新疆与美国早实核桃(Juglans regia L.)品种生长结实与成花特性比较
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摘要
以引进的美国核桃品种‘维纳’、‘艾米格’和新疆本地品种‘温185’和‘新新2号’为试材,研究分析不同品种的植物学和生物学特性、花芽分化过程以及相关成花基因的表达。结果显示:4个品种在树体生长、枝芽发育特性、物侯期以及成花率和坐果率方面没有明显差异。在结果枝同一坐果部位形成双果、三果或以上的能力‘新新2号’明显强于‘维纳’,但‘新新2号’的果实大小和单果重都低于‘维纳’,而‘维纳’平均结果母枝上形成侧花芽的数量和坐果率高于‘新新2号’。从萌芽到开花结束直至当年侧芽发育过程中‘维纳’混合芽中的JrFT和JrLFY基因表达也高于‘新新2号’,而同时期的JrFLC基因表达则相对较低,说明‘维纳’在花芽分化和花器官形成过程中通过高表达JrFT和JrLFY基因,低表达JrFLC基因促进结果母枝上形成大量侧花芽,并利用大量侧花芽抽生结果枝坐果,形成较大的果实,使得最终产量并不低于同一个坐果部位大量着生2个或多个果实的‘新新2号’品种。
American walnut industry occupies an important position in the world walnut industry with its large nut,thin shell and high yield.Xinjiang is one of the main producing areas of China's walnut and has a long history of cultivation.However,in recent years,cultivation and management techniques were not updated in time,which resulted in unstable yield,uneven quality,poor overall efficiency of walnut and other issues in the walnut production in Xinjiang.In this study,the introduced American walnut varieties‘vina',‘Amigo'and Xinjiang local varieties‘Wen 185'and‘Xinxin2'were used as materials to compare the botanical and biological characteristics of different varieties of flower buds differentiation process and expression of related flowering genes.The results showed that there was no significant difference among the four cultivars in tree growth,development characteristics of buds,astern stage,as well as flower rate and fruit setting rate.However,the ability to create double fruit,three fruit or more in the same branch of fertile branch of‘Xinxin2'was significantly higher than thatof‘vina'.The size and fruit weight of‘Xinxin2'were lower than‘vina''.The average flower buds on the shoots and fruit setting rate of‘vina was higher than that of‘Xinxin2'.At the same time,the expressions of JrFTand JrLFYgenes in mixed buds from bud to flower stage of‘vina'were also higher than those of‘Xinxin2',while the expression of JrFLC gene in the bud was relatively lower.The results showed that‘vina'increased the expressions of JrFTand JrLFYgenes during flower bud differentiation and flower organ formation,and formed a large number of lateral flower buds on the shoots.The use of a large number of lateral buds results fruiting branches,the formation of larger fruits with low expression of JrFLC gene.So its final yield produced was not lower than the‘Xinxin2'of the same set of fruit parts with two or more fruits.
American walnut industry occupies an important position in the world walnut industry with its large nut,thin shell and high yield.Xinjiang is one of the main producing areas of China's walnut and has a long history of cultivation.However,in recent years,cultivation and management techniques were not updated in time,which resulted in unstable yield,uneven quality,poor overall efficiency of walnut and other issues in the walnut production in Xinjiang.In this study,the introduced American walnut varieties‘vina',‘Amigo'and Xinjiang local varieties‘Wen 185'and‘Xinxin2'were used as materials to compare the botanical and biological characteristics of different varieties of flower buds differentiation process and expression of related flowering genes.The results showed that there was no significant difference among the four cultivars in tree growth,development characteristics of buds,astern stage,as well as flower rate and fruit setting rate.However,the ability to create double fruit,three fruit or more in the same branch of fertile branch of‘Xinxin2'was significantly higher than thatof‘vina'.The size and fruit weight of‘Xinxin2'were lower than‘vina''.The average flower buds on the shoots and fruit setting rate of‘vina was higher than that of‘Xinxin2'.At the same time,the expressions of JrFTand JrLFYgenes in mixed buds from bud to flower stage of‘vina'were also higher than those of‘Xinxin2',while the expression of JrFLC gene in the bud was relatively lower.The results showed that‘vina'increased the expressions of JrFTand JrLFYgenes during flower bud differentiation and flower organ formation,and formed a large number of lateral flower buds on the shoots.The use of a large number of lateral buds results fruiting branches,the formation of larger fruits with low expression of JrFLC gene.So its final yield produced was not lower than the‘Xinxin2'of the same set of fruit parts with two or more fruits.
