The effect of water stress on photosynthetic carbon metabolism in four species grown under field conditions

Abstract. The effect of gradually-developing water-stress has been studied in Lupinus albus L., Helianthus annuus L., Vitis vinifera cv. Rosaki and Eucalyptus globulus Labill. Water was withheld and diurnal rhythms were investigated 4–8d later, when the predawn water deficit was more negative than in watered plants, and the stomata closed almost completely early during the photoperiod. The contribution of ‘stomatal’ and ‘non-stomatal’ components to the decrease of photosynthetic rate was investigated by (1) comparing the changes of the rate of photosynthesis in air with the changes of stomatal conductance and (2) measuring photosynthetic capacity in saturating irradiance and 15% CO₂. Three species (lupin, eucalyptus and sunflower) showed larger changes of stomatal conductance than photosynthesis in air, and showed little or no decrease of photosynthetic capacity in saturating CO₂. Photosynthesis in air also recovered fully overnight after watering the plants in the evening. In grapevines, stomatal conductance and photosynthesis in air changed in parallel, there was a marked decrease of photosynthetic capacity, and photosynthesis and stomatal conductance did not recover overnight after watering water-stressed plants. Relative water content remained above 90% in grapevine. We conclude that non-stomatal components do not play a significant role in lupins, sunflower or eucalyptus, but could in grapevine. The effect of water-stress on partitioning of photosynthate was investigated by measuring the amounts of sucrose and starch in leaves during a diurnal rhythm, and by measuring the partitioning of ¹⁴C-carbon dioxide between sucrose and starch. In all four species, starch was depleted in water-stressed leaves but sucrose was maintained at amounts similar to, or higher than, those in watered plants. Partitioning into sucrose was increased in lupins and eucalyptus, and remained unchanged in grapevine and sunflower. It is concluded that water-stressed leaves in all four species maintain high levels of soluble sugars in their leaves, despite having lower rates of field photosynthesis, decreased rates of export, and low amounts of starch in their leaves.     

Decreased expression of sucrose phosphate synthase strongly inhibits the water stress-induced synthesis of sucrose in growing potato tubers

Water stress stimulates sucrose synthesis and inhibits starch synthesis in wild-type tubers. Antisense and co-suppression potato transformants with decreased expression of sucrose–phosphate synthase (SPS) have been used to analyse the importance of SPS for the regulation of this water-stress induced change in partitioning. (i) In the absence of water stress, a 70–80% decrease in SPS activity led to a 30–50% inhibition of sucrose synthesis and a slight (10–20%) increase of starch synthesis in tuber discs in short-term labelling experiments with low concentrations of labelled glucose. Similar changes were seen in short-term labelling experiments with intact tubers attached to well-watered plants. Provided plants were grown with ample light and water, transformant tubers had a slightly lower water and sucrose content and a similar or even marginally higher starch content than wild-type tubers. (ii) When wild-type tuber slices were incubated with labelled glucose in the presence of mannitol to generate a moderate water deficit (between –0.12 and –0.72 MPa), there was a marked stimulation of sucrose synthesis and inhibition of starch synthesis. A similar stimulation was seen in labelling experiments with wild-type tubers that were attached to water-stressed wild-type plants. These changes were almost completely suppressed in transformants with a 70–80% reduction of SPS activity. (iii) Decreased irrigation led to an increase in the fraction of the dry-matter allocated to tubers in wild-type plants. This shift in allocation was prevented in transformants with reduced expression of SPS. (iv) The results show that operation of SPS and the sucrose cycle in growing potato tubers may lead to a marginal decrease in starch accumulation in non-stressed plants. However, SPS becomes a crucial factor in water-stressed plants because it is required for adaptive changes in tuber metabolism and whole plant allocation.     

Tissue water relations and leaf growth of tropical corn cultivars under water deficits

Abstract This study reports on the effect of water deficit on the tissue water relations and leaf growth of six corn cultivars, growing in glasshouse conditions, in order to understand growth responses to drought of tropical corn. A mild water-stress treatment was imposed slowly; plants reached a minimum pre-dawn leaf water potential of about –1.5 MPa by day 12 after watering was withheld. Analysis of the water relation characteristics of growing leaves using the pressure–volume technique demonstrated that under water deficits all the cultivars changed their moisture-release curves compared with irrigated plants. Osmotic potential at full turgor was lowered in water-stressed plants of all the genotypes and the degree of such change was between 0.34 MPa and 0.58 MPa. Thus, turgor pressure was lost at a lower water potential in water-stressed plants than in irrigated plants of all the varieties. Volumetric elastic moduli were also increased under water deficits and the increase ranged between 10% and 141% among the cultivars. In all the genotypes, the stress imposed led to a reduction of leaf area and dry matter accumulation. Leaf expansion was very sensitive to low turgor pressure and it ceased when turgor reached 0.2 MPa. Thus, varieties able to maintain a higher degree of turgor pressure (i.e. by osmotic adjustment) under water deficits may be able to prolong leaf growth.     

