Photosynthetic responses to chilling in a chilling-tolerant and chilling-sensitive Miscanthus hybrid.

Miscanthus is a C4 perennial grass being developed for bioenergy production in temperate regions where chilling events are common. To evaluate chilling effects on Miscanthus, we assessed the processes controlling net CO2 assimilation rate (A) in Miscanthus x giganteus (M161) and a chilling-sensitive Miscanthus hybrid (M115) before and after a chilling treatment of 12/5 °C. The temperature response of A and maximum Rubisco activity in vitro were identical below 20 °C in chilled and unchilled M161, demonstrating Rubisco capacity limits or co-limits A at cooler temperatures. By contrast, A in M115 decreased at all measurement temperatures after growth at 12/5 °C. Rubisco activity in vitro declined in proportion to the reduction in A in chilled M115 plants, indicating Rubisco capacity is responsible in part for the decline in A. Pyruvate orthophosphate dikinase activities were also reduced by the chilling treatment when assayed at 28 °C, indicating this enzyme may also contribute to the reduction in A in M115. The maximum extractable activities of PEPCase and NADP-ME remained largely unchanged after chilling. The carboxylation efficiency of the C4 cycle was depressed in both genotypes to a similar extent after chilling. ΦP :ΦCO2 remained unchanged in both genotypes indicating the C3 and C4 cycles decline equivalently upon chilling.

Photorespiratory compensation: a driver for biological diversity.

This paper reviews how terrestrial plants reduce photorespiration and thus compensate for its inhibitory effects. As shown in the equation φ = (1/Sc/o )O/C, where φ is the ratio of oxygenation to carboxylation, Sc/o is the relative specificity of Rubisco, O is stromal O2 level and C is the stromal CO2 concentration, plants can reduce photorespiration by increasing Sc/o or C, or by reducing O. By far the most effective means of reducing φ is by concentrating CO2, as occurs in C4 and CAM plants, and to a lesser extent in plants using a glycine shuttle to concentrate CO2 into the bundle sheath. Trapping and refixation of photorespired CO2 by a sheath of chloroplasts around the mesophyll cell periphery in C3 plants also enhances C, particularly at low atmospheric CO2. O2 removal is not practical because high energy and protein investment is needed to have more than a negligible effect. Sc/o enhancement provides for modest reductions in φ, but at the potential cost of limiting the kcat of Rubisco. An effective means of decreasing φ and enhancing carbon gain is to lower leaf temperature by reducing absorbance of solar radiation, or where water is abundant, opening stomata. By using a combination of mechanisms, C3 plants can achieve substantial (>30%) reductions in φ. This may have allowed many C3 species to withstand severe competition from C4 plants in low CO2 atmospheres of recent geological time, thereby preserving some of the Earth's floristic diversity that accumulated over millions of years.

Engineering photorespiration: current state and future possibilities.

Reduction of flux through photorespiration has been viewed as a major way to improve crop carbon fixation and yield since the energy-consuming reactions associated with this pathway were discovered. This view has been supported by the biomasses increases observed in model species that expressed artificial bypass reactions to photorespiration. Here, we present an overview about the major current attempts to reduce photorespiratory losses in crop species and provide suggestions for future research priorities.

The response of the high altitude C(4) grass Muhlenbergia montana (Nutt.) A.S. Hitchc. to long- and short-term chilling.

