Kinetics of recovery of leaf hydraulic conductance and vein functionality from cavitation-induced embolism in sunflower

The kinetics of leaf vein recovery from cavitation-induced embolism was studied in plants of sunflower cv. Margot, together with the impact of vein embolism on the overall leaf hydraulic conductance (Kleaf). During the air-dehydration of leaves to leaf water potentials (Psi L) of -1.25 MPa, Kleaf was found to decrease by about 46% with respect to values recorded in well-hydrated leaves. When leaves, previously dehydrated to Psi L= -1.1 MPa (corresponding to the turgor loss point), were put in contact with water, Kleaf recovered completely in 10 min and so did leaf water potential. Functional vein density was estimated in both dehydrating and rehydrating leaves in terms of total length of red-stained veins infiltrated with a Phloxine B solution per unit leaf surface area. Veins were found to embolize (unstained) with kinetics showing a linear relationship with Kleaf so that about a 70% loss of functional veins corresponded with a Kleaf loss of 46%. Cavitated veins recovered from embolism within 10 min from the beginning of leaf rehydration. These data indicate that: (a) leaves of sunflower underwent substantial vein embolism during dehydration; (b) vein embolism and leaf hydraulic efficiency apparently recovered from dehydration completely and rapidly upon rehydration; (c) vein refilling occurred while conduits were still at more negative xylem pressures than those required for spontaneous bubble dissolution on the basis of Henry's law. The possible consistent contribution of vital mechanisms for vein refilling is discussed.     

Vein recovery from embolism occurs under negative pressure in leaves of sunflower (Helianthus annuus)

Leaf veins undergo cavitation at water potentials (Psi(leaf)) commonly experienced by field-growing plants. Theoretically, embolism reversal should not be possible until xylem pressures rise by several kilopascals of atmospheric pressure, but recent evidence suggests that embolized conduits can be refilled even when surrounded by others at substantial tension (novel refilling). The present study reports 'novel refilling' occurring in leaf veins of sunflower (Helianthus annuus L.) while at Psi(leaf) = -0.33 MPa. Sixty per cent loss of vein hydraulic conductance (K(vein)) was recorded at Psi(leaf) < -0.65 MPa, while stem hydraulic conductance (K(stem)) was unaffected even at Psi(leaf) = -1.1 MPa. Loss of K(vein) was accompanied by stomatal closure. Water-stressed plants (Psi(leaf) = -1.1 MPa) were rehydrated overnight to different target water potentials achieved by using PEG at different concentrations as irrigation medium. K(vein) recovered by 50% at Psi(leaf) = -0.47 MPa and vein refilling was complete at Psi(leaf) = -0.33 MPa, i.e. well below the theoretical limit for conduit refilling (-0.05 MPa as calculated for sunflower minor veins). Mercurials supplied to detached leaves had no effect on the refilling process. Upon rehydration, recovery of K(vein) was not paralleled by recovery of whole-plant hydraulic conductance or leaf conductance to water vapour (g(L)), as a likely consequence of hydraulic failure of other components of the water pathway (root system or extravascular leaf compartments) and/or root-to-leaf chemical signalling. This is the first study providing experimental evidence for 'novel refilling' in a herbaceous dicot and highlighting the importance of this process in the leaf.     

Declining hydraulic efficiency as transpiring leaves desiccate: two types of response

The conductance of transpiring leaves to liquid water (Kleaf) was measured across a range of steady-state leaf water potentials (Psileaf). Manipulating the transpiration rate in excised leaves enabled us to vary Psileaf in the range -0.1 MPa to less than -1.5 MPa while using a flowmeter to monitor the transpiration stream. Employing this technique to measure how desiccation affects Kleaf in 19 species, including lycophytes, ferns, gymnosperms and angiosperms, we found two characteristic responses. Three of the six angiosperm species sampled maintained a steady maximum Kleaf while Psileaf remained above -1.2 MPa, although desiccation of leaves beyond this point resulted in a rapid decline in Kleaf. In all other species measured, declining Psileaf led to a proportional decrease in Kleaf, such that midday Psileaf of unstressed plants in the field was sufficient to depress Kleaf by an average of 37%. It was found that maximum Kleaf was strongly correlated with maximum CO2 assimilation rate, while Kleaf = 0 occurred at a Psileaf slightly less negative than at leaf turgor loss. A strong linear correlation across species between Psileaf at turgor loss and Psileaf at Kleaf = 0 raises the possibility that declining Kleaf was related to declining cell turgor in the leaf prior to the onset of vein cavitation. The vulnerability of leaves rehydrating after desiccation was compared with vulnerability of leaves during steady-state evaporation, and differences between methods suggest that in many cases vein cavitation occurs only as Kleaf approaches zero.     

