Search

In some forest systems, stomata have also been observed to reopen rapidly on recovery from drought, with delays in gas exchange recovery only observed once embolism-induced declines in hydraulic conductance occur ([29]). (B) When relief from a drought occurs after embolism has formed in the xylem then gas exchange recovery can take many months in species that must construct new xylem and organs to regain pre-drought gas exchange capacity ([3], [1]). [2] report that on rewatering from a drought severe enough to close stomata, the complete recovery of leaf gas exchange took >7 days, yet water potentials recovered within 12 h of rewatering. [Extracted from the article]

Clarifying the mechanisms underlying the recovery of gas exchange following drought is the key to providing insights into plant drought adaptation and habitat distribution. However, the mechanisms are still largely unknown. Targeting processes known to inhibit gas exchange during drought recovery, we measured leaf water potential, the leaf hydraulic conductance, stomatal reopening, abscisic acid (ABA) and the ethylene emission rate (EER) following moderate drought stress in seedlings of the globally pervasive woody tree Fraxinus chinensis. We found strong evidence that the slow stomatal reopening after rehydration is regulated by a slow decrease in EER, rather than changes in leaf hydraulics or foliar ABA levels. This was supported by evidence of rapid gas exchange recovery in plants after treatment with the ethylene antagonist 1-methylcyclopropene. These findings provide evidence to rigorously support ethylene as a key factor constraining stomatal reopening from moderate drought directly, thereby potentially opening new windows for understanding species drought adaptation. [ABSTRACT FROM AUTHOR]

The study of ancient species provides valuable information concerning the evolution of specific adaptations to past and current environmental conditions. Araucaria araucana (Molina) K. Koch belongs to one of the oldest families of conifers in the world, but despite this, there are few studies focused on its physiology and responses to changes in environmental conditions. We used an integrated approach aimed at comprehensively characterizing the ecophysiology of this poorly known species, focusing in its stomatal, mesophyll and biochemical traits, hypothesizing that these traits govern the carbon assimilation of A. araucana under past and present levels of atmospheric CO2. Results indicated that A. araucana presents the typical traits of an ancient species, such as large stomata and low stomatal density, which trigger low stomatal conductance and slow stomatal responsiveness to changing environmental conditions. Interestingly, the quantitative analysis showed that photosynthetic rates were equally limited by both diffusive and biochemical components. The Rubisco catalytic properties proved to have a low Rubisco affinity for CO2 and O2, similar to other ancient species. This affinity for CO2, together with the low carboxylation turnover rate, are responsible for the low Rubisco catalytic efficiency of carboxylation. These traits could be the result of the diverse environmental selective pressures that A. araucana was exposed during its diversification. The increase in measured temperatures induced an increase in stomatal and biochemical limitations, which together with a lower Rubisco affinity for CO2 could explain the low photosynthetic capacity of A. araucana in warmer conditions. [ABSTRACT FROM AUTHOR]

Rehschuh and Ruehr did find lower F SB v sb /F SB M sb in the heatwave and droughted heatwave plants; however, it did not lead to long-term damage, as F SB v sb /F SB M sb slowly recovered post-heatwave and drought. Broader questions for ecosystem response to hot drought Rehschuh and Ruehr provide excellent insight into how coniferous seedlings might respond to a heatwave-drought event; however, it is still unclear how these results might scale seasonally, and with mature trees or other plant functional types. This scientific commentary refers to 'Diverging responses of water and carbon relations during and after heat and hot drought stress in Pinus sylvestris' by Rehschuh and Ruehr (doi: 10.1093/treephys/tpab141). [Extracted from the article]

The survival and performance of urban forests are increasingly challenged by urban drought, consequently compromising the sustainability and functionality of urban vegetation. Plant–water relations largely determine species drought tolerance, yet little is known about the hydraulics of urban forest species. Here, we report the leaf hydraulic and carbon traits that govern plant growth and drought resistance, including vulnerability to embolism, hydraulic conductivity and leaf gas exchange characteristics, as well as morphological traits that are potentially linked with these physiological attributes, with the aim of guiding species selection and management in urban forests. Plant materials were collected from mature shrubs and trees on our university campus in Beijing, representing 10 woody species common to urban forests in north China. We found that the leaf embolism resistance, represented by the water potential inducing 50% loss of hydraulic conductivity (P 50), as well as the hydraulic safety margin (HSM) defined by P 50 and the water potential threshold at the inception of embolism (P 12), varied remarkably across species, but was unrelated to growth form. Likewise, stem and leaf-specific hydraulic conductivity (K stem and k l) was also highly species-specific. Leaf P 50 was positively correlated with hydraulic conductivity. However, neither P 50 nor hydraulic conductivity was correlated with leaf gas exchange traits, including maximum photosynthetic rate (A max) and stomatal conductance (g s). Plant morphological and physiological traits were not related, except for specific leaf area, which showed a negative relationship with HSM. Traits influencing plant–water transport were primarily correlated with the mean annual precipitation of species climatic niche. Overall, current common woody species in urban forest environments differed widely in their drought resistance and did not have the capacity to modify these characteristics in response to a changing climate. Species morphology provides limited information regarding physiological drought resistance. Thus, screening urban forest species based on plant physiology is essential to sustain the ecological services of urban forests. [ABSTRACT FROM AUTHOR]

Extreme drought is one of the key climatic drivers of tree mortality on a global scale. However, it remains unclear whether the drought-induced tree mortality will increase under nocturnal climate warming. Here we exposed seedlings of two wide-ranging subtropical tree species, Castanopsis sclerophylla and Schima superba , with contrasting stomatal regulation strategies to prolonged drought under ambient and elevated night-time temperature by 2 °C. We quantified the seedling survival time since drought treatment by measuring multiple leaf traits such as leaf gas exchange, predawn leaf water potential and water-use efficiency. The results showed that all seedlings in the ambient temperature died within 180 days and 167 days of drought for C. sclerophylla and S. superba , respectively. Night warming significantly shortened the survival time of C. sclerophylla , by 31 days, and S. superba by 28 days, under the drought treatment. A survival analysis further showed that seedlings under night warming suffered a 1.6 times greater mortality risk than those under ambient temperature. Further analyses revealed that night warming suppressed net leaf carbon gain in both species by increasing the nocturnal respiratory rate of S. superba across the first 120 days of drought and decreasing the photosynthetic rate of both species generally after 46 days of drought. These effects on net carbon gain were more pronounced in S. superba than C. sclerophylla. After 60 days of drought, night warming decreased the predawn leaf water potential and leaf water-use efficiency of C. sclerophylla but not S. superba. These contrasting responses are partially due to variations in stomatal control between the two species. These findings suggest that stomatal traits can regulate the response of leaf gas exchange and plant water-use to nocturnal warming during drought. This study indicates that nocturnal warming can accelerate tree mortality during drought. Night warming accelerates the mortality of two subtropical seedlings under drought. Night warming differently affects the drought response of leaf gas exchange and plant water-use between the two species due to species-specific stomatal morphological traits. Carbon metabolism changes and hydraulic damage play differential roles in driving night-warming impacts on the drought-induced mortality between the two species. [ABSTRACT FROM AUTHOR]

The objective of this study was to quantify the effect of potassium (K) supply on osmotic adjustment and drought avoidance mechanisms of Eucalyptus seedlings growing under short-term water stress. The effects of K supply on plant growth, nutritional status, leaf gas exchange parameters, leaf water potential (Ψw), leaf area (LA), stomatal density (SD), leaf carbon (C) and nitrogen (N) isotopic compositions (δ13C and δ15N ‰) and leaf C/N ratio under polyethylene glycol (PEG)-induced water deficit were measured. Under both control (non-PEG) and osmotic stress (+PEG) conditions, K supply increased plant growth, boosting dry matter yield with decreased C/N leaf ratio and δ15N ‰ values. The +PEG significantly reduced LA, plant growth, dry matter yield, Ψw, number of stomata per plant and leaf gas exchange, relative to non-PEG condition. Potassium supply alleviated osmotic-induced alterations in Eucalyptus seedlings by better regulating leaf development as well as SD, thus improving the rate of leaf gas exchange parameters, mesophyll conductance to CO2 (lower δ13C ‰ values) and water use efficiency (WUE). Consequently, K-supplied plants under drought better acclimated to osmotic stress than K-deficient plants, which in turn induced lower CO2 assimilation and dry matter yield, as well as higher leaf δ13C ‰ and δ15N ‰ values. In conclusion, management practices should seek to optimize K-nutrition to improve WUE, photosynthesis-related parameters and plant growth under water deficit conditions. [ABSTRACT FROM AUTHOR]

