From leaf to whole-plant water use efficiency(WUE)in complex canopies:Limitations of leaf WUE as a selection target
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摘要
Plant water use efficiency(WUE) is becoming a key issue in semiarid areas, where crop production relies on the use of large volumes of water. Improving WUE is necessary for securing environmental sustainability of food production in these areas. Given that climate change predictions include increases in temperature and drought in semiarid regions,improving crop WUE is mandatory for global food production. WUE is commonly measured at the leaf level, because portable equipment for measuring leaf gas exchange rates facilitates the simultaneous measurement of photosynthesis and transpiration. However,when those measurements are compared with daily integrals or whole-plant estimates of WUE, the two sometimes do not agree. Scaling up from single-leaf to whole-plant WUE was tested in grapevines in different experiments by comparison of daily integrals of instantaneous water use efficiency [ratio between CO2assimilation(AN) and transpiration(E); AN/E] with midday AN/E measurements, showing a low correlation, being worse with increasing water stress. We sought to evaluate the importance of spatial and temporal variation in carbon and water balances at the leaf and plant levels. The leaf position(governing average light interception) in the canopy showed a marked effect on instantaneous and daily integrals of leaf WUE. Night transpiration and respiration rates were also evaluated, as well as respiration contributions to total carbon balance. Two main components were identified as filling the gap between leaf and whole plant WUE: the large effect of leaf position on daily carbon gain and water loss and the large flux of carbon losses by dark respiration. These results show that WUE evaluation among genotypes or treatments needs to be revised.
Plant water use efficiency(WUE) is becoming a key issue in semiarid areas, where crop production relies on the use of large volumes of water. Improving WUE is necessary for securing environmental sustainability of food production in these areas. Given that climate change predictions include increases in temperature and drought in semiarid regions,improving crop WUE is mandatory for global food production. WUE is commonly measured at the leaf level, because portable equipment for measuring leaf gas exchange rates facilitates the simultaneous measurement of photosynthesis and transpiration. However,when those measurements are compared with daily integrals or whole-plant estimates of WUE, the two sometimes do not agree. Scaling up from single-leaf to whole-plant WUE was tested in grapevines in different experiments by comparison of daily integrals of instantaneous water use efficiency [ratio between CO2assimilation(AN) and transpiration(E); AN/E] with midday AN/E measurements, showing a low correlation, being worse with increasing water stress. We sought to evaluate the importance of spatial and temporal variation in carbon and water balances at the leaf and plant levels. The leaf position(governing average light interception) in the canopy showed a marked effect on instantaneous and daily integrals of leaf WUE. Night transpiration and respiration rates were also evaluated, as well as respiration contributions to total carbon balance. Two main components were identified as filling the gap between leaf and whole plant WUE: the large effect of leaf position on daily carbon gain and water loss and the large flux of carbon losses by dark respiration. These results show that WUE evaluation among genotypes or treatments needs to be revised.
Plant water use efficiency(WUE) is becoming a key issue in semiarid areas, where crop production relies on the use of large volumes of water. Improving WUE is necessary for securing environmental sustainability of food production in these areas. Given that climate change predictions include increases in temperature and drought in semiarid regions,improving crop WUE is mandatory for global food production. WUE is commonly measured at the leaf level, because portable equipment for measuring leaf gas exchange rates facilitates the simultaneous measurement of photosynthesis and transpiration. However,when those measurements are compared with daily integrals or whole-plant estimates of WUE, the two sometimes do not agree. Scaling up from single-leaf to whole-plant WUE was tested in grapevines in different experiments by comparison of daily integrals of instantaneous water use efficiency [ratio between CO2assimilation(AN) and transpiration(E); AN/E] with midday AN/E measurements, showing a low correlation, being worse with increasing water stress. We sought to evaluate the importance of spatial and temporal variation in carbon and water balances at the leaf and plant levels. The leaf position(governing average light interception) in the canopy showed a marked effect on instantaneous and daily integrals of leaf WUE. Night transpiration and respiration rates were also evaluated, as well as respiration contributions to total carbon balance. Two main components were identified as filling the gap between leaf and whole plant WUE: the large effect of leaf position on daily carbon gain and water loss and the large flux of carbon losses by dark respiration. These results show that WUE evaluation among genotypes or treatments needs to be revised.
