Effect of panicle removal on photosynthetic acclimation under elevated CO<Subscript>2</Subscript> in rice

To examine the role of sink size on photosynthetic acclimation under elevated atmospheric CO₂ concentrations ([CO₂]), we tested the effects of panicle-removal (PR) treatment on photosynthesis in rice (Oryza sativa L.). Rice was grown at two [CO₂] levels (ambient and ambient + 200 μmol mol⁻¹) throughout the growing season, and at full-heading stage, at half the plants, a sink-limitation treatment was imposed by the removal of the panicles. The PR treatment alleviated the reduction of green leaf area, the contents of chlorophyll (Chl) and Rubisco after the full-heading stage, suggesting delay of senescence. Nonetheless, elevated [CO₂] decreased photosynthesis (measured at current [CO₂]) of plants exposed to the PR treatment. No significant [CO₂] × PR interaction on photosynthesis was observed. The decrease of photosynthesis by elevated [CO₂] of plants was associated with decreased leaf Rubisco content and N content. Leaf glucose content was increased by the PR treatment and also by elevated [CO₂]. In conclusion, a sink-limitation in rice improved N status in the leaves, but this did not prevent the photosynthetic down-regulation under elevated [CO₂].     

How is the relationship between the C4 cereal Sorghum bicolor and the C3 root hemi-parasites Striga hermonthica and Striga asiatica affected by elevated CO2?

The C₄ cereal Sorghum bicolor was grown under either ambient (350 μmol mol^?1) or elevated (700 μmol mol^?1) [CO₂] in either the presence or absence of the C₃ obligate root hemi-parasites Striga hermonthica or S. asiatica. Both uninfected and infected sorghum plants were taller and had greater biomass, photosynthetic rates, water-use efficiencies and leaf areas under elevated compared with ambient [CO₂]. There was no evidence of any downregula-tion of photosynthesis in sorghum grown at elevated [CO₂]. Biomass of infected sorghum was lower under both ambient and elevated [CO₂], and although infected plants were larger under elevated [CO₂] the relative impact of infection on host biomass was either the same (S. asiatica) or only slightly less (S. hermonthica) than under ambient [CO₂]. In contrast, biomass of S. hermonthica and S. asiatica per host was lower under elevated than ambient [CO₂], although rates of photosynthesis were higher at elevated [CO₂] and parasite stomatal conductance was not responsive to [CO₂]. Parasites emerged above-ground and flowered earlier under ambient compared with elevated [CO₂]. It appears that the mechanism(s) by which the parasites affect host growth is (are) relatively insensitive to increased atmospheric [CO₂], although the parasites themselves were adversely affected by growth at elevated [CO₂].     

Acclimation of photosynthesis in relation to Rubisco and non-structural carbohydrate contents and in situ carboxylase activity in Scirpus olneyi grown at elevated CO2 in the field

Stands of Scirpus olneyi, a native saltmarsh sedge with C₃ photosynthesis, had been exposed to normal ambient and elevated atmospheric CO₂ concentrations (C_a) in their native habitat since 1987. The objective of this investigation was to characterize the acclimation of photosynthesis of Scirpus olneyi stems, the photosynthesizing organs of this species, to long-term elevated C_a treatment in relation to the concentrations of Rubisco and non-structural carbohydrates. Measurements were made on intact stems in the Held under existing natural conditions and in the laboratory under controlled conditions on stems excised in the field early in the morning. Plants grown at elevated C_a had a significantly higher (30–59%) net CO₂ assimilation rate (A) than those grown at ambient C_a when measurements were performed on excised stems at the respective growth C_a. However, when measurements were made at normal ambient C_a, A was smaller (45–53%) in plants grown at elevated C_a than in those grown at ambient C_a. The reductions in A at normal ambient C_a, carboxylation efficiency and in situ carboxylase activity were caused by a decreased Rubisco concentration (30–58%) in plants grown at elevated C_a; these plants also contained less soluble protein (39–52%). The Rubisco content was 43 to 58% of soluble protein, and this relationship was not significantly altered by the growth CO₂ concentrations. The Rubisco activation state increased slightly, but the in situ carboxylase activity decreased substantially in plants grown at elevated C_a. When measurements were made on intact stems in the field, the elevated C_a treatment caused a greater stimulation of,A (100%) and a smaller reduction in carboxylation efficiency (which was not statistically significant) than when measurements were made on excised stems in the laboratory. The possible reasons for this arc discussed. Plants grown at elevated C_a contained more non-structural carbohydrates (25–53%) than those grown at ambient C_a. Plants grown at elevated C_a appear to have sufficient sink capacity to utilize the additional carbohydrates formed during photosynthesis. Overall, our results are in agreement with the hypothesis that elevated C_a leads to an increased carbohydrate concentration and the ensuing acclimation of the photo-synthetic apparatus in C₃ plants results in a reduction in the protein complement, especially Rubisco, which reduces the photosynthetic capacity in plants grown at elevated C_a, relative to plants grown at normal ambient C_a. Nevertheless, when compared at their respective growth C_a, Scirpus olneyi plants grown at elevated C_a in their native habitat maintained a substantially higher rate of photosynthesis than those grown at normal ambient C_a even after 8 years of growth at elevated C_a.     

