RESUMO
This study aimed to reconstruct the environmental conditions and the crop management practices and plant characteristics when agriculture appeared in western Europe. We analyzed oak charcoal and a large number of cereal caryopsides recovered from La Draga (Girona, Spain), an early (5300 to 4800 cal. BC) agricultural site from the Iberian Peninsula. The carbon isotope discrimination (Δ13C) values of oak, the dominant forest species in the region, indicates prevalence of a wet climate at the site. Further, we reconstructed crop management conditions, achievable yield, and crop characteristics through the analysis of Δ13C, nitrogen isotope composition (δ15N), nitrogen content, and the reconstructed weight of wheat and barley caryopsides, following protocols developed by our team [Araus et al., Nat. Commun. 5, 3953 (2014)] and comparison of these parameters with present-day organic agriculture in the region. In parallel, a regional perspective was achieved through the study of wheat and barley grains of seventeen Neolithic sites from the western Mediterranean. The results suggest that rather than small-garden cultivation, a more extensive agriculture was practiced under good water availability and moderate manuring. Moreover, results from La Draga evidence that grain weight and spike morphology were comparable to contemporary cereals. Growing conditions and the prevalence of improved crop traits indicate that agriculture was fairly consolidated at the time it reached the western edge of Europe.
Assuntos
Agricultura , Isótopos de Carbono , Hordeum , Isótopos de Nitrogênio , Triticum , Isótopos de Carbono/análise , Agricultura/métodos , Isótopos de Nitrogênio/análise , Produtos Agrícolas/crescimento & desenvolvimento , Europa (Continente) , Quercus , Espanha , Grão Comestível , História AntigaRESUMO
Although the straightening capacity of the stem is key for light capture and mechanical stability in forest trees, little is known about its adaptive implications. Assuming that stem straightening is costly, trade-offs are expected with competing processes such as growth, maintenance, and defence. We established a manipulative experiment in a common garden of Pinus pinaster including provenances typically showing either straight-stemmed or crooked-stemmed phenotypes. We imposed a bending up to 35º on plants aged 9 years of both provenance groups and followed the straightening kinetics and shoot elongation after releasing. Eight months later, we destructively assessed biomass partitioning, reaction wood, wood microdensity, xylem reserve carbohydrates, and phloem secondary metabolites. The experimental bending and release caused significant, complex changes with a marked difference between straight- and crooked-type plants. The straight-type recovered verticality faster and to a higher degree and developed more compression wood, while displaying a transitory delay in shoot elongation, reducing resource allocation to defence and maintaining the levels of non-structural carbohydrates compared with the crooked type. This combination of responses indicates the existence of intraspecific divergence in the reaction to mechanical stresses that may be related to different adaptive phenotypic plasticity.
Assuntos
Pinus , Pinus/crescimento & desenvolvimento , Árvores/crescimento & desenvolvimento , MadeiraRESUMO
BACKGROUND AND AIMS: Understanding the genetic basis of adaptation and plasticity in trees constitutes a knowledge gap. We linked dendrochronology and genomics [single nucleotide polymorphisms (SNPs)] for a widespread conifer (Pinus halepensis Mill.) to characterize intraspecific growth differences elicited by climate. METHODS: The analysis comprised 20-year tree-ring series of 130 trees structured in 23 populations evaluated in a common garden. We tested for genotype by environment interactions (Gâ ×â E) of indexed ring width (RWI) and early- to latewood ratios (ELI) using factorial regression, which describes Gâ ×â E as differential gene sensitivity to climate. KEY RESULTS: The species' annual growth was positively influenced by winter temperature and spring moisture and negatively influenced by previous autumn precipitation and warm springs. Four and five climate factors explained 10 % (RWI) and 16 % (ELI) of population-specific interannual variability, respectively, with populations from drought-prone areas and with uneven precipitation experiencing larger growth reductions during dry vegetative periods. Furthermore, four and two SNPs explained 14 % (RWI) and 10 % (ELI) of interannual variability among trees, respectively. Two SNPs played a putative role in adaptation to climate: one identified from transcriptome sequencing of P. halepensis and another involved in response regulation to environmental stressors. CONCLUSIONS: We highlight how tree-ring phenotypes, obtained from a common garden experiment, combined with a candidate-gene approach allow the quantification of genetic and environmental effects determining adaptation for a conifer with a large and complex genome.
