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Ecology and ecophysiology of tree stems: corticular and wood photosynthesis.

Below the outer peridermal or rhytidomal layers, most stems of woody plants possess greenish tissues. These chlorophyll-containing tissues (the chlorenchymes) within the stems are able to use the stem internal CO2 and the light penetrating the rhytidome to photoassimilate and produce sugars and starch. Although net photosynthetic uptake of CO2 is rarely found, stem internal re-fixation of CO2 in young twigs and branches may compensate for 60-90% of the potential respiratory carbon loss. Isolated chlorenchymal tissues reveal rather high rates of net photosynthesis (being up to 75% of the respective rates for leaf photosynthesis). Corticular photosynthesis is thus thought to be an effective mechanism for recapturing respiratory carbon dioxide before it diffuses out of the stem. Furthermore, chloroplasts of the proper wood or pith fraction also take part in stem internal photosynthesis. Although there has been no strong experimental evidence until now, we suggest that the oxygen evolved during wood or pith photosynthesis may play a decisive role in avoiding/reducing stem internal anaerobiosis.

Carbon Dioxide↗

Arabidopsis cue mutants with defective plastids are impaired primarily in the photocontrol of expression of photosynthesis-associated nuclear genes.

Plant photoreceptors detect light cues and initiate responses ranging from chloroplast differentiation to the control of morphogenesis and flowering. The photocontrol of photosynthesis-related nuclear genes appears closely related to 'retrograde plastid signals' by which the status of the organelle controls the expression of nuclear genes. However, what specific role, if any, plastid-originated signals play in light responses is poorly understood: it has in the past been proposed that plastid signals play a role in all responses to 'high fluence' far-red light perceived by the light-labile phytochrome A, irrespective of whether they involve photosynthesis-related genes. To explore this further, we have re-examined the phenotype of three cue (cab-underexpressed) Arabidopsis mutants, defective in chloroplast development. The mutants have underdeveloped etioplasts, with increasing impairments in cue6, cue8 and cue3. The mutants show only small defects in photocontrol of hypocotyl elongation and cotyledon opening under prolonged far-red or red light, and normal photocontrol under blue. On the other hand, the expression of photosynthesis-associated nuclear genes is much more impaired in the mutants in the dark and following red or far-red light short treatments or continuous light, than that of those phytochrome-dependent genes tested which are not associated with photosynthesis. Furthermore, red/far-red photoreversible responses involving photosynthesis-related genes (induction of Lhcb1-cab promoter activity, and photoreversible extent of greening) mediated by phytochrome B and other photo-stable phytochromes, both show a reduction in the cue mutants, which correlates with the etioplast defect. Our evidence demonstrates that plastid-derived signals need to be operational in order for the phytochrome control of photosynthetic nuclear genes to occur.

Arabidopsis↗

C3 photosynthesis in silico.

A computer model comprising light reactions, electron-proton transport, enzymatic reactions, and regulatory functions of C3 photosynthesis has been developed as a system of differential budget equations for intermediate compounds. The emphasis is on electron transport through PSII and PSI and on the modeling of Chl fluorescence and 810 nm absorptance signals. Non-photochemical quenching of PSII excitation is controlled by lumenal pH. Alternative electron transport is modeled as the Mehler type O2 reduction plus the malate-oxaloacetate shuttle based on the chloroplast malate dehydrogenase. Carbon reduction enzymes are redox-controlled by the ferredoxin-thioredoxin system, sucrose synthesis is controlled by the fructose 2,6-bisphosphate inhibition of cytosolic FBPase, and starch synthesis is controlled by ADP-glucose pyrophosphorylase. Photorespiratory glycolate pathway is included in an integrated way, sufficient to reproduce steady-state rates of photorespiration. Rate-equations are designed on principles of multisubstrate-multiproduct enzyme kinetics. The parameters of the model were adopted from literature or were estimated from fitting the photosynthetic rate and pool sizes to experimental data. The model provided good simulations for steady-state photosynthesis, Chl fluorescence, and 810 nm transmittance signals under varying light, CO2 and O2 concentrations, as well as for the transients of post-illumination CO2 uptake, Chl fluorescence induction and the 810 nm signal. The modeling shows that the present understanding of photosynthesis incorporated in the model is basically correct, but still insufficient to reproduce the dark-light induction of photosynthesis, the time kinetics of non-photochemical quenching, 'photosynthetic control' of plastoquinone oxidation, cyclic electron flow around PSI, oscillations in photosynthesis. The model may find application for predicting the results of gene transformations, the analysis of kinetic experimental data, the training of students.