Mechanisms underlying leaf photosynthetic acclimation to warming and elevated CO2 as inferred from least‐cost optimality theory

The mechanisms responsible for photosynthetic acclimation are not well understood, effectively limiting predictability under future conditions. Least‐cost optimality theory can be used to predict the acclimation of photosynthetic capacity based on the assumption that plants maximize carbon uptake while minimizing the associated costs. Here, we use this theory as a null model in combination with multiple datasets of C₃ plant photosynthetic traits to elucidate the mechanisms underlying photosynthetic acclimation to elevated temperature and carbon dioxide (CO₂). The model‐data comparison showed that leaves decrease the ratio of the maximum rate of electron transport to the maximum rate of Rubisco carboxylation (J_max/V_cmax) under higher temperatures. The comparison also indicated that resources used for Rubisco and electron transport are reduced under both elevated temperature and CO₂. Finally, our analysis suggested that plants underinvest in electron transport relative to carboxylation under elevated CO₂, limiting potential leaf‐level photosynthesis under future CO₂ concentrations. Altogether, our results show that acclimation to temperature and CO₂ is primarily related to resource conservation at the leaf level. Under future, warmer, high CO₂ conditions, plants are therefore likely to use less nutrients for leaf‐level photosynthesis, which may impact whole‐plant to ecosystem functioning.     

Photosynthetic downregulation in leaves of the Japanese white birch grown under elevated CO2 concentration does not change their temperature‐dependent susceptibility to photoinhibition

To determine the effects of elevated CO₂ concentration ([CO₂]) on the temperature‐dependent photosynthetic properties, we measured gas exchange and chlorophyll fluorescence at various leaf temperatures (15, 20, 25, 30, 35 and 40°C) in 1‐year‐old seedlings of the Japanese white birch (Betula platyphylla var. japonica), grown in a phytotron under natural daylight at two [CO₂] levels (ambient: 400 µmol mol^?1 and elevated: 800 µmol mol^?1) and limited N availability (90 mg N plant^?1). Plants grown under elevated [CO₂] exhibited photosynthetic downregulation, indicated by a decrease in the carboxylation capacity of Rubisco. At temperatures above 30°C, the net photosynthetic rates of elevated‐CO₂‐grown plants exceeded those grown under ambient [CO₂] when compared at their growth [CO₂]. Electron transport rates were significantly lower in elevated‐CO₂‐grown plants than ambient‐CO₂‐grown ones at temperatures below 25°C. However, no significant difference was observed in the fraction of excess light energy [(1 ? q_P)× F_v′/F_m′] between CO₂ treatments across the temperature range. The quantum yield of regulated non‐photochemical energy loss was significantly higher in elevated‐CO₂‐grown plants than ambient, when compared at their respective growth [CO₂] below 25°C. These results suggest that elevated‐CO₂‐induced downregulation might not exacerbate the temperature‐dependent susceptibility to photoinhibition, because reduced energy consumption by electron transport was compensated for by increased thermal energy dissipation at low temperatures.     

