大麦耐低温种质资源筛选和耐性机理研究
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摘要
低温是一种全球普遍发生、影响农作物生产的环境胁迫。作为世界上第四大谷类作物,大麦种植分布十分广泛。在冬大麦种植区,冬季极端低温一般并不导致苗期遭受严重冻害,而春季时常发生的低温寒潮常造成减产。防御这种灾害性天气,提高大麦生殖生长期低温耐性是最根本且有效的途径。本研究旨在探讨大麦耐低温种质资源筛选技术,并试图明确大麦营养生长期和生殖生长期低温耐性机制的差异,为大麦育种提供理论依据和种质筛选技术方法。主要研究结果如下:
     大麦等越冬作物耐低温的一个重要农艺特性是冷驯化。以苗期低温敏感性不同的两个大麦品种为材料,探讨了冷驯化和光强对大麦非致死冻害处理后恢复过程中叶片H_2O_2积累和ROS清除系统的影响,并利用叶绿素荧光技术研究恢复过程中光合器官功能的变化。结果表明:在冻害损伤发生后的恢复过程中测定叶绿素荧光参数可以直接了解植物体的生活力指标,从而明确低温和光照强度这两个重要的环境因子对相关指标的影响。冷驯化可能并不直接增强大麦的抗冻性,但它可显著提高大麦冻害损伤后的恢复能力,并显著增强冷驯化植株对过剩光能和光抑制的应对能力。
     细胞膜是植物受低温损伤的初始位点,且根系的稳定性在植物体抵御低温胁迫及其恢复生长过程中起着重要的作用。为检测大麦根系离子流对低温胁迫的反应,本研究在国内首次采用澳大利亚塔斯马尼亚大学Newman等研制的MIFE(Microelectrode Ion Flux Estimation)技术,比较了低温胁迫下3个大麦品种的H~+离子流和K~+离子流动力学,并研究不同温度胁迫对根系K~+离子吸收能力的影响。结果表明,利用MIFE技术进行离子流的测定,可以及时准确地监测到大麦根系离子流受低温胁迫而引起的细微变化,可为大麦低温胁迫生理研究和种质筛选提供一种潜在的方法。同时证明低温处理显著抑制大麦根系K~+的吸收,且在本实验的非致死低温胁迫后经过短暂的恢复,大麦根系K~+的吸收能力即可得到显著的恢复。
     光温变化显著影响植物光合器官的功能,从而抑制大麦生长和产量形成。本研究采用抽穗期低温耐性有一定差异的2个大麦品种,利用人工气候室进行低温肋、迫处理,分析低温胁迫后恢复过程中的光合作用相关参数,旨在明确大麦生殖生长期低温耐性的基因型差异及其生理机制。结果表明,在大麦抽穗期利用实验室(人工气候室)处理可以有效模拟自然低温条件,可作为大麦生殖生长期耐低温种质材料的鉴定方法之一。同时,本研究首次尝试利用离体穗培养的方法在大麦生殖生长期进行低温处理,通过研究低温对大麦离体穗的旗叶光合器官的影响,证明该方法可用于大麦生殖生长期耐低温种质的筛选,可克服因生育期差异而造成的大田筛选困难,也可减少整株植物体移栽需要的大量人力物力。同时,本研究采用这一方法对一个大麦DH群体进行了生殖生长期低温耐性相关基因的QTL分析,在3H染色体上检测到一个与低温耐性有关的QTL位点。
Low temperature is a major environmental stress for cereal crops worldwide, reducing yield potential and grain quality of barley in sensitive reproductive stages. Barley (Hordeum vulgare L.) is a major world crop ranked the fourth in planting areas among cereal crops. In general, extreme low temperature in winter does not cause considerable damage to barley seedlings in the winter-barley growing areas. However, frost or low temperature frequently occurring in spring affects barley grain production significantly, because the plants at reproductive stage are quite sensitive to low temperature. Developing of frost-tolerant barley in reproductive stages is one of basic and effective approaches to fignt against the low temperature during spring. This study is going to developing potential methods of screening barley germplasm with low temperature tolerance. Meanwhile a comprehensive study was undertaken to compare the difference in low temperature tolerance between vegetative and reproductive stage of barley plants.
     One of the most important agronomic traits in barley is freezing tolerance, which can be enhanced by cold acclimation. In order to clarify the mechanisms of alleviating injury by cold acclimation in vegetative stage, two winter barley cultivars with different cold tolerance in vegetative stage were used to study the effect of irradiance on leaf H_2O_2 accumulation and ROS scavenging systems after removal of unlethal freezing stress. Moreover, the functions of photosynthetic apparatus in leaves were measured by chlorophyll fluorescence during recovery. The results indicated that some chlorophyll fluorescence parameters during recovery from freezing treatment might be used as the indicators in identification and evaluation of cold tolerance in barley. The functions of photosynthetic apparatus were affected by irradiance after freezing stress. It can be concluded that cold acclimation may not directly enhance cold tolerance of barley plants, but it increased recovery ability from freezing injury, and the ability to cope with excess energy and photoinhibition was much improved in cold-acclimated plants.
     The primary sites of chilling action appear to be plant membranes, and the membrane stability in root plays a central role in cold stress tolerance and recovery from cold injury. MIFE technique, developed by Newman in University of Tasmania, Australia, was employed to determine the effect of low temperature stress on ion flux from barley roots. Root H~+ and K~+ flux of three barley cultivars were measured and the effect of low temperature on K~+ uptake by barley roots was studied. As a effective, non-invasive, and laboratory-based technique, MIFE technique can detect minimum fluctuation of ion flux caused by low temperature stress in barley roots. The possibility of using MIFE technique as a potential screening tool in identification and screening of barley genotypes with high frost tolerance was tested. The results showed the potential value of the technique. Meanwhile, it was proved that the ability of K~+ uptake by barley roots was significantly inhibited by low temperature stress, and it can be reversed dramatically after removal of un-lethal low temperature.
     The functions of photosynthetic apparatus were affected by irradiance and temperature. In order to clarify the variation of genotypes and mechanisms of frost tolerance at reproductive stage of barley, two winter barley cultivars with different frost tolerance at reproductive stage were used in this study. Photosynthetic apparatus were measured during recovery stage after low temperature stress. The results showed that growth chamber could be used to simulate low temperature stress at reproductive stage of barley, and to evaluate frost tolerance of barley genotypes. Moreover, In vitro culture method was employed to investigate the photosynthetic apparatus responses of detached flag leaves to low temperature stress. The evaluation of the differences among barley genotypes in their responses to low temperature occurring at reproductive stage is limited by uncertainty in a course of weather and different growth rate. The results indicated that In vitro culture may provide an effective method of screening and evaluating barley germplasm in low temperature tolerance at reproductive stage. Meanwhile, In vitro culture method was used to evaluate frost tolerance of a double haploid population, and the analysis of QTLs controlling frost tolerance was conducted. A QTL relevant to frost tolerance was located on chromosome 3H.
