Plantecophys - An R Package for Analysing and Modelling Leaf Gas Exchange Data

Here I present the R package ''plantecophys'', a toolkit to analyse and model leaf gas exchange data. Measurements of leaf photosynthesis and transpiration are routinely collected with portable gas exchange instruments, and analysed with a few key models. These models include the Farquhar-von Caemmerer-Berry (FvCB) model of leaf photosynthesis, the Ball-Berry models of stomatal conductance, and the coupled leaf gas exchange model which combines the supply and demand functions for CO₂ in the leaf. The ''plantecophys'' R package includes functions for fitting these models to measurements, as well as simulating from the fitted models to aid in interpreting experimental data. Here I describe the functionality and implementation of the new package, and give some examples of its use. I briefly describe functions for fitting the FvCB model of photosynthesis to measurements of photosynthesis-CO₂ response curves (''A-C_i curves''), fitting Ball-Berry type models, modelling C3 photosynthesis with the coupled photosynthesis-stomatal conductance model, modelling C4 photosynthesis, numerical solution of optimal stomatal behaviour, and energy balance calculations using the Penman-Monteith equation. This open-source package makes technically challenging calculations easily accessible for many users and is freely available on CRAN.     

Nano-TiO2 Improve the Photosynthesis of Tomato Leaves under Mild Heat Stress

Nano-TiO₂ has been reported to promote photosynthesis in some crops; however, the mechanism behind this action remains unknown. In this research, the effects of nano-TiO₂ on leaf photosynthesis under mild heat stress were investigated. Results showed that the net photosynthetic rate, conductance to H₂O, and transpiration rate of tomato leaves increased after application of an appropriate concentration of nano-TiO₂. Nano-TiO₂ also significantly decreased the minimum chlorophyll fluorescence and relative electron transport in leaves. Under mild heat stress, Nano-TiO₂ increased regulated photosystem II (PS II) energy dissipation and decreased non-regulated PS II energy dissipation. These results indicate that nano-TiO₂ plays a positive role in promoting photosynthesis in tomato leaves under mild heat stress.     

A stomatal optimization theory to describe the effects of atmospheric CO2 on leaf photosynthesis and transpiration

Background and Aims

Global climate models predict decreases in leaf stomatal conductance and transpiration due to increases in atmospheric CO₂. The consequences of these reductions are increases in soil moisture availability and continental scale run-off at decadal time-scales. Thus, a theory explaining the differential sensitivity of stomata to changing atmospheric CO₂ and other environmental conditions must be identified. Here, these responses are investigated using optimality theory applied to stomatal conductance.

Methods

An analytical model for stomatal conductance is proposed based on: (a) Fickian mass transfer of CO₂ and H₂O through stomata; (b) a biochemical photosynthesis model that relates intercellular CO₂ to net photosynthesis; and (c) a stomatal model based on optimization for maximizing carbon gains when water losses represent a cost. Comparisons between the optimization-based model and empirical relationships widely used in climate models were made using an extensive gas exchange dataset collected in a maturing pine (Pinus taeda) forest under ambient and enriched atmospheric CO₂.

Key Results and Conclusion

In this interpretation, it is proposed that an individual leaf optimally and autonomously regulates stomatal opening on short-term (approx. 10-min time-scale) rather than on daily or longer time-scales. The derived equations are analytical with explicit expressions for conductance, photosynthesis and intercellular CO₂, thereby making the approach useful for climate models. Using a gas exchange dataset collected in a pine forest, it is shown that (a) the cost of unit water loss λ (a measure of marginal water-use efficiency) increases with atmospheric CO₂; (b) the new formulation correctly predicts the condition under which CO₂-enriched atmosphere will cause increasing assimilation and decreasing stomatal conductance.     