Influence of nitrogen on growth and photosynthesis of a C3 cereal, Oryza sativa, infected with the root hemiparasite Striga hermonthica

Striga hermonthica is a root hemiparasitic angiosperm nativeto the African semi-arid tropics. It is a major weed of C₄ cerealsbut locally it is also an important weed of the C₃ plant, rice[Oryza sativa). Infected rice plants produced 17% and 42% ofthe total biomass of uninfected plants when grown at two differentammonium nitrate concentrations, 1 and 3 mol m^–3, respectively.S. hermonthica prevented grain production at both concentrationsof nitrogen. At the lower concentration no heads were produced.At the higher concentration head weight was only 6% of uninfectedcontrols. S. hermonthica also altered the partitioning of drymatter between plant parts, such that shoot growth was reducedto a greater extent than root growth. As a consequence the root-to-shootratio of infected plants was approximately five times greaterthan that of uninfected control plants. Light saturated ratesof photosynthesis In infected plants were 56% and 70% of thoseof uninfected controls, at low and high nitrogen, respectively.Infection also led to lower values of stomatal conductance althoughthe substom-atal CO₂ concentration was unaffected. Analysisof the response of photosynthesis to substomatal CO₂ concentration(A/C_I curves) demonstrated that lower rates of photosynthesiscould not be solely attributed to lower stomatal conductances.Lower initial slopes and asymptotic rates suggest that bothcarboxylation and processes controlling regeneration of ribulose-1,5-bisphosphate are reduced by infection. The data are discussedwith respect to the influence of S. hermonthica on the growthand photosynthesis of C₄ hosts, where in contrast to the situationwith rice, nitrogen feeding results in a marked alleviationof the effects of the parasite on the host. Key words: Rice, Striga, growth, photosynthesis, nitrogen     

Variable photosynthetic acclimation in consecutive cohorts of Scots pine needles during 3 years of growth at elevated CO2 and elevated temperature

In this experiment, the photosynthetic acclimation of successive needle cohorts of Scots pine were studied during 3 years of growth at elevated CO₂ and temperature. Naturally regenerated Scots pine (Pinus sylvestris L.) trees were subjected to elevated CO₂ concentration (+CO₂, 700 p.p.m), elevated temperature (+T, ambient +2 to +6 °C) and to a combination of elevated CO₂ and temperature (+CO₂ + T) in closed‐top chambers, starting in August 1996. Trees growing in chambers with ambient CO₂ and ambient temperature served as controls (AmbC). Elevated CO₂ influenced the dark reactions more than the light reactions of photosynthesis, as in the 1996 and 1997 cohorts the carboxylation capacity of Rubisco was reduced in the first and second year of exposure, but there was no consistent change in chlorophyll fluorescence. Net photosynthesis measured at growth concentration of CO₂ was higher at +CO₂ than at AmbC on only one measuring occasion, was generally lower at +T and was not changed at +CO₂ + T. However, trees grown at +T tended to invest more nitrogen (N) in Rubisco, as Rubisco/chlorophyll and the proportion of the total needle N bound to Rubisco occasionally increased. The interaction of +CO₂ and +T on Rubisco was mostly negative; consequently, in the second and third year of the experiment the carboxylation capacity decreased at +CO₂ + T. In the 1996, 1997 and 1998 cohorts, the structural N concentration of needles was lower at +CO₂ than at AmbC. Elevated CO₂ and elevated temperature generally had a positive interaction on N concentration; consequently, N concentration in needles decreased less at +CO₂ + T than at +CO₂. At +CO₂ + T, the acclimation response of needles varied between years and was more pronounced in the 1‐year‐old needles of the 1997 cohort than in those of the 1998 cohort. Thus, acclimation was not always greater in 1‐year‐old needles than in current‐year needles. In the +CO₂ + T treatment, elevated temperature had a greater effect on acclimation of needles than elevated CO₂.     