Drought is the most severe abiotic stress and hinders the normal growth and development of plants. Sea buckthorn (Hippophae rhamnoides Linn.) is a typical drought-resistant tree species. In this study, the leaves of the H. rhamnoides ssp. sinensis ('FN') and H. rhamnoides ssp. mongolica ('XY') were selected during drought-recovery cycles for RNA sequencing, and physiological and biochemical analyses. The results revealed that drought stress significantly decreased leaf water potential, net photosynthetic rate and stomatal conductance in both sea buckthorn subspecies. Similarly, the contents of flavone, flavonol, isoflavone and flavanone significantly decreased under drought stress in 'XY'. Conversely, in 'FN', the flavone and abscisic acid (ABA) contents were significantly higher under drought stress and recovered after rehydration. Meanwhile, 4618 and 6100 differentially expressed genes (DEGs) were identified under drought stress in 'FN' and 'XY', respectively. In total, 5164 DEGs were observed in the comparison between 'FN' and 'XY' under drought stress. This was more than the 3821 and 3387 DEGs found when comparing the subspecies under control and rehydration conditions, respectively. These DEGs were mainly associated with carotenoid biosynthesis, flavonoid biosynthesis, photosynthesis and plant hormone signal transduction. Six hub DEGs (ABCG5 , ABCG22 , ABCG32, ABCG36 , ABF2 and PYL4) were identified to respond to drought stress based on weighted gene co-expression network analysis and Basic Local Alignment Search Tool (BLAST) analysis using DroughtDB. These six DEGs were annotated to play roles in the ABA-dependent signaling pathway. Sixteen RNA sequencing results involving eight genes and similar expression patterns (12/16) were validated using quantitative real-time Polymerase Chain Reaction (PCR). The biochemical and molecular mechanisms underlying the regulation of drought responses by ABA and flavonoids in sea buckthorn were clarified. In this study, gene co-expression networks were constructed, and the results suggested that the mutual regulation of ABA and flavonoid signaling contributed to the difference in drought resistance between the different sea buckthorn subspecies. [ABSTRACT FROM AUTHOR]

Despite the economic importance of long-lived crop species in the Mediterranean Basin and their expansion to new warmer regions, their potential responses to prolonged temperature increases have not been adequately addressed. The objectives of this study were to: (i) assess leaf gas exchange responses to prolonged elevated temperature in young olive trees; (ii) evaluate some additional leaf traits such as stomatal density and size under these same conditions; and (iii) determine whether photosynthetic acclimation to temperature was apparent. A field experiment with two temperature levels was conducted using well-irrigated, potted olive trees (cvs. Arbequina, Coratina) grown in open-top chambers during the summer and early fall in two growing seasons. The temperature levels were a near-ambient control (T0) and a heated (T+) treatment (+4 °C). Maximum photosynthetic rate (A max), stomatal conductance (g s), transpiration (E) and chlorophyll fluorescence were measured. Stomatal size and density and trichome density were also determined. The A max, g s and chlorophyll fluorescence were little affected by heating. However, leaf E was higher at T+ than T0 in the summer in both seasons due in large part to the moderate increase in vapor pressure deficit that accompanied heating, and consequently water-use efficiency was reduced in heated leaves. When reciprocal temperature measurements were conducted in mid-summer of the second season, A max values of T0 and T+ leaves were higher under the temperature level at which they grew than when measured at the other temperature level, which suggests some thermal acclimation. Stomatal size and density were greater in T+ than in T0 grown leaves in some cases, which was consistent with a greater E in T+ leaves when measured at both temperature levels. These results suggest that acclimation to long-term changes in temperature must be carefully considered to help determine how olive trees will be influenced by global warming. [ABSTRACT FROM AUTHOR]

The overall coordination between gas exchanges and plant hydraulics may be affected by soil water availability and source-to-sink relationships. Here we evaluated how branch growth and mortality, leaf gas exchange and metabolism are affected in coffee (Coffea arabica L.) trees by drought and fruiting. Field-grown plants were irrigated or not, and maintained with full or no fruit load. Under mild water deficit, irrigation per se did not significantly impact growth but markedly reduced branch mortality in fruiting trees, despite similar leaf assimilate pools and water status. Fruiting increased net photosynthetic rate in parallel with an enhanced stomatal conductance, particularly in irrigated plants. Mesophyll conductance and maximum RuBisCO carboxylation rate remained unchanged across treatments. The increased stomatal conductance in fruiting trees over nonfruiting ones was unrelated to internal CO2 concentration, foliar abscisic acid (ABA) levels or differential ABA sensitivity. However, stomatal conductance was associated with higher stomatal density, lower stomatal sensitivity to vapor pressure deficit, and higher leaf hydraulic conductance and capacitance. Increased leaf transpiration rate in fruiting trees was supported by coordinated alterations in plant hydraulics, which explained the maintenance of plant water status. Finally, by preventing branch mortality, irrigation can mitigate biennial production fluctuations and improve the sustainability of coffee plantations. [ABSTRACT FROM AUTHOR]

Sexual dimorphism occurs regarding carbon and nitrogen metabolic processes in response to nitrogen supply. Differences in fixation and remobilization of carbon and allocation and assimilation of nitrogen between sexes may differ under severe defoliation. The dioecious species Populus cathayana was studied after two defoliation treatments with two N levels. Males had a higher capacity of carbon fixation because of higher gas exchange and fluorescence traits of leaves after severe long-term defoliation under deficient N. Males had higher leaf abscisic acid, stomatal conductance and leaf sucrose phosphate synthase activity increasing transport of sucrose to sinks. Males had a higher carbon sink than females, because under N-deficient conditions, males accumulated >131.10% and 90.65% root starch than males in the control, whereas females accumulated >40.55% and 52.81%, respectively, than females in the control group. Males allocated less non-protein N (NNon-p) to leaves, having higher nitrogen use efficiency (photosynthetic nitrogen use efficiency), higher glutamate dehydrogenase (GDH) and higher leaf GDH expression, even after long-term severe defoliation under deficient N. Females had higher leaf jasmonic acid concentration and NNon-p. The present study suggested that females allocated more carbon and nitrogen to defense chemicals than males after long-term severe defoliation under deficient N. [ABSTRACT FROM AUTHOR]

Elevational variations in the growing environment and sex differences in individuals drive the diversification of photosynthetic capacity of plants. However, photosynthetic response of dioecious plants to elevation gradients and the mechanisms that cause these responses are poorly understood. We measured foliar gas exchange, chlorophyll fluorescence and nitrogen allocations of male and female Seabuckthorn (Hippophae rhamnoides L.) at the elevation of 1900–3700 m above sea level (a.s.l.) on the eastern Qinghai-Tibetan Plateau, China. Male and female plants showed increased leaf photosynthetic capacity at higher elevation generally with no sex-specific difference. Photosynthetic photon flux density-saturated photosynthesis (A sat) was limited mostly by diffusional components (77 ± 1%), whereas biochemical components contributed minor limitations (22 ± 1%). Mesophyll conductance (g m) played an essential role in A sat variation, accounting for 40 ± 2% of the total photosynthetic limitations and had a significant positive correlation with A sat. Leaf nitrogen allocations to Rubisco (P R) and bioenergetics (P B) in the photosynthetic apparatus were major drivers for variations in photosynthetic nitrogen-use efficiency. The increase of these resource uptake capacities enables H. rhamnoides to maintain a high level of carbon assimilation and function efficiently to cope with the harsh conditions and shorter growing season at higher elevation. [ABSTRACT FROM AUTHOR]

The intensity of extreme heat and drought events has drastically risen in recent decades and will likely continue throughout the century. Northern forests have already seen increases in tree mortality and a lack of new recruitment, which is partially attributed to these extreme events. Boreal species, such as paper birch (Betula papyrifera) and white spruce (Picea glauca), appear to be more sensitive to these changes than lower-latitude species. Our objectives were to investigate the effects of repeated heatwaves and drought on young paper birch and white spruce trees by examining (i) responses in leaf gas exchange and plant growth and (ii) thermal acclimation of photosynthetic and respiratory traits to compare ecophysiological responses of two co-occurring, yet functionally dissimilar species. To address these objectives, we subjected greenhouse-grown seedlings to two consecutive summers of three 8-day long, +10 °C heatwaves in elevated atmospheric CO2 conditions with and without water restriction. The data show that heatwave stress reduced net photosynthesis, stomatal conductance and growth—more severely so when combined with drought. Acclimation of both photosynthesis and respiration did not occur in either species. The combination of heat and drought stress had a similar total effect on both species, but each species adjusted traits differently to the combined stress. Birch experienced greater declines in gas exchange across both years and showed moderate respiratory but not photosynthetic acclimation to heatwaves. In spruce, heatwave stress reduced the increase in basal area in both experimental years and had a minor effect on photosynthetic acclimation. The data suggest these species lack the ability to physiologically adjust to extreme heat events, which may limit their future distributions, thereby altering the composition of boreal forests. [ABSTRACT FROM AUTHOR]

