植物叶肉导度的测定及计算方法综述
|
摘要
叶肉导度(g_m)指叶肉细胞内部的CO_2扩散能力,它是叶肉细胞阻力的倒数.光合作用研究的早期,研究者们多将叶肉细胞对CO_2的扩散阻力视为零,即假定g_m无穷大,而忽略了其对光合作用的限制.但近来的研究表明,g_m是有限的,并随着环境条件的改变而发生变化,此外,g_m的大小直接决定了CO_2在叶片内的扩散量,进而影响到植物光合效率的高低.因此,g_m的估算对于植物光合能力的评估意义重大.目前,气体交换与叶绿素荧光相结合法、曲线拟合法以及瞬时碳同位素(~(13)CO_2)辨别法已经成为植物g_m估测的3种常用方法,但国内针对这3种方法原理及其优缺点介绍的文献极少,阐述这3种方法的原理、过程并比较分析其优缺点就显得尤为必要.本文综合了相关文献,从原理、推导过程及优缺点3方面对上述3种g_m估测方法进行了详细介绍.结果表明:曲线拟合法虽然易于理解,便于操作,但其拟合模型因光合作用的发生状态不同而不同,需要研究者对光合作用不同状态进行严格划分,不利于广泛推广应用;瞬时碳同位素(~(13)CO_2)辨别法虽然提高了结果的准确性,但其测定过程比较复杂,对试验操作的要求比较严格,同时该方法对试验误差的敏感性较差,可靠性不高.相较上述两种方法,气体交换与叶绿素荧光相结合法的可操作性更强,可靠性更高,更利于多处理多重复的大样本的观测分析,叶绿素荧光技术的使用,既简化了试验步骤,又降低了试验过程的偶然误差,增加了观测结果的科学性;此外,叶绿素荧光技术还能为叶片提供饱和脉冲活化能,从而最大限度地激发叶片的光合潜能,但该方法也存在很多问题,比如,为了提高叶片叶绿素荧光参数的准确程度,试验中需要使用较低的气体流速,而流速的降低又会增大气体扩散泄漏的风险,所以该方法对选择合理气体流速的要求很高.综合来看,气体交换与叶绿素荧光相结合法在植物g_m的实际测定中的认可度最高,使用最广泛.
Mesophyll conductance(g_m) refers to the diffusion capacity of CO_2 inside mesophyll cells, which is the reciprocal of resistance of mesophyll cells. In the early stage of photosynthesis research, mesophyll diffusion resistance to CO_2 was usually assumed to be zero, namely the g_m was infinite. In recent studies, however, the g_m was found to be limited and changed with external environments. As g_m directly determines CO_2 diffusion and affects leaf photosynthetic efficiency, it is of great significance to mechanestic research of photosynthesis. Presently, simultaneous chlorophyll fluorescence and gas exchange, the curve-fitting and instantaneous carbon isotope(~(13)CO_2) discrimination are commonly used to estimate g_m, but few literature have been introduced on those methods in China. Therefore, it is particularly necessary to elaborate the principles and processes of these methods and to compare their advantages and disadvantages. We synthesized the relevant literature, and introduced the three methods in detail from the aspects of principle, derivation process and advantages and disadvantages, aiming to provide a methodological basis to promote the research on g_m in China. The curve-fitting method was easy to understand and operate. Its fitting model varied with the status of photosynthesis, which is needed to be divided strictly by researchers. Consequently, it was not conducive to be widely used. Although the instantaneous carbon isotope(~(13)CO_2) discrimination method improved the accuracy of results, it was complex in measurement and strict in operation. Furthermore, it was less sensitive to test errors with low reliability. Compared with the above two methods, simultaneous chlorophyll fluorescence and gas exchange was more operable and reliable, and was more conducive to the observation and analysis for large samples with multi-processing and multi-repetition. In addition, the use of chlorophyll fluorescence technology not only simplified the test procedures, but also reduced the accidental errors, making the results more scientific. Chlorophyll fluorescence technology also provided saturated pulse activation energy to maximize leaf photosynthetic potential. But this method also had many problems, for instance, to improve the accuracy of chlorophyll fluorescence parameters, a lower gas flow rate was needed, which would increase the risk of gas diffusion and leakage. Thus, this method had a high requirement for a reasonable gas flow rate. In general, simultaneous chlorophyll fluorescence and gas exchange method was most widely used in the actual determination of plant g_m.
