Photosynthesis and Transpiration as a Function of Gaseous Diffusive Resistances in Orange Leaves

The CO₂ and H₂O vapour exchange of single attached orange, Citrus sinensis (L.), leaves was measured under laboratory conditions using infrared gas analysis. Gaseous diffusive resistances were derived from measurements at a saturating irradiance and at a leaf temperature optimum for photosynthesis. Variation in leaf resistance (within the range 1.6 to 60 s cm^-1) induced by moisture status, or by cyclic oscillations in stomatal aperture, was associated with changes in both photosynthesis and transpiration. At low leaf resistance (ri less than 10 s cm^-1) the ratio of transpiration to photosynthesis declined with reduced stomatal aperture, indicating a tighter stomatal control over H₂O vapour loss than over CO₂ assimilation. At higher leaf resistance (ri greater than 10 s cm^-1) changes in transpiration and photosynthesis were linearly related, but leaf resistance and mesophyll resistance were also positively correlated, so that strictly stomatal control of photosynthesis became more apparent than real. This evidence, combined with direct measurements of CO₂ diffusive resistances (in a -O₂ gas stream) emphasised the presence of a significant mesophyll resistance; i.e., an additional and rate limiting resistance to CO₂ assimilation over and above that encountered by H₂O vapour escaping from the leaf.     

Oxygen Inhibition of Photosynthesis: I. Temperature Dependence and Relation to O(2)/CO(2) Solubility Ratio

The magnitude of the percentage inhibition of photosynthesis by atmospheric levels of O₂ in the C₃ species Solanum tuberosum L., Medicago sativa L., Phaseolus vulgaris L., Glycine max L., and Triticum aestivum L. increases in a similar manner with an increase in the apparent solubility ratio of O₂/CO₂ in the leaf over a range of solubility ratios from 25 to 45. The solubility ratio is based on calculated levels of O₂ and CO₂ in the intercellular spaces of leaves as derived from whole leaf measurements of photosynthesis and transpiration. The solubility ratio of O₂/CO₂ can be increased by increased leaf temperature under constant atmospheric levels of O₂ and CO₂ (since O₂ is relatively more soluble than CO₂ with increasing temperature); by increasing the relative levels of O₂/CO₂ in the atmosphere at a given leaf temperature, or by increased stomatal resistance. If the solubility ratio of O₂/CO₂ is kept constant, as leaf temperature is increased, by varying the levels of O₂ or CO₂ in the atmosphere, then the percentage inhibition of photosynthesis by O₂ is similar. The decreased solubility of CO₂ relative to O₂ (decreased CO₂/O₂ ratio) may be partly responsible for the increased percentage inhibition of photosynthesis by O₂ under atmospheric conditions with increasing temperature.     

Effects of sulfur on the photosynthesis of intact leaves and isolated chloroplasts of sugar beets

Effects of sulfur on photosynthesis in sugar beets (Beta vulgaris L. cv. F58-554H1) were studied by inducing sulfur deficiency and determining changes in the photosynthesis of whole attached leaves and of isolated chloroplasts. The rates of photosynthetic CO₂ uptake by intact leaves, photoreduction of ferricyanide, cyclic and noncyclic photophosphorylation of isolated chloroplasts, and the rate of CO₂ assimilation by ribulose diphosphate carboxylase, decreased with decrease in total leaf sulfur from 2500 to about 500 μg g⁻¹ dry weight. Sulfur deficiency reduced photosynthesis through an effect on chlorophyll content, which decreased linearly with leaf sulfur, and by decreasing the rate of photosynthesis per unit chlorophyll. There was only a small effect of sulfur deficiency on stomatal diffusion resistance to CO₂ until leaf sulfur decreased below 1000 μg g⁻¹ when stomatal resistance became a more significant proportion of the total diffusion resistance to CO₂. Light respiration rates were positively correlated with photosynthesis rates and dark respiration was unchanged as leaf sulfur concentrations declined.     