引文
Allen RD(1995) Dissection of oxidative stress tolerance using transgenic plants.Plant Physiology 107:1049-1054.
    Apel K,Hirt H(2004) Reactive oxygen species:metabolism,oxidative stress,and signal transduction.Annual Review of Plant Biology 55:373-399.
    Aro EM,Virgin I,Andersson B(1993)Photoinhibition of Photosystem Ⅱ.Inactivation,protein damage and turnover.Biochimica et Biophysica Acta 1143:113-134.
    Baek KH,Skinner DZ(2003)Alteration of antioxidant enzyme gene expression during cold acclimation of near-isogenic wheat lines.Plant Science 165:1221-1227.
    Baker NR,Rosenqvist E(2004)Applications of chlorophyll fluorescence can improve crop production strategies:an examination of future possibilities.Journal of Experimental Botany 55:1607-1621.
    Barber J(1995)Molecular basis of the vulnerability of photosystem Ⅱ to damage by light.Australian Journal of Plant Physiology 22:201-208.
    Barber J,Andersson B(1992)Too much of a good thing:light can be bad for photosynthesis.Trends in Biochemical Science 17:61-66.
    Belkhodja R,Morales F,Abadia A,Gomez-Aparisi J,Abadia J(1994)Chlorophyll fluorescence as a possible tool for salinity tolerance screening in barley(Hordeum vulgare L.).Plant Physiology 104:667-673.
    Bertin P,Bouharmont J,Kinet JM(1997)Somaclonal variation and improvement of chilling tolerance in rice:Changes in chilling-induced chlorophyll fluorescence.Crop Science 37:1727-1735.
    Chen HH,Brenner ML,Li PH(1983a)Involvement of abscisic acid in potato cold acclimation.Plant Physiology 71:362-365.
    Cheng SH,Willmann MR,Chen HC,Sheen J(2002)Calcium signaling through protein kinases.The Arabidopsis calcium-dependent protein kinase gene family.Plant Physiology 129:469-485.
    Chen THH,Gusta LV,Fowler DB(1983b)Freezing injury and root development in winter cereals.Plant Physiology 73:773-777.
    Dexter ST(1933)Effects of several environmental factors on hardening of plants.Plant Physiology 8:123-139.
    Duman,JG,Olsen TM(1993)Thermal hysteresis protein activity in bacteria,fungi,and phylogenetically diverse plants.Cryobiology 30:322-328.
    Eagles CF,Williams J,Louis DV(1993)Recovery after freezing in Avena sativa L.,Lolium perenne L.and L.multiorum Lam.New Phytologist 123:477-483.
    Elster EF(1982)Oxygen activation and oxygen toxicity.Annual Review of Plant Biology 33:69-731.
    Ensminger I,Busch F,Huner NPA(2006)Photostasis and cold acclimation:sensing low temperature through photosynthesis.Physiologia Plantarum 126:28-44.
    Fan,Y,Liu B,Wang H,Wang S,Wang J(2002)Cloning of antifreeze protein gene from carrot and its influence on cold tolerance in transgenic tobacco plants.Plant Cell Reports 21:296-301.
    Fowler DB,Limin AE,Ritchie JT(1999)Low-temperature tolerance in cereals:Model and genetic interpretation.Crop Science 39:626-633.
    Fowler S,Thomashow MF(2002)Arabidopsis transcriptome profiling indicates that multiple regulatory pathways are activated during cold acclimation in addition to the CBF cold response pathway.Plant Cell 14:1675-1690.
    Galiba G,Quarrie SA,Tuberosa R,Veisz O,Sutka J(1995)Regulation of freezing tolerance by abscisic acid in wheat.Acta Phytopathologica et Entomologica Hungarica 30:123-124.
    Gilmour SJ,Thomashow MF(1991)Cold acclimation and cold-regulated gene expression in ABA mutants of Arabidopsis thaliana.Plant Molecular Biology 17:1233-1240.
    Gilmour SJ,Sebolt AM,Salazar MP,Everard JD,Thomashow MF(2000)Overexpression of the Arabidopsis CBF3 transcriptional activator mimicsmultiple biochemical changes associated with cold acclimation.Plant Physiology 124:1854-1865.
    Gray GR,Chauvin LP,Sarhan F,Huner NPA(1997)Cold acclimation and freezing tolerance:a complex interaction of light and temperature.Plant Physiology 114:467-474.
    Gray GR,Hope BJ,Qin XQ,Taylor BG,Whitehead CL(2003)The characterization of photoinhibition and recovery during cold acclimation in Arabidopsis thaliana using chlorophyll fluorescence imaging.Physiologia Plantarum 119:365-375.
    Greer GH,Berry JA,Bjorkman O(1986)Photoinhibition of photosynthesis in intact bean leaves:role of light and temperature and requirement for chloroplast-protein synthesis during recovery.Planta 168:253-257.
    Griffith M,Mclntyre HCH(1993)The interrelationship of growth and frost tolerance in winter rye.Physiologia Plantarum 87:335-344.
    Griffith M,Yaish MWF(2004)Antifreeze proteins in overwintering plants:a tale of two activities.Trends in Plant Science 9:399-405.
    Griffith M,Ala P,Yang DSC,Hon WC,Moffatt BA(1992)Antifreeze protein produced endogenously in winter rye leaves.Plant Physiology 100:593-596.
    Griffith M,Lumb C,Wiseman SB,Johnson RW,Wisniewski M,Loewen MC(2003)Winter rye antifreeze proteins are not cryoprotective,rather they modify the growth of ice in plants.Acta Horticultural 618:31-37.
    Gupta R,Luan S(2003)Redox control of protein tyrosine phosphatases and mitogen-activated protein kinases in plants.Plant Physiology 132:1149-1152.
    Halliwell B,Gutteridge JMC(1999)Free Radicals in Biology and Medicine,Oxford University Press,New York.
    Hayes PM,Blake T,Chen TTH,Tragoonrung S,Chen F,Pan A,Liu B(1993)Quantitative trait loci on barley(Hordeum vulgare L.)chromosome 7 associated components of winter hardiness.Genome 36:66-71.
    Hetherington SE,He J,Smillie RM(1989)Photoinhibition at low-temperature in chilling-sensitive and chilling-resistant plants.Plant Physiology 90:1609-1615.
    Hon WC,Griffith M,Mlynarz A,Kwok YC,Yang DSC(1995)Antifreeze proteins in winter rye are similar to pathogenesis-related proteins.Plant Physiology 109:879-889.
