Net photosynthesis and leaf conductance of loblolly pine seedlings in 2 and 21% oxygen as influenced by irradiance, temperature and provenance

Carbon dioxide assimilation and transpiration by secondary needles of two-year-old loblolly pines (Pinus taeda L.) were measured at 2 and 21% (ambient) oxygen. Measurements were made with a Georgia provenance at irradiances (photosynthetic photon flux density) of 150, 300, 700 and 1200 micromol m(-2) s(-1) and a constant temperature of 25 degrees C, and at temperatures of 15, 25 and 35 degrees C and a constant irradiance of 1200 micromol m(-2) s(-1). Measurements were made with provenances from North Carolina, Florida, Arkansas, and Georgia at 25 degrees C and an irradiance of 1200 micromol m(-2) s(-1). There was no significant interaction between the effects of irradiance and oxygen on either net photosynthesis or leaf conductance. Taking all irradiances together, photosynthesis was 16% less and leaf conductance 28% less in 2% oxygen than in 21% oxygen. There was a significant interaction between the effects of temperature and oxygen concentration on both net assimilation and leaf conductance. Net photosynthesis at 21% oxygen relative to that at 2% was significantly reduced at 25 and 35 degrees C, but not at 15 degrees C, whereas leaf conductance at 21% oxygen relative to that at 2% was significantly increased at 15 and 25 degrees C, but not at 35 degrees C. In the provenance study, net photosynthesis was 11% higher and leaf conductance 36% lower in 2% oxygen than in 21% oxygen. There was no significant interaction between the effects of provenance and oxygen on either net photosynthesis or leaf conductance.     

Root water flow and growth of aspen (Populus tremuloides) at low root temperatures

Effects of root zone temperature on growth, shoot water relations, and root water flow were studied in 1-year-old aspen (Populus tremuloides Michx.) seedlings. Seedlings were grown in solution culture and exposed to day/night air temperatures of 22/16 degrees C and solution culture temperatures of 5, 10, or 20 degrees C for 28 days after bud flush. Compared with root growth at 20 degrees C, root growth was completely inhibited at 5 degrees C and inhibited by 97% at 10 degrees C. The 5 and 10 degrees C treatments severely reduced shoot growth, leaf size, and total leaf area. Root water flow was inhibited by the 5 and 10 degrees C treatments. However, when seedlings were grown for 28 days at 5 degrees C and root water flow was measured at 20 degrees C, there was an increase in flow rate. This increase in root water flow was similar in magnitude to the decrease in root water flow observed when seedlings were grown for 28 days at 20 degrees C and root water flow was measured at 5 degrees C. Reduced root water flow of seedlings grown at 5 and 10 degrees C resulted in decreased stomatal conductance, net assimilation, and shoot water potentials. Root water flow was positively correlated with leaf size, total leaf area, shoot length, and new root growth. Transferring seedlings from 5 to 20 degrees C for 24 h significantly increased root water flow, shoot water potential, and net photosynthesis, whereas transferring seedlings from 10 to 20 degrees C resulted in only a slightly increased shoot water potential. Transferring seedlings from 20 to 5 degrees C greatly reduced root water flow, stomatal conductance, and net photosynthesis, whereas shoot water potential decreased only slightly.     

Inhibition and acclimation of C(3) photosynthesis to moderate heat: a perspective from thermally contrasting genotypes of Acer rubrum (red maple)