A chlorophyll fluorescence (ChlF) assessment was carried out on oak seedlings (Quercus ilex L. Quercus pubescens Willd. Quercus frainetto Ten.) of Italian and Greek provenance, during the years 2017 and 2018, in a common garden in central Italy planted in 2017. This trial aimed to test the relative performances of the oak species in the perspective of assisted migration as part of the actions for the adaptation of forests to climate change. The assessment of the photosynthetic performance of the tree species included the analysis of the prompt chlorophyll fluorescence (PF) transient and the modulated reflection (MR) at 820 nm, leaf chlorophyll content, leaf gas exchange (net photosynthesis, stomatal conductance), plant growth (i.e. height) and mortality rate after 2 years from the beginning of the experiment. The assessment of the performance of the three oak species was carried out 'in vivo'. Plants were generated from seeds and exposed to several environmental factors, including changing seasonal temperature, water availability, and soil biological and physical functionality. The results of PF indicate a stable functionality of the photosynthetic system PSII (expressed as F V/ F M) across species and provenances and a decline in photochemistry functionality at the I–P phase (Δ V IP) in Q. frainetto , thus indicating a decline of the content of PSI in this species. This result was confirmed by the findings of MR analysis, with the speed of reduction and subsequent oxidation of PSI (V RED and V OX) strongly correlated to the amplitude of Δ V IP. The photosynthetic rates (net photosynthesis, P N) and growth were correlated with the parameters associated with PSI content and function, rather than those related to PSII. The low performance of Q. frainetto in the common garden seems to be related to early foliar senescence with the depletion of nitrogen, due to suboptimal climatic and edaphic conditions. Chlorophyll fluorescence allowed discrimination of populations of oak species and individuation of the less (or/and best) suitable species for future forest ecology and management purposes. [ABSTRACT FROM AUTHOR]

Photosynthetic carbon uptake by tropical forests is of critical importance in regulating the earth's climate, but rising temperatures threaten this stabilizing influence of tropical forests. Most research on how temperature affects photosynthesis focuses on fully sun-exposed leaves, and little is known about shade leaves, even though shade leaves greatly outnumber sun leaves in lowland tropical forests. We measured temperature responses of light-saturated photosynthesis, stomatal conductance, and the biochemical parameters V CMax (maximum rate of RuBP carboxylation) and J Max (maximum rate of RuBP regeneration, or electron transport) on sun and shade leaves of mature tropical trees of three species in Panama. As expected, biochemical capacities and stomatal conductance were much lower in shade than in sun leaves, leading to lower net photosynthesis rates. However, the key temperature response traits of these parameters—the optimum temperature (T Opt) and the activation energy—did not differ systematically between sun and shade leaves. Consistency in the J Max to V CMax ratio further suggested that shade leaves are not acclimated to lower temperatures. For both sun and shade leaves, stomatal conductance had the lowest temperature optimum (~25  ° C), followed by net photosynthesis (~30 °C), J Max (~34  ° C) and V CMax (~38  ° C). Stomatal conductance of sun leaves decreased more strongly with increasing vapor pressure deficit than that of shade leaves. Consistent with this, modeled stomatal limitation of photosynthesis increased with increasing temperature in sun but not shade leaves. Collectively, these results suggest that modeling photosynthetic carbon uptake in multi-layered canopies does not require independent parameterization of the temperature responses of the biochemical controls over photosynthesis of sun and shade leaves. Nonetheless, to improve the representation of the shade fraction of carbon uptake dynamics in tropical forests, better understanding of stomatal sensitivity of shade leaves to temperature and vapor pressure deficit will be required. [ABSTRACT FROM AUTHOR]

Due to its ubiquity across northern latitudes, silver birch (Betula pendula Roth) is an attractive model species for studying geographical trait variation and acclimation capacity. Six birch provenances from 60 to 67°N across Finland were grown in a common garden and studied for provenance and genotype variation. We looked for differences in height growth, photosynthetic gas exchange and chlorophyll content index (CCI) and compared the gas exchange of early and late leaves on short and long shoots, respectively. The provenances stratified into southern and northern groups. Northern provenances attained less height growth increment and had higher stomatal conductance (g s) and lower intrinsic water-use efficiency (WUE, A net/ g s) than southern provenances, whereas net photosynthesis (A net) or CCI did not show clear grouping. Short shoot leaves had lower g s and higher WUE than long shoot leaves in all provenances, but there was no difference in A net between shoot types. The separation of the provenances into two groups according to their physiological responses might reflect the evolutionary history of B. pendula. Latitudinal differences in gas exchange and water use traits can have plausible consequences for global carbon and water fluxes in a warming climate. [ABSTRACT FROM AUTHOR]

Vapour pressure deficit (D) is projected to increase in the future as temperature rises. In response to increased D , stomatal conductance (gs) and photosynthesis (A) are reduced, which may result in significant reductions in terrestrial carbon, water and energy fluxes. It is thus important for gas exchange models to capture the observed responses of gs and A with increasing D. We tested a series of coupled A – gs models against leaf gas exchange measurements from the Cumberland Plain Woodland (Australia), where D regularly exceeds 2 kPa and can reach 8 kPa in summer. Two commonly used A – gs models were not able to capture the observed decrease in A and gs with increasing D at the leaf scale. To explain this decrease in A and gs , two alternative hypotheses were tested: hydraulic limitation (i.e., plants reduce gs and/or A due to insufficient water supply) and non-stomatal limitation (i.e., downregulation of photosynthetic capacity). We found that the model that incorporated a non-stomatal limitation captured the observations with high fidelity and required the fewest number of parameters. Whilst the model incorporating hydraulic limitation captured the observed A and gs , it did so via a physical mechanism that is incorrect. We then incorporated a non-stomatal limitation into the stand model, MAESPA, to examine its impact on canopy transpiration and gross primary production. Accounting for a non-stomatal limitation reduced the predicted transpiration by ~19%, improving the correspondence with sap flow measurements, and gross primary production by ~14%. Given the projected global increases in D associated with future warming, these findings suggest that models may need to incorporate non-stomatal limitation to accurately simulate A and gs in the future with high D. Further data on non-stomatal limitation at high D should be a priority, in order to determine the generality of our results and develop a widely applicable model. [ABSTRACT FROM AUTHOR]

We quantified seasonal CO2 assimilation capacities for seven dominant vascular species in a wet boreal forest peatland then applied data to a land surface model parametrized to the site (ELM-SPRUCE) to test if seasonality in photosynthetic parameters results in differences in simulated plant responses to elevated CO2 and temperature. We collected seasonal leaf-level gas exchange, nutrient content and stand allometric data from the field-layer community (i.e. Maianthemum trifolium (L.) Sloboda), understory shrubs (Rhododendron groenlandicum (Oeder) Kron and Judd, Chamaedaphne calyculata (L.) Moench. Kalmia polifolia Wangenh. and Vaccinium angustifolium Alton.) and overstory trees (Picea mariana (Mill.) B.S.P. and Larix laricina (Du Roi) K. Koch). We found significant interspecific seasonal differences in specific leaf area, nitrogen content (by area; Na) and photosynthetic parameters (i.e. maximum rates of Rubisco carboxylation (V cmax25°C), electron transport (J max25°C) and dark respiration (R d25°C)), but minimal correlation between foliar Na and V cmax25°C, J max25°C or R d25°C, which illustrates that nitrogen alone is not a good correlate for physiological processes such as Rubisco activity that can change seasonally in this system. ELM-SPRUCE was sensitive to the introduction of observed interspecific seasonality in V cmax25°C, J max25°C and R d25°C, leading to simulated enhancement of net primary production (NPP) using seasonally dynamic parameters as compared with use of static parameters. This pattern was particularly pronounced under simulations with higher temperature and elevated CO2, suggesting a key hypothesis to address with future empirical or observational studies as climate changes. Inclusion of species-specific seasonal photosynthetic parameters should improve estimates of boreal ecosystem-level NPP, especially if impacts of seasonal physiological ontogeny can be separated from seasonal thermal acclimation. [ABSTRACT FROM AUTHOR]

Long-term high CO2 exposure accelerates photosynthetic induction response due to rapid light increase. However, it is unclear whether the acceleration is caused by acclimation of photosynthetic components (long-term CO2 effect) and/or by the sufficient substrate under high CO2 at the measurement (short-term CO2 effect). Populus koreana × trichocarpa cv. Peace has wide-open stomata almost not responding to changes of photon flux density. Using this species, we examined the long- and short-term CO2 effects on photosynthetic induction by focusing on biochemical components. We grew the plants under [CO2] of 380, 700 and 1020 μmol CO2 mol−1 air and measured the photosynthetic induction response under [CO2] of 380 and 1020 μmol CO2 mol−1 air. Despite significant reduction in Rubisco content and light-saturated photosynthetic rate in the leaves from the high growth CO2, the photosynthetic induction time was similar in leaves from different growth CO2 plants when measurement [CO2] was the same. The induction, however, was significantly fast at the higher than at the lower measurement [CO2], regardless of growth CO2 of the plants. These results demonstrate that the acceleration of apparent photosynthetic induction under high CO2 environment was mainly contributed by a short-term CO2 effect rather than by a long-term acclimation effect when stomatal limitation is not the major factor. [ABSTRACT FROM AUTHOR]

