Limitations to photosynthesis of lettuce grown under tropical conditions: alleviation by root-zone cooling.

Aerial parts of lettuce plants were grown under natural tropical fluctuating ambient temperatures, but with their roots exposed to two different root-zone temperatures (RZTs): a constant 20 degrees C-RZT and a fluctuating ambient (A-) RZT from 23-40 degrees C. Plants grown at A-RZT showed lower photosynthetic CO2 assimilation (A), stomatal conductance (gs), midday leaf relative water content (RWC), and chlorophyll fluorescence ratio Fv/Fm than 20 degrees C-RZT plants on both sunny and cloudy days. Substantial midday depression of A and g(s) occurred on both sunny and cloudy days in both RZT treatments, although Fv/Fm did not vary diurnally on cloudy days. Reciprocal temperature transfer experiments investigated the occurrence and possible causes of stomatal and non-stomatal limitations of photosynthesis. For both temperature transfers, light-saturated stomatal conductance (gs sat) and photosynthetic CO2 assimilation (A(sat)) were highly correlated with each other and with midday RWC, suggesting that A was limited by water stress-mediated stomatal closure. However, prolonged growth at A-RZT reduced light- and CO2-saturated photosynthetic O2 evolution (Pmax), indicating non-stomatal limitation of photosynthesis. Tight temporal coupling of leaf nitrogen content and P(max) during both temperature transfers suggested that decreased nutrient status caused this non-stomatal limitation of photosynthesis.     

Physiological effects of kaolin applications in well-irrigated and water-stressed walnut and almond trees

BACKGROUND AND AIMS: Kaolin applications have been used to mitigate the negative effects of water and heat stress on plant physiology and productivity with variable results, ranging from increased to decreased yields and photosynthetic rates. The mechanisms of action of kaolin applications are not clear: although the increased albedo reduces leaf temperature and the consequent heat stress, it also reduces the light available for photosynthesis, possibly offsetting benefits of lower temperature. The objective of this study was to investigate which of these effects are prevalent and under which conditions. METHODS: A 6% kaolin suspension was applied on well-irrigated and water-stressed walnut (Juglans regia) and almond (Prunus dulcis) trees. Water status (i.e. stem water potential, psi(s)), gas exchange (i.e. light-saturated CO2 assimilation rate, Amax; stomatal conductance, g(s)), leaf temperature (T(l)) and physiological relationships in treated and control trees were then measured and compared. KEY RESULTS: In both species, kaolin did not affect the daily course of psi(s) whereas it reduced Amax by 1-4 micromol CO2 m(-2) s(-1) throughout the day in all combinations of species and irrigation treatments. Kaolin did not reduce g(s) in any situation. Consequently, intercellular CO2 concentration (C(i)) was always greater in treated trees than in controls, suggesting that the reduction of Amax with kaolin was not due to stomatal limitations. Kaolin reduced leaf temperature (T(l)) by about 1-3 degrees C and leaf-to-air vapour pressure difference (VPD(l)) by about 0.1-0.7 kPa. Amax was lower at all values of g(s), T(l) and VPD(l) in kaolin-treated trees. Kaolin affected the photosynthetic response to the photosynthetically active radiation (PAR) in almond leaves: kaolin-coated leaves had similar dark respiration rates and light-saturated photosynthesis, but a higher light compensation point and lower apparent quantum yield, while the photosynthetic light-response curve saturated at higher PAR. When these parameters were used to model the photosynthetic response curve to PAR, it was estimated that the kaolin film allowed 63% of the incident PAR to reach the leaf. CONCLUSIONS: The main effect of kaolin application was the reduction, albeit minor, of photosynthesis, which appeared to be related to the shading of the leaves. The reduction in T(l) and VPD(l) with kaolin did not suffice to mitigate the adverse effects of heat and water stress on Amax.     

