Regulation of transpirational water loss in Quercus suber trees in a Mediterranean-type ecosystem.

Sap flux density in branches, leaf transpiration, stomatal conductance and leaf water potentials were measured in 16-year-old Quercus suber L. trees growing in a plantation in southern Portugal to understand how evergreen Mediterranean trees regulate water loss during summer drought. Leaf specific hydraulic conductance and leaf gas exchange were monitored during the progressive summer drought to establish how changes along the hydraulic pathway influence shoot responses. As soil water became limiting, leaf water potential, stomatal conductance and leaf transpiration declined significantly. Predawn leaf water potential reflected soil water potential measured at 1-m depth in the rhizospheres of most trees. The lowest predawn leaf water potential recorded during this period was -1.8 MPa. Mean maximum stomatal conductance declined from 300 to 50 mmol m(-2) s(-1), reducing transpiration from 6 to 2 mmol m(-2) s(-1). Changes in leaf gas exchange were attributed to reduced soil water availability, increased resistances along the hydraulic pathway and, hence, reduced leaf water supply. There was a strong coupling between changes in soil water content and stomatal conductance as well as between stomatal conductance and leaf specific hydraulic conductance. Despite significant seasonal differences among trees in predawn leaf water potential, stomatal conductance, leaf transpiration and leaf specific hydraulic conductance, there were no differences in midday leaf water potentials. The strong regulation of changes in leaf water potential in Q. suber both diurnally and seasonally is achieved through stomatal closure, which is sensitive to changes in both liquid and vapor phase conductance. This sensitivity allows for optimization of carbon and water resource use without compromising the root-shoot hydraulic link.

Age-related effects on leaf area/sapwood area relationships, canopy transpiration and carbon gain of Norway spruce stands (Picea abies) in the Fichtelgebirge, Germany.

Stand age is an important structural determinant of canopy transpiration (E(c)) and carbon gain. Another more functional parameter of forest structure is the leaf area/sapwood area relationship, A(L)/A(S), which changes with site conditions and has been used to estimate leaf area index of forest canopies. The interpretation of age-related changes in A(L)/A(S) and the question of how A(L)/A(S) is related to forest functions are of current interest because they may help to explain forest canopy fluxes and growth. We conducted studies in mature stands of Picea abies (L.) Karst. varying in age from 40 to 140 years, in tree density from 1680 to 320 trees ha(-1), and in tree height from 15 to 30 m. Structural parameters were measured by biomass harvests of individual trees and stand biometry. We estimated E(c) from scaled-up xylem sap flux of trees, and canopy-level fluxes were predicted by a three-dimensional microclimate and gas exchange model (STANDFLUX). In contrast to pine species, A(L)/A(S) of P. abies increased with stand age from 0.26 to 0.48 m(2) cm(-2). Agreement between E(c) derived from scaled-up sap flux and modeled canopy transpiration was obtained with the same parameterization of needle physiology independent of stand age. Reduced light interception per leaf area and, as a consequence, reductions in net canopy photosynthesis (A(c)), canopy conductance (g(c)) and E(c) were predicted by the model in the older stands. Seasonal water-use efficiency (WUE = A(c)/E(c)), derived from scaled-up sap flux and stem growth as well as from model simulation, declined with increasing A(L)/A(S) and stand age. Based on the different behavior of age-related A(L)/A(S) in Norway spruce stands compared with other tree species, we conclude that WUE rather than A(L)/A(S) could represent a common age-related property of all species. We also conclude that, in addition to hydraulic limitations reducing carbon gain in old stands, a functional change in A(L)/A(S) that is related to reduced light interception per leaf area provides another potential explanation for reduced carbon gain in old stands of P. abies, even when hydraulic constraints increase in response to changes in canopy architecture and aging.

Site-specific water relations and stomatal response of Quercus ilex in a Mediterranean watershed.