Little is known about the ability of trees to recover hydraulic conductance (k) within a growing season by regrowth or refilling of embolized conduits. Recovery of k lost to drought or other causes would prevent chronic reductions in gas exchange and productivity. To test recovery ability we conducted a 2-year experiment (2014-15) on a cohort of aspen ramets (Populus tremuloides, Michx.). Whole-tree k was measured from mid-June through September from sapflow (Q) and pre-dawn and mid-day xylem pressure. We induced embolism in the treatment group with high air pressure delivered by a split pressure chamber sealed around the main trunk. Successful treatments reduced k and Q by 50% or more without causing rapid desiccation. The majority of trees recovered following treatment, rising to control levels of k and Q between 12 and 17 days. Failure to recover was correlated with drier climate conditions. The growing-season recovery of k was attributed to refilling of embolized vessels, based on the absence of diameter growth. Pre-dawn xylem pressures during recovery were similar to the threshold needed to passively collapse emboli. Successful recovery during the 2-year study was consistent with no reduction in cumulative Q or canopy area in treatment vs controls. However, non-recovering trees in 2014 exhibited lower basal area growth at the start of the 2015 growing season, suggesting a linkage between recovery ability and productivity. This study provides evidence for the potential of trees to recover xylem function by refilling during the growing season. [ABSTRACT FROM AUTHOR]

Climate change exposes vegetation to unusual levels of drought, risking a decline in productivity and an increase in mortality. It still remains unclear how trees and forests respond to such unusual drought, particularly Southeast Asian tropical rain forests. To understand leaf ecophysiological responses of tropical rain forest trees to soil drying, a rainfall exclusion experiment was conducted on mature canopy trees of Dryobalanops aromatica Gaertn.f. (Dipterocarpaceae) for 4 months in an aseasonal tropical rain forest in Sarawak, Malaysia. The rainfall was intercepted by using a soft vinyl chloride sheet. We compared the three control and three treatment trees with respect to leaf water use at the top of the crown, including stomatal conductance (gsmax), photosynthesis (Amax), leaf water potential (predawn: Ψpre; midday: Ψmid), leaf water potential at turgor loss point (πtlp), osmotic potential at full turgor (π100) and a bulk modulus of elasticity (ε). Measurements were taken using tree-tower and canopy-crane systems. During the experiment, the treatment trees suffered drought stress without evidence of canopy dieback in comparison with the control trees; e.g., Ψpre and Ψmid decreased with soil drying. Minimum values of Ψmid in the treatment trees decreased during the experiment, and were lower than πtlp in the control trees. However, the treatment trees also decreased their πtlp by osmotic adjustment, and the values were lower than the minimum values of their Ψmid. In addition, the treatment trees maintained gs and Amax especially in the morning, though at midday, values decreased to half those of the control trees. Decreasing leaf water potential by osmotic adjustment to maintain gs and Amax under soil drying in treatment trees was considered to represent anisohydric behavior. These results suggest that D. aromatica may have high leaf adaptability to drought by regulating leaf water consumption and maintaining turgor pressure to improve its leaf water relations. [ABSTRACT FROM AUTHOR]

Predicting the impact of drought on forest ecosystem processes requires an understanding of trees' species-specific responses to drought, especially in the Eastern USA, where species composition is highly dynamic due to historical changes in land use and fire regime. Here, we adapted a framework that classifies trees' water-use strategy along the spectrum of isohydric to anisohydric behavior to determine the responses of three canopy-dominant species to drought. We used a collection of leaf-level gas exchange, tree-level sap flux and stand-level eddy covariance data collected in south-central Indiana from 2011 to 2013, which included an unusually severe drought in the summer of 2012. Our goal was to assess how patterns in the radial profile of sap flux and reliance on hydraulic capacitance differed among species of contrasting water-use strategies. In isohydric species, which included sugar maple (Acer saccharum Marsh.) and tulip poplar (Liriodendron tulipifera L.), we found that the sap flux in the outer xylem experienced dramatic declines during drought, but sap flux at inner xylem was buffered from reductions in water availability. In contrast, for anisohydric oak species (Quercus alba L. and Quercus rubra L.), we observed relatively smaller variations in sap flux during drought in both inner and outer xylem, and higher nighttime refilling when compared with isohydric species. This reliance on nocturnal refilling, which occurred coincident with a decoupling between leaf- and tree-level water-use dynamics, suggests that anisohydric species may benefit from a reliance on hydraulic capacitance to mitigate the risk of hydraulic failure associated with maintaining high transpiration rates during drought. In the case of both isohydric and anisohydric species, our work demonstrates that failure to account for shifts in the radial profile of sap flux during drought could introduce substantial bias in estimates of tree water use during both drought and non-drought periods. [ABSTRACT FROM AUTHOR]

The article focuses in a study conducted on a whole-tree model to analyze the link between leaf gas exchange, sugar utilization and soil water uptake and xylem and phloem fluxes. According to the study, the photosynthesis rate is maximized by the plant within the constraints imposed by meteorological conditions, soil water availability and phloem's sugar transport.

The occurrence of amphistomatic leaves (stomata on both surfaces) versus hypostomatic leaves (stomata limited to the lower or abaxial surface) has strong associations with environment. Amphistomy provides the advantage of higher conductance of CO2 for photosynthesis, however, unless the stomata on both leaf surfaces can be independently controlled in response to environmental cues, amphistomy may lead to inefficient gas exchange. While previous studies have found evidence that stomata can operate independently across and between surfaces of dorsiventral leaves, we investigate whether an independent stomatal response can be induced for isobilateral leaves by largely natural conditions. Here, we exposed surfaces of isobilateral, amphistomatic Eucalyptus globulus Labill. leaves to natural diurnal variation in differential evaporative demand, using leaf orientation to drive differences in irradiance and heat load on leaf surfaces. We identified preferential closure of stomata on the surface exposed to higher irradiation (and therefore evaporative demand) during the afternoon under natural conditions and similarly induced differential stomatal closure under experimental conditions in the laboratory. The differential response confirms that sufficient hydraulic isolation exists for independent stomatal response to occur between surfaces of amphistomatic, isobilateral leaves, and importantly, we show that natural conditions can induce surface-specific stomatal closure. [ABSTRACT FROM AUTHOR]

Plant surface properties influence solid-liquid interactions and matter exchange between the organs and their surrounding environment. In the case of fruits, surface processes may be of relevance for seed production and dispersal. To gain insight into the relationship between surface structure, chemical composition and function of aerial reproductive organs, we performed diverse experiments with the dry, winged fruits, or samaras, of Ulmus laevis Pall. and Ulmus minor Mill. both at the time of full maturity (green samaras) and of samara dispersal (dry samaras). Samaras of both elm species showed positive photosynthetic rates and absorbed water through their epidermal surfaces. The surface wettability, free energy, polarity and solubility parameter were lower in U. laevis than in U. minor and decreased for dry samaras in both species. Ulmus laevis samaras had a high degree of surface nano-roughness mainly conferred by cell wall folds containing pectins that substantially increased after hydration. The samaras in this species also had a thicker cuticle that could be isolated by enzymatic digestion, whereas that of U. minor samaras had higher amounts of soluble lipids. Dry samaras of U. laevis had higher floatability and lower air sustentation than those of U. minor. We concluded that samaras contribute to seed development by participating in carbon and water exchange. This may be especially important for U. minor, whose samaras develop before leaf emergence. The trichomes present along U. laevis samara margin may enhance water absorption and samara floatability even in turbulent waters. In general, U. minor samaras show traits that are consistent with a more drought tolerant character than U. laevis samaras, in line with the resources available both at the tree and ecosystem level for these species. Samara features may additionally reflect different adaptive strategies for seed dispersal and niche differentiation between species, by favoring hydrochory for U. laevis and anemochory for U. minor. [ABSTRACT FROM AUTHOR]

