Non-structural Carbohydrate Content Research Articles

Impacts of groundwater depth and tree age on the non-structural carbohydrates of Haloxylon ammodendron

Nonstructural carbohydrates (NSCs) are important substrates for plant growth and metabolism, and their concentration reflects the plant's ability to adapt to environmental changes. Although the response of NSCs to changes in water availability has been extensively studied, it is still not fully understood whether this response is modulated by tree ages and organs. This study investigates Haloxylon ammodendron (C.A. Mey.) Bunge ex Fenzl, the dominant species in the Gurbantunggut Desert in the Uyghur Autonomous Region of China. Utilizing the natural topographic conditions characterized by a gradual increase in groundwater depth from the desert edge to the interior, we collected samples of different age classes of H. ammodendron along a groundwater depth gradient of 3, 7, 10, and 14 meters. We measured the total concentrations of non-structural carbohydrates (NSCs) and their components soluble sugar (SS) and starch (ST) in the assimilative twigs and stems. The results showed that the assimilative twigs of H. ammodendron exhibited higher NSC concentrations at the site with the deepest groundwater, while the other three sites showed similar NSC concentrations. Furthermore, as groundwater depth increased, the concentrations of SS in the assimilative twigs increased, whereas ST concentrations decreased. Similarly, the concentrations of SS in the stems also increased at sites with deeper groundwater. The NSC concentrations in the assimilative twigs were significantly affected by groundwater depth, while variations in stem NSC were primarily driven by plant age. In younger trees, higher soluble sugars concentrations in the stem may enhance water transport efficiency, whereas older trees tend to store more NSC to alleviate drought stress. Overall, elevated nonstructural carbohydrate concentrations contributed to greater drought resilience in H. ammodendron. These results suggest that different age classes of H. ammodendron exhibit distinct physiological responses to decreasing groundwater depth. The varying requirements for soluble sugars and starch in H. ammodendron help to partially mitigate the adverse effects of reduced groundwater accessibility. These findings provide important insights into the physiological adaptations of H. ammodendron in arid environments and offer a scientific basis for future ecological restoration and management strategies.

Analyzing the impact of phosphorous and nitrogen on Castanopsis sclerophylla early growth stages

Plant growth elements, particularly nitrogen (N) and phosphorus (P) are vital for their growth and development, particularly for understory vegetation and their excess limits the net productivity of terrestrial ecosystems. This study focuses on the understory vegetation responds and adaptation to key essential nutrients under changing climate scenarios in subtropical evergreen broad-leaved forests, still needs research attention. this, we set up an experiment taking four treatments in a 50-year-old Castanopsis sclerophylla secondary forest under (a) control (CK), (b) N, (c) P, and (d) combined N and P addition, applied to natural forest regeneration seedlings of C. sclerophylla attained similar growth parameters of diameter of 3 cm and 10 cm height. In addition, carbon, N, P, and non-structural carbohydrates (NSC) were determined through the anthrone colorimetric approach in different parts of seedlings. Results show that the combined N + P application enhanced the N and P by 14.48 %−140.55 % in the seedlings in both dry and wet seasons, respectively. However, during wet season, the content of NSC in the plant leaves significantly exceeded under P addition. Remarkably, CK showed increased P in the growing season but lower during the dry season. Furthermore, the root starch content of seedlings showed a significant increase under the application of N and P compared to combined N + P, ranging between 45.60 % and 58.70 %. Overall, the plant growth is attributed to N and P intake. The nutrient addition and seasonal variations have a coupled effect on seedling growth as proved in the in the natural open forest experiment. The study outcomes emphasize that the alterations in NSC allocation in the roots and leaves of C. sclerophylla seedlings under N + P addition could enhance their adaptation to future global climate changes, drought conditions, and high N concentrations.

