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

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

Journal Article↗

Low humidity can cause uneven photosynthesis in olive (Olea europea L.) leaves.

We examined the photosynthetic responses of olive (Olea europea L.) leaves exposed to either (a) two hours of high leaf-to-air vapor pressure difference (vpd) or (b) four 30-min cycles of high vpd separated by 15-min periods of recovery at low vpd. Neither treatment affected photosynthesis when vpd was less than 3.0 kPa. Photosynthesis by mature leaves was also insensitive to higher vpd, but photosynthesis of young leaves was reduced by both treatments at a vpd higher than 3.2 kPa. This effect of vpd was much smaller under high intercellular CO(2) pressure. Autoradiograms showed that under a vpd of 3.2 kPa, mature leaves photosynthesized uniformly, but patches of reduced CO(2) fixation occurred in the distal part of young leaves. We conclude that heterogeneities in photosynthesis along the length of the leaf caused the apparent reduction of photosynthesis in our experiments. This pattern of patchy photosynthesis was different from that observed in mesophytic herbs, but the effect on gas exchange analysis was the same. In this case, apparent biochemical effects of low humidity on photosynthesis of young olive leaves are likely an artifact.

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Effects of O2 and CO2 on Nonsteady-State Photosynthesis (Further Evidence for Ribulose-1,5-Bisphosphate Carboxylase/Oxygenase Limitation).

The effects of CO2 and O2 on nonsteady-state photosynthesis following an increase in photosynthetic photon flux density (PPFD) were examined in Spinacia oleracea to investigate the hypotheses that (a) a slow exponential phase (the ribulose-1,5-bisphosphate carboxylase/oxygenase [Rubisco] phase) of nonsteady-state photosynthesis is primarily limited by Rubisco activity and (b) Rubisco activation involves two sequential, light-dependent processes as described in a previous study (I.E. Woodrow, K.A. Mott [1992] Plant Physiol 99: 298-303). Photosynthesis was found to be sensitive to O2 during the Rubisco phase in the approach of photosynthesis to steady state. Analyses of this sensitivity to O2 showed that the control coefficient for Rubisco was approximately equal to 1 during this phase, suggesting that Rubisco was the primary limitation to photosynthesis. O2 had almost no effect on the kinetics (described using a relaxation time, [tau] of the Rubisco phase for leaves starting in darkness or for leaves starting in low PPFD, but [tau] was substantially higher in the former case. CO2 was found to affect both the rate of photosynthesis and the magnitude of [tau] for the Rubisco phase. The [tau] value for the Rubisco phase was found to be negatively correlated with intercellular CO2 concentration (ci), and leaves starting in darkness had higher values of [tau] at any ci than leaves starting in low PPFD. The effects of CO2 and O2 on the Rubisco phase are consistent with the existence of two sequential, light-dependent processes in the activation of Rubisco if neither process is sensitive to O2 and only the second process is sensitive to CO2. The implications of the data for the mechanism of Rubisco activation and for the effects of stomatal conductance on nonsteady-state photosynthesis are discussed.

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C4 Photosynthesis (The CO2-Concentrating Mechanism and Photorespiration).

Despite previous reports of no apparent photorespiration in C4 plants based on measurements of gas exchange under 2 versus 21% O2 at varying [CO2], photosynthesis in maize (Zea mays) shows a dual response to varying [O2]. The maximum rate of photosynthesis in maize is dependent on O2 (approximately 10%). This O2 dependence is not related to stomatal conductance, because measurements were made at constant intercellular CO2 concentration (Ci); it may be linked to respiration or pseudocyclic electron flow. At a given Ci, increasing [O2] above 10% inhibits both the rate of photosynthesis, measured under high light, and the maximum quantum yield, measured under limiting light ([phi]CO2). The dual effect of O2 is masked if measurements are made under only 2 versus 21% O2. The inhibition of both photosynthesis and [phi]CO2 by O2 (measured above 10% O2) with decreasing Ci increases in a very similar manner, characteristically of O2 inhibition due to photorespiration. There is a sharp increase in O2 inhibition when the Ci decreases below 50 [mu]bar of CO2. Also, increasing temperature, which favors photorespiration, causes a decrease in [phi]CO2 under limiting CO2 and 40% O2. By comparing the degree of inhibition of photosynthesis in maize with that in the C3 species wheat (Triticum aestivum) at varying Ci, the effectiveness of C4 photosynthesis in concentrating CO2 in the leaf was evaluated. Under high light, 30[deg]C, and atmospheric levels of CO2 (340 [mu]bar), where there is little inhibition of photosynthesis in maize by O2, the estimated level of CO2 around ribulose-1,5-bisphosphate carboxylase/oxygenase (Rubisco) in the bundle sheath compartment was 900 [mu]bar, which is about 3 times higher than the value around Rubisco in mesophyll cells of wheat. A high [CO2] is maintained in the bundle sheath compartment in maize until Ci decreases below approximately 100 [mu]bar. The results from these gas exchange measurements indicate that photorespiration occurs in maize but that the rate is low unless the intercellular [CO2] is severely limited by stress.