The acclimation of C(4) photosynthesis to low temperature was studied in the montane grass Muhlenbergia montana in order to evaluate inherent limitations in the C(4) photosynthetic pathway following chilling. Plants were grown in growth cabinets at 26 degrees C days, but at night temperatures of either 16 degrees C (the control treatment), 4 degrees C for at least 28 nights (the cold-acclimated treatment), or 1 night (the cold-stress treatment). Below a measurement temperature of 25 degrees C, little difference in the thermal response of the net CO(2) assimilation rate (A) was observed between the control and cold-acclimated treatment. By contrast, above 30 degrees C, A in the cold-acclimated treatment was 10% greater than in the control treatment. The temperature responses of Rubisco activity and net CO(2) assimilation rate were similar below 22 degrees C, indicating high metabolic control of Rubisco over the rate of photosynthesis at cool temperatures. Analysis of the response of A to intercellular CO(2) level further supported a major limiting role for Rubisco below 20 degrees C. As temperature declined, the CO(2) saturated plateau of A exhibited large reductions, while the initial slope of the CO(2) response was little affected. This type of response is consistent with a Rubisco limitation, rather than limitations in PEP carboxylase capacity. Stomatal limitations at low temperature were not apparent because photosynthesis was CO(2) saturated below 23 degrees C at air levels of CO(2). In contrast to the response of photosynthesis to temperature and CO(2) in plants acclimated for 4 weeks to low night temperature, plants exposed to 4 degrees C for one night showed substantial reduction in photosynthetic capacity at temperatures above 20 degrees C. Because these reductions were at both high and low CO(2), enzymes associated with the C(4) carbon cycle were implicated as the major mechanisms for the chilling inhibition. These results demonstrate that C(4) plants from climates with low temperature during the growing season can fully acclimate to cold stress given sufficient time. This acclimation appears to involve reversal of injury to the C(4) cycle following initial exposure to low temperature. By contrast, carbon gain at low temperatures generally appears to be constrained by the carboxylation capacity of Rubisco, regardless of acclimation time. The inability to overcome the Rubisco limitation at low temperature may be an inherent limitation restricting C(4) photosynthetic performance in cooler climates.

Effects of low atmospheric CO(2) on plants: more than a thing of the past.

In recent geological time, atmospheric CO(2) concentrations were 25-50% below the current level. Photosynthetic productivity of C(3) plants is substantially reduced at these low CO(2) levels, particularly at higher temperatures and during stress. Acclimation of photosynthesis to reduced CO(2) levels might compensate for some of this inhibition, but plants have a limited capacity to modulate Rubisco and other photosynthetic proteins following CO(2) reduction. Because of this, low CO(2) probably acted as a significant evolutionary agent, selecting plants adapted to CO(2) deficiency. Adaptations to low CO(2) might still exist in plants and might constrain responses to a rising CO(2) concentration.

Acclimation of photosynthesis to increasing atmospheric CO2: The gas exchange perspective.

The nature of photosynthetic acclimation to elevated CO2 is evaluated from the results of over 40 studies focusing on the effect of long-term CO2 enrichment on the short-term response of photosynthesis to intercellular CO2 (the A/Ci response). The effect of CO2 enrichment on the A/Ci response was dependent on growth conditions, with plants grown in small pots (< 5 L) or low nutrients usually exhibiting a reduction of A at a given Ci, while plants grown without nutrient deficiency in large pots or in the field tended to exhibit either little reduction or an enhancement of A at a given Ci following a doubling or tripling of atmospheric CO2 during growth. Using theoretical interpretations of A/Ci curves to assess acclimation, it was found that when pot size or nutrient deficiency was not a factor, changes in the shape of A/Ci curves which are indicative of a reallocation of resources within the photosynthetic apparatus typically were not observed. Long-term CO2 enrichment usually had little effect or increased the value of A at all Ci. However, a minority of species grown at elevated CO2 exhibited gas exchange responses indicative of a reduced amount of Rubisco and an enhanced capacity to metabolize photosynthetic products. This type of response was considered beneficial because it enhanced both photosynthetic capacity at high CO2 and reduced resource investment in excessive Rubisco capacity. The ratio of intercellular to ambient CO2 (the Ci/Ca ratio) was used to evaluate stomatal acclimation. Except under water and humidity stress, Ci/Ca exhibited no consistent change in a variety of C3 species, indicating no stomatal acclimation. Under drought or humidity stress, Ci/Ca declined in high-CO2 grown plants, indicating stomata will become more conservative during stress episodes in future high CO2 environments.

Regulation of Ribulose-1,5-Bisphosphate Carboxylase/Oxygenase Activity in Response to Reduced Light Intensity in C4 Plants.

