Impacts of long‐term precipitation manipulation on hydraulic architecture and xylem anatomy of piñon and juniper in Southwest USA

Hydraulic architecture imposes a fundamental control on water transport, underpinning plant productivity, and survival. The extent to which hydraulic architecture of mature trees acclimates to chronic drought is poorly understood, limiting accuracy in predictions of forest responses to future droughts. We measured seasonal shoot hydraulic performance for multiple years to assess xylem acclimation in mature piñon (Pinus edulis ) and juniper (Juniperus monosperma ) after 3+ years of precipitation manipulation. Our treatments consisted of water addition (+20% ambient precipitation), partial precipitation‐exclusion (?45% ambient precipitation), and exclusion‐structure control. Supplemental watering elevated leaf water potential, sapwood‐area specific hydraulic conductivity, and leaf‐area specific hydraulic conductivity relative to precipitation exclusion. Shifts in allocation of leaf area to sapwood area enhanced differences between irrigated and droughted K _L in piñon but not juniper. Piñon and juniper achieved similar K _L under ambient conditions, but juniper matched or outperformed piñon in all physiological measurements under both increased and decreased precipitation treatments. Embolism vulnerability and xylem anatomy were unaffected by treatments in either species. Absence of significant acclimation combined with inferior performance for both hydraulic transport and safety suggests piñon has greater risk of local extirpation if aridity increases as predicted in the southwestern USA.     

Prolonged experimental drought reduces plant hydraulic conductance and transpiration and increases mortality in a piñon–juniper woodland

Plant hydraulic conductance (k_s) is a critical control on whole‐plant water use and carbon uptake and, during drought, influences whether plants survive or die. To assess long‐term physiological and hydraulic responses of mature trees to water availability, we manipulated ecosystem‐scale water availability from 2007 to 2013 in a piñon pine (Pinus edulis) and juniper (Juniperus monosperma) woodland. We examined the relationship between k_s and subsequent mortality using more than 5 years of physiological observations, and the subsequent impact of reduced hydraulic function and mortality on total woody canopy transpiration (E_C) and conductance (G_C). For both species, we observed significant reductions in plant transpiration (E) and k_s under experimentally imposed drought. Conversely, supplemental water additions increased E and k_s in both species. Interestingly, both species exhibited similar declines in k_s under the imposed drought conditions, despite their differing stomatal responses and mortality patterns during drought. Reduced whole‐plant k_s also reduced carbon assimilation in both species, as leaf‐level stomatal conductance (g_s) and net photosynthesis (A_n) declined strongly with decreasing k_s. Finally, we observed that chronically low whole‐plant k_s was associated with greater canopy dieback and mortality for both piñon and juniper and that subsequent reductions in woody canopy biomass due to mortality had a significant impact on both daily and annual canopy E_C and G_C. Our data indicate that significant reductions in k_s precede drought‐related tree mortality events in this system, and the consequence is a significant reduction in canopy gas exchange and carbon fixation. Our results suggest that reductions in productivity and woody plant cover in piñon–juniper woodlands can be expected due to reduced plant hydraulic conductance and increased mortality of both piñon pine and juniper under anticipated future conditions of more frequent and persistent regional drought in the southwestern United States.     

Regulation and acclimation of leaf gas exchange in a piñon–juniper woodland exposed to three different precipitation regimes

Leaf gas‐exchange regulation plays a central role in the ability of trees to survive drought, but forecasting the future response of gas exchange to prolonged drought is hampered by our lack of knowledge regarding potential acclimation. To investigate whether leaf gas‐exchange rates and sensitivity to drought acclimate to precipitation regimes, we measured the seasonal variations of leaf gas exchange in a mature piñon–juniper Pinus edulis–Juniperus monosperma woodland after 3 years of precipitation manipulation. We compared trees receiving ambient precipitation with those in an irrigated treatment (+30% of ambient precipitation) and a partial rainfall exclusion (?45%). Treatments significantly affected leaf water potential, stomatal conductance and photosynthesis for both isohydric piñon and anisohydric juniper. Leaf gas exchange acclimated to the precipitation regimes in both species. Maximum gas‐exchange rates under well‐watered conditions, leaf‐specific hydraulic conductance and leaf water potential at zero photosynthetic assimilation all decreased with decreasing precipitation. Despite their distinct drought resistance and stomatal regulation strategies, both species experienced hydraulic limitation on leaf gas exchange when precipitation decreased, leading to an intraspecific trade‐off between maximum photosynthetic assimilation and resistance of photosynthesis to drought. This response will be most detrimental to the carbon balance of piñon under predicted increases in aridity in the southwestern USA.     

