C3 photosynthesis and high temperature acclimation of CAM in opuntia basilaris engelm. and bigel

Summary Carbon fixation by CAM at high night temperatures was examined in the stem succulent, Opuntia basilaris. Nighttime accumulation of titratable acids was uniformly high among plants growing naturally along an altitudinal temperature gradient in Death Valley, California during the hot summer period. Plants grown at high temperature regimes (40°/30°C) had rates of CAM and C₃ fixation similar to rates observed in plants maintained at a cool temperature (20°/10°C). C₃ fixation comprised 30% of the total carbon fixed by the potted, well watered plants. However, when pads were excised, C₃ fixation was suppressed while CAM continued unabated.     

Temperature effects on the carbon-isotope ratio of C3, C4 and crassulacean-acid-metabolism (CAM) plants

Planta - The temperature coefficient of the δ13C value over the temperature range 14 to 40° has been measured for plants with different photosynthetic pathways. C3 plants (6 species) had...     

The temperature response of C(3) and C(4) photosynthesis

We review the current understanding of the temperature responses of C(3) and C(4) photosynthesis across thermal ranges that do not harm the photosynthetic apparatus. In C(3) species, photosynthesis is classically considered to be limited by the capacities of ribulose 1.5-bisphosphate carboxylase/oxygenase (Rubisco), ribulose bisphosphate (RuBP) regeneration or P(i) regeneration. Using both theoretical and empirical evidence, we describe the temperature response of instantaneous net CO(2) assimilation rate (A) in terms of these limitations, and evaluate possible limitations on A at elevated temperatures arising from heat-induced lability of Rubisco activase. In C(3) plants, Rubisco capacity is the predominant limitation on A across a wide range of temperatures at low CO(2) (<300 microbar), while at elevated CO(2), the limitation shifts to P(i) regeneration capacity at suboptimal temperatures, and either electron transport capacity or Rubisco activase capacity at supraoptimal temperatures. In C(4) plants, Rubisco capacity limits A below 20 degrees C in chilling-tolerant species, but the control over A at elevated temperature remains uncertain. Acclimation of C(3) photosynthesis to suboptimal growth temperature is commonly associated with a disproportional enhancement of the P(i) regeneration capacity. Above the thermal optimum, acclimation of A to increasing growth temperature is associated with increased electron transport capacity and/or greater heat stability of Rubisco activase. In many C(4) species from warm habitats, acclimation to cooler growth conditions increases levels of Rubisco and C(4) cycle enzymes which then enhance A below the thermal optimum. By contrast, few C(4) species adapted to cooler habitats increase Rubisco content during acclimation to reduced growth temperature; as a result, A changes little at suboptimal temperatures. Global change is likely to cause a widespread shift in patterns of photosynthetic limitation in higher plants. Limitations in electron transport and Rubisco activase capacity should be more common in the warmer, high CO(2) conditions expected by the end of the century.     

Temperature and leaf osmotic potential as factors in the acclimation of photosynthesis to high temperature in desert plants

Seasonal changes in the high temperature limit for photosynthesis of desert winter annuals growing under natural conditions in Death Valley, California were studied using an assay based upon chlorophyll fluorescence. All species of this group were 6 to 9°C more tolerant of high temperature at the end of the growing season (May) than at its beginning (February). Over this same time period, the mean daily maximum air temperatures increased by 12°C. Laboratory studies have demonstrated that increases in thermal tolerance could be induced by increasing growth temperature alone. For plants growing under field conditions there was also a good correlation between the thermal tolerance of leaves and the osmotic potential of leaf water, indicating that increases in the concentrations of some small molecules might also confer increased thermal tolerance. Isolated chloroplast thylakoids subjected to increasing concentrations of sorbitol could be demonstrated to have increased thermal tolerance.     

High temperature acclimation of C4 photosynthesis is linked to changes in photosynthetic biochemistry