Assuntos
Pinus , Árvores , Clima , Secas , Interação Gene-Ambiente , Fenótipo , Pinus/fisiologiaRESUMO
Progress in high-throughput phenotyping and genomics provides the potential to understand the genetic basis of plant functional differentiation. We developed a semi-automatic methodology based on unmanned aerial vehicle (UAV) imagery for deriving tree-level phenotypes followed by genome-wide association study (GWAS). An RGB-based point cloud was used for tree crown identification in a common garden of Pinus halepensis in Spain. Crowns were combined with multispectral and thermal orthomosaics to retrieve growth traits, vegetation indices and canopy temperature. Thereafter, GWAS was performed to analyse the association between phenotypes and genomic variation at 235 single nucleotide polymorphisms (SNPs). Growth traits were associated with 12 SNPs involved in cellulose and carbohydrate metabolism. Indices related to transpiration and leaf water content were associated with six SNPs involved in stomata dynamics. Indices related to leaf pigments and leaf area were associated with 11 SNPs involved in signalling and peroxisome metabolism. About 16-20% of trait variance was explained by combinations of several SNPs, indicating polygenic control of morpho-physiological traits. Despite a limited availability of markers and individuals, this study is provides a successful proof-of-concept for the combination of high-throughput UAV-based phenotyping with cost-effective genotyping to disentangle the genetic architecture of phenotypic variation in a widespread conifer.
Assuntos
Estudo de Associação Genômica Ampla , Pinus , Genótipo , Fenótipo , Pinus/genética , EspanhaRESUMO
PREMISE: Persistence of tree populations in the face of global change relies on their capacity to respond to biotic and abiotic stressors through plastic or adaptive changes. Genetic adaptation will depend on the additive genetic variation within populations and the heritability of traits related to stress tolerance. Because traits can be genetically linked, selective pressure acting on one trait may lead to correlated responses in other traits. METHODS: To test direct and correlated responses to selection for growth and drought tolerance in Pinus halepensis, we selected trees in a parental population for higher growth and greater water-use efficiency (WUE) and compared their offspring with the offspring of random trees from the parental population in two contrasting common gardens. We estimated direct responses to selection for growth and WUE and correlated responses for growth and tolerance to abiotic and biotic stressors. RESULTS: We found a strong response to selection and high realized heritability for WUE, but no response to selection for growth. Correlated responses to selection in other life-history traits were not significant, except for concentration of some chemical defenses, which was greater in the offspring of mother trees selected for growth than in the offspring of unselected control trees. CONCLUSIONS: The empirical evidence of direct responses to selection for high WUE suggests that P. halepensis has the potential to evolve in response to increasing drought stress. Contrary to expectations, the results are not conclusive of a potential negative impact of WUE and growth selection on other key life-history traits.
Assuntos
Pinus , Água , Secas , Fenótipo , Pinus/genética , Árvores/genéticaRESUMO
Many ecologically important forest trees from dry areas have been insufficiently investigated for their ability to adapt to the challenges posed by climate change, which hampers the implementation of mitigation policies. We analyzed 14 common-garden experiments across the Mediterranean which studied the widespread thermophilic conifer Pinus halepensis and involved 157 populations categorized into five ecotypes. Ecotype-specific tree height responses to climate were applied to projected climate change (2071-2100 ad), to project potential growth patterns both locally and across the species' range. We found contrasting ecotypic sensitivities to annual precipitation but comparatively uniform responses to mean temperature, while evidence of local adaptation for tree height was limited to mesic ecotypes. We projected intriguing patterns of response range-wide, implying either height inhibition or stimulation of up to 75%, and deduced that the ecotype currently experiencing more favorable (wetter) conditions will show the largest inhibition. Extensive height reductions can be expected for coastal areas of France, Greece, Spain and northern Africa. Our findings underline the fact that intraspecific variations in sensitivity to precipitation must be considered when projecting tree height responses of dry forests to future climate. The ecotype-specific projected performances call for management activities to ensure forest resilience in the Mediterranean through, for example, tailored deployment strategies.