Hydraulic limits preceding mortality in a piñon-juniper woodland under experimental drought

Drought‐related tree mortality occurs globally and may increase in the future, but we lack sufficient mechanistic understanding to accurately predict it. Here we present the first field assessment of the physiological mechanisms leading to mortality in an ecosystem‐scale rainfall manipulation of a piñon–juniper (Pinus edulis–Juniperus monosperma) woodland. We measured transpiration (E) and modelled the transpiration rate initiating hydraulic failure (E_crit). We predicted that isohydric piñon would experience mortality after prolonged periods of severely limited gas exchange as required to avoid hydraulic failure; anisohydric juniper would also avoid hydraulic failure, but sustain gas exchange due to its greater cavitation resistance. After 1 year of treatment, 67% of droughted mature piñon died with concomitant infestation by bark beetles (Ips confusus) and bluestain fungus (Ophiostoma spp.); no mortality occurred in juniper or in control piñon. As predicted, both species avoided hydraulic failure, but safety margins from E_crit were much smaller in piñon, especially droughted piñon, which also experienced chronically low hydraulic conductance. The defining characteristic of trees that died was a 7 month period of near‐zero gas exchange, versus 2 months for surviving piñon. Hydraulic limits to gas exchange, not hydraulic failure per se, promoted drought‐related mortality in piñon pine.     

Importance of juniper to birds nesting in piñon–juniper woodlands in northwest New Mexico

Piñon–juniper (Pinus spp.–Juniperus spp.) woodlands are common throughout western North America, yet relatively little is known about the habitat use and requirements for many members of its avian community. During summer 2005–2007, we assessed avian nesting substrates within piñon (Pinus edulis)–juniper (Juniperus osteosperma) woodlands in northwestern New Mexico. Of all nests in live trees, 86% were in junipers. The selection of juniper as a nest tree was significantly higher than expected from the region's piñon–juniper ratio (1:1.06) for the community as a whole, for both open cup and cavity nesting species, and for 8 species (of which 6 are piñon–juniper obligate or semi-obligate species). Nest survival, however, was not higher in juniper than in piñon for the nesting community as a whole or for chipping sparrows (Spizella passerina), the single species that was well represented nesting in piñon. The high use of juniper as a nesting substrate differs from previous studies, which have suggested that a presence of piñon is among the most important habitat features for many piñon–juniper species. Because of their importance to nesting birds, managers should avoid preferential thinning of junipers within piñon–juniper woodlands. © 2011 The Wildlife Society.     

Plasticity in hydraulic architecture of Scots pine across Eurasia

Widespread tree species must show physiological and structural plasticity to deal with contrasting water balance conditions. To investigate these plasticity mechanisms, a meta-analysis of Pinus sylvestris L. sap flow and its response to environmental variables was conducted using datasets from across its whole geographical range. For each site, a Jarvis-type, multiplicative model was used to fit the relationship between sap flow and photosynthetically active radiation, vapour pressure deficit (D) and soil moisture deficit (SMD); and a logarithmic function was used to characterize the response of stomatal conductance (G _s) to D. The fitted parameters of those models were regressed against climatic variables to study the acclimation of Scots pine to dry/warm conditions. The absolute value of sap flow and its sensitivity to D and SMD increased with the average summer evaporative demand. However, relative sensitivity of G _s to D (m/G _s,ref, where m is the slope and G _s,ref is reference G _s at D = 1 kPa) did not increase with evaporative demand across populations, and transpiration per unit leaf area at a given D increased accordingly in drier/warmer climates. This physiological plasticity was linked to the previously reported climate- and size-related structural acclimation of leaf to sapwood area ratios. G _s,ref, and its absolute sensitivity to D (m), tended to decrease with age/height of the trees as previously reported for other pine species. It is unclear why Scots pines have higher transpiration rates at drier/warmer sites, at the expense of lower water-use efficiency. In any case, our results suggest that these structural adjustments may not be enough to prevent lower xylem tensions at the driest sites.     

