Midday stomatal conductance is more related to stem rather than leaf water status in subtropical deciduous and evergreen broadleaf trees

Midday depressions in stomatal conductance (g_s) and photosynthesis are common in plants. The aim of this study was to understand the hydraulic determinants of midday g_s, the coordination between leaf and stem hydraulics and whether regulation of midday g_s differed between deciduous and evergreen broadleaf tree species in a subtropical cloud forest of Southwest (SW) China. We investigated leaf and stem hydraulics, midday leaf and stem water potentials, as well as midday g_s of co‐occurring deciduous and evergreen tree species. Midday g_s was correlated positively with midday stem water potential across both groups of species, but not with midday leaf water potential. Species with higher stem hydraulic conductivity and greater daily reliance on stem hydraulic capacitance were able to maintain higher stem water potential and higher g_s at midday. Deciduous species exhibited significantly higher stem hydraulic conductivity, greater reliance on stem capacitance, higher stem water potential and g_s at midday than evergreen species. Our results suggest that midday g_s is more associated with midday stem than with leaf water status, and that the functional significance of stomatal regulation in these broadleaf tree species is probably for preventing stem xylem dysfunction.     

Water Balance of Birch and Larch Leaves and Their Resistance to Short and Progressive Soil Drought

In a greenhouse experiment, which imitated a short (4-day-long) and progressive (3-week-long) soil drought, an infrared gas analyzer was employed to assess transpiration (mol/(m² s)) and leaf transpiration conductivity (g _w, mol H₂O/(m² s)) in intact one-year-old plants of Betula platyphylla Suk. and Larix gmelini (Rupr.) Rupr. grown at the saturation levels of photosynthetically active radiation. The drought duration did not affect the adaptation of leaf water balance, it was determined only by leaf temperature and water supply. At ample water supply and with all other conditions being equal, the transpiration of birch exceeded that of larch by the factor of 3; however, birch was less tolerant to soil drought. The authors conclude that water supply and leaf temperature determine plant resistance to soil drought. The species-specific drought resistance depends on the g _w value.     

Photosynthesis,transpiration, and water use efficiency of <Emphasis Type="Italic">Leymus dasystachys</Emphasis> on the Hunshandake desert

Net photosynthetic rate (P _N), transpiration rate (E), and stomatal conductance (g _s) declined from upper leaves to the lower ones during dry and rainy seasons, indicating that long-term carbon budget should take into account P _N variations for different leaf types. Relatively greater P _N in the dry season suggested that this species is more able to maintain higher P _N under drought, but the relatively higher E in the dry season might reduce water use efficiency (P _N/E) for the species. Significant correlations between P _N and g _s indicated that g _s may be the critical factor for P _N variability in the desert region.     

The response of stomatal conductance to seasonal drought in tropical forests

Stomata regulate CO₂ uptake for photosynthesis and water loss through transpiration. The approaches used to represent stomatal conductance (g_s) in models vary. In particular, current understanding of drivers of the variation in a key parameter in those models, the slope parameter (i.e. a measure of intrinsic plant water‐use‐efficiency), is still limited, particularly in the tropics. Here we collected diurnal measurements of leaf gas exchange and leaf water potential (Ψ_leaf), and a suite of plant traits from the upper canopy of 15 tropical trees in two contrasting Panamanian forests throughout the dry season of the 2016 El Niño. The plant traits included wood density, leaf‐mass‐per‐area (LMA), leaf carboxylation capacity (V_c,max), leaf water content, the degree of isohydry, and predawn Ψ_leaf. We first investigated how the choice of four commonly used leaf‐level g_s models with and without the inclusion of Ψ_leaf as an additional predictor variable influence the ability to predict g_s, and then explored the abiotic (i.e. month, site‐month interaction) and biotic (i.e. tree‐species‐specific characteristics) drivers of slope parameter variation. Our results show that the inclusion of Ψ_leaf did not improve model performance and that the models that represent the response of g_s to vapor pressure deficit performed better than corresponding models that respond to relative humidity. Within each g_s model, we found large variation in the slope parameter, and this variation was attributable to the biotic driver, rather than abiotic drivers. We further investigated potential relationships between the slope parameter and the six available plant traits mentioned above, and found that only one trait, LMA, had a significant correlation with the slope parameter (R² = 0.66, n = 15), highlighting a potential path towards improved model parameterization. This study advances understanding of g_s dynamics over seasonal drought, and identifies a practical, trait‐based approach to improve modeling of carbon and water exchange in tropical forests.     