Dry periods are predicted to become more frequent and severe in the future in some parts of the tropics, including Amazonia, potentially causing reduced productivity, higher tree mortality and increased emissions of stored carbon. Using a long-term (12 year) through-fall exclusion (TFE) experiment in the tropics, we test the hypothesis that trees produce leaves adapted to cope with higher levels of water stress, by examining the following leaf characteristics: area, thickness, leaf mass per area, vein density, stomatal density, the thickness of palisade mesophyll, spongy mesophyll and both of the epidermal layers, internal cavity volume and the average cell sizes of the palisade and spongy mesophyll. We also test whether differences in leaf anatomy are consistent with observed differential drought-induced mortality responses among taxa, and look for relationships between leaf anatomy, and leaf water relations and gas exchange parameters. Our data show that trees do not produce leaves that are more xeromorphic in response to 12 years of soil moisture deficit. However, the drought treatment did result in increases in the thickness of the adaxial epidermis (TFE: 20.5 ± 1.5 μm, control: 16.7 ± 1.0 μm) and the internal cavity volume (TFE: 2.43 ± 0.50 mm3 cm-2, control: 1.77 ± 0.30 mm3cm-2). No consistent differences were detected between drought-resistant and drought-sensitive taxa, although interactions occurred between drought-sensitivity status and drought treatment for the palisade mesophyll thickness (P=0.034) and the cavity volume of the leaves (P=0.025). The limited response to water deficit probably reflects a tight coordination between leaf morphology, water relations and photosynthetic properties. This suggests that there is little plasticity in these aspects of plant anatomy in these taxa, and that phenotypic plasticity in leaf traits may not facilitate the acclimation of Amazonian trees to the predicted future reductions in dry season water availability. [ABSTRACT FROM AUTHOR]

The effects of warming on the temperature response of leaf photosynthesis have become an area of major concern in recent decades. Although growth temperature (GT) and day length (DL) affect leaf gas exchange characteristics, the way in which these factors influence the temperature dependence of photosynthesis remains uncertain. We established open-top canopy chambers at the canopy top of a deciduous forest, in which average daytime leaf temperature was increased by 1.0 °C. We conducted gas exchange measurements for the canopy leaves of deciduous trees exposed to artificial warming during different seasons. The carbon dioxide assimilation rate at 20 °C (A20) was not affected by warming, whereas that at 25 °C (A25) tended to be higher in leaves exposed to warming. Warming increased the optimal temperature of photosynthesis by increasing the activation energy for the maximum rate of carboxylation. Regression analysis indicated that both GT and DL strongly influenced gas exchange characteristics. Sensitivity analysis revealed that DL affected A without obvious effects on the temperature dependence of A, whereas GT almost maintained constant A20 and strongly influenced the temperature dependence. These results indicate that GT and DL have different influences on photosynthesis; GT and DL affect the 'slope' and intercept' of the temperature dependence of photosynthesis, respectively. [ABSTRACT FROM AUTHOR]

Forest functionality and productivity are directly related to canopy light interception and can be affected by potential damage from high irradiance. However, the mechanisms by which leaves adapt to the variable light environments along the multilayer canopy profile are still poorly known. We explored the leaf morphophysiological and metabolic responses to the natural light gradient in a pure European beech (Fagus sylvatica L.) forest at three different canopy heights (top, middle and bottom). Structural adjustment through light-dependent modifications in leaf mass per area was the reason for most of the variations in photosynthetic capacity. The different leaf morphology along the canopy influenced nitrogen (N) partitioning, water- and photosynthetic N-use efficiency, chlorophyll (Chl) fluorescence and quali-quantitative contents of photosynthetic pigments. The Chl a to Chl b ratio and the pool of xanthophyll-cycle pigments (VAZ) increased at the highest irradiance, as well as lutein and β-carotene. The total pool of ascorbate and phenols was higher in leaves of the top and middle canopy layers when compared with the bottom layer, where the ascorbate peroxidase was relatively more activated. The non-photochemical quenching was strongly and positively related to the VAZ/(Chl a + b) ratio, while Chl a/Chl b was related to the photochemical efficiency of photosystem II. Along the multilayer canopy profile, the high energy dissipation capacity of leaves was correlated to an elevated redox potential of antioxidants. The middle layer gave the most relevant contribution to leaf area index and carboxylation capacity of the canopy. In conclusion, a complex interplay among structural, physiological and biochemical traits drives the dynamic leaf acclimation to the natural gradients of variable light environments along the tree canopy profile. The relevant differences observed in leaf traits within the canopy positions of the beech forest should be considered for improving estimation of carbon fluxes in multilayer canopy models of temperate forests. [ABSTRACT FROM AUTHOR]

Ozone (O3) sensitivity varies greatly among plant species. Leaf traits such as stomatal conductance, antioxidant capacity and leaf morphology and anatomy may play important roles in controlling this variation, but the relative contributions of each trait remain elusive. In this study, we examined the differences in O3 sensitivity among 29 deciduous and evergreen woody species used for urban greening in China in an open-top chamber experiment. Elevated O3 caused visible injury and reductions in net photosynthesis, and these effects differed significantly among species. The deciduous species Sorbaria sorbifolia, Hibiscus syriacus and Fraxinus chinensis were the most sensitive, while evergreen species ranked among the most tolerant. O3 sensitivity was linked to both low leaf mass per area (LMA) and low leaf area-based antioxidant levels, but not to variation in leaf mass-based antioxidant levels or stomatal conductance. The well-known and easily measured leaf trait LMA thus represents a potentially useful metric for O3 risk assessment and for selecting appropriate species for urban greening in O3-polluted areas. [ABSTRACT FROM AUTHOR]

Clonal integration between ramets can be an ecological advantage of clonal plant species in environments where resources are patchily distributed. We investigated physiological integration among Populus balsamifera L. ramets under drought stress in order to demonstrate water sharing between connected ramets. Pairs of connected ramets were grown in separate pots in the greenhouse where half of ramets had the parental root connection severed and half were left intact. Drought stress was applied to one ramet, and growth, specific leaf area (SLA), net photosynthesis, stomatal conductance, leaf water potential and carbon isotopic composition (δ13C) were measured after an 8-week growing period. Droughted ramets connected to watered ramets were able to maintain high gas exchange activity and water potential, similar to watered ramets. Leaf water potential and SLA results showed that the root connection was more beneficial for proximal compared with distal ramets. The parental root connection also allowed droughted ramets to discriminate more against 13C compared with severed ramets. In conclusion, this study shows compelling evidence of physiological integration of connected P. balsamifera ramets through water sharing. [ABSTRACT FROM AUTHOR]

We employed the warm temperate conifer Cunninghamia lanceolata (Lamb.) Hook. as a model of plantation forest species to investigate ecophysiological responses to root treatments (control (0%), and ~25, 50 or 75% of the initial root mass) under well-watered and water-limited conditions. Our results indicated that total root dry mass accumulation was negatively associated with the severity of root pruning, but there was evidence of multiple compensatory responses. The plants exhibited higher instantaneous and long-term (assessed by carbon isotope composition, δ13C) water-use efficiency in pruning treatments, especially under low water availability. Root pruning also increased the fine root/total root mass ratio, specific root length and fine root vitality in both water availability treatments. As a result of the compensatory responses, under well-watered conditions, height, stem dry mass accumulation, leaf/fine root biomass ratio (L/FR), transpiration rate, photosynthetic capacity and photosynthetic nitrogen-use efficiency (EN) were the highest under 25% pruning. Yet, all these traits except L/FR and foliage nitrogen content were severely reduced under 75% pruning. Drought negatively affected growth and leaf gas exchange rates, and there was a greater negative effect on growth, water potential, gas exchange and EN when >25% of total root biomass was removed. The stem/aboveground mass ratio was the highest under 25% pruning in both watering conditions. These results indicate that the responses to root severance are related to the excision intensity and soil moisture content. A moderate root pruning proved to be an effective means to improve stem dry mass accumulation. [ABSTRACT FROM AUTHOR]

Projections of future climates suggest that droughts (Ds) may become more frequent and severe in many regions. Genetic variation, especially within populations in traits related to D resistance, is poorly investigated in forest trees, but this knowledge is necessary to better understand how forests will respond to water shortages. In this study, we investigated variability among seven open-pollinated half-sib families of a single population and two population-level progenies of Norway spruce (Picea abies (L.) H. Karst.) in their gas exchange response to imposed D and xylem vulnerability to embolism. During their third growing season, saplings were subjected to three treatments—control (C), D (for 19 weeks) and broken drought (BD, 54 days without watering starting in mid-July, then well-watered). In response to D, all families reduced their stomatal conductance (gs) and light-saturated rates of photosynthesis (Amax) in a similar way. After rewatering, the xylem water potential (Ψ) recovered in the BD treatment, but gs and Amax remained lower than in C. Needle starch concentration was altered in both D treatments compared with C. Xylem of D-exposed trees was more vulnerable to embolism than in C. The minimum attained safety margin remained positive for all families, indicating that no catastrophic hydraulic failure occurred in stem xylem during D. Significant family variation was found for Ψ early in the D (midday Ψ between -1.2 and -1.8 MPa), and for needle damage, but not for sapling mortality. Family variation found at the initial stages of D, and not afterward, suggests that all families responded similarly to greater D intensity, exhibiting the species-specific response. Limited variation at the family level indicates that the response to D and the traits we examined were conservative within the species. This may limit breeding opportunities for increased D resistance in Norway spruce in light of expected climatic changes. [ABSTRACT FROM AUTHOR]

