Transpiration by two poplar varieties grown as coppice for biomass production

Fast-growing tree clones selected for biomass plantations are highly productive and therefore likely to use more water than the agricultural crops they replace. We report field measurements of transpiration through the summer of 1994 from two poplar clones, Beaupré (Populus trichocarpa Torr. & A. Gray x P. deltoides Bartr. ex Marsh.) and Dorschkamp (P. deltoides x P. nigra L.), grown as unirrigated short-rotation coppice in southern England. Stand transpiration was quantified by scaling up from sap flow measurements made with the heat balance method in a sample of stems. Leaf conductances, leaf area development, meteorological variables and soil water deficit were also measured to investigate the response of the trees to the environment. High rates of transpiration were found for Beaupré. In June, when soil water was plentiful, the mean (+/- SD) transpiration rate over an 18-day period was 5.0 +/- 1.8 mm day(-1), reaching a maximum of 7.9 mm day(-1). Transpiration rates from Dorschkamp were lower, as a result of its lower leaf area index. High total leaf conductances were measured for both Beaupré (0.34 +/- 0.17 mol m(-2) s(-1)) and Dorschkamp (0.39 +/- 0.16 mol m(-2) s(-1)). Leaf conductance declined slightly with increasing atmospheric vapor pressure deficit in both clones, but only in Beaupré did leaf conductance decrease as soil water deficit increased.     

Changes in photosynthesis and water status of developing leaves of Brachystegia spiciformis Benth

Changes in net carbon assimilation and water status were studied during leaf development in the deciduous, tropical species Brachystegia spiciformis Benth. In this upland savanna African tree, bud-burst and leaf development occur approximately two months before the rainy season. The newly formed leaves synthesize anthocyanin until the fully expanded leaves of the whole canopy are red. This foliage is referred to as "spring flush" foliage. Subsequently, the anthocyanins are metabolized and the pre-rain leaves become green. Carbon dioxide assimilation exhibited a bimodal diurnal pattern and was similar for pre-rain green leaves and fully expanded flushing leaves, although pre-rain green leaves showed a net uptake of carbon throughout the daylight period, whereas flushing leaves exhibited only brief periods of net photosynthesis in the morning and early afternoon. Measurements of leaf water potential and relative water content showed a diurnal pattern with considerable variation throughout the day. Leaf water potential and relative water content values decreased soon after sunrise reaching a minimum at a time corresponding to the afternoon peak in CO(2) assimilation. Stomatal conductance was closely related to transpiration rate in both flushing and pre-rain green leaves, although flushing leaves had lower stomatal conductances than pre-rain green leaves. Pre-rain green leaves exhibited a compensation irradiance of approximately 180 micro mol m(-2) s(-1), whereas flushing leaves had positive net photosynthesis only at PPFDs greater than 300 micro mol m(-2) s(-1). Rate of photosynthesis (expressed per leaf area or chlorophyll unit) increased as anthocyanin concentration decreased, although the photosynthetic rate continued to increase long after the leaf anthocyanins had been degraded to low, visually undetectable amounts. Post-rain green leaves had chlorophyll concentrations, transpiration rates and stomatal conductances similar to those of pre-rain green leaves; however, photosynthetic rates in post-rain leaves were more than three times higher. Thus, during the early stages of the spring flush, carbon asimilation rates of the flushing leaves were inversely related to leaf anthocyanin concentrations. In pre-rain green leaves, photosynthesis was limited by other non-stomatal factors.     

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.     

