Contribution of Metabolites of Photosynthesis to Postillumination CO(2) Assimilation in Response to Lightflects

In the shade plant Alocasia macrorrhiza grown in low light, photosynthetic CO₂ assimilation during a 5 second lightfleck plus postillumination CO₂ assimilation can allow up to 60% more photosynthesis than that which occurs during 5 seconds of steady state light of the same intensity (RL Chazdon, RW Pearcy 1986 Oecologia. 69: 524-531). Metabolites of photosynthesis were measured to determine if the pool of ribulose 1,5-bisphosphate (RuBP) could account for all of the postillumination CO₂ assimilation following a lightfleck in Alocasia. It was found that the pool of triose-P was much larger than that of RuBP and could account for five times more postillumination CO₂ assimilation than could RuBP. The same trend was seen in the sun plant Phaseolus vulgaris when it was grown in the shade. In contrast, sun-grown Alocasia and Phasiolus did not have a large pool of triose-P relative to RuBP following a lightfleck. In sun plants, carbon may rapidly be converted to RuBP in the light whereas in shade plants there may be a restriction in the path between the triose-P and RuBP pools. It is hypothesized that in shade plants the buildup of triose-P rather than RuBP during the lightfleck prevents inhibition of electron transport which may otherwise occur because of competition for ATP between the two kinases of the photosynthetic carbon reduction cycle. Utilization of the triose-P for postillumination CO₂ fixation would require the capacity for significant postillumination ATP synthesis. The extensive grana stacking and large intrathylakoid space which accompanies the high level of chlorophyll in low-light-grown Alocasia could be an important contributing factor to postillumination ATP formation.     

Salinity and Nitrogen Effects on Photosynthesis, Ribulose-1,5-Bisphosphate Carboxylase and Metabolite Pool Sizes in Phaseolus vulgaris L

Salinity (100 millimolar NaCl) was found to reduce photosynthetic capacity independent of stomatal closure in Phaseolus vulgaris. This reduction was shown to be a consequence of a reduction in the efficiency of ribulose-1,5-bisphosphate (RuBP) carboxylase (RuBPCase) rather than a reduction in the leaf content of photosynthetic machinery. In control plants, photosynthesis became RuBP-limited at approximately 1.75 moles RuBP per mole 2-carboxyarabinitol bisphosphate binding sites. Salinization caused the RuBP pool size to reach this limiting value for CO₂ fixation at much lower values of intercellular CO₂. Plants grown at low nitrogen and ± NaCl became RuBP limited at similar RuBP pool sizes as the high nitrogen-grown plants. At limiting RuBP pool sizes and equal values of intercellular CO₂ photosynthetic capacity of salt-stressed plants was less than control plants. This effect of salinity on RuBPCase activity could not be explained by deactivation of the enzyme or inhibitor synthesis. Thus, salinity reduced photosynthetic capacity by reducing both the RuBP pool size by an effect on RuBP regeneration capacity and RuBPCase activity by an unknown mechanism when RuBP was limiting.     

Leaf phosphate status, photosynthesis, and carbon partitioning in sugar beet: I. Changes in growth, gas exchange, and calvin cycle enzymes

Sugar beets (Beta vulgaris L. cv F58-554H1) were cultured hydroponically for 2 weeks in growth chambers with two levels of orthophosphate (Pi) supplied in half strength Hoagland solution. Low-P plants were supplied with 1/20th of the Pi supplied to control plants. With low-P treatment, the acid soluble leaf phosphate and total leaf P decreased by about 88%. Low-P treatment had a much greater effect on leaf area than on photosynthesis. Low-P decreased total leaf area by 76%, dry weight per plant by 60%, and the rate of photosynthesis per area at light saturation by 35%. Low-P treatment significantly decreased the total extractable activity of phosphoglycerate kinase (by 18%) and NADP-glyceraldehyde-3-phosphate dehydrogenase (by 16%), but did not decrease the total activities of ribulose-1,5-bisphosphate (RuBP) carboxylase (RuBPCase) and ribulose-5-phosphate kinase. Low-P treatment decreased the initial activities of three rate-limiting Calvin cycle enzymes, but had no effect on the initial activity of RuBPCase. Furthermore, low-P treatment significantly increased the total extractable activities of fructose-1,6-bisphosphatase (by 61%), fructose-1,6-bisphosphate aldolase (by 53%), and transketolase (by 46%). The results suggest that low-P treatment affected photosynthetic rate through an effect on RuBP regeneration rather than through RuBPCase activity and that the changes in Calvin cycle enzymes with low-P resulted in an increased flow of carbon to starch.     