Carbon↗

Photosynthetic carbon metabolism in Panicum milioides, a C3-C4 intermediate species: evidence for a limited C4 dicarboxylic acid pathway of photosynthesis.

Panicum milioides, a naturally occurring species with C4-like Kranz leaf anatomy, is intermediate between C3 and C4 plants with respect to photo-respiration and the associated oxygen inhibition of photosynthesis. This paper presents direct evidence for a limited degree of C4 photosynthesis in this C3-C4 intermediate species based on: (a) the appearance of 24% of the total 14C fixed following 4 s photosynthesis in 14CO2-air by excised leaves in malate and aspartate and the complete transfer of label from the C4 acids to Calvin cycle intermediates within a 15 s chase in 12CO2-air; (b) pyruvate- or alanine-enhanced light-dependent CO2 fixation and pyruvate stimulation ote- or alanine-enhanced light-dependent CO2 fixation and pyruvate stimulation of oxaloacetate- or 3-phosphoglycerate-dependent O2 evolution by illuminated mesophyll protoplasts, but not bundle sheath strands; and (c) NAD-malic enzyme-dependent decarboxylation of C4 acids at the C-4 carboxyl position, C4 acid-dependent O2 evolution, and 14CO2 donation from (4-14C)C4 acids to Calvin cycle intermediates during photosynthesis by bundle sheath strands, but not mesophyll protoplasts. However, P. milloides differs from C4 plants in that the activity of the C4 cycle enzymes is only 15 to 30% of a C4 Panicum species and the Calvin cycle and phosphoenolpyruvate carboxylase are present in both cell types. From these and related studies (Rathnam, C.K.M. and Chollet, R. (1979) Arch. Biochem. Biophys. 193, 346-354; (1978) Biochem. Biophys. Res. Commun. 85, 801-808) we conclude that reduced photorespiration in P. milioides is due to a limited degree of NAD-malic enzyme-type C4 photosynthesis permitting an increase in pCO2 at the site of bundle sheath, but not mesophyll, ribulose-bisphosphate carboxylase-oxygenase.

Aspartic Acid↗

Interactions between photosynthesis and 'light-enhanced dark respiration' (LEDR) in the flagellate Euglena gracilis after irradiation with ultraviolet radiation.

The effects of ultraviolet radiation (UV-A, 315-400 nm plus UV-B, 280-315 nm) on photosynthesis and 'light-enhanced dark respiration' (LEDR) in Euglena gracilis have been investigated by using light pulses (80 s) with increasing photon fluence rates of 59, 163, 600, 1180, 2080 and 3340 micromol m(-2) s(-1) and dark periods between the light pulses. LEDR is estimated as the maximum rate of oxygen consumption after a period of light minus the rate of oxygen consumption 30 s after the maximum rate. Without any exposure to UV radiation, the photosynthetic rate and LEDR increase with increasing photon fluence rate. After 20 and 40 min exposures to UV radiation, the photosynthetic rate and LEDR as functions of photon fluence rate are reduced. After a 20 min UV treatment respiration is greater than photosynthesis after the first light pulse of 59 micromol m(-2) s(-1) radiation, and especially at higher photon fluence rates photosynthesis is lower than the control values. The inhibitory effects of UV radiation on photosynthetic rate and LEDR are greater after a 40 min UV exposure than after a 20 min exposure. Only at 600 micromol m(-2) s(-1) is the rate of oxygen evolution greater than that of oxygen consumption after a 40 min UV treatment. Both photosynthetic rate and LEDR are inhibited by the photosynthetic inhibitor DCMU (10(-5) M) in a similar way, which indicates close regulatory interactions between photosynthesis and LEDR. Potassium cyanide (KCN) inhibits dark respiration more than it inhibits LEDR. Dark respiration is not affected to the same degree by UV radiation as are photosynthesis and LEDR.