Intraspecific variations in the water relations and stomatal response of Quercus ilex L. were analyzed under field conditions by comparing trees at two locations within a Mediterranean watershed (l'Avic, Catalonia, NE Spain). Distinct environmental gradients exist between the two sites (referred to as ridge top at 975 m and valley bottom at 700 m) with greater soil depth for water storage, reduced radiation, reduced wind and higher water vapor pressure deficits at the valley bottom than at the ridge top. Osmotic adjustment and changes in tissue elasticiity did not significantly increase drought resistance in the trees studied. The leaf-to-air vapor pressure difference (Deltaw) threshold for inducing stomatal closure was higher at the ridge top (15.6 kPa MPa(-1) +/- 0.5 SE) than at the valley bottom (9.8 kPa MPa(-1) +/- 1.0 SE). However, increases in Deltaw beyond the threshold were followed by greater reductions in leaf conductance of trees at the ridge top than at the valley bottom. At both sites, maximum leaf conductance was related to predawn shoot water potential which, in turn, was related to watershed stream flow. The effects of water deficits during the dry summer of 1989 were more severe in trees at the valley bottom than at the ridge top. During periods of high evaporative demand, site-specific differences in the control of water loss led to more conservative water use by trees at the ridge top and, thus, to even greater drought avoidance (higher predawn water potentials) in late summer.

Above- and below-ground environmental influences on leaf conductance ofCeanothus thyrsiflorus growing in a chaparral environment: drought response and the role of abscisic acid.

Small shrubs ofCeanothus thyrsiflorus were grown in 19-1 pots irrigated under natural conditions in a chaparral region of Southern California and then subjected to soil drying. Characteristics of leaf gas exchange, leaf water potential, and concentrations of the stress hormone abscisic acid in the xylem sap, ABAxyl, were determined at various stages of drought. Diurnal changes in conductance were strongly correlated with leaf net photosynthesis rate, which provides an effective, integrative predictor of above-ground climate effects on conductance. In drought conditions, ABAxyl concentration increased. Increases in the concentration range of 50-500 nmol/l appeared to induce stomatal closure, restricting water loss and carbon dioxide uptake. When the momentary water potential is related to ABAxyl, ABA appeared to increase significantly only after a threshold of approximately -1.5 MPa was exceeded. At less negative water potentials, large variation in ABAxyl in the 50-1000 nmol/l range occurred for all water-potential values, because ABAxyl remains relatively constant over diurnal courses as water potentials decrease and then recover. When the water potential became more negative than -1.5 MPa, ABAxyl concentrations occurred between approximately 500 and 10 000 nmol/l and even greater in isolated cases. An approximately linear relationship is recognizable between ABAxyl and momentary water potential in this range because in plants under drought conditions, ABAxyl increases during the course of the day as water potential decreases. Increases in ABAxyl in the high concentration range were associated with relatively minor additional restrictions in gas exchange, but they might contribute to improved water use efficiency and explain diurnal changes in the potential for stomatal opening that have been observed in Mediterranean sclerophyllous species. When we examined long-term seasonal change in the response of irrigated plants, changes in average daily temperature greater than 10°C occurred (also associated with shifts in relative humidity and radiation input), which apparently led to small changes in predawn water potential in the -0.1 to -0.7 MPa range. Increases in ABAxyl occurred that were in turn negatively correlated with daily maximum leaf conductance. Thus, chaparral shrubs under non-drought conditions seem to sense even small changes in environmental conditions, in our opinion most probably due to initial drying of the uppermost soil and synthesis of ABA in the shallow roots. The results support the hypothesis that information of photosynthesis rate and predawn water potential may be used as primary variables to predict canopy conductance of Mediterranean sclerophyll shrub vegetation.

Photoinhibition as a control on photosynthesis and production of Sphagnum mosses.

The effect of high light intensity on photosynthesis and growth of Sphagnum moss species from Alaskan arctic tundra was studied under field and laboratory conditions. Field experiments consisted of experimental shading of mosses at sites normally exposed to full ambient irradiance, and removal of the vascular plant canopy from above mosses in tundra water track habitats. Moss growth was then monitored in the experimental plots and in adjacent control areas for 50 days from late June to early August 1988. In shaded plots total moss growth was 2-3 times higher than that measured in control plots, while significant reductions in moss growth were found in canopy removal plots. The possibility that photoinhibition of photosynthesis might occur under high-light conditions and affect growth was studied under controlled laboratory conditions with mosses collected from the arctic study site, as well as from a temperate location in the Sierra Nevada, California. After 2 days of high-light treatment (800 µmol photons m-2 s-1) in a controlled environmental chamber, moss photosynthetic capacity was significantly lowered in both arctic and temperate samples, and did not recover during the 14-day experimental period. The observed decrease in photosynthetic capacity was correlated (r 2=0.735, P<0.001) with a decrease in the ratio of variable to maximum chlorophyll fluorescence (F v/F m) in arctic and temperate mosses. This relationship indicates photoinhibition of photosynthesis in both arctic and temperate mosses at even moderately high light intensities. It is suggested that susceptibility to photoinhibition and failure to photoacclimate to higher light intensities in Sphagnum spp. may be related to low tissue nitrogen levels in these exclusively ombrotrophic plants. Photoinhibition of photosynthesis leading to lowered annual carbon gain in Sphagnum mosses may be an important factor affecting CO2 flux at the ecosystem level, given the abundance of these plants in Alaskan tussock tundra.

