CO2 and water vapour exchange in four alpine herbs at two altitudes and under varying light and temperature conditions

CO₂ and water vapour exchange rates of four alpine herbs namely: Rheum emodi, R. moorcroftianum, Megacarpaea polyandra and Rumex nepalensis were studied under field conditions at 3600 m (natural habitat) and 550 m altitudes. The effect of light and temperature on CO₂ and water vapour exchange was studied in the plants grown at lower altitude. In R. moorcroftianum and R. nepalensis, the average photosynthesis rates were found to be about three times higher at 550 m as compared to that under their natural habitat. However, in M. polyandra, the CO₂ exchange rates were two times higher at 3600 m than at 550 m but in R. emodi, there were virtually no differences at the two altitudes. These results indicate the variations in the CO₂ exchange rates are species specific. The change in growth altitude does not affect this process uniformly.The transpiration rates in R. emodi and M. polyandra were found to be very high at 3600 m compared to 550 m and are attributed to overall higher stomatal conductance in plants of these species, grown at higher altitude. The mid-day closure of stomata and therefore, restriction of transpirational losses of water were observed in all the species at 550 m altitude. In addition to the effect of temperature and relative humidity, the data also indicate some endogenous rhythmic control of stomatal conductance.The temperature optima for photosynthesis was close to 30°C in M. polyandra and around 20°C in the rest of the three species. High temperature and high light intensity, as well as low temperature and high light intensity, adversely affect the net rate of photosynthesis in these species.Both light compensation point and dark respiration rate increased with increasing temperature.The effect of light was more prominent on photosynthesis than the effect of temperature, however, on transpiration the effect of temperature was more prominent than the effect of light intensity.No definite trends were found in stomatal conductance with respect to light and temperature. Generally, the stomatal conductance was highest at 20°C.The study reveals that all these species can easily be cultivated at relatively lower altitudes. However, proper agronomical methodology will need to be developed for better yields.     

Daily and seasonal CO2 exchange in Scots pine grown under elevated O3 and CO2: experiment and simulation

Starting in early spring of 1994, naturally regenerated, 30-year-old Scots pine (Pinus sylvestris L.) trees were grown in open-top chambers and exposed in situ to doubled ambient O₃,doubled ambient CO₂ and a combination of O₃ and CO₂ from 15 April to 15 September. To investigate daily and seasonal responses of CO₂ exchange to elevated O₃ and CO₂, the CO₂ exchange of shoots was measured continuously by an automatic system for measuring gas exchange during the course of one year (from 1 Januray to 31 December 1996). A process-based model of shoot photosynthesis was constructed to quantify modifications in the intrinsic capacity of photosynthesis and stomatal conductance by simulating the daily CO₂ exchange data from the field. Results showed that on most days of the year the model simulated well the daily course of shoot photosynthesis. Elevated O₃ significantly decreased photosynthetic capacity and stomatal conductance during the whole photosynthetic period. Elevated O₃ also led to a delay in onset of photosynthetic recovery in early spring and an increase in the sensitivity of photosynthesis to environmental stress conditions. The combination of elevated O₃ and CO₂ had an effect on photosynthesis and stomatal conductance similar to that of elevated O₃ alone, but significantly reduced the O₃-induced depression of photosynthesis. Elevated CO₂ significantly increased the photosynthetic capacity of Scots pine during the main growing season but slightly decreased it in early spring and late autumn. The model calculation showed that, compared to the control treatment, elevated O₃ alone and the combination of elevated O₃ and CO₂ decreased the annual total of net photosynthesis per unit leaf area by 55% and 38%, respectively. Elevated CO₂ increased the annual total of net photosynthesis by 13%.     

Leaf cavity CO2 concentrations and CO2 exchange in onion,Allium cepa L.

