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.     

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.     

Stomatal responses to drought at a Mediterranean site: a comparative study of co-occurring woody species differing in leaf longevity

We studied stomatal responses to decreasing predawn water potential (Psipd) and increasing leaf-to-air water vapor pressure difference (VPD) of co-occurring woody Mediterranean species with contrasting leaf habits and growth form. The species included two evergreen oaks (Quercus ilex subsp. ballota (Desf.) Samp. and Q. suber L.), two deciduous oaks (Q. faginea Lam. and Q. pyrenaica Willd.) and two deciduous shrubs (Pyrus bourgaeana Decne. and Crataegus monogyna Jacq.). Our main objective was to determine if stomatal sensitivity is related to differences in leaf life span and leaf habit. The deciduous shrubs had the least conservative water-use characteristics, with relatively high stomatal conductance and low stomatal sensitivity to soil and atmospheric drought. As a result, Psipd decreased greatly in both species during the growing season, resulting in early leaf abscission in the summer. The deciduous oaks showed intermediate water-use characteristics, having maximum stomatal conductances and CO2 assimilation rates similar to or even higher than those of the deciduous shrubs. However, they had greater stomatal sensitivity to soil drying and showed less negative Psipd values than the deciduous shrubs. The evergreen oaks, and especially the species with the greatest leaf longevity, Q. ilex, exhibited the most conservative water-use behavior, having lower maximum stomatal conductances and greater sensitivity to VPD than the deciduous species. As a result, Psipd decreased less during the growing season in the evergreens than in the deciduous species, which may contribute to greater leaf longevity by avoiding irreversible damage during the summer drought. However, the combination of low maximum CO2 assimilation rates and high stomatal sensitivity to drought must have a negative impact on the final carbon budget of leaves with a long life span.     

Internal leaf anatomy and photosynthetic resource-use efficiency: interspecific and intraspecific comparisons

Leaf mass per unit area (LMA) and internal leaf anatomy often affect net gas exchange because of their effects on internal CO2 conductance to the site of carboxylation, internal shading, competition for CO2 among carboxylation sites, nitrogen concentration and its partitioning. To evaluate effects of LMA and leaf anatomy on CO2 assimilation, water-use efficiency (WUE) and nitrogen-use efficiency (NUE), we measured LMA, leaf thickness, the thickness of mesophyll components, and gas exchange rates at ambient CO2 concentration in leaves of six woody deciduous and evergreen species with different leaf life spans. In two species, CO2 assimilation was also estimated at saturating CO2 concentrations. There were interspecific differences in all morphological variables studied. Long-lived leaves had higher LMA and were thicker than short-lived leaves. Species with high LMA had low assimilation rates and NUE, both in ambient and saturating CO2 concentrations. Thus, in species with high LMA, assimilation was reduced by non-stomatal limitations, possibly because of a lower allocation of N to the photosynthetic machinery than in species with low LMA. Within a species, thicker leaves tended to have a lower tissue density. In intraspecific comparisons under field conditions, increasing internal air volume had positive effects on WUE, probably because of enhanced internal CO2 conductance to the site of carboxylation. We conclude that, in interspecific comparisons, different patterns of N partitioning strongly influence NUE, whereas in intraspecific comparisons, internal leaf anatomy is a key factor regulating resource-use efficiency.     

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.     

Photosynthetic acclimation of overstory Populus tremuloides and understory Acer saccharum to elevated atmospheric CO2 concentration: interactions with shade and soil nitrogen

We exposed Populus tremuloides Michx. and Acer saccharum Marsh. to a factorial combination of ambient and elevated atmospheric CO2 concentrations ([CO2]) and high-nitrogen (N) and low-N soil treatments in open-top chambers for 3 years. Our objective was to compare photosynthetic acclimation to elevated [CO2] between species of contrasting shade tolerance, and to determine if soil N or shading modify the acclimation response. Sun and shade leaf responses to elevated [CO2] and soil N were compared between upper and lower canopy leaves of P. tremuloides and between A. saccharum seedlings grown with and without shading by P. tremuloides. Both species had higher leaf N concentrations and photosynthetic rates in high-N soil than in low-N soil, and these characteristics were higher for P. tremuloides than for A. saccharum. Electron transport capacity (Jmax) and carboxylation capacity (Vcmax) generally decreased with atmospheric CO2 enrichment in all 3 years of the experiment, but there was no evidence that elevated [CO2] altered the relationship between them. On a leaf area basis, both Jmax and Vcmax acclimated to elevated [CO2] more strongly in shade leaves than in sun leaves of P. tremuloides. However, the apparent [CO2] x shade interaction was largely driven by differences in specific leaf area (m2 g-1) between sun and shade leaves. In A. saccharum, photosynthesis acclimated more strongly to elevated [CO2] in sun leaves than in shade leaves on both leaf area and mass bases. We conclude that trees rooted freely in the ground can exhibit photosynthetic acclimation to elevated [CO2], and the response may be modified by light environment. The hypothesis that photosynthesis acclimates more completely to elevated [CO2] in shade-tolerant species than in shade-intolerant species was not supported.     