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Interactions between Photosynthesis and Respiration in the Green Alga Chlamydomonas reinhardtii (Characterization of Light-Enhanced Dark Respiration).

The rate of respiratory O2 consumption by Chlamydomonas reinhardtii cell suspensions was greater after a period of photosynthesis than in the preceding dark period. This "light-enhanced dark respiration" (LEDR) was a function of both the duration of illumination and the photon fluence rate. Mass spectrometric measurements of gas exchange indicated that the rate of gross respiratory O2 consumption increased during photosynthesis, whereas gross respiratory CO2 production decreased in a photon fluence rate-dependent manner. The rate of postillumination O2 consumption provided a good measure of the O2 consumption rate in the light. LEDR was substantially decreased by the presence of 3-(3,4-dichlorophenyl)-1,1-dimethylurea or glycolaldehyde, suggesting that LEDR was photosynthesis-dependent. The onset of photosynthesis resulted in an increase in the cellular levels of phosphoglycerate, malate, and phosphoenolpyruvate, and a decrease in whole-cell ATP and citrate levels; all of these changes were rapidly reversed upon darkening. These results are consistent with a decrease in the rate of respiratory carbon flow during photosynthesis, whereas the increase in respiratory O2 consumption during photosynthesis may be mediated by the export of photogenerated reductant from the chloroplast. We suggest that photosynthesis interacts with respiration at more than one level, simultaneously decreasing the rate of respiratory carbon flow while increasing the rate of respiratory O2 consumption.

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Does Long-Term Elevation of CO2 Concentration Increase Photosynthesis in Forest Floor Vegetation? (Indiana Strawberry in a Maryland Forest).

As the partial pressure of CO2 (pCO2) in the atmosphere rises, photorespiratory loss of carbon in C3 photosynthesis will diminish and the net efficiency of light-limited photosynthetic carbon uptake should rise. We tested this expectation for Indiana strawberry (Duchesnea indica) growing on a Maryland forest floor. Open-top chambers were used to elevate the pCO2 of a forest floor habitat to 67 Pa and were paired with control chambers providing an ambient pCO2 of 38 Pa. After 3.5 years, D. indica leaves grown and measured in the elevated pCO2 showed a significantly greater maximum quantum efficiency of net photosynthesis (by 22%) and a lower light compensation point (by 42%) than leaves grown and measured in the control chambers. The quantum efficiency to minimize photorespiration, measured in 1% O2, was the same for controls and plants grown at elevated pCO2. This showed that the maximum efficiency of light-energy transduction into assimilated carbon was not altered by acclimation and that the increase in light-limited photosynthesis at elevated pCO2 was simply a function of the decrease in photorespiration. Acclimation did decrease the ribulose-1,5-bisphosphate carboxylase/oxygenase and light-harvesting chlorophyll protein content of the leaf by more than 30%. These changes were associated with a decreased capacity for light-saturated, but not light-limited, photosynthesis. Even so, leaves of D. indica grown and measured at elevated pCO2 showed greater light-saturated photosynthetic rates than leaves grown and measured at the current atmospheric pCO2. In situ measurements under natural forest floor lighting showed large increases in leaf photosynthesis at elevated pCO2, relative to controls, in both summer and fall. The increase in efficiency of light-limited photosynthesis with elevated pCO2 allowed positive net photosynthetic carbon uptake on days and at locations on the forest floor that light fluxes were insufficient for positive net photosynthesis in the current atmospheric pCO2.

