Differences In Drought Response Research Articles

Contrasting growth responses to drought in three tree species widely distributed in northern China

Tree species-specific responses to drought are urgently needed for assessing the impacts of current climate change on forest ecosystems. Here, we characterized the resistance, recovery, resilience, and growth recovery periods in response to different drought events based on tree-ring width index data (>30 years) for three tree species widely distributed in northern China, among which larch (Larix principis-rupprechtii Mayr) and Mongolian pine (Pinus sylvestris L. var. mongolica Litv.) are two major species used for afforestation, and birch (Betula platyphylla Suk.) is one of the most common natural tree species. Despite no significant effects of mild drought on tree growth, severe drought events significantly reduced the growth of all species, with contrasting species-specific responses. Larch trees had the lowest resistance and resilience among the three species, and Mongolian pine trees were more resistant but had a longer recovery period than birch trees. The drought responses varied with tree size. Large Mongolian pine and birch trees were more resistant but large larch trees were much more vulnerable than small trees during severe droughts. Smaller birch trees had higher resilience to severe droughts. Our study shows species-specific differences in drought responses and suggests that drought responses are tree-size dependent and drought-intensity associated, which further provides a guidance for selecting optimal cultivated tree species and designing forest managements in this region.

Effects of thinning on tradeoffs between drought resistance, drought resilience, and wood production in mature Douglas-fir in western Oregon, USA

Climate projections predict more frequent and severe drought in coastal Pseudotsuga menziesii forests of western North America, raising concerns over how to promote drought adaptation. Thinning often increases drought resistance (the ability to maintain growth during a drought) and resilience (the ability to recover growth after a drought), but these effects vary with thinning intensity, shift over time, and may have tradeoffs with fiber production. We collected tree cores from a long-term thinning study with four residual density levels replicated across both uniform thinning and thinning with gaps, and used annual growth data to investigate responses to droughts occurring 8 and 21 years after thinning. For the first drought, resistance and resilience were higher in treatments with lower residual densities. For the second drought, there were no differences in drought response between the lowest and highest residual density treatments, and all treatments had lower drought resistance and resilience than for the first drought. Spatial arrangement had little impact on drought resistance or resilience and residual density level had a significant effect on the periodic annual volume increment—drought resistance tradeoff. Our results suggest that thinning can promote drought adaptation in Pseudotsuga menziesii forests, but these effects dissipate over time.

Drought survival strategies differ between coastal and montane conifers in northern California

AbstractIncreasingly severe and prolonged droughts are contributing to tree stress and forest mortality across western North America. However, in many cases, we currently have poor information concerning how drought responses in forests vary in relation to competition, climate, and site and tree characteristics. We used annual tree ring evidence of 13C discrimination (Δ13C) and growth metrics to assess drought resistance and resilience for six conifer species at the intersection of several bioregions in northern California. Within each species' range in northern California, we collected competition and tree characteristics from 270 focal trees across sites that varied from wetter to drier habitat conditions (54 sites). Across sites, all six conifer species weathered the severe 2013–2015 drought with reasonably high resistance and post‐drought resilience. However, we found important differences in drought responses between coastal and montane species based on annual growth and Δ13C metrics. Broadly, the two coastal species showed consistent declines in drought resistance across successive drought years, whereas the four montane species maintained high drought resistance across drought years. More specifically, we found lower Δ13C and growth during drought years in coastal species, suggesting stomatal closure during drought with the potential for vulnerability to carbon depletion during long‐term drought. Conversely, Δ13C and growth were stable in montane species throughout the drought, which may contribute to hydraulic failure under increased drought frequency and/or severity. We also evaluated environmental factors that affect Δ13C using data from before and during the drought. These physiological models were consistent for the two coastal species, with a positive relationship between annual precipitation and Δ13C and a negative relationship between tree density and Δ13C. Conversely, the four montane models illustrated a greater importance of site conditions on drought responses for these species. Our findings show differential risk for drought stress across diverse conifers during severe drought. This work highlights the importance of site and tree characteristics in determining drought responses across cool, annually humid coastal habitats to seasonally dry montane habitats.

