Comparison of physiological responses of guard cell protoplasts of Nicotiana glauca isolated from leaves collected before dawn or at midday

We observed that guard cell protoplasts isolated from leaves collected at midday from Nicotiana glauca Graham (tree tobacco) did not give the same physiological responses to light as those isolated from leaves collected in early morning. Based on that observation, we attempted to determine whether there were significant differences between the physiological responses of guard cell protoplasts isolated from leaves collected before dawn (with closed stomata) and those isolated from leaves collected at midday (with open stomata). We isolated guard cell protoplasts from leaves collected before dawn and at midday and compared (1) rates of red and blue light-induced pH changes in weakly buffered media caused by changes in their metabolism, (2) their rates of oxygen consumption in darkness and oxygen evolution in light and (3) relative rates of decay of variable chlorophyll a fluorescence in their chloroplasts. Studies with the vital stain fluorescein diacetate failed to reveal any significant differences in the viabilities of protoplast preparations from leaves collected before dawn and at midday. Furthermore, protoplasts from leaves collected at these times swelled to similar extents in an osmotic medium containing 10 µM fusicoccin and 5 mM KCI. Nevertheless, rates of light-induced pH changes, rates of oxygen consumption and evolution and rates of decay of variable chlorophyll a fluorescence were all lower in preparations of guard cell protoplasts from leaves collected at midday than in preparations from leaves collected before dawn. Initial volumes of guard cell protoplasts isolated from leaves collected at midday were 150% of those of guard cell protoplasts isolated from leaves collected before dawn. We suggest that the differences in responses of guard cell protoplasts isolated from leaves collected before dawn and at midday may be caused by (1) nonoptimal isolation conditions for guard cell protoplasts prepared from leaves collected at midday, (2) the lower surface-to-volume ratio of guard cell protoplasts isolated from leaves collected at midday or (3) diurnal and/or circadian regulation of guard cell metabolism over the course of a day.     

Division of guard cell protoplasts of Nicotiana glauca (Graham) in liquid cultures

Abstract. Guard cells are uniquely differentiated to transduce signals into the metabolic and ion transport processes that result in turgor-driven stomatal movements. We tested the hypothesis that these highly specialized cells are terminally differentiated. Guard cell protoplasts were isolated from abaxial epidermal tissue of leaves of Nicotiana glauca (Graham) and cultured in a medium designed for culturing mesophyll protoplasts of Nicotiana tabacum. Protoplasts were incubated at densities of 2–5 × 10¹¹ cells m⁻³ in eight-well microchamber slides under 50μmol m⁻² s⁻¹ of photons of continuous fluorescent light at 25°C. When the medium was modified by the addition of 100mol m⁻³ of sucrose and by buffering with 10mol m⁻³ of MES buffer at pH 6.1, cell division began within 96h of the time the culture was initiated. After 9d of culture, 80% of surviving cells had synthesized new cell walls, had dedifferentiated, and were dividing to form small colonies. Callus tissue was visible after 4–5 weeks. We conclude that guard cells of Nicotiana glauca are not terminally differentiated, and that guard cell protoplasts of this species have the capacity to grow, synthesize cell walls and divide.     

Responses of stomatal conductance to light, humidity and temperature in winter wheat and barley grown at three concentrations of carbon dioxide in the field

Responses of leaf stomatal conductance to light, humidity and temperature were characterized for winter wheat and barely grown at ambient (about 350 μmol mol^?1 in the daytime), ambient + 175 and ambient + 350 μmol mol^?1 concentrations of carbon dioxide in open‐topped chambers in field plots over a three year period. Stomatal responses to environment were determined by direct manipulation of single environmental factors, and those results were compared with responses derived from natural day to day variation in mid‐day stomatal conductance. The purpose of these experiments was to determine the magnitude of reduction in stomatal conductance at elevated [CO₂], and to assess whether the relative response of conductance to elevated [CO₂] was constant across light, humidity and temperature conditions. The results indicated that light, humidity and temperature all significantly affected the relative decrease in stomatal conductance at elevated [CO₂]. The relative decrease in conductance with elevated [CO₂] was greater at low light, low water vapour pressure difference, and high temperature in both species. For measurements made at saturating light near mid‐day, the ratio of mid‐day stomatal conductances at doubled [CO₂] to that at ambient [CO₂] ranged from 0.42 to 0.86, with a mean of 0.66 in barley, and from 0.33 to 0.80, with a mean of 0.56 in wheat. Day‐to‐day variation in the relative effect of elevated [CO₂] on conductance was correlated with the relative stimulation of [CO₂] assimilation rate and with temperature. Some limitations of multiple linear regression, multiplicative, and ‘Ball–Berry' models as summaries of the data are discussed. In barley, a better fit to the models occurred in individual years than for the combined data, and in wheat a better fit to the models occurred when data from near the end of the season were removed.     