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Effect of Temperature, CO(2) Concentration, and Light Intensity on Oxygen Inhibition of Photosynthesis in Wheat Leaves.

The effect of 21% O(2) and 3% O(2) on the CO(2) exchange of detached wheat leaves was measured in a closed system with an infrared carbon dioxide analyzer. Temperature was varied between 2 degrees and 43 degrees , CO(2) concentration between 0.000% and 0.050% and light intensity between 40 ft-c and 1000 ft-c. In most conditions, the apparent rate of photosynthesis was inhibited in 21% O(2) compared to 3% O(2). The degree of inhibition increased with increasing temperature and decreasing CO(2) concentration. Light intensity did not alter the effect of O(2) except at light intensities or CO(2) concentrations near the compensation point. At high CO(2) concentrations and low temperature, O(2) inhibition of apparent photosynthesis was absent. At 3% O(2), wheat resembled tropical grasses in possessing a high rate of photosynthesis, a temperature optimum for photosynthesis above 30 degrees , and a CO(2) compensation point of less than 0.0005% CO(2). The effect of O(2) on apparent photosynthesis could be ascribed to a combination of stimulation of CO(2) production during photosynthesis, and inhibition of photosynthesis itself.

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Nonstomatal inhibition of photosynthesis in sunflower at low leaf water potentials and high light intensities.

The inhibition of photosynthesis at low leaf water potentials was studied in soil-grown sunflower to determine the degree to which photosynthesis under high light was affected by stomatal and nonstomatal factors. Below leaf water potentials of -11 to -12 bars, rates of photosynthesis at high light intensities were insensitive to external concentrations of CO(2) between 200 and 400 microliters per liter. Photosynthesis also was largely insensitive to leaf temperature between 10 and 30 C. Changes in CO(2) concentration and temperature had negligible effect on leaf diffusive resistance. The lack of CO(2) and temperature response for both photosynthesis and leaf diffuse resistance indicates that rates of photosynthesis were not limited by either CO(2) diffusion or a photosynthetic enzyme. It was concluded that photosynthesis under high light was probably limited by reduced photochemical activity of the leaves at water potentials below -11 to -12 bars.

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pH Dependence of Photosynthesis and Photorespiration in Soybean Leaf Cells.

The effect of pH on the kinetics of photosynthesis, O(2) inhibition of photosynthesis, and photorespiration was examined with mesophyll cells isolated from soybean (Glycine max [L.] Merr.) leaves. At constant, subsaturating bicarbonate concentration (0.5 mm), O(2) inhibition of photosynthesis increased with increasing pH because high pH shifts the CO(2)-bicarbonate equilibrium toward bicarbonate, thereby reducing the CO(2) concentration. At constant, substrating CO(2) concentrations, cell photorespiration decreased with increasing pH. This was indicated by decreases in the CO(2) compensation concentration, O(2) inhibition of photosynthesis, and glycine synthesis. Km(CO(2)) values for isolated cell photosynthesis and in vitro ribulose-1, 5-diphosphate carboxylase activity decreased with increasing pH, while the Ki(O(2)) for both systems was similar at all pH values. The responses to pH of the corresponding kinetic constants of cell photosynthesis and in vitro RuDP carboxylase with respect to CO(2) and O(2) were identical. This provides additional evidence that the relative rates of photosynthesis and photorespiration in C(3) plants are determined by the kinetic properties of RuDP carboxylase.

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Oxygen inhibition of photosynthesis and stimulation of photorespiration in soybean leaf cells.