Stomatal closure during leaf dehydration,correlation with other leaf physiological traits

The question as to what triggers stomatal closure during leaf desiccation remains controversial. This paper examines characteristics of the vascular and photosynthetic functions of the leaf to determine which responds most similarly to stomata during desiccation. Leaf hydraulic conductance (K(leaf)) was measured from the relaxation kinetics of leaf water potential (Psi(l)), and a novel application of this technique allowed the response of K(leaf) to Psi(l) to be determined. These "vulnerability curves" show that K(leaf) is highly sensitive to Psi(l) and that the response of stomatal conductance to Psi(l) is closely correlated with the response of K(leaf) to Psi(l). The turgor loss point of leaves was also correlated with K(leaf) and stomatal closure, whereas the decline in PSII quantum yield during leaf drying occurred at a lower Psi(l) than stomatal closure. These results indicate that stomatal closure is primarily coordinated with K(leaf). However, the close proximity of Psi(l) at initial stomatal closure and initial loss of K(leaf) suggest that partial loss of K(leaf) might occur regularly, presumably necessitating repair of embolisms.     

Leaf hydraulic capacity in ferns, conifers and angiosperms: impacts on photosynthetic maxima

* The hydraulic plumbing of vascular plant leaves varies considerably between major plant groups both in the spatial organization of veins, as well as their anatomical structure. * Five conifers, three ferns and 12 angiosperm trees were selected from tropical and temperate forests to investigate whether the profound differences in foliar morphology of these groups lead to correspondingly profound differences in leaf hydraulic efficiency. * We found that angiosperm leaves spanned a range of leaf hydraulic conductance from 3.9 to 36 mmol m2 s-1 MPa-1, whereas ferns (5.9-11.4 mmol m-2 s-1 MPa-1) and conifers (1.6-9.0 mmol m-2 s-1 MPa-1) were uniformly less conductive to liquid water. Leaf hydraulic conductance (Kleaf) correlated strongly with stomatal conductance indicating an internal leaf-level regulation of liquid and vapour conductances. Photosynthetic capacity also increased with Kleaf, however, it became saturated at values of Kleaf over 20 mmol m-2 s-1 MPa-1. * The data suggest that vessels in the leaves of the angiosperms studied provide them with the flexibility to produce highly conductive leaves with correspondingly high photosynthetic capacities relative to tracheid-bearing species.     

Chloroplast response to low leaf water potentials: I. Role of turgor

The effect of decreases in turgor on chloroplast activity was studied by measuring the photochemical activity of intact sunflower (Helianthus annuus L. cv. Russian Mammoth) leaves having low water potentials. Leaf turgor, calculated from leaf water potential and osmotic potential, was found to be affected by the dilution of cell contents by water in the cell walls, when osmotic potentials were measured with a thermocouple psychrometer. After the correction of measurements of leaf osmotic potential, both the thermocouple psychrometer and a pressure chamber indicated that turgor became zero in sunflower leaves at leaf water potentials of −10 bars. Since most of the loss in photochemical activity occurred at water potentials below −10 bars, it was concluded that turgor had little effect on the photochemical activity of the leaves.     

Nutrient availability constrains the hydraulic architecture and water relations of savannah trees

