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 I review studies published up to May 1994, and several unpublished works, which reported at least one of the following: net CO₂ assimilation (A), stomatal conductance (g_s), leaf dark respiration (R_d) leaf nitrogen or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.     

Seasonal trends of light-saturated net photosynthesis and stomatal conductance of loblolly pine trees grown in contrasting environments of nutrition, water and carbon dioxide

Repeated measures analysis was used to evaluate the effect of long-term CO₂ enhancement on seasonal trends of light-saturated rates of net photosynthesis (A_sat) and stomatal conductance to water vapour (g_sat) of 9-year-old loblolly pine (Pinus taeda L.) trees grown in a 2 × 2 factorial experimental design of nutrition and water. A significant interaction effect of CO₂ and nutrition on mean A_sat was observed for juvenile foliage. Also, juvenile foliage exposed to +350 μmol mol^?1 CO₂ had a higher rate of increase of A_sat between late summer and early autumn. This would lead to a greater potential for recharging carbohydrate reserves for winter. Mature foliage was affected by CO_sat, water and nutrient treatments in two ways. First, A_sat was significantly increased as a result of elevated CO₂ in January, a period when stomatal conductance was only 47% of the maximum observed rate. Secondly, the rate of increase of A_sat from winter to early spring was accelerated as a result of both nutrient + water and + 350 μmol mol^?1 CO₂ treatments. This accelerated response resulted in a greater potential for photosynthate production during the period when growth initiation occurred. Nutrient, water or carbon dioxide treatments did not significantly alter trends in g_sat for mature or juvenile foliage. A significant nutrition × CO₂ interaction was observed for the mature foliage, suggesting that g_sat increased with increasing CO₂ and nutrition. These results may have important consequences for the determination of the water use efficiency of loblolly pine. In spite of low g_sat in the winter to early spring period, there was a substantial gain in A_sat attributable to elevated CO₂ concentrations.     

Light and water-use efficiencies of pine shoots exposed to elevated carbon dioxide and temperature

An automatic gas exchange system was used to continuously measure water and carbon fluxes of attached shoots of Scots pine trees (Pinus sylvestris L.) grown in environment-controlled chambers for a 3-year period (1998-2000) and exposed to either normal ambient conditions (CON), elevated CO2 (+350 micro mol mol-1; EC), elevated temperature (+2-6 degrees C; ET) or a combination of EC and ET (ECT). EC treatment enhanced the mean daily total carbon flux per unit projected needle area (Fc.d) by 17-21 %, depending on the year. This corresponds to a 16-24 % increase in light-use efficiency (LUE) based on incident photosynthetically active radiation. The EC treatment reduced the mean daily total water flux (Fw.d) by 1-12 %, corresponding to a 13-35 % increase in water-use efficiency (WUE). The ET treatment increased Fc.d by 10-18 %, resulting in an 8-19 % increase in LUE, and Fw.d by 48-74 %, resulting in a reduction of WUE by 19-34 %. There was no interaction between CO2 and temperature elevation in connection with either carbon or water fluxes, as the carbon flux responded similarly in both ECT and EC, while the water flux in the ECT treatment was similar to that in ET. Regressions indicated that the increase in maximum LUE was greater with increasing air temperature, whereas changes in WUE were related only to high vapour pressure deficit. Furthermore, changes in LUE and WUE caused by ECT treatment displayed strong diurnal and seasonal variation.     

Decreased hydraulic conductance in plants at elevated carbon dioxide

Previous work indicated that long-term exposure to elevated carbon dioxide levels can reduce hydraulic conductance in some species, but the basis of the response was not determined. In this study, hydraulic conductance was measured at concentrations of both 350 and 700 cm³ m^–3 carbon dioxide for plants grown at both concentrations, to determine the reversibility of the response. In Zea mays and Amaranthus hypochondriacus , exposure to the higher carbon dioxide concentration for several hours reduced whole-plant transpiration rate by 22–40%, without any consistent change in leaf water potential, indicating reversible reductions in hydraulic conductance at elevated carbon dioxide levels. Hydraulic conductance in these species grown at both carbon dioxide concentrations responded similarly to measurement concentration of carbon dioxide, indicating that the response was reversible. In Glycine max , which in earlier work had shown a long-term decrease in hydraulic conductance at elevated carbon dioxide levels, and in Abutilon theophrasti , no short-term changes in hydraulic conductance with measurement concentration of carbon dioxide were found, despite lower transpiration rates at elevated carbon dioxide. In G. max and Medicago sativa , growth at high dew-point temperature reduced transpiration rate and decreased hydraulic conductance. The results indicate that both reversible and irreversible decreases in hydraulic conductance can occur at elevated carbon dioxide concentrations, and that both could be responses to reduced transpiration rate, rather than to carbon dioxide concentration itself.     