Assuntos
Pinus , Traqueófitas , Mudança Climática , Florestas , França , Espanha , ÁrvoresRESUMO
Resprouting is an ancestral trait in angiosperms that confers resilience after perturbations. As climate change increases stress, resprouting vigor is declining in many forest regions, but the underlying mechanism is poorly understood. Resprouting in woody plants is thought to be primarily limited by the availability of non-structural carbohydrate reserves (NSC), but hydraulic limitations could also be important. We conducted a multifactorial experiment with two levels of light (ambient, 2-3% of ambient) and three levels of water stress (0, 50 and 80 percent losses of hydraulic conductivity, PLC) on two Mediterranean oaks (Quercus ilex and Q. faginea) under a rain-out shelter (n = 360). The proportion of resprouting individuals after canopy clipping declined markedly as PLC increased for both species. NSC concentrations affected the response of Q. ilex, the species with higher leaf construction costs, and its effect depended on the PLC. The growth of resprouting individuals was largely dependent on photosynthetic rates for both species, while stored NSC availability and hydraulic limitations played minor and non-significant roles, respectively. Contrary to conventional wisdom, our results indicate that resprouting in oaks may be primarily driven by complex interactions between hydraulics and carbon sources, whereas stored NSC play a significant but secondary role.
Assuntos
Metabolismo dos Carboidratos/fisiologia , Raízes de Plantas/metabolismo , Quercus/fisiologia , Desidratação , Fotossíntese/fisiologia , Folhas de Planta/metabolismo , Quercus/crescimento & desenvolvimento , EspanhaRESUMO
BACKGROUND AND AIMS: Understanding inter-population variation in the allocation of resources to specific anatomical compartments and physiological processes is crucial to disentangle adaptive patterns in forest species. This work aims to evaluate phenotypic integration and trade-offs among functional traits as determinants of life history strategies in populations of a circum-Mediterranean pine that dwells in environments where water and other resources are in limited supply. METHODS: Adult individuals of 51 populations of Pinus halepensis grown in a common garden were characterized for 11 phenotypic traits, including direct and indirect measures of water uptake at different depths, leaf area, stomatal conductance, chlorophyll content, non-structural carbohydrates, stem diameter and tree height, age at first reproduction and cone production. The population differentiation in these traits was tested through analysis of variance (ANOVA). The resulting populations' means were carried forward to a structural equation model evaluating phenotypic integration between six latent variables (summer water uptake depth, summer transpiration, spring photosynthetic capacity, growth, reserve accumulation and reproduction). KEY RESULTS: Water uptake depth and transpiration covaried negatively among populations, as the likely result of a common selective pressure for drought resistance, while spring photosynthetic capacity was lower in populations originating from dry areas. Transpiration positively influenced growth, while growth was negatively related to reproduction and reserves among populations. Water uptake depth negatively influenced reproduction. CONCLUSIONS: The observed patterns indicate a differentiation in life cycle features between fast-growing and slow-growing populations, with the latter investing significantly more in reproduction and reserves. We speculate that such contrasting strategies result from different arrays of life history traits underlying the very different ecological conditions that the Aleppo pine must face across its distribution range. These comprise, principally, drought as the main stressor and fire as the main ecological disturbance of the Mediterranean basin.
