中国东北典型森林生态系统植物叶片的非结构性碳水化合物研究

植物叶片的非结构性碳水化合物(NSC)不仅可以反应植物的碳供应状况,也能反应植物对外界环境的适应策略。利用传统的蒽酮比色法测定了东北3个典型森林生态系统(呼中、凉水和长白山)242种常见植物叶片的非结构碳水化合物,探讨了温带主要森林植物叶片NSC沿纬度梯度的变化趋势及其在物种-生活型-群落间的分布规律。实验结果表明:3个典型森林生态系统植物叶片可溶性糖、淀粉和NSC含量均呈偏正态分布,多数物种的含量偏中低水平;242种植物叶片可溶性糖、淀粉和NSC的平均含量分别为63.31、65.66和128.96 mg/g。在所调查的森林生态系统中,叶片可溶性糖、淀粉和NSC含量在不同生活型中表现各异。此外,乔木植物叶片的可溶性糖、淀粉和NSC含量从北到南呈递增趋势,呼中最低,凉水次之,长白山最高。乔木淀粉含量均表现为落叶树种大于常绿树种,可溶性糖和NSC含量变化趋势复杂。研究结果不仅为阐明东北主要森林生态系统植被碳代谢和生长适应对策提供数据基础,而且对理解植物对未来气候变化的响应机理提供数据支撑。     

Patterns in nonstructural carbohydrate contents at the tree organ level in response to drought duration

Nonstructural carbohydrates (NSCs) facilitate the adaptation of trees to drought stress. There have been a large number of studies exploring NSC changes in individual plant species and individual organ under drought and showed different trends; however, an understanding of the universal pattern of the plant NSCs responses to drought, particularly to drought duration, is still lacking. Here, we compiled data from 47 experimental studies on 52 tree species and conducted a meta‐analysis to evaluate the responses of soluble sugars, starch, and TNSC (total nonstructural carbohydrates including both soluble sugars and starch) concentrations in different tree organs (leaf, stem, and root) to drought intensity and duration. We found that starch in all organs decreased and soluble sugars in leaf increased with prolonged experiment time, and the changes in soluble sugars in all organs were stronger under severe drought than under slight‐to‐moderate drought. Under slight‐to‐moderate drought, the NSC content of each organ varied with time, while with the extension of the drought duration, the NSCs gradually approached the control value (no drought stress); this trend remained in the late drought, which means that trees activated physiological regulation processes to increase carbon storage and reduce the risks of carbon starvation. In contrast, long‐term severe drought could lead to a net loss of carbohydrates, especially in the root, implying that prolonged severe drought could lead to NSC depletion in the whole plant. As prolonged drought duration has occurred in and is projected for many regions, this paper could shed light into studies on how trees respond and adapt extending drought duration through nonstructural carbon production, transportation, and reallocation.     

Homoeostatic maintenance of nonstructural carbohydrates during the 2015–2016 El Niño drought across a tropical forest precipitation gradient

Nonstructural carbohydrates (NSCs) are essential for maintenance of plant metabolism and may be sensitive to short‐ and long‐term climatic variation. NSC variation in moist tropical forests has rarely been studied, so regulation of NSCs in these systems is poorly understood. We measured foliar and branch NSC content in 23 tree species at three sites located across a large precipitation gradient in Panama during the 2015–2016 El Niño to examine how short‐ and long‐term climatic variation impact carbohydrate dynamics. There was no significant difference in total NSCs as the drought progressed (leaf P = 0.32, branch P = 0.30) nor across the rainfall gradient (leaf P = 0.91, branch P = 0.96). Foliar soluble sugars decreased while starch increased over the duration of the dry period, suggesting greater partitioning of NSCs to storage than metabolism or transport as drought progressed. There was a large variation across species at all sites, but total foliar NSCs were positively correlated with leaf mass per area, whereas branch sugars were positively related to leaf temperature and negatively correlated with daily photosynthesis and wood density. The NSC homoeostasis across a wide range of conditions suggests that NSCs are an allocation priority in moist tropical forests.     