Influence of leaf age on photosynthesis, enzyme activity, and metabolite levels in wheat

The rate of photosynthesis under high light (1000 micromole quanta per square meter per second) and at 25°C was measured during development of the third leaf on wheat plants and compared with the activity of several photosynthetic enzymes and the level of metabolites. The rate of photosynthesis reached a maximum the 7th day after leaf emergence and declined thereafter. There was a high and significant correlation between the rate of photosynthesis per leaf area and the activities of the enzymes ribulose 5-phosphate kinase (r = 0.91), ribulose 1,5-bisphosphate (RuBP) carboxylase (r = 0.94), 3-phosphoglycerate (PGA) kinase (r = 0.82), and fructose 1,6-bisphosphatase (r = 0.80) per leaf area. There was not a significant correlation of photosynthesis rate with chlorophyll content. The rate of photosynthesis was strongly correlated with the level of PGA (r = 0.85) and inversely correlated with the level of triose phosphate (dihydroxyacetone phosphate and glyceraldehyde 3-phosphate) (r = 0.92). RuBP levels did not change much during leaf development; therefore photosynthesis rate was not correlated with the level of RuBP. The rate of photosynthesis was at a maximum when the ratio of PGA/triose phosphate was high, and when the ratio of RuBP/PGA was low. Although several enzymes change in parallel with leaf development, the metabolite changes suggest the greatest degree of control may be through RuBP carboxylase. The sucrose content of the leaf was highest under high rates of photosynthesis. There was no evidence that later in leaf development, photosynthesis (measured under high light and at 25°C) was limited by utilization of photosynthate.     

Light adaptation/acclimation of photosynthesis and the regulation of ribulose-1,5-bisphosphate carboxylase activity in sun and shade plants

The consequences of light adaptation and acclimation of photosynthesis on photosynthetic nitrogen use efficiency (NUE), particularly as it relates to the efficiency of ribulose-1,5-bisphosphate carboxylase (Rubisco) use in photosynthetic CO₂ assimilation, was studied in the sun species Glycine max and the shade species Alocasia macrorrhiza. Both G. max and A. macrorrhiza were found to possess the capacity for light acclimation of CO₂ assimilation, but over distinctly different ranges of photon flux density (PFD). For each species, light acclimation of photosynthesis had little effect on the rate of photosynthesis per unit Rubisco protein or the light response of Rubisco carbamylation and CA 1P metabolism. In contrast, photosynthesis per unit Rubisco protein was significantly higher in G. max than in A. macrorrhiza, due in part to a lower total (fully carbamylated) molar activity (activity per unit enzyme) of A. macrorrhiza Rubisco than that of G. max. Comparison of the light response of Rubisco regulatory mechanisms between G. max and A. macrorrhiza indicated some degree of adaptation, such that carbamylation was higher and CA 1P levels lower at lower PFDs in the shade species than the sun species. However, this adjustment was not sufficient for Rubisco in low light grown A. macrorrhiza to be fully active at the growth PFD. Photosynthesis in A. macrorrhiza appeared to become RuBP regeneration-limited at lower PFDs than G. max, and this was probably the determinant of the light saturated rate of photosynthesis in the shade species. The low efficiency of Rubisco use in A. macrorrhiza was a major contributing factor to its five- to sixfold lower photosynthetic NUE than G. max. Shade species such as A. macrorrhiza appear to make far from maximal use of Rubisco protein N.     

Modelling Optimal Temperature Acclimation of the Photosynthetic Apparatus in C3Plants with Respect to Nitrogen Use