Contribution of adenosine 5′-diphosphoglucose pyrophosphorylase to the control of starch synthesis is decreased by water stress in growing potato tubers

Water stress stimulates sucrose synthesis and inhibits starch and cell-wall synthesis in tissue slices of growing potato (Solanum tuberosum L. cv. Desirée) tubers. Based on the analysis of fluxes and metabolites, Geigenberger et al. (1997, Planta 201: 502–518) proposed that water deficits up to −0.72 MPa stimulate sucrose synthesis, leading to decreased starch synthesis as a result of the resulting decline of phosphorylated metabolite levels, whereas more-severe water deficits directly inhibit the use of ADP-glucose. Potato plants with decreased expression of adenosine 5′-diphosphoglucose pyrophosphorylase (AGPase) have been used to test the prediction that the contribution of AGPase to the control of starch synthesis should decrease in severely water-stressed tuber material. Freshly cut slices from wild-type and antisense tubers were incubated at a range of mannitol concentrations (20, 300 and 500 mM) and the metabolism of [¹⁴C]glucose was analysed. A 86–97% reduction of AGPase activity led to a major but non-stoichiometric inhibition of starch accumulation in intact growing tubers attached to the plant (40–85%), and an inhibition of starch synthesis in non-stressed tuber slices incubated in 20 mM mannitol (60–80%). The inhibition of starch synthesis was accompanied by a 2- to 8-fold increase in the levels of sugars in intact tubers and a 2- to 3-fold stimulation of sucrose synthesis in tuber slices, whereas respiration and cell-wall synthesis were not significantly affected. The strong impact of AGPase on carbon partitioning in non-stressed tubers and tuber slices was retained in slices subjected to moderate water deficit (300 mM mannitol, corresponding to −0.72 MPa). In discs incubated in 500 mM mannitol (corresponding to −1.2 MPa) this response was modified. A 80–97% reduction of AGPase resulted in only a 0–40% inhibition of starch synthesis. Further, the water stress-induced stimulation of sucrose synthesis was abolished in the transformants. The results provide direct evidence that the contribution of AGPase to the control of starch synthesis can be modified by environmental factors, leading to a lower degree of control during severe water deficits. There was also a dramatic decrease in the labelling of cell-wall components in wild-type tuber slices incubated with 300 or 500 mM mannitol. The water stress-induced inhibition of cell-wall synthesis occurred independently of AGPase expression and the accompanying changes in starch and sucrose metabolism, indicating a direct inhibition of cell-wall synthesis in response to water stress. Received: 24 February 1999 / Accepted: 28 May 1999     

Rhythmic growth and carbon allocation in Quercus robur. Sucrose metabolizing enzymes in leaves

The high sucrose phosphate synthase (SPS) capacity and the low soluble acid invertase activity of mature leaves of the first flush of leaves remained stable during second flush development. Conversely, fluctuations of sucrose synthase (SS) activity were in parallel with the sucrose requirement of the second flush. Sucrose synthase activity (synthesis direction) in first flush leaves could increase in 'response' to sink demand constituted by the second flush growth. Only the ptotosynthates provided by flush mature leaves were translocated for a current flush, while the starch content of these leaves remained stable. After their emergence, second flush leaves showed an increase in SPS and SS (Synthetic direction) activities. The high sucrose synthesis in second flush leaves was used for leaf expansion. When young leaves were 30% fully expanded (stage II20), SPS activity showed little change whereas SS activity declined rapidly toward and after full leaf expansion. The starch accumulation in the young leaves occured simultaneously with their expansion. Developing leaves showed a high level of acid invertase activity until maximum leaf expansion (stage II1). In first and second flush leaves, changes in acid invertase activity correlated positively with changes in reducing sugar concentrations. Alkaline invertase and sucrose synthase (cleavage direction) activities showed similar changes with low values when compared with those of acid invertase activity, especially in second flush leaves. The present results suggest that soluble acid invertase was the primary enzyme responsible for sucrose catabolism in the expanding common oak leaf.     