    Hong Z(2000)Removal of feedback inhibited ion ofAl-pyrroling-5-carboxylate synthase results in increased proline accumulation and protection of plants from osmotic stress.Plant physiology 122:1129-1136.
    Hovenden MJ,Warren CR(1998)Photochemistry,energy dissipation and cold-hardening in Eucalyptus nitens and E.pauciflora.Australian Journal of Plant Physiology 25 581-589.
    Huang T,Nicodemus J,Zarka DG,Thomashow MF,Wisniewski M,Duman JG(2002)Expression of an insect(Dendroides canadensis)antifreeze protein in Arabidopsis thaliana results in a decrease in plant freezing temperature.Plant Molecular Biology 50:333-344.
    Hughes MA,Dunn MA(1996)The molecular biology of plant acclimation to low temperature.Journal of Experimental Botany 47:291-305.
    Huner NPA,Oquist G,Sarhan F(1998)Energy balance and acclimation to light and cold.Trends in Plant Science 3:224-230.
    Janda T,Szalai G,Gonzalez R K,Veisz O,Paldi E(2003)Comparative study of frost tolerance and antioxidant activity in cereals.Plant Science 164:301-306.
    Kendall EJ,McKersie BD(1989)Free radical and freezing injury to cell membranes of winter wheat.Physiologia Plantarum 76:86-94.
    Knight H,Trewavas AJ,Knight MR(1996)Cold calcium signaling in Arabidopsis involved two cellular pools and a change in calcium signature after acclimation.Plant Cell 8:489-503.
    Knight,CA,Wen D,Laursen RA(1995)Nonequilibrium antifreeze peptides and the recrystallization of ice.Cryobiology 32:23-34.
    Knowles AE,Shabala S(2004)Overcoming the Problem of Non-Ideal Liquid Ion Exchanger Selectivity in Microelectrode Ion Flux Measurements.Journal of Membrane Biology 202:51-59.
    Koch KE(1996)Carbohydrate-modulated gene expression in plants.Annual Review of Plant Physiology and Plant Molecular Biology 47:509-540.
    Koo BC,Bushman BS,Mott IW(2008)Transcripts associated with non-acclimated freezing response in two barley cultivars.The Plant Genome.1:21-32
    Koo BC,Park MW,Lee CW,Yoon EB,Kim KJ,An JG.(2003)Classification for types of damages caused by cold stress at different young spike development stages of barley and wheat.Korean Journal of Crop Science 48:252-261.
    Kornyeyev D,Logan BA,Payton P,Allen RD,Holaday AS(2001)Enhanced photochemical light utilization and decreased chilling-induced photoinhibition of photosystem Ⅱ in cotton overexpressing genes encoding chloroplast-targeted antioxidant enzymes.Physiologia Plantarum 113:323-331.
    Kurkela S,Franck M(1990)Cloning and characterization of a cold-and ABA-inducible Arabidopsis gene.Plant Molecular Biology 15:137-144.
    Lang V,Mantyla E,Welin B,Sundberg B,Palva ET(1994)Alterations in water status,endogenous abscisic acid content,and expression of rab18 gene during the development of freezing tolerance in Arabidopsis thaliana.Plant Physiology 104:1341-1349.
    Lucas WJ,Kochian LV(1986)Ion transport processes in corn roots:an approach utilizing microelectrode techniques.In:Gensler WG(ed.),Advanced Agricultural Instrumentation:Design and Use.Martinus Nijhoff,Dordrecht,The Netherlands.pp.402-425.
    Lyons JM(1973)Chilling injury in plants.Annual Review of Plant Physiology 24:445-466.
    Marcellos M(1988)Comments on frost injury in wheat.In:Frost injury in wheat:a workshop of the wheat research council.Haymarket,N.S.W.
    Marentes E,Griffith M,Mlynarz A,Brush RA(1993)Proteins accumulate in the apoplast of winter rye leaves during cold acclimation.Physiologia Plantarum 87:499-507.
    Martin M L,Busconi LA(2001)Rice membrane-bound calcium-dependent protein kinase is activated in response to low temperature.Plant Physiology 125:1442-1449.
    McKown R,Kuroki G,Warren G(1996)Cold response of Arabidopsis mutants imparled in freezing tolerance.Journal of Experiment Botany 107:1919-1925.
    Mittler R,Vanderauwera S,Gollery M,Breusegem FV(2004)Reactive oxygen gene network of plants.Trends in Plant Science 9:490-498,
    Mittler R(2002)Oxidative stress,antioxidants and stress tolerance.Trends in Plant Science 7:405-410.
    Monk LS,Fagerstedt KV,Crawford RMM(1989)Oxygen toxicity and superoxide dismutase as an antioxidant in physiological stress.Plant Physiology 76:456-459.
    Monroy AF,Castonguay Y,Laberge S,Sarhan F,Vezina LP,Dhindsa RS(1993)A new cold-induced alfalfa gene is associated with enhanced hardening at subzero temperature.Plant Physiology 102:873-879.
    Monroy AF,Dhindsa RS(1995)Low-temperature signal transduction:induction of cold acclimation-specific genes of alfalfa by calcium at 25 degrees C.Plant Cell 7:321-331.
    Monroy AF,Sangwan V,Dhindsa RS(1998)Low temperature signal transduction during cold acclimation:protein phosphatase 2A as an early target for cold-inactivation.The Plant Journal 13,653-660.
    NDong C,Danyluk J,Wilson KE,Pocock T,Huner NPA,Sarhan F(2002)Cold-Regulated cereal chloroplast late embryogenesis abundant-like proteins.Molecular characterization and functional analyses.Plant Physiology 129:1368-1381.
    Newman IA(2001)Ion transport in roots:measurement of fluxes using ion-selective microelectrodes to characterise transporter function.Plant,Cell and Environment 24:1-14.
    Noctor G,Foyer CH(1998)Ascorbate and glutathione:keeping active oxygen under control.Annual Review of Plant Physiology and Plant Molecular Biology 49:249-279.
    Nordin K,Vahala T Palva ET(1993)Differential expression of two related, low-temperature-induced gene in Arabidopsis thaliana(L.)Heynh.Plant Molecular Biology 21:641-653.
    Okuda T,Matsuda Y,Yamanaka A,Sagisaka S(1991)Abrupt increase in the level of hydrogen-peroxide in leaves of winter wheat is caused by cold treatment.Plant Physiology 97:1265-1267.
    Olien CR(1964)Freezing processes in the crown in“Hudson”barley.Crop Science 4:91-95.
    Oquist G,Hurry VM,Huner NPA(1993)Low-temperature effects on photosynthesis and correlation with freezing tolerance in spring and winter cultivars of wheat and rye.Plant Physiology 101:245-250.