A meta-analysis of leaf gas exchange and nitrogen in trees grown under elevated carbon dioxide

The response of trees to rising atmospheric CO₂ concentration ([CO₂]) is of concern to forest ecologists and global carbon modellers and is the focus of an increasing body of research work. I review studies published up to May 1994, and several unpublished works, which reported at least one of the following: net CO₂ assimilation (A), stomatal conductance (g_s), leaf dark respiration (R_d) leaf nitrogen or specific leaf area (SLA) in woody plants grown at <400 μmol mol^?1 CO₂ or at 600–800 μmol mol^?1 CO₂. The resulting data from 41 species were categorized according to growth conditions (unstressed versus stressed), length of CO₂ exposure, pot size and exposure facility [growth chamber (GC), greenhouse (GH), or open-top chamber (OTC)] and interpreted using meta-analytic methods. Overall, A showed a large and significant increase at elevated [CO₂] but length of CO₂ exposure and the exposure facility were important modifiers of this response. Plants exposed for < 50 d had a significantly greater response, and those from GCs had a significantly lower response than plants from longer exposures or from OTC studies. Negative acclimation of A was significant and general among stressed plants, but in unstressed plants was influenced by length of CO₂ exposure, the exposure facility and/or pot size. Growth at elevated [CO₂] resulted in moderate reductions in gs in unstressed plants, but there was no significant effect of CO₂ on gs in stressed plants. Leaf dark respiration (mass or area basis) was reduced strongly by growth at high [CO₂] > while leaf N was reduced only when expressed on a mass basis. This review is the first meta-analysis of elevated CO₂ studies and provides statistical confirmation of several general responses of trees to elevated [CO₂]. It also highlights important areas of continued uncertainty in our understanding of these responses.     

Soybean leaf growth and gas exchange response to drought under carbon dioxide enrichment

This study was conducted to determine the response in leaf growth and gas exchange of soybean (Glycine max Merr.) to the combined effects of water deficits and carbon dioxide (CO₂) enrichment. Plants grown in pots were allowed to develop initially in a glasshouse under ambient CO₂ and well-watered conditions. Four-week old plants were transferred into two different glasshouses with either ambient (360 μmol mol^-1) or elevated (700 μmol mol^-1) CO₂. Following a 2-day acclimation period, the soil of the drought-stressed pots was allowed to dry slowly over a 2-week period. The stressed pots were watered daily so that the soil dried at an equivalent rate under the two CO₂ levels. Elevated [CO₂] decreased water loss rate and increased leaf area development and photosynthetic rate under both well-watered and drought-stressed conditions. There was, however, no significant effect of [CO₂] in the response relative to soil water content of normalized leaf gas exchange and leaf area. The drought response based on soil water content for transpiration, leaf area, and photosynthesis provide an effective method for describing the responses of soybean physiological processes to the available soil water, independent of [CO₂].     

Control of photosynthetic carbon dioxide fixation by the boundary layer, stomata and ribulose 1,5-biphosphate carboxylase/oxygenase

Abstract Coefficients describing the sensitivity of the rate of photosynthetic carbon dioxide fixation to small changes in the stomatal conductance and boundary layer conductance are derived. These sensitivity or ‘control’ coefficients, together with those for the carboxylase and oxygenase activities of ribulose 1,5-bisphosphate carboxylase/oxygenase, are calculated from standard gas exchange data and apply under conditions where leaf temperature and water vapour concentration at the leaf surface remain largely constant. It is shown that the magnitude of the control coefficients depends on conditions such as photon flux density, ambient CO₂ concentration and relative humidity at the leaf surface. The extension of this analysis to encompass the sensitivity of the photosynthetic fluxes to changes in enzyme concentrations and kinetic properties is also discussed.     