A new hypothesis for temperature acclimation by the photosyntheticapparatus is presented. An optimization model is developed toexamined effects of changes in the organization of photosyntheticcomponents on leaf photosynthesis under various growth temperatureswhere the photosynthetic apparatus is not damaged. In this model,photosynthetic rate is limited either by the capacity of ribulosebisphosphate carboxylase (RuBPCase) to consume ribulose bisphosphate(RuBP), or by the capacity of RuBP regeneration. For temperaturedependence of the RuBPCase activity, data fromSpinacia oleraceaL.,which have a temperature optimum of 30 °C, are used. Fortemperature dependence of the capacity of RuBP regeneration,two contrasting curves that have temperature optima of 30 °C(Eucalyptus paucifloraSieb. ex Spreng) and 40 °C (LarreadivaricataCav.) are applied. The temperature dependence of eachprocess is fixed for respective species, but the rate of eachprocess varies with changes in the amounts of components. Thecost of proteins, in terms of nitrogen, required to carry outeach process is calculated when nitrogen is partitioned differentlyamong photosynthetic components. The optimal nitrogen partitioningthat maximizes daily photosynthesis at a given temperature isobtained. The predicted temperature optimum of the photosyntheticrate inLarrea divaricataexhibits large shifts with changes intarget temperature, while shifts are negligible inEucalyptuspauciflora. It is suggested that the shift in temperature optimumof photosynthetic rate is large when the temperature dependencesof the capacities of RuBPCase and RuBP regeneration differ fromeach other.Copyright 1997 Annals of Botany Company Optimization model; nitrogen use efficiency; photosynthetic acclimation; temperature dependence     

Variation in photosynthetic electron transport capacity in vivo and its effects on the light modulation of ribulose bisphosphate carboxylase

Photosynthetic electron transport capacity was varied in vivo in sugar beets using iron deficiency, and its effects on the light modulation of ribulose bisphosphate carboxylase (RuBPCase) studied. Three treatment groups corresponding to decreasing amounts of thylakoids per leaf area were examined: iron sufficient (control), moderately iron-stressed, and severely iron-stressed. Reduction in electron transport capacity in vivo was correlated with a substantial decrease in the level of RuBPCase activation, even at saturating irradiances. These results indicate a direct relationship between RuBPCase activation and photosynthetic electron transport. In addition, our data suggest that the activation of RuBPCase could not solely account for the increases in the photosynthetic rate at high irradiances; RuBPCase reached maximal activation at irradiances well below light saturation for net photosynthesis.Abbreviations Chl chlorophyll - FeCN ferricyanide - FBPase fructose 1,6-bisphosphatase - RuBP ribulose 1,5-bisphosphate - RuBPCase ribulose 1,5-bisphosphate carboxylase - SBPase sedoheptulose 1,7-bisphosphatase     

An improved dynamic model of photosynthesis for estimation of carbon gain in sunfleck light regimes

A dynamic model of leaf photosynthesis for C₃ plants has been developed for examination of the role of the dynamic properties of the photosynthetic apparatus in regulating CO₂ assimilation in variable light regimes. The model is modified from the Farquhar-von Caemmerer-Berry model by explicitly including metabolite pools and the effects of light activation and deactivation of Calvin cycle enzymes. It is coupled to a dynamic stomatal conductance model, with the assimilation rate at any time being determined by the joint effects of the dynamic biochemical model and the stomatal conductance model on the intercellular CO₂ pressure. When parametrized for each species, the model was shown to exhibit responses to step changes in photon flux density that agreed closely with the observed responses for both the understory plant Alocasia macrorrhiza and the crop plant Glycine max. Comparisons of measured and simulated photosynthesis under simulated light regimes having natural patterns of lightfleck frequencies and durations showed that the simulated total for Alocasia was within ±4% of the measured total assimilation, but that both were 12–50% less than the predictions from a steady–state solution of the model. Agreement was within ±10% for Glycine max, and only small differences were apparent between the dynamic and steady–state predictions. The model may therefore be parametrized for quite different species, and is shown to reflect more accurately the dynamics of photosynthesis than earlier dynamic models.     

Photosynthesis and Ribulose 1,5-Bisphosphate Concentrations in Intact Leaves of Xanthium strumarium L

The interacting effects of the rate of ribulose 1,5-bisphosphate (RuBP) regeneration and the rate of RuBP utilization as influenced by the amount and activation of RuBP carboxylase on photosynthesis and RuBP concentrations were resolved in experiments which examined the kinetics of the response of photosynthesis and RuBP concentrations after step changes from a rate-saturating to a rate-limiting light intensity in Xanthium strumarium. Because RuBP carboxylase requires several minutes to deactivate in vivo, it was possible to observe the effect of reducing the rate of RuBP regeneration on the RuBP concentration at constant enzyme activation state by sampling very soon after reducing the light intensity. Samples taken over longer time periods showed the effect of changes in enzyme activation at constant RuBP regeneration rate on RuBP concentration and photosynthetic rate. Within 15 s of lowering the light intensity from 1500 to 600 microEinsteins per square meter per second the RuBP concentration in the leaves dropped below the enzyme active site concentration, indicating that RuBP regeneration rate was limiting for photosynthesis. After longer intervals of time, the RuBP concentration in the leaf increased as the RuBP carboxylase assumed a new steady state activation level. No change in the rate of photosynthesis was observed during the interval that RuBP concentration increased. It is concluded that the rate of photosynthesis at the lower light intensity was limited by the rate of RuBP regeneration and that parallel changes in the activation of RuBP carboxylase occurred such that concentrations of RuBP at steady state were not altered by changes in light intensity.     