Mesophyll conductance(g_m) refers to the diffusion capacity of CO_2 inside mesophyll cells, which is the reciprocal of resistance of mesophyll cells. In the early stage of photosynthesis research, mesophyll diffusion resistance to CO_2 was usually assumed to be zero, namely the g_m was infinite. In recent studies, however, the g_m was found to be limited and changed with external environments. As g_m directly determines CO_2 diffusion and affects leaf photosynthetic efficiency, it is of great significance to mechanestic research of photosynthesis. Presently, simultaneous chlorophyll fluorescence and gas exchange, the curve-fitting and instantaneous carbon isotope(~(13)CO_2) discrimination are commonly used to estimate g_m, but few literature have been introduced on those methods in China. Therefore, it is particularly necessary to elaborate the principles and processes of these methods and to compare their advantages and disadvantages. We synthesized the relevant literature, and introduced the three methods in detail from the aspects of principle, derivation process and advantages and disadvantages, aiming to provide a methodological basis to promote the research on g_m in China. The curve-fitting method was easy to understand and operate. Its fitting model varied with the status of photosynthesis, which is needed to be divided strictly by researchers. Consequently, it was not conducive to be widely used. Although the instantaneous carbon isotope(~(13)CO_2) discrimination method improved the accuracy of results, it was complex in measurement and strict in operation. Furthermore, it was less sensitive to test errors with low reliability. Compared with the above two methods, simultaneous chlorophyll fluorescence and gas exchange was more operable and reliable, and was more conducive to the observation and analysis for large samples with multi-processing and multi-repetition. In addition, the use of chlorophyll fluorescence technology not only simplified the test procedures, but also reduced the accidental errors, making the results more scientific. Chlorophyll fluorescence technology also provided saturated pulse activation energy to maximize leaf photosynthetic potential. But this method also had many problems, for instance, to improve the accuracy of chlorophyll fluorescence parameters, a lower gas flow rate was needed, which would increase the risk of gas diffusion and leakage. Thus, this method had a high requirement for a reasonable gas flow rate. In general, simultaneous chlorophyll fluorescence and gas exchange method was most widely used in the actual determination of plant g_m.
引文
[1] Flexas J,Brugnoli E,Warren CR.Mesophyll conductance to CO2// Flexas J,Loreto F,Medrano H,eds,Terrestrial Photosynthesis in a Changing Environment:A Molecular,Physiological and Ecological Approach.Cambridge:Cambridge University Press,2012:169-185
[2] Flexas J,Ribas-Carbó M,Diaz-Espejo A,et al.Mesophyll conductance to CO2:Current knowledge and future prospects.Plant,Cell & Environment,2008,31:602-621
[3] Farquhar GD,von Caemmerer S,Berry JA.A biochemical model of photosynthetic CO2 assimilation in leaves of C3 species.Planta,1980,149:78-90
[4] Han J-M (韩吉梅),Zhang W-F (张旺锋),Xiong D-L (熊栋梁),et al.Mesophyll conductance and its limiting factors in plant leaves.Chinese Journal of Plant Ecology (植物生态学报),2017,41(8):914-924 (in Chinese)
[5] Loreto F,Centritto M,Chartzoulakis K.Photosynthetic limitations in olive cultivars with different sensitivity to salt stress.Plant,Cell & Environment,2003,26:595-601
[6] Bernacchi CJ,Morgan PB,Ort DR,et al.The growth of soybean under free air [CO2] enrichment (FACE) stimulates photosynthesis while decreasing in vivo Rubisco capacity.Planta,2005,220:434-446
[7] Flexas J,Ribas-Carb M,Bota J.Decreased Rubisco activity during water stress is not induced by decreased relative water content but related to conditions of low stomatal conductance and chloroplast CO2 concentration.New Phytologist,2006,172:73-82
[8] Galmés J,Medrano H,Flexas J.Acclimation of Rubisco specificity factor to drought in tobacco:Discrepancies between in vitro and in vivo estimations.Journal of Experimental Botany,2006,57:3659-3667
[9] Galmés J,Medrano H,Flexas J.Photosynthetic limitations in response to water stress and recovery in Mediterranean plants with different growth forms.New Phytologist,2007,175:81-93
[10] Flowers MD,Fiscus EL,Burkey KO,et al.Photosynthesis,chlorophyll fluorescence,and yield of snap bean (Phaseolus vulgaris L.) genotypes differing in sensitivity to ozone.Environmental and Experimental Botany,2007,61:190-198