Diurnal changes in leaf gas exchange characteristics in the uppermost canopy of a rain forest tree, Dryobalanops aromatica Gaertn. f

Dryobalanops aromatica Gaertn. f. is a major tropical canopy species in lowland tropical rain forests in Peninsular Malaysia. Diurnal changes in net photosynthetic rate (A) and stomatal conductance to water vapor (g(s)) were measured in fully expanded young and old leaves in the uppermost canopy (35 m above ground). Maximum A was 12 and 10 micro mol m(-2) s(-1) in young and old leaves, respectively; however, because of large variation in A among leaves, mean maximum A in young and old leaves was only 6.6 and 5.5 micro mol m(-2) s(-1), respectively. Both g(s) and A declined in young leaves when T(leaf) exceeded 34 degrees C and leaf-to-air vapor pressure deficit (DeltaW) exceeded 0.025, whereas in old leaves, g(s) and A did not start to decline until T(leaf) and DeltaW exceeded 36 degrees C and 0.035, respectively. Under saturating light conditions, A was linearly related to g(s). The coefficient of variation (CV) for the difference between the CO(2) concentrations of ambient air and the leaf intercellular air space (C(a) - C(i)) was smaller than the CV for A or g(s), suggesting that maximum g(s) was mainly controlled by mesophyll assimilation (A/C(i)). Minimum C(i)/C(a) ratios were relatively high (0.72-0.73), indicating a small drought-induced stomatal limitation to A and non-conservative water use in the uppermost canopy leaves.     

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.     

Gas exchange characteristics of a Canarian laurel forest tree species (Laurus azorica) in relation to environmental conditions and leaf canopy position

Diurnal courses of gas exchange were measured over a 1-year period in fully expanded current-year leaves in the upper (sun-exposed, 18 m above ground) and the lower (shaded, 12 m above ground) canopy of Laurus azorica (Seub.) Franco, a major canopy species of the Canarian laurel forest in Tenerife, Canary Islands, Spain. Laurus azorica exhibited high leaf plasticity in gas exchange characteristics, with a maximum carbon assimilation rate (Amax) of shade leaves about 50% that of sun leaves. This difference reflects the high leaf area index (LAI) of the stand and the correspondingly sharp light attenuation with increasing canopy depth. In sun leaves, Amax peaked at about 11 micromol m-2 s-1 and maximum transpiration (E) was about 8 mmol m-2 s-1, which corresponded with a maximum stomatal conductance (gs) of about 650 mmol m-2 s-1. Mean maximum instantaneous water-use efficiency (WUE) was 1.5 mmol mol-1 and the mean maximum A/gs was 20-35 micromol mol-1. Mean minimum internal CO2 concentration (Ci) was 225 micromol mol-1. Although high air vapor pressure deficit (VPD) caused a small decrease in gs, it remained high enough to maintain relatively high A and E. These gas exchange characteristics indicate a non-conservative use of water, which is appropriate for a species subject to droughts that are mild or of short duration. In this respect, Laurus azorica differs from its congener, L. nobilis L., of the Mediterranean region and other shrubs growing in Mediterranean-type climates in California and Chile that have to withstand more severe or more prolonged droughts.     

Leaf CO(2) exchange of Erythrina poeppigiana (Leguminosae: Phaseolae) in humid tropical field conditions

An idealized model was developed to describe leaf CO(2) exchange in the leguminous tree Erythrina poeppigiana (Walpers) O.F. Cook under well-watered field conditions. Photosynthetic rate in mature leaves (p) was modeled as a rectangular hyperbolic function of photon flux density (q) and ambient CO(2) concentration (c(a)), relative photosynthetic capacity (pi) was modeled as a logistic s-function of leaf age (l(a)), metabolic dark respiration rate (r(m)) was modeled as an exponential function of leaf temperature (T(l)), and photorespiration rate (r(p)) was modeled as a hyperbolic function of c(a). Assimilation rate (a(c)) was modeled as the difference between the product of p and pi and the sum of r(m) and r(p): a(c) = p(q,c(a))pi(l(a)) - [r(m)(T(l)) + r(p)(c(a))]. The model parameters were estimated separately for five sources of E. poeppigiana (Clones 2660, 2662, 2687 and 2693 and half-sib Family 2431) from field data measured with a portable closed-loop gas exchange system at a humid tropical site in Costa Rica. The between-source differences in leaf CO(2) exchange characteristics were small, but statistically significant. Aboveground biomass production was highest in sources that maintained high relative photosynthetic capacity throughout the leaf life span. Quantum yield varied between 0.046 and 0.067, and light-saturated assimilation rate (q = 2000 micro mol m(-2) s(-1) and T(l) = 28 degrees C) at natural atmospheric c(a) (350 micro mol mol(-1)) was 16.8-19.9 micro mol m(-2) s(-1). Increasing c(a) to 1000 micro mol mol(-1) resulted in an approximate doubling of the light-saturated assimilation rate. Foliole nitrogen concentration, which was 45.3-51.2 mg g(-1) in mature leaves, was positively correlated with relative photosynthetic capacity. Foliole nitrogen concentration, quantum yield and maximum assimilation rate of E. poeppigiana are among the highest values observed in tropical woody legumes.     