Photosynthesis and Ribulose 1,5-Bisphosphate Carboxylase in Rice Leaves: Changes in Photosynthesis and Enzymes Involved in Carbon Assimilation from Leaf Development through Senescence

Changes in photosynthesis and the ribulose 1,5-bisphosphate (RuBP) carboxylase level were examined in the 12th leaf blades of rice (Oryza sativa L.) grown under different N levels. Photosynthesis was determined using an open infrared gas analysis system. The level of RuBP carboxylase was measured by rocket immunoelectrophoresis. These changes were followed with respect to changes in the activities of RuBP carboxylase, ribulose 5-phosphate kinase, NADP-glyceraldehyde 3-phosphate dehydrogenase, and 3-phosphoglyceric acid kinase.

RuBP carboxylase activity was highly correlated with the net rate of photosynthesis (r = 0.968). Although high correlations between the activities of other enzymes and photosynthesis were also found, the activity per leaf of RuBP carboxylase was much lower than those of other enzymes throughout the leaf life. The specific activity of RuBP carboxylase on a milligram of the enzyme protein basis remained fairly constant (1.16 ± 0.07 micromoles of CO₂ per minute per milligram at 25°C) throughout the experimental period.

Kinetic parameters related to CO₂ fixation were examined using the purified carboxylase. The K_m(CO₂) and V_max values were 12 micromolar and 1.45 micromoles of CO₂ per minute per milligram, respectively (pH 8.2 and 25°C). The in vitro specific activity calculated at the atomospheric CO₂ level from the parameters was comparable to the in situ true photosynthetic rate per milligram of the carboxylase throughout the leaf life.

The results indicated that the level of RuBP carboxylase protein can be a limiting factor in photosynthesis throughout the life span of the leaf.

     

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 α-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 CO₂ diffusion. The concentration of CO₂ 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.     

Diaheliotropic leaf movement enhances leaf photosynthetic capacity and photosynthetic light and nitrogen use efficiency via optimising nitrogen partitioning among photosynthetic components in cotton (Gossypium hirsutum L.)

• Phototropic leaf movement of plants is an effective mechanism for adapting to light conditions. Light is the major driver of plant photosynthesis. Leaf N is also an important limiting factor on leaf photosynthetic potential. Cotton (Gossypium hirsutum L.) exhibits diaheliotropic leaf movement. Here, we compared the long‐term photosynthetic acclimation of fixed leaves (restrained) and free leaves (allowed free movement) in cotton.
• The fixed leaves and free leaves were used for determination of PAR, leaf chlorophyll concentration, leaf N content and leaf gas exchange. The measurements were conducted under clear sky conditions at 0, 7, 15 and 30 days after treatment (DAT).
• The results showed that leaf N allocation and partitioning among different components of the photosynthetic apparatus were significantly affected by diaheliotropic leaf movement. Diaheliotropic leaf movement significantly increased light interception per unit leaf area, which in turn affected leaf mass per area (LMA), leaf N content (N_A) and leaf N allocation to photosynthesis (N_P). In addition, cotton leaves optimised leaf N allocation to the photosynthetic apparatus by adjusting leaf mass per area and N_A in response to optimal light interception.
• In the presence of diaheliotropic leaf movement, cotton leaves optimised their structural tissue and photosynthetic characteristics, such as LMA, N_A and leaf N allocation to photosynthesis, so that leaf photosynthetic capacity was maximised to improve the photosynthetic use efficiency of light and N under high light conditions.