[11] Warren CR,L?w M,Matyssek R,et al.Internal conductance to CO2 transfer of adult Fagus sylvatica:Variation between sun and shade leaves and due to free-air ozone fumigation.Environmental and Experimental Botany,2007,59:130-138
[12] Diaz-Espejo A,Nicolás E,Fernández JE.Seasonal evolution of diffusional limitations and photosynthetic capacity in olive under drought.Plant,Cell & Environment,2007,30:922-933
[13] Warren CR.Does growth temperature affect the temperature response of photosynthesis and internal conductance to CO2?A test with Eucalyptus regnans.Tree Physiology,2008,28:11-19
[14] Warren CR,Livingston NJ,Turpin DH.Water stress decreases the transfer conductance of Douglas-fir (Pseudotsuga menziesii) seedlings.Tree Physiology,2004,24:971-979
[15] Evans JR,Sharkey TD,Berry JA,et al.Carbon isotope discrimination measured concurrently with gas exchange to investigate CO2 diffusion in leaves of higher plants.Australian Journal of Plant Physiology,1986,13:281-292
[16] Evans JR,Terashima I.Photosynthetic characteristics of spinach leaves grown with different treatments.Plant and Cell Physiology,1988,29:157-165
[17] Ren B (任博),Li J (李俊),Tong X-J (同小娟),et al.Simulation on photosynthetic-CO2 response of Quercus variabilis and Robinia pseudoacacia in the southern foot of the Taihang Mountain,China.Chinese Journal of Applied Ecology (应用生态学报),2018,29(1):1-10 (in Chinese)
[18] Monteith JL.Solar radiation and productivity in tropical ecosystem.Journal of Applied Ecology,1972,9:747-766
[19] Monteith JL,Moss CJ.Climate and the Efficiency of Crop Production in Britain.Philosophical Transactions of the Royal Society of London B,1977,281:277-294
[20] Yu Q (于强),Ren B-H (任保华),Wang T-D (王天铎),et al.Simulations of diurnal variation of photosynthesis in C3 Plants.Chinese Journal of Atmospheric Sciences (大气科学),1998,22(6):867-880 (in Chinese)
[21] Di Marco G,Manes F,Tricoli D,et al.Fluorescence parameters measured concurrently with net photosynthesis to investigate chloroplastic CO2 concentration in leaves of Quercus ilex L.Journal of Plant Physiology,1990,136:538-543
[22] Sharkey TD,Bernacchi CJ,Farquhar GD,et al.Fitting photosynthetic carbon dioxide response curves for C3 leaves.Plant,Cell & Environment,2007,30:1035-1040
[23] Sharkey TD,Vassey TL,Vanderveer PJ,et al.Carbon metabolism enzymes and photosynthesis in transgenic tobacco (Nicotiana tabacum L.) having excess phytochrome.Planta,1991,185:287-296
[24] Loreto F,Harley PC,Marco GD,et al.Estimation of mesophyll conductance to CO2 flux by three different methods.Plant Physiology,1992,98:1437-1443
[25] Sage RF.A model describing the regulation of ribulose-1,5-bisphosphate carboxylase,electron transport,and triose phosphate use in response to light intensity and CO2 in C3 plants.Plant Physiology,1990,94:1728-1734
[26] Manter DK,Kerrigan J.A/Ci curve analysis across a range of woody plant species:Influence of regression analysis parameters and mesophyll conductance.Journal of Experimental Botany,2004,55:2581-2588
[27] Li Y (李勇),Peng S-B (彭少兵),Huang J-L (黄见良),et al.Components and magnitude of mesophyll conductance and its responses to environmental variations.Plant Physiology Journal (植物生理学报),2013,49(11):1143-1154 (in Chinese)
[28] Guo S-W (郭世伟),Ran W (冉伟),Zhou Y (周毅),et al.On carbon and nitrogen metabolism of rice plants under elevated CO2 conditions.Chinese Journal of Rice Science (中国水稻科学),2006,20(5):560-566 (in Chinese)
[29] von Caemmerer S,Evans JR,Hudson GS,et al.The kinetics of ribulose-1,5-bisphosphate carboxylase/oxygenase in vivo inferred from measurements of photosynthesis in leaves of transgenic tobacco.Planta,1994,195:88-97
[30] von Caemmerer S.Biochemical Models of Leaf Photosynthesis.Collingwood,Victoria,Australia:CSIRO Publishing,2000
[31] Long SP,Bernacchi CJ.Gas exchange measurements,what can they tell us about the underlying limitations to photosynthesis?Procedures and sources of error.Journal of Experimental Botany,2003,54:2393-2401
[32] Warren C.Estimating the internal conductance to CO2 movement.Functional Plant Biology,2006,33:431-442
[33] Pons TL,Flexas J,von Caemmerer S,et al.Estimating mesophyll conductance to CO2:Methodology,potential errors,and recommendations.Journal of Experimental Botany,2009,60:2217-2234
[34] Genty B,Briantais JM,Baker NR.The relationship between the quantum yield of photosynthetic electron transport and quenching of chlorophyll fluorescence.Biochimica et Biophysica Acta,1989,990:87-92
[35] Laisk A,Loreto F.Determining photosynthetic parameters from leaf CO2 exchange and chlorophyll fluorescence.Ribulose-1,5-bisphosphate carboxylase/oxygenase specificity factor,dark respiration in the light,excitation distribution between photosystems,alternative electron transport rate,and mesophyll diffusion resistance.Plant Physiology,1996,110:903-912