Effects of moderate heat on growth and photosynthesis were investigated in two clonal genotypes of Acer rubrum L., originally collected from the thermally contrasting habitats of Florida and Minnesota, USA, and known in the horticultural trade for sensitivity and insensitivity to heat, respectively. Under both common garden and warm greenhouse conditions (day/night temperature of 33/25 degrees C), the Florida genotype exhibited more growth than the Minnesota genotype. To determine the physiological parameters associated with this response, plants were acclimated to ambient (27/25 degrees C) or moderately elevated (33/25 degrees C) temperatures for 21 days before measurement of net photosynthesis at temperatures ranging from 25 to 48 degrees C. In vivo measurements of gas exchange and chlorophyll a fluorescence of ambient-acclimated plants revealed that, compared with the Minnesota genotype, the Florida genotype maintained a higher photosynthetic rate, higher stomatal conductance, more open PSII reaction centers, a greater PSII quantum yield and a lower quantum requirement for photosystem II (phi(PSII)) per mole of CO(2) fixed (phi(CO(2) )) throughout the measurement temperature range. When both genotypes were acclimated at 33/25 degrees C and measured at 33 degrees C, analysis of the response of net photosynthesis to calculated intercellular CO(2) concentration indicated that the maximal rate of Rubisco carboxylation (V(cmax)) decreased more in the Minnesota genotype than in the Florida genotype in response to elevated temperature. Additionally, phi(PSII)/phi(CO(2) ) at 33 degrees C was markedly higher for Minnesota plants under photorespiratory conditions, but similar to Florida plants under non-photorespiratory conditions. The results indicate that the higher net photosynthetic rate at 33/25 degrees C of the Florida genotype compared with the Minnesota genotype could be a result of several mechanisms, including the maintenance of a higher V(cmax )and a more efficient quantum requirement of PSII per mole of CO(2) fixed, which is likely the result of lower photorespiration.     

Effects of atmospheric humidity and acclimation temperature on the temperature response of photosynthesis in young Larix decidua Mill

Larch (Larix decidua Mill.) seedlings of a low altitude (600 m) Austrian provenance were raised outdoors and acclimated in chambers for 14 to 24 days during August and September at either 8 degrees C and an atmospheric saturation vapor pressure deficit (DeltaW) of 2.5 Pa kPa(-1), or 24 degrees C and a DeltaW of 6.2 Pa kPa(-1). Subsequently, their rates of photosynthesis, dark respiration and transpiration were measured at temperatures between 5 and 30 degrees C with DeltaW either maintained below 10 Pa kPa(-1) or allowed to increase with temperature up to 38 Pa kPa(-1). Below 15 degrees C the photosynthetic rate of cold-acclimated plants was higher, but above 15 degrees C it was lower, than that of warm-acclimated plants. Temperature acclimation caused a greater shift in the temperature optimum for photosynthesis when DeltaW was kept small than when it was allowed to increase with temperature. When DeltaW was kept small, leaf conductance of cold-acclimated plants, unlike that of warm-acclimated plants, did not increase with temperature above 15 degrees C. When DeltaW increased with temperature, leaf conductance of cold-acclimated plants decreased more rapidly with temperature than that of warm-acclimated plants. Low temperature acclimation increased the rate of photosynthesis below 15 degrees C without affecting leaf conductance, which indicates that there was an adaptation in leaf internal processes. Further evidence of a metabolic adaptation to acclimation temperature is that dark respiration of cold-acclimated plants was twice that of warm-acclimated plants at all temperatures.     

Diurnal patterns of leaf photosynthesis, conductance and water potential at the top of a lowland rain forest canopy in Cameroon: measurements from the Radeau des Cimes

Diurnal patterns of leaf conductance, net photosynthesis and water potential of five tree species were measured at the top of the canopy in a tropical lowland rain forest in southwestern Cameroon. Access to the 40 m canopy was by a large canopy-supported raft, the Radeau des Cimes. The measurements were made under ambient conditions, but the raft altered the local energy balance at times, resulting in elevated leaf temperatures. Leaf water potential was equal to or greater than the gravitational potential at 40 m in the early morning, falling to values as low as -3.0 MPa near midday. Net photosynthesis and conductance were typically highest during midmorning, with values of about 10-12 micro mol CO(2) m(-2) s(-1) and 0.2-0.3 mol H(2)O m(-2) s(-1), respectively. Leaf conductance and net photosynthesis commonly declined through midday with occasional recovery late in the day. Photosynthesis was negatively related to leaf temperature above midday air temperature maxima. These patterns were similar to those observed in other seasonally droughted evergreen communities, such as Mediterranean-climate shrubs, and indicate that environmental factors may cause stomatal closure and limit photosynthesis in tropical rain forests during the midday period.     