     

Regulation of Ribulose-1,5-Bisphosphate Carboxylase Activity in Alocasia macrorrhiza in Response to Step Changes in Irradiance

The regulation of ribulose-1,5-bisphosphate (RuBP) carboxylase (Rubisco) activity and pool sizes of RuBP and P-glycerate were examined in the tropical understory species Alocasia macrorrhiza following step changes in photon flux density (PFD). Previous gas exchange analysis of this species following a step increase in PFD from 10 to 500 micromoles quanta per square meter per second suggested that the increase in photosynthetic rate was limited by the rate of increase of Rubisco activity for the first 5 to 10 minutes. We demonstrate here that the increase in photosynthetic rate was correlated with an increase in both the activation state of Rubisco and the total k_cat (fully activated specific activity) of the enzyme. Evidence presented here suggests that a change in the pool size of the naturally occurring tight binding inhibitor of Rubisco activity, 2-carboxyarabinitol 1-phosphate, was responsible for the PFD-dependent change in the total k_cat of the enzyme. RuBP pool size transiently increased after the increase in PFD, indicating that photosynthesis was limited by the capacity for carboxylation. After 5 to 10 minutes, RuBP pool size was again similar to the pool size at low PFD, presumably because of the increased activity of Rubisco. Following a step decrease in PFD from 500 to 10 micromoles quanta per square meter per second, Rubisco activity declined but at a much slower rate than it had increased in response to a step increase in PFD. This slower rate of activity decline than increase was apparently due to the slower rate of 2-carboxyarabinitol 1-phosphate synthesis than degradation and, to a lesser degree, to slower deactivation than activation. RuBP pool size initially declined following the decrease in PFD, indicating that RuBP regeneration was limiting photosynthesis. As Rubisco activity decreased, RuBP slowly increased to its original level at high PFD. The slow rate of activity loss by Rubisco in this species suggests a biochemical basis for the increased efficiency for CO₂ assimilation of successive lightfleck use by species such as A. macrorrhiza.     

Drought Stress and Elevated CO(2) Effects on Soybean Ribulose Bisphosphate Carboxylase Activity and Canopy Photosynthetic Rates

Soybean (Glycine max [L.] cv Bragg) was grown at 330 or 660 microliters CO₂ per liter in outdoor, controlled-environment chambers. When the plants were 50 days old, drought stress was imposed by gradually reducing irrigation each evening so that plants wilted earlier each succeeding day. On the ninth day, as the pots ran out of water CO₂ exchange rate (CER) decreased rapidly to near zero for the remainder of the day. Both CO₂-enrichment and drought stress reduced the total (HCO₃⁻/Mg²⁺-activated) extractable ribulose-1,5-bisphosphate carboxylase (RuBPCase) activity, as expressed on a chlorophyll basis. In addition, drought stress when canopy CER values and leaf water potentials were lowest, reduced the initial (nonactivated) RuBPCase activity by 50% compared to the corresponding unstressed treatments. This suggests that moderate to severe drought stress reduces the in vivo activation state of RuBPCase, as well as lowers the total activity. It is hypothesized that stromal acidification under drought stress causes the lowered initial RuBPCase activities. The K_m(CO₂) values of activated RuBPCase from stressed and unstressed plants were similar; 15.0 and 12.6 micromolar, respectively. RuBP levels were 10 to 30% lower in drought stressed as compared to unstressed treatments. However, RuBP levels increased from near zero at night to around 150 to 200 nanomoles per milligram chlorophyll during the day, even as water potentials and canopy CERs decreased. This suggests that the rapid decline in canopy CER cannot be attributed to drought stress induced limitations in the RuBP regeneration capability. Thus, in soybean leaves, a nonstomatal limitation of leaf photosynthesis under drought stress conditions appears due, in part, to a reduction of the in vivo activity of RuBPCase. Because initial RuBPCase activities were not reduced as much as canopy CER values, this enzymic effect does not explain entirely the response of soybean photosynthesis to drought stress.     

Effects of Light and Elevated Atmospheric CO(2) on the Ribulose Bisphosphate Carboxylase Activity and Ribulose Bisphosphate Level of Soybean Leaves

Soybean (Glycine max L. Merr. cv Bragg) was grown throughout its life cycle at 330, 450, and 800 microliters CO₂ per liter in outdoor controlled-environment chambers under solar irradiance. Leaf ribulose-1,5-bisphosphate carboxylase (RuBPCase) activities and ribulose-1,5-bisphosphate (RuBP) levels were measured at selected times after planting. Growth under the high CO₂ levels reduced the extractable RuBPCase activity by up to 22%, but increased the daytime RuBP levels by up to 20%.