Environmental effects on CO2 efflux from riparian tundra in the northern foothills of the Brooks Range, Alaska, USA.

Carbon dioxide efflux and soil microenvironmental factors were measured diurnally in Carex aquatilus-and Eriophorum angustifolium-dominated riparian tundra communities to determine the relative importance of soil environmental factors controlling ecosystem carbon dioxide exchange with the atmosphere. Measurements were made weekly between 18 June and 24 July 1990. Diurnal patterns in carbon dioxide efflux were best explained by changes in soil temperature, while seasonal changes in efflux were correlated with changes in depth to water table, depth to frozen soil and soil moisture. Carbon dioxide efflux rates were lowest early in the growing season when high water tables and low soil temperatures limited microbial and root activity. Individual rainfall events that raised the water table were found to strongly reduce carbon dioxide efflux. As the growing season progressed, rainfall was low and depth to water table and soil temperatures increased. In response, carbon dioxide efflux increased strongly, attaining rates late in the season of approximately 10 g CO2 m-2 day-1. These rates are as high as maxima recorded for other arctic sites. A mathematical model is developed which demonstrates that soil temperature and depth to water table may be used as efficient predictors of ecosystem CO2 efflux in this habitat. In parallel with the field measurements of CO2 efflux, microbial respiration was studied in the laboratory as a function of temperature and water content. Estimates of microbial respiration per square meter under field conditions were made by adjusting for potential respiring soil volume as water table changed and using measured soil temperatures. The results indicate that the effect of these factors on microbial respiration may explain a large part of the diurnal and seasonal variation observed in CO2 efflux. As in coastal tundra sites, environmental changes that alter water table depth in riparian tundra communities will have large effects on ecosystem CO2 efflux and carbon balance.

Factors influencing carbon fixation and water use by mediterranean sclerophyll shrubs during summer drought.

Mediterranean sclerophyll shrubs respond to seasonal drought by adjusting the amount of leaf area exposed and by reducing gas exchange via stomatal closure mechanisms. The degree to which each of these modifications can influence plant carbon and water balances under typical mediterranean-type climate conditions is examined. Leaf area changes are assessed in the context of a canopy structure and light microclimate model. Shifts in physiological response are examined with a mechanistically-based model of C3 leaf gas exchange that simulates progressive reduction of maximum photosynthesis and transpiration rates and increasingly strong midday stomatal closure over the course of drought. The results demonstrate that midday stomatal closure may effectively contribute to drought avoidance, increase water use efficiency, and strongly alter physiological efficiency in the conversion of intercepted light energy to photoproducts. Physiological adjustments lead to larger reductions in water use than occur when comparing leaf area index 3.5 to 1.5, extremes found for natural stands of sclerophyll shrubs in the California chaparral. Reductions in leaf area have the strongest effect on resource capture and use during non-water-stressed periods and the least effect under extreme drought conditions, while shifts in physiological response lead to large savings of water and efficient water use under extreme stress. An important model parameter termed GFAC (proportionality factor expressing the relation of conductance [g] to net photosynthesis rate) is utilized, which changes in response to the integrated water stress experimence of shrubs and alters the degree to which stomata may open for a given rate of carbon fixation. We attempt to interpret this parameter in terms of physiological mechanisms known to modify control of leaf gas exchange during drought. The analysis helps visualize means by which canopy gas exchange behavior may be coupled to physiological changes occurring in the root environment during soil drying.

Water content effects on photosynthetic response of Sphagnum mosses from the foothills of the Philip Smith Mountains, Alaska.