Photosynthetic response of Tempranillo grapevine to climate change scenarios

Photosynthesis in C₃ plants is CO₂ limited and therefore any increase in Rubisco carboxylation substrate may increase net CO₂ fixation, unless plants experience acclimation or other limitations. These aspects are largely unexplored in grapevine. Photosynthesis analysis was used to assess the stomatal, mesophyll, photochemical and biochemical contributions to the decreasing photosynthesis observed in Tempranillo grapevines (Vitis vinifera) from veraison to ripeness, modulated by CO₂, temperature and water availability. Photosynthesis and photosystem II photochemistry decreased from veraison to ripeness. The elevated CO₂ and temperature increased photosynthesis, but transiently, in both well irrigated (WI) and water‐stressed plants. Photosynthetic rates were maxima 1 week after the start of elevated CO₂ and temperature treatments, but differences with treatments of ambient conditions disappeared with time. There were not marked changes in leaf water status, leaf chlorophyll or leaf protein that could limit photosynthesis at ripeness. Leaf total soluble sugars remained at ripeness as high as 2 weeks after the start of treatments. On the other hand, and as expected, CO₂ diffusional limitations impaired photosynthesis in grapevine plants grown under water scarcity, stomatal and mesophyll conductances to CO₂ decreased and in turn low chloroplastic CO₂ concentrations limited photosynthetic CO₂ fixation. In summary, photochemistry and photosynthesis from veraison to ripeness in Tempranillo grapevine were dominated by a developmental‐related decreasing trend that was only transiently influenced by elevated CO₂ concentrations.     

Effects of elevated CO2 and temperature on photosynthesis and Rubisco in rice and soybean

Rice (Oryza sativa L. cv. IR-72) and soybean (Glycine max L. Merr. cv. Bragg), which have been reported to differ in acclimation to elevated CO₂, were grown for a season in sunlight at ambient and twice-ambient [CO₂], and under daytime temperature regimes ranging from 28 to 40°C. The objectives of the study were to test whether CO₂ enrichment could compensate for adverse effects of high growth temperatures on photosynthesis, and whether these two C₃ species differed in this regard. Leaf photosynthetic assimilation rates (A) of both species, when measured at the growth [CO₂], were increased by CO₂ enrichment, but decreased by supraoptimal temperatures. However, CO₂ enrichment more than compensated for the temperature-induced decline in A. For soybean, this CO₂ enhancement of A increased in a linear manner by 32–95% with increasing growth temperatures from 28 to 40°C, whereas with rice the degree of enhancement was relatively constant at about 60%, from 32 to 38°C. Both elevated CO₂ and temperature exerted coarse control on the Rubisco protein content, but the two species differed in the degree of responsiveness. CO₂ enrichment and high growth temperatures reduced the Rubisco content of rice by 22 and 23%, respectively, but only by 8 and 17% for soybean. The maximum degree of Rubisco down-regulation appeared to be limited, as in rice the substantial individual effects of these two variables, when combined, were less than additive. Fine control of Rubisco activation was also influenced by both elevated [CO₂] and temperature. In rice, total activity and activation were reduced, but in soybean only activation was lowered. The apparent catalytic turnover rate (K_cat) of rice Rubisco was unaffected by these variables, but in soybean elevated [CO₂] and temperature increased the apparent K_cat by 8 and 22%, respectively. Post-sunset declines in Rubisco activities were accelerated by elevated [CO₂] in rice, but by high temperature in soybean, suggesting that [CO₂] and growth temperature influenced the metabolism of 2-carboxyarabinitol-1-phosphate, and that the effects might be species-specific. The greater capacity of soybean for CO₂ enhancement of A at supraoptimal temperatures was probably not due to changes in stomatal conductance, but may be partially attributed to less down-regulation of Rubisco by elevated [CO₂] in soybean than in rice. However, unidentified species differences in the temperature optimum for photosynthesis also appeared to be important. The responses of photosynthesis and Rubisco in rice and soybean suggest that among C₃ plants species-specific differences will be encountered as a result of future increases in global [CO₂] and air temperatures.     

The temporal and species dynamics of photosynthetic acclimation in flag leaves of rice (Oryza sativa) and wheat (Triticum aestivum) under elevated carbon dioxide