Drought response in Arabidopsis displays synergistic coordination between stems and leaves.

The synergy between drought-responsive traits across different organs is crucial in the whole-plant mechanism influencing drought resilience. These organ interactions, however, are poorly understood, limiting our understanding of drought response strategies at the whole-plant level. Therefore, we need more integrative studies, especially on herbaceous species that represent many important food crops but remain underexplored in their drought response. We investigated inflorescence stems and rosette leaves of six Arabidopsis thaliana genotypes with contrasting drought tolerance, and combined anatomical observations with hydraulic measurements and gene expression studies to assess differences in drought response. The soc1ful double mutant was the most drought-tolerant genotype based on its synergistic combination of low stomatal conductance, largest stomatal safety margin, more stable leaf water potential during non-watering, reduced transcript levels of drought stress marker genes, and reduced loss of chlorophyll content in leaves, in combination with stems showing the highest embolism resistance, most pronounced lignification, and thickest intervessel pit membranes. In contrast, the most sensitive Cvi ecotype shows the opposite extreme of the same set of traits. The remaining four genotypes show variations in this drought syndrome. Our results reveal that anatomical, ecophysiological, and molecular adaptations across organs are intertwined, and multiple (differentially combined) strategies can be applied to acquire a certain level of drought tolerance.

Multispectral image analysis detects differences in drought responses in novel seeded Miscanthus sinensis hybrids

AbstractDrought tolerance in Miscanthus sinensis is a desirable trait because of its potential use to develop new varieties, namely intra‐specific hybrids, adapted to drought‐prone marginal lands. In this study, drought tolerance was evaluated on 8 M. sinensis intra‐specific hybrids (GRC1–GRC8). Plants were grown in the growth chamber (14/10 h, 25/20°C, 70% relative air humidity and 300 μmol m−2 s−1 irradiance). Drought was induced by withholding irrigation for 21 days (stress phase) and after re‐watering (recovery phase) for 7 days. Nondestructive multispectral 3D images for plant morphology, color, and chlorophyll fluorescence imaging were used to quantify drought‐induced changes on a weekly basis for the entire duration of the experiment. Total leaf area (TLA) and digital biomass (DB) responded most rapidly to water deficits (7–14 days), followed by leaf senescence (14–21 days), and finally, a drop in the maximum efficiency of PSII (Fv/Fm; 21 days). Traits measured on the last day of the drought treatment were used to calculate the phenotypic plasticity. Significant differences in drought susceptibility and phenotypic plasticity were found among the studied hybrids. Drought treatment (21 days) reduced DB and TLA on average by 50%, normalized difference vegetation index (NDVI) by 20% and Fv/Fm from 0.79 (in control) to 0.69 in the less drought‐susceptible hybrids (GRC4 and GRC5), whereas in drought‐sensitive hybrids (GRC2 and GRC3), 21 days of drought reduced DB and TLA on average by 80%, NDVI by 45% and Fv/Fm dropped from 0.79 (in control) to 0.35. The more drought‐resilient hybrids showed lower phenotypic plasticity than their more sensitive counterparts, a stay‐green strategy enabled through lower biomass accumulation and by extension, reduced water usage. Multispectral imaging and image analysis enabled fast and nondestructive quantification of plant morphological and physiological responses under drought conditions and could be used as an effective screening tool for drought susceptibility.

Limited hydraulic recovery in seedlings of six tree species with contrasting leaf habits in subtropical China.