With average global temperatures predicted to increase over the next century, it is important to understand the extent and mechanisms of C4 photosynthetic acclimation to modest increases in growth temperature. To this end, we compared the photosynthetic responses of two C4 grasses (Panicum coloratum and Cenchrus ciliaris) and one C4 dicot (Flaveria bidentis) to growth at moderate (25/20 degrees C, day/night) or high (35/30 degrees C, day/night) temperatures. In all three C4 species, CO2 assimilation rates (A) underwent significant thermal acclimation, such that when compared at growth temperatures, A increased less than what would be expected given the strong response of A to short-term changes in leaf temperature. Thermal photosynthetic acclimation was further manifested by an increase in the temperature optima of A, and a decrease in leaf nitrogen content and leaf mass per area in the high- relative to the moderate-temperature-grown plants. Reduced photosynthetic capacity at the higher growth temperature was underpinned by selective changes in photosynthetic components. Plants grown at the higher temperature had lower amounts of ribulose-1,5-bisphosphate carboxylase/oxygenase and cytochrome f and activity of carbonic anhydrase. The activities of photosystem II (PSII) and phosphoenolpyruvate carboxylase were not affected by growth temperature. Chlorophyll fluorescence measurements of F. bidentis showed a corresponding decrease in the quantum yield of PSII (phi(PSII)) and an increase in non-photochemical quenching (phi(NPQ)). It is concluded that through these biochemical changes, C4 plants maintain the balance between the various photosynthetic components at each growth temperature, despite the differing temperature dependence of each process. As such, at higher temperatures photosynthetic nitrogen use efficiency increases more than A. Our results suggest C4 plants will show only modest changes in photosynthetic rates in response to changes in growth temperature, such as those expected within or between seasons, or the warming anticipated as a result of global climate change.     

Enzyme regulation in C4 photosynthesis: purification, properties, and activities of thioredoxins from C4 and C3 plants

Procedures are described for the purification to homogeneity of chloroplast thioredoxins f and m from leaves of corn (Zea mays, a C4 plant) and spinach (Spinacea oleracea, a C3 plant). The C3 and C4f thioredoxins were similar immunologically and biochemically, but differed in certain of their physiochemical properties. The f thioredoxins from the two species were capable of activating both NADP-malate dehydrogenase (EC 1.1.1.37) and fructose-1,6-bisphosphatase (EC 3.1.3.11) when tested in standard thioredoxin assays. Relative to its spinach counterpart, corn thioredoxin f showed a greater molecular mass (15.0-16.0 kDa vs 10.5 kDa), lower isoelectric point (ca. 5.2 vs 6.0), and lower ability to form a stable noncovalent complex with its target fructose bisphosphatase enzyme. The C3 and C4 m thioredoxins were similar in their specificity (ability to activate NADP-malate dehydrogenase, and not fructose-1,6-bisphosphatase) and isoelectric points (ca. 4.8), but differed slightly in molecular mass (13.0 kDa for spinach vs 13.5 kDa for corn) and substantially in their immunological properties. Results obtained in conjunction with these studies demonstrated that the thioredoxin m-linked activation of NADP-malate dehydrogenase in selectively enhanced by the presence of halide ions (e.g., chloride) and by an organic solvent (e.g., 2-propanol). The results suggest that in vivo NADP-malate dehydrogenase interacts with thylakoid membranes and is regulated to a greater extent by thioredoxin m than thioredoxin f.     

Effects of irradiance and temperature on photosynthesis in C3, C4 and C3/C4 Panicum species

Species in the Laxa and Grandia groups of the genus Panicum are adapted to low, wet areas of tropical and subtropical America. Panicum milioides is a species with C₃ photosynthesis and low apparent photorespiration and has been classified as a C₃/C₄ intermediate. Other species in the Laxa group are C₃ with normal photorespiration. Panicum prionitis is a C₄ species in the Grandia group. Since P. milioides has some leaf characteristics intermediate to C₃ and C₄ species, its photosynthetic response to irradiance and temperature was compared to the closely related C₃ species, P. laxum and P. boliviense and to P. prionitis. The response of apparent photosynthesis to irradiance and temperature was similar to that of P. laxum and P. boliviense, with saturation at a photosynthetic photo flux density of about 1 mmol m^-2 s^-1 at 30°C and temperature optimum near 30°C. In contrast, P. prionitis showed no light saturation up to 2 mmol m^-2 s^-1 and an optimum temperature near 40°C. P. milioides exhibited low CO₂ loss into CO₂-free air in the light and this loss was nearly insensitive to temperature. Loss of CO₂ in the light in the C₃ species, P. laxum and P. boliviense, was several-fold higher than in P. milioides and increased 2- to 5-fold with increases in temperature from 10 to 40°C. The level of dark respiration and its response to temperature were similar in all four Panicum species examined. It is concluded that the low apparent photorespiration in P. milioides does not influence its response of apparent photosynthesis to irradiance and temperature in comparison to closely related C₃ Panicum species.Abbreviations AP apparent photosynthesis - I CO₂ compensation point - gl leaf conductance; gm, mesophyll conductance - PPFD photosynthetic photon flux density - PR apparent photorespiration rate - RuBPC sibulose bisphosphate carboxylase     