Vertical gradients in photosynthetic gas exchange characteristics and refixation of respired CO(2) within boreal forest canopies

We compared vertical gradients in leaf gas exchange, CO(2) concentrations, and refixation of respired CO(2) in stands of Populus tremuloides Michx., Pinus banksiana Lamb. and Picea mariana (Mill.) B.S.P. at the northern and southern boundaries of the central Canadian boreal forest. Midsummer gas exchange rates in Populus tremuloides were over twice those of the two conifer species, and Pinus banksiana rates were greater than Picea mariana rates. Gas exchange differences among the species were attributed to variation in leaf nitrogen concentration. Despite these differences, ratios of intercellular CO(2) to ambient CO(2) (c(i)/c(a)) were similar among species, indicating a common balance between photosynthesis and stomatal conductance in boreal trees. At night, CO(2) concentrations were high and vertically stratified within the canopy, with maximum concentrations near the soil surface. Daytime CO(2) gradients were reduced and concentrations throughout the canopy were similar to the CO(2) concentration in the well-mixed atmosphere above the canopy space. Photosynthesis had a diurnal pattern opposite to the CO(2) profile, with the highest rates of photosynthesis occurring when CO(2) concentrations and gradients were lowest. After accounting for this diurnal interaction, we determined that photosynthesizing leaves in the understory experienced greater daily CO(2) concentrations than leaves at the top of the canopy. These elevated CO(2) concentrations were the result of plant and soil respiration. We estimated that understory leaves in the Picea mariana and Pinus banksiana stands gained approximately 5 to 6% of their carbon from respired CO(2).     

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.     

Effects of CO2 enrichment on the photosynthetic light response of sun and shade leaves of canopy sweetgum (Liquidambar styraciflua) in a forest ecosystem

To investigate whether sun and shade leaves respond differently to CO2 enrichment, we examined photosynthetic light response of sun and shade leaves in canopy sweetgum (Liquidambar styraciflua L.) trees growing at ambient and elevated (ambient + 200 microliters per liter) atmospheric CO2 in the Brookhaven National Laboratory/Duke University Free Air CO2 Enrichment (FACE) experiment. The sweetgum trees were naturally established in a 15-year-old forest dominated by loblolly pine (Pinus taeda L.). Measurements were made in early June and late August 1997 during the first full year of CO2 fumigation in the Duke Forest FACE experiment. Sun leaves had a 68% greater leaf mass per unit area, 63% more leaf N per unit leaf area, 27% more chlorophyll per unit leaf area and 77% greater light-saturated photosynthetic rates than shade leaves. Elevated CO2 strongly stimulated light-saturated photosynthetic rates of sun and shade leaves in June and August; however, the relative photosynthetic enhancement by elevated CO2 for sun leaves was more than double the relative enhancement of shade leaves. Elevated CO2 stimulated apparent quantum yield by 30%, but there was no interaction between CO2 and leaf position. Daytime leaf-level carbon gain extrapolated from photosynthetic light response curves indicated that sun leaves were enhanced 98% by elevated CO2, whereas shade leaves were enhanced 41%. Elevated CO2 did not significantly affect leaf N per unit area in sun or shade leaves during either measurement period. Thus, the greater CO2 enhancement of light-saturated photosynthesis in sun leaves than in shade leaves was probably a result of a greater amount of nitrogen per unit leaf area in sun leaves. A full understanding of the effects of increasing atmospheric CO2 concentrations on forest ecosystems must take account of the complex nature of the light environment through the canopy and how light interacts with CO2 to affect photosynthesis.     