Why is liana abundance low in semiarid climates?

Lianas are abundant in seasonal tropical forests, where they avoid seasonal water stress presumably by accessing deep‐soil water reserves. Although lianas are favoured in seasonal environments, their occurrence and abundance are low in semiarid environments. We hypothesized that lianas do not tolerate the great water shortage in the soil and air characteristic of semiarid environments, which would increase the risk of embolism. We compared the rooting depth of coarse roots, leaf dynamics, leaf water potential (ψ_leaf), embolism resistance (P₅₀) and lethal levels of embolism (P₈₈) between congeneric lianas that occur with different abundances in two semiarid sites differing in soil characteristics and vapour pressure deficit in the air (VPD_air). Regardless of soil texture and depth, water availability was restricted to the rainy season. All liana species were drought deciduous and had superficial coarse roots (not deeper than 35 cm). P₅₀ varied from ?1.8 to ?2.49 MPa, and all species operated under narrow safety margins against catastrophic (P₅₀) and irreversible hydraulic failure (P₈₈), even during the rainy season. In short, lianas that occur in semiarid environments have lower resistance to cavitation and limit carbon fixation to the rainy season because of leaf fall in the early dry season. We suggest that leaf shedding and shallow roots impairing carbon gain and growth in the dry season may explain why liana abundance is lower in semiarid than in other seasonally dry environments.     

Water relations of Caatinga trees in the dry season

The Caatinga is one of the world's richest dry forests. This forest occurs only in Brazil, but is the least studied and protected Brazilian ecosystem. There are few reports about drought tolerance mechanisms in Caatinga trees. This work evaluates water relations of six adult species in the middle of the dry season to further understand water relations in this ecosystem, which will be important for future reforestation and management projects. Based on results, the trees were classified into four groups: (I), Mimosa caesalpiniifolia had low leaf water potential (Ψ_w) at predawn and no significant decrease at midday. Stomatal conductance (g_s) analyses indicates that plants have reached its lowest Ψ_w; (II), Caesalpinia pyramidalis and Auxemma oncocalyx had low Ψ_w at predawn and significant decrease at midday. For these species the recuperation of water status at night may have been sufficient for maintaining stomata open during the day; (III), Caesalpinia ferrea and Calliandra spinosa had relatively high Ψ_w at predawn and a significant decrease at midday. These species might maintain their water status similar to individuals of group II, but they might also have deeper root systems; and (IV), Tabebuia caraiba with the highest Ψ_w at predawn and no significant decrease at midday, possibly indicating a combination of good stomatal control of water loss and a deeper root system. Moreover, except for individuals of group I, both in species with lower and higher Ψ_w at predawn it was not observed strong inhibition of g_s.     

Stem water storage capacity and efficiency of water transport: their functional significance in a Hawaiian dry forest

We investigated the contribution of internal water storage and efficiency of water transport to the maintenance of water balance in six evergreen tree species in a Hawaiian dry forest. Wood‐saturated water content, a surrogate for relative water storage capacity, ranged from 70 to 105%, and was inversely related to its morphological correlate, wood density, which ranged between 0·51 and 0·65 g cm^?3. Leaf‐specific conductivity (k_L) measured in stem segments from terminal branches ranged from 3 to 18 mmol m^?1 s^?1 MPa^?1, and whole‐plant hydraulic efficiency calculated as stomatal conductance (g) divided by the difference between predawn and midday leaf water potential (Ψ_L), ranged from 70 to 150 mmol m^?2 s^?1 MPa^?1. Hydraulic efficiency was positively correlated with k_L (r² = 0·86). Minimum annual Ψ_L ranged from ? 1·5 to ? 4·1 MPa among the six species. Seasonal and diurnal variation in Ψ_L were associated with differences among species in wood‐saturated water content, wood density and k_L. The species with higher wood‐saturated water content were more efficient in terms of long‐distance water transport, exhibited smaller diurnal variation in Ψ_L and higher maximum photosynthetic rates. Smaller diurnal variation in Ψ_L in species with higher wood‐saturated water content, k_L and hydraulic efficiency was not associated with stomatal restriction of transpiration when soil water deficit was moderate, but avoidance of low minimum seasonal Ψ_L in these species was associated with a substantial seasonal decline in g. Low seasonal minimum Ψ_L in species with low k_L, hydraulic efficiency, and wood‐saturated water content was associated with higher leaf solute content and corresponding lower leaf turgor loss point. Despite the species‐specific differences in leaf water relations characteristics, all six evergreen tree species shared a common functional relationship defined primarily by k_L and stem water storage capacity.     