Simultaneous and independent effects of abscisic acid on stomata and the photosynthetic apparatus in whole leaves

(±)-Abscisic acid (ABA) at 10^-5 M was added to the transpiration stream of leaves of 16 species (C₃ and C₄, monocotyledons and dicotyledons). Stomatal responses followed one of three patterns: i) stomata that were wide and insensitive to CO₂ initially, closed partially and became sensitive to CO₂; ii) for stomata that were sensitive to CO₂ before the application of ABA, the range of highest sensitivity to CO₂ shifted from high to low intercellular partial pressures of CO₂, for instance in leaves of Zea mays from 170–350 to 70–140 bar; iii) when stomata responded strongly to ABA, their conductance was reduced to a small fraction of the initial conductance, and sensitivity to CO₂ was lost. The photosynthetic apparatus was affected by applications of ABA to various degrees, from no response at all (in agreement with several previous reports on the absence of effects of ABA on photosynthesis) through a temporary decrease of its activity to a lasting reduction. Saturation curves of photosynthesis with respect to the partial pressure of CO₂ in the intercellular spaces indicated that application of ABA could produce three phenomena: i) a reduction of the initial slope of the saturation curve (which indicates a diminished carboxylation efficiency); ii) a reduction of the level of the CO₂-saturated rate of assimilation (which indicates a reduction of the ribulose-1,5-bisphosphate regeneration capacity); and iii) an increase of the CO₂ compensation point. Photosynthesis of isolated mesophyll cells was not affected by ABA treatments. Responses of the stomatal and photosynthetic apparatus were usually synchronous and often proportional to each other, with the result that the partial pressure of CO₂ in the intercellular spaces frequently remained constant in spite of large changes in conductance and assimilation rate. Guard cells and the photosynthetic apparatus were able to recover from effects of ABA applications while the ABA supply continued. Recovery was usually partial, in the case of the photosynthetic apparatus occasionally complete. Abscisic acid did not cause stomatal closure or decreases in the rate of photosynthesis when it was applied during a phase of stomatal opening and induction of photosynthesis that followed a transition from darkness to light.Abbreviations and symbols A rate of CO₂ assimilation - ABA (±)-abscisic acid - c _a partial pressure of CO₂ in the ambient air or in the gas supplied to the leaf chambers - c _i partial pressure of CO₂ in the intercellular spaces of a leaf - e _a partial pressure of H₂O in the air - g conductance for water vapor - J quantum flux - T ₁ leaf temperature     

Effects of humidity on short-term responses of stomatal conductance to an increase in carbon dioxide concentration

The magnitude of the response of stomatal conductance to a change in the concentration of carbon dioxide external to the leaf from 350 to 700 cm³ m^–3 was found to be extremely variable from day to day in the field in Glycine max , Hordeum vulgare and Triticum aestivum . It was found that the leaf-to-air water vapour pressure difference (LAVPD) during the midday measurements of the stomatal response to carbon dioxide affected the magnitude of the response. On days when LAVPD was low, no significant change in conductance occurred with the increase in carbon dioxide concentration. When LAVPD was higher, conductance decreased by 24–52% with the increase in carbon dioxide within a few minutes. The sensitivity of conductance was approximately linearly related to LAVPD in wheat and barley. Experiments with G. max in the field indicated that, on days with low LAVPD, increasing the LAVPD just around the measured portion of a leaflet made stomatal conductance responsive to increased carbon dioxide. This result was also obtained under laboratory conditions with G. max , Helianthus annuus and Amaranthus retroflexus . In G. max , it was determined that leaves in which conductance was not responsive to the increase in carbon dioxide could be made responsive even at low LAVPD by the injection of abscisic acid into their petioles. Because it is known that abscisic acid sensitizes stomata to carbon dioxide, these results are consistent with the idea that abscisic acid may be involved in the response of stomatal conductance to changes in LAVPD.     

Distinct light responses of the adaxial and abaxial stomata in intact leaves of Helianthus annuus L

Using a laboratory-constructed system that can measure the gas exchange rates of two leaf surfaces separately, the light responses of the adaxial and abaxial stomata in intact leaves of sunflower ( Helianthus annuus L.) were investigated, keeping the intercellular CO₂ concentration ( C _i) at 300  µ L L⁻¹. When evenly illuminating both sides of the leaf, the stomatal conductance ( g _s) of the abaxial surface was higher than that of the adaxial surface at any light intensity. When each surface of the leaf was illuminated separately, both the adaxial and abaxial stomata were more sensitive to the light transmitted through the leaf (self-transmitted light) than to direct illumination. Relationships between the whole leaf photosynthetic rate ( A _n) and the g _s for each side highlighted a strong dependence of stomatal opening on mesophyll photosynthesis. Light transmitted through another leaf was more effective than the direct white light for the abaxial stomata, but not for the adaxial stomata. Moreover, green monochromatic light induced an opening of the abaxial stomata, but not of the adaxial stomata. As the proportion of blue light in the transmitted light is less than that in the white light, there may be some uncharacterized light responses, which are responsible for the opening of the abaxial stomata by the transmitted, green light.     