水分胁迫对棉花叶片中IAA从含量、IAA氧化酶和过氧化物酶活性的影响

整株干旱降低盐棉46号叶片中的IAA总量,叶龄愈小下降愈多。幼叶中IAA总量的下降主要是结合态IAA减少的结果。气干和-1.7MPa PEG溶液渗透胁迫处理也降低离体成熟叶片的IAA总量,其变化与叶片含水量呈直线相关(r=0.905)。整株干旱处理提高各叶片的过氧化物酶活性,叶龄愈小提高愈多,但IAA氧化酶活性无显著变化。离体和整株干旱时IAA总量的下降,可能是过氧化物酶活性增加所致。     

The effect of wilting on the selection of leaves by the leaf-cutting ant Atta laevigata

Using field assays of leaf preference, we tested the hypothesis that wilting affects the selection of leaves by the leaf-cutting ant Atta laevigata (Fr. Smith). Detached leaves were left to air-dry until noticeably wilted. The area removed by the ants from wilted leaves was significantly greater than the area removed from fresh leaves, this effect being observed in several plant species, in leaves of different age, and in assays with different ant colonies. Leaves collected from water-stressed plants were also preferred to leaves from non-stressed plants. A. laevigata was found to employ a two-stage, size-related, strategy when cutting plants. Larger workers climbed the plant stem and dropped whole leaves to the ground by severing their petioles; smaller workers cut the lamina of the dropped leaves. The ants frequently left dropped leaves on the ground, until the next foraging day or even later, when they were harvested in a wilted condition in preference to newly-dropped leaves.It is possible that during wilting some repellent substances evaporate or become less effective, thus enhancing leaf palatability. Alternatively or in addition, changes in nutrient and water content may have rendered wilted leaves more palatable to leaf-cutting ants.     

Effects of low concentrations of O3, leaf age and water stress on leaf diffusive conductance and water use efficiency in soybean

Ozone, leaf age and water stress each affected leaf conductance in soybean [ Glycine max (L.) Merr. Hodgson], but there were no interactions among these factors. Exposure to increased concentrations of O₃ (0.01, 0.05, 0.09. and 0.13 μl l⁻¹) resulted in linear declines in abaxial and adaxial conductances in leaves of all ages. There were no differences in relative response to O₃ between the two leaf surfaces. For well-watered plants, water use efficiency also decreased with exposure to increased O₃ concentrations (water-stressed plants were not tested). Abaxial conductance increased as leaves aged from 4 to 10 days and then declined with further aging. Adaxial conductance decreased with all increases in leaf age beyond 4 days, and the ratio of abaxial/adaxial conductance increased continuously throughout the leaf lifespan. During water-stress cycles (water withheld for 2–3 days) leaves of water-stressed plants had lower conductances than those from well-watered plants, and there was no difference in relative response between abaxial and adaxial stomata.     

Influence of young potted apple tree water status on European Red Mite (Acari: Tetranychidae) abundance

The effect of water-stress on population growth of the European Red Mite (ERM) Panonychus ulmi (Koch) was tested in a plastic shelter using young potted apple trees under three water treatment. Mite abundance was significantly greater with higher water supply. The relationship between plant water status and mite number was non-linear, both treatments with leaf water potential from-30 to about -5 bars bearing the lower mite populations. Over the threshold value of-5 bars, mite number sharply increased four times. A non-linear regression analysis showed that the stomatal conductance explained 49.7% of the variance in mite number. Low leaf surface humidity resulting from reduced leaf transpiration of water-stressed trees may increase mite egg and larval mortalities. Daily leaf surface temperatures on the water-stressed trees averaged 1.5°C higher than on the other trees, but did not result in an increase in mite populations, because of the negative influence of low leaf humidity.     

Influence of Phosphorus Nutrition on Growth and Carbon Partitioning in Glycine max

Soybean plants (Glycine max [L.] Merr var Amsoy 71) were grown in growth chambers with high-phosphorus (high-P) and low-phosphorus (low-P) culture solutions. Low-P treatment reduced shoot growth significantly 7 days after treatment began. Root growth was much less affected by low-P, there being no significant reduction in root growth rate until 17 days had elapsed. The results suggest that low-P treatment decreased soybean growth primarily through an effect on the expansion of the leaf surface which was diminished by 85%, the main effect of low-P being on the rate of expansion of individual leaves. Low-P had a lesser effect on photosynthesis; light saturated photosynthetic rates at ambient and saturating CO₂ levels were lowered by 55 and 45%, respectively, after 19 days of low-P treatment. Low-P treatment increased starch concentrations in mature leaves, expanding leaves and fibrous roots; sucrose concentrations, however, were reduced by low-P in leaves and increased in roots. Foliar F-2,6-BP levels were not affected by P treatment in the light but in darkness they increased with high-P and decreased with low-P. The increase in the starch/sucrose ratio in low-P leaves was correlated primarily with changes in the total activities of enzymes of starch and sucrose metabolism.     