抗松材线虫病马尾松无性系夏季光合作用日变化研究

利用LI6400光合仪测定抗松材线虫病马尾松针叶的净光合速率,研究其夏季变化与生理生态因子的关系。结果表明：（1）抗性马尾松净光合速率日变化趋势呈现“双峰”曲线,峰值出现在10时和15时,有明显的“午休”现象。这种现象不是由于气孔阻力的增大而引起叶肉细胞间CO2亏缺造成的,而是非气孔限制起着主要作用。（2）净光合速率（Pn）与胞间CO2浓度（Ci）、蒸腾速率（Tr）、空气温度（Tair）、叶片温度（Tleaf）、光合有效辐射（PAR）都有极显著的相关关系．但经过通径分析发现,对Pn起作用的最主要因子是PAR。     

Does growth temperature affect the temperature responses of photosynthesis and internal conductance to CO2? A test with Eucalyptus regnans

Internal conductance to CO(2) transfer from intercellular spaces to chloroplasts (g(i)) poses a major limitation to photosynthesis, but only three studies have investigated the temperature dependance of g(i). The aim of this study was to determine whether acclimation to 15 versus 30 degrees C affects the temperature response of photosynthesis and g(i) in seedlings of the evergreen tree species Eucalyptus regnans F. Muell. Six-month-old seedlings were acclimated to 15 or 30 degrees C for 6 weeks before g(i) was estimated by simultaneous measurements of gas exchange and chlorophyll fluorescence (variable J method). There was little evidence for acclimation of photosynthesis to growth temperature. In seedlings acclimated to either 15 or 30 degrees C, the maximum rate of net photosynthesis peaked at around 30 or 35 degrees C. Such lack of temperature acclimation may be related to the constant day and night temperature acclimation regime, which differed from most other studies in which night temperatures were lower than day temperatures. Internal conductance averaged 0.25 mol m(-2) s(-1) at 25 degrees C and increased threefold from 10 to 35 degrees C. There was some evidence that g(i) was greater in seedlings acclimated to 15 than to 30 degrees C, which resulted in seedlings acclimated to 15 degrees C having, if anything, a smaller relative limitation due to g(i) than seedlings acclimated to 30 degrees C. Stomatal limitations were also smaller in seedlings acclimated to 15 degrees C than in seedlings acclimated to 30 degrees C. Based on chloroplast CO(2) concentration, neither maximum rates of carboxylation nor RuBP-limited rate of electron transport peaked between 10 and 35 degrees C. Both were described well by an Arrhenius function and had similar activation energies (57-70 kJ mol(-1)). These findings confirm previous studies showing g(i) to be positively related to measurement temperature.     

Moderate shade can increase net gas exchange and reduce photoinhibition in citrus leaves

Daily variations in net gas exchange, chlorophyll a fluorescence and water relations of mature, sun-acclimated grapefruit (Citrus paradisi Macfady.) and orange (Citrus sinensis L. Osbeck) leaves were determined in tree canopies either shaded with 50% shade screens or left unshaded (sunlit). Mean daily maximum photosynthetic photon flux density (PPFD) under shade varied from 500 to 700 micromol m-2 s-1 and was sufficient to achieve maximum net CO2 assimilation rates (A CO2). Responses of grapefruit and orange leaves to shading were remarkably similar. At midday, on bright clear days, the temperatures of sunlit leaves were 2-6 degrees C above air temperature and 1-4 degrees C above the temperatures of shaded leaves. Although midday depressions of stomatal conductance (gs) and A CO2 were observed in both sunlit and shaded leaves, shaded leaves had lower leaf-to-air vapor pressure differences (D) along with higher gs, A CO2 and leaf water-use efficiency than sunlit leaves. Estimated stomatal limitation to A CO2 was generally less than 25% and did not differ between shaded and sunlit leaves. Leaf intercellular CO2 partial pressure was not altered by shade treatment and did not change substantially with increasing D. Radiation and high temperature stress-induced non-stomatal limitation to A CO2 in sunlit leaves was greater than 40%. Reversible photoinhibition of photosystem II efficiency was more pronounced in sunlit than in shaded leaves. Thus, non-stomatal factors play a major role in regulating A CO2 of citrus leaves during radiation and high temperature stress.     