沿干旱梯度樟子松人工林针叶和枝条非结构性碳水化合物及氮含量的变化

选取沿内蒙古科尔沁沙地自然干旱梯度分布的6个樟子松人工林样点(辉南、西丰、付家、章古台、奈曼和乌兰敖都)为研究对象,测定当年生和1年生针叶和枝条中非结构性碳水化合物(NSCs)和氮(N)含量的变化,以探究干旱条件下樟子松的碳供需状态和养分贮存策略。结果表明: 随干旱加剧,樟子松针叶和枝条中的NSCs和可溶性糖含量显著降低。从最湿润样点(辉南)到最干旱样点(乌兰敖都),樟子松当年生和1年生针叶中可溶性糖含量分别由12.8%和12.5%下降到9.0%和9.5%,而当年生枝条中可溶性糖含量由15.6%下降到9.2%。随干旱加剧,樟子松针叶和枝条中淀粉含量变化不显著,当年生和1年生针叶可溶性糖和淀粉的比值降低,当年生和1年生枝条中N含量显著增加。科尔沁沙地樟子松在干旱条件下显著消耗可溶性糖存储,存在“碳饥饿”致死风险。樟子松倾向维持稳定的淀粉含量以及在枝条中积累N以应对长期的干旱胁迫。     

Embolism recovery strategies and nocturnal water loss across species influenced by biogeographic origin

Drought‐induced tree mortality is expected to increase in future climates with the potential for significant consequences to global carbon, water, and energy cycles. Xylem embolism can accumulate to lethal levels during drought, but species that can refill embolized xylem and recover hydraulic function may be able to avoid mortality. Yet the potential controls of embolism recovery, including cross‐biome patterns and plant traits such as nonstructural carbohydrates (NSCs), hydraulic traits, and nocturnal stomatal conductance, are unknown. We exposed eight plant species, originating from mesic (tropical and temperate) and semi‐arid environments, to drought under ambient and elevated CO₂ levels, and assessed recovery from embolism following rewatering. We found a positive association between xylem recovery and NSCs, and, surprisingly, a positive relationship between xylem recovery and nocturnal stomatal conductance. Arid‐zone species exhibited greater embolism recovery than mesic zone species. Our results indicate that nighttime stomatal conductance often assumed to be a wasteful use of water, may in fact be a key part of plant drought responses, and contribute to drought survival. Findings suggested distinct biome‐specific responses that partially depended on species climate‐of‐origin precipitation or aridity index, which allowed some species to recover from xylem embolism. These findings provide improved understanding required to predict the response of diverse plant communities to drought. Our results provide a framework for predicting future vegetation shifts in response to climate change.     

木本植物非结构性碳水化合物变化及其影响因子研究进展

非结构性碳水化合物(NSC)是植物新陈代谢过程中重要的供能物质,NSC库的动态变化不仅反映了植物体内的碳收支状况,还影响着植物的生理代谢活动.为了预测树木甚至整个森林生态系统对未来气候变化的响应和适应能力,本文综述了树木NSC研究的最新进展,重点介绍了NSC库的季节和区域性变化,分析总结了影响树木NSC含量和分配格局变化的主要环境因素.最后还对未来气候变化背景下树木NSC库可能的响应策略和适应状况进行了讨论,展望了未来树木NSC领域的研究方向.     