Assuntos
Características de História de Vida , Pinus , Secas , Análise de Classes Latentes , FenótipoRESUMO
Forests play a key role in the carbon balance of terrestrial ecosystems. One of the main uncertainties in global change predictions lies in how the spatiotemporal dynamics of forest productivity will be affected by climate warming. Here we show an increasing influence of climate on the spatial variability of tree growth during the last 120 y, ultimately leading to unprecedented temporal coherence in ring-width records over wide geographical scales (spatial synchrony). Synchrony in growth patterns across cold-constrained (central Siberia) and drought-constrained (Spain) Eurasian conifer forests have peaked in the early 21st century at subcontinental scales (â¼ 1,000 km). Such enhanced synchrony is similar to that observed in trees co-occurring within a stand. In boreal forests, the combined effects of recent warming and increasing intensity of climate extremes are enhancing synchrony through an earlier start of wood formation and a stronger impact of year-to-year fluctuations of growing-season temperatures on growth. In Mediterranean forests, the impact of warming on synchrony is related mainly to an advanced onset of growth and the strengthening of drought-induced growth limitations. Spatial patterns of enhanced synchrony represent early warning signals of climate change impacts on forest ecosystems at subcontinental scales.
Assuntos
Mudança Climática , Florestas , Árvores/crescimento & desenvolvimento , Modelos Lineares , Sibéria , Espanha , Especificidade da Espécie , Fatores de TempoRESUMO
There is increasing evidence that the circadian clock is a significant driver of photosynthesis that becomes apparent when environmental cues are experimentally held constant. We studied whether the composition of photosynthetic pigments is under circadian regulation, and whether pigment oscillations lead to rhythmic changes in photochemical efficiency. To address these questions, we maintained canopies of bean and cotton, after an entrainment phase, under constant (light or darkness) conditions for 30-48 h. Photosynthesis and quantum yield peaked at subjective noon, and non-photochemical quenching peaked at night. These oscillations were not associated with parallel changes in carbohydrate content or xanthophyll cycle activity. We observed robust oscillations of Chl a/b during constant light in both species, and also under constant darkness in bean, peaking when it would have been night during the entrainment (subjective nights). These oscillations could be attributed to the synthesis and/or degradation of trimeric light-harvesting complex II (reflected by the rhythmic changes in Chl a/b), with the antenna size minimal at night and maximal around subjective noon. Considering together the oscillations of pigments and photochemistry, the observed pattern of changes is counterintuitive if we assume that the plant strategy is to avoid photodamage, but consistent with a strategy where non-stressed plants maximize photosynthesis.
Assuntos
Ritmo Circadiano/fisiologia , Gossypium/fisiologia , Phaseolus/fisiologia , Fotossíntese/fisiologia , Pigmentos Biológicos/metabolismo , Metabolismo dos Carboidratos , Clorofila/metabolismo , Clorofila A , Folhas de Planta/metabolismoRESUMO
The relevance of interspecific variation in the use of plant water sources has been recognized in drought-prone environments. By contrast, the characterization of intraspecific differences in water uptake patterns remains elusive, although preferential access to particular soil layers may be an important adaptive response for species along aridity gradients. Stable water isotopes were analysed in soil and xylem samples of 56 populations of the drought-avoidant conifer Pinus halepensis grown in a common garden test. We found that most populations reverted to deep soil layers as the main plant water source during seasonal summer droughts. More specifically, we detected a clear geographical differentiation among populations in water uptake patterns even under relatively mild drought conditions (early autumn), with populations originating from more arid regions taking up more water from deep soil layers. However, the preferential access to deep soil water was largely independent of aboveground growth. Our findings highlight the high plasticity and adaptive relevance of the differential access to soil water pools among Aleppo pine populations. The observed ecotypic patterns point to the adaptive relevance of resource investment in deep roots as a strategy towards securing a source of water in dry environments for P. halepensis.