Modelling photosynthetic responses to temperature of grapevine (Vitis vinifera cv. Semillon) leaves on vines grown in a hot climate

Field measurements of photosynthesis of Vitis vinifera cv. Semillon leaves in relation to a hot climate, and responses to photon flux densities (PFDs) and internal CO(2) concentrations (c(i) ) at leaf temperatures from 20 to 40 °C were undertaken. Average rates of photosynthesis measured in situ decreased with increasing temperature and were 60% inhibited at 45 °C compared with 25 °C. This reduction in photosynthesis was attributed to 15-30% stomatal closure. Light response curves at different temperatures revealed light-saturated photosynthesis optimal at 30 °C but also PFDs saturating photosynthesis increased from 550 to 1200 μmol (photons) m(-2)s(-1) as temperatures increased. Photosynthesis under saturating CO(2) concentrations was optimal at 36 °C while maximum rates of ribulose 1,5-bisphosphate (RuBP) carboxylation (V(cmax)) and potential maximum electron transport rates (J(max)) were also optimal at 39 and 36 °C, respectively. Furthermore, the high temperature-induced reduction in photosynthesis at ambient CO(2) was largely eliminated. The chloroplast CO(2) concentration at the transition from RuBP regeneration to RuBP carboxylation-limited assimilation increased steeply with an increase in leaf temperature. Semillon assimilation in situ was limited by RuBP regeneration below 30 °C and above limited by RuBP carboxylation, suggesting high temperatures are detrimental to carbon fixation in this species.     

Internal conductance to CO(2) diffusion and C(18)OO discrimination in C(3) leaves

(18)O discrimination in CO(2) stems from the oxygen exchange between (18)O-enriched water and CO(2) in the chloroplast, a process catalyzed by carbonic anhydrase (CA). A proportion of this (18)O-labeled CO(2) escapes back to the atmosphere, resulting in an effective discrimination against C(18)OO during photosynthesis (Delta(18)O). By constraining the delta(18)O of chloroplast water (delta(e)) by analysis of transpired water and the extent of CO(2)-H(2)O isotopic equilibrium (theta(eq)) by measurements of CA activity (theta(eq) = 0.75-1.0 for tobacco, soybean, and oak), we could apply measured Delta(18)O in a leaf cuvette attached to a mass spectrometer to derive the CO(2) concentration at the physical limit of CA activity, i.e. the chloroplast surface (c(cs)). From the CO(2) drawdown sequence between stomatal cavities from gas exchange (c(i)), from Delta(18)O (c(cs)), and at Rubisco sites from Delta(13)C (c(c)), the internal CO(2) conductance (g(i)) was partitioned into cell wall (g(w)) and chloroplast (g(ch)) components. The results indicated that g(ch) is variable (0.42-1.13 mol m(-2) s(-1)) and proportional to CA activity. We suggest that the influence of CA activity on the CO(2) assimilation rate should be important mainly in plants with low internal conductances.     

Growth and photosynthetic responses to salinity of the salt-marsh shrub Atriplex portulacoides

BACKGROUND AND AIMS: Atriplex (Halimione) portulacoides is a halophytic, C(3) shrub. It is virtually confined to coastal salt marshes, where it often dominates the vegetation. The aim of this study was to investigate its growth responses to salinity and the extent to which these could be explained by photosynthetic physiology. METHODS: The responses of young plants to salinity in the range 0-700 mol m(-3) NaCl were investigated in a glasshouse experiment. The performance of plants was examined using classical growth analysis, measurements of gas exchange (infrared gas analysis), determination of chlorophyll fluorescence characteristics (modulated fluorimeter) and photosynthetic pigment concentrations; total ash, sodium, potassium and nitrogen concentrations, and relative water content were also determined. KEY RESULTS: Plants accumulated Na(+) approximately in proportion to external salinity. Salt stimulated growth up to an external concentration of 200 mol m(-3) NaCl and some growth was maintained at higher salinities. The main determinant of growth response to salinity was unit leaf rate. This was itself reflected in rates of CO(2) assimilation, which were not affected by 200 mol m(-3) but were reduced at higher salinities. Reductions in net photosynthetic rate could be accounted for largely by lower stomatal conductance and intercellular CO(2) concentration. Apart from possible effects of osmotic shock at the beginning of the experiment, salinity did not have any adverse effect on photosystem II (PSII). Neither the quantum efficiency of PSII (Phi(PSII)) nor the chlorophyll fluorescence ratio (F(v)/F(m)) were reduced by salinity, and lower mid-day values recovered by dawn. Mid-day F(v)/F(m) was in fact depressed more at low external sodium concentration, by the end of the experiment. CONCLUSIONS: The growth responses of the hygro-halophyte A. portulacoides to salinity appear largely to depend on changes in its rate of photosynthetic gas exchange. Photosynthesis appears to be limited mainly through stomatal conductance and hence intercellular CO(2) concentration, rather than by effects on PSII; moderate salinity might stimulate carboxylation capacity. This is in contrast to more extreme halophytes, for which an ability to maintain leaf area can partially offset declining rates of carbon assimilation at high salinity.     

