The 'hydrology' of leaves: co-ordination of structure and function in temperate woody species

The hydraulic conductance of the leaf lamina (K_lamina) substantially constrains whole‐plant water transport, but little is known of its association with leaf structure and function. K_lamina was measured for sun and shade leaves of six woody temperate species growing in moist soil, and tested for correlation with the prevailing leaf irradiance, and with 22 other leaf traits. K_lamina varied from 7.40 × 10^?5 kg m^?2 s^?1 MPa^?1 for Acer saccharum shade leaves to 2.89 × 10^?4 kg m^?2 s^?1 MPa^?1 for Vitis labrusca sun leaves. Tree sun leaves had 15–67% higher K_lamina than shade leaves. K_lamina was co‐ordinated with traits associated with high water flux, including leaf irradiance, petiole hydraulic conductance, guard cell length, and stomatal pore area per lamina area. K_lamina was also co‐ordinated with lamina thickness, water storage capacitance, 1/mesophyll water transfer resistance, and, in five of the six species, with lamina perimeter/area. However, for the six species, K_lamina was independent of inter‐related leaf traits including leaf dry mass per area, density, modulus of elasticity, osmotic potential, and cuticular conductance. K_lamina was thus co‐ordinated with structural and functional traits relating to liquid‐phase water transport and to maximum rates of gas exchange, but independent of other traits relating to drought tolerance and to aspects of carbon economy.     

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.     

Are Sclerophylls and Malacophylls Hydraulically Different?

This work tests the hypothesis that sclerophylls (i.e. hard-leaved species) would be less efficient than malacophylls (i.e. soft-leaved species) in terms of water transport through the stem as well as within the leaf blade. Mean leaf surface area (A_L), leaf specific mass (LSM) as well as shoot (K_WL), stem (K_SL) and leaf (K_LL) hydraulic conductances were measured in eight Mediterranean evergreen sclerophylls and eight temperate deciduous malacophylls. No difference was observed between the two groups in terms of K_LL and of the contribution of leaves to the overall shoot hydraulic resistance. Leaves represented in all cases 48 to 90 % of the shoot hydraulic resistance, suggesting that the sclerophyllous habitus does not per se lead to low efficiency in water transport within the leaf blade. A weak negative relationship (r² = 0.252) appeared to exist between K_SL and LSM. This might provide an explanation for the lower growth rates of sclerophylls with respect to malacophylls.     

Functional relationships between hydraulic traits and the timing of diurnal depression of photosynthesis

The hydraulic coordination along the water transport pathway helps trees provide adequate water supply to the canopy, ensuring that water deficits are minimized and that stomata remain open for CO₂ uptake. We evaluated the stem and leaf hydraulic coordination and the linkages between hydraulic traits and the timing of diurnal depression of photosynthesis across seven evergreen tree species in the southern Andes. There was a positive correlation between stem hydraulic conductivity (k_s) and leaf hydraulic conductance (K_Leaf) across species. All species had similar maximum photosynthetic rates (A_max). The species with higher k_s and K_Leaf attained A_max in the morning, whereas the species with lower k_s and K_Leaf exhibited their A_max in the early afternoon concurrently with turgor loss. These latter species had very negative leaf water potentials, but far from the pressure at which the 88% of leaf hydraulic conductance is lost. Our results suggest that diurnal gas exchange dynamics may be determined by leaf hydraulic vulnerability such that a species more vulnerable to drought restrict water loss and carbon assimilation earlier than species less vulnerable. However, under stronger drought, species with earlier CO₂ uptake depression may increase the risk of hydraulic failure, as their safety margins are relatively narrow.     

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.     