The occurrence of photorespiration in soybean (Glycine max [L.] Merr.) leaf cells was demonstrated by the presence of an O(2)-dependent CO(2) compensation concentration, a nonlinear time course for photosynthetic (14)CO(2) uptake at low CO(2) and high O(2) concentrations, and an O(2) stimulation of glycine and serine synthesis which was reversed by high CO(2) concentration. The compensation concentration was a linear function of O(2) concentration and increased as temperature increased. At atmospheric CO(2) concentration, 21% O(2) inhibited photosynthesis at 25 C by 27%. Oxygen inhibition of photosynthesis was competitive with respect to CO(2) and increased with increasing temperature. The Km (CO(2)) of photosynthesis was also temperature-dependent, increasing from 12 mum CO(2) at 15 C to 38 mum at 35 C. In contrast, the Ki (O(2)) was similar at all temperatures. Oxygen inhibition of photosynthesis was independent of irradiance except at 10 mm bicarbonate and 100% O(2), where inhibition decreased with increasing irradiance up to the point of light saturation of photosynthesis. Concomitant with increasing O(2) inhibition of photosynthesis was an increased incorporation of carbon into glycine and serine, intermediates of the photorespiratory pathway, and a decreased incorporation into starch. The effects of CO(2) and O(2) concentration and temperature on soybean cell photosynthesis and photorespiration provide further evidence that these processes are regulated by the kinetic properties of ribulose-1,5-diphosphate carboxylase with respect to CO(2) and O(2).

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Utilization of Exogenous Inorganic Carbon Species in Photosynthesis by Chlorella pyrenoidosa.

The nature of the inorganic carbon utilized during photosynthesis by Chlorella pyrenoidosa was investigated using three experimental techniques (open gas analysis system with "artificial leaf" or "aqueous" chambers and O(2) electrode system) to measure carbon assimilation. Photosynthesis was studied as a function of pH and CO(2) concentration. The CO(2) concentration was inadequate to meet the requirements of photosynthesis only when HCO(3) (-) was added at high pH. Under all other conditions, the low and constant K(m) (CO(2)), in contrast to the highly variable K(m) (HCO(3) (-)), suggested that CO(2) was the major species utilized.Higher rates of photosynthesis were observed under limiting CO(2) conditions above pH 7.5 but rates of hydration of CO(2) were not consistent with the view that the stimulation in photosynthesis was supported by HCO(3) (-). In the same pH region lower rates of photosynthesis were observed under saturating CO(2) conditions. These conflicting changes seemed not to be related to pH but to some as yet undetermined effect of bicarbonate. No support was obtained for the view that the quantum efficiency was different under conditions where CO(2) was assimilated compared to conditions where CO(2) and HCO(3) (-) could be assimilated, although at saturating CO(2), lower maximal rates of photosynthesis were observed in the latter conditions.

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Photosynthesis, leaf resistances, and ribulose-1,5-bisphosphate carboxylase degradation in senescing barley leaves.

The relationship between loss of ribulose-1,5-bisphosphate carboxylase (RuBPCase) and the decline in photosynthesis during the senescence of barley primary leaves was assessed. Loss of RuBPCase accounted for about 85% of the decrease in soluble protein. RuBPCase was highly correlated with in vitro RuBPCase activity (r = 0.95) and gross photosynthesis (r = 0.96). However, the rate of photosynthesis per milligram RuBPCase increased during the early stages of leaf senescence. The concentration of nonreducing sugars was negatively correlated (1% level) with photosynthesis. Free alpha-amino N, in contrast to nonreducing sugars, declined markedly during senescence. A decrease in chlorophyll and an increase in in vitro protease activity was observed, but these changes did not appear to be closely related to the decline in photosynthesis and RuBPCase. Mesophyll resistance increased at the same rate that photosynthesis and RuBPCase declined. Stomatal resistance increased more rapidly than mesophyll resistance and accounted for about 24% of the total increase in resistance to CO(2) diffusion. The concentration of CO(2) in the intercellular air spaces decreased during the last stage of senescence. Although loss of RuBPCase probably is the primary event responsible for the decline in photosynthesis during leaf senescence, other factors such as in vivo regulation and stomatal aperture must also be considered.

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The Regulation of Photosynthesis in Leaves of Field-Grown Spring Wheat (Triticum aestivum L., cv Albis) at Different Levels of Ozone in Ambient Air.