Subtropical tree species may experience severe drought stress due to variable rainfall under future climates. However, the capacity to restore hydraulic function post-drought might differ among co-occurring species with contrasting leaf habits (e.g., evergreen and deciduous) and have implications for future forest composition. Moreover, the links between hydraulic recovery and physiological and morphological traits related to water-carbon availability are still not well understood. Here, potted seedlings of six tree species (four evergreen and two deciduous) were grown outdoors under a rainout shelter. They grew under favorable water conditions until they were experimentally subjected to a soil water deficit leading to losses of ca. 50% of hydraulic conductivity, and then soils were re-watered to field capacity. Traits related to carbon and water relations were measured. There were differences in drought responses and recovery between species, but not as a function of evergreen or deciduous groups. Sapindus mukorossi exhibited the most rapid drought response, which was associated with a suite of physiological and morphological traits (larger plant size, the lowest hydraulic capacitance (Cbranch), higher minimum conductance (gmin) and lower HV (Huber value)). Upon re-watering, xylem water potential exhibited fast recovery in 1–3 days among species, while photosynthesis at saturating light (Asat) and stomatal conductance (gs) recovery lagged behind water potential recovery depending on species, with gs recovery being more delayed than Asat in most species. Furthermore, none of the six species exhibited significant hydraulic recovery during the 7 days re-watering period, indicating that xylem refilling was apparently limited; in addition, NSC availability had a minimal role in facilitating hydraulic recovery during this short-term period. Collectively, if water supply is limited by insignificant hydraulic recovery post-drought, the observed carbon assimilation recovery of seedlings may not be sustained over the longer term, potentially altering seedling regeneration and shifting forest species composition in subtropical China under climate change.

Inferring the genetic responses to acute drought stress across an ecological gradient

BackgroundHow do xerophytic species thrive in environments that experience extreme annual drought? Although critical to the survival of many species, the genetic responses to drought stress in many non-model organisms has yet to be explored. We investigated this question in Mentzelia section Bartonia (Loasaceae), which occurs throughout western North America, including arid lands. To better understand the genetic responses to drought stress among species that occur in different habitats, the gene expression levels of three species from Mentzelia were compared across a precipitation gradient. Two de novo reference transcriptomes were generated and annotated. Leaf and root tissues were collected from control and drought shocked plants and compared to one another for differential expression. A target-gene approach was also implemented to better understand how drought-related genes from model and crop species function in non-model systems.ResultsWhen comparing the drought-shock treatment plants to their respective control plants, we identified 165 differentially expressed clusters across all three species. Differentially expressed genes including those associated with water movement, photosynthesis, and delayed senescence. The transcriptome profiling approach was coupled with a target genes approach that measured expression of 90 genes associated with drought tolerance in model organisms. Comparing differentially expressed genes with a ≥ 2 log-fold value between species and tissue types showed significant differences in drought response. In pairwise comparisons, species that occurred in drier environments differentially expressed greater genes in leaves when drought shocked than those from wetter environments, but expression in the roots mostly produced opposite results.ConclusionsArid-adapted species mount greater genetic responses compared to the mesophytic species, which has likely evolved in response to consistent annual drought exposure across generations. Drought responses also depended on organ type. Xerophytes, for example, mounted a larger response in leaves to downregulate photosynthesis and senescence, while mobilizing carbon and regulating water in the roots. The complexity of drought responses in Mentzelia suggest that whole organism responses need to be considered when studying drought and, in particular, the physiological mechanisms in which plants regulate water, carbon, cell death, metabolism, and secondary metabolites.

Drought tolerance traits of leaves of 20 tree species in temperate forest of Northeast China

The increases in frequency and intensity of drought worldwide has seriously affected tree growth, and even led to widespread forest mortality. Leaf traits estimated from pressure-volume (PV) curve provide key leaf physiological information that reflects the drought tolerance of trees. However, it is uncertain that which PV parameter performs the best at local scale. Here, we measured five PV traits (including TLP, π0, ε, Cleaf, and RWCtlp) and two leaf structural traits (specific leaf area and leaf density) in 20 tree species (16 angiosperms and 4 gymnosperms) in a temperate mixed forest at the Maoershan Forest Ecosystem Research Station, Northeast China. The objectives of this study were to search the best indicators of leaf drought tolerance at local scale, and to explore the correlation between PV traits and leaf structural traits. We found that angiosperms had significantly greater RWCtlp and lower Cleaf than gymnosperms, indicating that RWCtlp and Cleaf might be the good indicators of leaf drought tolerance in temperate mixed forest in Northeast China. Within angiosperm species, TLP and π0 were significantly and negatively correlated with leaf density, but positively correlated with specific leaf area; while ε was negatively correlated with specific leaf area. However, the opposite trends between PV traits and leaf structural traits were observed between gymnosperms and angiosperms, which might be attributed to their differences in drought response and adaptation strategies.