The light-dependent regulation of ribulose-1,5-bisphosphate carboxylase/oxygenase (Rubisco) activity was studied in 16 species of C4 plants representing all three biochemical subtypes and a variety of taxonomic groups. Rubisco regulation was assessed by measuring (a) the ratio of initial to total Rubisco activity, which reflects primarily the carbamylation state of the enzyme, and (b) total Rubisco activity per mol of Rubisco catalytic sites, which declines when 2-carboxyarabinitol 1-phosphate (CA1P) binds to carbamylated Rubisco. In all species examined, the activity ratio of Rubisco declined with a reduction in light intensity, although substantial variation was apparent between species in the degree of Rubisco deactivation. No relationship existed between the degree of Rubisco deactivation and C4 subtype. Dicots generally deactivated Rubisco to a greater degree than monocots. The total activity of Rubisco per catalytic site was generally independent of light intensity, indicating that CA1P and other inhibitors are not major contributors to the light-dependent regulation of Rubisco activity in C4 plants. The light response of the activity ratio of Rubisco was measured in detail in Amaranthus retroflexus, Brachiaria texana, and Zea mays. In A. retroflexus and B. texana, the activity ratio declined dramatically below a light intensity of 400 to 500 [mu]mol of photons m-2 s-1. In Z. mays, the activity ratio of Rubisco was relatively insensitive to light intensity compared with the other species. In A. retroflexus, the pool size of ribulose bisphosphate (RuBP) declined with reduced light intensity except between 50 and 500 [mu]mol m-2 s-1, when the activity ratio of Rubisco was light dependent. In Z. mays, by contrast, the pool size of RuBP was light dependent only below 350 [mu]mol m-2 s-1. These results indicate that, in response to changes in light intensity, most C4 species regulate Rubisco by reversible carbamylation of catalytic sites, as commonly observed in C3 plants. In a few species, notably Z. mays, Rubisco is not extensively regulated in response to changes in light intensity, possibly because the activity of the CO2 pump may become limiting for photosynthesis at subsaturating light intensity.

Light-dependent modulation of ribulose-1,5-bisphosphate carboxylase/oxygenase activity in the genus Phaseolus.

Modulation of the activity of ribulose-1,5-bisphosphate carboxylase/oxygenase (Rubisco) in low light and darkness was measured in A) 25 genotypes from the four cultivated species of Phaseolus (P. vulgaris, P. acutifolius, P. lunatus and P. coccineus), B) 8 non-cultivated Phaseolus species, and C) the related species Macroptileum atropurpureum. The activity ratio of Rubisco (the ratio of initial and total Rubisco activities, which reflects Rubisco carbamylation), and the molar activity of fully-activated Rubisco (which primarily reflects the inhibition of Rubisco activity by carboxyarabinitol 1-phosphate, CA1P) were assayed in leaves from the cultivated species sampled at midday in full sunlight, in low light at dusk (60 to 100 µmol photons m(-2)s(-1)), and after at least 4 h in darkness. Dark inhibition of Rubisco molar activity was compared in both cultivated and non-cultivated species. In all cultivated genotypes, a significant reduction of the activity ratio of Rubisco was measured in leaves sampled at low light; however, the molar activity of fully activated Rubisco was not greatly reduced in these low light samples. In darkened leaves, molar activities substantially declined in most Phaseolus species with 11 of 13 exhibiting greater than 60% reduction. In P. vulgaris, the reduction of molar activity was extensive (greater than 69%) in all genotypes studied, which included wild progenitors as well as ancient and advanced cultivars. These results indicate that at low light late in the day, modulation of Rubisco activity is primarily through changes in carbamylation state, with CA1P playing a more limited role. By contrast in the dark, binding of CA1P dominates the modulation of Rubisco activity in Phaseolus in a pattern that appears to be conserved within a species, but can vary significantly between species within a genus. The degree of CA1P inhibition in Phaseolus was associated with phylogenetic affinities within the genus, as the species with extensive dark-inhibition of Rubisco activity tended to be more closely related to each other than to species with reduced inhibition of Rubisco activity.

A Comparison of Dark Respiration between C(3) and C(4) Plants.