Integrating ecophysiology and forest landscape models to improve projections of drought effects under climate change

Fundamental drivers of ecosystem processes such as temperature and precipitation are rapidly changing and creating novel environmental conditions. Forest landscape models (FLM) are used by managers and policy‐makers to make projections of future ecosystem dynamics under alternative management or policy options, but the links between the fundamental drivers and projected responses are weak and indirect, limiting their reliability for projecting the impacts of climate change. We developed and tested a relatively mechanistic method to simulate the effects of changing precipitation on species competition within the LANDIS‐II FLM. Using data from a field precipitation manipulation experiment in a piñon pine (Pinus edulis) and juniper (Juniperus monosperma) ecosystem in New Mexico (USA), we calibrated our model to measurements from ambient control plots and tested predictions under the drought and irrigation treatments against empirical measurements. The model successfully predicted behavior of physiological variables under the treatments. Discrepancies between model output and empirical data occurred when the monthly time step of the model failed to capture the short‐term dynamics of the ecosystem as recorded by instantaneous field measurements. We applied the model to heuristically assess the effect of alternative climate scenarios on the piñon–juniper ecosystem and found that warmer and drier climate reduced productivity and increased the risk of drought‐induced mortality, especially for piñon. We concluded that the direct links between fundamental drivers and growth rates in our model hold great promise to improve our understanding of ecosystem processes under climate change and improve management decisions because of its greater reliance on first principles.     

Carbohydrate dynamics and mortality in a piñon‐juniper woodland under three future precipitation scenarios

Drought‐induced forest mortality is an increasing global problem with wide‐ranging consequences, yet mortality mechanisms remain poorly understood. Depletion of non‐structural carbohydrate (NSC) stores has been implicated as an important mechanism in drought‐induced mortality, but experimental field tests are rare. We used an ecosystem‐scale precipitation manipulation experiment to evaluate leaf and twig NSC dynamics of two co‐occurring conifers that differ in patterns of stomatal regulation of water loss and recent mortality: the relatively desiccation‐avoiding piñon pine (Pinus edulis) and the relatively desiccation‐tolerant one‐seed juniper (Juniperus monosperma). Piñon pine experienced 72% mortality after 13–25 months of experimental drought and juniper experienced 20% mortality after 32–47 months. Juniper maintained three times more NSC in the foliage than twigs, and converted NSC to glucose and fructose under drought, consistent with osmoregulation requirements to maintain higher stomatal conductance during drought than piñon. Despite these species differences, experimental drought caused decreased leaf starch content in dying trees of both species (P < 0.001). Average dry‐season leaf starch content was also a good predictor of drought‐survival time for both species (R² = 0.93). These results, along with observations of drought‐induced reductions to photosynthesis and growth, support carbon limitation as an important process during mortality of these two conifer species.     

Stomatal acclimation to vapour pressure deficit doubles transpiration of small tree seedlings with warming

Future climate change is expected to increase temperature (T) and atmospheric vapour pressure deficit (VPD) in many regions, but the effect of persistent warming on plant stomatal behaviour is highly uncertain. We investigated the effect of experimental warming of 1.9–5.1 °C and increased VPD of 0.5–1.3 kPa on transpiration and stomatal conductance (g_s) of tree seedlings in the temperate forest understory (Duke Forest, North Carolina, USA). We observed peaked responses of transpiration to VPD in all seedlings, and the optimum VPD for transpiration (D_opt) shifted proportionally with increasing chamber VPD. Warming increased mean water use of Carya by 140% and Quercus by 150%, but had no significant effect on water use of Acer. Increased water use of ring‐porous species was attributed to (1) higher air T and (2) stomatal acclimation to VPD resulting in higher g_s and more sensitive stomata, and thereby less efficient water use. Stomatal acclimation maintained homeostasis of leaf T and carbon gain despite increased VPD, revealing that short‐term stomatal responses to VPD may not be representative of long‐term exposure. Acclimation responses differ from expectations of decreasing g_s with increasing VPD and may necessitate revision of current models based on this assumption.     