Genotypic variation in transpiration of coppiced poplar during the third rotation of a short‐rotation bio‐energy culture

The productivity of short‐rotation coppice (SRC) plantations with poplar (Populus spp.) strongly depends on soil water availability, which limits the future development of its cultivation, and makes the study of the transpirational water loss particularly timely under the ongoing climate change (more frequent drought and floods). This study assesses the transpiration at different scales (leaf, tree and stand) of four poplar genotypes belonging to different species and from a different genetic background grown under an SRC regime. Measurements were performed for an entire growing season during the third year of the third rotation in a commercial scale multigenotype SRC plantation in Flanders (Belgium). Measurements at leaf level were performed on specific days with a contrasted evaporative demand, temperature and incoming shortwave radiation and included stomatal conductance, stem and leaf water potential. Leaf transpiration and leaf hydraulic conductance were obtained from these measurements. To determine the transpiration at the tree level, single‐stem sap flow using the stem heat balance (SHB) method and daily stem diameter variations were measured during the entire growing season. Sap flow‐based canopy transpiration (E_c), seasonal dry biomass yield, and water use efficiency (WUE; g aboveground dry matter/kg water transpired) of the four poplar genotypes were also calculated. The genotypes had contrasting physiological responses to environmental drivers and to soil conditions. Sap flow was tightly linked to the phenological stage of the trees and to the environmental variables (photosynthetically active radiation and vapor pressure deficit). The total E_c for the 2016 growing season was of 334, 350, 483 and 618 mm for the four poplar genotypes, Bakan, Koster, Oudenberg and Grimminge, respectively. The differences in physiological traits and in transpiration of the four genotypes resulted in different responses of WUE.     

Biomass production and water use efficiency in perennial grasses during and after drought stress

Drought is a great challenge to agricultural production, and cultivation of drought‐tolerant or water use‐efficient cultivars is important to ensure high biomass yields for bio‐refining and bioenergy. Here, we evaluated drought tolerance of four C₃ species, Dactylis glomerata cvs. Sevenop and Amba, Festuca arundinacea cvs. Jordane and Kora, Phalaris arundinacea cvs. Bamse and Chieftain and Festulolium pabulare cv. Hykor, and two C₄ species Miscanthus × giganteus and M. lutarioriparius. Control (irrigated) and drought‐treated plants were grown on coarse and loamy sand in 1 m² lysimeter plots where rain was excluded. Drought periods started after harvest and lasted until 80% of available soil water had been used. Drought caused a decrease in dry matter yield (DM; P < 0.001) for all species and cultivars during the drought period. Cultivars Sevenop, Kora and Jordane produced DM at equal levels and higher than the other C₃ cultivars in control and drought‐treated plots both during and after the drought period. Negative correlations were observed between stomatal conductance (g_s) and leaf water potential (P < 0.01) and positive correlations between g_s and DM (P < 0.05) indicating that g_s might be suitable for assessment of drought stress. There were indications of positive associations between plants carbon isotope composition and water use efficiency (WUE) as well as DM under well‐watered conditions. Compared to control, drought‐treated plots showed increased growth in the period after drought stress. Thus, the drought events did not affect total biomass production (DM_total) of the whole growing season. During drought stress and the whole growing season, WUE was higher in drought‐treated compared to control plots, so it seems possible to save water without loss of biomass. Across soil types, M. lutarioriparius had the highest DM_total (15.0 t ha^?1), WUE_total (3.6 g L^?1) and radiation use efficiency (2.3 g MJ^?1) of the evaluated grasses.     