Seasonal variation in respiration of 1-year-old shoots of scots pine exposed to elevated carbon dioxide and temperature for 4 years

Sixteen 20-year-old Scots pine (Pinus sylvestris L.) trees growing in the field were enclosed for 4 years in environment-controlled chambers that maintained: (1) ambient conditions (CON); (2) elevated atmospheric CO2 concentration (ambient + 350 micro mol mol-1; EC); (3) elevated temperature (ambient +2-6 degrees C; ET); or (4) elevated CO2 and elevated temperature (ECT). The dark respiration rates of 1-year-old shoots, from which needles had been partly removed, were measured over the growing season in the fourth year. In all treatments, the temperature coefficient of respiration, Q10, changed with season, being smaller during the growing season than at other times. Respiration rate varied diurnally and seasonally with temperature, being highest around mid-summer and declining gradually thereafter. When measurements were made at the temperature of the chamber, respiration rates were reduced by the EC treatment relative to CON, but were increased by ET and ECT treatments. However, respiration rates at a reference temperature of 15 degrees C were reduced by ET and ECT treatments, reflecting a decreased capacity for respiration at warmer temperatures (negative acclimation). The interaction between season and treatment was not significant. Growth respiration did not differ between treatments, but maintenance respiration did, and the differences in mean daily respiration rate between the treatments were attributable to the maintenance component. We conclude that maintenance respiration should be considered when modelling respiratory responses to elevated CO2 and elevated temperature, and that increased atmospheric temperature is more important than increasing CO2 when assessing the carbon budget of pine forests under conditions of climate change.     

Quantifying the impact of daily and seasonal variation in sap pH on xylem dissolved inorganic carbon estimates in plum trees

In studies on internal CO₂ transport, average xylem sap pH (pH_x) is one of the factors used for calculation of the concentration of dissolved inorganic carbon in the xylem sap ([]). Lack of detailed pH_x measurements at high temporal resolution could be a potential source of error when evaluating [] dynamics. In this experiment, we performed continuous measurements of CO₂ concentration ([CO₂]) and stem temperature (T_stem), complemented with pH_x measurements at 30‐min intervals during the day at various stages of the growing season (Day of the Year (DOY): 86 (late winter), 128 (mid‐spring) and 155 (early summer)) on a plum tree (Prunus domestica L. cv. Reine Claude d'Oullins). We used the recorded pH_x to calculate [] based on T_stem and the corresponding measured [CO₂]. No statistically significant difference was found between mean [] calculated with instantaneous pH_x and daily average pH_x. However, using an average pH_x value from a different part of the growing season than the measurements of [CO₂] and T_stem to estimate [] led to a statistically significant error. The error varied between 3.25 ± 0.01% under‐estimation and 3.97 ± 0.01% over‐estimation, relative to the true [] data. Measured pH_x did not show a significant daily variation, unlike [CO₂], which increased during the day and declined at night. As the growing season progressed, daily average [CO₂] (3.4%, 5.3%, 7.4%) increased and average pH_x (5.43, 5.29, 5.20) decreased. Increase in [CO₂] will increase its solubility in xylem sap according to Henry's law, and the dissociation of [] will negatively affect pH_x. Our results are the first quantifying the error in [] due to the interaction between [CO₂] and pH_x on a seasonal time scale. We found significant changes in pH_x across the growing season, but overall the effect on the calculation of [] remained within an error range of 4%. However, it is possible that the error could be more substantial for other tree species, particularly if pH_x is in the more sensitive range (pH_x > 6.5).     