Axial and radial profiles in conductivities, water storage and native embolism in trunks of young and old-growth ponderosa pine trees

Ponderosa pine has very wide sapwood, and yet the spatial and temporal use of that sapwood for water transport is poorly understood. Moreover, there have been few comparisons of function in tips of old-growth trees in comparison with young trees. In the present study, axial and radial specific conductivity (k_s), leaf specific conductivity (LSC), leaf specific conductance (k_l), native embolism and the compartmentalization of sapwood water storage were characterized in trunks of young and old-growth trees. Trunks of young trees had lower k_s, lower LSC and lower native embolism [corresponding to 5% loss of conductivity (PLC)] than trunks of old-growth trees. However, k_l in young trees was 3.5 times higher than in old-growth trees, supporting the hypothesis that tall trees have a reduced ability to transport water to their leaves. Water storage (capacitance) of young trees was not significantly different than at the base of old-growth trees. Although the top of the old-growth trees had similar k_s, LSC and k_l to the young trees for a given cambial age, they had higher native embolism and lower capacitance. There was no trade-off between k_s and native embolism at any height. In the tree crown, outer sapwood had 35–50% higher k_s than the inner sapwood and 17–25 PLC lower native embolism. At the base of the old trees, there was no significant difference in native embolism between the outer, middle and inner sapwood, showing that refilling of embolisms was complete despite the 130-year difference in wood age among these radial positions. Although during the dry season the inner sapwood tended to be more saturated than the outer sapwood, the outer part of the sapwood contributed up to 60% of the overall stored water. Safer xylem, higher capacitance and higher k_l would appear adaptive in the young trees for regulating their water resource, which is likely to be less reliable than the water availability of older trees with their more developed root system.     

Scaling CO2-photosynthesis relationships from the leaf to the canopy

Responses of individual leaves to short-term changes in CO₂ partial pressure have been relatively well studied. Whole-plant and plant community responses to elevated CO₂ are less well understood and scaling up from leaves to canopies will be complicated if feedbacks at the small scale differ from feedbacks at the large scale. Mathematical models of leaf, canopy, and ecosystem processes are important tools in the study of effects on plants and ecosystems of global environmental change, and in particular increasing atmospheric CO₂, and might be used to scale from leaves to canopies. Models are also important in assessing effects of the biosphere on the atmosphere. Presently, multilayer and big leaf models of canopy photosynthesis and energy exchange exist. Big leaf models — which are advocated here as being applicable to the evaluation of impacts of global change on the biosphere — simplify much of the underlying leaf-level physics, physiology, and biochemistry, yet can retain the important features of plant-environment interactions with respect to leaf CO₂ exchange processes and are able to make useful, quantitative predictions of canopy and community responses to environmental change. The basis of some big leaf models of photosynthesis, including a new model described herein, is that photosynthetic capacity and activity are scaled vertically within a canopy (by plants themselves) to match approximately the vertical profile of PPFD. The new big leaf model combines physically based models of leaf and canopy level transport processes with a biochemically based model of CO₂ assimilation. Predictions made by the model are consistent with canopy CO₂ exchange measurements, although a need exists for further testing of this and other canopy physiology models with independent measurements of canopy mass and energy exchange at the time scale of 1 h or less.Abbreviations LAI leaf area index - NIR near infrared (700–3000 nm) radiation - PAR photosynthetically active (400–700 nm) radiation - PI photosynthetic irradiance (400–700 nm) - PPFD photosynthetic photon flux area density (400–700 nm) - PS I Photosystem I - PS II Photosystem II - Rubisco ribulose-1,5-bisphosphate carboxylase/oxygenase - RuP₂ ribulose-1,5-bisphosphate     

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.     

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

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

Modelling the soil-plant-atmosphere continuum in a Quercus–Acer stand at Harvard Forest: the regulation of stomatal conductance by light, nitrogen and soil/plant hydraulic properties

Our objective is to describe a multi-layer model of C₃-canopy processes that effectively simulates hourly CO₂ and latent energy (LE) fluxes in a mixed deciduous Quercus-Acer (oak–maple) stand in central Massachusetts, USA. The key hypothesis governing the biological component of the model is that stomatal conductance (g_s) is varied so that daily carbon uptake per unit of foliar nitrogen is maximized within the limitations of canopy water availability. The hydraulic system is modelled as an analogue to simple electrical circuits in parallel, including a separate soil hydraulic resistance, plant resistance and plant capacitance for each canopy layer. Stomatal opening is initially controlled to conserve plant water stores and delay the onset of water stress. Stomatal closure at a threshold minimum leaf water potential prevents xylem cavitation and controls the maximum rate of water flux through the hydraulic system. We show a strong correlation between predicted hourly CO₂ exchange rate (r²= 0.86) and LE (r²= 0.87) with independent whole-forest measurements made by the eddy correlation method during the summer of 1992. Our theoretical derivation shows that observed relationships between CO₂ assimilation and LE flux can be explained on the basis of stomatal behaviour optimizing carbon gain, and provides an explicit link between canopy structure, soil properties, atmospheric conditions and stomatal conductance.     