Long-term photosynthetic acclimation to increased atmospheric CO(2) concentration in young birch (Betula pendula) trees

To study the long-term response of photosynthesis to elevated atmospheric CO(2) concentration in silver birch (Betula pendula Roth.), 18 trees were grown in the field in open-top chambers supplied with 350 or 700 &mgr;mol mol(-1) CO(2) for four consecutive growing seasons. Maximum photosynthetic rates, stomatal conductance and CO(2) response curves were measured over the fourth growing season with a portable photosynthesis system. The photosynthesis model developed by Farquhar et al. (1980) was fitted to the CO(2) response curves. Chlorophyll, soluble proteins, total nonstructural carbohydrates, nitrogen and Rubisco activity were determined monthly. Elevated CO(2) concentration stimulated photosynthesis by 33% on average over the fourth growing season. However, comparison of maximum photosynthetic rates at the same CO(2) concentration (350 or 700 &mgr;mol mol(-1)) revealed that the photosynthetic capacity of trees grown in an elevated CO(2) concentration was reduced. Analysis of the response curves showed that acclimation to elevated CO(2) concentration involved decreases in carboxylation efficiency and RuBP regeneration capacity. No clear evidence for a redistribution of nitrogen within the leaf was observed. Down-regulation of photosynthesis increased as the growing season progressed and appeared to be related to the source-sink balance of the trees. Analysis of the main leaf components revealed that the reduction in photosynthetic capacity was accompanied by an accumulation of starch in leaves (100%), which was probably responsible for the reduction in Rubisco activity (27%) and to a lesser extent for reductions in other photosynthetic components: chlorophyll (10%), soluble protein (9%), and N concentrations (12%) expressed on an area basis. Despite a 21% reduction in stomatal conductance in response to the elevated CO(2) treatment, stomatal limitation was significantly less in the elevated, than in the ambient, CO(2) treatment. Thus, after four growing seasons exposed to an elevated CO(2) concentration in the field, the trees maintained increased photosynthetic rates, although their photosynthetic capacity was reduced compared with trees grown in ambient CO(2).     

Fruit load and canopy shading affect leaf characteristics and net gas exchange of 'Spring' navel orange trees

Five-year-old 'Spring' navel (Citrus sinensis (L.) Osbeck) orange trees were completely defruited, 50% defruited or left fully laden to study effects of fruit load on concentrations of nitrogen (N) and carbohydrate, net assimilation of CO2 (Ac) and stomatal conductance (gs) of mature leaves on clear winter days just before fruit harvest. Leaves on defruited trees were larger, had higher starch concentrations and greater leaf dry mass per area (LDMa) than leaves on fruited trees. Both Ac and gs were more than 40% lower in sunlit leaves on defruited trees than in sunlit leaves on trees with fruit. Leaves immediately adjacent to fruit were smaller, had lower leaf nitrogen and carbohydrate concentrations, lower LDMa and lower Ac than leaves on non-fruiting branches of the same trees. Removing half the crop increased individual fruit mass, but reduced fruit color development. Half the trees were shaded with 50% shade cloth for 4 months before harvest to determine the effects of lower leaf temperature (Tl) and leaf-to-air vapor pressure difference on leaf responses. On relatively warm days when sunlit Tl > 25 degrees C, shade increased Ac and gs, but had no effect on the ratio of internal to ambient CO2 (Ci/Ca) concentration in leaves, implying that high mesophyll temperatures in sunlit leaves were more important than gs in limiting Ac. Sunlit leaves were more photoinhibited than shaded leaves on cooler days when Tl < 25 degrees C. Shade decreased total soluble sugar concentrations in leaves, but had no effect on leaf starch concentrations. Shading had no effects on canopy volume, yield or fruit size, but shaded fruit developed better external color than sun-exposed fruit. Overall, the presence of a normal fruit crop resulted in lower foliar carbohydrate concentrations and higher Ac compared with defruited trees, except on warm days when Ac was reduced by high leaf temperatures.     

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.     