Responses of trees to elevated carbon dioxide and climate change

The enhancement in photosynthesis at elevated concentration of carbon dioxide level than the ambient level existing in the atmosphere is widely known. However, many of the earlier studies were based on instantaneous responses of plants grown in pots. The availability of field chambers for growing trees, and long-term exposure studies of tree species to elevated carbon dioxide, has changed much of our views on carbon dioxide acting as a fertiliser. Several tree species showed acclimation or even down-regulation of photosynthetic responses while a few of them showed higher photosynthesis and better growth responses. Whether elevated levels of carbon dioxide can serve as a fertilizer in a changed climate scenario still remains an unresolved question. Forest-Air-Carbon dioxide-Enrichment (FACE) sites monitored at several locations have shown lately, that the acclimation or down regulation as reported in chamber studies is not as wide-spread as originally thought. FACE studies predict that there could be an increase of 23–28% productivity of trees at least till 2050. However, the increase in global temperature could also lead to increased respiration, and limitation of minerals in the soil could lead to reduced responses in growth. Elevated carbon dioxide induces partial closure of leaf stomata, which could lead to reduced transpiration and more economical use of water by the trees. Even if the carbon dioxide acts as a fertilizer, the responses are more pronounced only in young trees. And if there are variations in species responses to growth due to elevated carbon dioxide, only some species are going to dominate the natural vegetation. This will have serious implications on the biodiversity and the structure of the ecosystems. This paper reviews the research done on trees using elevated CO₂ and tries to draw conclusions based on different methods used for the study. It also discusses the possible functional variations in some tree species due to climate change.     

Response of respiration of soybean leaves grown at ambient and elevated carbon dioxide concentrations to day-to-day variation in light and temperature under field conditions

BACKGROUND AND AIMS: Respiration is an important component of plant carbon balance, but it remains uncertain how respiration will respond to increases in atmospheric carbon dioxide concentration, and there are few measurements of respiration for crop plants grown at elevated [CO(2)] under field conditions. The hypothesis that respiration of leaves of soybeans grown at elevated [CO(2)] is increased is tested; and the effects of photosynthesis and acclimation to temperature examined. METHODS: Net rates of carbon dioxide exchange were recorded every 10 min, 24 h per day for mature upper canopy leaves of soybeans grown in field plots at the current ambient [CO(2)] and at ambient plus 350 micromol mol(-1) [CO(2)] in open top chambers. Measurements were made on pairs of leaves from both [CO(2)] treatments on a total of 16 d during the middle of the growing seasons of two years. KEY RESULTS: Elevated [CO(2)] increased daytime net carbon dioxide fixation rates per unit of leaf area by an average of 48 %, but had no effect on night-time respiration expressed per unit of area, which averaged 53 mmol m(-2) d(-1) (1.4 micromol m(-2) s(-1)) for both the ambient and elevated [CO(2)] treatments. Leaf dry mass per unit of area was increased on average by 23 % by elevated [CO(2)], and respiration per unit of mass was significantly lower at elevated [CO(2)]. Respiration increased by a factor of 2.5 between 18 and 26 degrees C average night temperature, for both [CO(2)] treatments. CONCLUSIONS: These results do not support predictions that elevated [CO(2)] would increase respiration per unit of area by increasing photosynthesis or by increasing leaf mass per unit of area, nor the idea that acclimation of respiration to temperature would be rapid enough to make dark respiration insensitive to variation in temperature between nights.     

Stomatal conductance, photosynthesis and respiration of temperate deciduous tree seedlings grown outdoors at an elevated concentration of carbon dioxide

Seedlings of temperate deciduous tree species were grown outdoors at ambient and at an elevated concentration of carbon dioxide to examine how aspects of their gas exchange would be altered by growth at elevated carbon dioxide concentration. Leaf conductances to water vapour and net carbon dioxide exchange rates were determined periodically near midday. Whole-plant carbon dioxide efflux rates in darkness were also determined. The stomatal conductance of leaves of plants grown and measured at 700 cm³ m^?3 carbon dioxide did not differ from that of plants grown and measured at 350 cm³ m^?3 in Malus domestica, Quercus prinus and Quercus robur at any measurement time. In Acer saccharinum, lower conductances occurred for plants grown and measured at elevated carbon dioxide concentration only at measurement temperatures above 33°C. Photo-synthetic adjustment to elevated carbon dioxide concentration was evident only in Q. robur. All species examined had lower rates of dark respiration per unit of mass when grown and measured at elevated carbon dioxide concentration.     

Growth at elevated carbon dioxide concentration reduces hydraulic conductance in alfalfa and soybean

Hydraulic conductances of alfalfa and soybean plants grown in controlled environment chambers at the current ambient carbon dioxide concentration and at twice the current ambient concentration were determined from measurements of transpiration rate and leaf and stem water potentials in the growth conditions. Growth at elevated carbon dioxide concentration reduced both transpiration rate and hydraulic conductance from the soil to the leaf in both species. Hydraulic conductance from the soil to the base of the stem was also lower at elevated carbon dioxide in soybean, but not alfalfa. These measurements identified the stem to leaf hydraulic pathway as a major target of the carbon dioxide effect in both species. The conductance of excised stem segments was much less in plants grown at elevated carbon dioxide in soybeans.     