The effect of tree height on crown level stomatal conductance

Variation in stomatal conductance is typically explained in relation to environmental conditions. However, tree height may also contribute to the variability in mean stomatal conductance. Mean canopy stomatal conductance of individual tree crowns (G_Si) was estimated using sap flux measurements in Fagus sylvatica L., and the hypothesis that G_Si decreases with tree height was tested. Over 13 d of the growing season during which soil moisture was not limiting, G_Si decreased linearly with the natural logarithm of vapour pressure deficit (D), and increased exponentially to saturation with photosynthetic photon flux density (Q_o). Under conditions of D = 1 kPa and saturating Q_o, G_Si decreased by approximately 60% with 30 m increase in tree height. Over the same range in height, sapwood‐to‐leaf area ratio (A_S:A_L) doubled. A simple hydraulic model explained the variation in G_Si based on an inverse relationship with height, and a linear relationship with A_S:A_L. Thus, in F. sylvatica, adjustments in A_S:A_L partially compensate for the negative effect of increased flow‐path length on leaf conductance. Furthermore, because stomata with low conductance are less sensitive to D, gas exchange of tall trees is reduced less by high D. Despite these compensations, decreasing hydraulic conductance with tree height in F. sylvatica reduces carbon uptake through a corresponding decrease in stomatal conductance.     

Stomatal conductance and photosynthesis vary linearly with plant hydraulic conductance in ponderosa pine

Recent work has shown that stomatal conductance (g_s) and assimilation (A) are responsive to changes in the hydraulic conductance of the soil to leaf pathway (K_L), but no study has quantitatively described this relationship under controlled conditions where steady‐state flow is promoted. Under steady‐state conditions, the relationship between g_s, water potential (Ψ) and K_L can be assumed to follow the Ohm's law analogy for fluid flow. When boundary layer conductance is large relative to g_s, the Ohm's law analogy leads to g_s = K_L (Ψ_soil?Ψ_leaf)/D, where D is the vapour pressure deficit. Consequently, if stomata regulate Ψ_leaf and limit A, a reduction in K_L will cause g_s and A to decline. We evaluated the regulation of Ψ_leaf and A in response to changes in K_L in well‐watered ponderosa pine seedlings (Pinus ponderosa). To vary K_L, we systematically reduced stem hydraulic conductivity (k) using an air injection technique to induce cavitation while simultaneously measuring Ψ_leaf and canopy gas exchange in the laboratory under constant light and D. Short‐statured seedlings (< 1 m tall) and hour‐long equilibration times promoted steady‐state flow conditions. We found that Ψ_leaf remained constant near ? 1·5 MPa except at the extreme 99% reduction of k when Ψ_leaf fell to ? 2·1 MPa. Transpiration, g_s, A and K_L all declined with decreasing k (P < 0·001). As a result of the near homeostasis in bulk Ψ_leaf, g_s and A were directly proportional to K_L (R² > 0·90), indicating that changes in K_L may affect plant carbon gain.     

Characterization of photosynthesis of Populus euphratica grown in the arid region

Net photosynthetic rate (P _N), stomatal conductance (g _s), intercellular CO₂ concentration (C _i), transpiration rate (E), water use efficiency (WUE), and stomatal limitation (L_s) of Populus euphratica grown at different groundwater depths in the arid region were measured. g _s of the trees with groundwater depth at 4.74 m (D₄) and 5.82 m (D₅) were lower and a little higher than that at 3.82 m (D₃), respectively. Compared with C _i and L_s of the D₃ trees, C _i decreased and L_s increased at 4.74 m, however, Ci increased and L_s decreased at D₅. Hence photosynthetic reduction of P. euphratica was attributed to either stomatal closure or non-stomatal factors depending on the groundwater depths in the plant locations. P _N of the D₃ trees was significantly higher than those at D₄ or D₅. The trees of D₄ and D₅ did not show a significant difference in their P _N, indicating that there are mechanisms of P. euphratica tolerance to mild and moderate drought stress.     