    Pang CH,Zhang SJ,Gong ZZ,Wang BS(2005)NaCl treatment markedly enhances H_2O_2-scavenging system in leaves of halophyte Suaeda salsa.Physiologia Plantarum 125:490-499.
    Pang J,Cuin TA,Shabala L,Zhou M,Mendham N,Shabala S(2007)Effect of secondary metabolites associated with anaerobic soil conditions on ion fluxes and electrophysiology in barley roots.Plant Physiology 145:266-276.
    Pang J,Newman I,Mendham N,Zhou M,Shabala S(2006)Microelectrode ion and O_2 flux measurements reveal differential sensitivity of barley root tissues to hypoxia.Plant,Cell and Environment 29:1107-1121.
    Pihakaski-Maunsbach K,Tamminen I,Pietiainen M,Griffith M,(2003)Antifreeze proteins are secreted by winter rye cells in suspension culture.Physiologia Plantarum 118:390-398.
    Powles SB(1984)Photoinhibition of photosynthesis induced by visible light.Annual Review of Plant Biology 35:15-44.
    Reinheimer JL,Barr AR,Eglinton JK(2004)QTL mapping of chromosomal regions conferring reproductive frost tolerance in barley(Hordeum vulgare L.).Theoretical and Applied Genetics 109:1267-1274.
    Rizza F,Pagani D,Stanca AM,Cattivelli L(2001)Use of chlorophyll fluorescence to evaluate the cold acclimation and freezing tolerance of winter and spring oats.Plant Breeding 120:389-396.
    Saijo Y,Hata S,Kyozuka J,Shimamoto K,Izui K(2000)Over expression of a single Ca-dependent protein kinase confers both cold and salt/drought tolerance on rice plants.The Plant Journal 23:319-327.
    Schreiber U(1983)Chlorophyll fluorescence yield changes as a tool in plant physiology I.The measuring system.Photosynthesis Research 4:361-371.
    Schreiber U,Bilger W(1993)Progress in chlorophyll fluorescence research:major developments during the past years in retrospect.Progress in Botany 54:151-173.
    Shabala S(1996)Leaf temperature kinetics measure plant adaptation to extreme high temperatures.Australia Journal of Plant Physiology 23:445-452.
    Shabala S(2006)Non-invasive microelectrode ion flux measurements in plant stress physiology.In:Volkov A(ed.),Plant Electrophysiology-Theory and Methods.Springer-Verlag,Berlin,Germany,pp.35-71.
    Shabala L,Cuin TA,Newman I,Shabala S(2005a)Salinity-induced ion flux patterns from the excised roots of Arabidopsis sos mutants.Planta 222:1041-1050.
    Shabala S,Newman IA(1997)H+ flux kinetics around plant roots after short-term exposure to low temperature:indentifying critical temperatures for plant chilling tolerance.Plant,Cell and Environment 20:1401-1410.
    Shabala S,Newman IA(1998)Osmotic Sensitivity of Ca2+ and H+ Transporters in Corn Roots:Effect on Fluxes and Their Oscillations in the Elongation Region.Journal of Membrane Biology 161:45-54.
    Shabala S,Shabala L,Martynenko AI,Babourina O,Newman IA(1998)Salinity effect on bioelectric activity,growth,Na~+ accumulation and chlorophyll fluorescence of maize leaves:a comparative survey and prospects for screening.Australia Journal of Plant Physiology 25:609-616.
    Shabala S,Shabala L,Volkenburgh E(2003)Effect of calcium on root development and root ion fluxes in salinised barley seedlings.Functional Plant Biology 30:507-514.
    Shabala S,Shabala L,Volkenburgh E,Newman I(2005b)Effect of divalent cations on ion fluxes and leaf photochemistry in salinised barley leaves.Journal of Experimental Botany 56:13 69-1378.
    Shinozaki K,Yamaguchi-Shinozaki K(1996)Molecular responses to drought and cold stress.Current Opinion in Biotechnology 7:161-167.
    Single WV(1988)Resistance to frost injury during stem elongation and early heading.In:Frost injury in wheat:a workshop of the Wheat Research Council.Haymarket,N.S.W.
    Smallwood M,Worralt D,Byass L,Elias L,Ashford D,Doucet CJ,Holt C,Telford J,Lillford P,Bowles DJ(1999)Isolation and characterization of a novel antifreeze protein fromcarrot(Daucus carota).Biophysical Journal 340:385-391.
    Snape JW,Sarma R,Quarrie SA,Fish L,Galiba G,Sutka J(2001)Mapping genes for flowering time and frost tolerance in cereals using precise genetic stocks.Euphytica 120:309-315.
    Sowinski P,Rudzinska-Langwald A,Adamczyk J,Kubica I,Fronk J(2005)Recovery of maize seedling growth,development and photosynthetic efficiency after initial growth at low temperature.Journal of Plant Physiology 162:67-80.
    Sundbom E,Strand M,Hallgren JE(1982)Temperature-induced fluorescence changes:A screening method for frost tolerance of potato(Solanum sp.).Plant Physiology 70:1299-1302.
    Sutka J(2001)Genes for frost resistance in wheat.Euphytica 119:167-172.
    Suzuki I,Los AD,Kanesaki Y,Mikami K,Murata N(2000)The pathway for perception and transduction of low-temperature signals in Synechocystis.EMBO Journal 19:1327-1334.
    Tahtiharju S,Sangwan V,Monroy AF,Dhindsa RS,Borg M(1997)The induction of kin genes in cold-acclimating Arabidopsis thaliana.Evidence of a role for calcium.Planta 203:442-447.
    Terashima I,Huang LR,Osmond CB(1989)Effects of leaf chilling on thylakoid functions measured at room temperature in Cucumis sativus L.and Oryza sativa L.Plant Cell Physiology 30:841-850.
    Thomashow MF(2001)So what's new in the field of plant cold acclimation? Plant Physiology 125:89-93.
    Thomashow MF(1999)Plant cold tolerance:freezing tolerance genes and regulatory mechanisms.Annual Review of Plant Physiology and Plant Molecular Biology 50:571-599.
    Thomashow MF(1998)Role of cold-responsive genes in plant freezing tolerance.Plant Physiology 118:1-8.
    Toth B,Galiba G,Feher E,Sutka J,Snape JW(2003)Mapping genes affecting flowering time and frost resistance on chromosome 5B of wheat.Theoretical and Applied Genetics 107:509-514.
    Wanner LA,Junttila O(1999)Cold-induced freezing tolerance in Arabidopsis.Plant Physiology 120:391-400.
    Wherrett T(2006)Aluminium toxicity,tolerance,and amelioration in wheat.Thesis University of Tasmania.