Effects of elevated carbon dioxide and nitrogen fertilization on nitrate reductase activity in sweetgum and loblolly pine trees in two temperate forests

Nitrogen (N) availability is a major factor limiting plant production in many terrestrial ecosystems and is a key regulator of plant response to elevated CO₂. Plant N status is a function of both soil N availability and plant N uptake and assimilation capacity. As a rate-limiting step in nitrate assimilation, the reduction of nitrate is an important component of plant physiological response to elevated CO₂ and terrestrial carbon sequestration. We examine the effects of elevated CO₂ and N availability on the activity of nitrate reductase, the enzyme catalyzing the reduction of nitrate to nitrite, in two temperate forests—a closed canopy sweetgum (Liquidambar styraciflua) plantation in Tennessee (Oak Ridge National Laboratory (ORNL)) and a loblolly pine (Pinus taeda) stand in North Carolina (Duke). Both CO₂ and N enrichment had species specific impacts on nitrate reductase activity (NaR). Elevated CO₂ and N fertilization decreased foliar NaR in P. taeda, but there were no treatment effects on L. styraciflua NaR at ORNL or Duke. NaR in 1-year P. taeda needles was significantly greater than in 0-year old needles across treatments. P. taeda NaR was negatively correlated with bio-available molybdenum concentrations in soils, suggesting that CO₂ and N-mediated changes in soil nutrient status may be altering soil-plant N-dynamics. The variation in response among species may reflect different strategies for acquiring N and suggests that elevated CO₂ may alter plant N dynamics through changes in NaR.     

Scots pine root distribution derived from radial sap flow patterns in stems of large leaning trees

This study characterizes whole tree root system distribution in a non-destructive way based on its functional parameters, particularly the sap flow patterns in stems. This approach particularly considers sap flow variation across stems, both radial and circumferential patterns of flow that are usually used for a better integration of sap flow density at the whole tree level. We focused at: (1) Showing examples of sap flow variation across stems at a defined situation (high midday values at the period of non-limiting water supply; (2) Analyzing radial flow patterns in terms of root distribution; (3) Validating these results at the stand level (mean data of series of individual trees) using results of classical biometric methods used at the same site; and (4) Applying the results for evaluation of root distribution around leaning trees. Sap flow rate was measured by the heat deformation method on a set of 14 trees at an experimental pine forest stand in Brasschaat (Belgium) during the growing season of 2000. Sap flow variation across stems was measured at a total of 700 points. Amounts of water supplied by superficial (horizontally oriented) and sinker (vertically oriented) roots were estimated from sap flow patterns. The vertical distribution of absorbing roots as derived from the analysis of sap flow patterns in stem sapwood was very similar to the distribution determined by the classical biometric analysis of fine roots. Trees leaning to the East had stem radii at the stump level and crown radii enhanced in the leaning direction. Sinker roots showed higher absorption activities in the leaning direction, but superficial roots were more absorbing in the opposite direction. The application of the above-described method allows for a better evaluation of the whole-tree behavior and facilitates the evaluation of tree and stand properties in traditional forest stands, which are not equipped for detailed scientific research. This may also facilitate practical applications in landscape-level studies.     

Dynamics of daily height growth in Scots pine trees at elevated temperature and CO2

The aim of this study was to analyse and model the effects of elevated temperature and carbon dioxide concentration on daily height growth of 20-year-old Scots pines (Pinus sylvestris L.). The trees were grown with a low nitrogen supply in closed chambers with a factorial combination of two temperature regimes (ambient and elevated) and two carbon dioxide concentrations (ambient and twice ambient). The temperature elevation corresponded to the predicted increase at the site after a doubling in atmospheric CO₂. The height growth of Scots pines was first empirically studied in terms of its onset, cessation and duration, and the allocation of daily height growth within the growing period in 2000 and 2001, and then a model predicting daily height growth as a function of daily temperature and temperature sum was developed. The empirical results showed elevated temperature to be the dominant variable explaining variation in daily height growth. Elevated temperature also hastened both the onset and cessation of height growth, and the temperature sums for both of them were higher in the elevated than in the ambient temperature treatments. The daily variation in height growth could also be explained by the daily mean temperature in the model. Elevated CO₂ concentration had no effect on the onset, cessation or duration of height growth. The amount of height growth was not affected by any of the treatments.     