Inter-specific tolerance to recurrent droughts of pine species revealed in saplings rather than adult trees

Understanding species’ tolerance to recurrent extreme droughts is key to predict species’ performance and forest dynamics under ongoing climate change. Inter-specific differences in juvenile responses can largely shape forest composition and structure. However, the vulnerability of tree species is typically evaluated in adult canopy dominant individuals. The question of whether inter-specific differences in drought tolerance vary between adult and young trees remains largely unexplored. We sampled canopy dominant trees and saplings of three co-occurring pine species with different drought tolerance (Pinus pinaster, P. nigra and P. sylvestris) in a relict forest in the Northern Iberian plateau. Using basal area increment in canopy dominant trees and shoot elongation in saplings, we reconstructed annual growth for the period 2000–2015 and calculated resilience and resistance indices to two consecutive extreme droughts (2005 and 2012). We used generalized least square models to analyze species-specific differences in resistance and resilience in adults and saplings, respectively after accounting for tree size and stand basal area effects. In addition, we recorded living and dead saplings to assess inter-specific dissimilarities after the drought events. Drought events strongly reduced the growth of both adults and saplings (40% on average), but they were more resilient to the 2005 than to the 2012 drought. Increased magnitude and frequency of drought events led to inter-specific differences in drought responses, which were revealed in saplings rather than adults after the 2012 drought. P. pinaster saplings were more resilient than P. nigra and P. sylvestris, consistently to the drought stress experienced by each species in the dry-edge of their natural ranges. Moreover, a greater recruitment and similar sapling mortality confirm P. pinaster as the most successful pine species from a demographic perspective. In addition, tree size reduced resilience whereas basal area in the tree neighborhood did not show any significant effect on pine tolerance to drought. In conclusion, we showed that increased the magnitude and frequency of drought events uncovers species-specific differences in drought resilience in saplings rather than adults. Our results suggest potential shifts in the dominance of P. pinaster as the expense of P. nigra and P. sylvestris under climate change in this relict forest. This highlights the need to include juveniles and demographic rates in models aiming to describe and predict species responses to climate change in dry-edge forests.

Physiological drought responses improve predictions of live fuel moisture dynamics in a Mediterranean forest

The moisture content of live fuels is an important determinant of forest flammability. Current approaches for modelling live fuel moisture content typically focus on the use of drought indices. However, these have mixed success partly because of species-specific differences in drought responses. Here we seek to understand the physiological mechanisms driving changes in live fuel moisture content, and to investigate the potential for incorporating plant physiological traits into live fuel moisture models. We measured the dynamics of leaf moisture content, access to water resources (through stable isotope analyses) and physiological traits (including leaf water potential, stomatal conductance, and cellular osmotic and elastic adjustments) across a fire season in a Mediterranean mixed forest in Catalonia, NE Spain. We found that differences in both seasonal variation and minimum values of live fuel moisture content were a function of access to water resources and plant physiological traits. Specifically, those species with the lowest minimum moisture content and largest seasonal variation in moisture (Cistus albidus: 49–137% and Rosmarinus officinalis: 47–144%) were most reliant on shallow soil water and had the lowest values of predawn leaf water potential. Species with the smallest variation in live fuel moisture content (Pinus nigra: 96–116% and Quercus ilex: 56–91%) exhibited isohydric behaviour (little variation in midday leaf water potential, and relatively tight regulation of stomata in response to soil drying). Of the traits measured, predawn leaf water potential provided the strongest predictor of live fuel moisture content (R2 = 0.63, AIC = 249), outperforming two commonly used drought indices (both with R2 = 0.49, AIC = 258). This is the first study to explicitly link fuel moisture with plant physiology and our findings demonstrate the potential and importance of incorporating ecophysiological plant traits to investigating seasonal changes in fuel moisture and, more broadly, forest flammability.

