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.     

亚热带森林演替树种叶片气孔导度对环境水分的水力响应

利用LI-1600稳态气孔计和PMS压力室,在田间测定了群落演替早期强阳生性树种桃金娘(Rhodomyrtus tomentosa)和三叉苦(Evodia lepta)、偏中性的阳生性树种荷木(Schima superba)、群落演替后期的耐荫树种鸭脚木(Schefllera octophylla)和九节(Psychotrie rubra)的叶片气孔导度(gs)和叶片水势(ΨL),研究不同演替阶段树种的气孔导度对环境水分的响应.结果表明,早上叶片有较高的ΨL,随着时间推移ΨL逐渐降低,与此同时比叶水力导度(KL)随ΨL降低而下降,桃金娘、三叉苦、荷木、鸭脚木和九节水力导度初始最低值时的ΨL分别为-1.6、-1.42、-1.30、-0.9MPa和-1.05MPa.随着ΨL降低,田间测定的gs开始从上午的较低值上升至约中午时的最大值,随后开始降低,此时的ΨL分别为-1.58、-1.52、-1.35、-1.02MPa和-1.0MPa.不同植物种类有不同的KL初始最低值的ΨL和gs达到最大值的ΨL.但不论何种供试树种,KL最低值时的ΨL与gs开始从最大值下降时的ΨL相近;显示KL与gs在动态变化中存在协调关系.树种间的gs和KL对ΨL的不同响应显示桃金娘和三叉苦的KL最低值时和gs开始下降时的ΨL均较鸭脚木和九节对应的ΨL低(p<0.05),意味着演替早期树种能在较强水分胁迫下保持较高的气孔导度.这一水力特性保证树种在水分胁迫下维持叶片的光合速率,有利于其在群落中的生长和优势地位的维护,而演替后期树种在较高ΨL下气孔关闭,降低了光合速率.全球变暖和环境进一步干旱可能成为限制亚热带森林植物群落的正向演替进程的潜在因素之一.     

Leaf hydraulics and drought stress: response, recovery and survivorship in four woody temperate plant species

Efficient conduction of water inside leaves is essential for leaf function, yet the hydraulic-mediated impact of drought on gas exchange remains poorly understood. Here we examine the decline and subsequent recovery of leaf water potential ( Ψ _leaf), leaf hydraulic conductance ( K _leaf), and midday transpiration ( E ) in four temperate woody species exposed to controlled drought conditions ranging from mild to lethal. During drought the vulnerability of K _leaf to declining Ψ _leaf varied greatly among the species sampled. Following drought, plants were rewatered and the rate of E and K _leaf recovery was found to be strongly dependent on the severity of the drought imposed. Gas exchange recovery was strongly correlated with the relatively slow recovery of K _leaf for three of the four species, indicating conformity to a hydraulic-stomatal limitation model of plant recovery. However, there was also a shift in the sensitivity of stomata to Ψ _leaf suggesting that the plant hormone abscisic acid may be involved in limiting the rate of stomatal reopening. The level of drought tolerance varied among the four species and was correlated with leaf hydraulic vulnerability. These results suggest that species-specific variation in hydraulic properties plays a fundamental role in steering the dynamic response of plants during recovery.     

Functional coordination between leaf gas exchange and vulnerability to xylem cavitation in temperate forest trees