Animals↗

Development of Arabidopsis thaliana leaves at low temperatures releases the suppression of photosynthesis and photosynthetic gene expression despite the accumulation of soluble carbohydrates.

Arabidopsis thaliana plants were grown at 23 degrees C and changes in carbohydrate metabolism, photosynthesis and photosynthetic gene expression were studied after the plants were shifted to 5 degrees C. The responses of leaves shifted to 5 degrees C after development at 23 degrees C are compared to leaves that developed at 5 degrees C. Shifting warm developed leaves to 5 degrees C lead to a severe suppression of photosynthesis that correlated with a rapid and sustained accumulation of hexose phosphates and soluble sugars. Associated with the suppression of photosynthesis and the accumulation of soluble sugars was a reduction in the amount of transcript for genes encoding photosynthetic proteins (cab and rbcS). In contrast, leaves that developed at 5 degrees C showed an increase in photosynthesis and control levels of photosynthetic gene expression. This recovery occurred even though leaves that developed at 5 degrees C maintained large pools of soluble sugars. Leaves that developed at 5 degrees C also showed a strong upregulation of the cytosolic pathway for soluble sugar synthesis but not of the chloroplastic pathway for starch synthesis. This was shown at the level of both enzyme activity and the amount of transcript. Thus, development of Arabidopsis leaves at 5 degrees C resulted in metabolic changes that enabled them to produce and accumulate large soluble sugar pools without any associated suppression of photosynthesis or photosynthetic gene expression. These changes were also associated with enhanced freezing tolerance. We suggest that this reprogramming of carbohydrate metabolism associated with development at low temperature is essential to the development of full freezing tolerance and for winter survival of over-wintering herbaceous annuals.

Adaptation, Physiological↗

The responses of photosynthesis and oxygen consumption to short-term changes in temperature and irradiance in a cyanobacterial mat (Ebro Delta, Spain).

We have evaluated the effects of short-term changes in incident irradiance and temperature on oxygenic photosynthesis and oxygen consumption in a hypersaline cyanobacterial mat from the Ebro Delta, Spain, in which Microcoleus chthonoplastes was the dominant phototrophic organism. The mat was incubated in the laboratory at 15, 20, 25 and 30 degrees C at incident irradiances ranging from 0 to 1,000 micromol photons m(-2) s(-1). Oxygen microsensors were used to measure steady-state oxygen profiles and the rates of gross photosynthesis, which allowed the calculation of areal gross photosynthesis, areal net oxygen production, and oxygen consumption in the aphotic layer of the mat. The lowest surface irradiance that resulted in detectable rates of gross photosynthesis increased with increasing temperature from 50 micromol photons m(-2) s(-1) at 15 degrees C to 500 micromol photons m(-2) s(-1) at 30 degrees C. These threshold irradiances were also apparent from the areal rates of net oxygen production and point to the shift of M. chthonoplastes from anoxygenic to oxygenic photosynthesis and stimulation of sulphide production and oxidation rates at elevated temperatures. The rate of net oxygen production per unit area of mat at maximum irradiance, J0, did not change with temperature, whereas, JZphot, the flux of oxygen across the lower boundary of the euphotic zone increased linearly with temperature. The rate of oxygen consumption per volume of aphotic mat increased with temperature. This increase occurred in darkness, but was strongly enhanced at high irradiances, probably as a consequence of increased rates of photosynthate exudation, stimulating respiratory processes in the mat. The compensation irradiance (Ec) marking the change of the mat from a heterotrophic to an autotrophic community, increased exponentially in this range of temperatures.

Cyanobacteria↗

Impact of folivory on photosynthesis is greater than the sum of its holes.