In tussock tundra areas of the foothills north of the Brooks Range, Alaska, up to two-thirds of annual precipitation may occur during intermittent summer thunderstorms. The seasonal pattern in capitulum water content of Sphagnum spp. depends on the frequency and duration of these precipitation events, on the microtopography of the habitat including depth of thaw, and on morphological characteristics of the individual species. The response of net photosynthesis to varying water content in Sphagnum squarrosum and S. angustifolium growing under willow canopies in a tussock tundra area near the Dalton Highway on the North Slope of Alaska was examined in the field. After a period in June required to develop photosynthetic capability, capitula water content was essentially optimal for photosynthesis in the range from 6 to 10 g H2O/g DW. Above this range, the rate of CO2 uptake was reduced, presumably due to limitations on CO2 diffusion to the photosynthetically active sites. At water contents below the optimum, net photosynthesis fell rapidly until reaching compensation at approximately 1 g H2O/g DW. Dependent on changes in weather conditions, average water content of Sphagnum samples collected in the field occasionally fell below 5 g H2O/g DW. During a particularly dry period, water content of individual Sphagnum hummocks fell below 1 g H2O/g DW, indicating that water stress does limit Sphagnum photosynthetic production in this habitat.

Irradiance and temperature effects on photosynthesis of tussock tundra Sphagnum mosses from the foothills of the Philip Smith Mountains, Alaska.

Photosynthetic characteristics of three species of Sphagnum common in the foothills of the Brooks Range on the North Slope of Alaska were investigated. Generally, light-saturated rates of net photosynthesis decreased in the order S. squarrosum, S. angustifolium, and S. warnstorfii when plants were grown under common growth chamber conditions. For field-grown S. angustifolium, average light compensation point at 10°C was 37 µmol m-2s-1 photosynthetic photon flux density (PPFD), and light saturation occurred between 250 and 500 µmol m-2 s-1. At 20°C, compensation point increased to 127 µmol m-2s-1 and the PPFD required for light saturation increased to approximately 500 µmol m-2s-1, while maximum rates of CO2 uptake increased only slightly. Light response curves of chamber-grown plants exhibited substantially lower compensation points and higher light-saturated rates of CO2 assimilation than field-grown material, due perhaps to a higher percentage of green, photosynthetically competent tissue. All three species exhibited broad responses to temperature, with optima near 20°C, and maintained at least 75% of maximum assimilation between approx. 13° and 30°C. Rates at 5°C were approx. 50% of maximum. Studies of the microclimate of Sphagnum at the field research site suggest that CO2 uptake should occur at near light-saturated rates during the day in open tussock tundra but that PPFD may often be limiting under Salix and Betula canopies in a water track drainage. Simulations using a simple model provided a seasonal estimate of 0.78 g dry weight (DW) of S. angustifolium produced from each initial g of photosynthetic tissue under willow canopies, assuming no water limitations. Although the simulation model suggests that production would be 66% higher in open tussock tundra, S. angustifolium is rarely found in this potentially more stressful habitat. To explain the relative abundance of Sphagnum in shaded water track areas as compared to open tussock tundra, we postulate that the vascular plant canopies provide protection from adverse effects of high temperatures, excess irradiance and reduced water availability. Under conditions of normal water availability, removal of the vascular plant cover did not affect the tissue water content of S. squarrosum, but resulted in a strong decrease in photosynthetic capacity, accompanied by chlorophyll bleaching. These results suggest that photoinhibition may limit production under certain conditions.

Limitations on Sphagnum growth and net primary production in the foothills of the Philip Smith Mountains, Alaska.