Osmotic Response of Sugar Beet Source Leaves at CO(2) Compensation Point

As sugar beet source leaves lowered the CO₂ concentration to compensation point in a closed atmosphere, leaf thickness and relative water content decreased. Leaf water potential declined rapidly from −0.5 to −1.4 megapascals. At 340 microliters CO₂ per liter, water potential and sucrose, glucose, and fructose contents were steady in photosynthesizing source leaves. Within 90 minutes after leaves were exposed to a CO₂ concentration at the compensation point, leaf sucrose content declined to 60% of the preteatment level, rapidly in the first 30 minutes and then more slowly. During the subsequent 200 minutes, sucrose content increased to 180% of pretreatment level. Glucose and fructose remained unchanged during the treatment. Degradation of starch was sufficient to account for the additional sucrose that accumulated. Labeled carbon lost from starch appeared in sucrose and several other compounds that likely contributed to the recovery in leaf water content.     

Diurnal Pattern of Translocation and Carbohydrate Metabolism in Source Leaves of Beta vulgaris L

Transitions in carbohydrate metabolism and translocation rate were studied for evidence of control of export by the sugar beet (Beta vulgaris L. Klein E.) source leaf. Steady-state labeling was carried out for two consecutive 14-hour light periods and various quantities related to translocation were measured throughout two 24-hour periods. Starch accumulation following illumination was delayed. Near the end of the light period, starch stopped accumulating, whereas photosynthesis rate and sucrose level remained unchanged. At the beginning of the dark period there was a 75-minute delay before starch was mobilized. The rate of import to the developing sink leaves at night was similar to that during the day, whereas export decreased considerably at night.

Starch accumulation and degradation seemed to be initiated in response to the level of illumination. Cessation of starch accumulation before the end of the light period was initiated endogenously. Exogenous control appeared to be mediated by the level of sucrose in the source leaf while endogenous control seemed to be keyed to photoperiod or photosynthetic duration.

     

Contributions of soluble carbohydrates to the osmotic adjustment in the C4 grass Setaria sphacelata: a comparison between rapidly and slowly imposed water stress

Photosynthetic carbohydrate content in Setaria sphacelata var. splendida under rapidly and slowly induced water deficit and its contribution to osmotic adjustment were studied. In short-term stress experiments, a decrease in the total content of sucrose (Su) and starch (St) was observed in leaf discs submitted to stress. An increase in the ratio between free hexoses and sucrose was found in stressed leaves, but no significant differences were found in the amount of free hexoses nor in the ratio between soluble and insoluble sugars. In long-term stress experiments, a higher amount of soluble sugars and a lower amount of starch were found in stressed leaves, when compared to the control. The ratios of free hexoses to sucrose and of soluble to insoluble sugars were also higher in stressed leaves. The contribution of the accumulation of soluble sugars to osmotic adjustment was absent in rapidly stressed leaves and was of minor importance in slowly stressed leaves.     

Proline Accumulation in Water-stressed Barley Leaves in Relation to Translocation and the Nitrogen Budget

Mobilization of N from leaves of barley (Hordeum vulgare L.) during water stress, and the role of proline as a mobilized species, were examined in plants at the three-leaf stage. The plants responded to water stress by withdrawing about 25% of the total reduced N from the leaf blades via phloem translocation. Most of this N loss was during the first 2 days while translocation of ¹⁴C-photosynthate out of the stressed blade still remained active. Free proline accumulation in the blade was initially slow, and became more rapid during the 2nd day of stress. Although a major free amino acid, proline accounted for only about 5% of the total N (soluble + insoluble) retained in severely stressed blades. When the translocation pathway in water-stressed leaves was interrupted just below the blade by a heat girdle, a cold jacket, or by blade excision, N loss from the blade was prevented and proline began to accumulate rapidly on 1st day of stress. Little free proline accumulated in the blades until after the ability to translocate was lost. Proline was, however, probably not a major species of N translocated during stress, because proline N accumulation in heat-girdled stressed leaves was five times slower than the rate of total N export from intact blades.     