    Wherrett TC,Ryan PB,Delhaize E,Shabala S(2005) Effect of aluminium on membrane potential and ion fluxes at the apices of wheat roots.Functional Plant Biology 32:199-208.
    Xin Z,Browse J(2000) Cold comfort farm:the acclimation of plants to freezing temperatures.Plant Cell and Environment 23:893-902.
    Xin Z,Browse J(1998) Eskimol mutants of Arabidopsis are constitutively freezing tolerant.Proceedings of the National Academy of Sciences of the USA 95:7799-7804
    Yamaguchi-Shinozaki K Koizumi M,Urao S,Shinozaki K(1994) Molecular cloning and characterization of 9 cDNAs for genes that are responsive to desiccation in Arabidopsis thaliana:sequence analysis of one cDNA clone that encodes a putative gene transmembrane changel protein.Plant and Cell Physiology 33:217-224.
    Yang WJ,Nadolska-Orczyk A,Wood KV,Hahn DT,Rich PJ,Wood AJ,Saneoka H,Premachandra GS,Bonham CC,Rhodes JC(1995) Near-isogenic lines of maize differing for glycinbetaine.Plant Physiology 107:621-630.
    Yu,XM,Griffith M,Wiseman SB(2001) Ethylene induces antifreeze activity in winter rye leaves.Plant Physiology 126:1232-1240.
    Zhu JK(2002) Salt and drought stress signal transduction in plants.Annual Review of Plant Biology 53:247-273.
    赵军,王以柔(1997)低温锻炼对水稻叶片中Ruhisco的影响.植物生理学报 23:1-6.
    Alscher RG,Donaue JL,Cramer CL(1997)Reactive oxygen species and antioxidants:relationships in green cells.Physiologia Plantarum 100:224-233.
    Apel K,Hirt H(2004)Reactive oxygen species:metabolism,oxidative stress,and signal transduction.Annual Review of Plant Biology 55:373-399.
    Assad MT,Paulsen GM(2002)Genetic changes in resistance to environmental stresses by U.S.Great Plains wheat cultivars.Euphytica 128:87-96.
    Baek KH,Skinner DZ(2003)Alteration of antioxidant enzyme gene expression during cold acclimation of near-isogenic wheat lines.Plant Science 165:1221-1227.
    Belkhodja R,Morales F,Abadia A,Gomez-Aparisi J,Abadia J(1994)Chlorophyll fluorescence as a possible tool for salinity tolerance screening in barley(Hordeum vulgare L.).Plant Physiology 104:667-673.
    Bertin P,Bouharmont J,Kinet JM(1997)Somaclonal variation and improvement of chilling tolerance in rice:Changes in chilling-induced chlorophyll fluorescence.Crop Science 37:1727-1735.
    Bosco CD,Busconi M,Govoni C,Baldi P,Stanca AM,Crosatti C,Bassi R,Cattivelli L(2003)cor gene expression in barley mutants affected in chloroplast development and photosynthetic electron transport.Plant Physiology 131:793-802.
    Bradford MM(1976)A rapid and sensitive method for the quantitation of microgram quantities of protein utilizing the principle of protein-dye binding.Analytical Biochemistry 72:248-254.
    Cakmak I,Marschner H(1992)Magnesium deficiency and high light intensity enhance activities of superoxide dismutase ascorbate peroxidase,and glutathione reductase in bean leaves.Plant Physiology 98:1222-1227.
    Dau H(1994)Short-term adaptation of plants to changing light intensties and its relation to photosystem Ⅱ photochemistry and fluorescence emission.Journal of Photochemistry and Photobiology B:Biology 26:3-27.
    Foyer CH,Descourvieres P,Kunert KJ(1994a)Protection against oxygen radicals:an important defense mechanism studied in transgenic plants.Plant Cell and Environment 17:507-523.
    Foyer CH,Lelandais M,Kunert KJ(1994b)Special review:Photooxidative stress in plants.Physiologia Plantarum 92:696-717.
    Gilmore AM(1997)Mechanistic aspects of xanthophyll cycle-dependent photoprotection in higher plant chloroplasts and leaves.Physiologia Plantarum 99:197-209.
    Gilmore AM,Hazlett TL,Govindjee(1995)Xanthophyll cycle-dependent quenching of photosystem Ⅱ chlorophyll a fluorescence:formation of a quenching complex with a short fluorescence lifetime.Proceedings of the National Academy of Sciences of the USA 92:2273-2277.
    Giorni E,Crosatti C,Baldi P,Grossi M,Mare C,Stanca AM,Cattivelli L(1999)Cold-regulated gene expression during winter in frost tolerant and frost susceptible barley cultivars grown under field conditions.Euphytica 106:149-157.
    Gray GR,Chauvin LP,Sarhan F,Huner NPA(1997)Cold acclimation and freezing tolerance:a complex interaction of light and temperature.Plant Physiology 114:467-474.
    Gray GR,Hope BJ,Qin XQ,Taylor BG,Whitehead CL(2003)The characterization of photoinhibition and recovery during cold acclimation in Arabidopsis thaliana using chlorophyll fluorescence imaging.Physiologia Plantarum 119:365-375.
    Gray GR,Ivanov AG,Krol M,Huner NPA(1998)Adjustment of thylakoid plastoquinone content and photosystem I electron donor pool size in response to growth temperature and growth irradiance in winter rye(Secale cereale L.).Photosynthesis Research 56:209-221.
    Gray GR,Savitch LV,Ivanov AG,Huner NPA(1996)Photosystem Ⅱ excitation pressure and development of resistance to photoinhibition:Ⅱ.Adjustment of photosynthetic capacity in winter wheat and winter rye.Plant Physiology 110:61-71.
    Griffith OW(1980)Determination of glutathione and glutathione disulfide using glutathione
    Guy CL(1990)Cold acclimation and freezing stress tolerance:Role of protein metabolism.Annual Review of Plant Physiology and Plant Molecular Biology 41:187-223.
    Hughes MA,Dunn MA(1996)The molecular biology of plant acclimation to low temperature.Journal of Experimental Botany 47:291-305.
    Huner NPA,(O|¨)quist G,Sarhan F(1998)Energy balance and acclimation to light and cold.Trends in Plant Science 3:224-230.
    Janda T,Szalai G,Rios-Gonzalez K,Veisz O,P(?)ldi E(2003)Comparative study of frost tolerance and antioxidant activity in cereals.Plant Science 164:301-306.
    Kendall EJ,McKersie BD(1989)Free radical and freezing injury to cell membranes of winter wheat.Physiologia Plantarum 76:86-94.
    Kocsya G,Galiba G,Brunold C(2001)Role of glutathione in adaptation and signaling during chilling and cold acclimation in plants.Physiologia Plantarum 113:158-164.