Daily time course of whole-shoot gas exchange rates in two drought-exposed Mediterranean shrubs

Effects of drought on water relations, whole-shoot gas-exchange characteristics, and pigment and zeatin concentrations were investigated in the Mediterranean shrubs rosemary (Rosmarinus officinalis L.) and lavender (Lavandula stoechas L.). Two-year-old, greenhouse-grown plants were placed in a whole-shoot gas-exchange measurement system and subjected to 10 days of drought, resulting in severe water stress, and then re-watered for 5 days in order to study their recovery. Water stress resulted in a significant decline in maximum whole-shoot net CO2 assimilation rates (An) for both species that was associated with reductions in leaf area and stomatal conductance. Because shoot dark respiration rate (Rd) was less sensitive to water stress than An, shoot Rd/An ratio increased from about 15 to 95% during water stress. No major changes in chlorophyll and carotenoid concentrations of rosemary leaves were observed during the experiments, but chlorophyll and carotenoid concentrations fell significantly in water-stressed lavender leaves. Zeatin concentrations were higher in rosemary leaves than in lavender leaves during water stress. After re-watering, whole-shoot An and Rd rapidly recovered to their pre-drought rates.     

Photosynthetic gas exchange and accumulation of phytotoxins in mangrove seedlings in response to soil physico-chemical characteristics associated with waterlogging

We evaluated photosynthetic gas exchange and the accumulation of iron, manganese and sulfur in seedlings of five mangrove species (Aegiceras corniculatum (L.) Blanco, Avicennia marina (Forsk.) Vierh., Bruguiera gymnorrhiza (L.) Lamk., Hibiscus tiliaceus L. and Rhizophora stylosa Griff.) growing under anoxic soil conditions at low irradiance. Seedlings of the viviparous mangroves showed no significant responses to root anoxia. The presence of ferrous sulfate or manganous sulfate had a smaller effect on CO(2) assimilation, transpiration rate and stomatal behavior than the presence of sodium sulfide. Sodium sulfide inhibited photosynthetic gas exchange and caused complete stomatal closure in all species. Stomatal closure was probably the result of the damaging effect of sulfide ions on root cell membranes. Some leaf epinasty and wilting were also observed in response to the Na(2)S treatment in all species. A combination of FeSO(4), MnSO(4) and Na(2)S had a smaller effect on photosynthesis than Na(2)S alone, especially for Avicennia marina and Rhizophora stylosa seedlings, which maintained appreciable rates of CO(2) assimilation (2.49 and 3.84 &mgr;mol m(-2) s(-1), respectively) in the presence of all three phytotoxins. Roots of phytotoxin-treated seedlings of all species accumulated significant amounts of the corresponding toxin compared with roots of the control plants. The FeSO(4) or MnSO(4) treatments had no effect on foliar concentrations of iron or manganese, whereas the Na(2)S treatment resulted in an accumulation of S in the leaves of all species. Interactions between Fe(2+) and Mn(2+) and sulfide in the culture medium appeared to reduce their uptake by the seedlings. We conclude that high concentrations of sulfides in mangrove sediments may adversely affect both growth and survival of mangrove seedlings at low irradiances.     

Photosynthetic responses of Scots pine to elevated CO(2) and nitrogen supply: results of a branch-in-bag experiment

Naturally seeded Scots pine (Pinus sylvestris L.) trees, age 25-30 years, were subjected to two soil-nitrogen-supply regimes and to elevated atmospheric CO(2) concentrations by the branch-in-bag method from April 15 to September 15 for two or three years. Gas exchange in detached shoots was measured in a diffuse radiation field. Seven parameters associated with photosynthetic performance and two describing stomatal conductance were determined to assess the effects of treatments on photosynthetic components. An elevated concentration of CO(2) did not lead to a significant downward regulation in maximum carboxylation rate (V(cmax)) or maximum electron transport rate (J(max)), but it significantly decreased light-saturated stomatal conductance (g(sat)) and increased minimum stomatal conductance (g(min)). Light-saturated rates of CO(2) assimilation were higher (24-31%) in shoots grown and measured at elevated CO(2) concentration than in shoots grown and measured at ambient CO(2) concentration, regardless of treatment time or nitrogen-supply regime. High soil-nitrogen supply significantly increased photosynthetic capacity, corresponding to significant increases in V(cmax) and J(max). However, the combined elevated CO(2) + high nitrogen-supply treatment did not enhance the photosynthetic response above that observed in the elevated CO(2) treatment alone.     