We examined functional coordination among stem and root vulnerability to xylem cavitation, plant water transport characteristics and leaf traits in 14 co-occurring temperate tree species. Relationships were evaluated using both traditional cross-species correlations and phylogenetically independent contrast (PIC) correlations. For stems, the xylem tension at which 50% of hydraulic conductivity was lost (psi50) was positively associated (P < 0.001) with specific conductivity (K(S)) and with mean hydraulically weighted xylem conduit diameter (D(h-w)), but was only marginally (P = 0.06) associated with leaf specific conductivity (K(L)). The PIC correlation for each of these relationships, however, was not statistically significant. There was also no relationship between root psi50 and root K(S) in either cross-species or PIC analysis. Photosynthetic rate (A) and stomatal conductance (g(s)) were strongly and positively correlated with root psi50 in the cross-species analysis (P < 0.001), a relationship that was robust to phylogenetic correction (P < 0.01). A and g(s) were also positively correlated with stem psi50 in the cross-species analysis (P = 0.02 and 0.10, respectively). However, only A was associated with stem psi50 in the PIC analysis (P = 0.04). Although the relationship between vulnerability to cavitation and xylem conductivity traits within specific organs (i.e. stems and roots) was weak, the strong correlation between g(s) and root psi50 across species suggests that there is a trade-off between vulnerability to cavitation and water transport capacity at the whole-plant level. Our results were therefore consistent with the expectation of coordination between vulnerability to xylem cavitation and the regulation of stomatal conductance, and highlight the potential physiological and evolutionary significance of root hydraulic properties in controlling interspecific variation in leaf function.     

Stomatal dynamics are limited by leaf hydraulics in ferns and conifers: results from simultaneous measurements of liquid and vapour fluxes in leaves

Stomatal responsiveness to vapour pressure deficit (VPD) results in continuous regulation of daytime gas‐exchange directly influencing leaf water status and carbon gain. Current models can reasonably predict steady‐state stomatal conductance (g_s) to changes in VPD but the g_s dynamics between steady‐states are poorly known. Here, we used a diverse sample of conifers and ferns to show that leaf hydraulic architecture, in particular leaf capacitance, has a major role in determining the g_s response time to perturbations in VPD. By using simultaneous measurements of liquid and vapour fluxes into and out of leaves, the in situ fluctuations in leaf water balance were calculated and appeared to be closely tracked by changes in g_s thus supporting a passive model of stomatal control. Indeed, good agreement was found between observed and predicted g_s when using a hydropassive model based on hydraulic traits. We contend that a simple passive hydraulic control of stomata in response to changes in leaf water status provides for efficient stomatal responses to VPD in ferns and conifers, leading to closure rates as fast or faster than those seen in most angiosperms.     

Leaf hydraulic safety margin and safety–efficiency trade-off across angiosperm woody species

Leaf hydraulic conductance and the vulnerability to water deficits have profound effects on plant distribution and mortality. In this study, we compiled a leaf hydraulic trait dataset with 311 species-at-site combinations from biomes worldwide. These traits included maximum leaf hydraulic conductance (K_leaf), water potential at 50% loss of K_leaf (P50_leaf), and minimum leaf water potential (Ψ_min). Leaf hydraulic safety margin (HSM_leaf) was calculated as the difference between Ψ_min and P50_leaf. Our results indicated that 70% of the studied species had a narrow HSM_leaf (less than 1 MPa), which was consistent with the global pattern of stem hydraulic safety margin. There was a positive relationship between HSM_leaf and aridity index (the ratio of mean annual precipitation to potential evapotranspiration), as species from humid sites tended to have larger HSM_leaf. We found a significant relationship between K_leaf and P50_leaf across global angiosperm woody species and within each of the different plant groups. This global analysis of leaf hydraulic traits improves our understanding of plant hydraulic response to environmental change.     

The ecological significance of long-distance water transport: short-term regulation, long-term acclimation and the hydraulic costs of stature across plant life forms