[36] Harley PC,Loreto F,Di Marco G,et al.Theoretical considerations when estimating the mesophyll conductance to CO2 flux by the analysis of the response of photosynthesis to CO2.Plant Physiology,1992,98:1429-1436
[37] Pons TL,Westbeek MHM.Analysis of differences in photosynthetic nitrogen-use efficiency between four contrasting species.Physiologia Plantarum,2004,122:68-78
[38] Seaton GGR,Walker DA.Chlorophyll fluorescence as a measure of photosynthetic carbon assimilation.Proceedings of the Royal Society B:Biological Science,1990,242:29-35
[39] Pons TL,Welschen RAM.Midday depression of net photosynthesis in the tropical rainforest tree Eperua grandiflora:Contributions of stomatal and internal conductances,respiration and Rubisco functioning.Tree Physiology,2003,23:937-947
[40] Flexas J,Diaz-Espejo A,Galmes J,et al.Rapid variations of mesophyll conductance in response to changes in CO2 concentration around leaves.Plant,Cell & Environment,2007,30:1284-1298
[41] Bongi G,Loreto F.Gas-exchange properties of salt-stressed olive (Olea europea L) leaves.Plant Physiology,1989,90:1408-1416
[42] Hassiotou F,Ludwig M,Renton M,et al.Influence of leaf dry mass per area,CO2 and irradiance on mesophyll conductance in sclerophylls.Journal of Experimental Botany,2009,60:2303-2314
[43] Flexas J,Barón M,Bota J,et al.Photosynthesis limitations during water stress acclimation and recovery in the drought-adapted Vitis hybrid Richter-110 (V.berlandieri × V.rupestris).Journal of Experimental Botany,2009,60:2361-2377
[44] Vrabl D,Vaskova M,Hronkova M,et al.Mesophyll conductance to CO2 transport estimated by two independent methods:Effect of ambient CO2 concentration and abscisic acid.Journal of Experimental Botany,2009,60:2315-2323
[45] Yin X,Struik PC,Romero P,et al.Using combined measurements of gas exchange and chlorophyll fluorescence to estimate parameters of a biochemical C3 photosynthesis model:A critical appraisal and a new integrated approach applied to leaves in a wheat (Triticum aestivum) canopy.Plant,Cell & Environment,2009,32:448-464
[46] Martins SCV,Galmés J,Molins A,et al.Improving the estimation of mesophyll conductance to CO2:On the role of electron transport rate correction and respiration.Journal of Experimental Botany,2013,64:3285-3298
[47] Ehleringer J,Pearcy RW.Variation in quantum yield for CO2 uptake among C3 and C4 plants.Plant Physiology,1983,73:555-559
[48] Evans JR,Loreto F.Acquisition and diffusion of CO2 in higher plant leaves//Leegood RC,Sharkey TD,von Caemmerer S,eds,Photosynthesis:physiology and metabolism.Advances in photosynthesis.Dordrecht,The Netherlands:Kluwer Academic Publishers,2000:321-351
[49] Flexas J,Bota J,Escalona JM,et al.Effects of drought on photosynthesis in grapevines under field conditions:An evaluation of stomatal and mesophyll limitations.Functional Plant Biology,2002,29:461-471
[50] Grassi G and Magnani F.Stomatal,mesophyll conductance and biochemical limitations to photosynthesis as affected by drought and leaf ontogeny in ash and oak trees.Plant,Cell & Environment,2005,28:834-849
[51] Valentini R,Epron D,Angelis PDE,et al.In situ estimation of net CO2 assimilation,photosynthetic electron flow and photorespiration in Turkey oak (Q.cerris L.) leaves:Diurnal cycles under different levels of water supply.Plant,Cell & Environment,1995,18:631-640
[52] Evans JR,Poorter H.Photosynthetic acclimation of plants to growth irradiance:the relative importance of SLA and nitrogen partitioning in maximising carbon gain.Plant,Cell & Environment,2001,24:755-768
[53] Laisk AK.Kinetics of photosynthesis and photorespiration in C3 plants.Moscow,Russia:Nauka Publishing,1977
[54] von Caemmerer S,Evans JR.Determination of the average partial pressure of CO2 in chloroplasts from leaves of several C3 plants.Australian Journal Plant Physiology,1991,18:287-305
[55] Peisker M,Apel H.Inhibition by light of CO2 evolution from dark respiration:Comparison of two gas exchange methods.Photosynthesis Research,2001,70:291-298
[56] Sun JW,Guan DX,Wu JB,et al.Day and night respiration of three tree species in a temperate forest of northeastern China.iForest:Biogeosciences and Forestry,2015,8:25-32
[57] Bernacchi CJ,Singsaas EL,Pimentel C,et al.Improved temperature response functions for models of Rubisco-limited photosynthesis.Plant,Cell & Environment,2001,24:253-259
[58] Bernacchi CJ,Portis AR,Nakano H,et al.Temperature response of mesophyll conductance.Implications for the determination of Rubisco enzyme kinetics and for limitations to photosynthesis in vivo.Plant Physiology,2002,130:1992-1998