Assuntos
Adaptação Fisiológica/genética , Secas , Ecótipo , Variação Genética , Pinus/genética , Estresse Fisiológico , Água/metabolismo , Região do Mediterrâneo , Pinus/fisiologia , Estações do Ano , Solo/química , Xilema/metabolismoRESUMO
Stable isotopes are extensively used as tracers for the study of plant-water sources. Isotope-ratio infrared spectroscopy (IRIS) offers a cheaper alternative to isotope-ratio mass spectroscopy (IRMS), but its use in studying plant and soil water is limited by the spectral interference caused by organic contaminants. Here, we examine two approaches to cope with contaminated samples in IRIS: on-line oxidation of organic compounds (MCM) and post-processing correction. We assessed these methods compared to IRMS across 136 samples of xylem and soil water, and a set of ethanol- and methanol-water mixtures. A post-processing correction significantly improved IRIS accuracy in both natural samples and alcohol dilutions, being effective with concentrations up to 8% of ethanol and 0.4% of methanol. MCM outperformed the post-processing correction in removing methanol interference, but did not effectively remove interference for high concentrations of ethanol. By using both approaches, IRIS can overcome with reasonable accuracy the analytical uncertainties associated with most organic contaminants found in soil and xylem water. We recommend the post-processing correction as the first choice for analysis of samples of unknown contamination. Nevertheless, MCM can be more effective for evaluating samples containing contaminants responsible for strong spectral interferences at low concentrations, such as methanol.
Assuntos
Plantas/química , Espectrofotometria Infravermelho/métodos , Água/química , Deutério , Etanol/química , Funções Verossimilhança , Metanol/química , Compostos Orgânicos/isolamento & purificação , Isótopos de Oxigênio , Poluentes Químicos da Água/isolamento & purificaçãoRESUMO
Vegetation in water-limited ecosystems relies strongly on access to deep water reserves to withstand dry periods. Most of these ecosystems have shallow soils over deep groundwater reserves. Understanding the functioning and functional plasticity of species-specific root systems and the patterns of or differences in the use of water sources under more frequent or intense droughts is therefore necessary to properly predict the responses of seasonally dry ecosystems to future climate. We used stable isotopes to investigate the seasonal patterns of water uptake by a sclerophyll forest on sloped terrain with shallow soils. We assessed the effect of a long-term experimental drought (12 years) and the added impact of an extreme natural drought that produced widespread tree mortality and crown defoliation. The dominant species, Quercus ilex, Arbutus unedo and Phillyrea latifolia, all have dimorphic root systems enabling them to access different water sources in space and time. The plants extracted water mainly from the soil in the cold and wet seasons but increased their use of groundwater during the summer drought. Interestingly, the plants subjected to the long-term experimental drought shifted water uptake toward deeper (10-35 cm) soil layers during the wet season and reduced groundwater uptake in summer, indicating plasticity in the functional distribution of fine roots that dampened the effect of our experimental drought over the long term. An extreme drought in 2011, however, further reduced the contribution of deep soil layers and groundwater to transpiration, which resulted in greater crown defoliation in the drought-affected plants. This study suggests that extreme droughts aggravate moderate but persistent drier conditions (simulated by our manipulation) and may lead to the depletion of water from groundwater reservoirs and weathered bedrock, threatening the preservation of these Mediterranean ecosystems in their current structures and compositions.