In the foothills of the Philip Smith Mountains, Brooks Range, Alaska, tussock tundra occurs on rolling hills and in valleys that were shaped by Pleistocene glaciations. During the 1986 and 1987 summer seasons, Sphagnum growth and production were determined in "water tracks" on tundra slopes that acted to channel water flow to the valley bottom stream and in "intertrack tundra" areas that were relatively homogeneous with respect to downslope drainage. Measurements were made under ambient environmental conditions and on mosses receiving supplemental irrigation in each area. Growth rate for Sphagnum spp. (cm shoot length increase/day) was low and relatively constant in intertrack tundra and highest but quite variable in water tracks. A strong negative correlation was found between Sphagnum spp. growth rate and solar irradiance in the shady environment below Salix canopies in the water tracks. Estimates of net annual dry weight (DW) production for Sphagnum spp. ranged from 0.10 g DW dm-2 yr-1 in intertrack tundra vegetation to 1.64 g DW dm-2 yr-1 in well-shaded water tracks. Experimental water additions had little effect on growth and production in intertrack tundra and well-developed water tracks, but significantly increased growth in a weakly-developed water track community. Low production over large areas of tundra slopes may occur due to presence of slow growing species resistant to dessication in intertrack tundra as opposed to rapidly growing less compact species within the limited extent of water tracks. We hypothesize that species capable of rapid growth occur also in weakly-developed water tracks, and that these are water-limited more often than plants occurring in well-developed water track situations. Where experienced, high light intensity may additionally limit growth due to photoinhibition.

Effect of photosynthetic photon flux density on carboxylation efficiency.

The effect of photosynthetic photon flux density (PPFD) on photosynthetic response (A) to CO(2) partial pressures between 35 pascals and CO(2) compensation point (Gamma) was investigated, especially below PPFD saturation. Spinacia oleracea cv ;Atlanta,' Glycine max cv ;Clark,' and Arbutus unedo were studied in detail. The initial slope of the photosynthetic response to CO(2) ( partial differentialA/ partial differentialC[Gamma]) was constant above a PPFD of about 500 to 600 micromoles per square meter per second for all three species; but declined rapidly with PPFD below this critical level. For Gamma there was also a critical PPFD (approximately 200 micromoles per square meter per second for S. oleracea and G. max; 100 for A. unedo) above which Gamma was essentially constant, but below which Gamma increased with decreasing PPFD. All three species showed a dependence of partial differentialA/ partial differentialC(Gamma) on PPFD at low PPFD. Simulated photosynthetic responses obtained with a biochemically based model of whole-leaf photosynthesis were similar to measured responses.

Seasonal changes in net photosynthesis rates and photosynthetic capacity in leaves of Cistus salvifolius, a European mediterranean semi-deciduous shrub.

During five different periods between Nov. 1982 and Aug. 1983, the diurnal patterns exhibited in photosynthetic CO2 uptake and stomatal conductance were observed under natural conditions on twigs of Cistus salvifolius, a Mediterranean semi-deciduous shrub which retains a significant proportion of its leaves through the summer drought. During the same periods, net photosynthesis at saturating CO2 partial pressure was measured on the same twigs as a function of irradiance at different temperatures. From these data, photosynthetic capacity, defined here as the CO2- and light-saturated net photosynthesis rate, was obtained as a function of leaf temperature. C. salvifolius is a winter growing species, shoot growth being initiated in Nov. and continuing through May. Photosynthetic capacity was quite high in Nov., March and June, exceeding 40 µmol m-2 s-1 at optimum temperature. In Dec., photosynthetic capacity was somewhat reduced, perhaps due to low night-time temperatures (<5°C) during the measurement period. In Aug., capacity in oversummering shoots at optimum temperature fell to less than 8 µmol m-2 s-1, due to water trees and perhaps leaf aging. Seasonal changes in maximal photosynthetic rates under ambient conditions were similar, and like those found in co-occurring evergreen sclerophylls. Like the evergreens, Cistus demonstrated considerable stomatal control of transpirational water loss, particularly in oversummering leaves. During each measurement period except Aug. when capacity was quite low, the maximum rates of net photosynthesis measured under ambient conditions were less than half the measured photosynthetic capacities at comparable temperatures, suggesting an apparent excess nitrogen investment in the photosynthetic apparatus.

Use of an analytical model to study limitations on net photosynthesis in Arbutus unedo under field conditions.

From field gas-exchange measurements on Arbutus unedo growing in Portugal, parameter values necessary to apply an analytical, physiologicallybased model of C 3 photosynthesis were obtained. The model successfully simulated measured diurnal photosynthetic responses in Arbutus during periods without water stress, under both natural and CO2-saturating conditions. The model was used to analyze those factors limiting primary productivity during each of the experimental days. Due to a large investment in ribulose bisphosphate (RuBP) regeneration capacity, irradiance was rarely limiting, even during cloudy periods, but the limitation imposed by stomatal conductance was quite large, averaging over 30%. The fact that experimental leaves were maintained in a horizontal position is at least partially responsible for these results. Possible other reasons for this apparent excess of RuBP regeneration capacity visa-vis RuBP carboxylase-oxygenase concentration are discussed.