Plant hydraulic conductance, namely the rate of water flow inside plants per unit time and unit pressure difference, varies largely from plant to plant and under different environmental conditions. Herein the main factors affecting: (a) the scaling between whole‐plant hydraulic conductance and leaf area; (b) the relationship between gas exchange at the leaf level and leaf‐specific xylem hydraulic conductance; (c) the short‐term physiological regulation of plant hydraulic conductance under conditions of ample soil water, and (d) the long‐term structural acclimation of xylem hydraulic conductance to changes in environmental conditions are reviewed. It is shown that plant hydraulic conductance is a highly plastic character that varies as a result of multiple processes acting at several time scales. Across species ranging from coniferous and broad‐leaved trees to shrubs, crop and herbaceous species, and desert subshrubs, hydraulic conductance scaled linearly with leaf area, as expected from first principles. Despite considerable convergence in the scaling of hydraulic properties, significant differences were apparent across life forms that underlie their different abilities to conduct gas exchange at the leaf level. A simple model of carbon allocation between leaves and support tissues explained the observed patterns and correctly predicted the inverse relationships with plant height. Therefore, stature appears as a fundamental factor affecting gas exchange across plant life forms. Both short‐term physiological regulation and long‐term structural acclimation can change the levels of hydraulic conductance significantly. Based on a meta‐analysis of the existing literature, any change in environmental parameters that increases the availability of resources (either above‐ or below‐ground) results in the long‐term acclimation of a less efficient (per unit leaf area) hydraulic system.     

不同甘蔗品种叶片气孔对水分胁迫的响应

干旱是甘蔗面临最主要的环境胁迫之一,为了解不同甘蔗品种在干旱胁迫时的气孔响应,该研究以F172、GT21、YT93/159和 YL6四个抗旱性有显著差异的甘蔗品种为材料,采用桶栽,在伸长期进行四种不同程度的干旱胁迫(不浇水)处理：土壤持水量在①65％~70％为轻度干旱;②45％~50％为中度干旱;③25％~30％为重度干旱;④以土壤含水量为75％为对照(CK).检测不同品种不同处理甘蔗的叶片相对持水量变化,并利用扫描电镜技术观察甘蔗叶片下表皮气孔特性.结果表明：在干旱胁迫下,四个甘蔗品种叶片气孔导度急剧下降,重度干旱时耐旱性强的 F172和 GT21的气孔导度低于耐旱性弱的 YT93/159和 YL6的;复水后3 d,F172和 GT21的气孔导度上升至82．07和88．85 mmol·m-2·s-1,而 YT93/159和 YL6的仅有18．88和33．08 mmol·m-2·s-1.干旱还导致气孔下陷、闭合,气孔器的长、宽明显减小,且品种间气孔器长度变化差异显著;干旱胁迫下气孔密度增大,尤以耐旱性最强的 F172在重度干旱时达到显著差异.重度干旱时 F172与GT21的气孔闭合百分比是 YT93/159和 YL6近3~4倍.在水分胁迫下,叶片相对含水量降低,但 F172和GT21在重度干旱时仍可以保持相对较高的含水量,其它两个品种相对较低,尤以 YT93/159的最低.在复水后叶片含水量都有所恢复.这些研究结果表明不同甘蔗品种抗旱能力与叶片气孔特性和含水量密切相关.     

Stomatal response to air humidity and its relation to stomatal density in a wide range of warm climate species

The gas exchange of 19 widely different warm climate species was observed at different leaf to air vapour pressure deficits (VPD). In all species stomata tended to close as VPD increased resulting in a decrease in net photosynthesis. The absolute reduction in leaf conductance per unit increase in VPD was greatest in those species which had a large leaf conductance at low VPDs. This would be expected even if stomata of all species were equally sensitive. However the percentage reduction in net photosynthesis (used as a measure of the relative sensitivity of stomata of the different species) was also closely related to the maximal conductance at low VPD. Similarily the relative sensitivity of stomata to changes in VPD was closely related to the weighted stomatal density or crowding index.The hypothesis is presented that stomatal closure at different VPDs is related to peristomatal evaporation coupled with a high resistance between the epidermis and the mesophyll and low resistance between the stomatal apparatus and the epidermal cells. This hypothesis is consistent with the greater relative sensitivity of stomata on leaves with a high crowding index.The results and the hypothesis are discussed in the light of selection, for optimal productivity under differing conditions of relative humidity and soil water availablility, by observation of stomatal density and distribution on the two sides of the leaf.Visiting scientist, plant physiologist and research assitant of the Cassava Program     