Water deficits at anthesis reduce CO(2) assimilation and yield of lychee (Litchi chinensis Sonn.) trees

Ten-year-old 'Tai So' lychee (Litchi chinensis Sonn.) trees growing on a sandy loam soil in subtropical South Africa (latitude 25 degrees S) were watered weekly (well-watered treatment) or droughted from late July until January (drought treatment). After 16 weeks, at which time the trees obtained most of their water from below 150 cm, average soil water content at 0 to 150 cm depth was 14.5 +/- 0.1% in the well-watered treatment and reached a minimum of 7.6% in the drought treatment. At Week 7, minimum leaf water potential (Psi(L)) in the morning and early afternoon declined to -2.6 and -2.8 MPa, respectively, in droughted trees compared with -1.5 and -2.2 MPa, respectively, in well-watered trees. From Week 9, stomatal conductance and net CO(2) assimilation rate ranged from 70 to 300 mmol m(-2) s(-1) and 3 to 13 micro mol CO(2) m(-2) s(-1), respectively, in well-watered trees. The corresponding values for droughted trees were 50 to 180 mmol m(-2) s(-1) and 2 to 6 micro mol CO(2) m(-2) s(-1). Five weeks after rewatering the droughted trees, gas exchange had not recovered to the rate in well-watered trees, although tree water status recovered within a week of rewatering. In the well-watered trees, water use (E(t)) was 26 +/- 1 mm week(-1) with evaporation (E(p)) of 20 to 70 mm week(-1) indicating a crop factor (k(c) = E(t)/E(p)) of 0.4 to 1.2. Before anthesis, tree water status did not affect extension growth of floral panicles or leafy shoots. In contrast, no vegetative shoots were initiated after fruit set in the droughted trees when Psi(L) in the morning declined to -2.5 MPa. Water deficits reduced initial fruit set by 30% and final fruit set by 70% as a result of fruit splitting (41.2 +/- 4.0% versus 10.0 +/- 1.3%). Water deficits did not alter the sigmoidal pattern of fruit growth, but reduced yield from 51.4 +/- 5.5 kg tree(-1) in well-watered trees to 7.4 +/- 3.3 kg tree(-1) in droughted trees.     

Diurnal and seasonal changes in the impact of CO(2) enrichment on assimilation, stomatal conductance and growth in a long-term study of Mangifera indica in the wet-dry tropics of Australia