The effects of herbivores on plant production and fitness may not relate directly to the quantity of biomass removed because folivory may alter photosynthetic rates at a considerable distance from the damaged tissue [Welter, S. C. (1989) in Insect-Plant Interactions, ed. Bernays, E. A. (CRC, Boca Raton), pp. 135-151.]. An impediment to understanding the effects of leaf damage on photosynthesis has been an inability to map photosynthetic function within a single leaf. We developed an instrument for imaging chlorophyll fluorescence and used it to map the effects of caterpillar feeding on whole-leaf photosynthesis in wild parsnip. The adverse effects of caterpillar feeding on photosynthesis were found to extend well beyond the areas of the leaflet in which caterpillars removed tissue. These "indirectly" affected areas remained impaired for at least 3 days after the caterpillars were removed and were six times as large as the area directly damaged by the caterpillars. Although photosynthesis in indirectly affected areas was reduced and not eliminated, these areas accounted for three times as much of the overall reduction in photosynthesis as the area removed by the caterpillars. The size of the indirect effects was positively correlated with defense-related synthesis of furanocoumarins, suggesting that costs of chemical defense may be one factor that accounts for the indirect effects of herbivory on plants.

Animals↗

Differential use of two cyclic electron flows around photosystem I for driving CO2-concentration mechanism in C4 photosynthesis.

Whereas linear electron flow (LEF) in photosynthesis produces both ATP and NADPH, the cyclic electron flow (CEF) around photosystem I has been shown to produce only ATP. Two alternative routes have been shown for CEF; NAD(P)H dehydrogenase (NDH)- and ferredoxin:plastoquinone oxidoreductase (FQR)-dependent flows, but their physiological relevance has not been elucidated in detail. Meanwhile, because C(4) photosynthesis requires more ATP than does C(3) photosynthesis to concentrate CO(2), it has not been clear how the extra ATP is produced. In this study, to elucidate whether CEF contributes to the additional ATP needed in C(4) photosynthesis, we estimated the amounts of PGR5, which participates in FQR-dependent flow, and NDH-H, a subunit of NDH, in four C(4) species. Although the expression profiles of PGR5 did not correlate well with the additional ATP requirement, NDH was greatly expressed in mesophyll cells in the NAD-malic enzyme (ME) species, and in bundle-sheath cells in NADP-ME species, where there is a strong need for ATP in the respective cells. Our results indicate that CEF via NDH plays a central role in driving the CO(2)-concentrating mechanism in C(4) photosynthesis.

Amino Acid Sequence↗

Effect of 3D nitrogen, dry mass per area and local irradiance on canopy photosynthesis within leaves of contrasted heterogeneous maize crops.

BACKGROUND AND AIMS: Nitrogen partitioning within stands has been described fairly comprehensively, especially for C(3) plants in dense stands where the horizontal heterogeneity of foliage distribution is relatively small. Nitrogen has been shown to be distributed vertically and in parallel to light, maximizing carbon assimilation and stand productivity. Conversely, row crops such as maize (C(4) plants) are characterized by strong horizontal heterogeneity of foliage distribution, and a three-dimensional (3D) approach is required to investigate the combined effect of spatial distribution of nitrogen and light on canopy photosynthesis. MODEL: The 3D geometry of maize canopies was modelled with varying densities and at different developmental stages using plant digitizing under field conditions. For lamina parts, photosynthesis was measured and nitrogen content per unit area (N(a)) was described from analysis of nitrogen content per unit mass (N(m)) and dry mass per unit area (M(a)). Hyperbolic relationships between photosynthesis at irradiance saturation (P(max)) and N(a) were established as well as a linear relationship between dark respiration (R(d)) and N(a), whereas quantum efficiency (alpha) was found to be independent of N(a). KEY RESULTS AND CONCLUSIONS: N(m), M(a) and N(a) were shown to change over time vertically (i.e. between laminae), which has been largely reported previously, and horizontally (i.e. within laminae), which has scarcely been described previously. Even if M(a) played a major role in N(a), a strong relationship between N(a) and M(a) could not be demonstrated, whereas several previous studies have found that N(a) was essentially related to M(a) rather than N(m). From simulations of radiative exchange using a 3D volume-based approach and lamina photosynthesis using a hyperbola, it was shown that real patterns of N(a) partitioning could increase daily crop photosynthesis by up to 8 % compared with uniform patterns of N(a), especially for the earliest stages of stand development.