Metabolism of <Superscript>14</Superscript>C-glycine as a substrate for photorespiration of the leaf at different developmental stages of ephemeroides

The metabolism of ¹⁴C-glycine (a substrate for photorespiration) was studied in the light and in darkness under natural CO₂ concentration (0.03%) in the leaves of ephemeroides Scilla sibirica Haw. and Ficaria verna Huds. at different developmental stages. Using one and the same sample, potential photosynthesis (at 1% CO₂), true photosynthesis (at 0.03% CO₂), and leaf respiratory capacity were measured by the radiometric and manometric methods, respectively. All measurements were performed at 15°C, an average temperature during ephemer growth. It was found that, in the white zone of the Scilla leaf, the rate of CO₂ evolution resulting from metabolization of exogenous ¹⁴C-glycine was similar in the light and in darkness. In the green zone of the Scilla leaf and in the green leaf of Ficaria, both ¹⁴C-glycine absorption and ¹⁴CO₂ evolution were lower in the light as compared with darkness, which is explained by CO₂ reassimilation. In all treatments of both plant species, a specific inhibitor of glycine decarboxylase complex (GDC), aminoacetonitrile (5 mM) suppressed CO₂ evolution by 20–40%. It was concluded that in ephemeroides mitochondrial GDC, responsible for CO₂ evolution in photorespiration, is formed at the earliest stage of leaf development. This indicates that photorespiration can occur simultaneously with the development of the leaf photosynthetic activity. On the basis of the assumption that carbon losses in the form of CO₂ evolved during photorespiration comprise 25% of true photosynthesis, it was calculated that, in ephemer leaves, the highest rates of photorespiration and photosynthesis were attained during flowering when the leaf area was the largest and the rate of dark respiration was reduced by 1.5–2.0 times. The highest rates of dark respiration were observed in the beginning of growth. In senescing leaves by the end of the plant vegetation, potential photosynthesis and true photosynthesis were reduced, whereas dark respiration remained essentially unchanged. It is concluded that the high rates of potential and true photosynthesis are characteristic of ephemeroides when they complete their short developmental program in early spring (at 15°C); theoretically, photorespiration also occurs at a high rate during this period, when this process provides for a defense against the threat of photoinhibition at low temperature and high insolation.     

Changes in photosynthetic characteristics during leaf development in apple

A comprehensive developmental survey of leaf area, chlorophyll, photosynthetic rate, leaf resistance, transpiration ratio, CO₂ compensation point and photorespiration was conducted in apple. The largest changes in each of the photosynthetic characteristics studied took place during the earliest stages of leaf development, coinciding with the period of greatest leaf expansion and chlorophyll synthesis. During early development, photosynthesis increased 5-fold, reaching a maximum rate of 40 mg CO₂ dm^-2 hr^-1 at a leaf plastochron index (LPI) of 10. During this same period, leaf resistance, transpiration ratio, CO₂ compensation point and mesophyll resistance decreased, while carboxylation efficiency increased. Two especially interesting aspects of the data discussed are simultaneous changes that occur at a LPI of 10 and 12 in all of the photosynthetic characteristics examined and an apparent decrease in photorespiration as leaves age. From our results it is clear that stage of leaf development is an important factor affecting the rate of photosynthesis and photorespiration.Scientific Paper No. 5687, College of Agriculture, Washington State University, Pullman. This work is supported by the National Science Foundation Grant 80-10958 and the Columbia River Orchards Foundation.     

Decrease in net photosynthesis caused by respiration

CO₂ exchange between air and leaf is conceived as a current along a resistor with a respiratory current, unaffected by CO₂ concentration, entering the resistor. The conclusion follows that a plant placed in an atmosphere free of CO₂ will increase the CO₂ concentration to the compensation concentration along a curve determined by the resistor and the volume of the atmosphere. This was verified. Also a photosynthesis rate calculated from the parameters of the observed curve agreed with an independent observation of photosynthesis in CO₂-free air. The decrease in net photosynthesis caused by respiration is, according to the model, the CO₂ compensation concentration divided by the concentration in the atmosphere.     

Effects of atmospheric CO2 enrichment and foliar methanol application on net photosynthesis of sour orange tree (Citrus aurantium; Rutaceae) leaves

Foliar spray applications of 40% aqueous methanol were made to sunlit leaves of sour orange trees that had been grown continuously in clear-plastic-wall open-top enclosures maintained out-of-doors at Phoenix, Arizona, for over 5.5 years in ambient air of approximately 400 μmol mol^-1 CO₂ and in air enriched with CO₂ to a concentration of approximately 700 μmol mol^-1. No unambiguous effects of the methanol applications were detected in net photosynthesis measurements made on foliage in either of the two CO₂ treatments. The 75% increase in CO₂, however, raised the upper-limiting leaf temperature for positive net photosynthesis by approximately 7 C, which resulted in a 75% enhancement in net photosynthesis at a leaf temperature of 31 C, a 100% enhancement at a leaf temperature of 35 C, and a 200% enhancement at 42 C.     