Leaf and whole plant-level functional traits were studied in five dominant woody savannah species from Central Brazil (Cerrado) to determine whether reduction of nutrient limitations in oligotrophic Cerrado soils affects carbon allocation, water relations and hydraulic architecture. Four treatments were used: control, N additions, P additions and N plus P additions. Fertilizers were applied twice yearly, from October 1998 to March 2004. Sixty-three months after the first nutrient addition, the total leaf area increment was significantly greater across all species in the N- and the N + P-fertilized plots than in the control and in the P-fertilized plots. Nitrogen fertilization significantly altered several components of hydraulic architecture: specific conductivity of terminal stems increased with N additions, whereas leaf-specific conductivity and wood density decreased in most cases. Average daily sap flow per individual was consistently higher with N and N + P additions compared to the control, but its relative increase was not as great as that of leaf area. Long-term additions of N and N + P caused midday PsiL to decline significantly by a mean of 0.6 MPa across all species because N-induced relative reductions in soil-to-leaf hydraulic conductance were greater than those of stomatal conductance and transpiration on a leaf area basis. Phosphorus-fertilized trees did not exhibit significant changes in midday PsiL. Analysis of xylem vulnerability curves indicated that N-fertilized trees were significantly less vulnerable to embolism than trees in control and P-fertilized plots. Thus, N-induced decreases in midday PsiL appeared to be almost entirely compensated by increases in resistance to embolism. Leaf tissue water relations characteristics also changed as a result of N-induced declines in minimum PsiL: osmotic potential at full turgor decreased and symplastic solute content on a dry matter basis increased linearly with declining midday PsiL across species and treatments. Despite being adapted to chronic nutrient limitations, Cerrado woody species apparently have the capacity to exploit increases in nutrient availability by allocating resources to maximize carbon gain and enhance growth. The cost of increased allocation to leaf area relative to water transport capacity involved increased total water loss per plant and a decrease in minimum leaf water potentials. However, the risk of increased embolism and turgor loss was relatively low as xylem vulnerability to embolism and leaf osmotic characteristics changed in parallel with changes in plant water status induced by N fertilization.     

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.     

Roles of leaf water potential and soil-to-leaf hydraulic conductance in water use by understorey woody plants

Diurnal changes of leaf water potential and stomatal conductance were measured for 12 deciduous shrubs and tree saplings in the understorey of a temperate forest. Sunflecks raised the leaf temperature by 4°C, and vapor pressure deficit to 2 kPa. Although the duration of the sunflecks was only 17% of daytime, the photon flux density (PFD) of sunflecks was 52% of total PFD on a sunny summer day. Leaf osmotic potential at full turgor decreased in summer, except in some species that have low osmotic potential in the spring. Plants that endured low leaf water potential had rigid cell walls and low osmotic potential at full turgor. These plants did not have lower relative water content and turgor potential than plants with higher leaf water potential. There were three different responses to an increase in transpiration rate: (i) plants had low leaf water potential and slightly increased soil-to-leaf hydraulic conductance; (ii) plants decreased leaf water potential and increased the hydraulic conductance; and (iii) plants had high leaf water potential and largely increased the hydraulic conductance.     

Changes in water relations and in some physiological functions of bean under very light osmotic shock induced by polyethylene glycol

Bean plantlets ( Phaseolus vulgaris L. cv. Topcrop) were stressed at the age of 16–18 days by gradual (2–8%) or abrupt addition of 6% (w/v) polyethylene glycol Mw 6000 (PEG 6000) to Hoagland solution. Leaf conductance, photosynthesis, internal CO₂ partial pressure (C_i), relative water content (RWC), water content/dry weight (H₂O/DW), apoplastic PEG concentrations and weight of leaves, stems and roots were determined. Leaf conductance, photosynthesis and C_i were determined on non-detached primary leaves, and leaf potentials (water, osmotic and turgor potentials) were investigated in freshly detached (non-rehydrated) primary leaves, both in treated and control plants; RWC and osmotic potential were also assessed at the null turgor point. Low PEG 6000 concentrations induced early and evident decrease in leaf conductance and photosynthesis, whereas C_i decreased only moderately and tended to recover during advanced stress. There were moderate though significant decreases in RWC and H₂O/DW, no change or increases in water potential, no significant changes in osmotic potential and a moderate but significant increase in turgor potential. Even when referred to null turgor point, RWC significantly decreased and osmotic potential was unchanged. It was concluded that apoplastic PEG 6000 accumulation at evaporating sites would account for the early decrease in conductance which would also justify the unchanged or the prevalent increase in water potential and turgor potential. The subsequent PEG diffusion and concentration in the leaf apoplastic water would have induced the RWC and H₂O/DW decrease and the final turgor flexion documented.     

Heterogeneity of gas exchange rates over the leaf surface in tobacco: an effect of hydraulic architecture?