Non-structural carbohydrate dynamics associated with antecedent stem water potential and air temperature in a dominant desert shrub

Non-structural carbohydrates (NSCs) are necessary for plant growth and affected by plant water status, but the temporal dynamics of water stress impacts on NSC are not well understood. We evaluated how seasonal NSC concentrations varied with plant water status (predawn xylem water potential, Ψ) and air temperature (T) in the evergreen desert shrub Larrea tridentata. Aboveground sugar and starch concentrations were measured weekly or monthly for ~1.5 years on 6–12 shrubs simultaneously instrumented with automated stem psychrometers; leaf photosynthesis (A_net) was measured monthly for 1 year. Leaf sugar increased during the dry, premonsoon period, associated with lower Ψ (greater water stress) and high T. Leaf sugar accumulation coincided with declines in leaf starch and stem sugar, suggesting the prioritization of leaf sugar during low photosynthetic uptake. Leaf starch was strongly correlated with A_net and peaked during the spring and monsoon seasons, while stem starch remained relatively constant except for depletion during the monsoon. Recent photosynthate appeared sufficient to support spring growth, while monsoon growth required the remobilization of stem starch reserves. The coordinated responses of different NSC fractions to water status, photosynthesis, and growth demands suggest that NSCs serve multiple functions under extreme environmental conditions, including severe drought.     

Stress from cold and drought as drivers of functional trait spectra in North American angiosperm tree assemblages

Understanding how environmental change alters the composition of plant assemblages, and how this in turn affects ecosystem functioning is a major challenge in the face of global climate change. Assuming that values of plant traits express species adaptations to the environment, the trait‐based approach is a promising way to achieve this goal. Nevertheless, how functional traits are related to species’ environmental tolerances and how trait spectra respond to broad‐scale environmental gradients remains largely unexplored. Here, we identify the main trait spectra for US angiosperm trees by testing hypotheses for the relationships between functional traits and species’ environmental tolerances to environmental stresses, as well as quantifying the environmental drivers of assemblage means and variances of these traits. We analyzed >74,000 community assemblages from the US Forest Inventory and Analysis using 12 functional traits, five traits expressing species’ environmental tolerances and 10 environmental variables. Results indicated that leaf traits, dispersal traits, and traits related to stem hydraulics were related to cold or drought tolerance, and their assemblage means were best explained by minimum temperatures. Assemblage means of traits related to shade tolerance (tree growth rate, leaf phosphorus content, and bark thickness) were best explained by aridity index. Surprisingly, aridity index, rather than minimum temperature, was the best predictors of assemblage variances of most traits, although these relationships were variable and weak overall. We conclude that temperature is likely to be the most important driver of functional community structure of North American angiosperm trees by selecting for optimum strategies along the cold and drought stress trade‐off. In turn, water availability primarily affects traits related to shade tolerance through its effect on forest canopy structure and vegetation openness.     

Climate and vegetation structure determine plant diversity in Quercus ilex woodlands along an aridity and human-use gradient in Northern Algeria

We studied the influence of environmental factors relating to climate, soil and vegetation cover on total species richness, species richness of different life-forms and species composition of plant communities occurring in Quercus ilex woodlands, across a 450-km long transect in Northern Algeria constituting a gradient of aridity and human use. We sampled vegetation and collected environmental data in 81 10 m × 10 m plots in five zones representing the largest Q. ilex woodlands throughout the study area, analysing them within an a priori hypothesis framework with the use of Path Analysis. Changes in plant diversity were mainly influenced by environmental factors related to precipitation and temperature regimes, as well as by total plant cover. In particular, changes in species composition were determined by factors associated with the temperature regime through their influence on both woody and annual herbaceous plant richness, and by factors related to the precipitation regime through their influence on perennial herbaceous plant richness, likely due to the differential tolerances of these functional groups to cold and water stress. Our results emphasize the importance of differences in environmental adaptability of the most important life-forms with regard to explaining compositional change (beta diversity) along aridity gradients, and the mediator role of total plant cover in relation to the effects of soil conditions on plant diversity.     