    Kuk YI,Shin JS,Burgos NR,Hwang TE,Han O,Cho BH,Jung S,Guh JO(2003)Antioxidative enzymes offer protection from chilling damage in rice plants.Crop Science 43:2109-2117.
    Law MY,Charles SA,Halliwell B(1983)Glutathione and ascorbic acid in spinach(Spinacia oleracea)chloroplasts.The effect of hydrogen peroxide and paraquat.BiochemicalJournal210:899-903.
    Lichtenthaler HK,Buschmann C,Knapp M(2005a)How to correctly determine the different chlorophyll fluorescence parameters and the chlorophyll fluorescence decrease ratio R_(Fd)of leaves with the PAM fluorometer.Photosynthetica 43:379-393.
    Lichtenthaler HK,Langsdorf G,Lenk,S,Buschmann C(2005b)Chlorophyll fluorescence imaging of photosynthetic activity with the flash-lamp fluorescence imaging system.Photosynthetica 43:355-369.
    Mahajan S,Tuteja N,(2005)Cold,salinity and drought stresses:An overview.Archives of Biochemistry and Biophysics AAA:139-158.
    Nakano Y,Asada K.(1981)Hydrogen peroxide is scavenged by ascorbate specific peroxidase in spinach chloroplasts.Plant and Cell Physiology 22:679-690.
    Noctor G,Foyer CH(1998)Ascorbate and glutathione:keeping active oxygen under control.Annual Review of Plant Physiology and Plant Molecular Biology 49:249-279.
    O'Kane D,Gill V,Boyd P,Burdon R(1996)Chilling,oxidative stress and antioxidant responses in Arabidopsis thaliana callus.Planta 198:371-377.
    Okuda T,Matsuda Y,Yamanaka A,Sagisaka S(1991)Abrupt increase in the level of hydrogen-peroxide in leaves of winter wheat is caused by cold treatment.Plant Physiology 97:1265-1267.
    (O|¨)quist G,Huner NPA(1993)Cold hardening induced resistance to photoinhibition in winter rye is dependent upon increase capacity for photosynthesis.Planta 189:150-156.
    (O|¨)quist G,Hurry VM,Huner NPA(1993)Low-temperature effects on photosynthesis and correlation with freezing tolerance in spring and winter cultivars of wheat and rye.Plant Physiology 101:245-250.
    Pang CH,Zhang SJ,Gong ZZ,Wang BS(2005)NaCl treatment markedly enhances H_2O_2-scavenging system in leaves of halophyte Suaeda salsa.Physiologia Plantarum 125:490-499
    Patterson BD,Macrae EA,Ferguson IB(1984)Estimation of hydrogen peroxide in plant extracts using Titanium(IV).Analytical Biochemistry 139:487-492.
    Prasad TK,Anderson MD,Martin BA,Stewart CR(1994)Evidence for chilling-induced oxidative stress in maize seedlings and a regulatory role for hydrogen peroxide.Plant Cell 6:65-74.
    Pukacki PM,Kendall EJ,McKersie(1991)Membrane injury during freezing stress to winter wheat(Triticum aestivum L.)crowns.Journal of Plant Physiology 1385:16-521.
    Racz I,Kov(?)cs M,L(?)sztity D,Veisz O,Szalai G,P(?)ldi E(1996)Effect of short-term and long-term low temperature stress on polyamine biosynthesis in wheat genotypes with varying degrees of frost tolerance.Journal of Plant Physiology 148:368-373.
    Rao MV,Paliyath G,Ormarod DP(1996)Ultraviolet-B-and ozone-induced biochemical changes in antioxidant enzymes of Arabidopsis thaliana.Plant Physiology 110:125-136.
    Rizza F,Crosatti C,Stanca AM,Cattivelli L(1994)Studies for assessing the influence of hardening on cold tolerance of barley genotypes.Euphytica 75:131-138.
    Rizza F,Pagani D,Stanca AM,Cattivelli L(2001)Use of chlorophyll fluorescence to evaluate the cold acclimation and freezing tolerance of winter and spring oats.Plant Breeding 120:389-396.
    Tao DL,(O|¨)quist G.,Wingsle G(1998)Active oxygen scavengers during cold acclimation of Scots pine seedlings in relation to freezing tolerance.Cryobiology 37:38-45.
    Scebba F,Sebustiani L,Vitagliano C(1998)Changes in activity of antioxidant enzymes in wheat(Triticum aestivum L.)seedlings under cold acclimation.Physiologia Plantarum 104:747-752.
    Scebba F,Sebustiani L,Vitagliano C(1999)Protective enzymes against activated oxygen species in wheat(Triticum aestivum L.)seedlings:Responses to cold acclimation.Journal of Plant Physiology 155:762-768.
    Somersalo S,Krause GH(1990)Photoinhibition at chilling temperatures and effects of freezing stress on cold acclimated spinach leaves in the field.A fluorescence study.Physiologia Plantarum 79:617-622.
    Sowinski P,Rudzihska-Langwald A,Adamczyk J,Kubica I,Fronk J(2005)Recovery of maize seedling growth,development and photosynthetic efficiency after initial growth at low temperature.Journal of Plant Physiology 162:67-80.
    Strand A,Hurry V,Gustafsson P,Gardestrom P(1997)Development of Arabidopsis thaliana leaves at low temperature releases the suppression of photosynthesis and photosynthetic gene expression despite the accumulation of soluble carbohydrates.Plant Journal 12:605-614.
    Strand A,Hurry V,Henkes S,Huner N,Gustafsson P,Gardestrom P,Stitt M(1999)Acclimation of Arabidopsis leaves developing at low temperatures:Increasing cytoplasmic volume accompanies increased activities of enzymes in the Calvin cycle and in the sucrose-biosynthesis pathway.Plant Physiology 119:1387-1398.
    Sundbom E,Strand M,Hallgren JE(1982)Temperature-induced fluorescence changes:A screening method for frost tolerance of potato(Solanum sp.).Plant Physiology 70:1299-1302.
    Suzuki N,Mittler R(2006)Reactive oxygen species and temperature stresses:A delicate balance between signaling and destruction.Physiologia Plantarum 126:45-51.
    Thomashow MF(1999)Plant cold acclimation:freezing tolerance genes and regulatory mechanisms.Annual Review of Plant Physiology and Plant Molecular Biology 50:571-599.
    Uemura M,Steponkus PL(1994)A contrast of the plasma membrane lipid composition of oat and rye leaves in relation to freezing tolerance.Plant Physiology 104:479-496.
    Willekens H,Inze D,Van Montagu M,Camp WV(1995)Catalase in plants.Molecular Breeding 1:207-228.
    Arif I,Newman IA,Keenlyside N(1995) Proton flux measurements from tissues in buffered solution.Plant,Cell and Environment 18:1319-1324.