Leaf senescence and late-season net photosynthesis of sun and shade leaves of overstory sweetgum (Liquidambar styraciflua) grown in elevated and ambient carbon dioxide concentrations

We examined the effects of elevated CO2 concentration ([CO2]) on leaf demography, late-season photosynthesis and leaf N resorption of overstory sweetgum (Liquidambar styraciflua L.) trees in the Duke Forest Free Air CO2 Enrichment (FACE) experiment. Sun and shade leaves were subdivided into early leaves (formed in the overwintering bud) and late leaves (formed during the growing season). Overall, we found that leaf-level net photosynthetic rates were enhanced by atmospheric CO2 enrichment throughout the season until early November; however, sun leaves showed a greater response to atmospheric CO2 enrichment than shade leaves. Elevated [CO2] did not affect leaf longevity, emergence date or abscission date of sun leaves or shade leaves. Leaf number and leaf area per shoot were unaffected by CO2 treatment. A simple shoot photosynthesis model indicated that elevated [CO2] stimulated photosynthesis by 60% in sun shoots, but by only 3% in shade shoots. Whole-shoot photosynthetic rate was more than 12 times greater in sun shoots than in shade shoots. In senescent leaves, elevated [CO2] did not affect residual leaf nitrogen, and nitrogen resorption was largely unaffected by atmospheric CO2 enrichment, except for a small decrease in shade leaves. Overall, elevated [CO2] had little effect on the number of leaves per shoot at any time during the season and, therefore, did not change seasonal carbon gain by extending or shortening the growing season. Stimulation of carbon gain by atmospheric CO2 enrichment in sweetgum trees growing in the Duke Forest FACE experiment was the result of a strong stimulation of photosynthesis throughout the growing season.     

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.     

Genotypic variation in physiological and growth responses of Populus tremuloides to elevated atmospheric CO2 concentration

Physiological and biomass responses of six genotypes of Populus tremuloides Michx., grown in ambient t (357 micromol mol(-1)) or twice ambient (707 micromol mol(-1)) CO2 concentration ([CO2]) and in low-N or high-N soils, were studied in 1995 and 1996 in northern Lower Michigan, USA. There was a significant CO2 x genotype interaction in photosynthetic responses. Net CO2 assimilation (A) was significantly enhanced by elevated [CO2] for five genotypes in high-N soil and for four genotypes in low-N soil. Enhancement of A by elevated [CO2] ranged from 14 to 68%. Genotypes also differed in their biomass responses to elevated [CO2], but biomass responses were poorly correlated with A responses. There was a correlation between magnitude of A enhancement by elevated [CO2] and stomatal sensitivity to CO2. Genotypes with low stomatal sensitivity to CO2 had a significantly higher A at elevated [CO2] than at ambient [CO2], but elevated [CO2] did not affect the ratio of intercellular [CO2] to leaf surface [CO2]. Stomatal conductance and A of different genotypes responded differentially to recovery from drought stress. Photosynthetic quantum yield and light compensation point were unaffected by elevated [CO2]. We conclude that P. tremuloides genotypes will respond differentially to rising atmospheric [CO2], with the degree of response dependent on other abiotic factors, such as soil N and water availability. The observed genotypic variation in growth could result in altered genotypic representation within natural populations and could affect the composition and structure of plant communities in a higher [CO2] environment in the future.     

Partial shoot removal increases net CO(2) assimilation and alters water relations of Citrus seedlings

Effects of defoliation on partial shoot removal by decapitation on seedling growth, water use and net gas exchange of remaining basal leaves, were examined in Citrus spp. Shoot and root growth rates were manipulated to test for effects of growth demands on net gas exchange. Partially defoliated plants had higher leaf pressure potentials, root conductivities and rates of water use than intact control plants. Shoot regrowth occurred at the expense of root loss. Basal leaves on defoliated plants consistently had higher rates of CO(2) assimilation (A) than leaves on intact plants. Stomatal conductance (g(s)) changed little after defoliation so the higher A of leaves on defoliated plants lowered the ratio of intercellular to ambient CO(2) concentration (C(i)/C(a)) in the mesophyll. In some cases, g(s) increased with A in defoliated plants but C(i)/C(a) was not affected. Stomatal conductance only limited A when intact seedlings were stressed by root confinement in small pots or when leaves were exposed to high vapor pressure deficits during gas exchange measurements. Increased carbon demand for shoot regrowth increased photosynthetic capacity and was more important than stomatal responses in determining A after partial shoot loss.     

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.