Light↗

Sink feedback regulation of photosynthesis in vines: measurements and a model.

An experimental and modelling study of source-sink interactions in Vitis vinifera L., cv. Cabernet Sauvignon, rooted cuttings under non-limiting environmental conditions with a 12 h photoperiod is presented here. After 4 h, measured photosynthesis, stomatal conductance and leaf carbohydrate content reached maximum values. Over the remainder of the photoperiod, photosynthesis and stomatal conductance decreased continuously, whereas leaf carbohydrate content remained relatively constant. Because the experiment took place in a non-limiting environment, the results suggest that stomatal regulation of photosynthesis was mediated by an internal factor, possibly related to sink activity. A simple 1-source, 2-sink model was developed to examine the extent to which the data could be explained by a hypothetical sink-to-source feedback mechanism mediated by carbohydrate levels in either the mesophyll, the source phloem or the phloem of one of the two sinks. Model simulations reproduced the data well under the hypothesis of a phloem-based feedback signal, although the data were insufficient to elucidate the detailed nature of such a signal. In a sensitivity analysis, the steady-state response of photosynthesis to sink activity was explored and predictions made for the partitioning of photosynthate between the two sinks. The analysis highlights the effectiveness of a phloem-based feedback signal in regulating the balance between source and sink activities. However, other mechanisms for the observed decline in photosynthesis, such as photoinhibition, endogenous circadian rhythms or hydraulic signals in the leaf cannot be excluded. Nevertheless, it is concluded that the phloem-based feedback model developed here may provide a useful working hypothesis for incorporation into plant growth models and for further development and testing.

Biological Transport, Active↗

Carbon metabolite feedback regulation of leaf photosynthesis and development.

Photosynthesis is regulated as a two-way process. Light regulates the expression of genes for photosynthesis and the activity of the gene products (feedforward control). Rate of end-product use down-stream of the Calvin cycle, determined largely by nutrition and temperature, also affects photosynthetic activity and photosynthetic gene expression (feedback control). Whereas feedforward control ensures efficient light use, feedback mechanisms ensure that carbon flow is balanced through the pathways that produce and consume carbon, so that inorganic phosphate is recycled and nitrogen is distributed optimally to different processes to ensure growth and survival. Actual mechanisms are sketchy and complex, but carbon to nitrogen balance rather than carbon status per se is central to understanding carbon metabolite feedback control of photosynthesis. In addition to determining the activity of the metabolic machinery, carbon metabolite feedback mechanisms also regulate photosynthesis at the leaf level through the regulation of leaf development. This review summarizes the current sketchy, but growing, knowledge of the mechanisms through which carbon metabolite feedback mechanisms regulate leaf photosynthesis.

Carbon↗

Effect of activase level and isoform on the thermotolerance of photosynthesis in Arabidopsis.

Ribulose 1,5-bisphosphate carboxylase/oxygenase (Rubisco) activation decreases under moderate heat stress. This decrease is caused by an impairment of activase function, which is exacerbated by faster rates of Rubisco deactivation at elevated temperatures. To determine if stromal oxidation causes inhibition of activase, transgenic Arabidopsis plants expressing suboptimal amounts of either the redox-regulated 46 kDa alpha- or non-redox regulated 43 kDa beta-isoform of activase were examined. Photosynthesis, as measured by gas exchange and chlorophyll fluorescence, and Rubisco activation were inhibited to a much greater extent by moderately high temperatures in the two transgenic lines expressing suboptimal levels of the individual isoforms of activase compared with wild-type plants or transgenic plants expressing levels of the beta-isoform sufficient for wild-type rates of photosynthesis. Net photosynthesis and Rubisco activation in transgenic plants expressing suboptimal amounts of the beta-isoform of activase from the Antarctic hairgrass were even more sensitive to inhibition by moderate heat stress than in the transgenic plants containing Arabidopsis activase. The results demonstrate that photosynthesis exhibits a similar sensitivity to inhibition by moderately high temperature in plants expressing either of the two different isoforms of activase. Thus, impairment of activase function under heat stress is not caused by oxidation of the redox-sensitive sulphydryls of the alpha-isoform of activase. Instead, the results are consistent with thermal denaturation of activase under moderate heat stress, the effects of which on Rubisco activation would be enhanced when activase levels are suboptimal for photosynthesis.