Frost Induces Respiration and Accelerates Carbon Depletion in Trees

Cellular respiration depletes stored carbohydrates during extended periods of limited photosynthesis, e.g. winter dormancy or drought. As respiration rate is largely a function of temperature, the thermal conditions during such periods may affect non-structural carbohydrate (NSC) availability and, ultimately, recovery. Here, we surveyed stem responses to temperature changes in 15 woody species. For two species with divergent respirational response to frost, P. integerrima and P. trichocarpa, we also examined corresponding changes in NSC levels. Finally, we simulated respiration-induced NSC depletion using historical temperature data for the western US. We report a novel finding that tree stems significantly increase respiration in response to near freezing temperatures. We observed this excess respiration in 13 of 15 species, deviating 10% to 170% over values predicted by the Arrhenius equation. Excess respiration persisted at temperatures above 0°C during warming and reoccurred over multiple frost-warming cycles. A large adjustment of NSCs accompanied excess respiration in P. integerrima, whereas P. trichocarpa neither excessively respired nor adjusted NSCs. Over the course of the years included in our model, frost-induced respiration accelerated stem NSC consumption by 8.4 mg (glucose eq.) cm^-3 yr^-1 on average in the western US, a level of depletion that may continue to significantly affect spring NSC availability. This novel finding revises the current paradigm of low temperature respiration kinetics.     

Phenotypic effects of the nurse Thylacospermum caespitosum on dependent plant species along regional climate stress gradients

Contrasting phenotypes of alpine cushion species have been recurrently described in several mountain ranges along small‐scale topography gradients, with tight competitive phenotypes in stressful convex topography and loose facilitative phenotypes in sheltered concave topography. The consistency of phenotypic effects along large‐scale climate stress gradients have been proposed as a test of the likely genetic bases of the differences observed at small‐scale. Inversely, plastic phenotypic effects are more likely to vanish at some points along climate stress gradients. We tested this hypothesis for two phenotypes of the alpine cushion species Thylacospermum caespitosum at four points along regional gradients of cold and drought stress in northwest China. We measured the traits of the two cushion phenotypes and quantified their associated plant communities and environmental variables along the regional temperature and aridity gradients. Cushion height, convexity and stem density overall showed significant effect of phenotypes. Difference in tightness of cushions between phenotypes was consistent across climate conditions, whereas differences in cushion convexity and height between phenotypes increased with increasing cold stress. Phenotypic effects on species richness and abundance were consistent along both climate gradients but not effects on species composition, while there were no phenotypic effects on environmental variables. Additionally, RII (relative interaction index) curves were linear along the drought gradient but unimodal along the temperature gradient, likely due to the occurrence of contrasting species pools at the different sites. We conclude that the consistency of phenotypic effects of T. caespitosum was high for species richness and abundance and mainly explained by differences in interference mediated by likely heritable differences in cushion tightness. Additionally, our study shows that the shapes of the relationship between plant responses to neighbours and environmental stresses are not necessarily driven by niche‐based deterministic factors.     

Growth responses to climate and drought at the southernmost European limit of Mediterranean Pinus pinaster forests

Climate warming and increasing aridity may negatively impact forest productivity across southern Europe. A better understanding of growth responses to climate and drought in southernmost populations could provide insight on the vulnerability of those forests to aridification. Here we investigate growth responses to climate and drought in nine Pinus pinaster (maritime pine) stands situated in Andalusia, southern Europe. The effect of climatic variables (temperatures and precipitation) and drought on radial growth was studied using dendrochronology along biogeographic and ecological gradients. We analyzed old native stands with non-tapped and resin-tapped trees mixed, showing their usefulness in dendroclimatic studies. Our results indicate a high plasticity in the growth responses of maritime pine to climate and drought, suggesting that site aridity modulated these responses. The positive growth responses to spring precipitation and the negative responses to summer drought were stronger in the more xeric inland sites than in wet coastal ones, in particular from the 1980s onwards. The characterization of tree species’ responses to climate at the southern or dry limits in relation to site conditions allows improving conservation strategies in drought-prone forest ecosystems.     