[59] Atkin OK,Scheurwater I,Pons TL.High thermal acclimation potential of both photosynthesis and respiration in two lowland Plantago species in contrast to an alpine congeneric.Global Change Biology,2006,12:500-515
[60] Théroux-Rancourt G,éthier G,Pepin S.Threshold response of mesophyll CO2 conductance to leaf hydraulics in highly transpiring hybrid poplar clones exposed to soil drying.Journal of Experimental Botany,2014,65:741-753
[61] Théroux-Rancourt G,éthier G,Pepin S.Greater efficiency of water use in poplar clones having a delayed response of mesophyll conductance to drought.Tree Physiology,2015,35:172-184
[62] Galmés J,Flexas J,Keys AJ,et al.Rubisco specificity factor tends to be larger in plant species from drier habitats and in species with persistent leaves.Plant,Cell & Environment,2005,28:571-579
[63] Ethier GH,Livingston NJ.On the need to incorporate sensitivity to CO2 transfer conductance into the Farquhar-von Caemmerer-Berry leaf photosynthesis model.Plant,Cell & Environment,2004,27:137-153
[64] Ethier GH,Livingston NJ,Harrison DL,et al.Low stomatal and internal conductance to CO2 versus Rubisco de activation as determinants of the photosynthetic decline of ageing evergreen leaves.Plant,Cell & Environment,2006,29:2168-2184
[65] von Caemmerer S,Evans JR.Determination of the average partial pressure of CO2 in chloroplasts of several C3 species.Australian Journal of Plant Physiology,1991,18:287-305
[66] Lloyd J,Syvertsen JP,Kriedeman PE,et al.Low conductances for CO2 diffusion from stomata to the sites of carboxylation in leaves of woody species.Plant,Cell & Environment,1992,15:873-899
[67] Bowling DR,Sargent SD,Tanner BD,et al.Tunable diode laser absorption spectroscopy for stable isotope studies of ecosystem-atmosphere CO2 exchange.Agricultural and Forest Meteorology,2003,118:1-19
[68] Farquhar GD,O’Leary MH,Berry JA.On the relationship between carbon isotope discrimination and the intercellular carbon dioxide concentration in leaves.Australian Journal of Plant Physiology,1982,9:121-137
[69] Gillon JS,Griffiths H.The influence of photorespiration on carbon isotope discrimination in plants.Plant,Cell & Environment,1997,20:1217-1230
[70] Ghashghaie J,Badeck F,Lanigan G,et al.Carbon isotope fractionation during dark respiration and photorespiration in C3 plants.Phytochemistry Reviews,2003,2:145-161
[71] Igamberdiev AU,Mikkelsen TN,Ambus P,et al.Photorespiration contributes to stomatal regulation and carbon isotope fractionation:A study with barley,potato and Arabidopsis plants deficient in glycine decarboxylase.Photosynthesis Research,2004,81:139-152
[72] Lanigan G,Betson N,Griffiths H,et al.Carbon isotope fractionation during photorespiration and carboxylation in Senecio.Plant Physiology,2008,148:2013-2020
[73] Brooks A,Farquhar GD.Effect of temperature on the CO2/O2 specificity of ribulose-1,5-bisphosphate carboxylase/oxygenase and the rate of respiration in the light.Planta,1985,165:397-406
[74] Lin G,Ehleringer JR.Carbon isotope fractionation does not occur during dark respiration in C3 and C4 plants.Plant Physiology,1997,114:391-394
[75] Shi Z-M (史作民),Feng Q-H (冯秋红),Cheng R-M (程瑞梅),et al.The research progress of mesophyll conductance.Acta Ecologica Sinica (生态学报),2010,30(17):4792-4803 (in Chinese)
[76] Warren C.Temperature response of photosynthesis and internal conductance to CO2:Results from two independent approaches.Journal of Experimental Botany,2006,57:3057-3067
[77] Xiong DL,Xi L,Liu LM,et al.Rapid responses of mesophyll conductance to changes of CO2 concentration,temperature and irradiance are affected by N supplements in rice.Plant,Cell & Environment,2015,38:2541-2550
[78] Wang XX,Du TT,Huang JL,et al.Leaf hydraulic vulnerability triggers the decline in stomatal and mesophyll conductance during drought in rice.Journal of Experimental Botany,2018,16:4033-4045
[2] Flexas J,Ribas-Carbó M,Diaz-Espejo A,et al.Mesophyll conductance to CO2:Current knowledge and future prospects.Plant,Cell & Environment,2008,31:602-621
[3] Farquhar GD,von Caemmerer S,Berry JA.A biochemical model of photosynthetic CO2 assimilation in leaves of C3 species.Planta,1980,149:78-90
[4] Han J-M (韩吉梅),Zhang W-F (张旺锋),Xiong D-L (熊栋梁),et al.Mesophyll conductance and its limiting factors in plant leaves.Chinese Journal of Plant Ecology (植物生态学报),2017,41(8):914-924 (in Chinese)
[5] Loreto F,Centritto M,Chartzoulakis K.Photosynthetic limitations in olive cultivars with different sensitivity to salt stress.Plant,Cell & Environment,2003,26:595-601
[6] Bernacchi CJ,Morgan PB,Ort DR,et al.The growth of soybean under free air [CO2] enrichment (FACE) stimulates photosynthesis while decreasing in vivo Rubisco capacity.Planta,2005,220:434-446