Plant water use responses along secondary forest succession during the 2015–2016 El Niño drought in Panama

Tropical forests are increasingly being subjected to hotter, drier conditions as a result of global climate change. The effects of drought on forests along successional gradients remain poorly understood. We took advantage of the 2015-2016 El Niño event to test for differences in drought response along a successional gradient by measuring the sap flow in 76 trees, representing 42 different species, in 8-, 25- and 80-yr-old secondary forests in the 15-km2 'Agua Salud Project' study area, located in central Panama. Average sap velocities and sapwood-specific hydraulic conductivities were highest in the youngest forest. During the dry season drought, sap velocities increased significantly in the 80-yr-old forest as a result of higher evaporative demand, but not in younger forests. The main drivers of transpiration shifted from radiation to vapor pressure deficit with progressing forest succession. Soil volumetric water content was a limiting factor only in the youngest forest during the dry season, probably as a result of less root exploration in the soil. Trees in early-successional forests displayed stronger signs of regulatory responses to the 2015-2016 El Niño drought, and the limiting physiological processes for transpiration shifted from operating at the plant-soil interface to the plant-atmosphere interface with progressing forest succession.

Genotypic differences in red clover (Trifolium pratense L.) response under severe water deficit

AimsIn Ontario, Canada, acreage of red clover (Trifolium pratense L.) intercropped with winter wheat (Triticum aestivum L. em. Thell) has declined, despite well-documented soil and yield benefits. The decline has resulted from increasing prevalence of stand non-uniformity, which has been attributed in part to soil moisture deficits. We examined whether there are genotypic differences in drought response between red clover varieties.MethodsA double-cut (Belle) and a single-cut variety (Altaswede) were grown under four different durations of drought (4, 8, 12 and 16 days below 15% relative soil water content, RSWC). Shoot dry weight, shoot relative water content (RWC), leaf area and crown water content were measured in control, drought and drought + recovery treatments.ResultsBelle used significantly more water during soil moisture deficit and had greater leaf area, shoot dry weight and RWC compared to Altaswede. In contrast, Altaswede had significantly higher survival rates than Belle, attributed to maintenance of meristematic tissue viability in the crown where re-growth, after shoot tissue desiccation, can occur.ConclusionsBy demonstrating genotypic variation in survival strategies of red clover, traits can be identified for the development of improved varieties. Varieties with higher survival rates during drought will result in more uniform stands and increased utilization of red clover for environmental and yield benefits.

Physiological Response of Garry Oak (Quercus garryana) Seedlings to Drought

The purpose of this study was to determine physiological differences in drought response among Garry oak (Quercus garryana Douglas ex Hook. [Fagaceae]) seedlings from acorns collected at seven acorn collection sites in Washington state. Our objective was to aid in Garry oak restoration efforts by investigating potential differences in drought tolerance for seedlings grown from acorns collected at different sites. Acorns were collected from six sites east of the Cascades, and one site on Whidbey Island (Oak Harbor) west of the Cascades. Differences in morphological and photosynthetic characteristics observed for seedlings grown from acorns collected at these different sites became more pronounced after drought stress was induced by withholding water. Seedlings from acorns collected at the northernmost east-side site (Swauk Creek) were most susceptible to drought and had photosynthetic rates that were 35% lower under drought conditions than when well-watered. Seedlings from acorns collected at the site west of the cascades (Oak Harbor) were the least susceptible to drought and had assimilation rates under drought conditions that were similar to those when well-watered. Differences were also observed among seedlings from different sites in the occurrence of drought-induced loss of stem hydraulic conductivity and changes in leaf water potential and relative water content in response to drought. We conclude that seedlings grown from Garry oak trees at different sites across Washington State have distinct physiological responses to drought that may lead to differing survivorship when used in reforestation efforts and exposed to drought stress in the field.