Arabidopsis↗

Photosynthesis during desiccation in an intertidal alga and a land plant.

This study was undertaken to determine how photosynthesis tolerates desiccation in an intertidal alga Fucus vesiculosus L. and a terrestrial sunflower Helianthus annuus L. Photosynthetic O2 evolution generally was inhibited at low water potentials (psiw) but more in sunflower leaves than in Fucus fronds at the same psiw. As psiw decreased, less carbon accumulated in an organic carbon store in Fucus. The inhibition of photosynthesis appeared to be mostly biochemical because it could not be prevented by supplying additional CO2 or by supplying CO2 from the internal organic carbon store. The inhibition of photosynthesis and carbon storage occurred after turgor disappeared and thus when solute concentrations were increasing in the cells. Solute concentrations were much higher in Fucus than in sunflower. After desiccation to the air-dry state (psiw below - 10 MPa), photosynthesis could not recover in sunflower but it recovered rapidly when Fucus was exposed to seawater. The lack of recovery in sunflower was associated with inability to recover turgor probably because of breaks in cell membranes. The ability to recover in Fucus was gradually lost during 1.5 d of desiccation at 45% relative humidity. At lower humidities, recovery was lost sooner as small amounts of water were removed. We conclude that photosynthesis tolerated desiccation more in Fucus than in sunflower because of differences in the molecular environment around the photosynthetic enzymes. Important aspects of this environment were features that prevented membrane breakage but promoted the retention of small amounts of water that were critical for viability.

Carbon Dioxide↗

The mechanism to suppress photosynthesis through end-product inhibition in single-rooted soybean leaves during acclimation to CO(2) enrichment.

Single-rooted soybean leaves were used to investigate the suppression of photosynthesis through end-product inhibition during acclimation to CO(2 )enrichment. The photosynthetic activity was greater in leaves cultured at a CO(2) partial pressure of 70 Pa (high-CO(2)) than that in the leaves cultured at 35 Pa CO(2) (control) during the initial exposure to CO(2) enrichment but then decreased rapidly with a large accumulation of starch, to well below the level of the control leaves. The response curve of photosynthesis (A) to the intercellular CO(2) concentration (Ci) in the high-CO(2) leaves cultured long-term exhibited a significantly low initial gradient. However, on exposure to darkness for 48 h, the initial gradient of the A to Ci curve and rate of photosynthesis were completely restored, and almost all of the accumulated starch was expended. The ribulose bisphosphate carboxylase (RuBPcase) content and activation ratio in the high-CO(2) leaves remained high and roughly constant during the experiment, and were unchanged by the exposure, while this enzyme was slightly inactivated or inhibited after long-term exposure to CO(2) enrichment. The lower rate of photosynthesis in the high-CO(2) leaves could be linearly increased to a rate approaching the control level by increasing the external atmospheric [CO(2)], which thereby compensated for a reduced CO(2) transfer diffusion from the intercellular space to the stroma in chloroplasts. It is consequently concluded that, during the acclimation to CO(2 )enrichment, the suppression of photosynthesis through end-product inhibition was mainly caused by a lowering of the carboxylation efficiency of RuBPcase due to hindrance of CO(2) diffusion from the intercellular space to the stroma in chloroplasts brought about by the large accumulation of starch.

Adaptation, Physiological↗

Chill-induced inhibition of photosynthesis: genotypic variation within Cucumis sativus.