[7] Flexas J,Ribas-Carb M,Bota J.Decreased Rubisco activity during water stress is not induced by decreased relative water content but related to conditions of low stomatal conductance and chloroplast CO2 concentration.New Phytologist,2006,172:73-82
[8] Galmés J,Medrano H,Flexas J.Acclimation of Rubisco specificity factor to drought in tobacco:Discrepancies between in vitro and in vivo estimations.Journal of Experimental Botany,2006,57:3659-3667
[9] Galmés J,Medrano H,Flexas J.Photosynthetic limitations in response to water stress and recovery in Mediterranean plants with different growth forms.New Phytologist,2007,175:81-93
[10] Flowers MD,Fiscus EL,Burkey KO,et al.Photosynthesis,chlorophyll fluorescence,and yield of snap bean (Phaseolus vulgaris L.) genotypes differing in sensitivity to ozone.Environmental and Experimental Botany,2007,61:190-198
[11] Warren CR,L?w M,Matyssek R,et al.Internal conductance to CO2 transfer of adult Fagus sylvatica:Variation between sun and shade leaves and due to free-air ozone fumigation.Environmental and Experimental Botany,2007,59:130-138
[12] Diaz-Espejo A,Nicolás E,Fernández JE.Seasonal evolution of diffusional limitations and photosynthetic capacity in olive under drought.Plant,Cell & Environment,2007,30:922-933
[13] Warren CR.Does growth temperature affect the temperature response of photosynthesis and internal conductance to CO2?A test with Eucalyptus regnans.Tree Physiology,2008,28:11-19
[14] Warren CR,Livingston NJ,Turpin DH.Water stress decreases the transfer conductance of Douglas-fir (Pseudotsuga menziesii) seedlings.Tree Physiology,2004,24:971-979
[15] Evans JR,Sharkey TD,Berry JA,et al.Carbon isotope discrimination measured concurrently with gas exchange to investigate CO2 diffusion in leaves of higher plants.Australian Journal of Plant Physiology,1986,13:281-292
[16] Evans JR,Terashima I.Photosynthetic characteristics of spinach leaves grown with different treatments.Plant and Cell Physiology,1988,29:157-165
[17] Ren B (任博),Li J (李俊),Tong X-J (同小娟),et al.Simulation on photosynthetic-CO2 response of Quercus variabilis and Robinia pseudoacacia in the southern foot of the Taihang Mountain,China.Chinese Journal of Applied Ecology (应用生态学报),2018,29(1):1-10 (in Chinese)
[18] Monteith JL.Solar radiation and productivity in tropical ecosystem.Journal of Applied Ecology,1972,9:747-766
[19] Monteith JL,Moss CJ.Climate and the Efficiency of Crop Production in Britain.Philosophical Transactions of the Royal Society of London B,1977,281:277-294
[20] Yu Q (于强),Ren B-H (任保华),Wang T-D (王天铎),et al.Simulations of diurnal variation of photosynthesis in C3 Plants.Chinese Journal of Atmospheric Sciences (大气科学),1998,22(6):867-880 (in Chinese)
[21] Di Marco G,Manes F,Tricoli D,et al.Fluorescence parameters measured concurrently with net photosynthesis to investigate chloroplastic CO2 concentration in leaves of Quercus ilex L.Journal of Plant Physiology,1990,136:538-543
[22] Sharkey TD,Bernacchi CJ,Farquhar GD,et al.Fitting photosynthetic carbon dioxide response curves for C3 leaves.Plant,Cell & Environment,2007,30:1035-1040
[23] Sharkey TD,Vassey TL,Vanderveer PJ,et al.Carbon metabolism enzymes and photosynthesis in transgenic tobacco (Nicotiana tabacum L.) having excess phytochrome.Planta,1991,185:287-296
[24] Loreto F,Harley PC,Marco GD,et al.Estimation of mesophyll conductance to CO2 flux by three different methods.Plant Physiology,1992,98:1437-1443
[25] Sage RF.A model describing the regulation of ribulose-1,5-bisphosphate carboxylase,electron transport,and triose phosphate use in response to light intensity and CO2 in C3 plants.Plant Physiology,1990,94:1728-1734
[26] Manter DK,Kerrigan J.A/Ci curve analysis across a range of woody plant species:Influence of regression analysis parameters and mesophyll conductance.Journal of Experimental Botany,2004,55:2581-2588
[27] Li Y (李勇),Peng S-B (彭少兵),Huang J-L (黄见良),et al.Components and magnitude of mesophyll conductance and its responses to environmental variations.Plant Physiology Journal (植物生理学报),2013,49(11):1143-1154 (in Chinese)
[28] Guo S-W (郭世伟),Ran W (冉伟),Zhou Y (周毅),et al.On carbon and nitrogen metabolism of rice plants under elevated CO2 conditions.Chinese Journal of Rice Science (中国水稻科学),2006,20(5):560-566 (in Chinese)
[29] von Caemmerer S,Evans JR,Hudson GS,et al.The kinetics of ribulose-1,5-bisphosphate carboxylase/oxygenase in vivo inferred from measurements of photosynthesis in leaves of transgenic tobacco.Planta,1994,195:88-97
[30] von Caemmerer S.Biochemical Models of Leaf Photosynthesis.Collingwood,Victoria,Australia:CSIRO Publishing,2000
[31] Long SP,Bernacchi CJ.Gas exchange measurements,what can they tell us about the underlying limitations to photosynthesis?Procedures and sources of error.Journal of Experimental Botany,2003,54:2393-2401