Photosynthesis and water relations of the floodplain tree, boxelder (Acer negundo L.)

During summer, gas exchange and water relations were measured in mature boxelder (Acer negundo L.) trees growing on a floodplain in central Indiana, USA. A shallow (< 1.25-m deep) water table and repeated flooding kept the soil water potential above -0.5 MPa at all times. Net photosynthesis and stomatal conductance were influenced primarily by light and, to a lesser extent, by leaf temperature, but showed no relationships with leaf-to-air water vapor gradient or leaf water potential. Throughout the summer, there was no midday stomatal closure on any measurement day, and leaf water potential at dawn and minimum daily leaf water potential remained above -0.4 and -1.4 MPa, respectively. Nevertheless, there was a seasonal decline in leaf osmotic potentials at saturation and turgor-loss point. Seasonal changes in maximum daily net photosynthesis and stomatal conductance, minimum daily leaf water potential and soil-to-leaf hydraulic conductance were not related to seasonal changes in soil water potential, air or soil temperature, or water table depth. Seasonal responses of net photosynthesis to intercellular CO(2) indicated that net photosynthesis was controlled primarily by nonstomatal factors. High soil water and a shallow water table may have kept soil-to-leaf hydraulic conductance large (5-9 mmol m(-1) s(-1) MPa(-1)) throughout the summer, permitting the trees to keep their stomata open, yet maintain leaf turgor and high net photosynthesis during the hot, low-humidity afternoons. This could also account for the dominance of nonstomatal influences on net photosynthesis.     

Seedlings of five boreal tree species differ in acclimation of net photosynthesis to elevated CO(2) and temperature

Biochemical models of photosynthesis suggest that rising temperatures will increase rates of net carbon dioxide assimilation and enhance plant responses to increasing atmospheric concentrations of CO(2). We tested this hypothesis by evaluating acclimation and ontogenetic drift in net photosynthesis in seedlings of five boreal tree species grown at 370 and 580 &mgr;mol mol(-1) CO(2) in combination with day/night temperatures of 18/12, 21/15, 24/18, 27/21, and 30/24 degrees C. Leaf-area-based rates of net photosynthesis increased between 13 and 36% among species in plants grown and measured in elevated CO(2) compared to ambient CO(2). These CO(2)-induced increases in net photosynthesis were greater for slower-growing Picea mariana (Mill.) B.S.P., Pinus banksiana Lamb., and Larix laricina (Du Roi) K. Koch than for faster-growing Populus tremuloides Michx. and Betula papyrifera Marsh., paralleling longer-term growth differences between CO(2) treatments. Measures at common CO(2) concentrations revealed that net photosynthesis was down-regulated in plants grown at elevated CO(2). In situ leaf gas exchange rates varied minimally across temperature treatments and, contrary to predictions, increasing growth temperatures did not enhance the response of net photosynthesis to elevated CO(2) in four of the five species. Overall, the species exhibited declines in specific leaf area and leaf nitrogen concentration, and increases in total nonstructural carbohydrates in response to CO(2) enrichment. Consequently, the elevated CO(2) treatment enhanced rates of net photosynthesis much more when expressed on a leaf area basis (25%) than when expressed on a leaf mass basis (10%). In all species, rates of leaf net CO(2) exchange exhibited modest declines with increasing plant size through ontogeny. Among the conifers, enhancements of photosynthetic rates in elevated CO(2) were sustained through time across a wide range of plant sizes. In contrast, for Populus tremuloides and B. papyrifera, mass-based photosynthetic rates did not differ between CO(2) treatments. Overall, net photosynthetic rates were highly correlated with relative growth rate as it varied among species and treatment combinations through time. We conclude that interspecific variation may be a more important determinant of photosynthetic response to CO(2) than temperature.     