Measuring and modelling carbon dioxide and water vapour exchange over a temperate broad-leaved forest during the 1995 summer drought

Forests in the south-eastern United States experienced a prolonged dry spell and above-normal temperatures during the 1995 growing season. During this episode, nearly continuous, eddy covariance measurements of carbon dioxide and water vapour fluxes were acquired over a temperate, hardwood forest. These data are used to examine how environmental factors and accumulating soil moisture deficits affected the diurnal pattern and magnitude of canopy-scale carbon dioxide and water vapour fluxes. The field data are also used to test an integrative leaf-to-canopy scaling model (CANOAK), which uses micrometeorological and physiological theory, to calculate mass and energy fluxes. When soil moisture was ample in the spring, peak rates of net ecosystem CO₂ exchange (N_F) occurred around midday and exceeded 20 μmol m^?2 s^?1. Rates of N_K were near optimal when air temperature ranged between 22 and 25°C. The accumulation of soil moisture deficits and a co-occurrence of high temperatures caused peak rates of daytime carbon dioxide uptake to occur earlier in the morning. High air temperatures and soil moisture deficits were also correlated with a dramatic reduction in the magnitude of N_E. On average, the magnitude of N_E decreased from 20 to 7 μmol m^?2 s^?1 as air temperature increased from 24 to 30°C and the soil dried. The CANAOK model yielded accurate estimates of canopy-scale carbon dioxide and water vapour fluxes when the forest had an ample supply of soil moisture. During the drought and heat spell, a cumulative drought index was needed to adjust the proportionality constant of the stomatal conductance model to yield accurate estimates of canopy CO₂ exchange. The adoption of the drought index also enabled the CANOAK model to give improved estimates of evaporation until midday. On the other hand, the scheme failed to yield accurate estimates of evaporation during the afternoon.     

Effects of environment during growth on the sensitivity of leaf conductance to changes in humidity

Soybeans (Glycine max) and grain amaranth (Amaranthus hypochondriacus) were grown at a range of temperatures, carbon dioxide concentrations and light conditions in controlled environment chambers, and the response of leaf conductance to water vapour to changes in humidity was then measured under a standard set of conditions. The sensitivity of conductance was analysed in terms of (i) the absolute sensitivity of conductance to changes in leaf to air water vapour pressure difference (LAVPD), (ii) the sensitivity of conductance relative to the absolute value of conductance, and (iii) the slope of the relationship between conductance and an index incorporating assimilation rate, carbon dioxide concentration and relative humidity. The sensitivity of conductance varied substantially with growth conditions for all three analyses in both species. The growth temperature of 25 °C increased the sensitivity of conductance by all three measures compared with growth at 20 or 30 °C in amaranth, with little difference between 25 and 30 °C in soybean. Growth at elevated carbon dioxide decreased sensitivity in amaranth by all three measures, and decreased the absolute but not the relative sensitivity in soybean. Growth at reduced photon flux density and growth at high stand density reduced sensitivity in amaranth by all three measures. In soybean, growth at high stand density reduced sensitivity by all three measures, but growth at low photon flux density increased the relative sensitivity. The sensitivity of leaf conductance to changes in humidity varied by a factor of two or more with growth environment by all measures of sensitivity in both the C3 and the C4 species.     

Photosynthesis of cotton plants exposed to elevated levels of carbon dioxide in the field

The cotton (Gossypium hirsutum L.) plant responds to a doubling of atmospheric CO₂ with almost doubled yield. Gas exchange of leaves was monitored to discover the photosynthetic basis of this large response. Plants were grown in the field in open-top chambers with ambient (nominally 350 l/l) or enriched (nominally either 500 or 650 l/l) concentrations of atmospheric CO₂. During most of the season, in fully-irrigated plants the relationship between assimilation (A) and intercellular CO₂ concentration (c_i) was almost linear over an extremely wide range of c_i. CO₂ enrichment did not alter this relationship or diminish photosynthetic capacity (despite accumulation of starch to very high levels) until very late in the season, when temperature was somewhat lower than at midseason. Stomatal conductance at midseason was very high and insensitive to CO₂, leading to estimates of c_i above 85% of atmospheric CO₂ concentration in both ambient and enriched chambers. Water stress caused A to show a saturation response with respect to c_i, and it increased stomatal closure in response to CO₂ enrichment. In fully-irrigated plants CO₂ enrichment to 650 l/l increased A more than 70%, but in water-stressed plants enrichment increased A only about 52%. The non-saturating response of A to c_i, the failure of CO₂ enrichment to decrease photosynthetic capacity for most of the season, and the ability of the leaves to maintain very high c_i, form in part the basis for the very large response to CO₂ enrichment.Abbreviations c_a- atmospheric CO₂ concentration - c_i- intercellular CO₂ concentration - A- rate of assimilation of CO₂ - g_s- stomatal conductance to water vapor - g_b- boundary layer conductance to water vapor - g_m- mesophyll conductance to CO₂ - VPD- vapor pressure deficit - _w leaf water potential - L- stomatal limitation to CO₂ uptake     