Difference in sodium spatial distribution in the shoot of two canola cultivars under saline stress

Among different mechanisms of salt resistance, regulation of ion distribution among various tissues and intracellular compartmentation are of great importance. In this study, we investigated the effects of salt stress on growth, photosynthesis, and Na(+) accumulation and distribution in leaf apoplast and symplast of two canola (Brassica napus L.) cultivars (NYY 1 and BZY 1). The results showed that the declines in shoot dry mass, leaf water potential and net photosynthetic rate of BZY 1 (salt sensitive) were higher than those of NYY 1 (salt resistant) in response to salt stress. Stomatal limitation to photosynthesis was mainly affected under moderate salinity, whereas the reduction in assimilation rate under severe salt stress was due to both stomatal and non-stomatal limitations. We also found that more Na(+) was distributed to leaf veins in NYY 1 than in BZY 1; simultaneously, less Na(+) accumulated in the leaf blade in NYY 1 than in BZY 1. The percentage of Na(+) in the leaf symplast in NYY 1 was markedly lower than that in BZY 1. Also, Na(+) diffusion in leaves through apoplastic and symplastic pathways of BZY 1 was stronger than that in NYY 1, and the transpiration rate in BZY 1, especially at the leaf edges, decreased more than in NYY 1. Our results showed that NYY 1 accumulated less Na(+) in the shoot, especially in leaf blades, and confined Na(+) to the apoplast to avoid leaf salt toxicity, which could be one reason for the higher resistance of NYY 1 than BZY 1 plants to salt stress.     

Photosynthetic and stomatal responses of the halophyte,Plantago maritima L. to fluctuations in salinity

Abstract Measurements of photosynthesis as a function of intercellular CO₂ (A-C₁ curve) were made on single. attached leaves of Plantago maritima L. while plants were exposed to changes in salinity. Salinity was increased in steps from 50 to 500 mol m^-3 NaCl and then returned to 50 mol m^-3 NaCl at two rates, 75 mol m^-3 (NaCl) day^-1 (experiment 1) and 150 mol m^-3 (NaCl) day^-1 (experiment 2). In experiment one, the CO₂ assimilation rate declined at high CO₂ concentrations, but the initial slope of the A-C₁ curve was unaffected in young leaves after salinity was increased to 500 mol m^-3 NaCl. The insensitivity of photosynthesis to increases in CO₂ concentration above air levels was not associated with insensitivity to a reduction in oxygen concentration. In experiment two increasing the rate at which salinity was changed resulted in larger declines in photosynthesis and leaf conductance than were observed in experiment one. Both the initial slope and the CO₂ saturated region of the A-C₁ curve were substantially reduced at high salinity suggesting that mesophyll biochemical capacity had been inhibited. However, concurrent measurements of photosynthesis as oxygen evolution under 5% CO₂ indicated no effect of increased salinity on photosynthetic capacity. This suggests that the apparent non-stomatal limitations indicated by A-C₁ measurements were artifacts caused by strong, nonuniform stomatal closure.     