[32] Warren C.Estimating the internal conductance to CO2 movement.Functional Plant Biology,2006,33:431-442
[33] Pons TL,Flexas J,von Caemmerer S,et al.Estimating mesophyll conductance to CO2:Methodology,potential errors,and recommendations.Journal of Experimental Botany,2009,60:2217-2234
[34] Genty B,Briantais JM,Baker NR.The relationship between the quantum yield of photosynthetic electron transport and quenching of chlorophyll fluorescence.Biochimica et Biophysica Acta,1989,990:87-92
[35] Laisk A,Loreto F.Determining photosynthetic parameters from leaf CO2 exchange and chlorophyll fluorescence.Ribulose-1,5-bisphosphate carboxylase/oxygenase specificity factor,dark respiration in the light,excitation distribution between photosystems,alternative electron transport rate,and mesophyll diffusion resistance.Plant Physiology,1996,110:903-912
[36] Harley PC,Loreto F,Di Marco G,et al.Theoretical considerations when estimating the mesophyll conductance to CO2 flux by the analysis of the response of photosynthesis to CO2.Plant Physiology,1992,98:1429-1436
[37] Pons TL,Westbeek MHM.Analysis of differences in photosynthetic nitrogen-use efficiency between four contrasting species.Physiologia Plantarum,2004,122:68-78
[38] Seaton GGR,Walker DA.Chlorophyll fluorescence as a measure of photosynthetic carbon assimilation.Proceedings of the Royal Society B:Biological Science,1990,242:29-35
[39] Pons TL,Welschen RAM.Midday depression of net photosynthesis in the tropical rainforest tree Eperua grandiflora:Contributions of stomatal and internal conductances,respiration and Rubisco functioning.Tree Physiology,2003,23:937-947
[40] Flexas J,Diaz-Espejo A,Galmes J,et al.Rapid variations of mesophyll conductance in response to changes in CO2 concentration around leaves.Plant,Cell & Environment,2007,30:1284-1298
[41] Bongi G,Loreto F.Gas-exchange properties of salt-stressed olive (Olea europea L) leaves.Plant Physiology,1989,90:1408-1416
[42] Hassiotou F,Ludwig M,Renton M,et al.Influence of leaf dry mass per area,CO2 and irradiance on mesophyll conductance in sclerophylls.Journal of Experimental Botany,2009,60:2303-2314
[43] Flexas J,Barón M,Bota J,et al.Photosynthesis limitations during water stress acclimation and recovery in the drought-adapted Vitis hybrid Richter-110 (V.berlandieri × V.rupestris).Journal of Experimental Botany,2009,60:2361-2377
[44] Vrabl D,Vaskova M,Hronkova M,et al.Mesophyll conductance to CO2 transport estimated by two independent methods:Effect of ambient CO2 concentration and abscisic acid.Journal of Experimental Botany,2009,60:2315-2323
[45] Yin X,Struik PC,Romero P,et al.Using combined measurements of gas exchange and chlorophyll fluorescence to estimate parameters of a biochemical C3 photosynthesis model:A critical appraisal and a new integrated approach applied to leaves in a wheat (Triticum aestivum) canopy.Plant,Cell & Environment,2009,32:448-464
[46] Martins SCV,Galmés J,Molins A,et al.Improving the estimation of mesophyll conductance to CO2:On the role of electron transport rate correction and respiration.Journal of Experimental Botany,2013,64:3285-3298
[47] Ehleringer J,Pearcy RW.Variation in quantum yield for CO2 uptake among C3 and C4 plants.Plant Physiology,1983,73:555-559
[48] Evans JR,Loreto F.Acquisition and diffusion of CO2 in higher plant leaves//Leegood RC,Sharkey TD,von Caemmerer S,eds,Photosynthesis:physiology and metabolism.Advances in photosynthesis.Dordrecht,The Netherlands:Kluwer Academic Publishers,2000:321-351
[49] Flexas J,Bota J,Escalona JM,et al.Effects of drought on photosynthesis in grapevines under field conditions:An evaluation of stomatal and mesophyll limitations.Functional Plant Biology,2002,29:461-471
[50] Grassi G and Magnani F.Stomatal,mesophyll conductance and biochemical limitations to photosynthesis as affected by drought and leaf ontogeny in ash and oak trees.Plant,Cell & Environment,2005,28:834-849
[51] Valentini R,Epron D,Angelis PDE,et al.In situ estimation of net CO2 assimilation,photosynthetic electron flow and photorespiration in Turkey oak (Q.cerris L.) leaves:Diurnal cycles under different levels of water supply.Plant,Cell & Environment,1995,18:631-640
[52] Evans JR,Poorter H.Photosynthetic acclimation of plants to growth irradiance:the relative importance of SLA and nitrogen partitioning in maximising carbon gain.Plant,Cell & Environment,2001,24:755-768
[53] Laisk AK.Kinetics of photosynthesis and photorespiration in C3 plants.Moscow,Russia:Nauka Publishing,1977
[54] von Caemmerer S,Evans JR.Determination of the average partial pressure of CO2 in chloroplasts from leaves of several C3 plants.Australian Journal Plant Physiology,1991,18:287-305