Onion (Allium cepa L.) plants were examined to determine the photosynthetic role of CO₂ that accumulates within their leaf cavities. Leaf cavity CO₂ concentrations ranged from 2250 L L^–1 near the leaf base to below atmospheric (<350 L L^–1) near the leaf tip at midday. There was a daily fluctuation in the leaf cavity CO₂ concentrations with minimum values near midday and maximum values at night. Conductance to CO₂ from the leaf cavity ranged from 24 to 202 mol m^–2 s^–1 and was even lower for membranes of bulb scales. The capacity for onion leaves to recycle leaf cavity CO₂ was poor, only 0.2 to 2.2% of leaf photosynthesis based either on measured CO₂ concentrations and conductance values or as measured directly by ¹⁴CO₂ labeling experiments. The photosynthetic responses to CO₂ and O₂ were measured to determine whether onion leaves exhibited a typical C₃-type response. A linear increase in CO₂ uptake was observed in intact leaves up to 315 L L^–1 of external CO₂ and, at this external CO₂ concentration, uptake was inhibited 35.4±0.9% by 210 mL L^–1 O₂ compared to 20 mL L^–1 O₂. Scanning electron micrographs of the leaf cavity wall revealed degenerated tissue covered by a membrane. Onion leaf cavity membranes apparently are highly impermeable to CO₂ and greatly restrict the refixation of leaf cavity CO₂ by photosynthetic tissue.Abbreviations C_a external CO₂ concentration - C_i intercellular CO₂ concentration - CO₂ compensation concentration - PPFR photosynthetic photon fluence rate     

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/C_i response). The effect of CO₂ enrichment on the A/C_i 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 C_i, 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 C_i following a doubling or tripling of atmospheric CO₂ during growth. Using theoretical interpretations of A/C_i curves to assess acclimation, it was found that when pot size or nutrient deficiency was not a factor, changes in the shape of A/C_i 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 C_i. 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 C_i/C_a ratio) was used to evaluate stomatal acclimation. Except under water and humidity stress, C_i/C_a exhibited no consistent change in a variety of C₃ species, indicating no stomatal acclimation. Under drought or humidity stress, C_i/C_a declined in high-CO₂ grown plants, indicating stomata will become more conservative during stress episodes in future high CO₂ environments.Abbreviations A net CO₂ assimilation rate - C_i (C_a) intercellular (ambient) partial pressure of CO₂ - operational C_i intercellular partial pressure of CO₂ at a given ambient partial pressure of CO₂ - g_s stomatal conductance - normal CO₂ current atmospheric mole fraction of CO₂ (330 to 355 mol mol^–1) - Rubisco ribulose-1,5-bisphosphate carboxylase/oxygenase     

Photosynthetic characteristics of coffee (Coffea arabusta) plantlets in vitro in response to different CO2 concentrations and light intensities

The photosynthetic characteristics of coffee ( Coffea arabusta) plantlets cultured in vitro in response to different CO₂ concentrations inside the culture vessel and photosynthetic photon flux (PPF) were investigated preliminarily. The estimation of net photosynthetic rate (P_n) of coffee plantlets involved three methods: (1) estimating time courses of actual P_n in situ based on measuring CO₂ concentrations inside and outside the vessel during a 45-day period, (2) estimating P_n in situ at different CO₂ concentrations and PPFs using the above measuring approach for 10-day and 30-day old in vitro plantlets, and (3) estimating P_n of a single leaf at different CO₂ concentrations and PPFs by using a portable photosynthesis measurement system for 45-day old in vitro coffee plantlets. The results showed that coffee plantlets in vitro had relatively high photosynthetic ability and that the P_n increased with the increase in CO₂ concentration inside the vessel. The CO₂ saturation point of in vitro coffee plantlets was high (4500–5000 μmol mol^-1); on the other hand, the PPF saturation point was not so high as compared to some other species, though it increased with increasing CO₂ concentration inside the vessel. This revised version was published online in June 2006 with corrections to the Cover Date.     

Modelling the CO2 gas exchange of grassland vegetation from experimental data

Pseudo-cubic spline functions were applied to the two atmospheric CO₂ concentrations of 340 and 600 l·l^–1 CO₂ for describing the average daily CO₂ gas exchange rates of simplifed grassland model ecosystems. Measurements used were from daily means of photon flux density (PhAR), temperature and relative air humidity, phytomass of each day, leaf area indexes, average phenological states of the vegetation (1–15), and water exchange rates of the entire model ecosystems. The functions were validated with, data from the same experimental year. We also succeeded in verifying the functions with the response data from years other than those used for constructing the model.     