Lower respiratory costs were hypothesized as providing an additional benefit in C(4) plants compared to C(3) plants due to less investment in proteins in C(4) leaves. Therefore, photosynthesis and dark respiration of mature leaves were compared between a number of C(4) and C(3) species. Although photosynthetic rates were generally greater in C(4) when compared to C(3) species, no differences were found in dark respiration rates of individual leaves at either the beginning or after 16 h of the dark period. The effects of nitrogen on photosynthesis and respiration of individual leaves and whole plants were also investigated in two species that occupy similar habitats, Amaranthus retroflexus (C(4)) and Chenopodium album (C(3)). For mature leaves of both species, there was no relationship between leaf nitrogen and leaf respiration, with leaves of both species exhibiting a similar rate of decline after 16 h of darkness. In contrast, leaf photosynthesis increased with increasing leaf nitrogen in both species, with the C(4) species displaying a greater photosynthetic response to leaf nitrogen. For whole plants of both species grown at different nitrogen levels, there was a clear linear relationship between net CO(2) uptake and CO(2) efflux in the dark. The dependence of nightly CO(2) efflux on CO(2) uptake was similar for both species, although the response of CO(2) uptake to leaf nitrogen was much steeper in the C(4) species, Amaranthus retroflexus. Rates of growth and maintenance respiration by whole plants of both species were similar, with both species displaying higher rates at higher leaf nitrogen. There were no significant differences in leaf or whole plant maintenance respiration between species at any temperature between 18 and 42 degrees C. The data suggest no obvious differences in respiratory costs in C(4) and C(3) plants.

Climate change and the evolution of C(4) photosynthesis.

Plants assimilate carbon by one of three photosynthetic pathways, commonly called the C(3), C(4), and CAM pathways. The C(4) photosynthetic pathway, found only among the angiosperms, represents a modification of C(3) metabolism that is most effective at low concentrations of CO(2). Today, C(4) plants are most common in hot, open ecosystems, and it is commonly felt that they evolved under these conditions. However, high light and high temperature, by themselves, are not sufficient to favor the evolution of C(4) photosynthesis at atmospheric CO(2) levels significantly above the current ambient values. A review of evidence suggests that C(4) plants evolved in response to a reduction in atmospheric CO(2) levels that began during the Cretaceous and continued until the Miocene.

A Model Describing the Regulation of Ribulose-1,5-Bisphosphate Carboxylase, Electron Transport, and Triose Phosphate Use in Response to Light Intensity and CO(2) in C(3) Plants.

A model of the regulation of the activity of ribulose-1,5-bisphosphate carboxylase, electron transport, and the rate of orthophosphate regeneration by starch and sucrose synthesis in response to changes in light intensity and partial pressures of CO(2) and O(2) is presented. The key assumption behind the model is that nonlimiting processes of photosynthesis are regulated to balance the capacity of limiting processes. Thus, at CO(2) partial pressures below ambient, when a limitation on photosynthesis by the capacity of rubisco is postulated, the activities of electron transport and phosphate regeneration are down-regulated in order that the rate of RuBP regeneration matches the rate of RuBP consumption by rubisco. Similarly, at subsaturating light intensity or elevated CO(2), when electron transport or Pi regeneration may limit photosynthesis, the activity of rubisco is downregulated to balance the limitation in the rate of RuBP regeneration. Comparisons with published data demonstrate a general consistency between modelled predictions and measured results.

Regulation of Ribulose-1,5-Bisphosphate Carboxylase Activity in Response to Light Intensity and CO(2) in the C(3) Annuals Chenopodium album L. and Phaseolus vulgaris L.