In this study, we tested for the temporal occurrence of photosynthetic acclimation to elevated [CO₂] in the flag leaf of two important cereal crops, rice and wheat. In order to characterize the temporal onset of acclimation and the basis for any observed decline in photosynthetic rate, we characterized net photosynthesis, g_s, g_m, C_i/C_a, C_i/C_c, V_cmax, J_max, cell wall thickness, content of Rubisco, cytochrome (Cyt) f, N, chlorophyll and carbohydrate, mRNA expression for rbcL and petA, activity for Rubisco, sucrose phosphate synthase (SPS) and sucrose synthase (SS) at full flag expansion, mid‐anthesis and the late grain‐filling stage. No acclimation was observed for either crop at full flag leaf expansion. However, at the mid‐anthesis stage, photosynthetic acclimation in rice was associated with RuBP carboxylation and regeneration limitations, while wheat only had the carboxylation limitation. By grain maturation, the decline of Rubisco content and activity had contributed to RuBP carboxylation limitation of photosynthesis in both crops at elevated [CO₂]; however, the sharp decrease of Rubisco enzyme activity played a more important role in wheat. Although an increase in non‐structural carbohydrates did occur during these later stages, it was not consistently associated with changes in SPS and SS or photosynthetic acclimation. Rather, over time elevated [CO₂] appeared to enhance the rate of N degradation and senescence so that by late‐grain fill, photosynthetic acclimation to elevated [CO₂] in the flag leaf of either species was complete. These data suggest that the basis for photosynthetic acclimation with elevated [CO₂] may be more closely associated with enhanced rates of senescence, and, as a consequence, may be temporally dynamic, with significant species variation.     

Acclimation of peanut (Arachis hypogaea L.) leaf photosynthesis to elevated growth CO2 and temperature

Peanut (Arachis hypogaea L. cv. Florunner) was grown from seed sowing to plant maturity under two daytime CO₂ concentrations ([CO₂]) of 360 μmol mol⁻¹ (ambient) and 720 μmol mol⁻¹ (elevated) and at two temperatures of 1.5 and 6.0 °C above ambient temperature. The objectives were to characterize peanut leaf photosynthesis responses to long-term elevated growth [CO₂] and temperature, and to assess whether elevated [CO₂] regulated peanut leaf photosynthetic capacity, in terms of activity and protein content of ribulose bisphosphate carboxylase-oxygenase (Rubisco), Rubisco photosynthetic efficiency, and carbohydrate metabolism. At both growth temperatures, leaves of plants grown under elevated [CO₂] had higher midday photosynthetic CO₂ exchange rate (CER), lower transpiration and stomatal conductance and higher water-use efficiency, compared to those of plants grown at ambient [CO₂]. Both activity and protein content of Rubisco, expressed on a leaf area basis, were reduced at elevated growth [CO₂]. Declines in Rubisco under elevated growth [CO₂] were 27–30% for initial activity, 5–12% for total activity, and 9–20% for protein content. Although Rubisco protein content and activity were down-regulated by elevated [CO₂], Rubisco photosynthetic efficiency, the ratio of midday light-saturated CER to Rubisco initial or total activity, of the elevated-[CO₂] plants was 1.3- to 1.9-fold greater than that of the ambient-[CO₂] plants at both growth temperatures. Leaf soluble sugars and starch of plants grown at elevated [CO₂] were 1.3- and 2-fold higher, respectively, than those of plants grown at ambient [CO₂]. Under elevated [CO₂], leaf soluble sugars and starch, however, were not affected by high growth temperature. In contrast, high temperature reduced leaf soluble sugars and starch of the ambient-[CO₂] plants. Activity of sucrose-P synthase, but not adenosine 5′-diphosphoglucose pyrophosphorylase, was up-regulated under elevated growth [CO₂]. Thus, in the absence of other environmental stresses, peanut leaf photosynthesis would perform well under rising atmospheric [CO₂] and temperature as predicted for this century.     

Striga hermonthica reduces photosynthesis in sorghum: the importance of stomatal limitations and a potential role for ABA?