    Chen Z,Newman I,Zhou M,Mendham N,Zhang G,Shabala S(2005) Screening plants for salt tolerance by measuring K~+ flux:a case study for barley.Plant,Cell and Environment 28:1230-1246.
    Clarkson DT,Hanson JB(1980)The mineral nutrition of higher plants.Annual Review of Plant Physiology 31:239-298.
    Kochian LV,Lucas WJ(1988)Potassium transport in roots.Advances in Botanical Research 15:93-178.
    Lyons JM(1973)Chilling injury in plants.Annual Review of Plant Physiology 24:445-466.
    Maathuis FJM,Ichida AM,Sanders D,Schroeder JI(1997)Roles of higher plant K~+ channels.Plant Physiology 114:1141-1149.
    Maser P,Gierth M,Schroeder J(2002b)Molecular mechanisms of potassium and sodium uptake in plants.Plant Soil 247:43-54.
    Newman IA(2001)Ion transport in roots:measurement of fluxes using ion-selective microelectrodes to characterise transporter function.Plant,Cell and Environment 24:1-14.
    Newman IA,Kochian LV,Grusak MA,Lucas WJ(1987)Fluxes of H~+ and K~+ in com roots:characterisation and stoichiometrics using ionselective microelectrodes.Plant Physiology 84:1177-1184.
    Palmgren MG(2001)Plant plasma membrane H~+-ATPases:powerhouses for nutrient uptake.Annual Review of Plant Physiology and Plant Molecular Biology 52:817-845.
    Pineros MA,Shaff JE,Kochian LV(1998)Development,characterisation,and application of a cadmium-selective microelectrode for the measurement of cadmium fluxes in roots of Thlaspi species and wheat.Plant Physiology 116:1393-1401.
    Reinheimer JL,Barr AR,Eglinton JK(2004)QTL mapping of chromosomal regions conferring reproductive frost tolerance in barley(Hordeum vulgare L.).Theoretical and Applied Genetics 109:1267-1274.
    Reinhold L,Seiden A,Volokita M(1984)Is modulation of the rate of proton pumping a key event in osmoregulation? Plant Physiology 75:846-849.
    Ryan PR,Newman I A,Shields B(1990)Ion fluxes in com roots measured by microelectrodes with ion-specific liquid membranes.Journal of Membrane Science 53:59-69.
    Shabala S(2006)Non-invasive microelectrode ion flux measurements in plant stress physiology.In:Volkov A(ed.),Plant Electrophysiology-Theory and Methods. Springer-Verlag,Berlin,Germany,pp.35-71.
    Shabala SN,Newman IA(1997)H+ flux kinetics around plant roots after short-term exposure to low temperature:indentifying critical temperatures for plant chilling tolerance.Plant,Cell and Environment 20:1401-1410.
    Shabala S,Newman IA,Morris J(1997)Oscillations in H~+ and Ca~(2+)ion fluxes around the elongation region of corn roots and effects of external pH.Plant Physiology 113:111-118.
    Shabala S,Newman I,Whittington J,Juswono U(1998)Protoplast ion fluxes:their measurement and variation with time,position and osmoticum.Planta 204:146-152.
    Shabala S,Shabala L(2002)Kinetics of net H~+,Ca~(2+),K~+,Na~+ and Cl~-fluxes associated with post-chilling recovery of plasma membrane transporters in Zea mays leaf and root tissues.Physiologia Plantarum 114:47-56.
    Shabala S,Shabala L,Volkenburgh E(2003)Effect of calcium on root development and root ion fluxes in salinised barley seedlings.Functional Plant Biology 30:507-514.
    Stiles KA,Van Volkenburgh E(2004)Role of K~ + in leaf growth:K~+ uptake is required for light-stimulated H~+ efflux but not solute accumulation.Plant,Cell and Environment 27:315-325.
    Walker DJ,Black CR,Miller AJ(1998)The Role of Cytosolic Potassium and pH in the Growth of Barley Roots.Plant Physiology 118:957-964.
    Wyn JRG,Brady CJ,Speirs J(1979)Ionic and osmotic relations in plant cells.In:Laidman DL,Wyn Jones RG(eds.),Recent Advances in the Biochemistry of Cereals.Academic Press,London,UK.pp.63-103
    Yoshida S(1991)Chilling-induced inactivation and its recovery of tonoplast H-ATPase in mung bean cell suspension cultures.Plant Physiology 95:456-460.
    Yoshida S(1994)Low temperature-induced cytoplasmic acidosis in cultured mung bean(Vigna radiata[L.]Wilczek)cells.Plant Physiology 104:1131-1138.
    Zieschang HE,Kohler K,Sievers A(1993)Changing proton concentrations at the surfaces of gravistimulated Phleum roots.Planta 190:546-554.
    Galiba G.,Kerepesi I,Snape JW,Sutka J(1997) Location of a gene regulating cold-induced carbohydrate production on chromosome 5A of wheat.Theoretical and Applied Genetics 95:265-270.
    Galiba G,Kerepesi I,Vagujfalvi A,Kocsy G,Cattivelli L,Dubcovsky J,Snape SW,Sutka J(2001)Mapping of genes involved in glutathione,carbohydrate and COR14b cold induced protein accumulation during cold hardening in wheat.Euphytica 119:173-177.
    Gray GR,Hope BJ,Qin XQ,Taylor BG,Whitehead CL(2003)The characterization of photoinhibition and recovery during cold acclimation in Arabidopsis thaliana using chlorophyll fluorescence imaging.Physiologia Plantarum 119:365-375.
    Marcellos M(1988)Comments on frost injury in wheat.In:Frost injury in wheat:a workshop of the wheat research council.Haymarket,N.S.W.
    (O|¨)gren E(1991)Prediction of photoinhibition of photosynthesis from measurements of fluorescence quenching components.Planta 184:538-544.
    (O|¨)quist G,Hurry VM,and Huner NPA(1993)Low-Temperature Effects on Photosynthesis and Correlation with Freezing Tolerance in Spring and Winter Cultivars of Wheat and Rye.Plant Physiology 101:245-250.
    Reinheimer JL,Barr AR,Eglinton JK(2004)QTL mapping of chromosomal regions conferring reproductive frost tolerance in barley(Hordeum vulgare L.).Theoretical and Applied Genetics 109:1267-1274.
    Schreiber U,Bilger W,Neubauer C(1994)Chlorophyll fluorescence as a nonintrusive indicator for rapid assessment of in vivo photosynthesis.In:Schulze E-D,Caldwell MM(eds)Ecophysiology of Photosynthesis.Spinger-Verlag,Berlin,pp.49-70.
    Single WV(1988)Resistance to frost injury during stem elongation and early heading.In:Frost injury in wheat:a workshop of the Wheat Research Council.Haymarket,N.S.W.