Potassium mediates coordination of leaf photosynthesis and hydraulic conductance by modifications of leaf anatomy

Typical symptoms of potassium deficiency, characterized as chlorosis or withered necrosis, occur concomitantly with downregulated photosynthesis and impaired leaf water transport. However, the prominent limitations and mechanisms underlying the concerted decreases of leaf photosynthesis and hydraulic conductance are poorly understood. Monocots and dicots were investigated based on responses of photosynthesis and hydraulic conductance and their components and the correlated anatomical determinants to potassium deficiency. We found a conserved pattern in which leaf photosynthesis and hydraulic conductance concurrently decreased under potassium starvation. However, monocots and dicots showed two different hydraulic‐redesign strategies: Dicots tended to show a decreased minor vein density, whereas monocots reduced the size of the bundle sheath and its extensions, rather than the minor vein density; both of these strategies may restrain xylem and outside‐xylem hydraulic conductance. Additionally, potassium‐deprived leaves developed with fewer mesophyll cell‐to‐cell connections, leading to a reduced area being available for liquid‐phase flow. Further quantitative analysis revealed that mesophyll conductance to CO₂ and outside‐xylem hydraulic resistance were the major contributors to photosynthetic limitation and increased hydraulic resistance, at more than 50% and 60%, respectively. These results emphasize the importance of potassium in the coordinated regulation of leaf photosynthesis and hydraulic conductance through modifications of leaf anatomy.     

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.     

Alterations in photosynthesis and pigment distributions in pea leaves following UV-B exposure

We compared photosynthetic and UV-B-absorbing pigment concentrations, gas-exchange rates and photosystem II (PSII) electron transport rates in leaves of pea (Pisum sativum mutant Argenteum) grown without UV-B or under an enhanced UV-B treatment (18 kJ m^?2 biologically effective daily dose) in a greenhouse. We also compared the distribution of chlorophyll by depth within leaves of each treatment by using image analysis of chlorophyll autofluorescence. Ultraviolet-B treatment elicited putative protective responses such as an 80% increase in UV-B-absorbing compound concentrations (leaf-area basis), and a slight increase in mesophyll thickness (178 in controls compared to 191 μm in UV-B-treated leaves). However, photosynthetic rates of UV-B-treated leaves were only 80% of those of controls. This was paralleled by reductions in leaf conductance to water vapor (50% of controls) and intercellular CO₂ concentrations, suggesting that stomatal limitations were at least partly responsible for lower photosynthetic rates under the UV-B treatment. Total chlorophyll concentrations (leaf-area basis) in UV-B-treated leaves were only 70% of controls, and there was a shift in the relative distribution of chlorophyll with depth in UV-B-treated leaves. In control leaves chlorophyll concentrations were highest near the adaxial surface of the upper palisade, dropped with depth and then increased slightly in the bottom of the spongy mesophyll nearest the abaxial surface. In contrast, in UV-B-treated leaves chlorophyll concentrations were lowest at the adaxial surface of the upper palisade and increased with depth through the leaf. The most notable treatment difference in chlorophyll concentrations was in the upper palisade near the adaxial surface of leaves, where we estimate that chlorophyll concentrations in each 1-μm-thick paradermal layer were about 50% lower in UV-B-treated leaves than in controls. We found reduced electron transport capacity in UV-B-treated leaves, based on lower maximum fluorescence (F_m), variable to maximum fluorescence ratios (F,/F_m) and quantum yield of PSII electron transport (Y). However, the above were assessed from fluorometer measurements on the adaxial leaf surface and may reflect the markedly lower chlorophyll concentrations in the upper palisade of UV-B-treated leaves.     