沙质草地3种优势植物叶片光合生理对增温和降水减少的响应北大核心CSCD

以沙质草地优势物种猪毛蒿、胡枝子和糙隐子草为研究对象,利用开顶式生长室(OTC)模拟增温,研究降水减少20%、40%和60%与增温的交互作用对3种典型植物叶片光合气体交换特征及叶绿素荧光特征的影响,以揭示沙质草地3种优势植物对气候变化的响应规律。结果显示:(1)与自然温度相比,OTC模拟增温增加了猪毛蒿C_(i),显著降低了胡枝子G_(s)、P_(n)和T_(r)、糙隐子草G_(s)和P_(n)、猪毛蒿WUE和L_(s),也显著降低了猪毛蒿和胡枝子F_(v)/F_(m)和F_(v)/F_(o)。(2)无论增温与否,随着降水减少幅度的增加,猪毛蒿G_(s)和P_(n)呈下降趋势,且中度以上的干旱胁迫下(降水减少>40%)胡枝子和糙隐子草P_(n)显著低于对照。(3)在自然温度条件下,轻度干旱胁迫时(降水减少20%)猪毛蒿T_(r)显著低于对照,重度干旱胁迫时(降水减少60%)其WUE、F_(v)/F_(m)和F_(v)/F_(o)显著低于对照;重度干旱胁迫时,胡枝子C_(i)显著高于对照,差异幅度达10.7%,L_(s)显著低于对照,轻度干旱胁迫时(降水减少20%)其F_(v)/F_(m)和F_(v)/F_(o)显著高于中度以上的干旱胁迫;中度以上的干旱胁迫下糙隐子草T_(r)和G_(s)显著低于对照,重度干旱胁迫时,其C_(i)、F_(v)/F_(m)和F_(v)/F_(o)显著低于对照,WUE和L_(s)显著高于对照。(4)增温与降水减少交互作用下,所有处理猪毛蒿C_(i)均高于对照,差异幅度分别达4.5%,6.0%和8.4%;胡枝子T_(r)均显著低于对照,差异幅度达57.8%;重度干旱胁迫时猪毛蒿L_(s)和WUE显著低于对照,糙隐子草F_(v)/F_(m)和F_(v)/F_(o)随降水减少而降低,中度以上的干旱胁迫时其值显著低于对照。(5)相关性分析表明,3个优势物种的P_(n)与F_(v)/F_(m)和F_(v)/F_(o)均呈显著正相关关系,其中猪毛蒿和糙隐子草的P_(n)—F_(v)/F_(m)和P_(n)—F_(v)/F_(o)斜率明显高于胡枝子。研究表明,气候变暖会在一定程度上加剧降水减少对沙质草地3种群落优势物种光合作用的抑制。     

Survey and synthesis of intra- and interspecific variation in stomatal sensitivity to vapour pressure deficit

Responses of stomatal conductance (g_s) to increasing vapour pressure deficit (D) generally follow an exponential decrease described equally well by several empirical functions. However, the magnitude of the decrease – the stomatal sensitivity – varies considerably both within and between species. Here we analysed data from a variety of sources employing both porometric and sap flux estimates of g_s to evaluate the hypothesis that stomatal sensitivity is proportional to the magnitude of g_s at low D ( ≤ 1 kPa). To test this relationship we used the function g_s = g_sref–m· lnD where m is the stomatal sensitivity and g_sref = g_s at D = 1 kPa. Regardless of species or methodology, m was highly correlated with g_sref (average r² = 0·75) with a slope of approximately 0·6. We demonstrate that this empirical slope is consistent with the theoretical slope derived from a simple hydraulic model that assumes stomatal regulation of leaf water potential. The theoretical slope is robust to deviations from underlying assumptions and variation in model parameters. The relationships within and among species are close to theoretical predictions, regardless of whether the analysis is based on porometric measurements of g_s in relation to leaf-surface D (D_s), or on sap flux-based stomatal conductance of whole trees (G_Si), or stand-level stomatal conductance (G_S) in relation to D. Thus, individuals, species, and stands with high stomatal conductance at low D show a greater sensitivity to D, as required by the role of stomata in regulating leaf water potential.     