Most stomatal closure in woody species under moderate drought can be explained by stomatal responses to leaf turgor

Reduced stomatal conductance (g_s) during soil drought in angiosperms may result from effects of leaf turgor on stomata and/or factors that do not directly depend on leaf turgor, including root‐derived abscisic acid (ABA) signals. To quantify the roles of leaf turgor‐mediated and leaf turgor‐independent mechanisms in g_s decline during drought, we measured drought responses of g_s and water relations in three woody species (almond, grapevine and olive) under a range of conditions designed to generate independent variation in leaf and root turgor, including diurnal variation in evaporative demand and changes in plant hydraulic conductance and leaf osmotic pressure. We then applied these data to a process‐based g_s model and used a novel method to partition observed declines in g_s during drought into contributions from each parameter in the model. Soil drought reduced g_s by 63–84% across species, and the model reproduced these changes well (r² = 0.91, P < 0.0001, n = 44) despite having only a single fitted parameter. Our analysis concluded that responses mediated by leaf turgor could explain over 87% of the observed decline in g_s across species, adding to a growing body of evidence that challenges the root ABA‐centric model of stomatal responses to drought.     

Recovery of Populus tremuloides seedlings following severe drought causing total leaf mortality and extreme stem embolism

In contrast with other native Populus species in North America, Populus tremuloides (aspen) can successfully establish itself in drought‐prone areas, yet no comprehensive analysis has been performed on the ability of seedlings to withstand and recover from a severe drought resulting in complete leaf mortality. Here, we subjected 4‐month‐old aspen seedlings grown in two contrasting soil media to a progressive drought until total leaf mortality, followed by a rewatering cycle. Stomatal conductance (g_s), photosynthesis and transpiration followed a sigmoid decline with declining fraction of extractable soil water values. Cessation of leaf expansion occurred close to the end of the linear‐decrease phase, when g_s was reduced by 95%. Leaf mortality started after g_s reached the lowest values, which corresponded to a stem–xylem pressure potential (Ψ_xp) of ?2.0 MPa and a percent loss of stem hydraulic conductivity (PLC) of 50%. In plants with 50% leaf mortality, PLC values remained around 50%. Complete leaf mortality occurred at an average stem PLC of 90%, but all seedlings were able to resprout after 6–10 days of being rewatered. Plants decapitated at soil level before rewatering developed root suckers, while those left with a 4‐cm stump or with their stems intact resprouted exclusively from axillary buds. Resprouting was accompanied by recovery of stem hydraulic conductivity, with PLC values around 30%. The percentage of resprouted buds was negatively correlated with the stem %PLC. Thus, the recovery of stem hydraulic conductivity appears as an important factor in the resprouting capacity of aspen seedlings following a severe drought.     

Photosynthesis and Growth Responses of Parthenocissus quinquefolia (L.) Planch to Soil Water Availability

Photosynthesis and growth characteristics of Parthenocissus quinquefolia were measured under differing soil water availability within a pot. Decreased soil moisture significantly reduced the leaf relative water content (RWC) and the above- and below-ground biomass. However, more biomass was allocated to the root than to the leaf. Net photosynthetic rate (P _N), stomatal conductance (g _s), and transpiration rate (E) were also significantly decreased but water use efficiency (WUE) was increased. Midday depressions in P _N and g _s were not evident for the well-irrigated plants. With the lower water availability, midday reductions in P _N and g _s were much more marked and the duration of the depression was longer. Additionally, the P _N-irradiance response curves also indicated that water supply affected photosynthesis capacity. The growth and photosynthetic response of P. quinquefolia to water supply indicated that this species could resilient to water availabilities and adapt to Hunshandak conditions very well.     

Drought responses,phenotypic plasticity and survival of Mediterranean species in two different microclimatic sites