Photosynthetic responses of the tropical spiny shrub Lycium nodosum (Solanaceae) to drought, soil salinity and saline spray

Water relations and photosynthetic characteristics of plants of Lycium nodosum grown under increasing water deficit (WD), saline spray (SS) or saline irrigation (SI) were studied. Plants of this perennial, deciduous shrub growing in the coastal thorn scrubs of Venezuela show succulent leaves which persist for approx. 1 month after the beginning of the dry season; leaf succulence is higher in populations closer to the sea. These observations suggested that L. nodosum is tolerant both to WD and salinity. In the glasshouse, WD caused a marked decrease in the xylem water potential (psi), leaf osmotic potential (psi(s)) and relative water content (RWC) after 21 d; additionally, photosynthetic rate (A), carboxylation efficiency (CE) and stomatal conductance (gs) decreased by more than 90 %. In contrast, in plants treated for 21 d with a foliar spray with 35 per thousand NaCl or irrigation with a 10 % NaCl solution, psi and RWC remained nearly constant, while psi(s) decreased by 30 %, and A, CE and gs decreased by more than 80 %. An osmotic adjustment of 0.60 (SS) and 0.94 MPa (SI) was measured. Relative stomatal and mesophyll limitations to A increased with both WD and SS, but were not determined for SI-treated plants. No evidence of chronic photoinhibition due to any treatment was observed, since maximum quantum yield of PSII, Fv/Fm, did not change with either drought in the field or water or salinity stress in the glasshouse. Nevertheless, WD and SI treatments caused a decrease in the photochemical (qP) and an increase in the non-photochemical (qN) quenching coefficients relative to controls; qN was unaffected by the SS treatment. The occurrence of co-limitation of A by stomatal and non-stomatal factors in plants of L. nodosum may be associated with the extended leaf duration under water or saline stress. Additionally, osmotic adjustment may partly explain the relative maintenance of A and gs in the SS and SI treatments and the tolerance to salinity of plants of this species in coastal habitats.     

Effects of nitrogen deficiency on leaf photosynthetic response of tall fescue to water deficit

Abstract. The objective of the present work was to study the effect of nitrogen deficiency on drought sensitivity of tall fescue plants. The authors compared photosynthetic and stomatal behaviour of plants grown at either high (8 mol m⁻³) or low (0.5 mol m⁻³) nitrogen levels during a drought cycle followed by rehydration. Other processes investigated were stomatal and non-stomatal inhibition of leaf photosynthesis, water use efficiency and leaf rolling. Plants were grown in pots in controlled conditions on expanded clay. A Wescor in situ hygrometer placed on the leaf base outside the assimilation chamber permitted, simultaneously to leaf gas exchange measurements, monitoring of leaf water potential. Drought was imposed by withholding water from the pot. CO₂ uptake and stomatal conductance decreased and leaves started to roll at a lower leaf water potential in the high-N than in the low-N grown plants. Stomatal inhibition of leaf photosynthesis seemed larger in the low-N than in the high-N plants. Water-use efficiency increased more in the high-N than in the low-N grown plants during the drought. The decrease of photosynthesis was largely reversible after rehydration in low-N but not in high-N leaves. The authors suggest that low-N plants avoid water deficit rather than tolerate it.     

Salt stress and fluctuating light have separate effects on photosynthetic acclimation,but interactively affect biomass

In nature, soil salinity and fluctuating light (FL) often occur concomitantly. However, it is unknown whether salt stress interacts with FL on leaf photosynthesis, architecture, biochemistry, pigmentation, mineral concentrations, as well as whole-plant biomass. To elucidate this, tomato (Solanum lycopersicum) seedlings were grown under constant light (C, 200 μmol m⁻² s⁻¹) or FL (5–650 μmol m⁻² s⁻¹), in combination with no (0 mM NaCl) or moderate (80 mM NaCl) salinity, for 14 days, at identical photoperiods and daily light integrals. FL and salt stress had separate effects on leaf anatomy, biochemistry and photosynthetic capacity: FL reduced leaf thickness as well as nitrogen, chlorophyll and carotenoid contents per unit leaf area, but rarely affected steady-state and dynamic photosynthetic properties along with abundance of key proteins in the electron transport chain. Salt stress, meanwhile, mainly disorganized chloroplast grana stacking, reduced stomatal density, size and aperture as well as photosynthetic capacity. Plant biomass was affected interactively by light regime and salt stress: FL reduced biomass in salt stressed plants by 17%, but it did not affect biomass of non-stressed plants. Our results stress the importance of considering FL when inferring effects of salt-stress on photosynthesis and productivity under fluctuating light intensities.     