The light and CO(2) response of (a) photosynthesis, (b) the activation state and total catalytic efficiency (k(cat)) of ribulose-1,5-bisphosphate carboxylase (rubisco), and (c) the pool sizes of ribulose 1,5-bisphosphate, (RuBP), ATP, and ADP were studied in the C(3) annuals Chenopodium album and Phaseolus vulgaris at 25 degrees C. The initial slope of the photosynthetic CO(2) response curve was dependent on light intensity at reduced light levels only (less than 450 micromoles per square meter per second in C. album and below 200 micromoles per square meter per second in P. vulgaris). Modeled simulations indicated that the initial slope of the CO(2) response of photosynthesis exhibited light dependency when the rate of RuBP regeneration limited photosynthesis, but not when rubisco capacity limited photosynthesis. Measured observations closely matched modeled simulations. The activation state of rubisco was measured at three light intensities in C. album (1750, 550, and 150 micromoles per square meter per second) and at intercellular CO(2) partial pressures (C(1)) between the CO(2) compensation point and 500 microbars. Above a C(1) of 120 microbars, the activation state of rubisco was light dependent. At light intensities of 550 and 1750 micromoles per square meter per second, it was also dependent on C(1), decreasing as the C(1) was elevated above 120 microbars at 550 micromoles per square meter per second and above 300 microbars at 1750 micromoles per square meter per second. The pool size of RuBP was independent of C(1) only under conditions when the activation state of rubisco was dependent on C(1). Otherwise, RuBP pool sizes increased as C(1) was reduced. ATP pools in C. album tended to increase as C(1) was reduced. In P. vulgaris, decreasing C(1) at a subsaturating light intensity of 190 micromoles per square meter per second increased the activation state of rubisco but had little effect on the k(cat). These results support modelled simulations of the rubisco response to light and CO(2), where rubisco is assumed to be down-regulated when photosynthesis is limited by the rate of RuBP regeneration.

Acclimation of Photosynthesis to Elevated CO(2) in Five C(3) Species.

The effect of long-term (weeks to months) CO(2) enhancement on (a) the gas-exchange characteristics, (b) the content and activation state of ribulose-1,5-bisphosphate carboxylase (rubisco), and (c) leaf nitrogen, chlorophyll, and dry weight per area were studied in five C(3) species (Chenopodium album, Phaseolus vulgaris, Solanum tuberosum, Solanum melongena, and Brassica oleracea) grown at CO(2) partial pressures of 300 or 900 to 1000 microbars. Long-term exposure to elevated CO(2) affected the CO(2) response of photosynthesis in one of three ways: (a) the initial slope of the CO(2) response was unaffected, but the photosynthetic rate at high CO(2) increased (S. tuberosum); (b) the initial slope decreased but the CO(2)-saturated rate of photosynthesis was little affected (C. album, P. vulgaris); (c) both the initial slope and the CO(2)-saturated rate of photosynthesis decreased (B. oleracea, S. melongena). In all five species, growth at high CO(2) increased the extent to which photosynthesis was stimulated following a decrease in the partial pressure of O(2) or an increase in measurement CO(2) above 600 microbars. This stimulation indicates that a limitation on photosynthesis by the capacity to regenerate orthophosphate was reduced or absent after acclimation to high CO(2). Leaf nitrogen per area either increased (S. tuberosum, S. melongena) or was little changed by CO(2) enhancement. The content of rubisco was lower in only two of the five species, yet its activation state was 19% to 48% lower in all five species following long-term exposure to high CO(2). These results indicate that during growth in CO(2)-enriched air, leaf rubisco content remains in excess of that required to support the observed photosynthetic rates.

Boron toxicity in the rare serpentine plant, Streptanthus morrisonii.

The release of boron-laden mist from the cooling towers of some geothermal power stations in northern California potentially threatens nearby populations of the rare serpentine plant, Streptanthus morrisonii F. W. Hoffm. To assess the tolerance of S. morrisonii to high levels of boron, the effect of boron on leaf condition, life history, germination rate, growth rate, allocation and photosynthesis was measured on plants grown in a greenhouse. Relative to other species, S. morrisonii was tolerant of excess boron. On serpentine soil, mild to moderate toxicity symptoms (older leaves exhibiting chlorosis and necrosis, but few leaves killed) were apparent when the boron concentration in applied nutrient solutions was 240-650 microm. Severe toxicity symptoms (significant leaf loss, young leaves with toxicity symptoms) were apparent when the applied solution was over 1000 microm boron. Above 1000 microm boron, S. morrisonii appeared unable to complete its life cycle. On a tissue basis, boron toxicity was first observed when leaf boron content was 40-90 micromol g(-1) dry weight. In leaves with severe boron toxicity (> 35% injury), the boron content was generally above 130 micromol g(-1) dry weight. These levels were an order of magnitude above the tissue boron content of plants in the field. Prior to the onset of pronounced boron toxicity symptoms, growth rate, allocation patterns, and photosynthesis were unaffected by high boron. These results indicate that inhibition of growth and photosynthesis occurred because of a loss of viable tissue due to boron injury, rather than a progressive decline as leaf boron levels increased.