A model describing photosynthesis in terms of gas diffusion and enzyme kinetics

Summary A model predicting net photosynthesis of individual plant leaves for a variety of environmental conditions has been developed. It is based on an electrical analogue describing gas diffusion from the free atmosphere to the sites of CO₂ fixation and a Michaelis-Menten equation describing CO₂ fixation. The model is presented in two versions, a simplified form without respiration and a more complex form including respiration. Both versions include terms for light and temperature dependence of CO₂ fixation and light control of stomatal resistance. The second version also includes terms for temperature, light, and oxygen dependence of respiration and O₂ dependence of CO₂ fixation.The model is illustrated with curves based on representative values of the various environmental and biological parameters. These curves relate net photosynthesis to light intensity, [CO₂], [O₂], temperature, and resistances to CO₂ uptake. The shape of the [CO₂]-net photosynthesis curves depends on the total diffusion resistance to CO₂ uptake and the Michaelis constant for CO₂ uptake. The curves range from typical Michaelis-Menten to Blackman types.The model is combined with a model of leaf energy exchange permitting simultaneous estimation of net photosynthesis and transpiration. The combined model is illustrated with curves relating transpiration to photosynthesis under a wide variety of environmental conditions. Environmental regimes yielding maximum efficiency of water use are identified for the given assumptions and biological parameters.     

Induced leaf intercellular CO2, photosynthesis, potassium and nitrate retention and strawberry early fruit formation under macronutrient limitation

Relationships between induced high leaf intercellular CO₂ concentrations, leaf K⁺ and NO₃ ^? ion movement and early fruit formation under macronutrient limitation are not well understood. We examined the effects and interactions of reduced K/N input treatments on leaf intercellular CO₂, photosynthesis rate, carboxylation and water use efficiency, berry formation as well as leaf/fruit K⁺, NO₃ ^? and photosynthate retention of strawberry (Fragaria × ananassa Duch.) to enhance low-input agriculture. The field study was conducted in Nova Scotia, eastern Canada during 2009–2010. The experimental treatments consisted of five K₂O rates (0, 6, 12, 18, and 24 kg ha^?1) and five N rates (0, 5, 10, 15, and 20 kg ha^?1), representing respectively, 0, 25, 50, 75, and 100 % of regular macronutrient recommendations based on the soil testing. The treatments were arranged in a split-plot design with three blocks in the field. The cultivar was ‘Mira’, a June-bearing crop. The results showed that strawberry plants treated with 25 %-reduced inputs could induce significantly higher leaf intercellular CO₂ concentrations to improve plant photosynthesis, carboxylation and water use efficiency and translocation of leaf/fruit K⁺ and dissolved solids, which could advance berry formation by 6 days and produce significantly higher marketable yields (P < 0.05). Higher leaf intercellular CO₂ inhibited leaf/fruit NO₃ ^? ion retention, but this inhibition did not occur in leaf/fruit K⁺ retention. Linear interactions of the K/N treatments were significant on fruit marketable yields, intercellular CO₂, net photosynthesis, leaf transpiration rates, and leaf temperatures (P < 0.05). It was concluded that higher leaf CO₂ could enhance plant photosynthesis, promote plant carboxylation and water use efficiency, and advance berry formation, but it could inhibit leaf NO₃ ^? retention. This inhibition did not find in leaf K⁺ ion and dissolved solid retention. Overlay co-limitation of leaf intercellular CO₂ and translocation of leaf/fruit K⁺/NO₃ ^? and total dissolved solids could constrain more fruit formation attributes under full macronutrient supply than reduced inputs. It was suggested that low input would be an optimal and sustainable option for improving small fruit crop physiological development and dealing with macronutrient deficiency challenge.     

Changes in the Water Balance and Photosynthesis of Onion, Bean and Cotton Plants under Saline Conditions

The osmotic concentration (osmotic potential) of onion leaf sap did not adjust to chloride salinity, and consequently water potential, turgor, stomatal aperture and transpiration were reduced. Although osmotic concentration of bean and cotton leaf sap did adjust to a saline root medium and turgor was no less in the salinized plants than in the controls, stomata of the salinized plants remained only partly open and transpiration was reduced. Net photosynthesis of onion plants was reduced by salinity (this effect being much enhanced in a hot dry atmosphere) but it could be rapidly raised to the level of the controls by inducing elevated leaf turgor. Stomatal closure was initially responsible for most of the ～30 % reduction in photosynthesis of salinized beans. This was due to interference with CO₂ diffusion and could be overcome by raising the CO₂ concentration in the air. At a later stage of growth, salinity affected the light reaction of bean photosynthesis, and elevation of the air CO₂ had little effect. Closure of stomata of salinized cotton plants had only a relatively small effect on net photosynthesis. Light intensity and CO₂ concentration experiments showed that salinity was reducing the photosynthesis of cotton leaves mainly by affecting the light reaction of photosynthesis. It is concluded that chloride salinity does affect the water balance and rate of photosynthesis of plants and that the nature and degree of the effects will depend upon climatic conditions and may be very different between plant species and in the same species at different periods of growth.     