CAM photosynthesis in submerged aquatic plants

Crassulacean acid metabolism (CAM) is a CO₂-concentrating mechanism selected in response to aridity in terrestrial habitats, and, in aquatic environments, to ambient limitations of carbon. Evidence is reviewed for its presence in five genera of aquatic vascular plants, includingIsoëtes, Sagittaria, Vallisneria, Crassula, andLittorella. Initially, aquatic CAM was considered by some to be an oxymoron, but some aquatic species have been studied in sufficient detail to say definitively that they possess CAM photosynthesis. CO₂-concentrating mechanisms in photosynthetic organs require a barrier to leakage; e.g., terrestrial C₄ plants have suberized bundle sheath cells and terrestrial CAM plants high stomatal resistance. In aquatic CAM plants the primary barrier to CO₂ leakage is the extremely high difrusional resistance of water. This, coupled with the sink provided by extensive intercellular gas space, generates daytime CO₂(pi) comparable to terrestrial CAM plants. CAM contributes to the carbon budget by both net carbon gain and carbon recycling, and the magnitude of each is environmentally influenced. Aquatic CAM plants inhabit sites where photosynthesis is potentially limited by carbon. Many occupy moderately fertile shallow temporary pools that experience extreme diel fluctuations in carbon availability. CAM plants are able to take advantage of elevated nighttime CO2 levels in these habitats. This gives them a competitive advantage over non-CAM species that are carbon starved during the day and an advantage over species that expend energy in membrane transport of bicarbonate. Some aquatic CAM plants are distributed in highly infertile lakes, where extreme carbon limitation and light are important selective factors. Compilation of reports on diel changes in titratable acidity and malate show 69 out of 180 species have significant overnight accumulation, although evidence is presented discounting CAM in some. It is concluded that similar proportions of the aquatic and terrestrial floras have evolved CAM photosynthesis. AquaticIsoëtes (Lycophyta) represent the oldest lineage of CAM plants and cladistic analysis supports an origin for CAM in seasonal wetlands, from which it has radiated into oligotrophic lakes and into terrestrial habitats. Temperate Zone terrestrial species share many characteristics with amphibious ancestors, which in their temporary terrestrial stage, produce functional stomata and switch from CAM to C₃. Many lacustrineIsoëtes have retained the phenotypic plasticity of amphibious species and can adapt to an aerial environment by development of stomata and switching to C₃. However, in some neotropical alpine species, adaptations to the lacustrine environment are genetically fixed and these constitutive species fail to produce stomata or loose CAM when artificially maintained in an aerial environment. It is hypothesized that neotropical lacustrine species may be more ancient in origin and have given rise to terrestrial species, which have retained most of the characteristics of their aquatic ancestry, including astomatous leaves, CAM and sediment-based carbon nutrition.     

An analysis of photosynthetic response and adaptation to temperature in higher plants: temperature acclimation in the desert evergreen Nerium oleander L*

Abstract. Factors underlying the process of photosynthetic acclimation to temperature were investigated for the shrub Nerium oleander L. Ramets of a single clone were grown under day/night temperature regimes of 20°C/15°C or 45°C/32°C. Plants grown at the lower temperature regime possessed rates of photosynthesis twice that of the high-temperature grown plants when CO₂ fixation was measured at 20°C. In contrast, the plants grown at the high-temperature regime had twice the rate of CO₂ fixation of the 20°C/l 5°C-grown plants at a measurement temperature of 45° C. It was determined that the ability to acclimate to changes in temperature regime was present in fully mature leaves. A reciprocal transfer of plants between the two growth regimes resulted in the appearance of the CO₂ fixation characteristics appropriate to the new growth temperature after 12–14d. The response of CO₂ fixation to light, temperature, and CO₂ partial pressure and the temperature responses of soluble and membrane-bound photosynthetic enzyme systems were analysed to determine which components might be responsible for the superior photosynthetic performance of the 20°C/I5°C-grown plants at 20°C, and the enhanced high-temperature stability of the 45°C/32°C plants. The measured photosynthetic capacity of the 20°C/15°C plants could not be attributed to gross morphological, stomatal, or other physical changes, or to a general increase in the concentration of components of the photosynthetic process. Only a single enzyme, Fru-P₂ phosphatase, was affected to an extent similar to that of photosynthesis. The enhanced thermal stability of the 45°C/32°C plants may be attributed primarily to an enhanced stability of the chloroplast membrane-bound enzymatic activities and the stability of the photosynthetic carbon metabolism enzymes which require lighl for activation.     