We studied assimilation, stomatal conductance and growth of Mangifera indica L. saplings during long-term exposure to a CO(2)-enriched atmosphere in the seasonally wet-dry tropics of northern Australia. Grafted saplings of M. indica were planted in the ground in four air-conditioned, sunlit, plastic-covered chambers and exposed to CO(2) at the ambient or an elevated (700 micro mol mol(-1)) concentration for 28 months. Light-saturating assimilation (A(max)), stomatal conductance (g(s)), apparent quantum yield (phi), biomass and leaf area were measured periodically. After 28 months, the CO(2) treatments were changed in all four chambers from ambient to the elevated concentration or vice versa, and A(max) and g(s) were remeasured during a two-week exposure to the new regime. Throughout the 28-month period of exposure, A(max) and apparent quantum yield of leaves in the elevated CO(2) treatment were enhanced, whereas stomatal conductance and stomatal density of leaves were reduced. The relative impacts of atmospheric CO(2) enrichment on assimilation and stomatal conductance were significantly larger in the dry season than in the wet season. Total tree biomass was substantially increased in response to atmospheric CO(2) enrichment throughout the experimental period, but total canopy area did not differ between CO(2) treatments at either the first or the last harvest. During the two-week period following the change in CO(2) concentration, A(max) of plants grown in ambient air but measured in CO(2)-enriched air was significantly larger than that of trees grown and measured in CO(2)-enriched air. There was no difference in A(max) between trees grown and measured in ambient air compared to trees grown in CO(2)-enriched air but measured in ambient air. No evidence of down-regulation of assimilation in response to atmospheric CO(2) enrichment was observed when rates of assimilation were compared at a common intercellular CO(2) concentration. Reduced stomatal conductance in response to atmospheric CO(2) enrichment was attributed to a decline in both stomatal aperture and stomatal density.     

Influence of photosynthetic photon flux density on growth and transpiration in seedlings of Fagus sylvatica

Beech seedlings (Fagus sylvatica L.) were grown in various combinations of three photosynthetic photon flux densities (PPFD, 0.7, 7.3 or 14.5 mol m(-2) day(-1)) for two years in a controlled environmental chamber. Dry mass of leaves, stem and roots, leaf area and number of leaves, and unit leaf rate were affected by both previous-year and current-year PPFD. Number of shoots and length of the main shoot were affected by previous-year PPFD but not by current-year PPFD. Number of leaves per shoot did not change with PPFD, whereas leaf dry mass/leaf area ratio was mainly affected by current-year PPFD. During the first 10 days that newly emerged seedlings were grown at a PPFD of 0.7 or 14.5 mol m(-2) day(-1), transpiration rate per unit leaf area declined. Thereafter, transpiration increased to a constant new rate. Transpiration rate per seedling was closely related to leaf area but the relationship changed with time. In two-year-old seedlings grown at various PPFD combinations of 0.7, 7.3 and 14.5 mol m(-2) day(-1) during Years 1 and 2, leaf area and transpiration rate per seedling were closely correlated at Weeks 7 and 11 after bud burst. Weak correlations were found between root dry mass and transpiration rate per seedling. During Year 2, transpiration rate per leaf area was higher at a particular PPFD in seedlings grown at a previous-year PPFD of 0.7 mol m(-2) day(-1) than in seedlings grown at a previous-year PPFD of 14.5 mol m(-2) day(-1). After transfer of two-year-old seedlings at the end of the experiment to a new PPFD (7.3 or 14.5 mol m(-2) day(-1)) for one day, transpiration rates per leaf area, measured at the new PPFD, were correlated with leaf area and root dry mass, irrespective of former PPFD treatment.     

Effects of light availability on leaf gas exchange and expansion in lychee (Litchi chinensis)