Photosynthetic characteristics of Amaranthus tricolor,a C4 tropical leafy vegetable

The gas exchange characteristics are reported for Amaranthus tricolor, a C₄ vegetable amaranth of southeastern Asia. Maximum photosynthetic capacity was 48.3±1.0μmol CO₂ m^?2s^?1 and the temperature optimum was 35°C. The calculated intercellular CO₂ concentration at this leaf temperature and an incident photon flux (400–700 mm) of 2 mmol m^?2s^?1 averaged 208±14 μl l^?1, abnormally high for a C₄ species. The photosynthetic rate, intercellular CO₂ concentration, and leaf conductance all decreased with an increase in water vapor pressure deficit. However, the decrease in leaf conductance which resulted in a decrease in intercellular CO₂ concentration accounted for only one fourth of the observed decrease in photosynthetic rate as water vapor pressure deficit was increased. Subsequent measurements indicated that the depence of net photosynthesis on intercellular CO₂ concetration changed with water vapor pressure deficit.     

Growth and physiological responses of cotton (Gossypium hirsutum L.) to elevated carbon dioxide and ultraviolet-B radiation under controlled environmental conditions

Better understanding of crop responses to projected changes in climate is an important requirement. An experiment was conducted in sunlit, controlled environment chambers known as soil–plant–atmosphere–research units to determine the interactive effects of atmospheric carbon dioxide concentration [CO₂] and ultraviolet‐B (UV‐B) radiation on cotton (Gossypium hirsutum L.) growth, development and leaf photosynthetic characteristics. Six treatments were used, comprising two levels of [CO₂] (360 and 720 µmol mol^?1) and three levels of 0 (control), 7.7 and 15.1 kJ m^?2 d^?1 biologically effective UV‐B radiations within each CO₂ level. Treatments were imposed for 66 d from emergence until 3 weeks after the first flower stage. Plants grown in elevated [CO₂] had greater leaf area and higher leaf photosynthesis, non‐structural carbohydrates, and total biomass than plants in ambient [CO₂]. Neither dry matter partitioning among plant organs nor pigment concentrations was affected by elevated [CO₂]. On the other hand, high UV‐B (15.1 kJ m^?2 d^?1) radiation treatment altered growth resulting in shorter stem and branch lengths and smaller leaf area. Shorter plants at high UV‐B radiation were related to internode lengths rather than the number of mainstem nodes. Fruit dry matter accumulation was most sensitive to UV‐B radiation due to fruit abscission. Even under 7.7 kJ m^?2 d^?1 of UV‐B radiation, fruit dry weight was significantly lower than the control although total biomass and leaf photosynthesis did not differ from the control. The UV‐B radiation of 15.1 kJ m^?2 d^?1 reduced both total (43%) and fruit (88%) dry weights due to smaller leaf area and lower leaf net photosynthesis. Elevated [CO₂] did not ameliorate the adverse effects of UV‐B radiation on cotton growth and physiology, particularly the boll retention under UV‐B stress.     

Scaling carbon dioxide and water vapour exchange from leaf to canopy in a deciduous forest. II. Model testing and application

The scaling of CO₂ and water vapour transfer from leaf to canopy dimensions was achieved by integrating mechanistic models for physiological (photosynthesis, stomatal conductance and soil/root and bole respiration) and micrometeorological (radiative transfer, turbulent transfer and surface energy exchanges) processes. The main objectives of this paper are to describe a canopy photosynthesis and evaporation model for a temperate broadleaf forest and to test it against field measurements. The other goal of this paper is to use the validated model to address some contemporary ecological and physiological questions concerning the transfer of carbon and water between forest canopies and the atmosphere. In particular, we examine the role of simple versus complex radiative transfer models and the effect of environmental (solar radiation and CO₂) and ecophysiological (photo-synthetic capacity) variables on canopy-scale carbon and water vapour fluxes.     