Assuntos
Secas , Árvores/fisiologia , Água/metabolismo , Hidrologia , Longevidade , Estações do Ano , Espanha , Especificidade da Espécie , Árvores/crescimento & desenvolvimentoRESUMO
Forest dynamics will depend upon the physiological performance of individual tree species under more stressful conditions caused by climate change. In order to compare the idiosyncratic responses of Mediterranean tree species (Quercus faginea, Pinus nigra, Juniperus thurifera) coexisting in forests of central Spain, we evaluated the temporal changes in secondary growth (basal area increment; BAI) and intrinsic water-use efficiency (iWUE) during the last four decades, determined how coexisting species are responding to increases in atmospheric CO2 concentrations (C(a)) and drought stress, and assessed the relationship among iWUE and growth during climatically contrasting years. All species increased their iWUE (ca. +15 to +21%) between the 1970s and the 2000s. This increase was positively related to C(a) for J. thurifera and to higher C(a) and drought for Q. faginea and P. nigra. During climatically favourable years the study species either increased or maintained their growth at rising iWUE, suggesting a higher CO2 uptake. However, during unfavourable climatic years Q. faginea and especially P. nigra showed sharp declines in growth at enhanced iWUE, likely caused by a reduced stomatal conductance to save water under stressful dry conditions. In contrast, J. thurifera showed enhanced growth also during unfavourable years at increased iWUE, denoting a beneficial effect of C(a) even under climatically harsh conditions. Our results reveal significant inter-specific differences in growth driven by alternative physiological responses to increasing drought stress. Thus, forest composition in the Mediterranean region might be altered due to contrasting capacities of coexisting tree species to withstand increasingly stressful conditions.
Assuntos
Dióxido de Carbono/análise , Isótopos de Carbono/análise , Secas , Árvores/química , Árvores/crescimento & desenvolvimento , Mudança Climática , Juniperus/química , Juniperus/crescimento & desenvolvimento , Juniperus/fisiologia , Pinus/química , Pinus/crescimento & desenvolvimento , Pinus/fisiologia , Quercus/química , Quercus/crescimento & desenvolvimento , Quercus/fisiologia , Espanha , Árvores/fisiologia , ÁguaRESUMO
Increasing evidence indicates that tree growth processes, including reproduction, can be either sink- or source-limited, or simultaneously co-limited by sink and source, depending on the interplay between internal and environmental factors. We tested the hypothesis that the relative strengths of photosynthate supply and demand by stem growth and reproduction create variable competition for substrate that is imprinted in the tree-ring isotopes (C and O) of stone pine (Pinus pinea L.), a masting gymnosperm with large costs of reproduction, under warming-induced drought. Across five representative stands of the Spanish Northern Plateau, we also identified reproductive phases where weather drivers of cone yield (CY) have varied over a 60-year period (1960-2016). We found that these drivers gradually shifted from winter-spring conditions 3 years before seed rain (cone setting) to a combination of 3- and 1-year lagged effects (kernel filling). Additionally, we observed positive regional associations between carbon isotope discrimination (Δ13C) of the year of kernel filling and CY arising at the turn of this century, which progressively offset similarly positive relationships between Δ13C of the year of cone setting and CY found during the first half of the study period. Altogether, these results pinpoint the increasing dependence of reproduction on fresh assimilates and suggest sink and source co-limitation superseding the sink-limited functioning of reproduction dominant before 2000. Under climate warming, it could be expected that drier conditions reinforce the role of source limitation on reproduction and, hence, on regeneration, forest structure and economic profit of the nutlike seeds of the species.
Assuntos
Pinus , Árvores , Fatores de Tempo , Florestas , Isótopos de Carbono/análise , ReproduçãoRESUMO
Winter-drought induced forest diebacks in the low-latitude margins of species' distribution ranges can provide new insights into the mechanisms (carbon starvation, hydraulic failure) underlying contrasting tree reactions. We analysed a winter-drought induced dieback at the Scots pine's southern edge through a dual-isotope approach (Δ(13) C and δ(18) O in tree-ring cellulose). We hypothesized that a differential long-term performance, mediated by the interaction between CO(2) and climate, determined the fates of individuals during dieback. Declining trees showed a stronger coupling between climate, growth and intrinsic water-use efficiency (WUEi) than non-declining individuals that was noticeable for 25 years prior to dieback. The rising stomatal control of water losses with time in declining trees, indicated by negative Δ(13) C-δ(18) O relationships, was likely associated with their native aptitude to grow more and take up more water (suggested by larger tracheid lumen widths) than non-declining trees and, therefore, to exhibit a greater cavitation risk. Freeze-thaw episodes occurring in winter 2001 unveiled such physiological differences by triggering dieback in those trees more vulnerable to hydraulic failure. Thus, WUEi tightly modulated growth responses to long-term warming in declining trees, indicating that co-occurring individuals were differentially predisposed to winter-drought mortality. These different performances were unconnected to the depletion of stored carbohydrates.