Conservative decrease in water potential in existing leaves during new leaf expansion in temperate and tropical evergreen Quercus species

BACKGROUND AND AIMS: This study aimed at clarifying how the water potential gradient (deltapsi) is maintained in the shoots of evergreen trees with expanding leaves, whose leaf water potentials at the turgor loss point (psi(tlp)) are generally high. MATERIALS: The water relations were examined in current-year expanding (CEX) and 1-year-old (OLD) leaves on the same shoots in temperate (Osaka, Japan) and tropical (Bogor, Indonesia) areas. A temperate evergreen species, Quercus glauca growing in both sites, was compared with a temperate deciduous species, Q. serrata, in Osaka, and two tropical evergreen species, Q. gemelliflora and Q. subsericea, in Bogor. KEY RESULTS: (1) In Osaka, the midday leaf water potential (psi(midday)) was slightly higher in OLD (-0.5 MPa) than in CEX leaves (-0.6 MPa), whereas psi(tlp) was significantly lower in OLD (-2.9 MPa) than in CEX leaves (-1.0 MPa). In Bogor, psi(midday) was also higher in OLD leaves (-1.0 MPa) despite the low psi(tlp) (-1.9 MPa), although stomatal conductance was not always low in OLD leaves. In the branch bearing CEX and OLD leaves, most of the hydraulic resistance (86 %) exists in the current-year branch, leading to differences in water supply between CEX and OLD leaves. The removal of buds just before breaking did not affect the high psi(midday) in OLD leaves after 1 month. Psi(midday) in OLD leaves thus appears to be independent of that in CEX leaves. CONCLUSIONS: The moderate decrease in psi(midday) in OLD leaves would contribute to maintenance of deltapsi in the shoots during leaf expansion.     

Co-ordination of vapour and liquid phase water transport properties in plants

The pathway for water movement from the soil through plants to the atmosphere can be represented by a series of liquid and vapour phase resistances. Stomatal regulation of vapour phase resistance balances transpiration with the efficiency of water supply to the leaves, avoiding leaf desiccation at one extreme, and unnecessary restriction of carbon dioxide uptake at the other. In addition to maintaining a long-term balance between vapour and liquid phase water transport resistances in plants, stomata are exquisitely sensitive to short-term, dynamic perturbations of liquid water transport. In balancing vapour and liquid phase water transport, stomata do not seem to distinguish among potential sources of variation in the apparent efficiency of delivery of water per guard cell complex. Therefore, an apparent soil-to-leaf hydraulic conductance based on relationships between liquid water fluxes and driving forces in situ seems to be the most versatile for interpretation of stomatal regulatory behaviour that achieves relative homeostasis of leaf water status in intact plants. Components of dynamic variation in apparent hydraulic conductance in intact plants include, exchange of water between the transpiration stream and internal storage compartments via capacitive discharge and recharge, cavitation and its reversal, temperature-induced changes in the viscosity of water, direct effects of xylem sap composition on xylem hydraulic properties, and endogenous and environmentally induced variation in the activity of membrane water channels in the hydraulic pathway. Stomatal responses to humidity must also be considered in interpreting co-ordination of vapour and liquid phase water transport because homeostasis of bulk leaf water status can only be achieved through regulation of the actual transpirational flux. Results of studies conducted with multiple species point to considerable convergence with regard to co-ordination of stomatal and hydraulic properties. Because stomata apparently sense and respond to integrated and dynamic soil-to-leaf water transport properties, studies involving intact plants under both natural and controlled conditions are likely to yield the most useful new insights concerning stomatal co-ordination of transpiration with soil and plant hydraulic properties.     