引文
[1]J.L.Araus,The problems of sustainable water use in the Mediterranean and research requirements for agriculture,Ann.Appl.Biol.144(2004)259–272.
[2]X.P.Deng,L.Shan,H.Zhang,N.C.Turner,Improving agricultural water use efficiency in arid and semiarid areas of China,Agric.Water Manag.80(2006)23–40.
[3]S.Geerts,N.Raes,Deficit irrigation as an on-farm strategy to maximize crop water productivity in dry areas,Agric.Water Manag.96(2009)1275–1284.
[4]N.Katerji,M.Mastrorilli,G.Ranab,Water use efficiency of crops cultivated in the Mediterranean region:review and analysis,Eur.J.Agron.28(2008)493–507.
[5]J.I.L.Morison,N.R.Baker,P.M.Mullineaux,W.J.Davies,Improving water use in crop production,Philos.T.R.Soc,B363(2008)639–658.
[6]IPCC,T.F.Stocker,Q.Dahe,G.K.Plattner,M.Tignor,S.K.Allen,J.Boschung,A.Nauels,Y.Xia,V.Bex,P.M.Midgley,Climate change 2013:the physical science basis,Working Group I Contribution to the Fifth Assessment Report of the Intergovernmental Panel on Climate Change,Cambridge University Press,Cambridge and New York,2013.
[7]H.G.Jones,R.A.Vaughan,Remote Sensing of Vegetation:Principles,Techniques,and Applications,Oxford University Press,Oxford,2010.
[8]S.M.Vicente-Serrano,J.I.Lopez-Moreno,S.Beguería,J.Lorenzo-Lacruz,A.Sanchez-Lorenzo,J.M.García-Ruiz,C.Azorin-Molina,E.Morán-Tejeda,J.Revuelto,R.Trigo,F.Coelho,F.Espejo,Evidence of increasing drought severity caused by temperature rise in Southern Europe,Environ.Res.Lett.9(2014)044001.
[9]M.M.Chaves,T.P.Santos,C.R.Souza,M.F.Ortuno,M.L.Rodrigues,C.M.Lopes,J.P.Maroco,J.S.Pereira,Deficit irrigation in grapevine improves water-use-efficiency without controlling vigour and production quality,Ann.Appl.Biol.150(2007)237–252.
[10]J.Flexas,J.Galmés,A.Gallé,J.Gulías,A.Pou,M.Ribas-Carbó,M.Tomás,H.Medrano,Improving water use efficiency in grapevines:potential physiological targets for biotechnological improvement,Aust.J.Grape Wine Res.161(2010)106–121.
[11]L.E.Williams,J.E.Ayars,Grapevine water use and the crop coefficient are linear functions of the shaded area measured beneath the canopy,Agric.For.Meteorol.132(2005)201–211.
[12]V.O.Sadras,Does partial root-zone drying improve irrigation water productivity in the field?A meta-analysis,Irrig.Sci.27(2009)183–190.
[13]L.E.Williams,Interaction of rootstock and applied water amounts at various fractions of estimated evapotranspiration(ETc)on productivity of Cabernet sauvignon,Aust.J.Grape Wine Res.3(2010)434–444.
[14]M.M.Chaves,O.Zarrouk,R.Francisco,J.M.Costa,T.Santos,A.P.Regalado,L.Rodrigues,C.M.Lopes,Grapevine under deficit irrigation:hints from physiological and molecular data,Ann.Bot.105(2010)661–676.
[15]P.Romero,J.I.Fernández-Fernández,A.Martinez-Cutillas,Physiological thresholds for efficient regulated deficitirrigation management in wine grapes grown under semiarid conditions,Am.J.Enol.Vitic.61(2010)300–312.