The Arabidopsis thaliana aquaporin AtPIP1;2 is a physiologically relevant CO₂ transport facilitator

Cellular exchange of carbon dioxide (CO₂) is of extraordinary importance for life. Despite this significance, its molecular mechanisms are still unclear and a matter of controversy. In contrast to other living organisms, plants are physiologically limited by the availability of CO₂. In most plants, net photosynthesis is directly dependent on CO₂ diffusion from the atmosphere to the chloroplast. Thus, it is important to analyze CO₂ transport with regards to its effect on photosynthesis. A mutation of the Arabidopsis thaliana AtPIP1;2 gene, which was characterized as a non‐water transporting but CO₂ transport‐facilitating aquaporin in heterologous expression systems, correlated with a reduction in photosynthesis under a wide range of atmospheric CO₂ concentrations. Here, we could demonstrate that the effect was caused by reduced CO₂ conductivity in leaf tissue. It is concluded that the AtPIP1;2 gene product limits CO₂ diffusion and photosynthesis in leaves.     

Changes in Stem and Leaf Hydraulics Preceding Leaf Shedding in Castanea Sativa L.

This paper describes changes in leaf water status and in stem, petiole and leaf blade hydraulics preceding leaf senescence and shedding in Castanea sativa L. (chestnut). Measurements of maximum diurnal leaf conductance to water vapour (g_L), minimum water potential (_L), hydraulic conductance per unit leaf surface area of stems (K_SL), petioles (K_PL) and leaf blades (K_LL) and number of functional conduits and inside diameter distribution were performed in June, September and October 1999. In September, still green leaves had undergone some dehydration as indicated by decreased g_L (by 75 %) and _L with respect to June. In the same time, K_SL decreased by 88 %, while K_PL and K_LL decreased by 50 % and 20 % of the conduits of stems and 10 % of the petioles (all belonging to the widest diameter range) were no longer functioning, causing a decrease in the theoretical flow by 82 % in stems and 27 % in petioles. Stem xylem blockage was apparently due to tyloses growing into conduits. We advance the hypothesis that the entire process of leaf shedding and winter rest may be initiated by extensive stem embolism occurring during the summer.     

Diurnal depression of leaf hydraulic conductance in a tropical tree species

Diurnal patterns of hydraulic conductance of the leaf lamina (K_leaf) were monitored in a field‐grown tropical tree species in an attempt to ascertain whether the dynamics of stomatal conductance (g_s) and CO₂ uptake (A_leaf) were associated with short‐term changes in K_leaf. On days of high evaporative demand mid‐day depression of K_leaf to between 40 and 50% of pre‐dawn values was followed by a rapid recovery after 1500 h. Leaf water potential during the recovery stage was less than ?1 MPa implying a refilling mechanism, or that loss of K_leaf was not linked to cavitation. Laboratory measurement of the response of K_leaf to Ψ_leaf confirmed that leaves in the field were operating at water potentials within the depressed region of the leaf ‘vulnerability curve’. Diurnal courses of K_leaf and Ψ_leaf predicted from measured transpiration, xylem water potential and the K_leaf vulnerability function, yielded good agreement with observed trends in both leaf parameters. Close correlation between depression of K_leaf, g_s and A_leaf suggests that xylem dysfunction in the leaf may lead to mid‐day depression of gas exchange in this species.     

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.     

Catalogo delle Alghe Raccolte Lungo IL Litorale di Trani

Abstract

Canopy light interception (CPFD_Int), spectral irradiance, leaf water potential, gas- exchange and optical properties were measured in an irrigated vineyard (Vitis vinifera L. cv Montepulciano) trained to the so-called tendone system in which leaf area index (LAI) was varied by means of 50% (T50) or 75% (T75) cluster removal. The 20.5 t ha^?1 yield in the unthinned treatment (UT) decreased by only 36% in T50 and by 52% in T75. LAI and CPFD_Int similarly increased until summer pruning when LAI was 1.75 m² m^?2 in UT, and 25.6% or 62.2% higher in T50 and T75, respectively. The two thinned treatments had only 12.4% higher CPFD_Int than in UT (1167.1 μmol m^?2 s^?1) due to the increased leaf self-shading. The red-to-far red ratio (R: FR) was as low as 0.10 below the canopy. Light-saturated CO₂ assimilation (A _max) in June averaged 14.4 μmol m^?2 s^?1 in sun-exposed leaves, and 7.6 μmol m^?2 s^?1 in shade leaves. By contrast, the apparent quantum yield of CO₂ assimilation (φ_e) was not significantly affected by leaf position, averaging 0.029 and 0.070 mol mol^?1 in June and October, respectively. Middle and low canopy leaves had only 27 or 6%, respectively, of the top canopy leaves actual CO₂ assimilation rate.     