Development of a photosynthesis model with an emphasis on ecological applications

Summary A physiologically based steady-state model of whole leaf photosynthesis (WHOLEPHOT) is detailed which describes the functional dependence of net photosynthesis in C ₃ leaves on [CO₂], [O₂], incident radiant flux (PhAR), and leaf temperature. The model simulates among other phenomena a) observed [CO₂], [O₂], and temperature effects on the initial slope of light response curves, b) a C ₃ type temperature response curve of net photosynthesis, c) a shift of the optimum temperature of net photosynthesis to higher temperatures with increasing light intensity, and d) observed temperature and [O₂] effects on the CO₂ compensation point. Model parameters are derived from published response data of several C ₃ species. Simulations also demonstrate that parameter changes based on literature data result in acclimation-like changes in net photosynthesis response with respect to light intensity and temperature. The advantages of this model are that the number of parameters is minimized in order to focus on environmental effects and that all parameters can be determined from measured net photosynthesis responses.     

Seasonal response of photosynthesis to elevated CO2 in loblolly pine (Pinus taeda L.) over two growing seasons

Trees growing in natural systems undergo seasonal changes in environmental factors that generate seasonal differences in net photosynthetic rates. To examine how seasonal changes in the environment affect the response of net photosynthetic rates to elevated CO₂, we grew Pinus taeda L. seedlings for three growing seasons in open-top chambers continuously maintained at either ambient or ambient + 30 Pa CO₂. Seedlings were grown in the ground, under natural conditions of light, temperature nd nutrient and water availability. Photosynthetic capacity was measured bimonthly using net photosynthetic rate vs. intercellular CO₂ partial pressure (A-C_i) curves. Maximum Rubisco activity (Vc_max) and ribulose 1,5-bisphosphate regeneration capacity mediated by electron transport (J_max) and phosphate regeneration (PiRC) were calculated from A-C_i curves using a biochemically based model. Rubisco activity, activation state and content, and leaf carbohydrate, chlorophyll and nitrogen concentrations were measured concurrently with photosynthesis measurements. This paper presents results from the second and third years of treatment. Mean leaf nitrogen concentrations ranged from 13.7 to 23.8 mg g^?1, indicating that seedlings were not nitrogen deficient. Relative to ambient CO₂ seedlings, elevated CO₂ increased light-saturated net photosynthetic rates 60–110% during the summer, but < 30% during the winter. A relatively strong correlation between leaf temperature and the relative response of net photosynthetic rates to elevated CO₂ suggests a strong effect of leaf temperature. During the third growing season, elevated CO₂ reduced Rubisco activity 30% relative to ambient CO₂ seedlings, nearly completely balancing Rubisco and RuBP-regeneration regulation of photosynthesis. However, reductions in Rubisco activity did not eliminate the seasonal pattern in the relative response of net photosynthetic rates to elevated CO₂. These results indicate that seasonal differences in the relative response of net photosynthetic rates to elevated CO₂ are likely to occur in natural systems.     

Photosynthate supply and utilization in alfalfa : a developmental shift from a source to a sink limitation of photosynthesis

Long-term carbon dioxide enrichment, ¹⁴CO₂ feeding, and partial defoliation were employed as probes to investigate source/sink limitations of photosynthesis during the development of symbiotically grown alfalfa. In the mature crop, long-term CO₂ enrichment does not affect the rates of net photosynthesis, relative growth, ¹⁴C export to nonphotosynthetic organs, or the rates of ¹⁴C label incorporation into leaf sucrose, starch, or malate. The rate of glycolate labeling is, however, substantially reduced under these conditions. When the mature crop was partially defoliated, a considerable increase in net photosynthesis occurred in the remaining leaves. In the seedling crop, long-term CO₂ enrichment increased dry matter accumulation, primarily as a result of increases in leaf starch content. Although the higher rates of starch synthesis are not maintained, the growth enhancement of the enriched plants persisted throughout the experimental period. These results imply a source limitation of seedling photosynthesis and a sink limitation of photosynthesis in more mature plants. Consequently, both the supply and the utilization of photosynthate may limit seasonal photosynthesis in alfalfa.     