Effects of Biochar and Nitrogen Fertilizer Application Ratio on Leaf Physiology and Soil Characteristics of Bambusa tuldoides ‘Swolleninternode’

As one of the main bamboo species in coastal sandy land protection forests, Bambusa tuldoides ‘Swolleninternode’ can effectively improve the structure of forest tree species, increase the diversity of tree species and enhance the protection efficiency. However, research on the cultivation and utilization of B. tuldoides is still relatively scarce. Therefore, in this study, B. tuldoides was used as the research object. By applying biochar and nitrogen fertilizer, the effects of different biochar and nitrogen fertilizer ratios on the physiology and soil characteristics of bamboo were analyzed, and the optimal ratio scheme was identified. The results showed that the ratio of the T5 (A2B2C3) treatment had the best effect on the total chlorophyll, non-structural carbohydrate and nutrient contents of B. tuldoides leaves, and the contents under treatments was generally higher than under the control, CK. The activities of soil invertase, nitrate reductase and nitrite reductase were significantly increased under biochar treatment, and the effects of the T5 treatment were the best. The results of principal component analysis showed that the absolute values of the coefficient of leaf potassium, phosphorus content and soil total nitrogen were larger and more important, and the comprehensive evaluation of the T5 treatment was the highest.

Water, not carbon, drives drought-constraints on stem terpene defense against simulated bark beetle attack in Pinus edulis.

Drought predisposes forest trees to bark beetle-induced mortality, but the physiological mechanisms remain unclear. While drought-induced water and carbon limitations have been implicated in defensive failure and tree susceptibility, evidence demonstrating how these factors interact is scarce. We withheld water from mature, potted Pinus edulis and subsequently applied a double-stem girdle to inhibit carbohydrate transport from the crown and roots. Within this isolated segment we then elicited a defense response by inoculating trees with a bark beetle-fungal symbiont (Ophiostoma sp.). We quantified local mono- and sesquiterpenes (MST), nonstructural carbohydrates (NSC), and pressure potential of the inner bark. Both drought-stressed and watered trees had similar NSC concentrations just before inoculation and depleted NSC similarly following inoculation, yet MST induction (i.e. increased concentration and altered composition) was constrained only in drought-stressed trees. Thus, NSC consumption was largely unrelated to de novo MST synthesis. Instead, stoichiometric calculations show that induction originated largely from stored resin. Watered trees experiencing higher pressure potentials consistently induced higher MST concentrations. We demonstrate the importance of preformed resin toward an induced MST response in a semi-arid conifer where drought-constraints on defense occurred through biophysical limitations (i.e. reduced turgor hindering resin transport) rather than through substrate limitation.

Halophytic succulence is a driver of the leaf non-structural carbohydrate contents in plants in the arid and hyper-arid deserts of northwestern China.

Non-structural carbohydrates (NSC), primarily sugars and starch, play a crucial role in plant metabolic processes and a plant's ability to tolerate and recover from drought stress. Despite their importance, our understanding of NSC characteristics in the leaves of plants that thrive in hyper-arid and saline environments remains limited. To investigate the variations in leaf NSC across different species and spatial scales, and to explore their possible causes, we collected 488 leaf samples from 49 native plant species at 115 sites in the desert area of northwestern China. The contents of soluble sugars (SS), starch, and total NSC were then determined. The average contents of SS, starch, and total NSC were 26.99, 60.28, and 87.27mg g-1 respectively, which are much lower than those reported for Chinese forest plants and global terrestrial plants. Herbaceous and woody plants had similar NSC levels. In contrast, succulent halophytes, a key component of desert flora, showed significantly lower leaf SS and total NSC contents than non-succulent plants. We observed a strong negative correlation between leaf succulence and SS content, suggesting a role of halophytic succulence in driving multi-species NSC pools. Environmental factors explained a minor portion of the spatial variation in leaf NSC, possibly due to the narrow climatic variation in the study area, and soil properties, particularly soil salinity, emerged as more significant contributors. Our findings increase the understanding of plant adaptation to drought and salt stress, emphasizing the crucial role of halophytic succulence in shaping the intricate dynamics of leaf NSC across diverse plant species in arid and hyper-arid environments.

Combined transcriptome and physiological analysis reveals exogenous sucrose enhances photosynthesis and source capacity in foxtail millet