We report the effects of the root hemiparasite Striga hermonthica (Del.) Benth. on the growth and photosynthesis of two cultivars of sorghum: CSH-1, a susceptible variety, and Ochuti, which shows some tolerance to S. hermonthica in the field. Within 4 d of parasite attachment to the host roots, infected plants of both cultivars were significantly shorter than uninfected controls. At 55 d, infected plants of both cultivars had significantly less shoot and root biomass, and significantly smaller leaf areas than uninfected controls. The dry weight of S. hermonthica attached to host roots was insufficient at this stage to explain the decreased growth in terms of a competing sink for carbon and nitrogen. Leaf chlorophyll and nitrogen per unit area were greater in infected plants of both cultivars compared with control plants. However, whereas photosynthesis and transpiration in young leaves of infected CSH-1 plants declined with time when compared with controls, the rates in infected Ochuti plants were similar to those in uninfected controls throughout the time course of observation. In both cultivars, a strong correlation was observed between the rate of photosynthesis and stomatal conductance during photosynthetic induction, but infection resulted in a much slower induction than in controls. In CSH-1 plants, both steady-state photosynthesis and stomatal conductance were lower than in controls, whereas in leaves of Ochuti steady-state photosynthesis and stomatal conductance eventually reached the same values as in the control leaves. Results from AlC_i analysis and also from determination of ¹³C isotope discrimination were consistent with a stomatal limitation to photosynthesis in the leaves of Striga-infected plants. The concentration of the plant growth regulator abscisic acid (ABA) was measured in the xylem sap of infected CSH-1 plants only, and was found to be twice that of uninfected plants. A possible role of ABA in determining host response to infection by S. hermonthica is discussed.     

Will rising CO2 protect plants from the midday sun? A study of photoinhibition of Quercus myrtifolia in a scrub‐oak community in two seasons

Over a large part of the photoperiod, light energy absorbed by upper canopy leaves saturates photosynthesis and exceeds the energetic requirements for light‐saturated linear electron flow through photosystem II (J_PSII), so that photoinhibition results. From a theoretical consideration of the response of light‐saturated photosynthesis to elevated atmospheric CO₂ partial pressure (pCO₂) it may be predicted that, where light‐saturated photosynthesis is Rubisco‐limited, an increase in pCO₂ will stimulate J_PSII. Therefore, the proportion of absorbed quanta dissipated photochemically will increase and the potential for photoinhibition of photosynthesis will decrease. This was tested by measuring modulated chlorophyll a fluorescence from Quercus myrtifolia Willd. growing in the field in open‐top chambers, at either current ambient or elevated (ambient + 35 Pa) pCO₂ on Merritt Island, Florida, USA. During spring and summer, light‐saturated photosynthesis at current ambient pCO₂ was Rubisco‐limited. Consistent with theoretical prediction, J_PSII was increased and photoinhibition decreased by elevated pCO₂ in spring. In the summer, when growth had largely ceased, an acclimatory decrease in the maximum Ribulose 1,5 bisphosphate saturated carboxylation capacity (V_{c max}) removed the stimulation of J_PSII seen in the spring, and photoinhibition was increased in elevated pCO₂. It is concluded that, for Q. myrtifolia growing in the field, the effects of elevated pCO₂ on J_PSII and photoinhibition will reflect seasonal differences in photosynthetic acclimation to elevated pCO₂ in a predictable manner.     

Does growth under elevated CO2 moderate photoacclimation in rice?

Acclimation of plant photosynthesis to light irradiance (photoacclimation) involves adjustments in levels of pigments and proteins and larger scale changes in leaf morphology. To investigate the impact of rising atmospheric CO₂ on crop physiology, we hypothesize that elevated CO₂ interacts with photoacclimation in rice (Oryza sativa). Rice was grown under high light (HL: 700 µmol m^?2 s^?1), low light (LL: 200 µmol m^?2 s^?1), ambient CO₂ (400 µl l^?1) and elevated CO₂ (1000 µl l^?1). Leaf six was measured throughout. Obscuring meristem tissue during development did not alter leaf thickness indicating that mature leaves are responsible for sensing light during photoacclimation. Elevated CO₂ raised growth chamber photosynthesis and increased tiller formation at both light levels, while it increased leaf length under LL but not under HL. Elevated CO₂ always resulted in increased leaf growth rate and tiller production. Changes in leaf thickness, leaf area, Rubisco content, stem and leaf starch, sucrose and fructose content were all dominated by irradiance and unaffected by CO₂. However, stomata responded differently; they were significantly smaller in LL grown plants compared to HL but this effect was significantly suppressed under elevated CO₂. Stomatal density was lower under LL, but this required elevated CO₂ and the magnitude was adaxial or abaxial surface‐dependent. We conclude that photoacclimation in rice involves a systemic signal. Furthermore, extra carbohydrate produced under elevated CO₂ is utilized in enhancing leaf and tiller growth and does not enhance or inhibit any feature of photoacclimation with the exception of stomatal morphology.     