Temperature acclimation of photosynthesis: mechanisms involved in the changes in temperature dependence of photosynthetic rate

Growth temperature alters temperature dependence of the photosynthetic rate (temperature acclimation). In many species, the optimal temperature that maximizes the photosynthetic rate increases with increasing growth temperature. In this minireview, mechanisms involved in changes in the photosynthesis-temperature curve are discussed. Based on the biochemical model of photosynthesis, change in the photosynthesis-temperature curve is attributable to four factors: intercellular CO2 concentration, activation energy of the maximum rate of RuBP (ribulose-1,5-bisphosphate) carboxylation (Vc max), activation energy of the rate of RuBP regeneration (Jmax), and the ratio of Jmax to Vc max. In the survey, every species increased the activation energy of Vc max with increasing growth temperature. Other factors changed with growth temperature, but their responses were different among species. Among these factors, activation energy of Vc max may be the most important for the shift of optimal temperature of photosynthesis at ambient CO2 concentrations. Physiological and biochemical causes for the change in these parameters are discussed.     

The acclimation of photosynthesis and respiration to temperature in the C3–C4 intermediate Salsola divaricata: induction of high respiratory CO2 release under low temperature

Photosynthesis in C₃–C₄ intermediates reduces carbon loss by photorespiration through refixing photorespired CO₂ within bundle sheath cells. This is beneficial under warm temperatures where rates of photorespiration are high; however, it is unknown how photosynthesis in C₃–C₄ plants acclimates to growth under cold conditions. Therefore, the cold tolerance of the C₃–C₄ Salsola divaricata was tested to determine whether it reverts to C₃ photosynthesis when grown under low temperatures. Plants were grown under cold (15/10 °C), moderate (25/18 °C) or hot (35/25 °C) day/night temperatures and analysed to determine how photosynthesis, respiration and C₃–C₄ features acclimate to these growth conditions. The CO₂ compensation point and net rates of CO₂ assimilation in cold‐grown plants changed dramatically when measured in response to temperature. However, this was not due to the loss of C₃–C₄ intermediacy, but rather to a large increase in mitochondrial respiration supported primarily by the non‐phosphorylating alternative oxidative pathway (AOP) and, to a lesser degree, the cytochrome oxidative pathway (COP). The increase in respiration and AOP capacity in cold‐grown plants likely protects against reactive oxygen species (ROS) in mitochondria and photodamage in chloroplasts by consuming excess reductant via the alternative mitochondrial respiratory electron transport chain.     

Photostasis and cold acclimation: sensing low temperature through photosynthesis

Photosynthesis is a highly integrated and regulated process which is highly sensitive to any change in environmental conditions, because it needs to balance the light energy absorbed by the photosystems with the energy consumed by metabolic sinks of the plant. Low temperatures exacerbate an imbalance between the source of energy and the metabolic sink, thus requiring adjustments of photosynthesis to maintain the balance of energy flow. Photosynthesis itself functions as a sensor of this imbalance through the redox state of photosynthetic electron-transport components and regulates photophysical, photochemical and metabolic processes in the chloroplast. Recent progress has been made in understanding how plants sense the low temperature signal. It is clear that photosynthesis interacts with other processes during cold acclimation involving crosstalk between photosynthetic redox, cold acclimation and sugar-signalling pathways to regulate plant acclimation to low temperatures.     

Biochar amendment boosts photosynthesis and biomass in C3 but not C4 plants: A global synthesis

Biochar is a carbon (C)‐rich solid produced from the thermochemical pyrolysis of biomass. Its amendment to soils has been proposed as a promising mean to mitigate greenhouse gas emissions and simultaneously benefit agricultural crops. However, how biochar amendment affects plant photosynthesis and growth remains unclear, especially on a global scale. In this study, we conducted a global synthesis of 74 publications with 347 paired comparisons to acquire an overall tendency of plant photosynthesis and growth following biochar amendment. Overall, we found that biochar amendment significantly increased photosynthetic rate by 27.1%, and improved stomatal conductance, transpiration rate, water use efficiency, and chlorophyll concentration by 19.6%, 26.9%, 26.8%, and 16.1%, respectively. Meanwhile, plant total biomass, shoot biomass, and root biomass increased by 25.4%, 22.1%, and 34.4%, respectively. Interestingly, plant types (C₃ and C₄ plants) showed greater control over plant photosynthesis and biomass than a broad suite of soil and biochar factors. Biochar amendment largely boosted photosynthesis and biomass on C₃ plants, but had a limited effect on C₄ plants. Our results highlight the importance of the differential response of plant types to biochar amendment with respect to plant growth and photosynthesis, providing a scientific foundation for making reasonable strategies towards an extensive application of biochar for agricultural production management.     