Establishing physiological thresholds to drought-induced mortality in a range of plant species is crucial in understanding how plants respond to severe drought. Here, five common European tree species were selected (Acer campestre L., Acer pseudoplatanus L., Carpinus betulus L., Corylus avellana L. and Fraxinus excelsior L.) to study their hydraulic thresholds to mortality. Photosynthetic parameters during desiccation and the recovery of leaf gas exchange after rewatering were measured. Stem vulnerability curves and leaf pressure–volume curves were investigated to understand the hydraulic coordination of stem and leaf tissue traits. Stem and root samples from well-watered and severely drought-stressed plants of two species were observed using transmission electron microscopy to visualize mortality of cambial cells. The lethal water potential (ψlethal) correlated with stem P99 (i.e., the xylem water potential at 99% loss of hydraulic conductivity, PLC). However, several plants that were stressed beyond the water potential at 100% PLC showed complete recovery during the next spring, which suggests that the ψlethal values were underestimated. Moreover, we observed a 1 : 1 relationship between the xylem water potential at the onset of embolism and stomatal closure, confirming hydraulic coordination between leaf and stem tissues. Finally, ultrastructural changes in the cytoplasm of cambium tissue and mortality of cambial cells are proposed to provide an alternative approach to investigate the point of no return associated with plant death. [ABSTRACT FROM AUTHOR]

Recently, three types of cavitation: (i) expanding gradually; (ii) expanding-exploding, becoming a long-shaped bubble-lengthening by degrees; (iii) suddenly exploding and fully filling the conduit instantly, were proposed. Directed by this theory, experiments were performed using light microscopy to study the natural drying processes of xylem sections of Platycladus orientalis (L.) Franco. Three different phenomena of gas filling process in conduits were captured by replaying recorded videos. The first phenomenon is that a bubble emerging in a conduit expands and elongates gradually to fill the conduit. The second phenomenon is that a bubble emerging in a conduit expands gradually, and then suddenly becomes long-shaped, and extends continuously. The third phenomenon is that a bubble instantly fully fills a conduit. This paper suggests in these experiments that after losing the bulk water of a section, as the water stress of that section became more severe, the water pressures of different conduits of the section were not necessarily the same, and as time went on, the water pressures decreased constantly. Considering some practical factors, the three phenomena captured in our experiment are explained by our theory. [ABSTRACT FROM AUTHOR]

Boreal trees experience repeated freeze-thaw cycles annually. While freezing has been extensively studied in trees, the dynamic responses occurring during the freezing and thawing remain poorly understood. At freezing and thawing, rapid changes take place in the water relations of living cells in needles and in stem. While freezing is mostly limited to extracellular spaces, living cells dehydrate, shrink and their osmotic concentration increases. We studied how the freezing-thawing dynamics reflected on leaf gas exchange, chlorophyll fluorescence and xylem and living bark diameter changes of Scots pine (Pinus sylvestris L.) saplings in controlled experiments. Photosynthetic rate quickly declined following ice nucleation and extracellular freezing in xylem and needles, almost parallel to a rapid shrinking of xylem diameter, while that of living bark followed with a slightly longer delay. While xylem and living bark diameters responded well to decreasing temperature and water potential of ice, the relationship was less consistent in the case of increasing temperature. Xylem showed strong temporal swelling at thawing suggesting water movement from bark. After thawing xylem diameter recovered to a pre-freezing level but living bark remained shrunk. We found that freezing affected photosynthesis at multiple levels. The distinct dynamics of photosynthetic rate and stomatal conductance reveals that the decreased photosynthetic rate reflects impaired dark reactions rather than stomatal closure. Freezing also inhibited the capacity of the light reactions to dissipate excess energy as heat, via non-photochemical quenching, whereas photochemical quenching of excitation energy decreased gradually with temperature in agreement with the gas exchange data. [ABSTRACT FROM AUTHOR]

First-year tree seedlings represent a particularly vulnerable life stage and successful seedling establishment is crucial for forest regeneration. We investigated the extent to which Pinus ponderosa P. & C. Lawson populations from different climate zones exhibit differential expression of functional traits that may facilitate their establishment. Seeds from two populations from sites with contrasting precipitation and temperature regimes east (PIPOdry) and west (PIPOmesic) of the Oregon Cascade mountains were sown in a common garden experiment and grown under two water availability treatments (control and drought). Aboveground biomass accumulation, vegetative phenology, xylem anatomy, plant hydraulic architecture, foliar stable carbon isotope ratios (δ13C), gas exchange and leaf water relations characteristics were measured. No treatment or population-related differences in leaf water potential were detected. At the end of the first growing season, aboveground biomass was 74 and 44% greater in PIPOmesic in the control and drought treatments, respectively. By early October, 73% of PIPOdry seedlings had formed dormant buds compared with only 15% of PIPOmesic seedlings. Stem theoretical specific conductivity, calculated from tracheid dimensions and packing density, declined from June through September and was nearly twice as high in PIPOmesic during most of the growing season, consistent with measured values of specific conductivity. Intrinsic water-use efficiency based on δ13C values was higher in PIPOdry seedlings for both treatments across all sampling dates. There was a negative relationship between values of δ13C and leaf-specific hydraulic conductivity across populations and treatments, consistent with greater stomatal constraints on gas exchange with declining seedling hydraulic capacity. Integrated growing season assimilation and stomatal conductance estimated from foliar δ13C values and photosynthetic CO2-response curves were 6 and 28% lower, respectively, in PIPOdry seedlings. Leaf water potential at the turgor loss point was 0.33 MPa more negative in PIPOdry, independent of treatment. Overall, PIPOdry seedlings exhibited more conservative behavior, suggesting reduced growth is traded off for increased resistance to drought and extreme temperatures. [ABSTRACT FROM AUTHOR]

Stable oxygen isotope techniques may be a useful tool to investigate the pathways of water movement within leaves. However, implementation of such methods is limited due to uncertainty in the effective path length (L) for the Péclet effect in leaf water enrichment models. Previous studies have found relationships between L and physiological parameters such as transpiration rate (E) and leaf hydraulic conductance (kleaf) both within and between species. However, these studies relied on assumptions in their calculation of L, which were not directly tested. Eucalyptus paniculata Smith was used to evaluate the relationships between L, kleaf and E under differing water availability and a range of leaf temperatures. Coupled gas exchange and transpiration isotope measurements allowed previous assumptions to be directly tested. L was significantly and negatively related to both kleaf and E when the isotopic signature of water vapour was assumed to be in equilibrium with source water, was equivalent to the room vapour or equal to source water. However, the relationship between L and kleaf was non-significant when measured δ 18O of transpired vapour was used and disappeared entirely when non-steady-state leaves were excluded, and when evaporation site water was calculated from coupled gas exchange and transpiration isotope values. These results suggest that great care must be taken when calculating L, particularly regarding assumptions of isotopic steady state and δ 18O of vapour. Previous suggestions of changes in pathways for water movement as transpiration rate varied need to be reassessed in light of these observations. [ABSTRACT FROM AUTHOR]

The frequency of extreme weather has been rising in recent years. A 3-year study of street trees was undertaken in Tokyo to determine whether: (i) street trees suffer from severe water stress in unusually hot summer; (ii) species respond differently to such climatic fluctuations; and (iii) street trees are also affected by nitrogen (N) deficiency, photoinhibition and aerosol pollution. During the study period (2010-12), midsummers of 2010 and 2012 were unusually hot (2.4-2.8 °C higher maximum temperature than the long-term mean) and dry (6-56% precipitation of the mean). In all species, street trees exhibited substantially decreased photosynthetic rate in the extremely hot summer in 2012 compared with the average summer in 2011. However, because of a more conservative stomatal regulation (stomatal closure at higher leaf water potential) in the hot summer, apparent symptoms of hydraulic failure were not observed in street trees even in 2012. Compared with Prunus × yedoensis and Zelkova serrata, Ginkgo biloba, a gymnosperm, was high in stomatal conductance and midday leaf water potential even under street conditions in the unusually hot summer, suggesting that the species had higher drought resistance than the other species and was less susceptible to urban street conditions. This lower susceptibility might be ascribed to the combination of higher soil-to-leaf hydraulic conductance and more conservative water use. Aside from meteorological conditions, N deficiency affected street trees significantly, whereas photoinhibition and aerosol pollution had little effect. The internal CO² and δ1³C suggested that both water and N limited the net photosynthetic rate of street trees simultaneously, but water was more limiting. From these results, we concluded that the potential risk of hydraulic failure caused by climatic extremes could be low in urban street trees in temperate regions. However, the size of the safety margin might be different between species. [ABSTRACT FROM AUTHOR]