Net CO(2) assimilation of cacao seedlings following dark chilling

Overnight exposure of cacao (Theobroma cacao L.) seedlings to chilling temperatures between 4.7 and 15.8 degrees C reduced net CO(2) assimilation rate (A) and stomatal conductance to water vapor (g(s)), with temperatures below 10 degrees C causing severe inhibition. Net CO(2) assimilation rates of chilled seedlings recovered to those of nonchilled plants within 7 days. No differences in daytime intercellular CO(2) concentration (c(i)) with overnight temperature were observed on the first day after the chilling treatment, which indicates that the reduction in photosynthesis was not caused by the reduction in stomatal conductance. However, c(i) of chilled plants was much less than that of nonchilled plants on the second day after treatment, which suggests that chilling caused a change in stomatal response to CO(2) concentration. Even 7 days after treatment, when A had recovered to control values, g(s) of chilled leaves was only approximately 70% that of controls. Chilling did not inhibit A through an effect on leaf water potential, which was higher in chilled plants than in unchilled plants.     

Parameterization and testing of a biochemically based photosynthesis model for walnut (Juglans regia) trees and seedlings

The biochemically based leaf photosynthesis model proposed by Farquhar et al. (1980) and the stomatal conductance model proposed by Jarvis (1976) were parameterized for walnut. Responses of photosynthesis to CO(2) and irradiance were used to determine the key parameters of the photosynthesis model. Concurrently, stomatal conductance responses to leaf irradiance (Q), leaf temperature (T(l)), water vapor pressure deficit at the leaf surface (D), and air CO(2) concentration at the leaf surface (C(s)) were used to parameterize the stomatal conductance model. To test the generality of the model parameters, measurements were made on leaves from a 20-year-old tree growing in the field, and from sunlit and shaded greenhouse-grown seedlings. The three key parameters of the photosynthesis model (maximum carboxylation rate V(cmax), electron transport capacity J(max), and dark respiration rate R(d)) and the key parameter of the conductance model (reference stomatal conductance, g(sref)) were linearly correlated with the amount of leaf nitrogen per unit leaf area. Unique relationships could be used to describe nitrogen effects on these parameters for leaves from both the tree and the seedlings. Our data allowed separation of the effects of increasing total photosynthetic apparatus per unit leaf area from the effects of partitioning nitrogen among different pools of this apparatus for foliage acclimation to leaf irradiance. Strong correlations were found between stomatal conductance g(s) and Q, D and C(s), whereas the relationship between g(s) and T(l) was weak. Based on these parameterizations, the model adequately predicted leaf photosynthesis and stomatal conductance when tested with an independent set of data obtained for the tree and seedlings. Total light-driven electron flows derived from chlorophyll fluorescence data obtained at different leaf temperatures were consistent with values computed by the model. The model was also tested with branch bag data acquired from a three-year-old potted walnut tree. Despite a relatively large variance between observed and simulated values, the model predicted stomatal conductance and photosynthesis reasonably well at the branch scale. The results indicate that the photosynthesis-conductance model developed here is robust and can be applied to walnut trees and seedlings under various environmental conditions where water is non-limiting.     

Influence of climate and fertilization on net photosynthesis of mature slash pine

Net photosynthesis was measured under field conditions in 23-year-old slash pine (Pinus elliottii Engelm. var. elliottii) trees to determine how it was affected by fertilization and climate. There was only a small decrease in rates of net photosynthesis from late summer through winter demonstrating that appreciable carbon gain occurs throughout the year in slash pine. Although fertilization substantially increased leaf area and aboveground biomass, it only slightly increased the rate of net photosynthesis. Simultaneous measurements of gas exchange in fertilized and unfertilized (control) plots allowed the detection of a small, but statistically significant difference in average net photosynthesis of 0.14 micro mol m(-2) s(-1). Irradiance, and to a lesser extent air temperature, were the environmental factors that exerted the most control on net photosynthesis. The highest rates of net photosynthesis occurred between air temperatures of 25 and 35 degrees C. Because air temperatures were within this range for 46% of all daylight hours during the year, air temperature was not often a significant limitation. Soil and atmospheric water deficits had less effect on photosynthesis than irradiance or air temperature. Although the depth to the water table changed during the year from 10 to 160 cm, predawn and midday xylem pressure potentials only changed slightly throughout the year. Predawn values ranged from -0.63 to -0.88 MPa in the control plot and from -0.51 to -0.87 MPa in the fertilized plot and were not correlated with water table depth. There was no correlation between xylem pressure potentials and net photosynthesis, presumably because water uptake was adequate. Although vapor pressure deficits reached 3.5 kPa during the summer, they had little effect on net photosynthesis. Over a vapor pressure deficit range from 1.0 to 3.0 kPa, net photosynthesis only decreased 21%. No differences in responses to these environmental factors could be attributed to fertilization.     