Effects of temperature at constant air dew point on leaf carboxylation efficiency and CO2 compensation point of different leaf types.

The effect of temperature on photosynthesis at constant water-vapor pressure in the air was investigated using two sclerophyll species, Arbutus unedo and Quercus suber, and one mesophytic species, Spinacia oleracea. Photosynthesis and transpiration were measured over a range of temperatures, 20-39° C. The external concentration of CO2 was varied from 340 µbar to near CO2 compensation. The initial slope (carboxylation efficiency, CE) of the photosynthetic response to intercellular CO2 concentration, the CO2 compensation point (Γ), and the extrapolated rate of CO2 released into CO2-free air (R i) were calculated. At an external CO2 concentration of 320-340 µbar CO2, photosynthesis decreased with temperature in all species. The effect of temperature on Γ was similar in all species. While CE in S. oleracea changed little with temperature, CE decreased by 50% in Q. suber as temperature increased from 25 to 34° C. Arbutus unedo also exhibited a decrease in CE at higher temperatures but not as marked as Q. suber. The absolut value of R i increased with temperature in S. oleracea, while changing little or decreasing in the sclerophylls. Variations in Γ and R i of the sclerophyll species are not consistent with greater increase of respiration with temperature in the light in these species compared with S. oleracea.

Limitations due to water stress on leaf net photosynthesis of Quercus coccifera in the Portuguese evergreen scrub.

Gas exchange characteristics in leaves of the sclerophyll shrub Quercus coccifera were studied in the natural habitat in Portugal during spring and during the summer dry period. Compared to other sclerophyll species growing at the same site, photosynthesis in leaves of Quercus coccifera was less affected by water stress. Moderate water stress after six weeks of drought led to large decreases in stomatal conductance but no change in mesophyll photosynthetic capacity as compared to late spring. Leaf internal CO2 pressure remained near 220 µbar during diurnal courses in the spring. On midsummer days, leaf internal CO2 decreased from a late morning value of 200 µbar to a late afternoon value of approximately 150 µbar. In contrast to Quercus suber (Tenhunen et al. 1984), restriction of CO2 supply due to stomatal closure reduced net CO2 uptake at midday and in the afternoon during midsummer. A decrease in leaf carboxylation efficiency and an increase in CO2 compensation point at midday also played an important role in determining the diurnal course of net photosynthesis. During the late stages of drought in September, severe water stress led to reduction in mesophyll photosynthetic capacity and further reduction in leaf conductance. The observed decrease in mesophyll photosynthetic capacity was correlated with decrease in the daily minimum leaf water potential to greater negative values than-30 bar. At this time, CO2 saturated photosynthetic rates decreased as much as 50% over the course of a day when measured at constant saturating light, 32° C leaf temperature, and a water vapor mole fraction difference between leaf and air of 30 mbar bar-1.

Analysis of gas exchange in seedlings of Acer saccharum: integration of field and laboratory studies.

In the field, photosynthesis of Acer saccharum seedlings was rarely light saturated, even though light saturation occurs at about 100 µmol quanta m-2 s-1 photosynthetic photon flux density (PPFD). PPFD during more than 75% of the daylight period was 50 µmol m-2 s-1 or less. At these low PPFD's there is a marked interaction of PPFD with the initial slope (CE) of the CO2 response. At PPFD-saturation CE was 0.018 µmol m-2 s-1/(µl/l). The apparent quantum efficiency (incident PPFD) at saturating CO2 was 0.05-0.08 mol/mol. [Formula: see text] and PPFD-saturated CO2 exchange was 6-8 µmol m-2 s-1. The ratio of internal CO2 concentration to external (C i /C a ) was 0.7 to 0.8 except during sunflecks when it decreased to 0.5. The decrease in C i /C a during sunflecks was the result of the slow response of stomates to increased PPFD compared to the response of net photosynthesis. An empirical model, which included the above parameters was used to simulate the measured CO2 exchange rate for portions of two days. Parameter values for the model were determined in experiments separate from the daily time courses being sumulated. Analysis of the field data, partly through the use of simulations, indicate that the elimination of sunflecks would reduce net carbon gain by 5-10%.