Wheat (Triticum aestivum L. cv Albis) was grown in open-top chambers in the field and fumigated daily with charcoal-filtered air (0.015 microliters per liter O(3)), nonfiltered air (0.03 microliters per liter O(3)), and air enriched with either 0.07 or 0.10 microliters per liter ozone (seasonal 8 hour/day [9 am-5 pm] mean ozone concentration from June 1 until July 10, 1987). Photosynthetic (14)CO(2) uptake was measured in situ. Net photosynthesis, dark respiration, and CO(2) compensation concentration at 2 and 21% O(2) were measured in the laboratory. Leaf segments were freeze-clamped in situ for the determination of the steady state levels of ribulose 1,5-bisphosphate, 3-phosphoglycerate, triose-phosphate, ATP, ADP, AMP, and activity of ribulose, 1,5-bisphosphate carboxylase/oxygenase. Photosynthesis of flag leaves was highest in filtered air and decreased in response to increasing mean ozone concentration. CO(2) compensation concentration and the ratio of dark respiration to net photosynthesis increased with ozone concentration. The decrease in photosynthesis was associated with a decrease in chlorophyll, soluble protein, ribulose bisphosphate carboxylase/oxygenase activity, ribulose bisphosphate, and adenylates. No decrease was found for triose-phosphate and 3-phosphoglycerate. The ratio of ATP to ADP and of triosephosphate to 3-phosphoglycerate were increased suggesting that photosynthesis was limited by pentose phosphate reductive cycle activity. No limitation occurred due to decreased access of CO(2) to photosynthetic cells since the decrease in stomatal conductance with increasing ozone concentration did not account for the decrease in photosynthesis. Ozonestressed leaves showed an increased degree of activation of ribulose bisphosphate carboxylase/oxygenase and a decreased ratio of ribulose bisphosphate to initial activity of ribulose bisphosphate carboxylase/oxygenase. Nevertheless, it is suggested that photosynthesis in ozone stressed leaves is limited by ribulose bisphosphate carboxylation possibly due to an effect of ozone on the catalysis by ribulose bisphosphate carboxylase/oxygenase.

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Photosynthesis, Carbohydrate Metabolism, and Export in Beta vulgaris L. and Phaseolus vulgaris L. during Square and Sinusoidal Light Regimes.

Rates of photosynthesis, sucrose synthesis, starch accumulation and degradation were measured in sugar beet (Beta vulgaris L.) and bean (Phaseolus vulgaris L.) plants under a square-wave light regime and under a sinusoidal regime that simulated the natural daylight period. Photosynthesis rate increased in a measured manner in direct proportion to the increasing light level. In contrast to this close correspondence between photosynthesis and light, a lag in photosynthesis rate was seen during the initial hour under square-wave illumination. The leaf appeared to be responding to limits set by carbon metabolism rather than by gas exchange or light reactions. Under the sinusoidal regime starch degradation occurred during the first and last 2 hours of the photoperiod, likely in response to photosynthesis rate rather than directly to light level. Starch broke down when photosynthesis was below a threshold rate and accumulated above this rate. Under square-wave illumination, accumulation of starch did not begin until irradiance was at full level for an hour or more and photosynthesis was at or near its maximum. Under a sinusoidal light regime, sucrose synthesis rate comprised carbon that was newly fixed throughout the day plus that from starch degradation at the beginning and end of the day. Synthesis of sucrose from recently fixed carbon increased with increasing net carbon fixation rate while its formation from degradation of starch decreased correspondingly. The complementary sources of carbon maintained a relatively steady rate of sucrose synthesis under the changing daytime irradiance.

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Modeling size-dependent photosynthesis: light absorption and the allometric rule.

Microalgal photosynthesis can be predicted using empirical allometric or mechanistic bio-optic models. These two descriptions are usually considered independently. We compare the size scaling of photosynthesis predicted by these two models. Size scaling exponents for phytoplankton often deviate from the allometric 3/4 rule. This may be because the allometric model does not account for the size dependence of light absorption and its effect on the size scaling of photosynthesis. In contrast to the allometric model and experimental data, the bio-optic model predicts photosynthesis should be independent of cell size when intracellular pigment concentrations are low or inversely related to cell diameter. A composite of the allometric and bio-optic models is described and compared to laboratory data of light-limited nutrient-saturated diatom photosynthesis. The allo-bio-optic model provides a mechanistic explanation for the anomalous size scaling found in laboratory and field studies of microalgal photosynthesis and growth.

Eukaryota↗

Long-term temperature acclimation of photosynthesis in steady-state cultures of the polar diatom Fragilariopsis cylindrus.