[16]J.Satisha,G.S.Prakash,R.Venugopalan,Statistical modeling of the effect of physio-biochemical parameters on water use efficiency of grape varieties,rootstocks and their stionic combinations under moisture stress conditions,Turk.J.Agric.For.30(2006)261–271.
[17]M.M.Alsina,F.de Herralde,X.Aranda,R.Save,C.Biel,Water relations and vulnerability to embolism are not related:experiments with eight grapevine cultivars,Vitis 46(2007)1–6.
[18]M.Tomás,M.Medrano,A.Pou,J.M.Escalona,S.Martorell,M.Ribas-Carbó,J.Flexas,Water use efficiency in grapevine cultivars:effects of water stress at leaf and whole plant level,Aust.J.Grape Wine Res.18(2012)164–172.
[19]P.Romero,R.Munoz,F.Del Amor,E.Valdes,J.I.Fernández,A.Martinez-Cutillas,Regulated deficit irrigation based upon optimum water status improves phenolic composition in Monastrell grapes in wines,Agric.Water Manag.121(2013)85–101.
[20]L.E.Williams,M.A.Matthews,Irrigation of agricultural crops.Agronomy monographs no.30,in:B.J.Stewart,D.R.Nielsen(Eds.),Experimental Agriculture,Cambridge University Press,Cambridge 1990,pp.1019–1055.
[21]H.Medrano,J.Gulías,M.Chaves,J.Galmés,J.Flexas,Photosynthesis water-use efficiency,in:J.Flexas,F.Loreto,H.Medrano(Eds.),Terrestrial Photosynthesis in a Changing Environment,A Molecular,Physiological and Ecological Approach Cambridge University Press,Cambridge 2012,pp.529–543.
[22]H.Medrano,J.Flexas,M.Ribas-Carbó,J.Gulías,Measuring water use efficiency in grapevines,in:S.Delrot,H.Medrano,E.Or,L.Bavaresco,S.Grando(Eds.),Methodologies and Results Grapevine Research,Springer,Germany 2010,pp.57–60.
[23]M.R.Gibberd,R.R.Walker,D.H.Blackmore,A.G.Condon,Transpiration efficiency and carbon-isotope discrimination of grapevines grown under well-watered conditions in either glasshouse or vineyard,Aust.J.Grape Wine Res.7(2001)110–117.
[24]S.Poni,F.Bernizzoni,S.Civardi,M.Gatti,D.Porro,F.Camin,Performance and water-use efficiency(single-leaf vs.wholecanopy)of well-watered and half-stressed split-root Lambrusco grapevines grown in Po Valley(Italy),Agric.Ecosyst.Environ.129(2009)97–106.
[25]J.M.Tarara,J.E.Pérez-Pena,R.P.Schreiner,M.Keller,P.Smithyman,Net carbon exchange in grapevine canopies responds rapidly to timing and extent of regulated deficit irrigation,Funct.Plant Biol.38(2011)386–400.
[26]S.Martorell,H.Medrano,M.Tomás,J.M.Escalona,J.Flexas,A.Díaz-Espejo,Plasticity of vulnerability to leaf hydraulic dysfunction during acclimation to drought in grapevines:an osmotic-mediated process,Physiol.Plant.153(2015)381–391.
[27]J.M.Escalona,S.Fuentes,M.Tomás,S.Martorell,J.Flexas,H.Medrano,Responses of leaf night transpiration to drought stress in Vitis vinifera L,Agric.Water Manag.118(2013)50–58.
[28]J.M.Escalona,M.Tomás,S.Martorell,H.Medrano,M.Ribas-Carbó,J.Flexas,Carbon balance in grapevines under different soil water supply:importance of whole plant respiration,Aust.J.Grape Wine Res.18(2012)308–318.