Water-deficit stress effects on pistil biochemistry and leaf physiology in cotton (Gossypium hirsutum,L.)

Flowering in cotton (Gossypium hirsutum L.) is a sensitive stage to water-deficit stress, but the effects on metabolism are not well understood. The objective of this study was to monitor gas exchange responses of cotton plants under conditions of limited water supply and evaluate the effects on the carbohydrate concentrations and glutathione reductase levels in the cotton flower. Growth chamber experiments were conducted in 2008 and 2009, with normal day/night conditions of 32/24 °C and optimum quantities of Hoagland's nutrient solution applied until flowering. Treatments were imposed at flowering and consisted of control (Control), where optimum quantities of water were applied, and water stress (WS) where 50% of optimum quantity of water was supplied. Water-deficit stress resulted in a significant decrease in leaf stomatal conductance. Leaf photosynthetic and respiration rates were similarly decreased compared to the control. Ovary and style water potential of water-stressed leaves were significantly higher compared to the water potential of water stressed leaves, indicating that cotton flowers are fairly resistant to changes in the water status of the plant. However, carbohydrate concentrations of water-stressed pistils (ovary and style) were significantly increased compared to the control and a similar pattern was observed in the levels of glutathione reductase of water-stressed pistils. In conclusion, water-deficit stress during flowering resulted in significant decreases in leaf gas exchange functions as well as leaf water potential. Cotton pistils appeared to be less sensitive since they were able to maintain water potential similar to the control under limited water supply and increase glutathione reductase levels. However, pistil carbohydrate metabolism was significantly affected resulting in accumulation of both hexose and sucrose indicating a perturbation in sucrose cleaving and hexose utilizing enzymes that could potentially have as a consequence a decrease in fertilization and seed set efficiency.     

木薯蔗糖合成、转运与块根淀粉积累关系研究

该研究以淀粉含量不同的两个木薯品种(辐选01和华南124)为材料,通过测定各品种不同生育期叶、茎和根的蔗糖含量及块根淀粉含量,分析了蔗糖合成、转运和块根淀粉积累过程的相关性.结果表明:与华南124相比,在整个生育期内辐选01叶、茎的蔗糖含量均较高,块根蔗糖含量在块根膨大初期以前高于华南124,块根膨大初期以后则相反.在木薯的整个生育期,与辐选01相比,华南124的淀粉合成量和淀粉合成速率均较低.叶和茎蔗糖含量的变化规律与淀粉合成速率的变化规律相反,即块根淀粉积累明显加快时叶和茎的蔗糖含量略呈下降趋势,而块根淀粉合成减慢时叶、茎的蔗糖含量又开始上升.随着生育期的延后,块根蔗糖含量越来越低.在块根形成初期,蔗糖含量最高的组织部位为块根,其次为茎秆,最低的是叶片;而在块根成熟期时则相反,即蔗糖含量最高的部位是叶片,其次为茎杆,块根的蔗糖含量最小.相关性分析结果表明,木薯叶片蔗糖含量与块根淀粉含量呈显著的正相关;茎秆蔗糖含量与块根淀粉积累量呈不显著的正相关;块根蔗糖含量与淀粉积累量呈显著的负相关.由此可见,木薯叶、茎和根蔗糖与块根淀粉积累过程密切相关,其中叶片合成蔗糖的能力与块根利用蔗糖的能力在淀粉的积累过程中发挥关键作用.该研究结果为木薯的生产选育与高效栽培提供了理论依据.     

Changes in carbohydrate metabolism and assimilate export in starch-excess mutants of Arabidopsis

The aim of this work was to investigate the effects on carbohydrate metabolism of a reduction in the capacity to degrade leaf starch in Arabidopsis. The major roles of leaf starch are to provide carbon for sucrose synthesis, respiration and, in developing leaves, for biosynthesis and growth. Wild-type plants were compared with plants of a starch-excess mutant line (sex4) deficient in a chloroplastic isoform of endoamylase. This mutant has a reduced capacity for starch degradation, leading to an imbalance between starch synthesis and degradation and the gradual accretion of starch as the leaves age. During the night the conversion of starch into sucrose in the mutant is impaired; the leaves of the mutant contained less sucrose than those of the wild type and there was less movement of ¹⁴C-label from starch to sucrose in radio-labelling experiments. Furthermore, the rate of assimilate export to the roots during the night was reduced in the mutant compared with the wild type. During the day however, photosynthetic partitioning was altered in the mutant, with less photosynthate partitioned into starch and more into sugars. Although the sucrose content of the leaves of the mutant was similar to the wild type during the day, the rate of export of sucrose to the roots was increased more than two-fold. The changes in carbohydrate metabolism in the mutant leaves during the day compensate partly for its reduced capacity to synthesize sucrose from starch during the night.     