Cultures of the obligate psychrophilic diatom Fragilariopsis cylindrus (Grunow) were grown for 4 months under steady-state conditions at -1 degrees C and +7 degrees C (50 micromol photons m(-2) s(-1)) prior to measurements in order to investigate long-term acclimation of photosynthesis to both temperatures. No differences in maximum intrinsic quantum yield of PS II (F(V)/F(M)) and relative electron transport rates could be detected at either temperature after 4 months of acclimation. Measurements of photosynthesis (relative electron transport rates) vs. irradiance (P vs. E curves) revealed similar values for relative light utilization efficiency (alpha = 0.57 at -1 degrees C, alpha = 0.60 at +7 degrees C) but higher values for irradiance levels at which photosynthesis saturates (E(K)) at -1 degrees C and, therefore, higher maximum photosynthesis (P(MAX) = 54 (relative units) at -1 degrees C, P(MAX) = 49 at +7 degrees C). Nonphotochemical quenching (NPQ) measurements at 385 mumol photons m(-2) s(-1) indicated higher (37%) NPQ for diatoms grown at -1 degrees C compared to +7 degrees C, which was possibly related to a 2-fold increase in the concentration of the pigment diatoxanthin and a 9-fold up-regulation of a gene encoding a fucoxanthin chlorophyll a,c-binding protein. Expression of the D1 protein encoding gene psbA was ca. 1.5-fold up-regulated at -1 degrees C, whereas expression levels of other genes from Photosystem II (psbC, psbU, psbO), as well as rbcL, the gene encoding the Rubisco large subunit were similar at both temperatures. However, a 2-fold up-regulation of a plastid glyceraldehyde-P dehydrogenase at -1 degrees C indicated enhanced Calvin cycle activity. This study revealed for the first time that a polar diatom could efficiently acclimate photosynthesis over a wide range of polar temperatures given enough time. Acclimation of photosynthesis at -1 degrees C was probably regulated similarly to high light acclimation.

Acclimatization↗

From genes to photosynthesis in Arabidopsis thaliana.

Although photosynthesis in higher plants is of cyanobacterial descent, it differs strikingly in organization and regulation from the prokaryotic process. Genomics, proteomics, and comparative genome analysis are now providing powerful new tools for the molecular dissection of photosynthesis in higher plants. Mutant screens and reverse genetics identify an increasing number of gene-function relationships that have a bearing on photosynthesis, revealing a marked interdependency between photosynthesis and other cellular processes. Photosynthesis-related functions are mostly located in the chloroplast, but can also be located in other compartments of the plant cell. The analysis by DNA-array hybridization of mRNA expression patterns both in the chloroplast and the nucleus, under various environmental conditions and/or in different genetic backgrounds that affect the function of the plastid, is rapidly improving our understanding of how photosynthesis is regulated, and it reveals that plastid-to-nucleus signaling plays a central role in its control.

Arabidopsis↗

Kranz anatomy is not essential for terrestrial C4 plant photosynthesis.

An important adaptation to CO2-limited photosynthesis in cyanobacteria, algae and some plants was development of CO2-concentrating mechanisms (CCM). Evolution of a CCM occurred many times in flowering plants, beginning at least 15-20 million years ago, in response to atmospheric CO2 reduction, climate change, geological trends, and evolutionary diversification of species. In plants, this is achieved through a biochemical inorganic carbon pump called C4 photosynthesis, discovered 35 years ago. C4 photosynthesis is advantageous when limitations on carbon acquisition are imposed by high temperature, drought and saline conditions. It has been thought that a specialized leaf anatomy, composed of two, distinctive photosynthetic cell types (Kranz anatomy), is required for C4 photosynthesis. We provide evidence that C4 photosynthesis can function within a single photosynthetic cell in terrestrial plants. Borszczowia aralocaspica (Chenopodiaceae) has the photosynthetic features of C4 plants, yet lacks Kranz anatomy. This species accomplishes C4 photosynthesis through spatial compartmentation of photosynthetic enzymes, and by separation of two types of chloroplasts and other organelles in distinct positions within the chlorenchyma cell cytoplasm.

Carbon Dioxide↗