小麦旗叶衰老过程不同数学模型拟合比较及衰老特征分析
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摘要
为了准确了解小麦叶片的衰老特征,筛选适合描述小麦叶片衰老过程的数学模型, 2011年和2012年分别以91个和105个小麦品种(系)为材料,用"S"型曲线中的Logistic、Gompertz和Richards模型拟合了试验品种叶片的衰老过程,解析了其叶片衰老特征。结果表明,试验品种可分为延绿、中等延绿、中等早衰和早衰4种类型。其旗叶衰老过程可分为衰老起始期、快速衰老期和衰老结束期3个阶段,3个阶段旗叶的衰老速度表现为"慢–快–慢",不同延绿类型品种开花后旗叶的绿色叶面积百分比下降主要在衰老过程的中后期。3种模型对不同延绿类型品种旗叶衰老过程均可以拟合,Gompertz和Richards模型拟合度接近,高于Logistic模型。Gompertz模型的拟合度以早衰>中等早衰>中等延绿>延绿类型。不同延绿类型品种旗叶衰老曲线特征参数达到最大衰老速度时间(TMRS)、平均衰老速度(ARS)和绿色叶面积持续期(GLAD)存在显著差异, TMRS和GLAD以延绿>中等延绿>中等早衰>早衰, ARS以早衰>中等早衰>中等延绿>延绿。Gompertz模型对小麦叶片衰老过程的拟合度优于Logistic模型。叶片衰老过程特征参数可以用于品种延绿性差异评价。
To clarity leaf senescence patterns and characteristics of wheat, we used 91 and 105 varieties or advanced lines in 2011 and 2012 respectively, to fit the leaf senescence process by Logistic, Gompertz, and Richards models. The varieties tested were classified into stay green(SG), moderately stay green(MSG), moderately non-stay green(MNSG) and non-stay green(NSG) types. Development of the flag leaf senescence was divided into initial senescence stage, rapid senescence stage and terminal senescence stage in all variety types. The senescence rates of the three stages showed a slow–fast–slow changing pattern. The decrease of green leaf area(% GLA) after anthesis occurred mainly in the middle and late senescence stages. Logistic, Gompertz and Richards models were able to simulate the leaf senescence process of all variety types and the fitness of Richards and Gompertz models was better than that of Logistic model. The fitting goodness of Gompertz model ranked as NSG type > MNSG type > MSG type > SG type. There was a significant difference in time of maximum rate of senescence, green leaf area duration and average rate of senescence derived from Gompertz equation in all varieties. The average rate of senescence showed a trend of NSG type > MNSG type > MSG type > SG type, while the remaining two parameters showed a trend of SG type > MSG type > MNSG type > NSG type. Our results indicate that Gompertz model is more suitable than Logistic model to describe the leaf senescence process of wheat and the characteristic parameters for leaf senescence process can be used to evaluate stay green difference between wheat varieties.
To clarity leaf senescence patterns and characteristics of wheat, we used 91 and 105 varieties or advanced lines in 2011 and 2012 respectively, to fit the leaf senescence process by Logistic, Gompertz, and Richards models. The varieties tested were classified into stay green(SG), moderately stay green(MSG), moderately non-stay green(MNSG) and non-stay green(NSG) types. Development of the flag leaf senescence was divided into initial senescence stage, rapid senescence stage and terminal senescence stage in all variety types. The senescence rates of the three stages showed a slow–fast–slow changing pattern. The decrease of green leaf area(% GLA) after anthesis occurred mainly in the middle and late senescence stages. Logistic, Gompertz and Richards models were able to simulate the leaf senescence process of all variety types and the fitness of Richards and Gompertz models was better than that of Logistic model. The fitting goodness of Gompertz model ranked as NSG type > MNSG type > MSG type > SG type. There was a significant difference in time of maximum rate of senescence, green leaf area duration and average rate of senescence derived from Gompertz equation in all varieties. The average rate of senescence showed a trend of NSG type > MNSG type > MSG type > SG type, while the remaining two parameters showed a trend of SG type > MSG type > MNSG type > NSG type. Our results indicate that Gompertz model is more suitable than Logistic model to describe the leaf senescence process of wheat and the characteristic parameters for leaf senescence process can be used to evaluate stay green difference between wheat varieties.