Interplay between abiotic (drought) and biotic (virus) stresses in tomato plants

With climate warming, drought becomes a vital challenge for agriculture. Extended drought periods affect plant–pathogen interactions. We demonstrate an interplay in tomato between drought and infection with tomato yellow leaf curl virus (TYLCV). Infected plants became more tolerant to drought, showing plant readiness to water scarcity by reducing metabolic activity in leaves and increasing it in roots. Reallocation of osmolytes, such as carbohydrates and amino acids, from shoots to roots suggested a role of roots in protecting infected tomatoes against drought. To avoid an acute response possibly lethal for the host organism, TYLCV down-regulated the drought-induced activation of stress response proteins and metabolites. Simultaneously, TYLCV promoted the stabilization of osmoprotectants' patterns and water balance parameters, resulting in the development of buffering conditions in infected plants subjected to prolonged stress. Drought-dependent decline of TYLCV amounts was correlated with HSFA1-controlled activation of autophagy, mostly in the roots. The tomato response to combined drought and TYLCV infection points to a mutual interaction between the plant host and its viral pathogen.     

Earlier summer drought affects leaf functioning of the Mediterranean species Cistus monspeliensis L.

The Mediterranean vegetation is characterized by a high diversity of growth forms, habits and phenology that enable it to endure under harsh environmental conditions. It is however unclear whether these adaptations may allow plant survival under more extreme conditions, as predicted by climatic models under the perspective of climate change. A manipulative experiment aiming at anticipating summer aridity has been run to analyse the effects of the experimental drought on spring-leaf functioning and characteristics of the leaf-dimorphic Mediterranean shrub Cistus monspeliensis L.Assimilation rates were reduced under anticipated summer aridity due to a decrease of stomatal conductance, but only before morphological adaptations to drought (an increase of leaf mass per area) occurred. These adaptations were anticipated under experimental dry conditions, and causes photosynthetic performances to recover compared to previous dates. When natural summer aridity occurred, the leaf mass per area also changed in the control. However, this causes no recovery of the photosynthetic performances, because of the decrease of stomatal conductance due to low soil water content and leaf water potential values. Moreover, under experimental drought, leaf shedding was anticipated to reduce water losses, causing an overall reduction of leaf lifespan.     

Long‐term forest resilience to climate change indicated by mortality,regeneration, and growth in semiarid southern Siberia

Several studies have documented that regional climate warming and the resulting increase in drought stress have triggered increased tree mortality in semiarid forests with unavoidable impacts on regional and global carbon sequestration. Although climate warming is projected to continue into the future, studies examining long‐term resilience of semiarid forests against climate change are limited. In this study, long‐term forest resilience was defined as the capacity of forest recruitment to compensate for losses from mortality. We observed an obvious change in long‐term forest resilience along a local aridity gradient by reconstructing tree growth trend and disturbance history and investigating postdisturbance regeneration in semiarid forests in southern Siberia. In our study, with increased severity of local aridity, forests became vulnerable to drought stress, and regeneration first accelerated and then ceased. Radial growth of trees during 1900–2012 was also relatively stable on the moderately arid site. Furthermore, we found that smaller forest patches always have relatively weaker resilience under the same climatic conditions. Our results imply a relatively higher resilience in arid timberline forest patches than in continuous forests; however, further climate warming and increased drought could possibly cause the disappearance of small forest patches around the arid tree line. This study sheds light on climate change adaptation and provides insight into managing vulnerable semiarid forests.     

Drought and shade deplete nonstructural carbohydrate reserves in seedlings of five temperate tree species

Plants that store nonstructural carbohydrates (NSC) may rely on carbon reserves to survive carbon‐limiting stress, assuming that reserves can be mobilized. We asked whether carbon reserves decrease in resource stressed seedlings, and if NSC allocation is related to species' relative stress tolerances. We tested the effects of stress (shade, drought, and defoliation) on NSC in seedlings of five temperate tree species (Acer rubrum Marsh., Betula papyrifera Marsh., Fraxinus americana L., Quercus rubra L., and Quercus velutina Lam.). In a greenhouse experiment, seedlings were subjected to combinations of shade, drought, and defoliation. We harvested seedlings over 32–97 days and measured biomass and NSC concentrations in stems and roots to estimate depletion rates. For all species and treatments, except for defoliation, seedling growth and NSC accumulation ceased. Shade and drought combined caused total NSC decreases in all species. For shade or drought alone, only some species experienced decreases. Starch followed similar patterns as total NSC, but soluble sugars increased under drought for drought‐tolerant species. These results provide evidence that species deplete stored carbon in response to carbon limiting stress and that species differences in NSC response may be important for understanding carbon depletion as a buffer against shade‐ and drought‐induced mortality.     

