敖汉和维多利亚紫花苜蓿对低磷环境应激机制的比较
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摘要
为了探索紫花苜蓿在低磷环境下的表现及响应机制,以期为提高苜蓿磷利用效率及磷吸收效率提供参考。试验以耐低磷较强的敖汉苜蓿和耐低磷较弱的维多利亚苜蓿为研究对象,以正常磷(500μmol·L~(-1) KH2PO4)处理为对照,低磷(5μmol·L~(-1) KH2PO4)胁迫处理10、15、20d后,分别观测其形态特征及生理指标的变化,比较两个品种在低磷胁迫下的差异,综合分析其耐低磷机制。结果表明,低磷胁迫下,敖汉和维多利亚的株高、地上生物量及磷含量有所下降,地下生物量、根冠比、总根长、根表面积及磷利用率有所提高,二者净光合速率、气孔导度和蒸腾速率显著下降(P<0.05),但是敖汉的下降幅度都远小于维多利亚;二者的酸性磷酸酶活性都极显著增强(P<0.01),但敖汉的酸性磷酸酶活性极显著高于维多利亚(P<0.01)。综合表明,低磷胁迫抑制了苜蓿的地上部分生长,促进了地下部分生长,敖汉较维多利亚耐低磷主要表现在光合作用受低磷影响较小、磷酸酶活性增强幅度较大。通过两个品种在低磷胁迫下的不同表现,为紫花苜蓿的耐低磷机制研究提供一定的理论依据。
In order to compare the performance and adaptive mechanisms of two varieties of alfalfa under low inorganic phosphate(low-Pi)conditions,and to provide more information for improvement of Pi utilization efficiency and absorption efficiency in alfalfa,the varieties Aohan and Victoria were grown under low-Pi(5μmol·L~(-1) KH2PO4)and normal Pi(500μmol·L~(-1) KH2PO4)conditions.Morphological and physiological traits were measured at 10,15 and 20days.It was found that,the height,aboveground biomass and Pi content of both Aohan and Victoria were decreased under low-Pi stress;while the underground biomass,root-shoot ratio,total root length,root surface area and Pi utilization were increased.The net photosynthetic rate,stomatal conductance and transpiration rate were decreased significantly under low-Pi(P<0.05),but the decrease was much smaller in cultivar Aohan than in cultivar Victoria.The acid phosphatase activity of both was significantly enhanced(P<0.01),and for Aohan was significantly higher than that of Victoria(P<0.01).To sum up,lowPi stress inhibited the growth of the above ground plant organs and promoted growth of the underground parts.The greater low-Pi tolerance of Aohan compared to Victoria is associated with greater increase in acid phosphatase activity under low-Pi in Aohan,and a lower reduction in photosynthesis under low-Pi.The contrasting responses of the two varieties under low-Pi stress indicate a potential mechanism for alfalfa tolerance to low-Pi.
In order to compare the performance and adaptive mechanisms of two varieties of alfalfa under low inorganic phosphate(low-Pi)conditions,and to provide more information for improvement of Pi utilization efficiency and absorption efficiency in alfalfa,the varieties Aohan and Victoria were grown under low-Pi(5μmol·L~(-1) KH2PO4)and normal Pi(500μmol·L~(-1) KH2PO4)conditions.Morphological and physiological traits were measured at 10,15 and 20days.It was found that,the height,aboveground biomass and Pi content of both Aohan and Victoria were decreased under low-Pi stress;while the underground biomass,root-shoot ratio,total root length,root surface area and Pi utilization were increased.The net photosynthetic rate,stomatal conductance and transpiration rate were decreased significantly under low-Pi(P<0.05),but the decrease was much smaller in cultivar Aohan than in cultivar Victoria.The acid phosphatase activity of both was significantly enhanced(P<0.01),and for Aohan was significantly higher than that of Victoria(P<0.01).To sum up,lowPi stress inhibited the growth of the above ground plant organs and promoted growth of the underground parts.The greater low-Pi tolerance of Aohan compared to Victoria is associated with greater increase in acid phosphatase activity under low-Pi in Aohan,and a lower reduction in photosynthesis under low-Pi.The contrasting responses of the two varieties under low-Pi stress indicate a potential mechanism for alfalfa tolerance to low-Pi.
引文
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[5] Ryan P,Delhaize E,Jones D.Function and mechanism of organic anion exudation from plant roots.Annual Review of Plant Physiology and Plant Molecular Biology,2001,52:527-560.
[6] Kou C L,Wang Q J,Ren L X,et al.Study on morphological differences of phosphorus utilization in wheat and peanut.Chinese Journal of Soil Science,1999,30(4):181-184.寇长林,王秋杰,任丽轩,等.小麦和花生利用磷形态差异的研究.土壤通报,1999,30(4):181-184.
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[8] Yan X L,Liao H,Ge Z Y,et al.Root architectural characteristics and phosphorus acquisition efficiency in plants.Chinese Bulletin of Botany,2000,17:511-519.严小龙,廖红,戈振扬,等.植物根构型特性与磷吸收效率.植物学通报,2000,17:511-519.
[9] Neumann G,Martinoia E.Cluster roots-an under-ground adaptation for survival in extreme environments.Trends in Plant Science,2002,7:162-167.
[10] Tang H L,Shen J B,Zhang F S,et al.Interactive effects of phosphorus deficiency and exogenous auxin on root morphological traits in white lupin(Lupinus albus L.).Science China Life Sciences,2013,43(3):201-212.唐宏亮,申建波,张福锁,等.磷和外源生长素对白羽扇豆(Lupinus albus L.)根形态和生理特性的影响.中国科学生命科学,2013,43(3):201-212.