Assuntos
Pinus sylvestris/crescimento & desenvolvimento , Água/metabolismo , Madeira/crescimento & desenvolvimento , Isótopos de Carbono/análise , Clima , Secas , Geografia , Isótopos de Oxigênio/análise , Pinus sylvestris/anatomia & histologia , Pinus sylvestris/fisiologia , Caules de Planta/anatomia & histologia , Caules de Planta/crescimento & desenvolvimento , Caules de Planta/fisiologia , Transpiração Vegetal , Chuva , Estudos Retrospectivos , Estações do Ano , Espanha , Temperatura , Fatores de Tempo , Árvores , Madeira/anatomia & histologia , Madeira/fisiologiaRESUMO
⢠Accurate phenotyping remains a bottleneck in breeding for salinity and drought resistance. Here the combined use of stable isotope compositions of carbon (δ¹³C), oxygen (δ¹8O) and nitrogen (δ¹5N) in dry matter is aimed at assessing genotypic responses of durum wheat under different combinations of these stresses. ⢠Two tolerant and two susceptible genotypes to salinity were grown under five combinations of salinity and irrigation regimes. Plant biomass, δ¹³C, δ¹8O and δ¹5N, gas-exchange parameters, ion and N concentrations, and nitrate reductase (NR) and glutamine synthetase (GS) activities were measured. ⢠Stresses significantly affected all traits studied. However, only δ¹³C, δ¹8O, δ¹5N, GS and NR activities, and N concentration allowed for clear differentiation between tolerant and susceptible genotypes. Further, a conceptual model explaining differences in biomass based on such traits was developed for each growing condition. ⢠Differences in acclimation responses among durum wheat genotypes under different stress treatments were associated with δ¹³C. However, except for the most severe stress, δ¹³C did not have a direct (negative) relationship to biomass, being mediated through factors affecting δ¹8O or N metabolism. Based upon these results, the key role of N metabolism in durum wheat adaptation to salinity and water stress is highlighted.
Assuntos
Adaptação Fisiológica/genética , Secas , Nitrogênio/metabolismo , Salinidade , Triticum/genética , Triticum/fisiologia , Água/metabolismo , Biomassa , Isótopos de Carbono , Análise por Conglomerados , Genótipo , Glutamato-Amônia Ligase/metabolismo , Íons , Modelos Lineares , Nitrato Redutase/metabolismo , Isótopos de Nitrogênio , Isótopos de Oxigênio , Fotossíntese , Brotos de Planta/crescimento & desenvolvimento , Característica Quantitativa Herdável , Triticum/enzimologia , Triticum/crescimento & desenvolvimentoRESUMO
The physiological performance of durum wheat and two related amphiploids was studied during the reproductive stage under different combinations of salinity and irrigation. One triticale, one tritordeum, and four durum wheat genotypes were grown in pots in the absence of stress until heading, when six different treatments were imposed progressively. Treatments resulted from the combination of two irrigation regimes (100% and 35% of container water capacity) with three levels of water salinity (1.8, 12, and 17 dS m(-1)), and were maintained for nearly 3 weeks. Gas exchange and chlorophyll fluorescence and content were measured prior to harvest; afterwards shoot biomass and height were recorded, and Delta(13)C, delta(15)N, and the concentration of nitrogen (N), phosphorus, and several ions (K(+), Na(+), Ca(2+), Mg(2+)) were analysed in shoot material. Compared with control conditions (full irrigation with Hoagland normal) all other treatments inhibited photosynthesis through stomatal closure, accelerated senescence, and decreased biomass. Full irrigation with 12 dS m(-1) outperformed other stress treatments in terms of biomass production and physiological performance. Biomass correlated positively with N and delta(15)N, and negatively with Na(+) across genotypes and fully irrigated treatments, while relationships across deficit irrigation conditions were weaker or absent. Delta(13)C did not correlate with biomass across treatments, but it was the best trait correlating with phenotypic differences in biomass within treatments. Tritordeum produced more biomass than durum wheat in all treatments. Its low Delta(13)C and high K(+)/Na(+) ratio, together with a high potential growth, may underlie this finding. Mechanisms relating delta(15)N and Delta(13)C to biomass are discussed.