Cucumber is generally a thermophilic species; however, cultivars have been selected for higher yield during winter cultivation in unheated glasshouses in temperate regions. We tested whether photosynthesis in these varieties had greater chilling tolerance. There was no difference in the instantaneous reduction of photosynthesis at low temperature between four winter glasshouse and four summer field cultivars. After 5 d of 10 degrees C and 100 micro mol m(-2) s(-1) photon flux density, the four field cultivars had a sustained depression of photosynthesis after returning to clement conditions. This inhibition was associated with reduced rates of CO(2) fixation and photosystem II (PSII) electron transport in the light, but not with sustained PSII photoinhibition. However, photosynthesis in the glasshouse genotypes was nearly identical to the pre-chill rates. Chill impacts on light-adapted chlorophyll fluorescence parameters, such as the quantum yield of PSII electron transport (phi(PSII)), correlated well with overall photosynthesis. This demonstrates the potential for using these fast and non-invasive techniques to screen for chill-tolerant genotypes, with the potential to further improve winter cucumber yield in unheated glasshouses.

Acclimatization↗

Relationships between net photosynthesis and foliar nitrogen concentrations in a loblolly pine forest ecosystem grown in elevated atmospheric carbon dioxide.

We examined the effects of elevated carbon dioxide concentration ([CO2]) on the relationship between light-saturated net photosynthesis (A(sat)) and area-based foliar nitrogen (N) concentration (N(a)) in the canopy of the Duke Forest FACE experiment. Measurements of A(sat) and N(a) were made on two tree species growing in the forest overstory and four tree species growing in the forest understory, in ambient and elevated [CO2] FACE rings, during early and late summer of 1999, 2001 and 2002, corresponding to years three, five and six of CO2 treatment. When measured at the growth [CO2], net photosynthetic rates of each species examined in the forest overstory and understory were stimulated by elevated [CO2] at each measurement date. We found no effect of elevated [CO2] on N(a) in any of the species. The slope of the A(sat)-N relationship was 81% greater in elevated [CO2] than in ambient [CO2] when averaged across all sample dates, reflecting a differential CO2 effect on photosynthesis at the top and bottom of the canopy. We compared A(sat)-N relationships in trees grown in ambient and elevated [CO2] at two common CO2 concentrations, during late summer 2001 and both early and late 2002, to determine if the stimulatory effect of elevated [CO2] on photosynthesis diminishes over time. At all three sample times, neither the slopes nor the y-intercepts of the A(sat)-N relationships of trees grown in ambient or elevated [CO2] differed when measured at common CO2 concentrations, indicating that the responses of photosynthesis to long-term elevated [CO2] did not differ from the responses to a short-term increase in [CO2]. This finding, together with the observation that N(a) was unaffected by growth in elevated [CO2], indicates that these overstory and understory trees growing at the Duke Forest FACE experiment continue to show a strong stimulation of photosynthesis by elevated [CO2].

Atmosphere↗

Photosynthesis of the grapevine (Vitis vinifera) inflorescence.

To further characterize carbohydrate physiology in grapevine flowers, we examined inflorescence autotrophy in the 'Gewurztraminer' and 'Pinot noir' cultivars, which differ in sensitivity to flower abscission. In both cultivars, positive net photosynthesis occurred in inflorescences. The rate of photosynthesis gradually decreased throughout flower development and there was no net carbon assimilation at fruit set. The rate of photosynthesis was positively correlated with chlorophyll concentration but not to stomatal conductance. Throughout flower development, the internal CO2 concentration increased in inflorescence tissues, suggesting that assimilates are also formed through refixation of respiratory CO2 by the phosphoenolpyruvate carboxylase (PEPC) pathway. Significant differences between the two cultivars were recorded during meiosis, when photosynthesis was higher in 'Gewurztraminer'. We conclude that the inflorescence of grapevine contributes to its own carbon nutrition by photosynthesizing throughout flower development. Moreover, the differential patterns of photosynthesis in the inflorescences of 'Gewurztraminer' and 'Pinot noir' might account for their differing fertilization rates and sensitivity to flower abscission.

Carbohydrate Metabolism↗