Genome-wide transcriptional and physiological responses to drought stress in leaves and roots of two willow genotypes.

BackgroundDrought is a major environmental stress that can have severe impacts on plant productivity and survival. Understanding molecular mechanisms of drought responses is crucial in order to breed for drought adapted plant cultivars. The aim of the present study was to investigate phenotypic and transcriptional drought responses in two willow genotypes (520 and 592) originating from an experimental cross between S. viminalis × (S. viminalis × S. schwerinii). Willows are woody perennials in the Salicaceae plant family that are grown as bioenergy crops worldwide.MethodsAn experiment was conducted where plants were exposed to drought and different eco-physiological parameters were assessed. RNA-seq data was furthermore generated with the Illumina technology from root tips and leaves from plants grown in drought and well-watered (WW) conditions. The RNA-seq data was assembled de novo with the Trinity assembler to create a reference gene set to which the reads were mapped in order to obtain differentially expressed genes (DEGs) between the drought and WW conditions. To investigate molecular mechanisms involved in the drought response, GO enrichment analyses were conducted. Candidate genes with a putative function in the drought response were also identified.ResultsA total of 52,599 gene models were obtained and after filtering on gene expression (FPKM ≥ 1), 35,733 gene models remained, of which 24,421 contained open reading frames. A total of 5,112 unique DEGs were identified between drought and WW conditions, of which the majority were found in the root tips. Phenotypically, genotype 592 displayed less growth reduction in response to drought compared to genotype 520. At the transcriptional level, genotype 520 displayed a greater response in the leaves as more DEGs were found in genotype 520 compared to genotype 592. In contrast, the transcriptional responses in the root tips were rather similar between the two genotypes. A core set of candidate genes encoding proteins with a putative function in drought response was identified, for example MYBs and bZIPs as well as chlorophyll a/b binding proteins.DiscussionWe found substantial differences in drought responses between the genotypes, both at the phenotypic and transcriptional levels. In addition to the genotypic variation in several traits, we also found indications for genotypic variation in trait plasticity, which could play a role in drought adaptation. Furthermore, the two genotypes displayed overall similar transcriptional responses in the root tips, but more variation in the leaves. It is thus possible that the observed phenotypic differences could be a result of transcriptional differences mostly at the leaf level.ConclusionsThis study has contributed to a better general understanding of drought responses in woody plants, specifically in willows, and has implications for breeding research towards more drought adapted plants.Electronic supplementary materialThe online version of this article (doi:10.1186/s12870-015-0630-2) contains supplementary material, which is available to authorized users.

Creeping Bentgrass Responses to Drought Stress and Polyamine Application

Polyamines (PAs) such as spermidine (Spd), spermine (Spm), and putrescine are involved in various biological functions including abiotic stress response. Whether PAs play an important role in cool-season turfgrass tolerance of drought stress is not well investigated. We have conducted a series of growth chamber (GC) studies including one hydroponic and two soil-based GC studies with creeping bentgrass (Agrostis stolonifera) ‘Penncross’ and ‘Penn-G2’ to determine whether exogenous application of PAs may affect plant growth and stress tolerance. Application of relatively low concentrations of Spd (500 or 750 μM) or Spm (500 μM) promoted tillering rates under optimal growth conditions in hydroponics. The same levels of PA treatments moderated the damages associated with drought stress in the soil-based GC studies. The most notable differences in drought response associated with PA treatment were increased membrane health. This was observed as greater photochemical efficiency, higher quantum yield, less electrolyte leakage, and less lipid peroxidation (malondialdehyde content) in PA-treated plants compared with control plants. The relatively low level of exogenous PAs used in this study did not have a major effect on plant water relations under drought stress. Canopy temperatures and soil moisture content were unaffected by any PA treatment; however, on some days during early drought stress, relative water content was significantly higher in PA-treated plants compared with controls. PA could play a major role in protecting photosynthetic and cellular membranes during drought stress of creeping bentgrass.