STEM AND LEAF GAS EXCHANGE OF TWO ARID LAND SHRUBS

Gas exchange studies were conducted on two shrub species found in cool shrub-steppe communities of the American West, big sagebrush (Artemisia tridentata subsp. tridentata Nutt.) and broom snakeweed (Gutierrezia sarothrae [Pursh] Britt. and Rusby), with a goal of evaluating characteristics and relative contributions of green stem and leaf material to total shoot CO₂ exchange at different temperatures. Variations in tissue temperature exerted a pronounced effect on CO₂ exchange—net photosynthesis and dark respiration—of green stems and leaves of both species. Definite temperature optima of net photosynthesis were noted, and dark respiration rates consistently increased with increases in temperature. Green stems of both species exhibited sizable dark respiration rates, although stem rates at all temperatures were lower than corresponding leaf rates. Artemisia tridentata did not exhibit sizeable green stem net photosynthesis even under conditions of optimal temperature and water availability, and leaf net photosynthesis rates were much lower than those of G. sarothrae. However, A. tridentata in general possessed a greater leaf biomass than G. sarothrae. Green stems of G. sarothrae exhibited considerable rates of net photosynthesis under both optimal and sub-optimal temperature and water availability conditions. A higher optimum temperature of net photosynthesis was noted for stems than for leaves of G. sarothrae. The adaptive significance of these interspecific differences in CO₂ exchange characteristics is discussed.     

Comparative gas exchange and growth responses of C3 and C4 beach species grown at different salinities

Summary Comparative laboratory gas exchange and relative growth rate experiments were conducted on three native California coastal strand species at four salinity treatment levels. Relative mesophyll conductance sensitivities to salinity of Atriplex leucophylla (Moq.) D. Dietr. (C₄) and Atriplex californica Moq. in D.C. (C₃) were nearly identical. Mesophyll conductances of both species were stimulated by moderate levels of salinity. Mesophyll conductances of Abronia maritima Nutt. ex Wats. (C₃) were highest in the absence of salinity and depressed by increasing levels of salinity. Increasing levels of salinity generally decreased net photosynthesis and leaf conductances but increased water use efficiencies. The C₄ species, Atriplex leucophylla, had higher mesophyll conductances and water use efficiencies at all salinity levels than the C₃ species. The effects of salinity on relative growth and net assimilation rates of greenhouse grown plants were not directly correlated with the effects on net photosynthesis measured in the laboratory. Growth of Abronia maritima was greatly stimulated by low levels of salinity whereas photosynthesis was substantially inhibited. The possible significance of C₄ photosynthesis in relation to salinity is discussed.     

Modelling C3 and C4 photosynthesis under water-stressed conditions

Despite the observed impact of water stress on photosynthesis, some of the most used models of CO₂ assimilation in C₃ and C₄ functional types do not directly account for it. We discuss an extension of these models, which explicitly includes the metabolic and diffusive limitations due to water stress on photosynthesis. Functional relationships describing the photosynthetic processes and CO₂ diffusion inside leaves are modified to account for leaf water status on the basis of experimental results available in the literature. Extensive comparison with data shows that the model is suitable to describe the reduction in CO₂ assimilation rate with decreasing leaf water potentials in various species. A simultaneous analysis of photosynthesis, transpiration and soil moisture dynamics is then carried out to explore the actual impact of drought on different photosynthesis processes and on the overall plant activity. The model well reproduces measured CO₂ assimilation rate as a function of soil moisture and could be useful to formulate hypotheses for detailed experiments as well as to simulate in detail transpiration and photosynthesis dynamics under water stress.     

LIGHT,TEMPERATURE AND SALINITY EFFECTS ON GROWTH,LEAF ANATOMY AND PHOTOSYNTHESIS OF DISTICHLIS SPICATA (L.) GREENE

The effects of light, temperature, and salinity on growth, net CO₂ exchange and leaf anatomy of Distichlis spicata were investigated in controlled environment chambers. When plants were grown at low light, growth rates were significantly reduced by high substrate salinity or low temperature. However, when plants were grown at high light, growth rates were not significantly affected by temperature or salinity. The capacity for high light to overcome depressed growth at high salinity cannot be explained completely by rates of net photosynthesis, since high salinity caused decreases in net photosynthesis at all environmental conditions. This salinity-induced decrease in net photosynthesis was caused largely by stomatal closure, although plants grown at low temperature and low light showed significant increases in internal leaf resistance to CO₂ exchange. Increased salinity resulted in generally thicker leaves with lower stomatal density but no significant differences in the ratio of mesophyll cell surface area to leaf area. Salinity and light during growth did not significantly affect rates of dark respiration. The mechanisms by which Distichlis spicata tolerates salt appear to be closely coulpled to the utilization of light energy. Salt-induced leaf succulence is of questionable importance to gas exchange at high salinity in this C₄ species.     