Seasonal CO(2) assimilation and stomatal limitations in a Pinus taeda canopy

Net CO(2) assimilation (A(net)) of canopy leaves is the principal process governing carbon storage from the atmosphere in forests, but it has rarely been measured over multiple seasons and multiple years. I measured midday A(net) in the upper canopy of maturing loblolly pine (Pinus taeda L.) trees in the piedmont region of the southeastern USA on 146 sunny days over 36 months. Concurrent data for leaf conductance and photosynthetic CO(2) response curves (A(net)-C(i) curves) were used to estimate the relative importance of stomatal limitations to CO(2) assimilation in the field. In fully expanded current-year and 1-year-old needles, midday light-saturated A(net) was constant over much of the growing season (5-6 &mgr;mol CO(2) m(-2) s(-1)), except during drought periods. During the winter season (November-March), midday A(net) of overwintering needles varied in proportion to leaf temperature. Net CO(2) assimilation at light saturation occurred when daytime air temperatures exceeded 5-6 degrees C, as happened on more than 90% of the sunny winter days. In both age classes of foliage, winter carbon assimilation accounted for approximately 15% of the daily carbon assimilation on sunny days throughout the year, and was relatively insensitive to year-to-year differences in temperature during this season. However, strong stomatal limitations to A(net) occurred as a result of water stress associated with freezing cycles in winter. During the growing season, drought-induced water stress produced the largest year-to-year differences in seasonal CO(2) assimilation on sunny days. Seasonal A(net) was more drought sensitive in current-year needles than in 1-year-old needles. Relative stomatal limitations to daily integrated A(net) were approximately 40% over the growing season, and summer drought rather than high temperatures had the largest impact on summer A(net) and integrated annual CO(2) uptake in the upper crown. Despite significant stomatal limitations, a long duration of near-peak A(net) in the upper crown, particularly in 1-year-old needles, conferred high seasonal and annual carbon gain.     

Temperature response of leaf photosynthetic capacity in seedlings from seven temperate tree species

Seedlings of seven temperate tree species (Acer pseudoplatanus L., Betula pendula Roth, Fagus sylvatica L., Fraxinus excelsior L., Juglans regia L., Quercus petraea Matt. Liebl. and Quercus robur L.) were grown in a nursery under neutral filters transmitting 45% of incident global irradiance. During the second or third year of growth, leaf photosynthetic capacity (i.e., maximal carboxylation rate, Vcmax, maximal photosynthetic electron transport rate, Jmax, and dark respiration, Rd) was estimated for five leaves from each species at five or six leaf temperatures (10, 18, 25, 32, 36 and 40 degrees C). Values of Vcmax and Jmax were obtained by fitting the equations of the Farquhar model on response curves of net CO2 assimilation (A) to sub-stomatal CO2 mole fraction (ci), at high irradiance. Primary parameters describing the kinetic properties of Rubisco (specificity factor, affinity for CO2 and for O2, and their temperature responses) were taken from published data obtained with spinach and tobacco, and were used for all species. The temperature responses of Vcmax and Jmax, which were fitted to a thermodynamic model, differed. Mean values of Vcmax and Jmax at a reference temperature of 25 degrees C were 77.3 and 139 micromol m(-2) s(-1), respectively. The activation energy was higher for Vcmax than for Jmax (mean values of 73.1 versus 57.9 kJ mol(-1)) resulting in a decrease in Jmax/Vcmax ratio with increasing temperature. The mean optimal temperature was higher for Vcmax than for Jmax (38.9 versus 35.9 degrees C). In addition, differences in these temperature responses were observed among species. Temperature optima ranged between 35.9 and above 45 degrees C for Vcmax and between 31.7 and 43.3 degrees C for Jmax, but because of data scatter and the limited range of temperatures tested (10 to 40 degrees C), there were few statistically significant differences among species. The optimal temperature for Jmax was highest in Q. robur, Q. petraea and J. regia, and lowest in A. pseudoplatanus and F. excelsior. Measurements of chlorophyll a fluorescence revealed that the critical temperature at which basal fluorescence begins to increase was close to 47 degrees C, with no difference among species. These results should improve the parameterization of photosynthesis models, and be of particular interest when adapted to heterogeneous forests comprising mixtures of species with diverse ecological requirements.     