Irrigation effects on daily and seasonal variations of trunk sap flow and leaf water relations in olive trees

Irrigation effects on whole-plant sap flow and leaf-level water relations were characterised throughout a growing season in an experimental olive (Olea europaea L.) orchard. Atmospheric evaporative demand and soil moisture conditions for irrigated and non-irrigated olive trees were also monitored. Whole-plant water use in field-grown irrigated and rain fed olive trees was determined using a xylem sap flow method (compensation heat-pulse velocity). Foliage gas exchange and water potentials were determined throughout the experimental period. Physiological parameters responded diurnally and seasonally to variations in tree water status, soil moisture conditions and atmospheric evaporative demand. There was a considerable degree of agreement between daily transpiration deduced from heat-pulse velocity and that determined by calibration using the Penman–Monteith equation in the field. Summer drought caused decreasing leaf gas exchange and water potentials, and a progressive increase in hydraulic conductance (stronger in non-irrigated than irrigated trees), probably attributable to modifications in hydraulic properties at the soil-root interface. Negligible hysteresis, attributable to low plant capacitance, was observed in the relationship between leaf water potential and sap flow. A proportional decrease in maximum daily leaf conductance with increasing vapour pressure deficit was observed, while mean daytime canopy stomatal conductance decreased with the season. As a result, plant water use was limited and excessive drought stress prevented. Non-irrigated olive trees recovered after the summer drought, showing a physiological behaviour similar to that of irrigated trees. In addition to physiological and environmental factors, there are endogenous keys (chemical signals) influencing leaf level parameters. Olive trees are confirmed to be economical and sparing users of soil water, with an efficient xylem sap transport, maintenance of significant gas exchange and transpiration, even during drought stress.     

Up-regulation of sucrose metabolizing enzymes in Oncidium goldiana grown under elevated carbon dioxide

Experiments were conducted in controlled growth chambers to evaluate how increase in CO₂ concentration affected sucrose metabolizing enzymes, especially sucrose phosphate synthase (SPS; EC 2.4.1.14) and sucrose synthase (SS; EC 2.4.1.13), as well as carbon metabolism and partitioning in a tropical epiphytic orchid species (Oncidium goldiana). Response of ribulose‐1,5‐bisphosphate carboxylase/oxygenase (Rubisco; EC 4.1.1.39) to elevated CO₂ was determined along with dry mass production, photosynthesis rate, chlorophyll content, total nitrogen and total soluble protein content. After 60 days of growth, there was a 80% and 150% increase in dry mass production in plants grown at 750 and 1 100 μl l^?1 CO₂, respectively, compared with those grown at ambient CO₂ (about 370 μl l^?1). A similar increase in photosynthesis rate was detected throughout the growth period when measured under growth CO₂ conditions. Concomitantly, there was a decline in leaf Rubisco activity in plants in elevated CO₂ after 10 days of growth. Over the growth period, leaf SPS and SS activities were up‐regulated by an average of 20% and 40% for plants grown at 750 and 1100 μl l^?1 CO₂, respectively. Leaf sucrose content and starch content were significantly higher throughout the growth period in plants grown at elevated CO₂ than those at ambient CO₂. The partitioning of photosynthetically fixed carbon between sucrose and starch appeared to be unaffected by the 750 μl l^?1 CO₂ treatment, but it was favored into starch under the 1 100 μl l^?1 CO₂ condition. The activities of SPS and SS in leaf extracts were closely associated with photosynthetic rates and with partitioning of carbon between starch and sucrose in leaves. The data are consistent with the hypothesis that the up‐regulation of leaf SPS and SS might be an acclimation response to optimize the utilization and export of organic carbon with the increased rate of inorganic‐carbon fixation in elevated CO₂ conditions.     