Diurnal measurements of RuBPCase (expressed in micromoles CO₂ per milligram chlorophyll per hour) showed that the enzyme values were low (230) when sampled before sunrise, even when activated in vitro with saturating HCO₃⁻ and Mg²⁺, but increased to 590 during the day as the solar quantum irradiance (photosynthetically active radiation or PAR, in micromoles per square meter per second) rose to 600. The nonactivated RuBPCase values, which averaged 20% lower than the corresponding HCO₃⁻ and Mg²⁺-activated values, increased in a similar manner with increasing solar PAR. The per cent RuBPCase activation (the ratio of nonactivated to maximum-activated values) increased from 40% before dawn to 80% during the day. Leaf RuBP levels (expressed in nanomoles per milligram chlorophyll) were close to zero before sunrise but increased to a maximum of 220 as the solar PAR rose beyond 1200. In a chamber kept dark throughout the morning, leaf RuBPCase activities and RuBP levels remained at the predawn values. Upon removal of the cover at noon, the HCO₃⁻ and Mg²⁺-activated RuBPCase values and the RuBP levels rose to 465 and 122, respectively, after only 5 minutes of leaf exposure to solar PAR at 1500.

These results indicate that, in soybean leaves, light may exert a regulatory effect on extractable RuBPCase in addition to the well-established activation by CO₂ and Mg²⁺.

     

The Nitrogen Use Efficiency of C(3) and C(4) Plants: II. Leaf Nitrogen Effects on the Gas Exchange Characteristics of Chenopodium album (L.) and Amaranthus retroflexus (L.)

The effect of leaf nitrogen (N) on the photosynthetic capacity and the light and temperature response of photosynthesis was studied in the ecologically similar annuals Chenopodium album (C₃) and Amaranthus retroflexus (C₄). Photosynthesis was linearly dependent on leaf N per unit area (N_a) in both species. A. retroflexus exhibited a greater dependence of photosynthesis on N_a than C. album and at any given N_a, it had a greater light saturated photosynthesis rate than C. album. The difference between the species became larger as N_a increased. These results demonstrate a greater photosynthetic N use efficiency in A. retroflexus than C. album. However, at a given applied N level, C. album allocated more N to a unit of leaf area so that photosynthetic rates were similar in the two species. Leaf conductance to water vapor increased linearly with N_a in both species, but at a given photosynthetic rate, leaf conductance was higher in C. album. Thus, A. retroflexus had a greater water use efficiency than C. album. Water use efficiency was independent of leaf N in C. album, but declined with decreasing N in A. retroflexus.     

Branching, nitrogen distribution, and carbon gain in solitary plants of an annual herb, Xanthium canadense

The theory of optimal nitrogen (N) distribution predicts that the carbon gain of plants will be maximised when leaves of higher irradiance have higher N content per area (N _area). Most previous studies have examined optimal N distribution without explicitly considering the branching status of plants. I investigated light environment, N distribution and photosynthetic traits of individual leaves of an herbaceous species, Xanthium canadense. X. canadense was grown solitary under high (HN) and low nutrients (LN). Light availability, leaf mass per unit area and N _area were measured for all leaves within plants. Daily photosynthesis of the plants of actual N distribution was compared with those of optimal and constant N distribution. Branch production was facilitated in HN but not in LN plants. N _area was correlated more with leaf order than with leaf light environment. Although N was more limited and the light environment was less heterogeneous within crowns in LN than in HN plants, leaf N distribution was closer to optimal in the latter. These results suggest that leaf N distribution was not optimised in solitary plants of X. canadense. Because this species often regenerates in a dense stand, leaf N distribution might be selected to maximise carbon gain only in such a stand. Leaf N distribution might thus be constrained by the regeneration strategy of the species.     