Xylem hydraulic conductance varies in response to changes in sap solute content, and in particular of potassium (K+) ion concentration. This phenomenon, known as the ‘ionic effect’, is enhanced in embolized stems, where it can compensate for cavitation-induced loss of hydraulic conductance. Previous studies have shown that in well-watered laurel plants (Laurus nobilis L.), potassium concentration of the xylem sap and plant hydraulic conductance increased 24 h after fertilization with KCl. The aim of this work was to test whether water-stressed laurel plants, grown under low potassium availability, could recover earlier from stress when irrigated with a KCl solution instead of potassium-free water. Two-year-old potted laurel seedlings were subjected to water stress by suspending irrigation until leaf conductance to water vapour (gL) dropped to ∼30% of its initial value and leaf water potential (ψL) reached the turgor loss point (ψTLP). Plants were then irrigated either with water or with 25 mM KCl and monitored for water status, gas exchange and plant hydraulics recovery at 3, 6 and 24 h after irrigation. No significant differences were found between the two experimental groups in terms of ψL, gL, plant transpiration, plant hydraulic conductance or leaf-specific shoot hydraulic conductivity. Analysis of xylem sap potassium concentration showed that there were no significant differences between treatments, and potassium levels were similar to those of potassium-starved but well-watered plants. In conclusion, potassium uptake from the soil solution and/or potassium release to the xylem appeared to be impaired in water-stressed plants, at least up to 24 h after relief from water stress, so that fertilization after the onset of stress did not result in any short-term advantage for recovery from drought. [ABSTRACT FROM AUTHOR]

The article discusses the study of Karst and colleagues on the effects of stress and root sugar concentrations on exudation rates in trees. The authors mention that the study was able to demonstrate that specific exudation rates can be correlated with photosynthesis during abiotic stress. The authors discuss the importance that the models used in the study should include exudation and not assumptions.

Dwarf mistletoes, obligate, parasitic plants with diminutive aerial shoots, have long-term effects on host tree water relations, hydraulic architecture and photosynthetic gas exchange and can eventually induce tree death. To investigate the long-term (1886–2010) impacts of dwarf mistletoe on the growth and gas exchange characteristics of host western hemlock, we compared the diameter growth and tree-ring cellulose stable carbon (C) and oxygen (O) isotope ratios (δ13Ccell, δ18Ocell) of heavily infected and uninfected trees. The relative basal area growth of infected trees was significantly greater than that of uninfected trees in 1886–90, but declined more rapidly in infected than uninfected trees through time and became significantly lower in infected than uninfected trees in 2006–10. Infected trees had significantly lower δ13Ccell and δ18Ocell than uninfected trees. Differences in δ18Ocell between infected and uninfected trees were unexpected given that stomatal conductance and environmental variables that were expected to influence the δ18O values of leaf water were similar for both groups. However, estimates of mesophyll conductance (gm) were significantly lower and estimates of effective path length for water movement (L) were significantly higher in leaves of infected trees, consistent with their lower values of δ18Ocell. This study reconstructs the long-term physiological responses of western hemlock to dwarf mistletoe infection. The long-term diameter growth and δ13Ccell trajectories suggested that infected trees were growing faster than uninfected trees prior to becoming infected and subsequently declined in growth and leaf-level photosynthetic capacity compared with uninfected trees as the dwarf mistletoe infection became severe. This study further points to limitations of the dual-isotope approach for identifying sources of variation in δ13Ccell and indicates that changes in leaf internal properties such as gm and L that affect δ18Ocell must be considered. [ABSTRACT FROM PUBLISHER]

The growth rate of triploid European aspen (Populus tremula L.) and hybrid aspen (P. tremula × Populus tremuloides Michx.) significantly exceeds that of diploid aspen, but the underlying physiological controls of the superior growth rates of these genotypes are not known. We tested the hypothesis that the superior growth rate of triploid and hybrid aspen reflects their greater net photosynthesis rate. Micropropagated clonal plants varying in age from 2.5 to 19 months were used to investigate the ploidy and plant age interaction. The quantum yield of net CO2 fixation (Φ) in leaves of young 2.5-month-old hybrid aspen was lower than that of diploid and triploid trees. However, Φ in 19-month-old hybrid aspen was equal to that in triploid aspen and higher than that in diploid aspen. Φ and the rate of light-saturated net photosynthesis (ANS) increased with plant age, largely due to higher leaf dry mass per unit area in older plants. ANS in leaves of 19-month-old trees was highest in hybrid, medium in triploid and lowest in diploid aspen. Light-saturated photosynthesis had a broad temperature optimum between 20 and 35 °C. Rate of respiration in the dark (RDS) did not vary among the genotypes in 2.5-month-old plants, and the shape of the temperature response was also similar. RDS increased with plant age, but RDS was still not significantly different among the leaves of 19-month-old diploid and triploid aspen, but it was significantly lower in leaves of 19-month-old hybrid plants. The initial differences in the growth of plants with different ploidy were minor up to the age of 19 months, but during the next 2 years, the growth rate of hybrid aspen exceeded that of triploid plants by 2.7 times and of diploid plants by five times, in line with differences in ANS of 19-month-old plants of these species. It is suggested that differences in photosynthesis and growth became more pronounced with tree aging, indicating that ontogeny plays a key role in the expression of superior traits determining the productivity of given genotypes. [ABSTRACT FROM PUBLISHER]

Boron (B) deficiency is a widespread problem in many crops, including Citrus. The effects of B-deficiency on gas exchange, carbohydrates, organic acids, amino acids, total soluble proteins and phenolics, and the activities of key enzymes involved in organic acid and amino acid metabolism in ‘Xuegan’ [Citrus sinensis (L.) Osbeck] leaves and roots were investigated. Boron-deficient leaves displayed excessive accumulation of nonstructural carbohydrates and much lower CO2 assimilation, demonstrating feedback inhibition of photosynthesis. Dark respiration, concentrations of most organic acids [i.e., malate, citrate, oxaloacetate (OAA), pyruvate and phosphoenolpyruvate] and activities of enzymes [i.e., phosphoenolpyruvate carboxylase (PEPC), NAD-malate dehydrogenase, NAD-malic enzyme (NAD-ME), NADP-ME, pyruvate kinase (PK), phosphoenolpyruvate phosphatase (PEPP), citrate synthase (CS), aconitase (ACO), NADP-isocitrate dehydrogenase (NADP-IDH) and hexokinase] involved in glycolysis, the tricarboxylic acid (TCA) cycle and the anapleurotic reaction were higher in B-deficient leaves than in controls. Also, total free amino acid (TFAA) concentration and related enzyme [i.e., NADH-dependent glutamate 2-oxoglutarate aminotransferase (NADH-GOGAT) and glutamate OAA transaminase (GOT)] activities were enhanced in B-deficient leaves. By contrast, respiration, concentrations of nonstructural carbohydrates and three organic acids (malate, citrate and pyruvate), and activities of most enzymes [i.e., PEPC, NADP-ME, PK, PEPP, CS, ACO, NAD-isocitrate dehydrogenase, NADP-IDH and hexokinase] involved in glycolysis, the TCA cycle and the anapleurotic reaction, as well as concentration of TFAA and activities of related enzymes (i.e., nitrate reductase, NADH-GOGAT, glutamate pyruvate transaminase and glutamine synthetase) were lower in B-deficient roots than in controls. Interestingly, leaf and root concentration of total phenolics increased, whereas that of total soluble protein decreased, in response to B-deficiency. In conclusion, respiration, organic acid (i.e., glycolysis and the TCA cycle) metabolism, the anapleurotic pathway and amino acid biosynthesis were upregulated in B-deficient leaves with excessive accumulation of carbohydrates to ‘consume’ the excessive carbon available, but downregulated in B-deficient roots with less accumulation of carbohydrates to maintain the net carbon balance. [ABSTRACT FROM PUBLISHER]

Gas exchange and hydraulic features were measured in leaves of three different Citrus species (Citrus aurantium L., Citrus limon L., Citrus × paradisii Macfad) infested by Phyllocnistis citrella Staiton, with the aim to quantify the impact of this pest on leaf hydraulics and, ultimately, on plant fitness. Infested leaves were characterized by the presence on the leaf blade of typical snake-shaped mines and, in some cases, of a crumpled leaf blade. Light microscopy showed that leaf crumpling was induced by damage to the cuticular layer. In all three Citrus species examined: (a) the degree of infestation did not exceed 10% of the total surface area of infested plants; (b) control and infested leaves showed similar values of minimum diurnal leaf water potential, leaf hydraulic conductance and functional vein density; and (c) maximum diurnal values of stomatal conductance to water vapour, transpiration rate and photosynthetic rate (An) were similar in both control leaves and the green areas of infested leaves. A strong reduction of An was recorded only in mined leaf areas. Our data suggest that infestation with P. citrella does not cause conspicuous plant productivity reductions in young Citrus plants, at least not in the three Citrus species studied here. [ABSTRACT FROM AUTHOR]