The effect of exogenous abscisic acid on stomatal development, stomatal mechanics, and leaf gas exchange in Tradescantia virginiana

Gas exchange parameters and stomatal physical properties were measured in Tradescantia virginiana plants grown under well-watered conditions and treated daily with either distilled water (control) or 3.0 mM abscisic acid (ABA). Photosynthetic capacity (CO(2) assimilation rate for any given leaf intercellular CO(2) concentration [c(i)]) and relative stomatal sensitivity to leaf-to-air vapor-pressure difference were unaffected by the ABA treatment. However, at an ambient CO(2) concentration (c(a)) of 350 micromol mol(-1), ABA-treated plants operated with significantly lower c(i). ABA-treated plants had significantly smaller stomata and higher stomatal density in their lower epidermis. Stomatal aperture versus guard cell pressure (P(g)) characteristics measured with a cell pressure probe showed that although the form of the relationship was similar in control and ABA-treated plants, stomata of ABA-treated plants exhibited more complete closure at P(g) = 0 MPa and less than half the aperture of stomata in control plants at any given P(g). Scaling from stomatal aperture versus P(g) to stomatal conductance versus P(g) showed that plants grown under ABA treatment would have had significantly lower maximum stomatal conductance and would have operated with lower stomatal conductance for any given guard cell turgor. This is consistent with the observation of lower c(i)/c(a) in ABA-treated plants with a c(a) of 350 micromol mol(-1). It is proposed that the ABA-induced changes in stomatal mechanics and stomatal conductance versus P(g) characteristics constitute an improvement in water-use efficiency that may be invoked under prolonged drought conditions.     

Relationships between growth and gas exchange characteristics in some salt-tolerant amphidiploid Brassica species in relation to their diploid parents

Relationships between growth and different gas exchange characteristics of two amphidiploid salt tolerant species, Brassica napus, and B. carinata with respect to their salt sensitive parents, B. oleracea, and B. nigra were investigated. Twenty three-day old plants of these four species along with those of another amphidiploid moderately salt tolerant B. juncea (developed by hybridization of diploids, B. campestris and B. nigra), and a diploid moderately salt tolerant, B. campestris, were subjected for 28 days to salinized sand culture containing 0, 100 or 200 mol NaCl m(-3) in Hoagland's nutrient solution. The species B. napus and B. carinata produced significantly greater shoot fresh and dry matters than their parents under saline conditions. A close association was found between growth, and assimilation rate for all species differing in degree of salt tolerance. Stomatal conductance (g(s)) was reduced due to salt stress in all species but this variable had no significant correlation with assimilation rate (A). However, the amphidiploid salt tolerant species, B. napus and B. carinata had significantly greater photosynthetic rate, water use efficiency (A/E), intrinsic water use efficiency (A/g(s)) than those of their diploid parents. In conclusion, high salt tolerance of the two amphidiploid species, B. napus and B. carinata was associated with a high assimilation rate, water use efficiency and intrinsic water use efficiency but there was little association of the tolerance of these species with stomatal conductance, leaf water potential or transpiration rate (E).     