Stomatal regulation in Douglas fir following a fungal-mediated chronic reduction in leaf area

Pathogens can cause chronic premature needle abscission in coniferous species. To assess the potential impacts on tree productivity, stomatal regulation was investigated in Douglas fir with chronic stomatal occlusion and defoliation from varying levels of the Swiss needle cast (SNC) fungus, Phaeocryptopus gaeumannii. Levels of SNC disease and subsequent defoliation were manipulated by choosing six sites with varying levels of disease and by foliar applications of fungicides on six trees per site. Diurnal measurements of leaf water potential (Ψ_leaf), stomatal conductance (g _s) and vapor pressure deficit (D) were made on six fungicide treated and six control trees per site. In addition, leaf specific hydraulic conductance was calculated on a single branch (K _{L_B}) from three trees per treatment per site. Stomatal conductance at D=1 kPa (g _sref) was negatively correlated with fungal colonization (number of fruiting bodies present in needle stomata) and positively correlated with K _{L_B}. Despite reduced needle retention in diseased trees, K _L declined due to a reduction in sapwood area and permeability (i.e., increasing presence of latewood in functional sapwood). In general, stomatal sensitivity to D for all foliage was consistent with stomatal regulation based on a simple hydraulic model [g _s=K _L(Ψ_soil?Ψ_leaf)/ D], which assumes strict stomatal regulation of Ψ_leaf. However, when fungal presence reduced maximum g _s below the potential maximum supported by hydraulic architecture, stomatal sensitivity was lower than expected based on the theoretical relationship: dg _s/dlnD=0.6·g _sref. The results indicate that losses in productivity associated with physical blockage of stomata and defoliation are compounded by additional losses in K _L and a reduction in g _s in remaining functional stomata.     

Postfire soil water repellency in piñon–juniper woodlands: Extent,severity, and thickness relative to ecological site characteristics and climate

Wildfires can create or intensify water repellency in soil, limiting the soil's capacity to wet and retain water. The objective of this research was to quantify soil water repellency characteristics within burned piñon–juniper woodlands and relate this information to ecological site characteristics. We sampled soil water repellency across forty‐one 1,000 m² study plots within three major wildfires that burned in piñon–juniper woodlands. Water repellency was found to be extensive—present at 37% of the total points sampled—and strongly related to piñon–juniper canopy cover. Models developed for predicting SWR extent and severity had R²_adj values of 0.67 and 0.61, respectively; both models included piñon–juniper canopy cover and relative humidity the month before the fire as coefficient terms. These results are important as they suggest that postfire water repellency will increase in the coming years as infilling processes enhance piñon–juniper canopy cover. Furthermore, reductions in relative humidity brought about by a changing climate have the potential to link additively with infilling processes to increase the frequency and intensity of wildfires and produce stronger water repellency over a greater spatial extent. In working through these challenges, land managers can apply the predictive models developed in this study to prioritize fuel control and postfire restoration treatments.     

Stem radial growth and water storage responses to heat and drought vary between conifers with differing hydraulic strategies

We investigated stem radial growth and water storage dynamics of 2 conifer species differing in hydraulic carbon strategies, Juniperus monosperma and Pinus edulis, under conditions of ambient, drought (～45% reduction in precipitation), heat (～4.8 °C temperature increase), and the combination of drought + heat, in 2013 and 2014. Juniper maintained low growth across all treatments. Overall, the relatively isohydric piñon pine showed significantly greater growth and water storage recharge than the relatively anisohydric juniper across all treatments in the average climate year (2014) but no differences in the regionally dry year (2013). Piñon pine ceased growth at a constant predawn water potential across all treatments and at a less negative water potential threshold than juniper. Heat has a greater negative impact on piñon pines' growth and water storage than drought, whereas juniper was, in contrast, unaffected by heat but strongly impacted by drought. The whole‐plant hydraulic carbon strategies, in this case captured using the isohydric/anisohydric concept, translate into alternative growth and water storage strategies under drought and heat conditions.     