Limiting Factors in Photosynthesis: VI. Regeneration of Ribulose 1,5-Bisphosphate Limits Photosynthesis at Low Photochemical Capacity

Earlier work (SE Taylor, N Terry [1984] Plant Physiol 75: 82-86) has shown that the rate of photosynthesis may be colimited by photosynthetic electron transport capacity, even at low intercellular CO₂ concentrations. Here we monitored leaf metabolites diurnally and the activities of key Calvin cycle enzymes in the leaves of three treatment groups of sugar beet (Beta vulgaris L.) plants representing three different in vivo photochemical capacities, i.e. Fe-sufficient (control) plants, moderately Fe-deficient, and severely Fe-deficient plants. The results show that the decrease in photosynthesis with Fe deficiency mediated reduction in photochemical capacity was through a reduction in ribulose 1,5-bisphosphate (RuBP) regeneration and not through a decrease in ribulose 1,5-bisphosphate carboxylase/oxygenase activity. Based on measurements of ATP and NADPH and triose phosphate/3-phosphoglycerate ratios in leaves, there was little evidence that photosynthesis and RuBP regeneration in Fe-deficient leaves were limited directly by the supply of ATP and NADPH. It appeared more likely that photochemical capacity influenced RuBP regeneration through modulation of enzymes in the photosynthetic carbon reduction cycle between fructose-6-phosphate and RuBP; in particular, the initial activity of ribulose-5-phosphate kinase was strongly diminished by Fe deficiency. Starch and sucrose levels changed independently of one another to some extent during the diurnal period (both increasing in the day and decreasing at night) but the average rates of starch or sucrose accumulation over the light period were each proportional to photochemical capacity and photosynthetic rate.     

Relationships between CO(2) Exchange Rates and Activities of Pyruvate,Pi Dikinase and Ribulose Bisphosphate Carboxylase, Chlorophyll Concentration, and Cell Volume in Maize Leaves

The relationships between CO₂-exchange rate (CER), DNA and chlorophyll (Chl) concentrations, pyruvate,Pi dikinase (PPDK) and ribulose bisphosphate carboxylase (RuBPCase) activities in ten maize (Zea mays L.) genotypes were investigated. The in vivo degrees of activation of PPDK and RuBPCase were estimated to make meaningful comparisons with CER. In leaves at a photosynthetic photon flux density (PPFD) of 720 micromoles per square meter per second, in vivo PPDK degree of activation was 80% of that of PPDK fully activated in vitro, whereas RuBPCase could not be further activated in vitro, suggesting that RuBPCase was fully activated in vivo. CER varied about 50% among the genotypes tested. Significant genetic differences were observed for the average weight of a cell (estimated by gram fresh weight per milligram DNA), but this character was not correlated with CER expressed on a fresh weight basis. CER was correlated with Chl concentration, and with estimates of the in vivo degree of activation of PPDK and RuBPCase. We concluded that in maize, CER is controlled by the metabolic components of photosynthesis rather than by membrane resistances to CO₂. If the latter factor were controlling CER, then smaller cells with higher amounts of exposed cell surface area per unit cell volume would have lower resistance to CO₂ diffusion, and therefore higher CER. When data were expressed on a DNA basis (proportional to a per cell basis), results indicated that larger cells (i.e. those with higher fresh weight per milligram DNA) have a higher content of Chl, and higher PPDK and RuBPCase activities, resulting in higher CER than in smaller cells.     

The relationship between carbon-dioxide-limited photosynthetic rate and ribulose-1,5-bisphosphate-carboxylase content in two nuclear-cytoplasm substitution lines of wheat,and the coordination of ribulose-bisphosphate-carboxylation and electron-transport capacities

Photosynthesis in two cultivars of Triticum aestivum was compared with photosynthesis in two lines having the same nuclear genomes but with cytoplasms derived from T. boeoticum. The in-vitro specific activity of ribulose-1,5-bisphosphate carboxylase (RuBPCase; EC 4.1.1.39) isolated from lines with T. boeoticum cytoplasm was only 71% of that of normal T. aestivum. By contrast, the RuBPCase activities calculated from the CO₂-assimilation rate at low partial pressures of CO₂, p(CO₂), were the same for all lines for a given RuBPCase content. This indicates that both types of RuBPCase have the same turnover numbers in-vivo of 27.5 mol CO₂·(mol enzyme)^–1·s^–1 (23°). The rate of CO₂ assimilation measured at normal p(CO₂), p _a =340 bar, and high irradiance could be quantitatively predicted from the amount of RuBPCase protein. The maximum rate of RuBP regeneration could also predict the rate of CO₂ assimilation at normal ambient conditions. Therefore, the maximum capacities for RuBP carboxylation and RuBP regeneration appear to be well-balanced for normal ambient conditions. As photosynthetic capacity declined with increasing leaf age, the capacities for RuBP carboxylation and RuBP regeneration declined in parallel.Abbreviations PAR photosynthetically active radiation - RuBP(Case) ribulose-1,5-bisphosphate (carboxylase)