Foxtail millet (Setaria italica (L.) P. Beauv.) is an environmentally friendly crop that meets the current requirements of international food security and is widely accepted as a photosynthesis research model. However, whether exogenous sucrose treatment has a positive effect on foxtail millet growth remains unknown. Here, we employed physiological and molecular approaches to identify photosynthesis and source capacity associated with exogenous sucrose during the growth of Jingu 21 seedlings. RNA-seq analysis showed that some differentially expressed genes (DEGs) related to photosynthesis and carotenoid biosynthesis were induced by exogenous sucrose and that most of these genes were up-regulated. An increase in gas exchange parameters, chlorophyll content, and chlorophyll fluorescence of Jingu 21 was noted after exogenous sucrose addition. Furthermore, exogenous sucrose up-regulated genes encoding sucrose and hexose transporters and enhanced starch and sucrose metabolism. More DEGs were up-regulated by sucrose, the nonstructural carbohydrate (NSC) content in the leaves increased and energy metabolism and sucrose loading subsequently improved, ultimately enhancing photosynthesis under normal and dark conditions. Further analysis revealed that WRKYs, ERFs, HY5, RAP2, and ABI5 could be key transcription factors involved in growth regulation. These results indicate that exogenous sucrose affects the normal photosynthetic performance of foxtail millet by increasing NSC transport and loading. They improve our understanding of the molecular mechanisms of the effects of exogenous sucrose on photosynthesis in foxtail millet, providing an effective measure to enhance source–sink relationships and improve yield.

Megafire smoke exposure jeopardizes tree carbohydrate reserves and yield.

The global incidence of megafires is on the rise, leading to extensive areas being shrouded in dense smoke for prolonged periods, spanning days or weeks1. Here, by integrating long-term regional observations of non-structural carbohydrate content in trees across California's Central Valley with spatiotemporal satellite data, we present compelling evidence that dense smoke plumes negatively impact carbohydrate stores in three tree species: Prunus dulcis, Pistacia vera and Juglans regia. Our findings show that the presence of smoke causes a significant decrease in total non-structural carbohydrates, with reductions in the accumulation of both soluble sugar and starch reserves. This decline in carbohydrate levels persists through the trees' dormancy period into the next season's bloom, culminating in a reduced yield. Our results highlight a previously unrecognized wildfire threat that could affect plant health and ecosystem stability in both agricultural and natural environments.

Adjustment of storage capacity for non-structural carbohydrates in response to limited water availability in two temperate woody species.

Reserves of non-structural carbohydrates (NSC) stored in living cells are essential for drought tolerance of trees. However, little is known about the phenotypic plasticity of living storage compartments (SC) and their interactions with NSC reserves under changing water availability. Here, we examined adjustments of SC and NSC reserves in stems and roots of seedlings of two temperate tree species, Acer negundo L. and Betula pendula Roth., cultivated under different substrate water availability. We found that relative contents of soluble NSC, starch and total NSC increased with decreasing water availability in stems of both species, and similar tendencies were also observed in roots of A. negundo. In the roots of B. pendula, soluble NSC contents decreased along with the decreasing water availability, possibly due to phloem decoupling or NSC translocation to shoots. Despite the contrast in organ responses, NSC contents (namely starch) positively correlated with proportions of total organ SC. Individual types of SC showed markedly distinct plasticity upon decreasing water availability, suggesting that water availability changes the partitioning of organ storage capacity. We found an increasing contribution of parenchyma-rich bark to the total organ NSC storage capacity under decreasing water availability. However, xylem SC showed substantially greater plasticity than those in bark. Axial storage cells, namely living fibers in A. negundo, responded more sensitively to decreasing water availability than radial parenchyma. Our results demonstrate that drought-induced changes in carbon balance affect the organ storage capacity provided by living cells, whose proportions are sensitively coordinated along with changing NSC reserves.

The Legacy Effect of Mountain Pine Beetle Outbreaks on the Chemical and Anatomical Defences of Surviving Lodgepole Pine Trees.

The recent mountain pine beetle outbreaks have caused widespread mortality among lodgepole pine trees in western North America, resulting in a reduced population of surviving trees. While previous studies have focused on the cascading impacts of these outbreaks on the physiology and growth of the surviving trees, there remains a need for a comprehensive study into the interactions among various physiological traits and the growth in post-outbreak stands. Specifically, the relationship between chemical (primarily terpenes) and anatomical (mainly resin ducts) defences, as well as the allocation of non-structural carbohydrates (NSCs) to support these defence modalities, is poorly understood. To address these gaps, we conducted a field survey of surviving lodgepole pine trees in post-mountain pine beetle outbreak stands in western Canada. Our retrospective analysis aimed at determining correlations between the post-outbreak concentrations of monoterpenes, diterpenes, and NSCs in the phloem and the historical resin duct characteristics and growth traits before and after the outbreak. We detected strong correlations between the post-outbreak concentrations of monoterpenes and historical resin duct characteristics, suggesting a possible link between these two defence modalities. Additionally, we found a positive relationship between the NSCs and the total concentrations of monoterpenes and diterpenes, suggesting that NSCs likely influence the production of these terpenes in lodgepole pine. Furthermore, historical tree growth patterns showed strong positive correlations with many individual monoterpenes and diterpenes. Interestingly, while surviving trees had enhanced anatomical defences after the outbreak, their growth patterns did not vary before and after the outbreak conditions. The complexity of these relationships emphasizes the dynamics of post-outbreak stand dynamics and resource allocations in lodgepole pine forests, highlighting the need for further research. These findings contribute to a broader understanding of conifer defences and their coordinated responses to forest insect outbreaks, with implications for forest management and conservation strategies.