Diurnal and seasonal variation in photosynthesis of peach palms grown under subtropical conditions

The Amazonian peach palm (Bactris gasipaes Kunth) has been grown for heart-of-palm production under subtropical conditions. As we did not see any substantial study on its photosynthesis under Amazonian or subtropical conditions, we carried out an investigation on the diurnal and seasonal variations in photosynthesis of peach palms until the first heart-of-palm harvest, considering their relationship with key environmental factors. Spineless peach palms were grown in 80-L plastic pots, under irrigation. Gas exchange and chlorophyll fluorescence emission measurements were taken in late winter, mid spring, mid summer and early autumn, from 7:00 to 18:00 h, with an additional chlorophyll fluorescence measurement at 6:00 h. The highest net CO₂ assimilation (P _N), observed in mid summer, reached about 15 μmol m⁻² s⁻¹, which was about 20% higher than the maximum values found in autumn and spring, and 60% higher than that in winter The same pattern of diurnal course for P _N was observed in all seasons, showing higher values from 8:00 to 9:00 h and declining gradually from 11:00 h toward late afternoon. The diurnal course of stomatal conductance (g _s) followed the same pattern of P _N, with the highest value of 0.6 mol m⁻² s⁻¹ being observed in February and the lowest one (0.23 mol m⁻² s⁻¹) in September. The maximal quantum yield of photosystem II (F_v/F_m) was above 0.75 in the early morning in all the months. The reversible decrease was observed around midday in September and October, suggesting the occurrence of dynamic photoinhibition. A significant negative correlation between the leaf-air vapour pressure difference (VPD_leaf-air) and P _N and a positive correlation between P _N and g _s were observed. The photosynthesis of peach palm was likely modulated mainly by the stomatal control that was quite sensible to atmospheric environmental conditions. Under subtropical conditions, air temperature (T _air) and VPD_leaf-air impose more significant effects over P _N of peach palm than an excessive photosynthetic photon flux density (PPFD). The occurrence of dynamic photoinhibition indicates that under irrigation, peach palms appeared to be acclimated to the full-sunlight conditions under which they have been grown.     

Water relations and stomatal characteristics of Mediterranean plants with different growth forms and leaf habits: responses to water stress and recovery

The aim of this study was to extent the range of knowledge about water relations and stomatal responses to water stress to ten Mediterranean plants with different growth forms and leaf habits. Plants were subjected to different levels of water stress and a treatment of recovery. Stomatal attributes (stomatal density, StoD), stomatal conductance (g _s), stomatal responsiveness to water stress (SR), leaf water relations (pre-dawn and midday leaf water potential and relative water content), soil to leaf apparent hydraulic conductance (K _L) and bulk modulus of elasticity (ε) were determined. The observed wide range of water relations and stomatal characteristics was found to be partially depended on the growth form. Maximum g _s was related to StoD and the stomatal area index (SAI), while g _s evolution after water stress and recovery was highly correlated with K _L. Relationships between SR to water deficit and other morphological leaf traits, such as StoD, LMA or ε, provided no general correlations when including all species. It is concluded that a high variability is present among Mediterranean plants reflecting a continuum of leaf water relations and stomatal behaviour in response to water stress.     

Stomatal behaviour in a beech canopy: an analysis of Bowen ratio measurements compared with porometer data

On the basis of measurements or stand transpiration and microclimate, the bulk stomatal or bulk leaf conductance (g_L) of a beech forest in northern Germany was calculated for periods in which leaves were fully expanded and the canopy was dry. This conductance depends strongly on light and humidity conditions above the forest. During periods with photosynthetic photon flux densities Q > 1200 μmol m^?2s^?1, g_L was reduced from 1500mmol m^?2s^?1 at a vapour pressure deficit D= 0.5kPa to 500 mmol m^?2s^?1 at D= 2kPa. Light saturation of g_L was not reached until Q= 1200 μmol m^?2s^?1 at low D, or until even higher Q at higher D. The dependence of g_L, on Q and D was described mathematically by a non-linear equation that requires two empirical parameters. Values for g_L as simulated by this equation provided a satisfactory agreement with independent porometer data collected on single leaves and scaled up to the canopy. A comparison of stomatal and aerodynamic conductances showed a strong coupling between the forest canopy and the atmosphere, indicating that transpiration of the beech forest is controlled mainly by the stomata.