Limitation of leaf carbon gain by stomatal and photochemical processes in the top canopy of Macaranga conifera,a tropical pioneer tree

Diurnal changes in gas exchange and chlorophyll fluorescence were measured in the top canopy leaves of the tropical rainforest tree species, Macaranga conifera (Zoll.) Muell. Arg. during a drought year. Maximum values of net photosynthetic rate (P(n), 10 &mgr;mol m(-2) s(-1)) and stomatal conductance (g(s), 0.2 mol m(-2) s(-1)) were found in east-facing leaves in early morning. After 1000 h, both P(n) and g(s) decreased. Minimum daytime values of P(n), g(s), and photosystem II (PSII) quantum yield (DeltaF/F(m)') were found in horizontally fixed leaves. At a given electron transport rate through PSII (ETR), P(n) was higher in early morning than at midday, suggesting a high rate of photorespiration at midday. We tested the hypothesis that the effect of low leaf temperature (T(leaf)) on P(n) is significant in the early morning, whereas the effect of low g(s) on P(n) predominates at midday. In the early morning, when T(leaf) was increased from 32 to 38 degrees C by artificial heating, P(n) at a given ETR decreased 29%, suggesting that the low T(leaf) was associated with a high P(n). When T(leaf) at midday was decreased from 37 to 32 degrees C by artificial cooling, P(n) increased 22%, but P(n) at a given ETR was higher in early morning than at midday, even at the same low T(leaf) (32 degrees C). This suggests that the rate of photorespiration was higher at midday than in early morning because low g(s) at midday caused a reduction in leaf intercellular CO(2) concentration. We conclude that low P(n) at midday was the result of both a reduction in the photochemical process and an increase in stomatal limitation.     

Thermal optima of photosynthetic functions and thermostability of photochemistry in cork oak seedlings

Temperature effects on photosynthesis were studied in seedlings of evergreen Mediterranean cork oak (Quercus suber L.). Responses to changes in temperature and the temperature optima of maximal carboxylation rate (V(cmax)) and maximal light-driven electron flux (J(max)) were estimated from gas exchange measurements and a leaf-level photosynthesis model. The estimated temperature optima were approximately 34 and 33 degrees C for V(cmax) and J(max), respectively, which fall within the lower range of temperature optima previously observed in deciduous tree species. The thermostability of the photosynthetic apparatus was estimated according to the temperature at which basal chlorophyll a fluorescence begins to increase (T(c)). The T(c) was highly variable, increasing from 42 to 51 degrees C when ambient temperature rose from 10 to 40 degrees C, and increasing from 44 to 54 degrees C with decreasing soil water availability while net CO(2) assimilation rate dropped to almost zero. When a heat shock was imposed, an additional small increase in T(c) was observed in drought-stressed and control seedlings. Maximal T(c) values following heat shock were about 56 degrees C, which, to our knowledge, are the highest values that have been observed in tree species. In conclusion, the intrinsic temperature responses of cork oak did not differ from those of other species (similar T(c) under ambient temperature and water availability, and relatively low thermal optima for photosynthetic capacity in seedlings grown at cool temperatures). However, the large ability of cork oak to acclimate to drought and elevated temperature may be an important factor in the tolerance of this evergreen Mediterranean species to summer drought and high temperatures.     