Divergent responses of woody plant leaf and root non-structural carbohydrates to nitrogen addition in China: Seasonal variations and ecological implications

Plant non-structural carbohydrates (NSCs), which largely comprise starch and soluble sugars, are essential energy reserves to support plant growth and physiological functions. While it is known that increasing global deposition of nitrogen (N) affects plant concentration of NSCs, quantification of seasonal responses and drivers of woody species leaf and root NSCs to N addition at larger spatial scales remains lacking. Here, we systematically analyzed data from 53 field experiments distributed across China, comprising 1202 observations, to test for effects of N addition on woody plant leaf and root NSCs across and within growing and non-growing seasons. We found (1) no overall effects of N addition on the concentrations of leaf and root NSCs, soluble sugars or starch during the growing season or the non-growing season for leaves. However, N addition decreased root NSC and starch concentrations by 13.8 % and 39.0 %, respectively, and increased soluble sugars concentration by 15.0 % during the non-growing season. (2) Shifts in leaf NSC concentration under N addition were driven by responses by soluble sugars in both seasons, while shifts in root NSC were driven by soluble sugars in the non-growing season and starch and soluble sugars in the growing season. (3) Relationships between N, carbon, and phosphorus stoichiometry with leaf and root NSCs indicated effects of N addition on woody plant NSCs allocation through impacts on plant photosynthesis, respiration, and growth. (4) Effects of N addition on leaf and root NSCs varied with plant functional types, where effects were more pronounced in roots than in leaves during the non-growing season. Overall, our results reveal divergent responses of woody plant leaf and root NSCs to N addition within non-growing season and highlight the role of ecological stoichiometry and plant functional types in woody plant allocation patterns of NSCs in response to ongoing N deposition under global change.

Physiological and biochemical changes of Picea abies (L.) during acute drought stress and their correlation with susceptibility to Ips typographus (L.) and I. duplicatus (Sahlberg)

IntroductionIn recent years, Norway spruce (Picea abies L.) forests in Central Europe have faced escalating threats from bark beetles, primarily Ips typographus (L.), and other species, such as I. duplicatus (Sahlberg). Outbreaks are partially attributed to weakened tree defense resulting from drought periods induced by climate change. This study examines Norway spruce’s physiological and metabolic reactions to acute drought stress during the growing season and evaluates its susceptibility to I. typographus and I. duplicatus.MethodologyIn order to induce drought stress, mature Norway spruces had their roots covered with a roof in April 2021, depriving them of water. Control trees were left with free access to natural rainwater. Over 5 months of the growing season, soil water potential, bark temperature, tree trunk circumference, and sap flow were monitored. Roofed trees and controls were sampled in July, August, and September and analyzed for non-structural carbohydrates and the two classes of defensive compounds, phenolics and terpenes. Furthermore, two different bioassays in tubes and boxes were performed using adult I. typographus and I. duplicatus beetles to assess host choice and acceptance.ResultsRoofed trees exhibited signs of stress as early as July, resulting in decreased tree trunk and a consequent increase in non-structural carbohydrate content. Defensive metabolites remained largely unaffected except for an increase in diterpenes in September. In bioassays, I. typographus preferred boring into the bark of roofed trees in August in tubes and in September in boxes. This increased tree acceptance correlated with increased levels of soluble carbohydrates in the phloem. I. typographus and I. duplicatus beetles showed higher mobility in boxes in August and September on roofed trees but not in July, even though bark surface temperatures were elevated in roof-covered trees during all three bioassay periods.ConclusionThe study revealed rapid physiological responses of trees to acute drought stress, although not many changes were observed in defense traits. Despite the absence of natural bark beetle attacks, drought trees were more accepted by I. typographus than naturally watered trees. This response may indicate the beetles’ preference for trees with phloem of higher nutritional quality induced by the acute drought stress conditions.