Photosynthetic and stomatal responses of the halophyte,Plantago maritima L. to fluctuations in salinity

Abstract Measurements of photosynthesis as a function of intercellular CO₂ (A-C₁ curve) were made on single. attached leaves of Plantago maritima L. while plants were exposed to changes in salinity. Salinity was increased in steps from 50 to 500 mol m^-3 NaCl and then returned to 50 mol m^-3 NaCl at two rates, 75 mol m^-3 (NaCl) day^-1 (experiment 1) and 150 mol m^-3 (NaCl) day^-1 (experiment 2). In experiment one, the CO₂ assimilation rate declined at high CO₂ concentrations, but the initial slope of the A-C₁ curve was unaffected in young leaves after salinity was increased to 500 mol m^-3 NaCl. The insensitivity of photosynthesis to increases in CO₂ concentration above air levels was not associated with insensitivity to a reduction in oxygen concentration. In experiment two increasing the rate at which salinity was changed resulted in larger declines in photosynthesis and leaf conductance than were observed in experiment one. Both the initial slope and the CO₂ saturated region of the A-C₁ curve were substantially reduced at high salinity suggesting that mesophyll biochemical capacity had been inhibited. However, concurrent measurements of photosynthesis as oxygen evolution under 5% CO₂ indicated no effect of increased salinity on photosynthetic capacity. This suggests that the apparent non-stomatal limitations indicated by A-C₁ measurements were artifacts caused by strong, nonuniform stomatal closure.     

Photosynthetic characteristics of Amaranthus tricolor,a C4 tropical leafy vegetable

The gas exchange characteristics are reported for Amaranthus tricolor, a C₄ vegetable amaranth of southeastern Asia. Maximum photosynthetic capacity was 48.3±1.0 μmol CO₂ m^-2 s^-1 and the temperature optimum was 35°C. The calculated intercellular CO₂ concentration at this leaf temperature and an incident photon flux (400–700 mm) of 2 mmol m^-2 s^-1 averaged 208±14 μl l^-1, abnormally high for a C₄ species. The photosynthetic rate, intercellular CO₂ concentration, and leaf conductance all decreased with an increase in water vapor pressure deficit. However, the decrease in leaf conductance which resulted in a decrease in intercellular CO₂ concentration accounted for only one fourth of the observed decrease in photosynthetic rate as water vapor pressure deficit was increased. Subsequent measurements indicated that the dependence of net photosynthesis on intercellular CO₂ concentration changed with water vapor pressure deficit.     

Temporal expression of effects of varying nitrogen supply on canopy growth, photosynthesis and fruit production for Actinidia deliciosa vines in the field

The effects of varying nitrogen supply on canopy leaf area, response of leaf net photosynthesis (A_n) to quantum flux density (Q), and fruit yields of kiwifruit vines (Actinidia deliciosa var. deliciosa) were examined in a two-year field experiment. Vines were grown with 0, 250 or 750 kg N ha^?1 year^?1. The responses to nitrogen supply were compared with responses to shade, to examine the impact of reduced carbon assimilation on canopy leaf area and fruit yields. Nitrogen supply did not affect significantly any of the measured variables during the first season of the experiment. In the second season, canopy leaf area was reduced significantly where nitrogen supply was limited. The quantum efficiency of photosynthesis (φ_q) increased from 0. 03 mol CO₂ mol^?1 Q soon after leaf emergence to more than 0. 05 mol CO₂ mol^?1 Q during the middle of the growing season. The quantum saturated rate of A_n (A_sat) also increased during the season, from 7–10 μmol CO₂ m^?2 s^?1 soon after leaf emergence, to 15–20 (μmol CO₂ m^?2 s^?1 during the middle of the growing season. φ_q and A_sat increased significantly with nitrogen supply at all measurement times during the second season. For vines with high nitrogen, fruit yields in both seasons were similar, averaging 3. 05 kg m^?2. Fruit yields in the second season were reduced significantly where nitrogen supply was limited, due to reduced fruit numbers. The relative effects of reduced leaf area and reduced leaf photosynthesis for carbon assimilation by nitrogen deficient vines were examined using a mathematical model of canopy photosynthesis for kiwifruit vines. Simulations of canopy photosynthesis indicated that effects on leaf area and on leaf photosynthesis were of similar importance in the overall effects of nitrogen deficiency on carbon assimilation. The effects of nitrogen supply on fruit numbers (i. e. flower development) preceded the measured effects on carbon assimilation, indicating that the nitrogen supply affected carbon partitioning to reserves in the first season.     