Spatial heterogeneity of gas exchange rates in the leaves of Nicotiana tabacum L. (tobacco) was investigated. Leaf conductance to water vapour was higher (by about 18%) at the apical regions of leaves than at the basal ones. Local, small-scale measurements of pressure-volume (PV) parameters and water status (performed with a dewpoint hygrometer) revealed that bulk leaf water potential, osmotic potential, turgor pressure and bulk modulus of elasticity were not significantly different in the leaf apex or base. Hydraulic measurements showed that the apical regions of the leaf blade were about 30% more conductive than the basal regions. Such differences were explained by analogous differences in terms of venation patterns. In fact, vein density turned out to be higher (by about 13%) near the leaf apex with respect to the leaf base. On the contrary, stomatal density was the same both in the apical and basal leaf portions. Our data suggest that spatial stomatal heterogeneity may arise from heterogenous distribution of local hydraulic resistances and would be addressed to maintaining local water potential above critical values, possibly triggering vein cavitation.     

Root pressurization affects growth-induced water potentials and growth in dehydrated maize leaves

Profiles of water potential (Psi w) were measured from the soil to the tips of growing leaves of maize (Zea mays L.) when pressure (P) was applied to the soil/root system. At moderately low soil Psi w, leaf elongation was somewhat inhibited, large tensions existed in the xylem, and Psi w were slightly lower in the elongating leaf tissues than in the xylem, i.e. a growth-induced Psi w was present but small. With P, the tension was relieved, enlarging the difference in Psi w between the xylem and the elongating tissues, i.e. enlarging the growth-induced Psi w, which is critical for growth. Guttation occurred, confirming the high Psi w of the xylem, and the mature leaf tissue rehydrated. Water uptake increased and met the requirements of transpiration. Leaf elongation recovered to control rates. Under more severe conditions at lower soil Psi w, P induced only a brief elongation and the growth-induced Psi w responded only slightly. Guttation did not occur, water flow did not meet the requirements of transpiration, and the mature leaf tissues did not rehydrate. A rewatering experiment indicated that a low conductance existed in the severely dehydrated soil, which limited water delivery to the root and shoot. Therefore, the initial growth inhibition appeared to be hydraulic because the enlargement of the growth-induced Psi w by P together with rehydration of the mature leaf tissue were essential for growth recovery. In more severe conditions, P was ineffective because the soil could not supply water at the required rate, and metabolic factors began to contribute to the inhibition.     

Leaf Elongation in Relation to Leaf Water Potential in Soybean

Leaf water potential, turgor pressure, and leaf elongation ratewere measured in soybeans growing in controlled environmentchambers, greenhouses, and outdoors. Plants in chambers hadthe highest water potentials and turgor pressures, and plantsoutdoors the lowest. In all three environments there was a linearrelationship between elongation rate and turgor pressure. Leavesof plants in drier environments required less turgor for elongation,and showed a greater increase in elongation rate per unit increasein turgor. Elongation rates over a 72 h period were equal inthe three environments. Leaves reached the largest final sizein the greenhouse (intermediate in water potential). Epidermalcells were larger in chamber- and greenhouse-grown leaves thanin leaves of plants grown outdoors. The number of epidermalcells per leaf was greater in the greenhouse and outdoors thanin the chamber. Leaf elongation characteristics of greenhouseplants were duplicated by mildly stressing chamber plants, andleaf elongation characteristics of field plants were duplicatedby more severely stressing chamber plants. Leaves of mildlystressed chamber plants also reached a larger final size thanleaves of more severely stressed chamber plants, or leaves ofcontrol plants in the chamber. Water stress in the chamber increasedthe number of epidermal cells per leaf. More severe water stressin the chamber reduced epidermal cell size. Based on the waterstress experiments it is concluded that the differences in plantwater status in the chamber, greenhouse, and field caused differencesin elongation characteristics, and were responsible for thedifferences in leaf size.     