Effects of photosynthetic photon flux density (PPFD) on leaf gas exchange of lychee (Litchi chinensis Sonn.) were studied in field-grown "Kwai May Pink" and "Salathiel" orchard trees and young potted "Kwai May Pink" plants during summer in subtropical Queensland (27 degrees S). Variations in PPFD were achieved by shading the trees or plants 1 h before measurement at 0800 h. In a second experiment, potted seedlings of "Kwai May Pink" were grown in a heated greenhouse in 20% of full sun (equivalent to maximum noon PPFD of 200 micromol m(-2)xs(-1)) and their growth over three flush cycles was compared with seedlings grown in full sun (1080 micromol m(-2)xs(-1)). Young potted plants of "Kwai May Pink" were also grown outdoors in artificial shade that provided 20, 40, 70 or 100% of full sun (equivalent to maximum PPFDs of 500, 900, 1400 and 2000 micromol m(-2)xs(-1)) and measured for shoot extension and leaf area development over one flush cycle. Net CO2 assimilation increased asymptotically in response to increasing PPFD in both orchard trees and young potted plants. Maximum rates of CO2 assimilation (11.9 +/- 0.5 versus 6.3 +/- 0.2 micromol CO2 m(-2) s(-1)), dark respiration (1.7 +/- 0.3 versus 0.6 +/- 0.2 micromol CO2 m(-2) s(-1)), quantum yield (0.042 +/- 0.005 versus 0.027 +/- 0.003 mol CO2 mol(-1)) and light saturation point (1155 versus 959 micromol m(-2) s(-1)) were higher in orchard trees than in young potted plants. In potted seedlings grown in a heated greenhouse, shoots and leaves exposed to full sun expanded in a sigmoidal pattern to 69 +/- 12 mm and 497 +/- 105 cm(2) for each flush, compared with 27 +/- 7 mm and 189 +/- 88 cm(2) in shaded seedlings. Shaded seedlings were smaller and had higher shoot:root ratios (3.7 versus 3.1) than seedlings grown in full sun. In the potted plants grown outdoors in 20, 40, 70 or 100% of full sun, final leaf area per shoot was 44 +/- 1, 143 +/- 3, 251 +/- 7 and 362 +/- 8 cm(2), respectively. Shoots were also shorter in plants grown in shade than in plants grown in full sun (66 +/- 5 mm versus 101 +/- 2 mm). Photosynthesis in individual leaves of lychee appeared to be saturated at about half full sun, whereas maximum leaf expansion occurred at higher PPFDs. We conclude that lychee plants can persist as seedlings on the forest floor, but require high PPFDs for optimum growth.     

Leaf physiology, production, water use, and nitrogen dynamics of the grassland invader Acacia smallii at elevated CO(2) concentrations

Invasion by woody legumes can alter hydrology, nutrient accumulation and cycling, and carbon sequestration on grasslands. The rate and magnitude of these changes are likely to be sensitive to the effects of atmospheric CO(2) enrichment on growth and water and nitrogen dynamics of leguminous shrubs. To assess potential effects of increased atmospheric CO(2) concentrations on plant growth and acquisition and utilization of water and nitrogen, seedlings of Acacia smallii Isely (huisache) were grown for 13 months at CO(2) concentrations of 385 (ambient), 690, and 980 micro mol mol(-1). Seedlings grown at elevated CO(2) concentrations exhibited parallel declines in leaf N concentration and photosynthetic capacity; however, at the highest CO(2) concentration, biomass production increased more than 2.5-fold as a result of increased leaf photosynthetic rates, leaf area, and N(2) fixation. Measurements of leaf gas exchange and aboveground biomass production and soil water balance indicated that water use efficiency increased in proportion to the increase in atmospheric CO(2) concentration. The effects on transpiration of an accompanying decline in leaf conductance were offset by an increase in leaf area, and total water loss was similar across CO(2) treatments. Plants grown at elevated CO(2) fixed three to four times as much N as plants grown at ambient CO(2) concentration. The increase in N(2) fixation resulted from an increase in fixation per unit of nodule mass in the 690 micro mol mol(-1) CO(2) treatment and from a large increase in the number and mass of nodules in plants in the 980 micro mol mol(-1) CO(2) treatment. Increased symbiotic N(2) fixation by woody invaders in response to CO(2) enrichment may result in increased N deposition in litterfall, and thus increased productivity on many grasslands.     

Influence of irrigation and fertilization on transpiration and hydraulic properties of Populus deltoides