Novel technique for component monitoring of CO2 exchange in plants

A novel technique designed for component monitoring of CO₂ exchange in plants is described. The system is based on application of self-clamping leaf chambers connected to an open gas-exchange measuring system and on automatic recording of CO₂ concentration. This technique was implemented in a commercially available instrument, PTM-48A Photosynthesis Monitor, which provides for long-term measurements of gas exchange and for discrimination of its separate components. Furthermore, many other plant functions can be monitored during plant growth and development under laboratory, greenhouse, and field conditions.     

The response of CO2 exchange rate to photosynthetic photon flux density for several Populus clones under laboratory conditions

No significant differences were found between four mathematical equations describing the response of CO₂ exchange rate to photosynthetic photon flux density in seven poplar clones under laboratory conditions. Choice of an optimal equation for poplar may be based on the contemplated aims. High significant differences (at p<0.001) were found among the clones.Research was supported by the Instituut tot Aanmoediging van het Wetenschappelijk Onderzoek in Nijverheid en Landbouw (I.W.O.N.L.), Brussels.     

A study on the relationship between leaf conductance,CO2 concentration and carboxylation rate in various species

Leaf conductance g_L is strongly influenced by environmental factors like CO₂, irradiance and air humidity. According to Ball et al. (1987), g_L is correlated with an index calculated as the product of net CO₂ exchange rate A and ambient water vapour concentration W_a, divided by ambient CO₂ concentration c_a. However, this empirical model does not apply to high values of g_L observed at c_a below CO₂ compensation concentration . Therefore, we applied modified indices in which A is replaced by estimates for the rate of carboxylation. Such estimates, P₁ and P₂, were determined by adding to A the quotient of and the sum of gas phase resistance r_g and intracellular resistance for CO₂ exchange r_i, P₁ = A+/(r_g + r_i), or the quotient of and r_i, P₂ = A + /r_i. If P₂ is chosen, c_a in the Ball index has to be replaced by the intercellular CO₂ concentration c_i. By using the modified indices P₁·W_a/c_a and P₂·W_a/c_i, we analysed data from the C₃ species Nicotiana tabacum and Nicotiana plumbaginifolia, the C₃–C₄ intermediate species Diplotaxis tenuifolia, and the C₄ species Zea mays. The data were collected at widely varying levels of irradiance and CO₂ concentration. For all species uniform relationships between g_L and the new indices were found for the whole range of CO₂ concentrations below and above . Correlations between g_L and P₁·W_a/c_a were closer than those between g_L and P₂·W_a/c_i because P₁/c_a implicitly contains g_L. Highly significant correlations were also obtained for the relationships between g_L and the ratios P₁/c_a and P₂/c_i.     

Effect of photon flux density on inorganic carbon accumulation and net CO2 exchange in a high-CO2-requiring mutant of Chlamydomonas reinhardtii

The effect of photon flux density on inorganic carbon accumulation and photosynthetic CO₂ assimilation was determined by CO₂ exchange studies at three, limiting CO₂ concentrations with a ca-1 mutant of Chlamydomonas reinhardiii. This mutant accumulates a large internal inorganic carbon pool in the light which apparently is unavailable for photosynthetic assimilation. Although steady-state photosynthetic CO₂ assimilation did not respond to the varying photon flux densities because of CO₂ limitation, components of inorganic-carbon accumulation were not clearly light saturated even at 1100 mol photons m^-2 s^-1, indicating a substantial energy requirement for inorganic carbon transport and accumulation. Steady-state photosynthetic CO₂ assimilation responded to external CO₂ concentrations but not to changing internal inorganic carbon concentrations, confirming that diffusion of CO₂ into the cells supplies most of the CO₂ for photosynthetic assimilation and that the internal inorganic carbon pool is essentially unavailable for photosynthetic assimilation. The estimated concentration of the internal inorganic carbon pool was found to be relatively insensitive to the external CO₂ concentration over the small range tested, as would be expected if the concentration of this pool is limited by the internal to external inorganic carbon gradient. An attempt to use this CO₂ exchange method to determine whether inorganic carbon accumulation and photosynthetic CO₂ assimilation compete for energy at low photon flux densities proved inconclusive.     