Short-term water stress impacts on stomatal, mesophyll and biochemical limitations to photosynthesis differ consistently among tree species from contrasting climates.

Predicting the large-scale consequences of drought in contrasting environments requires that we understand how drought effects differ among species originating from those environments. A previous meta-analysis of published experiments suggested that the effects of drought on both stomatal and non-stomatal limitations to photosynthesis may vary consistently among species from different hydroclimates. Here, we explicitly tested this hypothesis with two short-term water stress experiments on congeneric mesic and xeric species. One experiment was run in Australia using Eucalyptus species and the second was run in Spain using Quercus species as well as two more mesic species. In each experiment, plants were grown under moist conditions in a glasshouse, then deprived of water, and gas exchange was monitored. The stomatal response was analysed with a recently developed stomatal model, whose single parameter g1 represents the slope of the relationship between stomatal conductance and photosynthesis. The non-stomatal response was partitioned into effects on mesophyll conductance (gm), the maximum Rubisco activity (Vcmax) and the maximum electron transport rate (Jmax). We found consistency among the drought responses of g1, gm, Vcmax and Jmax, suggesting that drought imposes limitations on Rubisco activity and RuBP regeneration capacity concurrently with declines in stomatal and mesophyll conductance. Within each experiment, the more xeric species showed relatively high g1 under moist conditions, low drought sensitivity of g1, gm, Vcmax and Jmax, and more negative values of the critical pre-dawn water potential at which Vcmax declines most steeply, compared with the more mesic species. These results indicate adaptive interspecific differences in drought responses that allow xeric tree species to continue transpiration and photosynthesis for longer during periods without rain.

Physiological and biochemical responses to severe drought stress of nine Eucalyptus globulus clones: a multivariate approach.

Seasonal drought, typical of temperate and Mediterranean environments, creates problems in establishing plantations and affects development and yield, and it has been widely studied in numerous species. Forestry fast-growing species such as Eucalyptus spp. are an important resource in such environments, selected clones being generally used for production purposes in plantations in these areas. However, use of mono-specific plantations increases risk of plant loss due to abiotic stresses, making it essential to understand differences in an individual clone's physiological responses to drought stress. In order to study clonal differences in drought responses, nine Eucalyptus globulus (Labill.) clones (C14, C46, C97, C120, C222, C371, C405, C491 and C601) were gradually subjected to severe drought stress (<14% of field capacity). A total of 31 parameters, physiological (e.g., photosynthesis, gas exchange), biochemical (e.g., chlorophyll content) and hormonal (abscisic acid [ABA] content), were analysed by classic and multivariate techniques. Relationships between parameters were established, allowing related measurements to be grouped into functional units (pigment, growth, water and ABA). Differences in these units showed that there were two distinct groups of E. globulus clones on the basis of their different strategies when faced with drought stress. The C14 group (C14, C120, C405, C491 and C601) clones behave as water savers, maintaining high water content and showing high stomatal adjustment, and reducing their aerial growth to a great extent. The C46 group (C46, C97, C222 and C371) clones behave as water spenders, reducing their water content drastically and presenting osmotic adjustment. The latter maintains the highest growth rate under the conditions tested. The method presented here can be used to identify appropriate E. globulus clones for drought environments, facilitating the selection of material for production and repopulation environments.