Long-term hydraulic acclimation to resource availability was explored in 3-year-old Populus deltoides Bartr. ex Marsh. clones by examining transpiration, leaf-specific hydraulic conductance (G(L)), canopy stomatal conductance (G(S)) and leaf to sapwood area ratio (A(L):A(S)) in response to irrigation (13 and 551 mm year(-1) in addition to ambient precipitation) and fertilization (0 and 120 kg N ha(-1) year(-1)). Sap flow was measured continuously over one growing season with thermal dissipation probes. Fertilization had a greater effect on growth and hydraulic properties than irrigation, and fertilization effects were independent of irrigation treatment. Transpiration on a ground area basis (E) ranged between 0.3 and 1.8 mm day(-1), and increased 66% and 90% in response to irrigation and fertilization, respectively. Increases in G(L), G(S) at a reference vapor pressure deficit of 1 kPa, and transpiration per unit leaf area in response to increases in resource availability were associated with reductions in A(L):A(S) and consequently a minimal change in the water potential gradient from soil to leaf. Irrigation and fertilization increased leaf area index similarly, from an average 1.16 in control stands to 1.45, but sapwood area was increased from 4.0 to 6.3 m(2) ha(-1) by irrigation and from 3.7 to 6.7 m(2) ha(-1) by fertilization. The balance between leaf area and sapwood area was important in understanding long-term hydraulic acclimation to resource availability and mechanisms controlling maximum productivity in Populus deltoides.     

The effect of aqueous transport of CO(2) in xylem sap on gas exchange in woody plants

The influence of CO(2) transported in the transpiration stream on measurements of leaf photosynthesis and stem respiration was investigated. Measurements were made on trees in a temperate forest in Scotland and in a tropical rain forest in Cameroon, and on shrubs in the Sahelian zone in Niger. A chamber was designed to measure the CO(2) partial pressure in the gas phase within the woody stems of trees. High CO(2) partial pressures were found, ranging from 3000 to 9200 Pa. Henry's Law was used to estimate the CO(2) concentration of xylem sap, assuming that it was in equilibrium with the measured gas phase partial pressures. The transport of CO(2) in the xylem sap was calculated by multiplying sap CO(2) concentration by transpiration rate. The magnitude of aqueous transport in the studied species ranged from 0.03 to 0.35 &mgr;mol CO(2) m(-2) s(-1), representing 0.5 to 7.1% of typical leaf photosynthetic rates. These values strongly depend on sap pH. To examine the influence of aqueous transport of CO(2) on stem gas exchange, we made simultaneous measurements of stem CO(2) efflux and sap flow on the same stem. After removing the effect of temperature, stem CO(2) efflux was positively related to sap flow. The apparent effect on measurements of stem respiration was up to 0.7 &mgr;mol m(-2) s(-1), representing ~12% of peak stem respiration rates.     

Carbon dioxide enrichment improves growth, water relations and survival of droughted honey mesquite (Prosopis glandulosa) seedlings

Low water availability reduces the establishment of the invasive shrub Prosopis on some grasslands. Water deficit survival and traits that may contribute to the postponement or tolerance of plant dehydration were measured on seedlings of P. glandulosa Torr. var. glandulosa (honey mesquite) grown at CO(2) concentrations of 370 (ambient), 710, and 1050 micro mol mol(-1). Because elevated CO(2) decreases stomatal conductance, the number of seedlings per container in the elevated CO(2) treatments was increased to ensure that soil water content was depleted at similar rates in all treatments. Seedlings grown at elevated CO(2) had a greater root biomass and a higher ratio of lateral root to total root biomass than those grown at ambient CO(2) concentration; however, these seedlings also shed more leaves and retained smaller leaves. These changes, together with a reduced transpiration/leaf area ratio at elevated CO(2), may have contributed to a slight increase in xylem pressure potentials of seedlings in the 1050 micro mol mol(-1) CO(2) treatment during the first 37 days of growth (0.26 to 0.40 MPa). Osmotic potential was not affected by CO(2) treatment. Increasing the CO(2) concentration to 710 and 1050 micro mol mol(-1) more than doubled the percentage survival of seedlings from which water was withheld for 65 days. Carbon dioxide enrichment significantly increased survival from 0% to about 40% among seedlings that experienced the lowest soil water content. By increasing seedling survival of drought, rising atmospheric CO(2) concentration may increase abundance of P. glandulosa on grasslands where low water availability limits its establishment.     

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.     