Photosynthesis, water relations, and growth of planted Pinus strobus L. on burned sites in the southern Appalachians

We measured net photosynthesis, leaf conductance, xylem water potential, and growth of Pinus strobus L. seedlings two years after planting on two clear-cut and burned sites in the southern Appalachians. Multiple regression analysis was used to relate seedling net photosynthesis to vapor pressure deficit, seedling crown temperature, photosynthetically active radiation (PAR), needle N, xylem water potential, and soil water, and to relate seedling size and growth to physiological measurements (average net photosynthesis, leaf conductance, and cumulative xylem water potential), soil water, needle N, seedling temperature, and PAR. Seedling net photosynthesis was significantly related to vapor pressure deficit, midday water potential, crown temperature, and PAR (r(2) = 0.70) early in the growing season (May 1992) with vapor pressure deficit alone explaining 42% of the variation. As neighboring vegetation developed, light became more limiting and significantly reduced seedling net photosynthesis later in the growing season (July, August, and September). Final seedling diameter was significantly related to competitor biomass, average photosynthetic rate, and needle N (r(2) = 0.68).     

Diurnal Course of Gas Exchange and Water Use Efficiency of Platycladus orientalis in the Semiarid Region of the Loess Plateau

The daily gas exchange, stomatal conductance, and water use efficiency of Platycladus orientalis in the test field of Fangshan county of Shanxi Province in the semiarid region of the Loess Plateau were measured with portable Li-6200 gas analysis system in natural conditions. The results showed that the diurnal course of net photosynthetic rate displayed a two-peak pattern, that of stomatal conductance displayed a hollow pattern, and that of transpiration rate displayed a signal-peak pattern. Water use efficiency culminated in the early morning. On the basis of two criteria of stomatal limitation of photosynthesis suggested by Farquhar and Sharkey, the predominant limiting factor of photosynthesis was the stomatal conductance of stomatal limitation in the morning (10:00–12:00). However, the midday depression of photosynthesis at noon (12:00–14:00) and the decrease in photosynthesis in the afternoon (16:00–18:00) were the results of nonstomatal limitation, such as low carboxylation capacity of the leaf mesophyll.     

Higher growth temperatures decreased net carbon assimilation and biomass accumulation of northern red oak seedlings near the southern limit of the species range

If an increase in temperature will limit the growth of a species, it will be in the warmest portion of the species distribution. Therefore, in this study we examined the effects of elevated temperature on net carbon assimilation and biomass production of northern red oak (Quercus rubra L.) seedlings grown near the southern limit of the species distribution. Seedlings were grown in chambers in elevated CO(2) (700 μmol mol(-1)) at three temperature conditions, ambient (tracking diurnal and seasonal variation in outdoor temperature), ambient +3 °C and ambient +6 °C, which produced mean growing season temperatures of 23, 26 and 29 °C, respectively. A group of seedlings was also grown in ambient [CO(2)] and ambient temperature as a check of the growth response to elevated [CO(2)]. Net photosynthesis and leaf respiration, photosynthetic capacity (V(cmax), J(max) and triose phosphate utilization (TPU)) and chlorophyll fluorescence, as well as seedling height, diameter and biomass, were measured during one growing season. Higher growth temperatures reduced net photosynthesis, increased respiration and reduced height, diameter and biomass production. Maximum net photosynthesis at saturating [CO(2)] and maximum rate of electron transport (J(max)) were lowest throughout the growing season in seedlings grown in the highest temperature regime. These parameters were also lower in June, but not in July or September, in seedlings grown at +3 °C above ambient, compared with those grown in ambient temperature, indicating no impairment of photosynthetic capacity with a moderate increase in air temperature. An unusual and potentially important observation was that foliar respiration did not acclimate to growth temperature, resulting in substantially higher leaf respiration at the higher growth temperatures. Lower net carbon assimilation was correlated with lower growth at higher temperatures. Total biomass at the end of the growing season decreased in direct proportion to the increase in growth temperature, declining by 6% per 1 °C increase in mean growing season temperature. Our observations suggest that increases in air temperature above current ambient conditions will be detrimental to Q. rubra seedlings growing near the southern limit of the species range.