引文
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[12]Hu J Y,Zhou Y,He F,Dong X,Liu L Y,Coupland G,TurckF.Juliette de Meaux.miR824-regulated AGAMOUSLIKE16 contributes toflowering time repression in Arabidopsis[J].Plant Cell,2014,26(5):2024-2037
[13]Kardailsky I,Shukla V K,Ahn J H,Dagenais N,Christensen S K,Nguyen J T,Chory J,Harrison M J,Weigel D.Activation tagging of the floral inducer FT[J].Science,1999,286:1962-1965
[14]Fornara F,Panigrahi K C,Gissot L,Sauerbrunn N,Rühl M,Jarillo J A,Coupland G.Arabidopsis DOF transcription factors act redundantly to reduce CONSTANS expression and are essential for a photoperiodic flowering response[J].Dev Cell,2009,17:75-86
[15]温腾健,马凯,韩立群,王继勋,胡建芳.新疆核桃‘温185’和‘新新2号’花芽分化特点与成花基因表达分析[J].中国农业大学学报,2017,22(4):28-36Wen T J,Ma K,Han L Q,Wang J X,Hu J F.Differentiation of Xinjiang walnut flower bud and analysis of related flowering gene expression[J].Journals of China Agricultural University,2017,22(4):28-36(in Chinese)
[16]郗荣庭.中国果树科学与实践[M].西安:陕西科学技术出版社,2015Xi R T.TheScience and Practice of Chinese Fruit Tree[M].Xi'an:Shanxi Science and Technology Press,2015(in Chinese)
[17]赵宝军,宫永红.早实核桃产量及其构成因子研究[C]//干果研究进展.北京:中国农业科技出版社,2001,71-73Zhao B J,Gong Y H.Study on the yield and its constituent factors of early real walnut[C].In:Research Progress of Dried Fruit.Beijing:China Agricultural Science and Technology Press,2001,71-73(in Chinese)
[18]高英,董宁光,张志宏,张俊佩,裴东.早实核桃雌花芽分化外部形态与内部结构关系的研究[J].林业科学研究,2010,23(2):241-245Gao Y,Dong N G,Zhang Z H,Zhang J P,Pei D.Relationship between external morphology and anatomical structure of precocious walnut during pistillate flower bud differentiation[J].Forest Research,2010,23(2):241-245(in Chinese)
[19]李永涛,赵勇刚,杨克强,房瑶瑶,侯立群.早实核桃花器官发育的解剖学研究[J].园艺学报,2011,38(3):434-440Li Y T,Zhao Y G,Yang K Q,Fang Y Y,Hou L Q.Studies on floral organ development process of precocious walnut(Juglansregia)by Anatomia[J].ActaHorticulturae Sinica,2011,38(3):434-440(in Chinese)
[20]Srikanth A,Schmid M.Regulation of flowering time:All roads lead to Rome[J].Cellular and Molecular Life Sciences:CMLS,2011,68(12):2013-2037
[21]宋杨,窦连登,张红军.高等植物成花诱导调控的分子和遗传机制[J].植物生理学报,2014,50(10):1459-1468Song Y,Dou L D,Zhang H J.Molecular and genetic mechanisms of control of floral induction in higher plants[J].Plant Physiology Journal,2014,50(10):1459-1468(in Chinese)
[2]新疆维吾尔自治区统计局.新疆统计年鉴:2016[M].北京:中国统计出版社,2016:396Xinjiang Uygur Autonomous Region Bureau of Statistics.Xinjiang Statistical Yearbook:2016[M].Beijing:China Stiatistics Press,2013(in Chinese)
[3]王国安,张强,黄闽敏,阿卜杜许库尔·牙合甫.新疆核桃产业现状与分析[C]//干果研究进展.北京:中国农业科技出版社,2013(11):5-8Wang G A,Zhang Q,Huang M M,Yakup Abduxukur.The status and analysis of the walnut industry in Xinjiang[C].In:Research Progress of Dried Fruit.Beijing:China Agricultural Science and Technology Press,2013(11):5-8(in Chinese)
[4]陆斌,宁德鲁.美国核桃产业发展综述及其借鉴[J].林业调查规划,2011,36(3):98-105Lu B,Ning D L.American walnut industry overview and its reference function[J].Forest Inventory and Planning,2011,36(3):98-105(in Chinese)