The underlying mechanisms by which boron mitigates copper toxicity in Citrus sinensis leaves revealed by integrated analysis of transcriptome, metabolome and physiology.

Both copper (Cu) excess and boron (B) deficiency are often observed in some citrus orchard soils. The molecular mechanisms by which B alleviates excessive Cu in citrus are poorly understood. Seedlings of sweet orange (Citrus sinensis (L.) Osbeck cv. Xuegan) were treated with 0.5 (Cu0.5) or 350 (Cu350 or Cu excess) μM CuCl2 and 2.5 (B2.5) or 25 (B25) μM HBO3 for 24wk. Thereafter, this study examined the effects of Cu and B treatments on gene expression levels revealed by RNA-Seq, metabolite profiles revealed by a widely targeted metabolome, and related physiological parameters in leaves. Cu350 upregulated 564 genes and 170 metabolites, and downregulated 598 genes and 58 metabolites in leaves of 2.5μM B-treated seedlings (LB2.5), but it only upregulated 281 genes and 100 metabolites, and downregulated 136 genes and 40 metabolites in leaves of 25μM B-treated seedlings (LB25). Cu350 decreased the concentrations of sucrose and total soluble sugars and increased the concentrations of starch, glucose, fructose and total nonstructural carbohydrates in LB2.5, but it only increased the glucose concentration in LB25. Further analysis demonstrated that B addition reduced the oxidative damage and alterations in primary and secondary metabolisms caused by Cu350, and alleviated the impairment of Cu350 to photosynthesis and cell wall metabolism, thus improving leaf growth. LB2.5 exhibited some adaptive responses to Cu350 to meet the increasing need for the dissipation of excessive excitation energy (EEE) and the detoxification of reactive oxygen species (reactive aldehydes) and Cu. Cu350 increased photorespiration, xanthophyll cycle-dependent thermal dissipation, nonstructural carbohydrate accumulation, and secondary metabolite biosynthesis and abundances; and upregulated tryptophan metabolism and related metabolite abundances, some antioxidant-related gene expression, and some antioxidant abundances. Additionally, this study identified some metabolic pathways, metabolites and genes that might lead to Cu tolerance in leaves.

The rates of starch depletion and hydraulic failure both play a role in drought-induced seedling mortality

Key messageThe elapsed times to deplete starch concentrations and to reach a null hydraulic safety margin were related to tree seedling mortality under experimental drought. Starch concentration showed an accelerated decline across all species during the early stages of dehydration, while the concentrations of soluble sugars and total nonstructural carbohydrates remained stable. Concomitant carbohydrate depletion and hydraulic failure drive seedling mortality under drought.ContextCurrent upsurges of drought events are provoking impacts on tree physiology, resulting in forest mortality. Hydraulic dysfunction and nonstructural carbohydrate (NSC) depletion have been posited as the main mechanisms leading to plant mortality under drought.AimsThis study explores the dynamics of the two mortality-inducing processes during drought stress using an experimental approach with 12 evergreen tree species.MethodsSeedlings were subjected to drought until 100% mortality was observed. Midday (ΨMD) and predawn (ΨPD) water potentials, xylem pressure leading to a 50% loss of hydraulic conductivity (Ψ50), along with NSC concentrations in different organs (leaves, stems, and roots) were measured regularly during drought.ResultsTotal NSC concentrations and soluble sugar pools did not decline during drought. However, starch pools showed strong reductions early during drought stress as ΨPD decreased, and the time leading to starch depletion emerged as a strong mortality predictor. Ψ50 alone did not provide an accurate estimate of mortality, while the elapsed time to reach a null hydraulic safety margin (ΨMD—Ψ50 = 0) was related to seedling mortality.ConclusionAdopting a dynamic approach by estimating the times to consume both starch reserves and hydraulic safety margins is highly relevant to improve predictions of tree mortality under the current context of increasing global drought.