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.     

Evaluation of the light/dark C assay of photorespiration: tobacco leaf disk studies with glycidate and glyoxylate

Preincubation of illuminated tobacco (Nicotiana tabacum L.) leaf disks in glycidate (2,3-epoxypropionate) or glyoxylate inhibited photorespiration by about 40% as determined by the ratio of ¹⁴CO₂ evolved into CO₂-free air in light and in darkness. However, under identical preincubation conditions used for the light/dark ¹⁴C assays, the compounds failed to reduce photorespiration or stimulate net photosynthesis in tobacco leaf disks based on other CO₂ exchange parameters, including the CO₂ compensation concentration in 21% O₂, the inhibitory effect of 21% O₂ on net photosynthesis in 360 microliters per liter of CO₂ and the rate of net photosynthetic ¹⁴CO₂ uptake in air.

The effects of both glycidate and glyoxylate on the ¹⁴C assay are inconsistent with other measures of photorespiratory CO₂ exchange in tobacco leaf disks, and thus these data question the validity of the light to dark ratio of ¹⁴CO₂ efflux as an assay for relative rates of photorespiration (Zelitch 1968, Plant Physiol 43: 1829-1837). The results of this study specifically indicate that neither glycidate nor glyoxylate reduces photorespiration or stimulates net photosynthesis by tobacco leaf disks under physiological conditions of pO₂ and pCO₂, contrary to previous reports.

     

Long-term effects of drought on wild and cultivated plants in the Negev desert

Summary The relation between daily maximal rates of net photosynthesis and plant water status was studied during a dry season on irrigated and non-irrigated, naturally growing, perennial wild plants.Species were examined which differ in phenology, leaf anatomy and morphology: Hammada scoparia, Artemisia herba-alba, Zygophyllum dumosum, and Reaumuria negevensis. Prumus armeniaca which was growing in the run-off farm at Avdat and which has mosomorphic leaves was included in the comparison. All plants differed in their seasonal change in plant water status, and in their seasonal change in daily maximal net photosynthesis. Rates of CO₂ uptake were not uniquely related to simultanously measured leaf water potentials. Daily maximal rates of net photosynthesis of non-irrigated plants, and the difference between maximal CO₂ uptake of irrigated and non-irrigated plants were examined in relation to pre-dawn water potential. Maximal net photosynthesis rates decreased very rapidly with decrease in pre-dawn water potential or, for Hammada scoparia, they decreased even with a constant level of pre-dawn water potential. Consequently, it was considered necessary to include both time and water potential in a parameter bar day describing the accumulated drought stress of the plants. All species showed the same relation between relative maximal net photosynthesis and drought experience as determined by cumulative daily addition of pre-dawn water potentials for the non-irrigated plants since the last rain.     

C4 photosynthesis in C3 rice: a theoretical analysis of biochemical and anatomical factors

Engineering C₄ photosynthesis into rice has been considered a promising strategy to increase photosynthesis and yield. A question that remains to be answered is whether expressing a C₄ metabolic cycle into a C₃ leaf structure and without removing the C₃ background metabolism improves photosynthetic efficiency. To explore this question, we developed a 3D reaction diffusion model of bundle‐sheath and connected mesophyll cells in a C₃ rice leaf. Our results show that integrating a C₄ metabolic pathway into rice leaves with a C₃ metabolism and mesophyll structure may lead to an improved photosynthesis under current ambient CO₂ concentration. We analysed a number of physiological factors that influence the CO₂ uptake rate, which include the chloroplast surface area exposed to intercellular air space, bundle‐sheath cell wall thickness, bundle‐sheath chloroplast envelope permeability, Rubisco concentration and the energy partitioning between C₃ and C₄ cycles. Among these, partitioning of energy between C₃ and C₄ photosynthesis and the partitioning of Rubisco between mesophyll and bundle‐sheath cells are decisive factors controlling photosynthetic efficiency in an engineered C₃–C₄ leaf. The implications of the results for the sequence of C₄ evolution are also discussed.     