Changes in photosynthetic capacity, carboxylation efficiency, and CO2 compensation point associated with midday stomatal closure and midday depression of net CO2 exchange of leaves of Quercus suber.

The carbon-dioxide response of photosynthesis of leaves of Quercus suber, a sclerophyllous species of the European Mediterranean region, was studied as a function of time of day at the end of the summer dry season in the natural habitat. To examine the response experimentally, a "standard" time course for temperature and humidity, which resembled natural conditions, was imposed on the leaves, and the CO2 pressure external to the leaves on subsequent days was varied. The particular temperature and humidity conditions chosen were those which elicited a strong stomatal closure at midday and the simultaneous depression of net CO2 uptake. Midday depression of CO2 uptake is the result of i) a decrease in CO2-saturated photosynthetic capacity after light saturation is reached in the early morning, ii) a decrease in the initial slope of the CO2 response curve (carboxylation efficiency), and iii) a substantial increase in the CO2 compensation point caused by an increase in leaf temperature and a decrease in humidity. As a consequence of the changes in photosynthesis, the internal leaf CO2 pressure remained essentially constant despite stomatal closure. The effects on capacity, slope, and compensation point were reversed by lowering the temperature and increasing the humidity in the afternoon. Constant internal CO2 may aid in minimizing photoinhibition during stomatal closure at midday. The results are discussed in terms of possible temperature, humidity, and hormonal effects on photosynthesis.

Diurnal variations in abscisic acid content and stomatal response to applied abscisic acid in leaves of irrigated and non-irrigated Arbutus unedo plants under naturally fluctuating environmental conditions.

Endogenous abscisic acid content (ABA) of Arbutus unedo leaves growing under natural conditions in a macchia near Sobreda, Portugal, was very high (0.25 to 2.3 µg g1 fresh weight). Highest concentrations were found during the very early morning hours and at midday. During the late morning hours and in the late afternoon ABA concentrations decreased to between one-third and one-fourth of peak values. The samples for ABA content were obtained from both irrigated (Ψ between-10 and-25 bar) and non-irrigated plants experiencing natural water stress during the dry season (Ψ of-50 bar). During the course of the measurement day, stomatal conductance was relatively constant and conductance of watered plants was 50 to 100% greater than that of unwatered plants. No clear correlations between ABA content and stomatal conductance and/or xylem water potential were observed. Despite large differences in water potential and differences in degree of stomatal opening, absolute concentrations of ABA were not found to differ.Small quantities (8-14 pmoles cm2 leaf area) of ABA were applied to leaves of irrigated and non-irrigated Arbutus unedo plants by injection into the petiole. These extremely small ABA doses resulted in transient reductions in stomatal conductance. The effectiveness with which injected ABA closed stomata was highest during the morning and decreased substantially at midday. Increased sensitivity to injected ABA may again occur in the late afternoon but recent measurements suggest that this may depend on long-term drought experience of the plants. The characteristics of the response to injected ABA were similar in irrigated and non-irrigated plants although irrigated plants responded in general more strongly.

Light-Stimulated Burst of Carbon Dioxide Uptake following Nocturnal Acidification in the Crassulacean Acid Metabolism Plant Kalanchoë diagremontiana.

CO(2) exchange characteristics were studied during the light-stimulated burst of CO(2) uptake (MB) immediately following a period of nocturnal CO(2) fixation in the Crassulacean acid metabolism plant Kalanchoë daigremontiana. During the early parts of the MB, stimulation of net CO(2) uptake by low ambient O(2) concentration (1.5%) was small, and leaves showed the capacity for net CO(2) uptake at low ambient CO(2) partial pressure (30 microbars) and when the MB was interrupted by darkness. During the later phase of the MB, stimulation of net CO(2) uptake by 1.5% O(2) was increased, and net CO(2) loss was recorded both at 30 microbars CO(2) and during dark interruptions. These results suggest that CO(2) fixation during the MB occurs simultaneously via phosphoenolpyruvate carboxylase (predominant during the early phase of the MB) and via ribulose bisphosphate carboxylase (predominant during the later phase of the burst). The magnitude and duration of the MB was increased by a reduction in the length of the dark period and by low (15 degrees C) compared to high (30 degrees C) leaf temperatures.