Photosynthesis of individual field-grown cotton leaves during ontogeny

Photosynthetic characteristics of field-grown cotton (Gossypium hirsutum L.) leaves were determined at several insertion levels within the canopy during the growing season. Single-leaf measurements of net photosynthesis (Pn), stomatal conductance to CO₂ (g_s·CO₂), substomatal CO₂, leaf area expansion, leaf nitrogen, and light intensity (PPFD) were recorded for undisturbed leaves within the crop canopy at 3–4 day intervals during the development of all leaves at main-stem nodes 8, 10, and 12. Patterns of Pn during leaf ontogeny exhibited three distinct phases; a rapid increase to maximum at 16–20 days after leaf unfolding, a relatively short plateau, and a period of linear decline to negligible Pn at 60–65 days. Analysis of the parameters which contributed to the rise and fall pattern of Pn with leaf age indicated the primary involvement of leaf area expansion, leaf nitrogen, PPFD, and g_s·CO₂ in this process. The response of Pn and g_s·CO₂ to incident PPFD conditions during canopy development was highly age dependent. For leaves less than 16 days old, the patterns of Pn and g_s·CO₂ were largely controlled by non-PPFD factors, while for older leaves Pn and g_s·CO₂ were more closely coupled to PPFD-mediated processes. Maximum values of Pn were not significantly different for any of the leaves monitored in this study, however, those leaves at main-stem node 8 did possess a significantly diminished photosynthetic capacity with age compared to upper canopy leaves. This accelerated decline in Pn could not be explained by age-related variations in g_s·CO₂ since all leaves showed similar changes in g_s·CO₂ with leaf age.Abbreviations g_s·CO₂ stomatal conductance to CO₂ - Pn net photosynthesis - PPFD photosynthetic photon flux density     

Photosynthesis, leaf conductance and water relations of in vitro cultured grapevine rootstock in relation to acclimatisation

Leaf net CO₂ uptake and leaf photosynthetic capacity were investigated in micropropagated 41B grapevine rootstock (Vitis vinifera‘Chasselas’×Vitis berlandieri, Mill. De Gr.) plants grown in the presence of four sucrose concentrations (6.25, 12.5, 25.0 or 37.5 g l^?1). Sucrose concentration in the medium during growth in vitro did not affect the leaf photosynthetic performance of plants neither before nor after transplantation. The maximum photosynthetic rate, measured as CO₂-dependent O₂ evolution, was 7.3 µmol m^?2 s^?1 before transplanting and 15.4 µmol m^?2 s^?1 one month after transplantation. The maximum quantum yield of O₂ evolution (on the basis of incident light) was about 0.07 for all sucrose treatments both before and after transplantation. Dry biomass before transplanting was highest in plants grown with 25.0 or 37.5 g l^?1 sucrose in the medium. One month after transplantation the highest dry biomass was also observed for the same treatments. Survival of plants was 100% for all treatments. Leaf conductance to water vapour was always higher in plants before than after transplantation. Both before and after transplanting it increased with increasing light intensity and decreased slightly with increasing CO₂ molar ratio in in vitro plants. Stomata of plants before transplantation were unresponsive to vapour pressure deficit. In vitro plants experience an acute water stress when they are maintained with the whole root system in water and exposed to ambient controlled conditions in a growth chamber. However, there was no wilting of the leaves when similar plants with roots cut off were left in the same conditions. Hydraulic conductivity was low at both root and shoot-root connection levels. It is likely that water supply could be limiting during transplantation because of the low root and root-stem connection conductivity. Water uptake by roots rather than water loss from the shoots would be of primary importance for the maintenance of water balance during acclimatisation.