    Stitt,M(1986)Limitation of photosynthesis by carbon metabolism:I.Evidence for excess electron transport capacity in leaves carrying out photosynthesis in saturating light and C02.Plant Physiology 81:1115-1122.
    Tabaei-Aghdaei SR,Pearcel RS,Harrison P(2003)Sugars regulate cold-induced gene expression and freezing-tolerance in barley cell cultures.Journal of Experimental Botany 387:1565-1575.
    Bertin P,Bouharmont J,Kinet JM(1997) Somaclonal variation and improvement of chilling tolerance in rice:Changes in chilling-induced chlorophyll fluorescence.Crop Science 37:1727-1735.
    Butler WL(1978) Energy distribution in the photochemical apparatus of photosynthesis.Annual Review of Plant Physiology 29:345-378.
    Corke H,Atsmon D(1988)Effect of nitrogen nutrition on endosperm protein synthesis in wild and cultivated barley grown in spike culture.Plant Physiology 87:523-528.
    Galiba G,Kerepesi I,Snape JW,Sutka J(1997)Location of a gene regulating cold-induced carbohydrate production on chromosome 5A of wheat.Theoretical and Applied Genetics 95:265-270.
    Galiba G,Kerepesi I,Vagujfalvi A,Kocsy G,Cattivelli L,Dubcovsky J,Snape SW,Sutka J(2001)Mapping of genes involved in glutathione,carbohydrate and COR 14b cold induced protein accumulation during cold hardening in wheat.Euphytica 119:173-177.
    Galiba G,Quarrie SA,Tuberosa R,Veisz O,Sutka J(1995)Regulation of freezing tolerance by abscisic acid in wheat.Acta Phytopathologica et Entomologica Hungarica 30:123-124.
    Gray GR,Hope BJ,Qin XQ,Taylor BG,Whitehead CL(2003)The characterization of photoinhibition and recovery during cold acclimation in Arabidopsis thaliana using chlorophyll fluorescence imaging.Physiologia Plantarum 119:365-375.
    Gray GR,Chauvin LP,Sarhan F,Huner NPA(1997)Cold acclimation and freezing tolerance:a complex interaction of light and temperature.Plant Physiology 114:467-474.
    Havaux M,Kloppstech K(2001)The protective functions of carotenoid and flavonoid pigments against excess visible radiation at chilling temperature investigated in Arabidopsis npq and tt mutants.Planta 213:953-966.
    Hayes PM,Blake T,Chen TTH,Tragoonrung S,Chen F,Pan A,Liu B(1993)Quantitative trait loci on barley(Hordeum vulgare L.)chromosome 7 associated components of winter hardiness.Genome 36:66-71.
    Janda T,Szalai G,Rios-Gonzalez K,Veisz O,Paldi E(2003)Comparative study of frost tolerance and antioxidant activity in cereals.Plant Science 164:301-306.
    Krause GH,Weis E(1991)Chlorophyll fluorescence and photosynthesis:the basics.Annual Review of Plant Physiology and Plant Molecular Biology 42:313-349.
    Li HB,Vaillancourt R,Mendham NJ,Zhou MX(2008)Comparative mapping of quantitative trait loci associated with waterlogging tolerance in barley(Hordeum vulgare L.).BMC Genomics 9:401. Lichtenthaler HK,Buschmann C,Knapp M(2005)How to correctly determine the different chlorophyll fluorescence parameters and the chlorophyll fluorescence decrease ratio R_(Fd)of leaves with the PAM fluorometer.Photosynthetica 43:379-393.
    (O|¨)gren E(1991)Prediction of photoinhibition of photosynthesis from measurements of fluorescence quenching components.Planta 184:538-544.
    Qi JC,Zhang GP,Zhou MX(2006)Protein and hordein content in barley seeds as affected by nitrogen level and their relationship to beta-amylase activity.Journal of Cereal Science A3:102-107.
    Ralph PJ,Gademann R(2005)Rapid light curves:A powerful tool to assess photosynthetic activity.Aquatic Botany 82:222-237.
    Rascher U,Liebig M,Lüttge U(2000)Evaluation of instant light-response curves of chlorophyll fluorescence parameters obtained with a portable chlorophyll fluorometer on site in the field.Plant cell and environment 23:1397-1405.
    Reinheimer JL,Barr AR,Eglinton JK(2004)QTL mapping of chromosomal regions conferring reproductive frost tolerance in barley(Hordeum vulgare L.)Theoretical and Applied Genetics 109:1267-1274.
    Rizza F,Pagani D,Stanca AM,Cattivelli L(2001)Use of chlorophyll fluorescence to evaluate the cold acclimation and freezing tolerance of winter and spring oats.Plant Breeding 120:389-396.
    Schreiber U,Gademann R,Ralph PJ,Larkum AWD(1997)Assessment of photosynthetic performance of Prochloron in Lissoclinum patella in hospite by chlorophyll fluorescence measurements.Plant and Cell Physiology 38:945-951.
    Snape JW,Sarma R,Quarrie SA,Fish L,Galiba G,Sutka J(2001)Mapping genes for flowering time and frost tolerance in cereals using precise genetic stocks.Euphytica 120:309-315.
    Sowinski P,Rudzinska-Langwald A,Adamczyk J,Kubica I,Fronk J(2005)Recovery of maize seedling growth development and photosynthetic efficiency after initial growth at low temperature.Journal of Plant Physiology 162:67-80.
    Sundbom E,Strand M,Hallgren JE(1982)Temperature-induced fluorescence changes:A screening method for frost tolerance of potato(Solanum sp.).Plant Physiology 70:1299-1302.
    Sutka J(2001)Genes for frost resistance in wheat.Euphytica 119:167-172.
    Toth B,Galiba G,Feher E,Sutka J,Snape JW(2003)Mapping genes affecting flowering time and frost resistance on chromosome 5B of wheat.Theoretical and Applied Genetics 107:509-514.
    Walker MA,Mckersie BD,Pauls KP(1991)Effects of chilling on the biochemical and functional properties of thylakoid membranes.Plant Physiology 97:663-669.
    Wenzl P,Li HB,Carling J,Zhou MX,Raman H,Paul E,Hearnden P,Maier C,Xia L,Caig V,Ovesn(?)J,Cakir M,Poulsen D,Wang JP,Raman R,Smith K,Muehlbauer G,Chalmers K,Kleinhofs A,Huttner E,Kilian A(2006)A high-density consensus map of barley linking DArT markers to SSR,RFLP and STS loci and agricultural traits.BMC Genomics 7:206-228.
    White A J,Critchley C(1999)Rapid light curves:A new fluorescence method to assess the state of the photo synthetic apparatus.Photosynthesis Research 59:63-72.

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