黑河流域荒漠地区水分胁迫对梭梭苗木影响的试验研究(英文)

梭梭(HaloxylonAmmodendronBge,一种C4灌木)苗种植在15升的容器中,给予不同的水分胁迫处理,研究了其水分关系和气体交换特征。结果表明:当土壤水分含量大于11%时,梭梭苗有高的蒸腾量;土壤水分含量低于6%时,苗木就不能从土壤中吸取水分;很好供水的苗木的蒸腾量与潜在蒸发量成线型相关。气体交换测定发现,随着土壤水分含量的下降,造成了不同程度的气孔导度、叶蒸腾强度和光合作用的下降。对同一苗木而言,由于这个地区有高的水气压亏缺(VPD),很好和中度供水的苗木在气孔反应方面有较宽的范围,气孔在决定光合作用方面起着较小的作用,二者没有明显的线型相关关系。虽然水分胁迫使蒸腾速率比光合速率下降的更快,提高了水分利用效率,而较高的蒸发需求增加了蒸腾量,限制了光合作用,但是总的趋势是光合作用和蒸腾强度成线型相关。图6表2参15。     

Characteristics of photosynthesis and stomatal conductance in the shrubland species manuka (Leptospermum scoparium) and kanuka (Kunzea ericoides) for the estimation of annual canopy carbon uptake

Responses of photosynthesis to carbon dioxide (CO2) partial pressure and irradiance were measured on leaves of 39-year-old trees of manuka (Leptospermum scoparium J. R. Forst. & G. Forst.) and kanuka (Kunzea ericoides var. ericoides (A. Rich.) J. Thompson) at a field site, and on leaves of young trees grown at three nitrogen supply rates in a nursery, to determine values for parameters in a model to estimate annual net carbon uptake. These secondary successional species belong to the same family and commonly co-occur. Mean (+/- standard error) values of the maximum rate of carboxylation (hemi-surface area basis) (Vcmax) and the maximum rate of electron transport (Jmax) at the field site were 47.3 +/- 1.9 micromol m(-2) s(-1) and 94.2 +/- 3.7 micromol m(-2) s(-1), respectively, with no significant differences between species. Both Vcmax and Jmax were positively related to leaf nitrogen concentration on a unit leaf area basis, and the slopes of these relationships did not differ significantly between species or between the trees in the field and young trees grown in the nursery. Mean values of Jmax/Vcmax measured at 20 degrees C were significantly lower (P < 0.01) for trees in the field (2.00 +/- 0.05) than for young trees in the nursery with similar leaf nitrogen concentrations (2.32 +/- 0.08). Stomatal conductance decreased sharply with increasing air saturation deficit, but the sensitivity of the response did not differ between species. These data were used to derive parameters for a coupled photosynthesis-stomatal conductance model to scale estimates of photosynthesis from leaves to the canopy, incorporating leaf respiration at night, site energy and water balances, to estimate net canopy carbon uptake. Over the course of a year, 76% of incident irradiance (400-700 nm) was absorbed by the canopy, annual net photosynthesis per unit ground area was 164.5 mol m(-2) (equivalent to 1.97 kg C m(-2)) and respiration loss from leaves at night was 37.5 mol m(-2) (equivalent to 0.45 kg m(-2)), or 23% of net carbon uptake. When modeled annual net carbon uptake for the trees was combined with annual respiration from the soil surface, estimated net primary productivity for the ecosystem (0.30 kg C m(-2)) was reasonably close to the annual estimate obtained from independent mensurational and biomass measurements made at the site (0.22 +/- 0.03 kg C m(-2)). The mean annual value for light-use efficiency calculated from the ratio of net carbon uptake (net photosynthesis minus respiration of leaves at night) and absorbed irradiance was 13.0 mmol C mol(-1) (equivalent to 0.72 kg C GJ(-1)). This is low compared with values reported for other temperate forests, but is consistent with limitations to photosynthesis in the canopy attributable mainly to low nitrogen availability and associated low leaf area index.     