A Gene-Phenotype Network Based on Genetic Variability for Drought Responses Reveals Key Physiological Processes in Controlled and Natural Environments

Identifying the connections between molecular and physiological processes underlying the diversity of drought stress responses in plants is key for basic and applied science. Drought stress response involves a large number of molecular pathways and subsequent physiological processes. Therefore, it constitutes an archetypical systems biology model. We first inferred a gene-phenotype network exploiting differences in drought responses of eight sunflower (Helianthus annuus) genotypes to two drought stress scenarios. Large transcriptomic data were obtained with the sunflower Affymetrix microarray, comprising 32423 probesets, and were associated to nine morpho-physiological traits (integrated transpired water, leaf transpiration rate, osmotic potential, relative water content, leaf mass per area, carbon isotope discrimination, plant height, number of leaves and collar diameter) using sPLS regression. Overall, we could associate the expression patterns of 1263 probesets to six phenotypic traits and identify if correlations were due to treatment, genotype and/or their interaction. We also identified genes whose expression is affected at moderate and/or intense drought stress together with genes whose expression variation could explain phenotypic and drought tolerance variability among our genetic material. We then used the network model to study phenotypic changes in less tractable agronomical conditions, i.e. sunflower hybrids subjected to different watering regimes in field trials. Mapping this new dataset in the gene-phenotype network allowed us to identify genes whose expression was robustly affected by water deprivation in both controlled and field conditions. The enrichment in genes correlated to relative water content and osmotic potential provides evidence of the importance of these traits in agronomical conditions.

Water uptake dynamics under progressive drought stress in diverse accessions of the OryzaSNP panel of rice (Oryza sativa).

In addition to characterising root architecture, evaluating root water uptake ability is important for understanding drought response. A series of three lysimeter studies were conducted using the OryzaSNP panel, which consists of 20 diverse rice (Oryza sativa L.) genotypes. Large genotypic differences in drought response were observed in this genotype panel in terms of plant growth and water uptake. Total water uptake and daily water uptake rates in the drought-stress treatment were correlated with root length density, especially at depths below 30cm. Patterns of water uptake among genotypes remained consistent throughout the stress treatments: genotypes that initially extracted more water were the same genotypes that extracted more water at the end of the study. These results suggest that response to drought by deep root growth, rather than a conservative soil water pattern, seems to be important for lowland rice. Genotypes in the O. sativa type aus group showed some of the greatest water uptake and root growth values. Since the OryzaSNP panel has been genotyped in detail with SNP markers, we expect that these results will be useful for understanding the genetics of rice root growth and function for water uptake in response to drought.

Mortality of Australian alpine grasses (Poa spp.) after drought: species differences and ecological patterns

Aims Australian alpine ecosystems currently experience high precipitation in the snow-free season, but they are predicted to experience drier conditions under climate change. We observed high mortality of the dominant alpine grasses following drought in 2007. Our aims were as follows: to test the involvement of plant-available water (PAW) and other environmental variables in grass mortality in the field; to detect possible species differences in drought response and to link soil moisture to precipitation using soil properties and climate data. Methods The dominant tussock grasses of the Australian alpine zone, Poa hothamensis var. hothamensis N.G. Walsh, Poa hiemata Vickery and Poa phillipsiana Vickery (Poaceae), all exhibited mortality following drought in the Bogong High Plains, Victoria, Australia in 2007. PAW was calculated using soil water potential measurements, and past drought occurrence was modelled using climate data. We then tested the effects of PAW and soil depth on grass survival both at a large spatial scale spanning the elevational range of the alpine zone and at a smaller scale. Poa hothamensis and P. phillipsiana were compared in a common-garden experiment to test drought tolerance. Important Findings Poa hothamensis survival was predicted by dry-season PAW at the small spatial scale; at the large scale, soil depth and elevation were more important predictors of P. hothamensis survival, but dry-season PAW predicted P. hiemata survival. Common-garden experiments supported field observations that P. hothamensis is more droughtsensitive than is P. phillipsiana. We also present a simple polynomial relationship between rainfall and field soil moisture, which predicts that the alpine soils dry below wilting point several times a year. We suggest the timing of long rain-free periods may be more important than their duration.