Effects of water stress and rewatering on leaf water relations of lemon plants

Potted two-year-old lemon plants (Citrus limon (L.) Burm. fil.) cv. Fino, growing under field conditions were subjected to drought by withholding irrigation for 13 d. After that, plants were re-irrigated and the recovery was studied for 5 d. Control plants were daily irrigated maintaining the soil matric potential at about -30 kPa. Young leaves of control plants presented higher leaf conductance (g1) and lower midday leaf water potential (Ψmd) than mature ones. Young leaves also showed higher leaf water potential at the turgor loss point (Ψtlp) than mature leaves. In both leaf types g1 decreased with increased vapour pressure deficit of the atmosphere. From day 1 of the withholding water, predawn and midday leaf water potentials (Ψpd and Ψmd) decreased, reaching in both cases minimum values of -5.5 MPa, with no significant differences between mature and young leaves. Water stress induced stomatal closure, leaf rolling and partial defoliation. No osmotic adjustment was found in response to water stress in either leaf type, but both were able to enhance the cell wall elasticity (elastic adjustment). After rewatering, leaf water potential recovered quickly (within 2 d) but g1 did not. This revised version was published online in July 2006 with corrections to the Cover Date.     

立地条件和树龄对刺槐和小叶杨叶水力性状及抗旱性的影响

以形成黄土高原“小老树”的2种典型树种刺槐和小叶杨为对象,研究了立地条件(沟谷台地和沟间坡地)和树龄对两种树木叶水力学性质和抗旱性的影响,探讨“小老树”形成的水力生理机制.结果表明:水分较好的沟谷台地上生长的两种树木的叶最大水力导度(Kmax)明显大于水分较差的沟间坡地,叶水力脆弱性(P50)也较高;随树龄增加,两种树木的Kmax明显下降,但P50差异不大.台地上生长的两种树木的叶表皮导度和PV曲线参数(膨压损失点时的相对含水量RWCtlp、膨压损失点时的水势ψtlp饱和含水量时的渗透势ψsat)均大于 坡地;随树龄增加,两种树木的叶表皮导度显著下降,PV曲线参数出现不同程度的下降.两种树木Kmax与ψtlp呈显著正相关,P50与PV曲线参数之间存在一定的相关性,表明Kmax与抗旱性之间存在一种权衡关系,P50是反映两种树木的抗旱性特征之一.     

立地条件和树龄对刺槐和小叶杨叶水力性状及抗旱性的影响

以形成黄土高原“小老树”的2种典型树种刺槐和小叶杨为对象,研究了立地条件(沟谷台地和沟间坡地)和树龄对两种树木叶水力学性质和抗旱性的影响,探讨“小老树”形成的水力生理机制.结果表明: 水分较好的沟谷台地上生长的两种树木的叶最大水力导度(K_max)明显大于水分较差的沟间坡地,叶水力脆弱性(P₅₀)也较高;随树龄增加,两种树木的K_max明显下降,但P₅₀差异不大.台地上生长的两种树木的叶表皮导度和PV曲线参数(膨压损失点时的相对含水量RWC_tlp、膨压损失点时的水势ψ_tlp、饱和含水量时的渗透势ψ_sat)均大于坡地;随树龄增加,两种树木的叶表皮导度显著下降,PV曲线参数出现不同程度的下降.两种树木K_max与ψ_tlp呈显著正相关,P₅₀与PV曲线参数之间存在一定的相关性,表明K_max与抗旱性之间存在一种权衡关系,P₅₀是反映两种树木的抗旱性特征之一.     

Dynamics of leaf hydraulic conductance with water status: quantification and analysis of species differences under steady state

Leaf hydraulic conductance (K(leaf)) is a major determinant of photosynthetic rate in well-watered and drought-stressed plants. Previous work assessed the decline of K(leaf) with decreasing leaf water potential (Ψ(leaf)), most typically using rehydration kinetics methods, and found that species varied in the shape of their vulnerability curve, and that hydraulic vulnerability correlated with other leaf functional traits and with drought sensitivity. These findings were tested and extended, using a new steady-state evaporative flux method under high irradiance, and the function for the vulnerability curve of each species was determined individually using maximum likelihood for 10 species varying strongly in drought tolerance. Additionally, the ability of excised leaves to recover in K(leaf) with rehydration was assessed, and a new theoretical framework was developed to estimate how rehydration of measured leaves may affect estimation of hydraulic parameters. As hypothesized, species differed in their vulnerability function. Drought-tolerant species showed shallow linear declines and more negative Ψ(leaf) at 80% loss of K(leaf) (P(80)), whereas drought-sensitive species showed steeper, non-linear declines, and less negative P(80). Across species, the maximum K(leaf) was independent of hydraulic vulnerability. Recovery of K(leaf) after 1 h rehydration of leaves dehydrated below their turgor loss point occurred only for four of 10 species. Across species without recovery, a more negative P(80) correlated with the ability to maintain K(leaf) through both dehydration and rehydration. These findings indicate that resistance to K(leaf) decline is important not only in maintaining open stomata during the onset of drought, but also in enabling sustained function during drought recovery.     