Coloration and phenology manifest nutrient variability in senesced leaves of 46 temperate deciduous woody species

颜色和物候表明46种温带落叶木本植物衰老叶片的养分变异 不同共生植物的叶片养分含量差异显著,反映了不同的叶片养分利用策略。然而,衰老叶片养分的种间变异及其驱动因素尚不清楚。本研究旨在探讨衰老叶片养分的种间变异及其驱动因素。我们在中国东北的帽儿山森林生态系统研究站测定了46种共存温带落叶木本植物新鲜落叶的碳、氮、磷浓度。 采用随机森林模型量化10个生物因素(菌根类型、固氮类型、生长形态、耐阴性、叶片质地、变色程度、变色类型、叶片变色峰期、落叶峰期和落叶末期)的相对重要性。研究结果表明,落叶氮浓度种间变化为4倍,磷浓度变化达9倍。较高的氮和磷平均浓度(15.38和1.24 mg g⁻¹)表明该森林氮和磷限制较弱。功能群仅对特定养分及其比值有显著影响。磷浓度、氮磷比与外生菌根树种的落叶高峰日和落叶结束日呈负相关。颜色鲜艳的叶片(红色＞棕色＞黄色＞黄绿色＞绿色)倾向于比绿色叶片氮和磷浓度更低而碳氮比和碳磷比较高。随机森林模型表明,秋季叶变色和落叶物候贡献了80%的种间变异解释量。这些结果增加了我们对温带森林木本植物营养策略之衰老叶片养分变异性的理解。     

Photosynthetic water use efficiency in tree crowns of <Emphasis Type="Italic">Shorea beccariana</Emphasis> and <Emphasis Type="Italic">Dryobalanops aromatica</Emphasis> in a tropical rain forest in Sarawak,East Malaysia

Photosynthetic water use efficiency (PWUE), stomatal conductance (g _s), and water potential were measured at two different positions in the tree crown of two emergent tropical tree species (Shorea beccariana Burck, Dryobalanops aromatica Gaertn. f.). The trees were about 50 m high, in a tropical rain forest in Sarawak, East Malaysia. In both species, g _s at the upper crown position at midday was lower than at the lower crown position, even though both positions were exposed to full sunlight; the difference was greater in S. beccariana. Hydraulic limitation occurs in the upper crown position in both species. A midday depression was observed in the photon saturated photosynthetic rate in both species, especially at the upper crown. However, PWUE was markedly higher in the upper crown than the lower crown at midday, even though no morphological adjustment was observed in the leaves; this difference was greater in S. beccariana.     

Differential coordination of stomatal conductance,mesophyll conductance,and leaf hydraulic conductance in response to changing light across species

Stomatal conductance (g_s) and mesophyll conductance (g_m) represent major constraints to photosynthetic rate (A), and these traits are expected to coordinate with leaf hydraulic conductance (K_leaf) across species, under both steady‐state and dynamic conditions. However, empirical information about their coordination is scarce. In this study, K_leaf, gas exchange, stomatal kinetics, and leaf anatomy in 10 species including ferns, gymnosperms, and angiosperms were investigated to elucidate the correlation of H₂O and CO₂ diffusion inside leaves under varying light conditions. Gas exchange, K_leaf, and anatomical traits varied widely across species. Under light‐saturated conditions, the A, g_s, g_m, and K_leaf were strongly correlated across species. However, the response patterns of A, g_s, g_m, and K_leaf to varying light intensities were highly species dependent. Moreover, stomatal opening upon light exposure of dark‐adapted leaves in the studied ferns and gymnosperms was generally faster than in the angiosperms; however, stomatal closing in light‐adapted leaves after darkening was faster in angiosperms. The present results show that there is a large variability in the coordination of leaf hydraulic and gas exchange parameters across terrestrial plant species, as well as in their responses to changing light.     