Stomatal and non-stomatal limitations to carbon assimilation: an evaluation of the path-dependent method

Abstract Environmental stresses can decrease photosynthesis by a direct effect on photosynthetic capacity of the mesophyll or by a CO₂ limitation resulting from stomatal closure. In the present study, a ‘path-dependent method’ (Jones, 1985) for the partitioning of a stress-related decline in assimilation rate between non-stomatal and stomatal factors was evaluated, using light quality as a ‘stress’. Kinetic data on assimilation rate and conductance of Phragmipedium longifolium following a change in light quality from 95 μmol m^?2s^?1 white light to 95 μmol m^?2s^?1 red light failed to generate a smooth response curve for conductance. Partitioning of limitations on assimilation by a path-dependent method that utilizes the actual trajectories of conductance and assimilation was therefore not feasible. A simplified path-dependent method (Jones, 1985) which assumes that either mesophyll cells or guard cells respond first to a stress was applied to steady-state measurements of assimilation and conductance under red and white illumination. Either 5% or 23% of the observed reduction in assimilation rate under white light was attributable to stomatal factors, depending on whether the ‘stomatal first’ or the ‘mesophyll first’ path was assumed. In the absence of additional information indicating the appropriate choice of path, arbitrary choice may therefore lead to widely divergent estimates, and potentially erroneous conclusions. An alternative approach to the evaluation of the importance to carbon assimilation of stomatal and non-stomatal factors is suggested.     

Salicylic acid treatment via the rooting medium interferes with stomatal response, CO2 fixation rate and carbohydrate metabolism in tomato, and decreases harmful effects of subsequent salt stress

Salicylic acid (SA) applied at 10(-3) m in hydroponic culture decreased stomatal conductance (g(s)), maximal CO(2) fixation rate (A(max) ) and initial slopes of the CO(2) (A/C(i)) and light response (A/PPFD) curves, carboxylation efficiency of Rubisco (CE) and photosynthetic quantum efficiency (Q), resulting in the death of tomato plants. However, plants could acclimate to lower concentrations of SA (10(-7) -10(-4) m) and, after 3 weeks, returned to control levels of g(s), photosynthetic performance and soluble sugar content. In response to high salinity (100 mm NaCl), the pre-treated plants exhibited higher A(max) as a function of internal CO(2) concentration (C(i) ) or photosynthetic photon flux density (PPFD), and higher CE and Q values than salt-treated controls, suggesting more effective photosynthesis after SA treatment. Growth in 10(-7) or 10(-4) m SA-containing solution led to accumulation of soluble sugars in both leaf and root tissues, which remained higher in both plant parts during salt stress at 10(-4) m SA. The activity of hexokinase (HXK) with glucose, but not fructose, as substrate was reduced by SA treatment in leaf and root samples, leading to accumulation of glucose and fructose in leaf tissues. HXK activity decreased further under high salinity in both plant organs. The accumulation of soluble sugars and sucrose in roots of plants growing in the presence of 10(-4) m SA contributed to osmotic adjustment and improved tolerance to subsequent salt stress. Apart from its putative role in delaying senescence, decreased HXK activity may divert hexoses from catabolic reactions to osmotic adaptation.     

Drought-inhibition of photosynthesis in C3 plants: stomatal and non-stomatal limitations revisited

There is a long-standing controversy as to whether drought limits photosynthetic CO2 assimilation through stomatal closure or by metabolic impairment in C3 plants. Comparing results from different studies is difficult due to interspecific differences in the response of photosynthesis to leaf water potential and/or relative water content (RWC), the most commonly used parameters to assess the severity of drought. Therefore, we have used stomatal conductance (g) as a basis for comparison of metabolic processes in different studies. The logic is that, as there is a strong link between g and photosynthesis (perhaps co-regulation between them), so different relationships between RWC or water potential and photosynthetic rate and changes in metabolism in different species and studies may be 'normalized' by relating them to g. Re-analysing data from the literature using light-saturated g as a parameter indicative of water deficits in plants shows that there is good correspondence between the onset of drought-induced inhibition of different photosynthetic sub-processes and g. Contents of ribulose bisphosphate (RuBP) and adenosine triphosphate (ATP) decrease early in drought development, at still relatively high g (higher than 150 mmol H20 m(-2) s(-1)). This suggests that RuBP regeneration and ATP synthesis are impaired. Decreased photochemistry and Rubisco activity typically occur at lower g (<100 mmol H20 m(-2) s(-1)), whereas permanent photoinhibition is only occasional, occurring at very low g (<50 mmol H20 m(-2) s(-1)). Sub-stomatal CO2 concentration decreases as g becomes smaller, but increases again at small g. The analysis suggests that stomatal closure is the earliest response to drought and the dominant limitation to photosynthesis at mild to moderate drought. However, in parallel, progressive down-regulation or inhibition of metabolic processes leads to decreased RuBP content, which becomes the dominant limitation at severe drought, and thereby inhibits photosynthetic CO2 assimilation.     