C4 plants use fluctuating light less efficiently than do C3 plants: a study of growth,photosynthesis and carbon isotope discrimination

Plants in the field are commonly exposed to fluctuating light intensity, caused by variable cloud cover, self‐shading of leaves in the canopy and/or leaf movement due to turbulence. In contrast to C₃ plant species, only little is known about the effects of dynamic light (DL) on photosynthesis and growth in C₄ plants. Two C₄ and two C₃ monocot and eudicot species were grown under steady light or DL conditions with equal sum of daily incident photon flux. We measured leaf gas exchange, plant growth and dry matter carbon isotope discrimination to infer CO₂ bundle sheath leakiness in C₄ plants. The growth of all species was reduced by DL, despite only small changes in steady‐state gas exchange characteristics, and this effect was more pronounced in C₄ than C₃ species due to lower assimilation at light transitions. This was partially attributed to increased bundle sheath leakiness in C₄ plants under the simulated lightfleck conditions. We hypothesize that DL leads to imbalances in the coordination of C₄ and C₃ cycles and increasing leakiness, thereby decreasing the quantum efficiency of photosynthesis. In addition to their other constraints, the inability of C₄ plants to efficiently utilize fluctuating light likely contributes to their absence in such environments as forest understoreys.     

Effects of elevated CO2 on the tolerance of photosynthesis to acute heat stress in C3, C4, and CAM species

Determining the effect of elevated CO(2) on the tolerance of photosynthesis to acute heat stress (AHS) is necessary for predicting plant responses to global warming because photosynthesis is heat sensitive and AHS and atmospheric CO(2) will increase in the future. Few studies have examined this effect, and past results were variable, which may be related to methodological variation among studies. In this study, we grew 11 species that included cool and warm season and C(3), C(4), and CAM species at current or elevated (370 or 700 ppm) CO(2) and at species-specific optimal growth temperatures and at 30°C (if optimal ≠ 30°C). We then assessed thermotolerance of net photosynthesis (P(n)), stomatal conductance (g(st)), leaf internal [CO(2)], and photosystem II (PSII) and post-PSII electron transport during AHS. Thermotolerance of P(n) in elevated (vs. ambient) CO(2) increased in C(3), but decreased in C(4) (especially) and CAM (high growth temperature only), species. In contrast, elevated CO(2) decreased electron transport in 10 of 11 species. High CO(2) decreased g(st) in five of nine species, but stomatal limitations to P(n) increased during AHS in only two cool-season C(3) species. Thus, benefits of elevated CO(2) to photosynthesis at normal temperatures may be partly offset by negative effects during AHS, especially for C(4) species, so effects of elevated CO(2) on acute heat tolerance may contribute to future changes in plant productivity, distribution, and diversity.     

Effects of root applications of gibberellic acid on photosynthesis and growth in C3 and C4 plants

The effects of root applications of gibberellic acid (GA₃) on growth and photosynthesis of 12 species of plants including C₃ monocots (Triticum aestivum L., wheat, Hordeum vulgare L., barley and Avena sativa L., oat), C₃ dicots (Vigna radiata L., mung bean, Cucurbita moschata L., squash and Capsicum annuum L., pepper), C₄ monocots (Zea mays L., corn, Sorghum vulgare L., sorghum and Panicum ramosum L., millet) and C₄ dicots (Amaranthus retroflexus L., pigweed, Kochia scoparis L., kochia and Gomphrena celosoides L., gomphrena) were evaluated. Relative growth rates (RGR) of barley, oat, squash, pepper, corn, sorghum, millet, pigweed and kochia were increased above the control by 12.7%, 9.9%, 11.3%, 10.7%, 19.2% 10.1%, 11.5%, 16.4% and 32.7% respectively, four days following optimum GA₃ treatments. There was no effect of GA₃ on RGR in wheat, mung bean, and gomphrena. Gibberellic acid decreased the chlorophyll content expressed on an area basis by 20.0%, 13.9%, 20.9%, 17.1%, 11.9% and 28.0% in barley, squash, pepper, sorghum, pigweed and kochia, respectively, while that of oat, wheat, mung bean, corn, millet and gomphrena remained unchanged. When photosynthetic rates were expressed per mg of chlorophyll, it showed that GA₃ could stimulate photosynthesis in barley, squash, pepper, sorghum, millet, pigweed and kochia by 20.4%, 20.6%, 16.5%, 17.4%, 10.4%, 24.2%, and 29.4%; while there was no effect in oat, wheat, mung bean, corn and gomphrena. An increase in leaf blade area and/or length of sheath were observed in GA₃ treated plants of oat, barley, mung bean, squash, pepper, corn, sorghum, millet and kochia. The transpiration rate remained unchanged following GA₃ treatment in all 12 species.This work was supported in part by the Fair Funds administered by the Pennsylvania Department of Agriculture. Contribution No. 39, Department of Horticulture, The Pennsylvania State University. Authorized for publications as paper no. 6886 in the journal series of the Pennsylvania Agricultual Experiment Station.Research assistant and assistant professor respectively.     