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

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

The impact of elevated CO2 on growth and photosynthesis in Agrostis canina L. ssp. monteluccii adapted to contrasting atmospheric CO2 concentrations

 The aim of this study was to characterise growth and photosynthetic capacity in plants adapted to long-term contrasting atmospheric CO₂ concentrations (C _a). Seeds of Agrostis canina L. ssp. monteluccii were collected from a natural CO₂ transect in central-western Italy and plants grown in controlled environment chambers at both ambient and elevated CO₂ (350 and 700 μmol mol⁻¹) in nutrient-rich soil. Seasonal mean C _a at the source of the plant material ranged from 610 to 451 μmol CO₂ mol⁻¹, derived from C₄ leaf stable carbon isotope discrimination (δ¹³C). Under chamber conditions, CO₂ enrichment stimulated the growth of all populations. However, plants originating from elevated C _a exhibited higher initial relative growth rates (RGRs) irrespective of chamber CO₂ concentrations and a positive relationship was found between RGR and C _a at the seed source. Seed weight was positively correlated with C _a, but differences in seed weight were found to explain no more than 34% of the variation in RGRs at elevated CO₂. Longer-term experiments (over 98 days) on two populations originating from the extremes of the transect (451 and 610 μmol CO₂ mol⁻¹) indicated that differences in growth between populations were maintained when plants were grown at both 350 and 700 μmol CO₂ mol⁻¹. Analysis of leaf material revealed an increase in the cell wall fraction (CWF) in plants grown at elevated CO₂, with plants originating from high C _a exhibiting constitutively lower levels but a variable response in terms of the degree of lignification. In vivo gas exchange measurements revealed no significant differences in light and CO₂ saturated rates of photosynthesis and carboxylation efficiency between populations or with CO₂ treatment. Moreover, SDS-PAGE/ LISA quantification of leaf ribulose bisphosphate carboxylase/oxygenase (Rubisco) showed no difference in Rubisco content between populations or CO₂ treatments. These findings suggest that long-term adaptation to growth at elevated CO₂ may be associated with a potential for increased growth, but this does not appear to be linked with differences in the intrinsic capacity for photosynthesis. Received: 16 August 1996 / Accepted: 19 October 1996     

Striga hermonthica decreases photosynthesis in Zea mays through effects on leaf cell structure

Maize seedlings were grown in pots either with or without preconditionedseeds of the parasitic weed, Striga hermonthica. After between4 and 8 weeks, net photosynthesis in the leaves of maize plantsinfected with Striga decreased compared to leaves of uninfectedcontrol plants. The activities of four enzymes of photosyntheticmetabolism were, however, little affected by infection. A pulse-chaseexperiment using ¹⁴CO₂ showed that C₄ acids were the main earlyproducts of assimilation even when the rate of photosynthesiswas much decreased by infection, but more radio-activity appearedin glycine and serine than in leaves of healthy maize plants.Leaves of infected maize required longer to reach a steady rateof photosynthesis upon enclosure in a leaf chamber than leavesof uninfected plants after similar treatment. Electron microscopy of transverse sections of the leaves ofinfected maize indicated that the cell walls in the bundle sheathand vascular tissue were less robust than in leaves of healthyplants. The results suggest that infection with Striga causesan increase in the permeability of cell walls in the bundlesheath, leakage of CO₂ from the bundle sheath cells and decreasedeffectiveness of C₄ photosynthesis in host leaves. Key words: Zea mays, Striga hermonthica, photosynthesis, photorespiration, enzyme activity     

Photosynthetic capacity of loblolly pine (Pinus taeda L.) trees during the first year of carbon dioxide enrichment in a forest ecosystem