The photosynthetic limitation posed by internal conductance to CO2 movement is increased by nutrient supply

The internal conductance to CO(2) supply from substomatal cavities to sites of carboxylation may pose a large limitation to photosynthesis, but little is known of how it is affected by nutrient supply. Knowing how internal conductance responds to nutrient supply is critical for interpreting the biochemical responses from A-C(i) curves. The aim of this paper was to examine the response of g(i) and photosynthetic parameters to nutrient supply in glasshouse-grown seedlings of the evergreen perennial Eucalyptus globulus Labill. Seedlings were grown with five different nutrient treatments and g(i) was estimated from concurrent measurements of gas exchange and fluorescence. Internal conductance varied between 0.12 and 0.19 mol m(-2) s(-1) and the relative limitation of photosynthesis due to internal conductance was greater than the stomatal limitation. In most species these two limitations are rather similar, but in E. globulus stomatal limitations were abnormally low due to high stomatal conductance (0.31 to 0.39 mol m(-2) s(-1)). The large positive response of photosynthesis to nutrient supply was not matched by changes in internal conductance, and thus the relative limitation of photosynthesis due to internal conductance increased with increasing nutrient supply. Failure to account for finite internal conductance led to estimates of V(cmax) that were 60% of the true value, which, in turn, led to an underestimation of in vivo Rubisco specific activity (as V(cmax)/Rubisco content). The specific activity of Rubisco in E. globulus (21 mol mol(-1) s(-1)) was close to the maximum published estimates, and thus, despite these leaves containing a large fraction of N as Rubisco (38-44%) there was no evidence that Rubisco activity was down-regulated or that the enzyme was in excess.     

Higher rates of leaf gas exchange are associated with higher leaf hydrodynamic pressure gradients

Steady-state leaf gas-exchange parameters and leaf hydraulic conductance were measured on 10 vascular plant species, grown under high light and well-watered conditions, in order to test for evidence of a departure from hydraulic homeostasis within leaves as hydraulic conductance varied across species. The plants ranged from herbaceous crop plants to mature forest trees. Across species, under standardized environmental conditions (saturating light, well watered), mean steady-state stomatal conductance to water vapour (g(w)) was highly correlated with mean rate of CO2 assimilation (A) and mean leaf hydraulic conductance normalized to leaf area (k(leaf)). The relationship between A and g(w) was well described by a power function, while that between A and k(leaf) was highly linear. Non-linearity in the relationship between g(w) and k(leaf) contributed to an increase in the hydrodynamic (transpiration-induced) water potential drawdown across the leaf (delta psi(leaf)) as k(leaf) increased across species, although across the 10 species the total increase in delta psi(leaf) was slightly more than twofold for an almost 30-fold increase in g(w). Higher rates of leaf gas exchange were therefore associated with higher k(leaf) and higher leaf hydrodynamic pressure gradients. A mechanistic model incorporating the stomatal hydromechanical feedback loop is used to predict the relationship between delta psi(leaf) and k(leaf), and to explore the coordination of stomatal and leaf hydraulic properties in supporting higher rates of leaf gas exchange.     