Water movement through Quercus rubra I. Leaf water potential and conductance during polycyclic growth

Two experiments examined simultaneous changes in leaf area (A_L), root length (L_r), stomatal conductance (g_s), leaf water potential (Ψ_L), transpiration and hydraulic plant conductance per unit leaf area (G) during the first three shoot cycles of northern red oak (Quercus rubra L.) grown under favourable and controlled conditions. Each shoot cycle consisted of bud swell, stem elongation, leaf expansion and rest; roots grew almost continuously. The g_s of all leaves decreased substantially while leaves of the newest flush were expanding and increased modestly when seedling leaf area remained constant. Overall, g_s decreased. The Ψ_L of mature leaves decreased during leaf expansion and increased by an equivalent amount during intervening periods. Possible explanations for the paired changes in g_s and Ψ_L are considered. Changes in G closely paralleled those of canopy g_s. These parallel changes during polycyclic seedling growth should act to keep seedling Ψ_L relatively constant as plant size increases and thereby help prevent Ψ_L from dropping to levels that would cause runaway embolism.     

Coordination and trade‐offs between leaf and stem hydraulic traits and stomatal regulation along a spectrum of isohydry to anisohydry

The degree of plant iso/anisohydry, a widely used framework for classifying species‐specific hydraulic strategies, integrates multiple components of the whole‐plant hydraulic pathway. However, little is known about how it associates with coordination of functional and structural traits within and across different organs. We examined stem and leaf hydraulic capacitance and conductivity/conductance, stem xylem anatomical features, stomatal regulation of daily minimum leaf and stem water potential (Ψ), and the kinetics of stomatal responses to vapour pressure deficit (VPD) in six diverse woody species differing markedly in their degree of iso/anisohydry. At the stem level, more anisohydric species had higher wood density and lower native capacitance and conductivity. Like stems, leaves of more anisohydric species had lower hydraulic conductance; however, unlike stems, their leaves had higher native capacitance at their daily minimum values of leaf Ψ. Moreover, rates of VPD‐induced stomatal closure were related to intrinsic rather than native leaf capacitance and were not associated with species' degree of iso/anisohydry. Our results suggest a trade‐off between hydraulic storage and efficiency in the leaf, but a coordination between hydraulic storage and efficiency in the stem along a spectrum of plant iso/anisohydry.     

Canopy structure,vertical radiation profile and photosynthetic function in a Quercus ilex evergreen forest

The studied evergreen forest dominated by Quercus ilex showed a leaf area index (LAI) of 4.5, of which 61 % was accumulated within the tree layer, 30 % within the shrub layer, and 9 % within the herb layer. The leaves of all the species were ± horizontally oriented (41°), absorbing a relevant percentage of incident irradiance. The high LAI drastically modified the quality and quantity of solar radiation on the forest underground. The spectral distribution of the radiation under the forest was markedly deficient in blue and red wavelengths. The maximum absorption in these spectral bands was found in spring, when net photosynthetic rate (P _N ) was at its maximum, and in summer, when new leaves reached 90 % of their definitive structure. The vertical radiation profile showed an evident reduction of the red-far red ratio (R/FR). Radiation quality and quantity influenced leaf physiology and morphology. Clear differences in leaf size, leaf water content per area (LWC) and specific leaf area (SLA) on the vertical profile of the forest were observed. All the shrub species showed similar SLA (12.02 m2 kg-1, mean value). The ability to increase SLA whilst simultaneously reducing leaf thickness maximized the carbon economy. The high chlorophyll (Chl) content of shrub layer leaves (1.41 g kg-1, mean value) was an expression of shade adaptation. Both leaf morphology and leaf physiology expressed the phenotypic plasticity. Q. ilex, Phillyrea latifolia and Pistacia lentiscus of the forest shrub layer showed wide differences in leaf structure and function with respect to the same species developing under strong irradiance (low maquis): a 57 % mean increase of SLA and a 86 % mean decrease of PN. They showed high leaf plasticity. Leaf plasticity implies that the considered sclerophyllous species has an optimum developmental pattern achieving adaptation to environments. This revised version was published online in September 2006 with corrections to the Cover Date.