Effect of varying co2 and light levels on growth of hedyotis and sugarcane shoot cultures

Summary Shoot cultures of Hedyotis corymbosa, a C3 species, and sugarcane, a C4 species, were used to examine the effects of various CO₂ concentrations and two light intensities on growth and photosynthetic rates. The fresh and dry weights of new growth of Hedyotis shoots were higher when grown under the higher light intensity, while differences among shoots grown under different CO₂ levels were marginal. After 14 d of growth in various CO₂ concentrations, no significant differences could be observed in the newly produced leaves of Hedyotis with respect to stomatal distribution and number of mesophyll cell layers. Shoots grown under high light intensity did not show higher rates of photosynthesis than those grown under low light intensity. Also, sugarcane shoots grown in a CO₂-enriched environment did not have higher photosynthetic rates, perhaps because the C4 pathway is less sensitive to the ambient CO₂ concentration. The quantum yield of Hedyotis shoots grown on medium with 20 g l⁻¹ sucrose was lower than that of shoots on lower sucrose concentrations, supporting the view that photosynthesis is inhibited by high levels of sucrose. Our results suggest that Hedyotis shoots in culture exhibit some form of acclimation to high CO₂. so that there is no net gain in productivity by photosynthesis.     

Effect of a change in photoperiod on subsequent CO2 exchange

The effect of a shift from a long to a short photoperiod on CO₂ exchange of Chenopodium polyspermum was studied. Equal quantities of photosynthetic energy were given daily to the plants, long photoperiods being produced by low intensity red light extension. A change in the photoperiod was shown to affect the pattern of CO₂ loss at the beginning of the night period and the onset of CO₂ intake at the beginning of day time. These events seem to be under phytochrome control.The photoperiod had an effect on the slope of the CO₂ curve of photosynthesis, efficiency of photosynthesis being increased after a short day. This effect was not due to a variation in the stomatal resistance.The action of O₂ concentration on photosynthesis (Warburg effect) was affected by the photoperiodic treatment, being less important after a long day than after a short day.Involvement of phytochrome in photosynthetic efficiency and photorespiration is discussed.

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Résumé On étudie l'effet d'une variation de photopériode sur les échanges de CO₂ de Chenopodium polyspermum. Les plantes reçoivent la même quantité d'énergie utilisable par la photosynthèse, l'allongement de la photopériode étant obtenu par addition au cours de la nuit de lumière rouge de faible intensité.Un changement de photopériode affecte à court terme le déroulement de la chute de respiration en début de nuit ainsi que la mise en route de la photosynthèse le jour suivant.Une variation de traitement photopériodique modifie l'efficience de la photosynthèse: la pente de la courbe de photosynthèse en fonction du CO₂ est plus élevée après un jour court. Cet effet n'est pas dú à une variation de résistance stomatique.L'action de la concentration en oxygène de l'air sur la photosynthèse (effet Warburg) est également affectée par le traitement photopériodique: elle est moins importante après un jour long qu'après un jour court.On discute l'influence éventuelle du phytochrome sur l'efficience de la photosynthèse et la photorespiration.

     

Multichannel automated chamber system for continuous monitoring of CO2 exchange between the agro-ecosystem or soil and the atmosphere

A multichannel automated chamber system was developed for continuous monitoring of CO₂ exchange at multiple points between agro-ecosystem or soil and atmosphere. This system consisted of an automated chamber subsystem with a CO₂ concentration analyzer and a data logging subsystem. Both subsystems were under the control of a programmable logic controller (PLC). The automated chamber subsystem contained 18 chambers (50 cm × 50 cm × 50 cm) and a compressor. The chamber lids were closed and can be automatically opened. During measurement, one of the 18 chambers was kept closed for three minutes for measuring and the other chambers were kept open to maintain the natural soil conditions to the maximum extent. Environmental variables were simultaneously measured using sensors and recorded by the data logger. The reliability of the multichannel automated chamber system was tested and the results showed that the turbulence of the fans had no significant effect on the CO₂ exchange. The changes in the air and the temperature of soil and soil moisture inside the chambers, caused by the enclosure of the chambers, were not significant. The net ecosystem CO₂ exchange for the wheat ecosystem was ?2.35 μmol·m^?2·s^>?1 and the soil respiration was 3.87 μmol·m^?2·s^>?1 in the wheat field, and 6.61 μmol·m^?2·s^>?1 in the apple orchard.     