Photosynthesis and Transpiration as a Function of Gaseous Diffusive Resistances in Orange Leaves

The CO₂ and H₂O vapour exchange of single attached orange, Citrus sinensis (L.), leaves was measured under laboratory conditions using infrared gas analysis. Gaseous diffusive resistances were derived from measurements at a saturating irradiance and at a leaf temperature optimum for photosynthesis. Variation in leaf resistance (within the range 1.6 to 60 s cm^-1) induced by moisture status, or by cyclic oscillations in stomatal aperture, was associated with changes in both photosynthesis and transpiration. At low leaf resistance (ri less than 10 s cm^-1) the ratio of transpiration to photosynthesis declined with reduced stomatal aperture, indicating a tighter stomatal control over H₂O vapour loss than over CO₂ assimilation. At higher leaf resistance (ri greater than 10 s cm^-1) changes in transpiration and photosynthesis were linearly related, but leaf resistance and mesophyll resistance were also positively correlated, so that strictly stomatal control of photosynthesis became more apparent than real. This evidence, combined with direct measurements of CO₂ diffusive resistances (in a -O₂ gas stream) emphasised the presence of a significant mesophyll resistance; i.e., an additional and rate limiting resistance to CO₂ assimilation over and above that encountered by H₂O vapour escaping from the leaf.     

The role of stomatal acclimation in modelling tree adaptation to high CO2

Carbon dioxide enrichment changes the balance of photosynthetic limitations due to water, nitrogen, and light. This paper examines the role of stomata in these changes by comparing enrichment responses predicted by an optimality-based tree growth model, DESPOT, using three alternative 'setpoints' for stomatal acclimation: leaf water potential (psi(l)-setpoint), the ratio of intercellular to ambient CO(2) mole fraction (c(i)/c(a)-setpoint), and the parameters in a simple model in which stomata are controlled by H(2)O and CO(2) supply and demand (linked feedback). In each scenario, stomatal conductance (g(s)) and photosynthetic capacity (V(m)) declined, productivity and leaf area index (LAI) increased, and c(i)/c(a) remained within 5% of its pre-enrichment value. Height growth preceded the LAI response in the psi(l)-setpoint and linked feedback scenarios, but not in the c(i)/c(a)-setpoint scenario. These trends were explained in terms of photosynthetic resource substitution using the equimarginal principle of production theory, which controls carbon allocation in DESPOT: enrichment initially increased the marginal product for light, driving substitution towards light; height growth also drove substitution towards N in the psi(l) and feedback scenarios, but the inflexibility of c(i)/c(a) prevented that substitution in the c(i)/c(a) scenario, explaining the lack of height response. Each scenario, however, predicted similar behaviour for c(i)/c(a) and carbon and water flux. These results suggest that 'setpoints' may be robust tools for linking and constraining carbon and water fluxes, but that they should be used more cautiously in predicting or interpreting how those fluxes arise from changes in tree structure and physiology.     

The contribution of photosynthesis to the red light response of stomatal conductance

To determine the contribution of photosynthesis on stomatal conductance, we contrasted the stomatal red light response of wild-type tobacco (Nicotiana tabacum 'W38') with that of plants impaired in photosynthesis by antisense reductions in the content of either cytochrome b(6)f complex (anti-b/f plants) or Rubisco (anti-SSU plants). Both transgenic genotypes showed a lowered content of the antisense target proteins in guard cells as well as in the mesophyll. In the anti-b/f plants, CO(2) assimilation rates were proportional to leaf cytochrome b(6)f content, but there was little effect on stomatal conductance and the rate of stomatal opening. To compare the relationship between photosynthesis and stomatal conductance, wild-type plants and anti-SSU plants were grown at 30 and 300 micromol photon m(-2) s(-1) irradiance (low light and medium light [ML], respectively). Growth in ML increased CO(2) assimilation rates and stomatal conductance in both genotypes. Despite the significantly lower CO(2) assimilation rate in the anti-SSU plants, the differences in stomatal conductance between the genotypes were nonsignificant at either growth irradiance. Irrespective of plant genotype, stomatal density in the two leaf surfaces was 2-fold higher in ML-grown plants than in low-light-grown plants and conductance normalized to stomatal density was unaffected by growth irradiance. We conclude that the red light response of stomatal conductance is independent of the concurrent photosynthetic rate of the guard cells or of that of the underlying mesophyll. Furthermore, we suggest that the correlation of photosynthetic capacity and stomatal conductance observed under different light environments is caused by signals largely independent of photosynthesis.