Regulation of photosynthetic electron-transport in Phaseolus vulgaris L., as determined by room-temperature chlorophyll a fluorescence.

The regulation of photosystem II (PSII) by light-, CO2-, and O2-dependent changes in the capacity for carbon metabolism was studied. Estimates of the rate of electron transport through PSII were made from gas-exchange data and from measurements of chlorophyll fluorescence. At subsaturating photon-flux density (PFD), the rate of electron transport was independent of O2 and CO2. Feedback on electron transport was observed under two conditions. At saturating PFD and low partial pressure of CO2, p(CO2), the rate of electron transport increased with p(CO2). However, at high p(CO2), switching from normal to low p(O2) did not affect the net rate of photosynthetic CO2 assimilation but the rate of electron-transport decreased by an amount related to the change in the rate of photorespiration. We interpret these effects as 1) regulation of ribulose-1,5-bisphosphatecarboxylase (RuBPCase, EC 4.1.1.39) activity to match the rate of electron transport at limiting PFD, 2) regulation of electron-transport rate to match the rate of RuBPCase at low p(CO2), and 3) regulation of the electron-transport rate to match the capacity for starch and sucrose synthesis at high p(CO2) and PFD. These studies provide evidence that PSII is regulated so that the capacity for electron transport is matched to the capacity for other processes required by photosynthesis, such as ribulose-bisphosphate carboxylation and starch and sucrose synthesis. We show that at least two mechanisms contribute to the regulation of PSII activity and that the relative engagement of these mechanisms varies with time following a step change in the capacity for ribulose-bisphosphate carboxylation and starch and sucrose synthesis. Finally, we take advantage of the relatively slow activation of deactivated RuBPCase in vivo to show that the activation level of this enzyme can limit the rate of electron transport as evidenced by increased feedback on PSII following a step change in p(CO2). As RuBPCase as activated, the feedback on PSII declined.

The in-vivo response of the ribulose-1,5-bisphosphate carboxylase activation state and the pool sizes of photosynthetic metabolites to elevated CO2 inPhaseolus vulgaris L.

The short-term, in-vivo response to elevated CO2 of ribulose-1,5-bisphosphate carboxylase (RuBPCase, EC 4.1.1.39) activity, and the pool sizes of ribulose 1,5-bisphosphate, 3-phosphoglyceric acid, triose phosphates, fructose 1,6-bisphosphate, glucose 6-phosphate and fructose 6-phosphate in bean were studied. Increasing CO2 from an ambient partial pressure of 360-1600 µbar induced a substantial deactivation of RuBPCase at both saturating and subsaturating photon flux densities. Activation of RuBPCase declined for 30 min following the CO2 increase. However, the rate of photosynthesis re-equilibrated within 6 min of the switch to high CO2, indicating that RuBPCase activity did not limit photosynthesis at high CO2. Following a return to low CO2, RuBPCase activation increased to control levels within 10 min. The photosynthetic rate fell immediately after the return to low CO2, and then increased in parallel with the increase in RuBPCase activation to the initial rate observed prior to the CO2 increase. This indicated that RuBPCase activity limited photosynthesis while RuBPCase activation increased. Metabolite pools were temporarily affected during the first 10 min after either a CO2 increase or decrease. However, they returned to their original level as the change in the activation state of RuBPCase neared completion. This result indicates that one role for changes in the activation state of RuBPCase is to regulate the pool sizes of photosynthetic intermediates.

The Nitrogen Use Efficiency of C(3) and C(4) Plants : III. Leaf Nitrogen Effects on the Activity of Carboxylating Enzymes in Chenopodium album (L.) and Amaranthus retroflexus (L.).