Seasonal patterns of photosynthetic response and acclimation to elevated carbon dioxide in field-grown strawberry

Strawberry (Fragaria × ananassa) plants were grown in field plots at the current ambient [CO₂], and at ambient + 300 and ambient + 600 μmol mol⁻¹ [CO₂]. Approximately weekly measurements were made of single leaf gas exchange of upper canopy leaves from early spring through fall of two years, in order to determine the temperature dependence of the stimulation of photosynthesis by elevated [CO₂], whether growth at elevated [CO₂] resulted in acclimation of photosynthesis, and whether any photosynthetic acclimation was reduced when fruiting created additional demand for the products of photosynthesis. Stimulation of photosynthetic CO₂ assimilation by short-term increases in [CO₂] increased strongly with measurement temperature. The stimulation exceeded that predicted from the kinetic characteristics of ribulose-1,5-bisphosphate carboxylase at all temperatures. Acclimation of photosynthesis to growth at elevated [CO₂] was evident from early spring through summer, including the fruiting period in early summer, with lower rates under standard measurement conditions in plants grown at elevated [CO₂]. The degree of acclimation increased with growth [CO₂]. However, there were no significant differences between [CO₂] treatments in total nitrogen per leaf area, and photosynthetic acclimation was reversed one day after switching the [CO₂] treatments. Tests showed that acclimation did not result from a limitation of photosynthesis by triose phosphate utilization rate at elevated [CO₂]. Photosynthetic acclimation was not evident during dry periods in midsummer, when the elevated [CO₂] treatments conserved soil water and photosynthesis declined more at ambient than at elevated [CO₂]. Acclimation was also not evident during the fall, when plants were vegetative, despite wet conditions and continued higher leaf starch content at elevated [CO₂]. Stomatal conductance responded little to short-term changes in [CO₂] except during drought, and changed in parallel with photosynthetic acclimation through the seasons in response to the long-term [CO₂] treatments. The data do not support the hypothesis that source-sink balance controls the seasonal occurrence of photosynthetic acclimation to elevated [CO₂] in this species. This revised version was published online in June 2006 with corrections to the Cover Date.     

Responses of nitrogen metabolism in N-sufficient barley primary leaves to plant growth in elevated atmospheric carbon dioxide

Effects of atmospheric carbon dioxide enrichment on nitrogen metabolism were studied in barley primary leaves (Hordeum vulgare L. cv. Brant). Seedlings were grown in chambers under ambient (36 Pa) and elevated (100 Pa) carbon dioxide and were fertilized daily with complete nutrient solution providing 12 millimolar nitrate and 2.5 millimolar ammonium. Foliar nitrate and ammonium were 27% and 42% lower (P ≤ 0.01) in the elevated compared to ambient carbon dioxide treatments, respectively. Enhanced carbon dioxide affected leaf ammonium levels by inhibiting photorespiration. Diurnal variations of total nitrate were not observed in either treatment. Total and Mg²⁺inhibited nitrate reductase activities per gram fresh weight were slightly lower (P ≤ 0.01) in enhanced compared to ambient carbon dioxide between 8 and 15 DAS. Diurnal variations of total nitrate reductase activity in barley primary leaves were similar in either treatment except between 7 and 10 h of the photoperiod when enzyme activities were decreased (P ≤ 0.05) by carbon dioxide enrichment. Glutamate was similar and glutamine levels were increased by carbon dioxide enrichment between 8 and 13 DAS. However, both glutamate and glutamine were negatively impacted by elevated carbon dioxide when leaf yellowing was observed 15 and 17 DAS. The above findings showed that carbon dioxide enrichment produced only slight modifications in leaf nitrogen metabolism and that the chlorosis of barley primary leaves observed under enhanced carbon dioxide was probably not attributable to a nutritionally induced nitrogen limitation. This revised version was published online in June 2006 with corrections to the Cover Date.     