Comparative responses of model C3 and C4 plants to drought in low and elevated CO2

Interactive effects of CO₂ and water availability have been predicted to alter the competitive relationships between C3 and C4 species over geological and contemporary time scales. We tested the effects of drought and CO₂ partial pressures (pCO₂) ranging from values of the Pleistocene to those predicted for the future on the physiology and growth of model C3 and C4 species. We grew co-occurring Abutilon theophrasti (C3) and Amaranthus retroflexus (C4) in monoculture at 18 (Pleistocene), 27 (preindustrial), 35 (current), and 70 (future) Pa CO₂ under conditions of high light and nutrient availability. After 27 days of growth, water was withheld from randomly chosen plants of each species until visible wilting occurred. Under well-watered conditions, low pCO₂ that occurred during the Pleistocene was highly limiting to C3 photosynthesis and growth, and C3 plants showed increased photosynthesis and growth with increasing pCO₂ between the Pleistocene and future CO₂ values. Well-watered C4 plants exhibited increased photosynthesis in response to increasing pCO₂, but total mass and leaf area were unaffected by pCO₂. In response to drought, C3 plants dropped a large amount of leaf area and maintained relatively high leaf water potential in remaining leaves, whereas C4 plants retained greater leaf area, but at a lower leaf water potential. Furthermore, drought-treated C3 plants grown at 18 Pa CO₂ retained relatively greater leaf area than C3 plants grown at higher pCO₂ and exhibited a delay in the reduction of stomatal conductance that may have occurred in response to severe carbon limitations. The C4 plants grown at 70 Pa CO₂ showed lower relative reductions in net photosynthesis by the end of the drought compared to plants at lower pCO₂, indicating that CO₂ enrichment may alleviate drought effects in C4 plants. At the Pleistocene pCO₂, C3 and C4 plants showed similar relative recovery from drought for leaf area and biomass production, whereas C4 plants showed higher recovery than C3 plants at current and elevated pCO₂. Based on these model systems, we conclude that C3 species may not have been at a disadvantage relative to C4 species in response to low CO₂ and severe drought during the Pleistocene. Furthermore, C4 species may have an advantage over C3 species in response to increasing atmospheric CO₂ and more frequent and severe droughts.     

夏玉米三叶期持续干旱下不同叶位叶片含水量变化及其与光合作用的关系

植物干物质的累积依赖于群体光合速率,而群体光合速率又与单叶的光合能力密切有关。叶片光合作用与其含水量密切相关,目前关于不同叶位叶片含水量对持续干旱的响应及其与光合作用的关系还未见报道。以华北夏玉米郑单958为材料,设置6个不同灌水处理,模拟不同灌溉量下持续干旱对夏玉米不同叶位叶片生理特性的影响,分析夏玉米顶部开始的第一、三、五叶位叶片的水分变化及其与净光合速率的关系。结果表明:夏玉米不同叶位的叶片最大含水量不同,且随干旱进程的推进叶片含水量的变化速率也不同,第一叶的叶片含水量下降速率高于第三、第五叶,第一叶的最大含水量高于第三、五叶,且可进行光合产物积累的叶片含水量下限随叶位的增加而增大。同时,第一叶的叶片含水量与土壤水分呈显著相关,且与净光合速率的相关性也非常强。第一叶可进行光合产物积累的叶片水分下限(净光合速率为零时的叶片含水量)最小,表明其耐旱性最强,对干旱具有指导意义。研究结果可为提高冠层光合作用模拟的准确性及夏玉米干旱发生发展的监测预警提供参考。     

Validation of a Photosynthesis Model through the Use of the CO2Balance of a Greenhouse Tomato Canopy