[29]H.Medrano,A.Pou,M.Tomás,S.Martorell,J.Gulias,J.Flexas,J.M.Escalona,Average daily light interception determines leaf water use efficiency among different canopy locations in grapevine,Agric.Water Manag.114(2012)4–10.
[30]J.M.Escalona,J.Flexas,J.Bota,H.Medrano,Distribution of leaf photosynthesis and transpiration within grapevine canopies under different drought conditions,Vitis 42(2003)57–64.
[31]T.N.Buckley,S.Martorell,A.Díaz-Espejo,M.Tomás,H.Medrano,Is stomatal conductance optimized over both time and space in plant crowns?A field test in grapevine(Vitis vinifera),Plant Cell Environ.27(2014)1–15.
[32]S.Fuentes,R.De Bei,M.Collins,J.M.Escalona,H.Medrano,S.D.Tyerman,Night time responses to water supply in grapevines(Vitis vinifera L.)under deficit irrigation and partial root-zone drying,Agric.Water Manag.138(2014)1–9.
[33]K.Morinaga,S.Imai,H.Yakushiji,Y.Koshita,Effects of fruit load on partitioning of15N and13C,respiration,and growth of grapevine roots at different fruit stages,Sci.Hortic.97(2003)239–253.
[34]Z.W.Dai,P.Vivin,F.Barrieu,N.Ollat,S.Delrot,Physiological and modelling approaches to understand water and carbon fluxes during grape berry growth and quality development:a review,Aust.J.Grape Wine Res.16(2010)70–85.
[35]N.Franck,J.P.Morales,D.Arancibia-Avendano,V.G.de Cortazar,J.F.Pérez-Quezada,A.Zurita-Silva,C.Pastenes,Seasonal fluctuations in Vitis vinifera root respiration in the field,New Phytol.192(2011)939–951.
[36]M.Tomás,H.Medrano,J.M.Escalona,S.Martorell,A.Pou,M.Ribas-Carbó,J.Flexas,Genetic variability in water use efficiency in grapevines,Environ.Exp.Bot.103(2014)148–157.
[37]H.Medrano,M.Tomás,S.Martorell,J.M.Escalona,A.Pou,S.Fuentes,J.Flexas,J.Bota,Improving water use efficiency of vineyards in semiarid regions:a review,Agron.Sustain.Dev.35(2015)499–517.
[2]X.P.Deng,L.Shan,H.Zhang,N.C.Turner,Improving agricultural water use efficiency in arid and semiarid areas of China,Agric.Water Manag.80(2006)23–40.
[3]S.Geerts,N.Raes,Deficit irrigation as an on-farm strategy to maximize crop water productivity in dry areas,Agric.Water Manag.96(2009)1275–1284.
[4]N.Katerji,M.Mastrorilli,G.Ranab,Water use efficiency of crops cultivated in the Mediterranean region:review and analysis,Eur.J.Agron.28(2008)493–507.
[5]J.I.L.Morison,N.R.Baker,P.M.Mullineaux,W.J.Davies,Improving water use in crop production,Philos.T.R.Soc,B363(2008)639–658.
[6]IPCC,T.F.Stocker,Q.Dahe,G.K.Plattner,M.Tignor,S.K.Allen,J.Boschung,A.Nauels,Y.Xia,V.Bex,P.M.Midgley,Climate change 2013:the physical science basis,Working Group I Contribution to the Fifth Assessment Report of the Intergovernmental Panel on Climate Change,Cambridge University Press,Cambridge and New York,2013.
[7]H.G.Jones,R.A.Vaughan,Remote Sensing of Vegetation:Principles,Techniques,and Applications,Oxford University Press,Oxford,2010.
[8]S.M.Vicente-Serrano,J.I.Lopez-Moreno,S.Beguería,J.Lorenzo-Lacruz,A.Sanchez-Lorenzo,J.M.García-Ruiz,C.Azorin-Molina,E.Morán-Tejeda,J.Revuelto,R.Trigo,F.Coelho,F.Espejo,Evidence of increasing drought severity caused by temperature rise in Southern Europe,Environ.Res.Lett.9(2014)044001.