Contribution of stem CO2 fixation to whole-plant carbon balance in nonsucculent species

In many plant species that remain leafless part of the year, CO₂ fixation occurring in green stems represents an important carbon gain. Traditionally, a distinction has been made between stem photosynthesis and corticular photosynthesis. All stem photosynthesis is, sensu stricto, cortical, since it is carried out largely by the stem cortex. We proposed the following nomenclature: stem net photosynthesis (SNP), which includes net CO₂ fixation by stems with stomata in the epidermis and net corticular CO₂ fixation in suberized stems, and stem recycling photosynthesis (SRP), which defines CO₂ ling in suberized stems. The proposed terms should reflect differences in anatomical and physiological traits. SNP takes place in the chlorenchyma below the epidermis with stomata, where the net CO₂ uptake occurs, and it resembles leaf photosynthesis in many characteristics. SRP is found in species where the chlorenchyma is beneath a well-developed stomata-free periderm and where reassimilation of internally respired CO₂ occurs. SNP is common in plants from desert ecosystems, rates reaching up to 60% of the leaf photosynthetic rate. SRP has been demonstrated in trees from temperate forests and it offsets partially a carbon loss by respiration of stem nonphotosynthetic tissues. Reassimilation can vary between 7 and 123% of respired CO₂, the latter figure implying net CO₂ uptake from the atmosphere. Both types of stem photosynthesis contribute positively to the carbon economy of the species, in which they occur; they are advantageous to the plant because they allow the maintenance of physiological activity during stress, an increase of integrated water use efficiency, and they provide the carbon source used in the production of new organs.     

Nonstomatal inhibition of photosynthesis in sunflower at low leaf water potentials and high light intensities

The inhibition of photosynthesis at low leaf water potentials was studied in soil-grown sunflower to determine the degree to which photosynthesis under high light was affected by stomatal and nonstomatal factors. Below leaf water potentials of −11 to −12 bars, rates of photosynthesis at high light intensities were insensitive to external concentrations of CO₂ between 200 and 400 microliters per liter. Photosynthesis also was largely insensitive to leaf temperature between 10 and 30 C. Changes in CO₂ concentration and temperature had negligible effect on leaf diffusive resistance. The lack of CO₂ and temperature response for both photosynthesis and leaf diffuse resistance indicates that rates of photosynthesis were not limited by either CO₂ diffusion or a photosynthetic enzyme. It was concluded that photosynthesis under high light was probably limited by reduced photochemical activity of the leaves at water potentials below −11 to −12 bars.     

Long-term carbon exchange in a sparse, seasonally dry tussock grassland

Rainfall and its seasonal distribution can alter carbon dioxide (CO₂) exchange and the sustainability of grassland ecosystems. Using eddy covariance, CO₂ exchange between the atmosphere and a sparse grassland was measured for 2 years at Twizel, New Zealand. The years had contrasting distributions of rain and falls (446 mm followed by 933 mm; long‐term mean=646 mm). The vegetation was sparse with total above‐ground biomass of only 1410 g m^?2. During the dry year, leaf area index peaked in spring (November) at 0.7, but it was <0.2 by early summer. The maximum daily net CO₂ uptake rate was only 1.5 g C m^?2 day^?1, and it occurred before mid‐summer in both years. On an annual basis, for the dry year, 9 g C m^?2 was lost to the atmosphere. During the wet year, 41 g C m^?2 was sequestered from the atmosphere. The net exchange rates were determined mostly by the timing and intensity of spring rainfall. The components of ecosystem respiration were measured using chambers. Combining scaled‐up measurements with the eddy CO₂ effluxes, it was estimated that 85% of ecosystem respiration emanated from the soil surface. Under well‐watered conditions, 26% of the soil surface CO₂ efflux came from soil microbial activity. Rates of soil microbial CO₂ production and net mineral‐N production were low and indicative of substrate limitation. Soil respiration declined by a factor of four as the soil water content declined from field capacity (0.21 m³ m^?3) to the driest value obtained (0.04 m³ m^?3). Rainfall after periods of drought resulted in large, but short‐lived, respiration pulses that were curvilinearly related to the increase in root‐zone water content. Coupled with the low leaf area and high root : shoot ratio, this sparse grassland had a limited capacity to sequester and store carbon. Assuming a proportionality between carbon gain and rainfall during the summer, rainfall distribution statistics suggest that the ecosystem is sustainable in the long term.