Rates of tissue-level function have been hypothesized to decline as trees grow older and larger, but relevant evidence to assess such changes remains limited, especially across a wide range of sizes from saplings to large trees. We measured functional traits of leaves and twigs of three cold-temperate deciduous tree species in Minnesota, USA, to assess how these vary with tree height. Individuals ranging from 0.13 to 20 m in height were sampled in both relatively open and closed canopy environments to minimize light differences as a potential driver of size-related differences in leaf and twig properties. We hypothesized that (H1) gas-exchange rates, tissue N concentration and leaf mass per unit area (LMA) would vary with tree size in a pattern reflecting declining function in taller trees, yet maintaining (H2) bivariate trait relations, common among species as characterized by the leaf economics spectrum. Taking these two ideas together yielded a third, integrated hypothesis that (H3) nitrogen (N) content and gas-exchange rates should decrease monotonically with tree size and LMA should increase. We observed increasing LMA and decreasing leaf and twig Rd with increasing size, which matched predictions from H1 and H3. However, opposite to our predictions, leaf and twig N generally increased with size, and thus had inverse relations with respiration, rather than the predicted positive relations. Two exceptions were area-based leaf N of Prunus serotina Ehrh. in gaps and mass-based leaf N of Quercus ellipsoidalis E. J. Hill in gaps, both of which showed qualitatively hump-shaped patterns. Finally, we observed hump-shaped relationships between photosynthetic capacity and tree height, not mirroring any of the other traits, except in the two cases highlighted above. Bivariate trait relations were weak intra-specifically, but were generally significant and positive for area-based traits using the pooled dataset. Results suggest that different traits vary with tree size in different ways that are not consistent with a universal shift towards a lower ‘return on investment’ strategy. Instead, species traits vary with size in patterns that likely reflect complex variation in water, light, nitrogen and carbon availability, storage and use. [ABSTRACT FROM AUTHOR]

In the present study, biogenic volatile organic compound (BVOC) emissions and photosynthetic gas exchange of salt-sensitive (Populus x canescens (Aiton) Sm.) and salt-tolerant (Populus euphratica Oliv.) isoprene-emitting and non-isoprene-emitting poplars were examined under controlled high-salinity and high-temperature and -light episode (‘sunfleck’) treatments. Combined treatment with salt and sunflecks led to an increased isoprene emission capacity in both poplar species, although the photosynthetic performance of P. × canescens was reduced. Indeed, different allocations of isoprene precursors between the cytosol and the chloroplast in the two species were uncovered by means of 13CO2 labeling. Populus × canescens leaves, moreover, increased their use of ‘alternative’ carbon (C) sources in comparison with recently fixed C for isoprene biosynthesis under salinity. Our studies show, however, that isoprene itself does not have a function in poplar survival under salt stress: the non-isoprene-emitting leaves showed only a slightly decreased photosynthetic performance compared with wild type under salt treatment. Lipid composition analysis revealed differences in the double bond index between the isoprene-emitting and non-isoprene-emitting poplars. Four clear metabolomics patterns were recognized, reflecting systemic changes in flavonoids, sterols and C fixation metabolites due to the lack/presence of isoprene and the absence/presence of salt stress. The studies were complemented by long-term temperature stress experiments, which revealed the thermotolerance role of isoprene as the non-isoprene-emitting leaves collapsed under high temperature, releasing a burst of BVOCs. Engineered plants with a low isoprene emission potential might therefore not be capable of resisting high-temperature episodes. [ABSTRACT FROM PUBLISHER]

Woody species hydraulically vulnerable to xylem cavitation may experience daily xylem embolism. How such species cope with the possibility of accumulated embolism is unclear. In this study, we examined seven temperate woody species to assess the hypothesis that low cavitation resistance (high vulnerability to cavitation) is compensated by high recovery performance via vessel refilling. We also evaluated leaf functional and xylem structural traits. The xylem recovery index (XRI), defined as the ratio of xylem hydraulic conductivity in plants rewatered after soil drought to that in plants under moist conditions, varied among species. The xylem water potential causing 50% loss of hydraulic conductivity (Ψ50) varied among the species studied, whereas only a slight difference was detected with respect to midday xylem water potential (Ψmin), indicating smaller hydraulic safety margins (Ψmin - Ψ50) for species more vulnerable to cavitation. Cavitation resistance (|Ψ50|) was negatively correlated with XRI across species, with cavitation-vulnerable species showing a higher performance in xylem recovery. Wood density was positively correlated with cavitation resistance and was negatively correlated with XRI. These novel results reveal that coordination exists between cavitation resistance and xylem recovery performance, in association with wood functional traits such as denser wood for cavitation-resistant xylem and less-dense but water-storable wood for refillable xylem. These findings provide insights into long-term maintenance of water transport in tree species growing under variable environmental conditions. [ABSTRACT FROM AUTHOR]

Predawn leaf water potential (LWP), the LWP between 09:00 and 10:30 h (termed minimum LWP), stem xylem hydraulic conductivity, foliar nitrogen, leaf gas exchange and leaf traits were measured on the same days in adults and 1-year-old to 7-year-old resprouts that had regrown after removing all the aboveground shoots. Height growth and accumulation of aboveground biomass quickly decreased with resprout age and there was no difference between 7-year-old resprouts and the uncut adults. Predawn LWP showed no significant difference between resprouts and adults, but the minimum LWP decreased gradually from −2.0 MPa in 1-year-old resprouts to −3.0 MPa in 7-year-old resprouts. The decrease in minimum LWP was associated with increased hydraulic resistance, as indicated by the gradual decrease in leaf area-specific hydraulic conductivity (KL) and sapwood area-specific hydraulic conductivity (KS) and the associated increase in stem native percentage loss of hydraulic conductivity in older than 2-year-old resprouts. The leaf nitrogen content per unit area (Narea) also decreased steadily from 3.6 g m−2 in 1-year-old resprouts to 1.7 g m−2 in 7-year-old resprouts. With the decline in LWP and Narea, the rate of leaf photosynthesis per unit area (Aarea) decreased from 20 μ mol m−2 s−1 in 1-year-old resprouts to 11 μ mol m−2 s−1 in 7-year-old resprouts. In adults, although KS decreased further compared with 7-year-old resprouts, the minimum LWP, KL, Narea and the rate of photosynthesis increased by 0.3 MPa, 29, 34 and 23%, respectively. The results show that a progressive loss of stem hydraulic conductivity and a steady decrease in foliar nitrogen with age were associated with a decrease in the photosynthetic rate of Caragana korshinskii Kom. resprouts, possibly changing the allocation of photosynthetic assimilates and slowing resprout height growth. [ABSTRACT FROM AUTHOR]

In this article the author discusses patterns of stomatal conductance in response to long-term environmental changes. The author states that the models of stomatal conductance based on the old concepts of multiplicative constraints and semi-mechanistic parameterizations. The author further discusses the Bayesian Hierarchical model of stomatal conductance and two measurement patterns of estimating stomatal conductance including gas exchange and sap flux.

Northern forests are currently experiencing increasing mean temperatures, especially during autumn and spring. Consequently, alterations in carbon sequestration, leaf biochemical quality and freezing tolerance (FT) are likely to occur. The interactive effects of elevated temperature and ozone (O3), the most harmful phytotoxic air pollutant, on Norway spruce (Picea abies (L.) Karst.) seedlings were studied by analysing phenology, metabolite concentrations in the needles, FT and gas exchange. Sampling was performed in September and May. The seedlings were exposed to a year-round elevated temperature (+1.3 °C), and to 1.4× ambient O3 concentration during the growing season in the field. Elevated temperature increased the concentrations of amino acids, organic acids of the citric acid cycle and some carbohydrates, and reduced the concentrations of phenolic compounds, some organic acids of the shikimic acid pathway, sucrose, cyclitols and steroids, depending on the timing of the sampling. Although growth onset occurred earlier at elevated temperature, the temperature of 50% lethality (LT50) was similar in the treatments. Photosynthesis and the ratio of photosynthesis to dark respiration were reduced by elevated temperature. Elevated concentrations of O3 reduced the total concentration of soluble sugars, and tended to reduce LT50 of the needles in September. These results show that alterations in needle chemical quality can be expected at elevated temperatures, but the seedlings' sensitivity to autumn and spring frosts is not altered. Elevated O3 has the potential to disturb cold hardening of Norway spruce seedlings in autumn, and to alter the water balance of the seedling through changes in stomatal conductance (gs), while elevated temperature is likely to reduce gs and consequently reduce the O3-flux inside the leaves. [ABSTRACT FROM AUTHOR]