引文
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[2]李永攀,罗培高,任正隆.小麦持绿性与产量关系研究.西南农业学报,2008,21:1221-1225.Li Y P,Luo P G,Ren Z L.Study on the relation between the yield and trait of green-keeping wheat.Southwest China JAgric Sci,2008,21:1221-1225(in Chinese with English abstract).
[3]Christopher J T,Manschadi A M,Hammer G L,Borrell A K.Developmental and physiological traits associated with high yield and stay-green phenotype in wheat.Aust J Agric Res,2008,59:354-364.
[4]武永胜,薛晖,刘洋,龚月桦.持绿型小麦叶片衰老和叶绿素荧光特征的研究.干旱地区农业研究,2010,28(4):117-122.Wu Y S,Xue C,Liu H,Gong Y H.Study on senescence and chlorophyll fluorescence traits of stay-green leaf in wheat.Agric Res Arid Areas,2010,28(4):117-122(in Chinese with English abstract).
[5]Gaju O,Allard V,Martre P,Snape J W,Heumez E,Le Gouis J,Moreau D,Bogard M,Griffiths S,Orford S,Hubbart S,Foulkes M J.Identification of traits to improve the nitrogen-use efficiency of wheat genotypes.Field Crops Res,2011,23:139-152.
[6]Derkx A P,Orford S,Griffiths S,Foulkes M J,Hawkesford MJ.Identification of differentially senescing mutants of wheat and impacts on yield,biomass and nitrogen partitioning.JIntegr Plant Biol,2012,54:555-566.
[7]Verma V,Foulkes M J,Worland A J,Sylvester-Bradley R,Caligari P D S,Snape J W.Mapping quantitative trait loci for flag leaf senescence as a yield determinant in winter wheat under optimal and drought-stressed environments.Euphytica,2004,135:255-263.
[8]Luche H S L,Silva J A G,Nornberg R,Zimmer C M,Arenhardt E G,Caetano V R,Maia L C,Oliveira A C.Stay-green effects on adaptability and stability in wheat.Afr J Agric Res,2015,10:1142-1145.
[9]石慧清,龚月桦,张东武.花后高温对持绿型小麦叶片衰老及籽粒淀粉合成相关酶的影响.植物生态学报,2011,35:769-778.Shi H Q,Gong Y H,Zhang D W.Effect of high temperature on leaf senescence and related enzymes of grain starch synthesis in stay-green wheat after anthesis.Chin J Plant Ecol,2011,35:769-778(in Chinese with English abstract).
[10]Kumari M,Pudake R N,Singh V P,Joshi A K.Association of stay green trait with canopy temperature depression and yield traits under terminal heat stress in wheat(Triticum aestivum L.).Euphytica,2012,190:87-97.
[11]赵致,李家修,张成琦.贵州高原夏秋麦籽粒灌浆特性的研究.作物学报,1998,24:110-116.Zhao Z,Li J X,Zhang C Q.Studies on some characteristics of grain filling of wheat with late summer and early-autumn sowing in Guizhou plateau.Acta Agron Sin,1998,24:110-116(in Chinese with English abstract).
[12]冯素伟,胡铁柱,李淦,董娜,李笑慧,茹振钢,程自华.不同小麦品种籽粒灌浆特性分析.麦类作物学报,2009,29:643-646.Feng S W,Hu T Z,Li G,Dong N,Li X H,Ru Z G,Cheng Z H.Analysis on grain filling characteristics of different wheat varieties.J Triticeae Crops,2009,29:643-646(in Chinese with English abstract).
[13]陈朝儒,奚亚军,王竹林,王永朋,刘曙东,王秀波.冬小麦持绿和灌浆特征及其抗早衰特性评价.西北植物学报,2011,31:715-723.Chen C R,Xi Y J,Wang Z L,Wang Y P,Liu S D,Wang X P.Characteristics of stay-green and grain-filling and the evaluation of anti-senescence properties of winter wheat.Acta Bot Boreali-Occident Sin,2011,31:715-723(in Chinese with English abstract).
[14]张耀兰,曹承富,杜世州,赵竹,乔玉强,刘永华,张四华.淮北地区高产小麦籽粒灌浆特性分析.华北农学报,2010,25(增刊):84-87.Zhang Y L,Cao C F,Du S Z,Zhao Z,Qiao Y Q,Liu Y H,Zhang S H.Analysis on grain filling characteristics of high-yielding wheat in Huaibei area.Acta Agric Boreali-Sin,2010,25(suppl):84-87(in Chinese with English abstract).