Shoots, roots and ectomycorrhiza formation of pine seedlings at elevated atmospheric carbon dioxide

The effect of elevated atmospheric CO₂ concentration on the growth of shoots, roots, mycorrhizas and extraradical mycorrhizal mycelia of pine (Pinus silvestris L.) was examined. Two and a half-month-old seedlings were inoculated axenically with the mycorrhizal fungus Pisolithus tincto-rius (Pers.) by a method allowing rapid mycorrhiza formation in Petri dishes. The plants were then cultivated for 3 months in growth chambers with daily concentrations of 350 and 600 μmol mol^?1 CO₂ during the day. Whereas plants harvested after 1 and 2 months did not differ appreciably between ambient and increased CO₂ concentrations, after 3 months they developed a considerably higher root biomass (%57%) at elevated CO₂, but did not increase significantly in root length. The mycorrhizal fungus Pisolithus tinctorius, which depended entirely on the plant assimilates in the model system, grew much faster at increased CO₂: 3 times more mycorrhizal root clusters were formed and the extraradical mycelium produced had twice the biomass at elevated as at ambient CO₂. No difference in shoot biomass was found between the two treatments after 91 d. However, since the total water consumption of seedlings was similar in the two treatments, the water use efficiency was appreciably higher for the seedlings at increased CO₂ because of the higher below-ground biomass.     

Photosynthesis and nutrient composition of spinach and fenugreek grown under elevated carbon dioxide concentration

The effect of elevated carbon dioxide concentration on the changes in the biomass, photosynthesis and nutrient composition was investigated in two leafy vegetables. Spinach (Spinacia oleracea L.) and fenugreek (Trigonella foenum-graecum L.) plants were grown in open top chambers under either ambient (ACO₂, 350 ± 50 μmol mol⁻¹) or elevated (ECO₂, 600 ± 50 μmol mol⁻¹) CO₂ concentration and analyzed 40, 60 and 80 days after exposure. The plants grown in ECO₂ had higher net photosynthetic rate and lower stomatal conductance when compared with the plants grown in ACO₂. ECO₂ also changed the nutrient composition: a lower N, Mg and Fe contents and higher C and Ca contents were observed in the leaves of plants exposed to ECO₂ than in those grown at ACO₂.     

Transpiration and whole-tree conductance in ponderosa pine trees of different heights

Changes in leaf physiology with tree age and size could alter forest growth, water yield, and carbon fluxes. We measured tree water flux (Q) for 14 ponderosa pine trees in two size classes (12 m tall and ∼40 years old, and 36 m tall and ∼ 290 years old) to determine if transpiration (E) and whole-tree conductance (g _t) differed between the two sizes of trees. For both size classes, E was approximately equal to Q measured 2 m above the ground: Q was most highly correlated with current, not lagged, water vapor pressure deficit, and night Q was <12% of total daily flux. E for days 165–195 and 240–260 averaged 0.97 mmol m^–2 (leaf area, projected) s^–1 for the 12-m trees and 0.57 mmol m^–2 (leaf area) s^–1 for the 36-m trees. When photosynthetically active radiation (I _P) exceeded the light saturation for photosynthesis in ponderosa pine (900 μmol m^–2 (ground) s^–1), differences in E were more pronounced: 2.4 mmol m^–2 (leaf area) s^–1 for the 12-m trees and 1.2 mmol m^–2 s^–1 for the 36-m trees, yielding g _t of 140 mmol m^–2 (leaf area) s^–1 for the 12-m trees and 72 mmol m^–2 s^–1 for the 36-m trees. Extrapolated to forests with leaf area index =1, the 36-m trees would transpire 117 mm between 1 June and 31 August compared to 170 mm for the 12-m trees, a difference of 15% of average annual precipitation. Lower g _t in the taller trees also likely lowers photosynthesis during the growing season. Received: 19 April 1999 / Accepted: 23 March 2000     

Growth and maintenance components of leaf respiration of cotton grown in elevated carbon dioxide partial pressure

Elevated atmospheric carbon dioxide partial pressures have been shown to have variable direct and indirect effects on plant respiration rates. In this study, growth, leaf respiration, and leaf nitrogen and carbohydrate partitioning were measured in Gossypium hirsutum L. grown in 35 and 65 Pa CO₂ for 30d. Growth and maintenance coefficients of leaf respiration were estimated using gas exchange techniques both at night and during the day. Elevated CO₂ stimulated biomass production (107%) and net photo-synthetic rates (35–50%). Total day-time respiration (R_d) was not significantly affected by growth CO₂ partial pressure. However, night respiration (R_n) of leaves grown in 65 Pa CO₂ was significantly greater than that of plants grown in 35 Pa CO₂. Correlation of R_d and R_n with leaf expansion rates indicated that plants in both CO₂ treatments had equivalent growth respiration coefficients but maintenance respiration was significantly greater in elevated CO₂. Increased maintenance coefficients in elevated CO₂ appeared to be related to increased starch accumulation rather than to changes in leaf nitrogen.