Stomatal action directly feeds back on leaf turgor: new insights into the regulation of the plant water status from non‐invasive pressure probe measurements

Uptake of CO₂ by the leaf is associated with loss of water. Control of stomatal aperture by volume changes of guard cell pairs optimizes the efficiency of water use. Under water stress, the protein kinase OPEN STOMATA 1 (OST1) activates the guard‐cell anion release channel SLOW ANION CHANNEL‐ASSOCIATED 1 (SLAC1), and thereby triggers stomatal closure. Plants with mutated OST1 and SLAC1 are defective in guard‐cell turgor regulation. To study the effect of stomatal movement on leaf turgor using intact leaves of Arabidopsis, we used a new pressure probe to monitor transpiration and turgor pressure simultaneously and non‐invasively. This probe permits routine easy access to parameters related to water status and stomatal conductance under physiological conditions using the model plant Arabidopsis thaliana. Long‐term leaf turgor pressure recordings over several weeks showed a drop in turgor during the day and recovery at night. Thus pressure changes directly correlated with the degree of plant transpiration. Leaf turgor of wild‐type plants responded to CO₂, light, humidity, ozone and abscisic acid (ABA) in a guard cell‐specific manner. Pressure probe measurements of mutants lacking OST1 and SLAC1 function indicated impairment in stomatal responses to light and humidity. In contrast to wild‐type plants, leaves from well‐watered ost1 plants exposed to a dry atmosphere wilted after light‐induced stomatal opening. Experiments with open stomata mutants indicated that the hydraulic conductance of leaf stomata is higher than that of the root–shoot continuum. Thus leaf turgor appears to rely to a large extent on the anion channel activity of autonomously regulated stomatal guard cells.     

Response to Saturation Deficit of Leaf Extension in a Stand of Pearl Millet (Pennisetum typhoides S. & H.)II: II. DEPENDENCE ON LEAF WATER STATUS AND IRRADIANCE

This paper describes how the dominant relation between leafextension and temperature in pearl millet is modified by atmosphericsaturation vapour pressure deficit (SD) and irradiance. Standsof plants were grown at two levels of SD and soil moisture content.Leaf extension, water potential (₁) and stomatal conductancewere all reduced at high SD, ₁ was more closely related to transpirationrate than to SD itself. Leaf extension rate (R) was poorly correlatedwith ₁, even after correction for temperature differences, owingto variation in solute potential between leaves. However, Rin individual leaves was linearly related to turgor potential,except after periods of low irradiance. The thermal time conceptwas modified to incorporate turgor potential and used to showthat the ‘turgor thermal rate of extension’ decreasedsharply at low irradiances, presumably due to assimilate shortage. Key words: Extension, Saturation deficit, Millet     

Leaf water relations during summer water deficit: differential responses in turgor maintenance and variation in leaf structure among different plant communities in south-western Australia

We measured leaf water relations and leaf structural traits of 20 species from three communities growing along a topographical gradient. Our aim was to assess variation in seasonal responses in leaf water status and leaf tissue physiology between sites and among species in response to summer water deficit. Species from a ridge-top heath community showed the greatest reductions in pre-dawn leaf water potentials (Psi(leaf)) and stomatal conductance during summer; species from a valley-floor woodland and a midslope mallee community showed less reductions in these parameters. Heath species also displayed greater seasonal reduction in turgor-loss point (Psi(TLP)) than species from woodland or mallee communities. In general, species that had larger reductions in Psi(leaf) during summer showed significant shifts in either their osmotic potential at full turgor (Psi(pi 100); osmotic adjustment) or in tissue elasticity (epsilon(max)). Psi(pi 100) and epsilon(max) were negatively correlated, during both spring and summer, suggesting a trade-off between these different mechanisms to cope with water stress. Specific leaf area varied greatly among species, and was significantly correlated with seasonal changes in Psi(TLP) and pre-dawn Psi(leaf). These correlations suggest that leaf structure is a prerequisite for cellular mechanisms to be effective in adjusting to water deficit.