The relationships between leaf nitrogen content per unit area (N(a)) and (a) the initial slope of the photosynthetic CO(2) response curve, (b) activity and amount of ribulose-1,5-bisphosphate carboxylase/oxygenase (Rubisco) and phosphoenolpyruvate carboxylase (PEPC), and (c) chlorophyll content were studied in the ecologically similar weeds Chenopodium album (C(3)) and Amaranthus retroflexus (C(4)). In both species, all parameters were linearly dependent upon leaf N(a). The dependence of the initial slope of the CO(2) response of photosynthesis on N(a) was four times greater in A. retroflexus than in C. album. At equivalent leaf N(a) contents, C. album had 1.5 to 2.6 times more CO(2) saturated Rubisco activity than A. retroflexus. At equal assimilation capacities, C. album had four times the Rubisco activity as A. retroflexus. In A. retroflexus, a one to one ratio between Rubisco activity and photosynthesis was observed, whereas in C. album, the CO(2) saturated Rubisco activity was three to four times the corresponding photosynthetic rate. The ratio of PEPC to Rubisco activity in A. retroflexus ranged from four at low N(a) to seven at high N(a). The fraction of organic N invested in carboxylation enzymes increased with increased N(a) in both species. The fraction of N invested in Rubisco ranged from 10 to 27% in C. album. In A. retroflexus, the fraction of N(a) invested in Rubisco ranged from 5 to 9% and the fraction invested in PEPC ranged from 2 to 5%.

The Effect of Temperature on the Occurrence of O(2) and CO(2) Insensitive Photosynthesis in Field Grown Plants.

The sensitivity of photosynthesis to O(2) and CO(2) was measured in leaves from field grown plants of six species (Phaseolus vulgaris, Capsicum annuum, Lycopersicon esculentum, Scrophularia desertorum, Cardaria draba, and Populus fremontii) from 5 degrees C to 35 degrees C using gas-exchange techniques. In all species but Phaseolus, photosynthesis was insensitive to O(2) in normal air below a species dependent temperature. CO(2) insensitivity occurred under the same conditions that resulted in O(2) insensitivity. A complete loss of O(2) sensitivity occurred up to 22 degrees C in Lycopersicon but only up to 6 degrees C in Scrophularia. In Lycopersicon and Populus, O(2) and CO(2) insensitivity occurred under conditions regularly encountered during the cooler portions of the day. Because O(2) insensitivity is an indicator of feedback limited photosynthesis, these results indicate that feedback limitations can play a role in determining the diurnal carbon gain in the field. At higher partial pressures of CO(2) the temperature at which O(2) insensitivity occurred was higher, indicating that feedback limitations in the field will become more important as the CO(2) concentration in the atmosphere increases.

The Nitrogen Use Efficiency of C(3) and C(4) Plants: I. Leaf Nitrogen, Growth, and Biomass Partitioning in Chenopodium album (L.) and Amaranthus retroflexus (L.).

The effect of applied nitrogen (N) on the growth, leaf expansion rate, biomass partitioning and leaf N levels of Chenopodium album (C(3)) and Amaranthus retroflexus (C(4)) were investigated. At a given applied N level, C. album had 50% greater leaf N per unit area (N(a)) than A. retroflexus. Nitrate accumulated at lower N(a) in A. retroflexus than C. album. A. retroflexus was more productive than C. album at high N, but C. album was more productive at low N. At high applied N, nitrogen use efficiency (NUE), expressed either as net assimilation rate (NAR) per unit N or relative growth rate per unit N, was greater in A. retroflexus than C. album. However, at low applied N, C. album had a greater NUE on both an NAR and growth basis than A. retroflexus. The leaf area partitioning coefficient was similar in the species at high N, but was greater in A. retroflexus than C. album at low N. At low N, greater leaf area partitioning apparently lowered leaf N in A. retroflexus to levels at which necrosis occurred. In C. album by contrast, leaf area partitioning declined to a greater degree with declining N than it did in A. retroflexus, so that leaf N did not decline as much. Consequently, low N C. album plants did not lose leaf area to necrosis and had a greater NAR and NUE at low applied N than A. retroflexus.

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 (C(3)) and Amaranthus retroflexus (C(4)). 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.