[11] Wang Y,Sun J,Wang R P,et al.Correlation of biological characteristics in relation to P efficiency in maize genotypes in jointing stage.Journal of Shanxi Agricultural University,2003,23(1):28-31.王艳,孙杰,王荣萍,等.玉米自交系苗期生物学性状与磷效率的相关性.山西农业大学学报,2003,23(1):28-31.
[12] Jacob J,Lawlor D W.Stomatal and mesophyll limitations of photosynthesis in deficient sunflower,maize and wheat plants.Journal of Experimental Botany,1991,42(8):1003-1011.
[13] Zhang K W,Wang X L,Li K M,et al.Effect of low-level phosphorus stress on photosynthetic characteristics of maize inbred line seedling with low-level phosphorus tolerance.Journal of Shandong University,2007,(3):89-94.张可炜,王贤丽,李坤明,等.低磷胁迫对耐低磷玉米自交系幼苗光合特性的影响.山东大学学报,2007,(3):89-94.
[14] Pieters A J,Paul M J,Lawlor D W.Low sink demand limits photosynthesis under Pi deficiency.Journal of Experimental Botany,2001,52(358):1083-1091.
[15] Cao L M,Pan X H.Analysis of some indexes used for evaluating tolerance of different rice genotypes to low phosphorus treatment in sand culture.Acta Agriculturae Shanghai,2000,4:31-34.曹黎明,潘晓华.水稻不同耐低磷基因型的评价指标分析.上海农业学报,2000,4:31-34.
[16] Richardson A E.Regulating the phosphorus nutrition of plants:Molecular biology meeting agronomic needs.Plant and Soil,2009,322:17-24.
[17] Chen L S,Chen Y Z,Wang R,et al.Effects of P-deficiency on protective enzyme activities of different fine clones of Camellia oleifera Abel.Journal of Northeast Forestry University,2013,41(9):23-25.陈隆升,陈永忠,王瑞,等.低磷胁迫对不同油茶优良无性系酶活性的影响.东北林业大学学报,2013,41(9):23-25.
[18] Vance C P,Uh der-Stone C,Allan D L.Phosphorus acquisition and use:Critical adaptations by plants for securing a nonrenewable resource.New Phytologist,2003,157:423-447.
[19] Wang S Q,Han X Z,Qiao Y F,et al.Characteristics of organic acids exudated from soybean(Glycine max L.)roots under P deficiency stress.Soybean Science,2009,28:409-414.王树起,韩晓增,乔云发,等.缺磷胁迫条件下大豆根系有机酸的分泌特性.大豆科学,2009,28:409-414.
[20] Chen K,Ma J,Cao Y P,et al.Exudation of organic acids by the roots of different plant species under phosphorus deficiency.Journal of China Agricultural University,1999,4(3):58-62.陈凯,马敬,曹一平,等.磷亏缺下不同植物根系有机酸的分泌.中国农业大学学报,1999,4(3):58-62.
[21] Zhou L L.Proton and organic acids exudated by faba bean,soybean and maize and their significance in interspecific facilitation on phosphorus uptake by intercropping.Beijing:China Agricultural University,2005.周丽莉.蚕豆、大豆、玉米根系质子和有机酸分泌差异及其在间作磷营养中的意义.北京:中国农业大学,2005.
[22] LüY,Cheng W D,Huang K,et al.Comparison of rhizosphere processes of Vicia sativa and Vicia villosa in response to phosphorus deficiency.Plant Nutrition and Fertilizer Science,2011,17(3):674-679.吕阳,程文达,黄珂,等.低磷胁迫下箭筈豌豆和毛叶苕子根际过程的差异比较.植物营养与肥料学报,2011,17(3):674-679.
[23] Raghothama K G.Phosphate acquisition.Annual Review of Plant Physiology and Plant Molecular Biology,2004,50:665-693.
[24] Hernandez G,Ramirez M,Valdes-Lopez O,et al.Phosphorus stress in common bean:Root transcript and metabolic responses.Plant Physiological,2007,144(2):752-767.
[25] Zhao Y,Xie K Y,Wan J C,et al.Development and prospects of‘grain-forage supply’in modern animal husbandry.Pratacultural Science,2017,34(3):653-660.赵云,谢开云,万江春,等.粮草兼顾型畜牧业饲草料发展现状及展望.草业科学,2017,34(3):653-660.
[26] Berg W K,Cunningham S M,Brouder S M,et al.Influence of phosphorus and potassium on alfalfa yield and yield components.Crop Science,2005,45:297-304.
[27] He F,Han D M,Wan L Q,et al.The nutrient situation in the major alfalfa producing areas of China.Journal of Plant Nutrition and Fertilizer,2014,20(2):503-509.何峰,韩冬梅,万里强,等.我国主产区紫花苜蓿营养状况分析.植物营养与肥料学报,2014,20(2):503-509.
[28] Ren L F,Zhang W H,Li Y S.Effect of phosphorus deficiency on physiological properties of Medicago falcate.Acta Prataculturae Sinica,2012,21(3):242-249.任立飞,张文浩,李衍素.低磷胁迫对黄花苜蓿生理特性的影响.草业学报,2012,21(3):242-249.
[29] James D W,Hurst C J,Tindall T A.Alfalfa cultivar response to phosphorus and potassium deficiency:Elemental composition of the herbage.Journal of Plant Nutrition,1995,18(11):2447-2464.
[30] Ranjan S,Pandey R M,Srivastava R K,et al.Effect of phosphorus deficiency on the metabolic changes in free amino-acids in certain leguminous crop plants.Nature,1962,193(4819):997-998.
[31] Gleiter M E,Parker H E.The effect of phosphorus deficiency on the free amino acids of alfalfa.Archives of Biochemistry and Biophysics,1957,71(2):430-436.
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