

     

Neglecting acclimation of photosynthesis under drought can cause significant errors in predicting leaf photosynthesis in wheat

Extreme climatic events, such as heat waves, cold snaps and drought spells, related to global climate change, have become more frequent and intense in recent years. Acclimation of plant physiological processes to changes in environmental conditions is a key component of plant adaptation to climate change. We assessed the temperature response of leaf photosynthetic parameters in wheat grown under contrasting water regimes and growth temperatures (T_growth). Two independent experiments were conducted under controlled conditions. In Experiment 1, two wheat genotypes were subjected to well-watered or drought-stressed treatments; in Experiment 2, the two water regimes combined with high, medium and low T_growth were imposed on one genotype. Parameters of a biochemical C₃-photosynthesis model were estimated at six leaf temperatures for each factor combination. Photosynthesis acclimated more to drought than to T_growth. Drought affected photosynthesis by lowering its optimum temperature (T_opt) and the values at T_opt of light-saturated net photosynthesis, stomatal conductance, mesophyll conductance, the maximum rate of electron transport (J_max) and the maximum rate of carboxylation by Rubisco (V_cmax). T_opt for V_cmax was up to 40°C under well-watered conditions but 24–34°C under drought. The decrease in photosynthesis under drought varied among T_growth but was similar between genotypes. The temperature response of photosynthetic quantum yield under drought was partly attributed to photorespiration but more to alternative electron transport. All these changes in biochemical parameters could not be fully explained by the changed leaf nitrogen content. Further model analysis showed that both diffusional and biochemical parameters of photosynthesis and their thermal sensitivity acclimate little to T_growth, but acclimate considerably to drought and the combination of drought and T_growth. The commonly used modelling approaches, which typically consider the response of diffusional parameters, but ignore acclimation responses of biochemical parameters to drought and T_growth, strongly overestimate leaf photosynthesis under variable temperature and drought.     

Evidence of maternal effects on temperature preference in side‐blotched lizards: implications for evolutionary response to climate change

Natural populations respond to selection pressures like increasing local temperatures in many ways, including plasticity and adaptation. To predict the response of ectotherms like lizards to local temperature increase, it is essential to estimate phenotypic variation in and determine the heritability of temperature‐related traits like average field body temperature (T_b) and preferred temperature (T_p). We measured T_p of Uta stansburiana in a laboratory thermal gradient and assessed the contribution of sex, reproductive status and throat color genotype to phenotypic variation in T_b of adult lizards. Females had higher T_p than males. However, they temporarily preferred lower temperature when gravid than when nongravid. Using a nested half‐sib design for genetic crosses in the laboratory, we estimated relative contributions of additive genetic variation and maternal effects to T_p of hatchlings. Our results show that maternal effects, but not additive genetic variation, influence T_p of hatchlings in U. stansburiana. Maternal T_p and the presence or absence of blue throat color alleles significantly influenced T_p of hatchlings. We discuss ecological and evolutionary consequences of these maternal effects in the context of rapid climate change and natural selection that we measure on progeny survival to maturity as a function of maternal T_p.     

Variation in relative growth rate and its components in the annual Polygonum aviculare in relation to habitat disturbance and seed size

Polygonum aviculare is an annual weedy species showing extensive genetic variation in seed and leaf size and colonizing various types of man-disturbed habitats. A growth analysis was conducted on 12 genotypes representative of three regimes of disturbance of natural habitat (trampling, weeding, and no disturbance in the course of the growing season), grown under productive conditions in order to test whether relative growth rate (RGR) varies at the intraspecific level and, if so, which growth parameters may explain its variation. RGR showed significant genotypic variation (0.355–0.452 g g^-1 day^-1), positively correlated with specific leaf area (SLA) and leaf mass ratio (LMR) and negatively correlated with unit leaf rate per unit leaf area (ULR_A). Thus, the paramount importance of leaf area ratio (LAR=SLA×LMR) in determining growth rate variation between different herbaceous species is confirmed at the intraspecific level in this species. Genotypes originating from trampled habitats had smaller seeds and smaller leaves than genotypes from habitats subject to other disturbance regimes. Additionally, they showed a lower LAR, not entirely compensated for by a higher ULR_A, which resulted in a positive allometric relationship between seed size and RGR. It is hypothesized that their lower SLA, correlated with a higher leaf dry matter content (possibly a consequence of a higher cell wall content per unit leaf area) and their lower LMR have been co-selected with small leaf size as adaptations promoting resistance to trampling stress. It is suggested that variation in cell size and/or gibberellin content might be the mediators of the correlation found between seed size, leaf size and growth parameters within this species.     

Intraspecific variation in salt tolerance and morphology in the coastal grass spartina patens (poaceae)

Clones of Spartina patens were collected from 19 locations throughout Gulf coast marshes of Texas, Louisiana, and Florida. Following three vegetative generations of de-acclimation from field conditions, genotypes were subjected to a salinity screening protocol in which salinity was increased in weekly increments of 5‰ (gram salt/kilogram solution). Plants were harvested when there was 50% death of aboveground tissue, which we defined as the lethal salinity level. The genotypes displayed highly significant intraspecific variation in lethal salinity level, which ranged from 63‰ to 93‰. Significant intraspecific variation was also observed in all plant morphological variables, as well as leaf rolling, leaf expansion rates at 2‰ and 20‰ salinity, aboveground, belowground, and total biomass, and belowground-to-aboveground biomass ratio. An ANOVA of principal component scores from a PCA of lethal salinity level and covariable-adjusted total plant dry mass further illustrated intraspecific variation within this species in these two traits expressed as one principal component. Correlation analysis revealed that intraspecific variation in salt tolerance was not strongly associated with intraspecific variation in plant morphological traits, leaf rolling, or leaf expansion rates.     

Global patterns of intraspecific leaf trait responses to elevation

Elevational gradients are often used to quantify how traits of plant species respond to abiotic and biotic environmental variations. Yet, such analyses are frequently restricted spatially and applied along single slopes or mountain ranges. Since we know little on the response of intraspecific leaf traits to elevation across the globe, we here perform a global meta‐analysis of leaf traits in 109 plant species located in 4 continents and reported in 71 studies published between 1983 and 2018. We quantified the intraspecific change in seven morpho‐ecophysiological leaf traits along global elevational gradients: specific leaf area (SLA), leaf mass per area (LMA), leaf area (LA), nitrogen concentration per unit of area (Narea), nitrogen concentration per unit mass (Nmass), phosphorous concentration per unit mass (Pmass) and carbon isotope composition (δ¹³C). We found LMA, Narea, Nmass and δ¹³C to significantly increase and SLA to decrease with increasing elevation. Conversely, LA and Pmass showed no significant pattern with elevation worldwide. We found significantly larger increase in Narea, Nmass, Pmass and δ¹³C with elevation in warmer regions. Larger responses to increasing elevation were apparent for SLA of herbaceous compared to woody species, but not for the other traits. Finally, we also detected evidences of covariation across morphological and physiological traits within the same elevational gradient. In sum, we demonstrate that there are common cross‐species patterns of intraspecific leaf trait variation across elevational gradients worldwide. Irrespective of whether such variation is genetically determined via local adaptation or attributed to phenotypic plasticity, the leaf trait patterns quantified here suggest that plant species are adapted to live on a range of temperature conditions. Since the distribution of mountain biota is predominantly shifting upslope in response to changes in environmental conditions, our results are important to further our understanding of how plants species of mountain ecosystems adapt to global environmental change.