[55] Peisker M,Apel H.Inhibition by light of CO2 evolution from dark respiration:Comparison of two gas exchange methods.Photosynthesis Research,2001,70:291-298
[56] Sun JW,Guan DX,Wu JB,et al.Day and night respiration of three tree species in a temperate forest of northeastern China.iForest:Biogeosciences and Forestry,2015,8:25-32
[57] Bernacchi CJ,Singsaas EL,Pimentel C,et al.Improved temperature response functions for models of Rubisco-limited photosynthesis.Plant,Cell & Environment,2001,24:253-259
[58] Bernacchi CJ,Portis AR,Nakano H,et al.Temperature response of mesophyll conductance.Implications for the determination of Rubisco enzyme kinetics and for limitations to photosynthesis in vivo.Plant Physiology,2002,130:1992-1998
[59] Atkin OK,Scheurwater I,Pons TL.High thermal acclimation potential of both photosynthesis and respiration in two lowland Plantago species in contrast to an alpine congeneric.Global Change Biology,2006,12:500-515
[60] Théroux-Rancourt G,éthier G,Pepin S.Threshold response of mesophyll CO2 conductance to leaf hydraulics in highly transpiring hybrid poplar clones exposed to soil drying.Journal of Experimental Botany,2014,65:741-753
[61] Théroux-Rancourt G,éthier G,Pepin S.Greater efficiency of water use in poplar clones having a delayed response of mesophyll conductance to drought.Tree Physiology,2015,35:172-184
[62] Galmés J,Flexas J,Keys AJ,et al.Rubisco specificity factor tends to be larger in plant species from drier habitats and in species with persistent leaves.Plant,Cell & Environment,2005,28:571-579
[63] Ethier GH,Livingston NJ.On the need to incorporate sensitivity to CO2 transfer conductance into the Farquhar-von Caemmerer-Berry leaf photosynthesis model.Plant,Cell & Environment,2004,27:137-153
[64] Ethier GH,Livingston NJ,Harrison DL,et al.Low stomatal and internal conductance to CO2 versus Rubisco de activation as determinants of the photosynthetic decline of ageing evergreen leaves.Plant,Cell & Environment,2006,29:2168-2184
[65] von Caemmerer S,Evans JR.Determination of the average partial pressure of CO2 in chloroplasts of several C3 species.Australian Journal of Plant Physiology,1991,18:287-305
[66] Lloyd J,Syvertsen JP,Kriedeman PE,et al.Low conductances for CO2 diffusion from stomata to the sites of carboxylation in leaves of woody species.Plant,Cell & Environment,1992,15:873-899
[67] Bowling DR,Sargent SD,Tanner BD,et al.Tunable diode laser absorption spectroscopy for stable isotope studies of ecosystem-atmosphere CO2 exchange.Agricultural and Forest Meteorology,2003,118:1-19
[68] Farquhar GD,O’Leary MH,Berry JA.On the relationship between carbon isotope discrimination and the intercellular carbon dioxide concentration in leaves.Australian Journal of Plant Physiology,1982,9:121-137
[69] Gillon JS,Griffiths H.The influence of photorespiration on carbon isotope discrimination in plants.Plant,Cell & Environment,1997,20:1217-1230
[70] Ghashghaie J,Badeck F,Lanigan G,et al.Carbon isotope fractionation during dark respiration and photorespiration in C3 plants.Phytochemistry Reviews,2003,2:145-161
[71] Igamberdiev AU,Mikkelsen TN,Ambus P,et al.Photorespiration contributes to stomatal regulation and carbon isotope fractionation:A study with barley,potato and Arabidopsis plants deficient in glycine decarboxylase.Photosynthesis Research,2004,81:139-152
[72] Lanigan G,Betson N,Griffiths H,et al.Carbon isotope fractionation during photorespiration and carboxylation in Senecio.Plant Physiology,2008,148:2013-2020
[73] Brooks A,Farquhar GD.Effect of temperature on the CO2/O2 specificity of ribulose-1,5-bisphosphate carboxylase/oxygenase and the rate of respiration in the light.Planta,1985,165:397-406
[74] Lin G,Ehleringer JR.Carbon isotope fractionation does not occur during dark respiration in C3 and C4 plants.Plant Physiology,1997,114:391-394
[75] Shi Z-M (史作民),Feng Q-H (冯秋红),Cheng R-M (程瑞梅),et al.The research progress of mesophyll conductance.Acta Ecologica Sinica (生态学报),2010,30(17):4792-4803 (in Chinese)
[76] Warren C.Temperature response of photosynthesis and internal conductance to CO2:Results from two independent approaches.Journal of Experimental Botany,2006,57:3057-3067
[77] Xiong DL,Xi L,Liu LM,et al.Rapid responses of mesophyll conductance to changes of CO2 concentration,temperature and irradiance are affected by N supplements in rice.Plant,Cell & Environment,2015,38:2541-2550
[78] Wang XX,Du TT,Huang JL,et al.Leaf hydraulic vulnerability triggers the decline in stomatal and mesophyll conductance during drought in rice.Journal of Experimental Botany,2018,16:4033-4045