Our objective was to assess the photosynthetic responses of loblolly pine trees (Pinus taeda L.) during the first full growth season (1997) at the Brookhaven National Lab/Duke University Free Air CO₂ Enrichment (FACE) experiment. Gas exchange, fluorescence characteristics, and leaf biochemistry of ambient CO₂ (control) needles and ambient + 20 Pa CO₂ (elevated) needles were examined five times during the year. The enhancement of photosynthesis by elevated CO₂ in mature loblolly pine trees varied across the season and was influenced by abiotic and biotic factors. Photosynthetic enhancement by elevated CO₂ was strongly correlated with leaf temperature. The magnitude of photosynthetic enhancement was zero in March but was as great as 52% later in the season. In March, reduced sink demand and lower temperatures resulted in lower net photosynthesis, lower carboxylation rates and higher excess energy dissipation from the elevated CO₂ needles than from control needles. The greatest photosynthetic enhancement by CO₂ enrichment was observed in July during a period of high temperature and low precipitation, and in September during recovery from this period of low precipitation. In July, loblolly pine trees in the control rings exhibited lower net photosynthetic rates, lower maximum rates of photosynthesis at saturating CO₂ and light, lower values of carboxylation and electron transport rates (modelled from A–C_i curves), lower total Rubisco activity, and lower photochemical quenching of fluorescence in comparison to other measurement periods. During this period of low precipitation trees in the elevated CO₂ rings exhibited reduced net photosynthesis and photochemical quenching of fluorescence, but there was little effect on light- and CO₂-saturated rates of photosynthesis, modelled rates of carboxylation or electron transport, or Rubisco activity. These first-year data will be used to compare with similar measurements from subsequent years of the FACE experiment in order to determine whether photosynthetic acclimation to CO₂ occurs in these canopy loblolly pine trees growing in a forest ecosystem.     

Photosynthetic responses to CO2 enrichment of four hardwood species in a forest understory

We compared the CO₂- and light-dependence of photosynthesis of four tree species (Acer rubrum, Carya glabra, Cercis canadensis, Liquidambar styraciflua) growing in the understory of a loblolly pine plantation under ambient or ambient plus 200 μl l^–1 CO₂. Naturally-established saplings were fumigated with a free-air CO₂ enrichment system. Light-saturated photosynthetic rates were 159–190% greater for Ce. canadensis saplings grown and measured under elevated CO₂. This species had the greatest CO₂ stimulation of photosynthesis. Photosynthetic rates were only 59% greater for A. rubrum saplings under CO₂ enrichment and Ca. glabra and L. styraciflua had intermediate responses. Elevated CO₂ stimulated light-saturated photosynthesis more than the apparent quantum yield. The maximum rate of carboxylation of ribulose-1,5-bisphosphate carboxylase, estimated from gas-exchange measurements, was not consistently affected by growth in elevated CO₂. However, the maximum electron transport rate estimated from gas- exchange measurements and from chlorophyll fluorescence, when averaged across species and dates, was approximately 10% higher for saplings in elevated CO₂. The proportionately greater stimulation of light-saturated photosynthesis than the apparent quantum yield and elevated rates of maximum electron transport suggests that saplings growing under elevated CO₂ make more efficient use of sunflecks. The stimulation of light-saturated photosynthesis by CO₂ did not appear to correlate with shade-tolerance ranking of the individual species. However, the species with the greatest enhancement of photosynthesis, Ce. canadensis and L. styraciflua, also invested the greatest proportion of soluble protein in Rubisco. Environmental and endogenous factors affecting N partitioning may partially explain interspecific variation in the photosynthetic response to elevated CO₂. Received: 16 February 1999 / Accepted: 30 August 1999     

Down co-regulation of light absorption, photochemistry, and carboxylation in Fe-deficient plants growing in different environments

The regulation of photosynthesis through changes in light absorption, photochemistry, and carboxylation efficiency has been studied in plants grown in different environments. Iron deficiency was induced in sugar beet (Beta vulgaris L.) by growing plants hydroponically in controlled growth chambers in the absence of Fe in the nutrient solution. Pear (Pyrus communis L.) and peach (Prunus persica L. Batsch) trees were grown in field conditions on calcareous soils, in orchards with Fe deficiency-chlorosis. Gas exchange parameters were measured in situ with actual ambient conditions. Iron deficiency decreased photosynthetic and transpiration rates, instantaneous transpiration efficiencies and stomatal conductances, and increased sub-stomatal CO₂ concentrations in the three species investigated. Photosynthesis versus CO₂ sub-stomatal concentration response curves and chlorophyll fluorescence quenching analysis revealed a non-stomatal limitation of photosynthetic rates under Fe deficiency in the three species investigated. Light absorption, photosystem II, and Rubisco carboxylation efficiencies were down-regulated in response to Fe deficiency in a coordinated manner, optimizing the use of the remaining photosynthetic pigments, electron transport carriers, and Rubisco.