Non-Structural Carbohydrate Content and C:N:P Stoichiometry in Houpoea officinalis Flowers in Response to Development Stages

Non-Structural Carbohydrate Content and C:N:P Stoichiometry in Houpoea officinalis Flowers in Response to Development Stages

Snowpack permanence shapes the growth and dynamic of non-structural carbohydrates in Juniperus communis in alpine tundra

Climate warming is altering snowpack permanence in alpine tundra, modifying shrub growth and distribution. Plant acclimation to snowpack changes depends on the capability to guarantee growth and carbon storage, suggesting that the content of non-structural carbohydrates (NSC) in plant organs can be a key trait to depict the plant response under different snow regimes.To test this hypothesis, we designed a 3-years long manipulative experiment aimed at evaluating the effect of snow melt timing (i.e., early, control, and late) on NSC content in needles, bark and wood of Juniperus communis L. growing at high elevation in the Alps.Starch evidenced a general decrease from late spring to summer in control and early melting, while starch was low but stable in plants subjected to a late snow melt. Leaves, bark and wood have different level of soluble NSC changing during growing season: in bark, sugars content decreased significantly in late summer, while there was no seasonal effect in needles and wood. Soluble NSC and starch were differently related with the plant growth, when considering different tissues and snow treatment. In leaf and bark we observed a starch depletion in control and early melting plants, consistently to a higher growth (i.e., twig elongation), while in late snow melt, we did not find any significant relationship between growth and NSC concentration.Our findings confirmed that snowpack duration affects the onset of the growing season promoting a change in carbon allocation in plant organs and, between bark and wood in twigs. Finally, our results suggest that plants, at this elevation, could take advantage from an early snow melt caused by climate warming, most likely due to photosynthetic activity by maintaining the level of reserves and enhancing the carbon investment for growth.

Forage legume responses to climate change factors

AbstractIncorporating forage legumes into grasslands is a recommended climate change mitigation strategy, but accruing desired benefits from legumes is contingent upon their resilience when exposed to climate change factors (CCF). Our objective was to synthesize literature describing responses to CCF of a broad array of forage legume species, including annuals and perennials from both temperate and tropical/subtropical regions. Most‐represented species in the related literature include alfalfa (Medicago sativa L.), various Trifolium and Lotus species, rhizoma peanut (Arachis glabrata Benth.), and capitata stylo (Stylosanthes capitata Vogel). The data indicate that while CCF effects on forage legumes can be generalized, interactions among CCF, species, and site‐specific soil/climate conditions may cause variation from expected responses. In most instances, exposing forage legumes to CO2 enrichment (eCO2) increased forage accumulation (FA), but elevated temperature (eT) often reduced the magnitude of the positive response to eCO2. Photosynthetic acclimation to eCO2 occurs in non‐N‐fixing plants, but legume nodules are large C sinks and drive sustained increases in legume FA to eCO2. Legume N2 fixation and the proportion of legume N derived from N2 fixation increase with eCO2, but the magnitude of the increases lessens with eT. Legume nutritive value (NV) responses to eCO2 are less pronounced than FA responses, but decreased herbage N and greater nonstructural carbohydrate concentrations are common. Exposure to eT negatively affects NV unless intervals between defoliation events are shortened. Limited data on reproductive performance show CCF affect flower number, pollen grain morphology and viability, and behavior of pollinators, potentially influencing legume reproductive success.

Coping with extremes: Responses of Quercus robur L. and Fagus sylvatica L. to soil drought and elevated vapour pressure deficit

Climate change, particularly droughts and heat waves, significantly impacts global photosynthesis and forest ecosystem sustainability. To understand how trees respond to and recover from hydrological stress, we investigated the combined effects of soil moisture and atmospheric vapour pressure deficit (VPD) on seedlings of the two major European broadleaved tree species Fagus sylvatica (FS) and Quercus robur (QR). The experiment was conducted under natural forest gap conditions, while soil water availability was strictly manipulated. We monitored gas exchange (net photosynthesis, stomatal conductance and transpiration rates), nonstructural carbohydrates (NSC) concentration in roots and stomatal morphometry (size and density) during a drought period and recovery. Our comparative empirical study allowed us to distinguish and quantify the effects of soil drought and VPD on stomatal behavior, going beyond theoretical models. We found that QR conserved water more conservatively than FS by reducing transpiration and regulating stomatal conductance under drought. FS maintained higher stomatal conductance and transpiration at elevated VPD until soil moisture became critically low. QR showed higher intrinsic water use efficiency than FS. Stomata density and size also likely played a role in photosynthetic rate and speed of recovery, especially since QR with its seasonal adjustments in stomatal traits (smaller, more numerous stomata in summer leaves) responded and recovered faster compared to FS. Our focal species showed different responses in NSC content under drought stress and recovery, suggesting possible different evolutionary pathways in coping with stress. QR mobilized soluble sugars, while FS relied on starch mobilization to resist drought. Although our focal species often co-occur in mixed forests, our study showed that they have evolved different physiological, morphological and biochemical strategies to cope with drought stress. This suggests that ongoing climate change may alter their competitive ability and adaptive potential in favor of one of the species studied.