Response of 12 weedy species to elevated CO2 in low-phosphorus-availability soil

We conducted an experiment on responses of weedy species from an orchard ecosystem to elevated CO₂ (700–800 μmol mol⁻¹) under low phosphorus (P) soil in an environment-controlled growth chamber. Twelve local weedy species, Poa annua L., Lolium perenne L., Avena fatua L., Vicia cracca L., Medicago lupulina L., Kummerowia striata (Thunb.) Schindl., Veronica didyma Ten., Plantago virginica L., Gnaphalium affine D.Don., Echinochloa crusgalli var. mitis (L.) Beauv., Eleusine indica (L.) Gaertn. and Setaria glauca (L.) P. Beauv., grouped into four functional groups (C₃ grass, C₃ forb, legume and C₄ grass), were used in the experiment. The total plant biomass, P uptake, and mycorrhizal colonization were measured. The results showed that the total biomass of the 12 weedy species tended to increase under elevated CO₂. But changes in the total biomass under elevated CO₂ significantly differed among functional groups: legumes showed the greatest increase in the total biomass of all functional groups, following the order C₃ forbs > C₄ grasses > C₃ grasses. Elevated CO₂ significantly increased mycorrhizal colonization and P uptake of legumes, C₃ forbs and C₄ grasses but did not change C₃ grasses. Positive correlations between mycorrhizal colonization and shoot P concentration, and between total P uptake and total biomass were found under elevated CO₂. The results suggested that the interspecific difference in CO₂ response at low P availability was caused by the difference in CO₂ response in mycorrhizae and P uptake. These differences among species imply that plant interaction in orchard ecosystems may change under future CO₂ enrichment.     

Effect of O2 and CO2 on net CO2 exchange in a high-CO2-requiring mutant of Chlamydomonas reinhardtii during dark-light-dark transitions

Net CO₂ exchange was monitored through a dark-light-dark transition, under 2% and 21% O₂ in the presence and absence of CO₂, in Chlamydomonas reinhardtii wild type and the high-CO₂-requiring mutant ca-1-12-1C. Upon illumination at 350 l/l CO₂, ca-1-12-1C cell exhibited a large decrease in net CO₂ uptake following an initial surge of CO₂ uptake. Net CO₂ uptake subsequently attained a steady-state rate substantially lower than the maximum. A large, O₂-enchanced post-illumination burst of CO₂ efflux was observed after a 10-min illumination period, corresponding to a minimum in the net CO₂ uptake rate. A smaller, but O₂-insensitive post-illumination burst was observed following a 30-min illumination period, when net CO₂ uptake was at a steady-state rate. These post-illumination bursts appeared to reflect the release of an intracellular pool of inorganic carbon, which was much larger following the initial surge of net CO₂ uptake than during the subsequent steady-state CO₂ uptake period.With the mutant in CO₂-free gas, O₂-stimulated, net CO₂ efflux was observed in the light, and a small, O₂-dependent post-illumination burst was observed. With wild-type cells no CO₂ efflux was observed in the light in CO₂-free gas under either 2% or 21% O₂, but a small, O₂-dependent post-illumination burst was observed. These results were interpreted as indicating that photorespiratory rates were similar in the mutant and wild-type cells in the absence of CO₂, but that the wild-type cells were better able to scavenge the photorespiratory CO₂.     

Open top chambers for exposing plant canopies to elevated CO2 concentration and for measuring net gas exchange

Open top chamber design and function are reviewed. All of the chambers described maintain CO₂ concentrations measured at a central location within ±30 ppm of a desired target when averaged over the growing season, but the spatial and temporal range within any chamber may be closer to 100 ppm. Compared with unchambered companion plots, open top chambers modify the microenvironment in the following ways: temperatures are increased up to 3°C depending on the chamber design and location of the measurement; light intensity is typically diminished by as much as 20%; wind velocity is lower and constant; and relative humidity is higher. The chamber environment may significantly alter plant growth when compared with unchambered controls, but the chamber effect on growth has not been clearly attributed to a single or even a few environmental factors.A method for modifying an open top chamber for tracking gas exchange between natural vegetation and the ambient air is described. This modification consists of the addition of a top with exit chimney to reduce dilution of chamber CO₂ by external ambient air, is quickly made and permits estimation of the effects of elevated CO₂ and water vapor exchange.The relatively simple design and construction of open top chambers make them the most likely method to be used in the near future for long-term elevated CO₂ exposure of small trees, crops and grassland ecosystems. Improvements in the basic geometry to improve control of temperature, reduce the variation of CO₂ concentrations, and increase the turbulence and wind speed in the canopy boundary layer are desirable objectives. Similarly, modifications for measuring water vapor and carbon dioxide gas exchange will extend the usefulness of open top chambers to include non-destructive monitoring of the responses of ecosystems to rising atmospheric CO₂.     