• Climate models predict a further drying of the Mediterranean summer. One way for plant species to persist during such climate changes is through acclimation. Here, we determine the extent to which trait plasticity in response to drought differs between species and between sites, and address the question whether there is a trade‐off between drought survival and phenotypic plasticity.
• Throughout the summer we measured physiological traits (photosynthesis – A_max, stomatal conductance – g_s, transpiration – E, leaf water potential – ψl) and structural traits (specific leaf area – SLA, leaf density – LD, leaf dry matter content – LDMC, leaf relative water content – LRWC) of leaves of eight woody species in two sites with slightly different microclimate (north‐ versus south‐facing slopes) in southern Spain. Plant recovery and survival was estimated after the summer drought period.
• We found high trait variability between species. In most variables, phenotypic plasticity was lower in the drier site. Phenotypic plasticity of SLA and LDMC correlated negatively with drought survival, which suggests a trade‐off between them. On the other hand, high phenotypic plasticity of SLA and LDMC was positively related to traits associated with rapid recovery and growth after the drought period.
• Although phenotypic plasticity is generally seen as favourable during stress conditions, here it seemed beneficial for favourable conditions. We propose that in environments with fluctuating drought periods there can be a trade‐off between drought survival and growth during favourable conditions. When climate become drier, species with high drought survival but low phenotypic plasticity might be selected for.

     

Disentangling the contributions of ontogeny and water stress to photosynthetic limitations in almond trees

Very few studies have attempted to disentangle the respective role of ontogeny and water stress on leaf photosynthetic attributes. The relative significance of both effects on photosynthetic attributes has been investigated in leaves of field‐grown almond trees [Prunus dulcis (Mill.) D. A. Webb] during four growth cycles. Leaf ontogeny resulted in enhanced leaf dry weight per unit area (W_a), greater leaf dry‐to‐fresh weight ratio and lower N content per unit of leaf dry weight (N_w). Concomitantly, area‐based maximum carboxylation rate (V_cmax), maximum electron transport rate (J_max), mesophyll conductance to CO₂ diffusion (gm)′ and light‐saturated net photosynthesis (A_max) declined in both well‐watered and water‐stressed almond leaves. Although g_m and stomatal conductance (g_s) seemed to be co‐ordinated, a much stronger coordination in response to ontogeny and prolonged water stress was observed between g_m and the leaf photosynthetic capacity. Under unrestricted water supply, the leaf age‐related decline of A_max was equally driven by diffusional and biochemical limitations. Under restricted soil water availability, A_max was mainly limited by g_s and, to a lesser extent, by photosynthetic capacity and g_m. When both ontogeny and water stress effects were combined, diffusional limitations was the main determinant of photosynthesis limitation, while stomatal and biochemical limitations contributed similarly.     

Variations in photosynthetic rate and associated parameters with age of oil palm leaves under irrigation

Net photosynthetic rate (P _N), transpiration rate (E), and stomatal conductance (g _s) in an adult oil palm (Elaeis guineensis) canopy were highest in the 9^th leaf and progressively declined with leaf age. Larger leaf area (LA) and leaf dry mass (LDM) were recorded in middle leaves. P _N showed a significant positive correlation with g _s and a negative relationship with leaf mass per area (ALM). The oil palm leaf remains photosynthetically active for a longer time in the canopy which contributes significantly to larger dry matter production in general and greater fresh fruit bunch yields in particular.     

Effect of drought on leaf gland secretion of the mangrove Avicennia germinans L.

This study assessed the effects of salinity changes over space and time upon leaf gland secretion in Avicennia germinans trees growing naturally in an area featuring markedly seasonal rainfall. Soil ? (, soil N MPa) during the wet season was -0.95ǂ.05 and -2.12ǂ.08 at low and high salinity sites, respectively. During the dry season, these values decreased to -3.24ǂ.09 at low salinity and to -5.75ǂ.06 at high salinity. Consequently, predawn and midday plant water potential were lowered during drought at both sites. The rates of secretion (mmol m^-2 h^-1 ) increased during drought from 0.91ǂ.12 during the wet season to 1.93ǂ.12 at low salinity, and from 1.69ǂ.12 during the wet season to 2.81ǂ.15 at high salinity. Conversely, stomatal conductance (g_s) was lowered by both salinity and drought. As xylem osmolality increased during drought, secretion tended to rise exponentially, and g_s decayed hyperbolically. Thus, a trade-off is obtained between enhancement in salt secretion and control of water loss suggested by g_s.     