Elevated ozone reduces photosynthetic carbon gain by accelerating leaf senescence of inbred and hybrid maize in a genotype‐specific manner

Exposure to elevated tropospheric ozone concentration ([O₃]) accelerates leaf senescence in many C₃ crops. However, the effects of elevated [O₃] on C₄ crops including maize (Zea mays L.) are poorly understood in terms of physiological mechanism and genetic variation in sensitivity. Using free air gas concentration enrichment, we investigated the photosynthetic response of 18 diverse maize inbred and hybrid lines to season‐long exposure to elevated [O₃] (~100 nl L^?1) in the field. Gas exchange was measured on the leaf subtending the ear throughout the grain filling period. On average over the lifetime of the leaf, elevated [O₃] led to reductions in photosynthetic CO₂ assimilation of both inbred (?22%) and hybrid (?33%) genotypes. There was significant variation among both inbred and hybrid lines in the sensitivity of photosynthesis to elevated [O₃], with some lines showing no change in photosynthesis at elevated [O₃]. Based on analysis of inbred line B73, the reduced CO₂ assimilation at elevated [O₃] was associated with accelerated senescence decreasing photosynthetic capacity and not altered stomatal limitation. These findings across diverse maize genotypes could advance the development of more O₃ tolerant maize and provide experimental data for parameterization and validation of studies modeling how O₃ impacts crop performance.     

Control of the circadian rhythm of carbon dioxide assimilation in Bryophyllum leaves by exposure to darkness and high carbon dioxide concentrations

The circadian rhythm of CO₂ assimilation in detached leaves of Bryophyllum fedtschenkoi at 15° C in normal air and continuous illumination is inhibited both by exposure to darkness, and to an atmosphere enriched with 5% CO₂. During such exposures substantial fixation of CO₂ takes place, and the malate concentration in the cell sap increases from about 20 mM to a constant value of 40–50 mM after 16 h. On transferring the darkened leaves to light, and those exposed to 5% CO₂ to normal air, a circadian rhythm of CO₂ assimilation begins again. The phase of this rhythm is determined by the time the transfer is made since the first peak occurs about 24 h afterwards. This finding indicates that the circadian oscillator is driven to, and held at, an identical, fixed phase point in its cycle after 16 h exposure to darkness or to 5% CO₂, and it is from this phase point that oscillation begins after the inhibiting condition is removed. This fixed phase point is characterised by the leaves having acquired a high malate content. The rhythm therefore begins with a period of malate decarboxylation which lasts for about 8 h, during which time the malate content of the leaf cells must be reduced to a value that allows phosphoenolpyruvate carboxylase to become active. Inhibition of the rhythm in darkness, and on exposure to 5% CO₂ in continuous illumination, appears to be due to the presence of a high concentration of CO₂ within the leaf inhibiting malic enzyme which leads to the accumulation of high concentrations of malate in the leaf cells. The malate then allosterically inhibits phosphoenolpyruvate carboxylase upon which the rhythm depends. The results give support to the view that malate synthesis and breakdown form an integral part of the circadian oscillator in this tissue.Abbreviations B. Bryophyllum - PEPCase phosphoenolpyruvate carboxylase     

Photosynthesis as related to xylem water potential and carbon dioxide concentration in Sitka spruce

Current-year shoots of Sitka spruce ( Picea sitchensis (Bong.) Carr.) were removed from the forest canopy. After steady-state rates of net photosynthesis were obtained in a leaf chamber, the shoots were excised in air and removed at different times to establish a relationship between net photosynthesis and xylem water potential. The experiment was repeated at five ambient carbon dioxide concentrations.
Net photosynthesis remained constant over a wide range of xylem water potential and increased linearly with ambient carbon dioxide concentration between 20 and 300 cm³ m⁻³. At low water potential net photosynthesis declined at each ambient carbon dioxide concentration and there was little difference in the potential (±0.05 MPa) at which zero photosynthesis was observed.
There was a small increase in the CO₂ compensation concentration at low xylem water potentials, but calculated mesophyll conductance still declined at low water potential after correction for this change in compensation concentration. Mesophyll conductance reached zero within the same range of water potential as net photosynthesis. The results suggested that the non-stomatal contribution to the decline of photosynthesis was approximately 30% until almost complete stomatal closure occurred.