CO2-induced growth enhancements of co-occurring tree species decline at different rates

To elucidate how enriched CO₂ atmospheres, soil fertility, and light availability interact to influence the long-term growth of tree seedlings, six co-occurring members of temperate forest communities including ash (Fraxinus americana L.), gray birch (Betula populifolia), red maple (Acer rubrum), yellow birch (Betula alleghaniensis), striped maple (Acer pensylvanicum), and red oak (Quercus rubra L.) were raised in a glasshouse for three years in a complete factorial design. After three years of growth, plants growing in elevated CO₂ atmospheres were generally larger than those in ambient CO₂ atmospheres, however, magnitudes of CO₂-induced growth enhancements were contingent on the availability of nitrogen and light, as well as species identity. For all species, magnitudes of CO₂-induced growth enhancements after one year of growth were greater than after three years of growth, though species' growth enhancements over the three years declined at different rates. These results suggest that CO₂-induced enhancements in forest productivity may not be sustained for long periods of time. Additionally, species' differential growth responses to elevated CO₂ may indirectly influence forest productivity via long-term species compositional changes in forests.     

The influence of plant species,fertilization and elevated CO2 on soil aggregate stability

We tested the effects of plant species, fertilization and elevated CO₂ on water-stable soil aggregation. Five annual grassland species and a plant community were grown in outdoor mesocosms for 4 years, with and without NPK fertilization, at ambient or elevated atmospheric CO₂ concentrations. Aggregate stability (resistance of aggregates to slaking) in the top 0.15 m of soil differed among plant species. However, the more diverse plant community did not enhance aggregate stability relative to most monocultures. Species differences in aggregate stability were positively correlated with soil active bacterial biomass, but did not correlate with root biomass or fungal length. Plant species did not affect aggregate stability lower in the soil profile (0.15–0.45 m), where soil biological activity is generally decreased. Elevated CO₂ and NPK fertilization altered many of the factors known to influence aggregation, but did not affect water-stable aggregation at either depth, in any of the plant treatments. These results suggest that global changes will alter soil structure primarily due to shifts in vegetation composition.     

In situ CO2 gas-exchange in fruits of a tropical tree,Durio zibethinus Murray

We examined the in situ CO₂ gas-exchange of fruits of a tropical tree, Durio zibethinus Murray, growing in an experimental field station of the Universiti Pertanian Malaysia. Day and night dark respiration rates were exponentially related to air temperature. The temperature dependent dark respiration rate showed a clockwise loop as time progressed from morning to night, and the rate was higher in the daytime than at night. The gross photosynthetic rate was estimated by summing the rates of daytime dark respiration and net photosynthesis. Photosynthetic CO₂ refixation, which is defined as the ratio of gross photosynthetic rate to dark respiration rate in the daytime, ranged between 15 and 45%. The photosynthetic CO₂ refixation increased rapidly as the temperature increased in the lower range of air temperature T _c (T _c <28.5 °C), while it decreased gradually as the temperature increased in the higher range (T _c 28.5 °C). Light dependence of photosynthetic CO₂ refixation was approximated by a hyperbolic formula, where light saturation was achieved at 100 mol m^–2 s^–1 and the asymptotic CO₂ refixation was determined to be 37.4%. The estimated gross photosynthesis and dark respiration per day were 1.15 and 4.90 g CO₂ fruit^–1, respectively. Thus the CO₂ refixation reduced the respiration loss per day by 23%. The effect of fruit size on night respiration rate satisfied a power function, where the exponent was larger than unity.