Daily variations in water relations of apricot trees under different irrigation regimes

Mature apricot (Prunus armeniaca L. cv. Búlida) trees, growing under field conditions, were submitted to two drip irrigation treatments: a control (T1), irrigated to 100 % of seasonal crop evapotranspiration (ETc), and a continuous deficit (T2), irrigated to 50 % of the control throughout the year. The behaviour of leaf water potential and its components, leaf conductance and net photosynthesis were studied at three different times during the growing season, when they revealed a diurnal and seasonal pattern in response to water stress, evaporative demand of the atmosphere and leaf age. The deficit-irrigated trees showed, among other effects, a pronounced decrease in leaf water potential (ψ_w), decreased in leaf conductance (g_s) and no osmotic adjustment. For this reason, g_l and ψ_w can be considered good indicators of mature apricot tree water status and can therefore be used for irrigation scheduling.     

Stomatal sensitivity to vapour pressure difference over a subambient to elevated CO2 gradient in a C3/C4 grassland

In the present study the response of stomatal conductance (g_s) to increasing leaf‐to‐air vapour pressure difference (D) in early season C₃ (Bromus japonicus) and late season C₄ (Bothriochloa ischaemum) grasses grown in the field across a range of CO₂ (200–550 µmol mol^?1) was examined. Stomatal sensitivity to D was calculated as the slope of the response of g_s to the natural log of externally manipulated D (dg_s/dlnD). Increasing D and CO₂ significantly reduced g_s in both species. Increasing CO₂ caused a significant decrease in stomatal sensitivity to D in Br. japonicus, but not in Bo. ischaemum. The decrease in stomatal sensitivity to D at high CO₂ for Br. japonicus fit theoretical expectations of a hydraulic model of stomatal regulation, in which g_s varies to maintain constant transpiration and leaf water potential. The weaker stomatal sensitivity to D in Bo. ischaemum suggested that stomatal regulation of leaf water potential was poor in this species, or that non‐hydraulic signals influenced guard cell behaviour. Photosynthesis (A) declined with increasing D in both species, but analyses of the ratio of intercellular to atmospheric CO₂ (C_i/C_a) suggested that stomatal limitation of A occurred only in Br. japonicus. Rising CO₂ had the greatest effect on g_s and A in Br. japonicus at low D. In contrast, the strength of stomatal and photosynthetic responses to CO₂ were not affected by D in Bo. ischaemum. Carbon and water dynamics in this grassland are dominated by a seasonal transition from C₃ to C₄ photosynthesis. Interspecific variation in the response of g_s to D therefore has implications for predicting seasonal ecosystem responses to CO₂.     

Seasonal variations on water relations ofAmygdalus communis L. under drip irrigated and non irrigated conditions

Almond plants (Amygdalus communis L.) of the Garrigues variety were grown in the field drip irrigated and rainfed. Leaf water potential (Ψ) and leaf conductance (g₁) were determined throughout one growing season. Pre-dawn measurement for Ψ in the irrigated treatment was consistent through the growing season, whereas in the rainfed treatment it decreased gradually. Ψ values at midday (Ψ minimum) was closely dependent on atmospheric evaporative demand, and their recovery was quicker in the wet treatment than in the dry. The g₁ values were higher in the wet than dry treatments, decreasing in both cases by leaf ageing. Maximum values for g₁ were reached when evaporative demand was highest in the day. The relationship between Ψ and g₁ revealed a decrease in the hysteresis throughout the growing season, being most marked in the dry treatment. The results highlight the close dependence of Ψ and g₁ on evaporative demand, leaf ageing and irrigtion treatment during the growing season.     

Xylem cavitation and hydraulic control of stomatal conductance in Laurel (Laurus nobilis L.)

The possible link between stomatal conductance (g_L), leaf water potential ( Ψ _L) and xylem cavitation was studied in leaves and shoots of detached branches as well as of whole plants of Laurus nobilis L. (Laurel). Shoot cavitation induced complete stomatal closure in air‐dehydrated detached branches in less than 10 min. By contrast, a fine regulation of g_L in whole plants was the consequence of Ψ _L reaching the cavitation threshold ( Ψ _CAV) for shoots. A pulse of xylem cavitation in the shoots was paralleled by a decrease in g_L of about 50%, while Ψ _L stabilized at values preventing further xylem cavitation. In these experiments, no root signals were likely to be sent to the leaves from the roots in response to soil dryness because branches were either detached or whole plants were growing in constantly wet soil. The stomatal response to increasing evaporative demand appeared therefore to be the result of hydraulic signals generated during shoot cavitation. A negative feedback link is proposed between g_L and Ψ _CAV rather than with Ψ _L itself.