Short-term water stress leads to a stimulation of sucrose synthesis by activating sucrose-phosphate synthase

The aim of this work was to identify which aspects of photosynthetic metabolism respond most sensitively to leaf water deficit. Spinach (Spinacia oleracea L.) leaf discs were floated on sorbitol concentrations of increasing molarity and changes of the protoplast volume were estimated using [¹⁴C]sorbitol and ³H₂O penetration. Detached leaves were also wilted until 10% of their fresh weight was lost. Photosynthesis was studied at very high external CO₂ concentrations, to eliminate the effect of closing stomata. There was no large inhibition of CO₂ fixation after wilting leaves, or until the external water deficit was greater than-1.2 MPa. However, partitioning changed markedly at these moderate water deficits: more sucrose and less starch was made. When an inhibition of CO₂-saturated photosynthesis did appear at a water deficit of-2.0 MPa and above, measurements of chlorophyll-fluorescence quenching and metabolite levels showed the thylakoid reactions were not especially susceptible to short-term water stress. The inhibition was accompanied by a small increase of the triose phosphate: ribulose-1,5-bisphosphate ratio, showing regeneration of ribulose-1,5-bisphosphate was affected. However, there was also a general increase of the estimated concentrations of most metabolites, indicating that there is no specific site for the inhibition of photosynthesis. Increasing water deficit led to a large increase of fructose-2,6-bisphosphate. This is explained in terms of a simultaneous increase of fructose-6-phosphate and inorganic phosphate as the cell shrinks. The high fructose-2,6-bisphosphate led to the accumulation of triose phosphates, and the potential significance of this for protection against photoinhibition is discussed. There was an increase in the extractable activity of sucrose-phosphate synthase. This was only detected when the enzyme was assayed in conditions which distinguish between different kinetic forms which have previously been identified in spinach leaves. It is proposed that activation of sucrose-phosphate synthase is one of the first sites at which spinach leaves respond to a rising water deficit. This could be of importance for osmoregulation.Abbreviations Chl chlorophyll - Fru1,6bisP fructose-1,6-bisphosphate - Fru2,6bisP fructose-2,6-bisphosphate - Fru6P fructose-6-phosphate - Glc6P glucose-6-phosphate - PGA glycerate-3-phosphate - Pi inorgamic phosphate - Ru1,5bisP ribulose-1,5-bisphosphate - SPS sucrose-phosphate synthase - triose-P sum of glyceraldehyde-3-phosphate and dehydroxyacetone phosphate - UDPGlc uridine diphosphoglucose     

Up-regulation of sucrose phosphate synthase in rice grown under elevated CO2 and temperature

Rice (Oryza sativa L. cv. IR-30) was grown season-long in outdoor, controlled-environment chambers at 33 Pa CO₂ with day/night/paddy-water temperatures of 28/21/25 °C, and at 66 Pa CO₂ with five different day/night/paddy-water temperature regimes (25/18/21, 28/21/25, 31/24/28, 34/27/31 and 37/30/34 °C). Sucrose phosphate synthase (SPS) activities in leaf extracts at 21, 48 and 81 days after planting (DAP) were assayed under saturating and selective (limiting) conditions. Diel SPS activity data indicated that rice SPS was light regulated; with up to 2.2-fold higher rates during the day. Throughout the growth season, leaf SPS activities were up-regulated in the CO₂-enriched plants, averaging 20 and 12% higher than in ambient-CO₂ grown plants in selective and saturating assays, respectively. Similarly, SPS activities increased 2.4% for each 1 °C rise in growth temperature from 25 to 34 °C, but de creased 11.5% at 37 °C. Leaf sucrose content was higher, and mirrored SPS activity better, than starch, although starch was more responsive to CO₂ treatment. Leaf sucrose and starch contents were significantly higher throughout the season in plants at elevated CO₂, but the N content averaged 6.5% lower. Increasing growth temperatures from 25 to 37 °C caused a linear decrease (62%) in leaf starch content, but not in sucrose. Consequently, the starch:sucrose ratio declined with growth temperature. The data are consistent with the hypothesis that the up-regulation of leaf SPS may be an acclimation response of rice to optimize the utilization and export of organic-C with the increased rates of inorganic-C fixation in elevated CO₂ or temperature growth regimes.