Seasonal Variability in Non-Structural Carbohydrate Content of Warm-Adapted Zostera noltei and Zostera marina Populations

Non-structural carbohydrates (NSCs) are energetic compounds that can be accumulated in tissues and mobilized during periods of unfavorable conditions to maintain the biological functions of plants. The balance of these biochemical compounds is controlled by environmental factors such as temperature and irradiance. Zostera noltei and Zostera marina find one of their southern distribution limits in southern Spain, where relatively high seawater temperatures are reached during summer (23–24 °C). To better understand the effects of elevated temperatures on the concentration of NSCs, we conducted a seasonal study at Cadiz Bay, representing warm-adapted populations of these species. Our results showed a bimodal pattern in both species, with the highest NSC content observed in December and June, followed by a depletion in March and August. In addition, the NSC content observed in the leaves of Z. noltei (71.26 ± 30.77 mg g−1 dry weight) was higher than in the rhizomes and roots (52.14 ± 38.86 mg g−1 DW). The observed patterns suggest that these species accumulated NSCs to cope with periods of unfavorable environmental conditions. We also suggest that the limited concentration of NSCs in Z. noltei rhizomes and roots indicates that this population may be suffering physiological stress.

Leaf membrane leakage and xylem hydraulic failure define the point of no return in drought-induced tree mortality in Cupressus sempervirens.

Measurements of resistance to embolism suggest that Cupressus sempervirens has a stem xylem that resists embolism at very negative water potentials, with 50% embolism (P50) at water potentials of approximately -10 MPa. However, field observations in a semi-arid region suggest tree mortality occurs before 10% embolism. To explore the interplay between embolism and plant mortality, we conducted a controlled drought experiment involving two types of CS seedlings: a local seed source (S-type) and a drought-resistant clone propagated from a semi-arid forest (C-type). We measured resistance to embolism, leaf relative water content (RWC), water potential, photosynthesis, electrolyte leakage (EL), plant water loss, leaf hydraulic conductivity, and leaf non-structural carbohydrate (NSC) content during plant dehydration and before rewatering. All measured individuals were monitored for survival or mortality. While the S- and C-types differed in P50, transpiration, and mortality rates, both displayed seedling mortality corresponding to threshold values of 52-55% leaf RWC, 55% and 18.5% percent loss of conductivity (PLC) in the xylem, which corresponds to 48% and 37% average EL values for S and C types, respectively. Although C-type C. sempervirens NSC content increased in response to drought, no differences were observed in NSC content between live and dead seedlings of both types. Our findings do not fully explain tree mortality in the field but they do indicate that loss of membrane integrity occurs before or at xylem water potential, leading to hydraulic failure.

Seasonal plasticity of stem embolism resistance and its potential driving factors in six temperate woody species.

The seasonal plasticity of resistance to xylem embolism has been demonstrated in leaves of some tree species, but is controversial in stems. In this study, we investigated the seasonality of stem xylem resistance to embolism in six temperate woody species (four deciduous and two evergreen tree species) that were grown at the same site. The xylem conduit anatomy, the concentrations, and ratios of the main cation in the xylem sap, as well as the content of nonstructural carbohydrates (including soluble sugars and starch) were measured in each species under each season to reveal the potential mechanisms of seasonal change in embolism resistance. The stem of all species showed increasing resistance to embolism as seasons progressed, with more vulnerable xylem in spring, but no significant adjustment in the other three seasons. The seasonal plasticity of stem embolism resistance was greater in deciduous species than in evergreen. On a seasonal scale, conduit diameter and conduit implosion resistance, the ratios of K+/Ca2+ and K+/Na+, and starch content were generally not correlated with embolism resistance, suggesting that these are probably not the main drivers of seasonal plasticity of stem embolism resistance. The seasonality of embolism resistance provides critical information for better understanding plant hydraulics in response to seasonal environments, especially under climate change.