[9]M.M.Chaves,T.P.Santos,C.R.Souza,M.F.Ortuno,M.L.Rodrigues,C.M.Lopes,J.P.Maroco,J.S.Pereira,Deficit irrigation in grapevine improves water-use-efficiency without controlling vigour and production quality,Ann.Appl.Biol.150(2007)237–252.
[10]J.Flexas,J.Galmés,A.Gallé,J.Gulías,A.Pou,M.Ribas-Carbó,M.Tomás,H.Medrano,Improving water use efficiency in grapevines:potential physiological targets for biotechnological improvement,Aust.J.Grape Wine Res.161(2010)106–121.
[11]L.E.Williams,J.E.Ayars,Grapevine water use and the crop coefficient are linear functions of the shaded area measured beneath the canopy,Agric.For.Meteorol.132(2005)201–211.
[12]V.O.Sadras,Does partial root-zone drying improve irrigation water productivity in the field?A meta-analysis,Irrig.Sci.27(2009)183–190.
[13]L.E.Williams,Interaction of rootstock and applied water amounts at various fractions of estimated evapotranspiration(ETc)on productivity of Cabernet sauvignon,Aust.J.Grape Wine Res.3(2010)434–444.
[14]M.M.Chaves,O.Zarrouk,R.Francisco,J.M.Costa,T.Santos,A.P.Regalado,L.Rodrigues,C.M.Lopes,Grapevine under deficit irrigation:hints from physiological and molecular data,Ann.Bot.105(2010)661–676.
[15]P.Romero,J.I.Fernández-Fernández,A.Martinez-Cutillas,Physiological thresholds for efficient regulated deficitirrigation management in wine grapes grown under semiarid conditions,Am.J.Enol.Vitic.61(2010)300–312.
[16]J.Satisha,G.S.Prakash,R.Venugopalan,Statistical modeling of the effect of physio-biochemical parameters on water use efficiency of grape varieties,rootstocks and their stionic combinations under moisture stress conditions,Turk.J.Agric.For.30(2006)261–271.
[17]M.M.Alsina,F.de Herralde,X.Aranda,R.Save,C.Biel,Water relations and vulnerability to embolism are not related:experiments with eight grapevine cultivars,Vitis 46(2007)1–6.
[18]M.Tomás,M.Medrano,A.Pou,J.M.Escalona,S.Martorell,M.Ribas-Carbó,J.Flexas,Water use efficiency in grapevine cultivars:effects of water stress at leaf and whole plant level,Aust.J.Grape Wine Res.18(2012)164–172.
[19]P.Romero,R.Munoz,F.Del Amor,E.Valdes,J.I.Fernández,A.Martinez-Cutillas,Regulated deficit irrigation based upon optimum water status improves phenolic composition in Monastrell grapes in wines,Agric.Water Manag.121(2013)85–101.
[20]L.E.Williams,M.A.Matthews,Irrigation of agricultural crops.Agronomy monographs no.30,in:B.J.Stewart,D.R.Nielsen(Eds.),Experimental Agriculture,Cambridge University Press,Cambridge 1990,pp.1019–1055.
[21]H.Medrano,J.Gulías,M.Chaves,J.Galmés,J.Flexas,Photosynthesis water-use efficiency,in:J.Flexas,F.Loreto,H.Medrano(Eds.),Terrestrial Photosynthesis in a Changing Environment,A Molecular,Physiological and Ecological Approach Cambridge University Press,Cambridge 2012,pp.529–543.
[22]H.Medrano,J.Flexas,M.Ribas-Carbó,J.Gulías,Measuring water use efficiency in grapevines,in:S.Delrot,H.Medrano,E.Or,L.Bavaresco,S.Grando(Eds.),Methodologies and Results Grapevine Research,Springer,Germany 2010,pp.57–60.