Environmental aridity is associated with cytotype segregation and polyploidy occurrence in Brachypodium distachyon (Poaceae)

? The ecological and adaptive significance of plant polyploidization is not well understood and no clear pattern of association between polyploid frequency and environment has emerged. Climatic factors are expected to predict cytotype distribution. However, the relationship among climate, cytotype distribution and variation of abiotic stress tolerance traits has rarely been examined. ? Here, we use flow cytometry and root-tip squashes to examine the cytotype distribution in the temperate annual grass Brachypodium distachyon in 57 natural populations distributed across an aridity gradient in the Iberian Peninsula. We further investigate the link between environmental aridity, ploidy, and variation of drought tolerance and drought avoidance (flowering time) traits. ? Distribution of diploids (2n = 10) and allotetraploids (2n = 30) in this species is geographically structured throughout its range in the Iberian Peninsula, and is associated with aridity gradients. Importantly, after controlling for geographic and altitudinal effects, the link between aridity and polyploidization occurrence persisted. Water-use efficiency varied between ploidy levels, with tetraploids being more efficient in the use of water than diploids under water-restricted growing conditions. ? Our results indicate that aridity is an important predictor of polyploid occurrence in B. distachyon, suggesting a possible adaptive origin of the cytotype segregation.     

Response of plant species richness and primary productivity in shrublands along a north–south gradient in Europe to seven years of experimental warming and drought: reductions in primary productivity in the heat and drought year of 2003

We used a nonintrusive field experiment carried out at six sites – Wales (UK), Denmark (DK), the Netherlands (NL), Hungary (HU), Sardinia (Italy – IT), and Catalonia (Spain – SP) – along a climatic and latitudinal gradient to examine the response of plant species richness and primary productivity to warming and drought in shrubland ecosystems. The warming treatment raised the plot daily temperature by ca. 1 °C, while the drought treatment led to a reduction in soil moisture at the peak of the growing season that ranged from 26% at the SP site to 82% in the NL site. During the 7 years the experiment lasted (1999–2005), we used the pin‐point method to measure the species composition of plant communities and plant biomass, litterfall, and shoot growth of the dominant plant species at each site. A significantly lower increase in the number of species pin‐pointed per transect was found in the drought plots at the SP site, where the plant community was still in a process of recovering from a forest fire in 1994. No changes in species richness were found at the other sites, which were at a more mature and stable state of succession and, thus less liable to recruitment of new species. The relationship between annual biomass accumulation and temperature of the growing season was positive at the coldest site and negative at the warmest site. The warming treatment tended to increase the aboveground net primary productivity (ANPP) at the northern sites. The relationship between annual biomass accumulation and soil moisture during the growing season was not significant at the wettest sites, but was positive at the driest sites. The drought treatment tended to reduce the ANPP in the NL, HU, IT, and SP sites. The responses to warming were very strongly related to the Gaussen aridity index (stronger responses the lower the aridity), whereas the responses to drought were not. Changes in the annual aboveground biomass accumulation, litterfall, and, thus, the ANPP, mirrored the interannual variation in climate conditions: the most outstanding change was a decrease in biomass accumulation and an increase in litterfall at most sites during the abnormally hot year of 2003. Species richness also tended to decrease in 2003 at all sites except the cold and wet UK site. Species‐specific responses to warming were found in shoot growth: at the SP site, Globularia alypum was not affected, while the other dominant species, Erica multiflora, grew 30% more; at the UK site, Calluna vulgaris tended to grow more in the warming plots, while Empetrum nigrum tended to grow less. Drought treatment decreased plant growth in several studied species, although there were some species such as Pinus halepensis at the SP site or C. vulgaris at the UK site that were not affected. The magnitude of responses to warming and drought thus depended greatly on the differences between sites, years, and species and these multiple plant responses may be expected to have consequences at ecosystem and community level. Decreases in biodiversity and the increase in E. multiflora growth at the SP site as a response to warming challenge the assumption that sensitivity to warming may be less well developed at more southerly latitudes; likewise, the fact that one of the studied shrublands presented negative ANPP as a response to the 2003 heat wave also challenges the hypothesis that future climate warming will lead to an enhancement of plant growth and carbon sequestration in temperate ecosystems. Extreme events may thus change the general trend of increased productivity in response to warming in the colder sites.     