[15]吴晓丽,汤永禄,李朝苏,吴春,黄钢,马蓉.四川盆地小麦籽粒的灌浆特性.作物学报,2014,40:337-345.Wu X L,Tang Y L,Li C S,Wu C,Huang G,Ma R.Characteristics of grain filling in wheat growing in Sichuan basin.Acta Agron Sin,2014,40:337-345(in Chinese with English abstract).
[16]周竹青,朱旭彤,王维金.用Richards方程模拟三个典型小麦品种的灌浆过程.湖北农业科学,2001,(6):26-28.Zhou Z Q,Zhu X T,Wang W J.Modeling the filling process of three typical wheat cultivars with Richards equation.Hubei Agric Sci,2001,(6):26-28(in Chinese with English abstract).
[17]任红松,朱家辉,艾比布拉,宋羽,崔新菊,曹连莆.小麦籽粒灌浆特性分析.西北农林科技大学学报(自然科学版),2006,34(3):55-60.Ren H S,Zhu J H,Aibibula,Song Y,Cui X J,Cao L P.Analysis on grain filling characteristics of wheat variety.JNorthwest A&F Univ(Nat Sci Edn),2006,34(3):55-60(in Chinese with English abstract).
[18]薛香,吴玉娥,陈荣江,韩占江,郜庆炉.小麦籽粒灌浆过程的不同数学模型模拟比较.麦类作物学报,2006,26:169-171.Xue X,Wu Y E,Chen R J,Han Z J,Gao Q L.Comparison of different mathematical equations for simulating the grain filling process of wheat.J Triticeae Crops,2006,26:169-171(in Chinese with English abstract).
[19]Van Oosterom E J,Jayachandran R,Bidinger F R.Diallel analysis of the stay-green trait and its components in sorghum.Crop Sci,1996,36:549-555.
[20]Mahalakshmi V,Bidinger F R.Evaluation of stay green sorghum germplasm lines at ICRISAT.Crop Sci,2002,42:965-974.
[21]Kassahun B,Bidinger F R,Hash C T,Kuruvinashetti M S.Stay-green expression in early generation sorghum[Sorghum bicolor(L.)Moench]QTL introgression lines.Euphytica,2010,172:351-362.
[22]刘开昌,董树亭,赵海军,王庆成,李宗新,刘霞,张慧.我国玉米自交系叶片保绿性及其与产量的关系.作物学报,2009,35:1662-1671.Liu K C,Dong S T,Zhao H J,Wang Q C,Li Z X,Liu X,Zhang H.Leaf stay-green traits in Chinese maize inbred lines and their relationship with grain yield.Acta Agron Sin,2009,35:1662-1671(in Chinese with English abstract).
[23]赵延明,张海燕.基于叶片面积玉米叶片保绿度开花后衰减特性初步研究.中国农学通报,2009,25(12):91-94.Zhao Y M,Zhang H Y.The primary studies on reducing characteristics of leaf stay-green based on leaf area after flowering in maize.Chin Agric Sci Bull,2009,25(12):91-94(in Chinese with English abstract).
[24]薛晖,贾丽,龚月桦,刘赢洲,吴养会.冬小麦叶片持绿能力及其衰老特征研究.西北植物学报,2010,30:336-343.Xue H,Jia L,Gong Y H,Liu Y Z,Wu Y H.Study on the stay-green capacity and leaf senescence of winter wheat.Acta Bot Boreali-Occident Sin,2010,30:336-343(in Chinese with English abstract).
[25]Vijayalakshmi K,Fritz A K,Paulsen G M,Bai G,Pandravada S,Gill B S.Modeling and mapping QTL for senescencerelated traits in winter wheat under high temperature.Mol Breed,2010,26:163-175.
[26]梁增浩.不同水分条件下小麦持绿相关性状的QTL定位.山西农业大学硕士学位论文,山西太谷,2013.Liang Z H.Mapping QTLs for Traits Associated with Stay Green in Wheat(Triticum aestivum L.)under Two Water Regimes.MS Thesis of Shanxi Agriculture University,Taigu,Shanxi,China,2013(in Chinese with English abstract).
[27]Pask A J D,Pietragalla J,Mullan D M,Reynolds M P.Physiological Breeding:II.A Field Guide to Wheat Phenotyping.Chapter 12:Leaf area,green crop area and senescence.Mexico:CIMMYT,2012.pp 58-62.
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