Seasonal fluctuations and temperature dependence of leaf gas exchange parameters of co-occurring evergreen and deciduous trees in a temperate broad-leaved forest

Seasonal fluctuations in leaf gas exchange parameters were investigated in three evergreen (Quercus glauca Thunb., Cinnamomum camphora Sieb. and Castanopsis cuspidata Schottky) and one deciduous (Quercus serrata Thunb.) co-occurring, dominant tree species in a temperate broad-leaved forest. Dark respiration rate (Rn), maximum carboxylation rate (Vcmax) and stomatal coefficient (m), the ratio of stomatal conductance to net assimilation rate after adjustment to the vapor pressure deficit and internal carbon dioxide (CO2) concentration, were derived inversely from instantaneous field gas exchange data (one-point method). The normalized values of Rn and Vcmax at the reference temperature of 25 degrees C (Rn25, Vcmax25) and their temperature dependencies (Delta Ha(Rn), Delta Ha(Vcmax)) were analyzed. Parameter Vcmax25 ranged from 24.0-40.3 micromol m(-2) s(-1) and Delta Ha(Vcmax) ranged from 29.1- 67.0 kJ mol(-1). Parameter Rn25 ranged from 0.6-1.4 micromol m(-2) s(-1) and Delta Ha(Rn) ranged from 47.4-95.4 kJ mol(-1). The stomatal coefficient ranged from 7.2-8.2. For the three evergreen trees, a single set of Vcmax25 and Rn25 parameters and temperature dependence curves produced satisfactory estimates of carbon uptake throughout the year, except during the period of simultaneous leaf fall and leaf expansion, which occurs in April and May. In the deciduous oak, declines in Vcmax25 were observed after summer, along with changes in Vcmax25 and Rn25 during the leaf expansion period. In all species, variation in m during periods of leaf expansion and drought should be considered in modeling studies. We conclude that the changes in normalized gas exchange parameters during periods of leaf expansion and drought need to be considered when modeling carbon uptake of evergreen broad-leaved species.     

Interaction of nutrient limitation and elevated CO2 concentration on carbon assimilation of a tropical tree seedling (Cedrela odorata)

Carbon assimilation by Cedrela odorata L. (Meliaceae) seedlings was investigated in ambient and elevated CO2 concentrations ([CO2]) for 119 days, using small fumigation chambers. A solution containing macro- and micronutrients was supplied at two rates. The 5% rate (high rate) was designed to avoid nutrient limitation and allow a maximum rate of growth. The 1% rate (low rate) allowed examination of the effect of the nutrient limitation-elevated CO2 interaction on carbon assimilation. Root growth was stimulated by 23% in elevated [CO2] at a high rate of nutrient supply, but this did not lead to a change in the root:shoot ratio. Total biomass did not change at either rate of nutrient supply, despite an increase in relative growth rate at the low nutrient supply rate. Net assimilation rates and relative growth rates were stimulated by the high rate of nutrient addition, irrespective of [CO2]. We used a biochemical model of photosynthesis to investigate assimilation at the leaf level. Maximum rate of electron transport (Jmax) and maximum velocity of carboxylation (Vcmax) did not differ significantly with CO2 treatment, but showed a substantial reduction at the low rate of nutrient supply. Across both CO2 treatments, mean Jmax for seedlings grown at a high rate of nutrient supply was 75 micromol m(-2) s(-1) and mean Vcmax was 27 micromol m(-2) s(-1). The corresponding mean values for seedlings grown at a low rate of nutrient supply were 36 micromol m(-2) s(-1) and 15 micromol m(-2) s(-1), respectively. Concentrations of leaf nitrogen, on a mass basis, were significantly decreased by the low nutrient supply rate, in proportion to the observed decrease in photosynthetic parameters. Chlorophyll and carbohydrate concentrations of leaves were unaffected by growth [CO2]. Because there was no net increase in growth in response to elevated [CO2], despite increased assimilation of carbon at the leaf level, we hypothesize that the rate of respiration of non-photosynthetic organs was increased.