Temperature acclimation of photosynthesis in spinach leaves: analyses of photosynthetic components and temperature dependencies of photosynthetic partial reactions

Spinach (Spinacia oleracea) plants were grown under the day/night temperature regime of 15/10 °C (LT) or 30/25 °C (HT). The plants were also transferred from HT to LT when the sample leaves were at particular developmental stages (HL-transfer). With fully mature leaves, the light-saturated photosynthetic rate (A) at the ambient CO₂ concentration (C_a) of 1500 µL L⁻¹ (A₁₅₀₀) and the initial slope of A versus intercellular CO₂ concentration (C_i) at low C_i region (IS) were obtained to assess capacities of RuBP regeneration and carboxylation. Photosynthetic components including Rubisco and cytochrome f (Cyt f) were also determined. The optimum temperatures for A at C_a of 360 µL L⁻¹ (A₃₆₀), A₁₅₀₀ and IS in HT leaves were 27, 36 and 24 °C, whereas those in LT leaves were 18, 30 and 18 °C. The optimum temperatures in HL-transfer leaves approached those of LT leaves with the increase in the duration at LT. The shift in the optimum temperature was greater and quicker for IS than A₁₅₀₀. By the HL-transfer, the maximum values of A₁₅₀₀ and IS also increased. The maximum A₁₅₀₀ and Cyt f content increased more promptly than IS and Rubisco content. Changes in the Cyt f/Rubisco ratio were reflected to those in the A₁₅₀₀/IS ratio. Taken together, photosynthetic acclimation to low temperature in spinach leaves was due not only to the change in the balance of the absolute rates of RuBP regeneration and carboxylation but also to the large change in the optimum temperature of RuBP carboxylation.     

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.     

Effect of phosphinothricin (glufosinate) on photosynthesis and photorespiration of C3 and C4 plants

Phosphinothricin (glufosinate), an irreversible inhibitor of glutamine synthetase, causes an inhibition of photosynthesis in C₃ (Sinapis alba) and C₄ (Zea mays) plants under atmospheric conditions (400 ppm CO₂, 21% O₂). This photosynthesis inhibition is proceeding slower in C₄ leaves. Under non-photorespiratory conditions (1000 ppm CO₂, 2% O₂) there is no inhibition of photosynthesis. The inhibition of glutamine synthetase by phosphinothricin results in an accumulation of NH₄ ⁺. The NH₄ ⁺-accumulation is lower in C₄ plants than in C₃ plants. The inhibition of glutamine synthetase through phosphinothricin in mustard leaves results in a decrease in glutamine, glutamate, aspartate, asparagine, serine, and glycine. In contrast to this, a considerable increase in leucine and valine following phosphinothricin treatment is measured. With the addition of either glutamine, glutamate, aspartate, glycine or serine, photosynthesis inhibition by phosphinothricin can be reduced, although the NH₄ ⁺-accumulation is greatly increased. This indicates that NH₄ ⁺-accumulation cannot be the primary cause for photosynthesis inhibition by phosphinothricin. The investigations demonstrate the inhibition of transmination of glyoxylate to glycine in photorespiration through the total lack of amino donors. This could result in a glyoxylate accumulation inhibiting ribulose-1,5-bisphosphate-carboxylase and consequently CO₂-fixation.Abbreviations GOGAT glutamine-2-oxoglutarate-amidotransferase - GS glutamine synthetase - PPT phosphinothricin - MSO methionine sulfoximine - RuBP ribulose-1,5-bisphosphate