Several leaf photosynthesis models were developed from wellcontrolled experiments in growth chambers. However, only a fewhave been validated under greenhouse conditions for their quantitativeand qualitative adequacy. In this paper, rates of net photosynthesisfor a tomato crop (Lycopersicon esculentum Mill) were measuredin a semi-commercial greenhouse (615 m³) for a significant timeperiod. Concomitant measurements of climatic conditions andLAI were used for simulation of net photosynthesis using theTOMGRO model which integrates Acock's model for photosynthesiscalculations. From simulations and from sensitivity analysis,the prediction of net photosynthesis appeared to be very sensitiveto the quantum use efficiency. The Acock model with originalparameters underestimated the net photosynthesis rate, but anincrease in the quantum use efficiency by 10% gave a good fit.In an effort to generalize the validity of the model, a residualanalysis was performed and showed a systematic bias relatedto light intensity intercepted by the canopy. The Marquardtalgorithm was used to adjust our data to the model but did noteliminate residual heterogeneity of variance with new parametervalues. On the basis of collected data, the criteria of goodnessof fit used showed that the photosynthesis model is inadequatein describing the CO₂-balance of the greenhouse agrosystem.However, it was determined that it could be used as a submodelwithin a more complex model for predicting growth and development.Copyright 1999 Annals of Botany Company Greenhouse, CO₂-balance, photosynthesis, TOMGRO model, Acock's model, residuals, tomato Lycopersicon esculentum Mill.     

On the Reason for the Different Photosynthetic Rates of Seedlings of Pinus silvestris and Betula verrucosa

The growth and net photosynthetic properties of seedlings of Pinus silvestris L. and Betula verrucosa Ehrh., grown under identical conditions in a controlled environment chamber, were compared. The relative growth rate of birch was about twice that of pine. The rates of in situ net photosynthesis were 1.50 and 2.30 micromoles CO₂ meter⁻² second⁻¹ and the photosynthetic quantum yields under light-limiting conditions were 0.022 and 0.032 for pine and birch, respectively. The total leaf surface areas were used for calculating the CO₂ flux densities. The difference in the rates of in situ net photosynthesis depended equally on morphological and metabolic factors. It was assumed that a pronounced mutual shading and an unfavorable leaf inclination made the pine seedlings less efficient in absorbing the unidirectional light of the climate chamber than the broadleaved seedlings of birch. Both pine and birch were adapted to the growth conditions so the flux densities of absorbed quanta were rate-limiting for in situ net photosynthesis. It was concluded that the difference in the photosynthetic quantum yields (i.e. the linear slope of the photosynthetic light curve) of the two species defined the metabolically controlled part of the difference in the rate of in situ net photosynthesis. The quantum yield of pine was lower than that of birch and was partly explained by pine having a higher rate of photorespiration than birch. The remaining difference was most likely controlled by the properties of the chloroplast thylakoids, e.g. energy transfer efficiency between pigments, photosynthetic electron transport, or coupling between electron transport and photophosphorylation.     

Seasonal changes in photosynthetic characteristics of Anemone raddeana,a spring-active geophyte,in the temperate region of Japan

Summary Seasonal changes in the photosynthetic characteristics of intact involucral leaves of Anemone raddeana were investigated under laboratory conditions. Net photosynthesis and constant water vapor pressure deficit showed almost the same seasonal trend. They increased rapidly from mid-April immediately after unfolding of the leaves and reached the maximum in late-April, before the maximum expansion of the leaves. They retained the maximum values until early-May and then decreased toward late-May with a progress of leaf senescence. The calculated values of intercellular CO₂ concentration and relative stomatal limitation of photosynthesis showed no significant change throughout the season. The carboxylation efficiency as assessed by the initial slope of Ci-photosynthesis curve and the net photosynthesis under a high Ci regime varied seasonally in parallel with the change of the light-saturated photosynthesis. The results indicate that the seasonal changes in light-saturated net photosynthesis are not due to a change of stomatal conductance, but to a change in the photosynthetic capacity of mesophyll. Nevertheless, leaf conductance changed concomitantly with photosynthetic capacity, indicating that the seasonal change in stomatal conductance is modulated by the mesophyll photosynthetic capacity such that the intercellular CO₂ concentrations is maintained constant. The shape of light-photosynthesis curve was similar to that of sun-leaf type. The quantum yield also changed simultaneously with the photosynthetic capacity throughout the season.Contribution No. 2965 from the Institute of Low Temperature Science