Assuntos
Isótopos de Carbono/análise , Desidratação/metabolismo , Íons/análise , Isótopos de Nitrogênio/análise , Salinidade , Triticum/fisiologia , Água/metabolismo , Biomassa , Grão Comestível/química , Grão Comestível/genética , Grão Comestível/crescimento & desenvolvimento , Grão Comestível/fisiologia , Genótipo , Modelos Lineares , Fotossíntese/fisiologia , Brotos de Planta/química , Brotos de Planta/crescimento & desenvolvimento , Brotos de Planta/fisiologia , Poaceae/química , Poaceae/genética , Poaceae/crescimento & desenvolvimento , Poaceae/fisiologia , Triticum/química , Triticum/genética , Triticum/crescimento & desenvolvimentoRESUMO
Stable isotopes and their potential for detecting various and complex ecosystem processes are attracting an increasing number of scientists. Progress is challenging, particularly under global change scenarios, but some established views have been challenged. The IX meeting of the Spanish Association of Terrestrial Ecology (AAET, Ubeda, 18-22 October 2009) hosted a symposium on the ecology of stable isotopes where the linear mixing model approach of partitioning sinks and sources of carbon and water fluxes within an ecosystem was challenged, and new applications of stable isotopes for the study of plant interactions were evaluated. Discussion was also centred on the need for networks that monitor ecological processes using stable isotopes and key ideas for fostering future research with isotopes.
Assuntos
Ecossistema , Isótopos , Dióxido de Carbono , Ecologia , Isótopos de Oxigênio , Plantas/metabolismo , ÁguaRESUMO
Individual differences in the access to deep soil water pools may explain the differential damage among coexisting, conspecific trees as a consequence of drought-induced dieback. We addressed this issue by comparing the responses to a severe drought of three Mediterranean oak species with different drought tolerance, Quercus pubescens L. and Quercus frainetto Ten., mainly thriving at xeric and mesic sites, respectively, and Quercus cerris L., which dominates at intermediate sites. For each species, we compared coexisting declining (D) and non-declining (ND) trees. The stable isotope composition (δ2H, δ18O) of xylem and soil water was used to infer a differential use of soil water sources. We also measured tree size and radial growth to quantify the long-term divergence of wood production between D and ND trees and non-structural carbohydrates (NSCs) in sapwood to evaluate if D trees presented lower NSC values. The ND trees had access to deeper soil water than D trees except in Q. frainetto, as indicated by significantly more depleted xylem water values. However, a strong δ2H offset between soil and xylem water isotopes observed in peak summer could suggest that both tree types were not physiologically active under extreme drought conditions. Alternative processes causing deuterium fractionation, however, could not be ruled out. Tree height and recent (last 15-25 years) growth rates in all species studied were lower in D than in ND trees by 22 and 44%, respectively. Lastly, there was not a consistent pattern of NSC sapwood concentration; in Q. pubescens, it was higher in ND trees while in Q. frainetto, the D trees were the ones exhibiting the higher NSC concentration. We conclude that the vulnerability to drought among conspecific Mediterranean oaks depends on the differential access to deep soil water pools, which may be related to differences in rooting depth, tree size and growth rate.