Seasonal changes in cold tolerance,water relations and accumulation of cations and compatible solutes in Atriplex halimus L.

We determined the cold (freezing) tolerance for field-grown plants of Atriplex halimus L. (Chenopodiaceae) in relation to plant ploidy level, leaf water relations and accumulation of osmolytes. Plants were grown at two sites in Murcia (Spain), having average minimum temperatures in the coldest month of 0.6 and 12.1 °C, respectively. LT₅₀ values derived from laboratory freezing tests, using leaves taken from the plants in early winter and in spring, showed greater tolerance for winter-harvested leaves; the acclimation was more pronounced at the cold-winter site. Cold tolerance was related positively with leaf K and/or Na accumulation. Analysis of compatible organic solutes (soluble sugars, total amino acids and quaternary ammonium compounds) showed that cold tolerance (measured both as LT₅₀ and as winter freezing damage in situ) was related most closely with leaf concentrations of soluble sugars. The leaf percentage dry matter content was related to both in vitro and in vivo tolerance, while tolerance in vitro was correlated also with the osmotic (potential ψ_s) and the relative water content. The two diploid (2n = 2x = 18) populations, from Spain, showed greater cold tolerance than the three tetraploid (2n = 4x = 36) populations, from North Africa and Syria, which may be related to the latter's greater cell size and consequent dilution of osmolytes. In this halophytic species, cold tolerance, like salinity and drought tolerance, seems to depend on osmotic adjustment, driven by vacuolar accumulation of K and Na and cytoplasmic accumulation of compatible solutes.     

Seasonal changes in cold tolerance,water relations and accumulation of cations and compatible solutes in Atriplex halimus L.

We determined the cold (freezing) tolerance for field-grown plants of Atriplex halimus L. (Chenopodiaceae) in relation to plant ploidy level, leaf water relations and accumulation of osmolytes. Plants were grown at two sites in Murcia (Spain), having average minimum temperatures in the coldest month of 0.6 and 12.1 °C, respectively. LT₅₀ values derived from laboratory freezing tests, using leaves taken from the plants in early winter and in spring, showed greater tolerance for winter-harvested leaves; the acclimation was more pronounced at the cold-winter site. Cold tolerance was related positively with leaf K and/or Na accumulation. Analysis of compatible organic solutes (soluble sugars, total amino acids and quaternary ammonium compounds) showed that cold tolerance (measured both as LT₅₀ and as winter freezing damage in situ) was related most closely with leaf concentrations of soluble sugars. The leaf percentage dry matter content was related to both in vitro and in vivo tolerance, while tolerance in vitro was correlated also with the osmotic (potential ψ_s) and the relative water content. The two diploid (2n = 2x = 18) populations, from Spain, showed greater cold tolerance than the three tetraploid (2n = 4x = 36) populations, from North Africa and Syria, which may be related to the latter's greater cell size and consequent dilution of osmolytes. In this halophytic species, cold tolerance, like salinity and drought tolerance, seems to depend on osmotic adjustment, driven by vacuolar accumulation of K and Na and cytoplasmic accumulation of compatible solutes.