[23]M.R.Gibberd,R.R.Walker,D.H.Blackmore,A.G.Condon,Transpiration efficiency and carbon-isotope discrimination of grapevines grown under well-watered conditions in either glasshouse or vineyard,Aust.J.Grape Wine Res.7(2001)110–117.
[24]S.Poni,F.Bernizzoni,S.Civardi,M.Gatti,D.Porro,F.Camin,Performance and water-use efficiency(single-leaf vs.wholecanopy)of well-watered and half-stressed split-root Lambrusco grapevines grown in Po Valley(Italy),Agric.Ecosyst.Environ.129(2009)97–106.
[25]J.M.Tarara,J.E.Pérez-Pena,R.P.Schreiner,M.Keller,P.Smithyman,Net carbon exchange in grapevine canopies responds rapidly to timing and extent of regulated deficit irrigation,Funct.Plant Biol.38(2011)386–400.
[26]S.Martorell,H.Medrano,M.Tomás,J.M.Escalona,J.Flexas,A.Díaz-Espejo,Plasticity of vulnerability to leaf hydraulic dysfunction during acclimation to drought in grapevines:an osmotic-mediated process,Physiol.Plant.153(2015)381–391.
[27]J.M.Escalona,S.Fuentes,M.Tomás,S.Martorell,J.Flexas,H.Medrano,Responses of leaf night transpiration to drought stress in Vitis vinifera L,Agric.Water Manag.118(2013)50–58.
[28]J.M.Escalona,M.Tomás,S.Martorell,H.Medrano,M.Ribas-Carbó,J.Flexas,Carbon balance in grapevines under different soil water supply:importance of whole plant respiration,Aust.J.Grape Wine Res.18(2012)308–318.
[29]H.Medrano,A.Pou,M.Tomás,S.Martorell,J.Gulias,J.Flexas,J.M.Escalona,Average daily light interception determines leaf water use efficiency among different canopy locations in grapevine,Agric.Water Manag.114(2012)4–10.
[30]J.M.Escalona,J.Flexas,J.Bota,H.Medrano,Distribution of leaf photosynthesis and transpiration within grapevine canopies under different drought conditions,Vitis 42(2003)57–64.
[31]T.N.Buckley,S.Martorell,A.Díaz-Espejo,M.Tomás,H.Medrano,Is stomatal conductance optimized over both time and space in plant crowns?A field test in grapevine(Vitis vinifera),Plant Cell Environ.27(2014)1–15.
[32]S.Fuentes,R.De Bei,M.Collins,J.M.Escalona,H.Medrano,S.D.Tyerman,Night time responses to water supply in grapevines(Vitis vinifera L.)under deficit irrigation and partial root-zone drying,Agric.Water Manag.138(2014)1–9.
[33]K.Morinaga,S.Imai,H.Yakushiji,Y.Koshita,Effects of fruit load on partitioning of15N and13C,respiration,and growth of grapevine roots at different fruit stages,Sci.Hortic.97(2003)239–253.
[34]Z.W.Dai,P.Vivin,F.Barrieu,N.Ollat,S.Delrot,Physiological and modelling approaches to understand water and carbon fluxes during grape berry growth and quality development:a review,Aust.J.Grape Wine Res.16(2010)70–85.
[35]N.Franck,J.P.Morales,D.Arancibia-Avendano,V.G.de Cortazar,J.F.Pérez-Quezada,A.Zurita-Silva,C.Pastenes,Seasonal fluctuations in Vitis vinifera root respiration in the field,New Phytol.192(2011)939–951.
[36]M.Tomás,H.Medrano,J.M.Escalona,S.Martorell,A.Pou,M.Ribas-Carbó,J.Flexas,Genetic variability in water use efficiency in grapevines,Environ.Exp.Bot.103(2014)148–157.
[37]H.Medrano,M.Tomás,S.Martorell,J.M.Escalona,A.Pou,S.Fuentes,J.Flexas,J.Bota,Improving water use efficiency of vineyards in semiarid regions:a review,Agron.Sustain.Dev.35(2015)499–517.