[5]奚声坷,王哲理,游应天.美国核桃,黑核桃引种试验[J].林业科学研究,1995,8(3):285-290Xi S K,Wang Z L,You Y T.Introduction of american walnuts(JuglanL)in China[J].Forest Research,1995,8(3):285-290(in Chinese)
[6]马贯羊,杨建学,乔保水,赖政武,葛向珂.8个美国核桃品种的引种表现[J].落叶果树,2009,2:33-35Ma G Y,Yang J X,Qiao B S,Lai Z W,Ge X K.Introduction of 8american walnut varieties[J].Deciduous Fruits,2009,2:33-35(in Chinese)
[7]Y Akca,Ozongun S.Selection of late leafing,late flowering,laterally fruitful walnut(Juglansregia)types in Turkey[J].New Zealand Journal of Crop&Horticultural Science,2004,32(4):337-342
[8]Sabatier S,Barthelemy D.Bud structure in relation to shoot morphology and position on thevegetative annual shoots of JuglansregiaL(Juglandaceae)[J].Annals of Botany,2001,87(1):117-123
[9]Amasino,R.Seasonal and developmental timing of flowering[J].Plant Journal,2010,61:1001-1013
[10]Wang JW Regulation of flowering time by the miR156-mediated age pathway[J].Journal Experimental Biology,2014 65:4723-4730
[11]Han P,Garcia-Ponce B,Fonseca-Salazar G,Alvarez-Buylla E R,Yu H.AGAMOUS-LIKE 17,a novel flowering promoter,acts in a FT independent photoperiod pathway[J].Plant Journal,2008,55:253-265
[12]Hu J Y,Zhou Y,He F,Dong X,Liu L Y,Coupland G,TurckF.Juliette de Meaux.miR824-regulated AGAMOUSLIKE16 contributes toflowering time repression in Arabidopsis[J].Plant Cell,2014,26(5):2024-2037
[13]Kardailsky I,Shukla V K,Ahn J H,Dagenais N,Christensen S K,Nguyen J T,Chory J,Harrison M J,Weigel D.Activation tagging of the floral inducer FT[J].Science,1999,286:1962-1965
[14]Fornara F,Panigrahi K C,Gissot L,Sauerbrunn N,Rühl M,Jarillo J A,Coupland G.Arabidopsis DOF transcription factors act redundantly to reduce CONSTANS expression and are essential for a photoperiodic flowering response[J].Dev Cell,2009,17:75-86
[15]温腾健,马凯,韩立群,王继勋,胡建芳.新疆核桃‘温185’和‘新新2号’花芽分化特点与成花基因表达分析[J].中国农业大学学报,2017,22(4):28-36Wen T J,Ma K,Han L Q,Wang J X,Hu J F.Differentiation of Xinjiang walnut flower bud and analysis of related flowering gene expression[J].Journals of China Agricultural University,2017,22(4):28-36(in Chinese)
[16]郗荣庭.中国果树科学与实践[M].西安:陕西科学技术出版社,2015Xi R T.TheScience and Practice of Chinese Fruit Tree[M].Xi'an:Shanxi Science and Technology Press,2015(in Chinese)
[17]赵宝军,宫永红.早实核桃产量及其构成因子研究[C]//干果研究进展.北京:中国农业科技出版社,2001,71-73Zhao B J,Gong Y H.Study on the yield and its constituent factors of early real walnut[C].In:Research Progress of Dried Fruit.Beijing:China Agricultural Science and Technology Press,2001,71-73(in Chinese)
[18]高英,董宁光,张志宏,张俊佩,裴东.早实核桃雌花芽分化外部形态与内部结构关系的研究[J].林业科学研究,2010,23(2):241-245Gao Y,Dong N G,Zhang Z H,Zhang J P,Pei D.Relationship between external morphology and anatomical structure of precocious walnut during pistillate flower bud differentiation[J].Forest Research,2010,23(2):241-245(in Chinese)
[19]李永涛,赵勇刚,杨克强,房瑶瑶,侯立群.早实核桃花器官发育的解剖学研究[J].园艺学报,2011,38(3):434-440Li Y T,Zhao Y G,Yang K Q,Fang Y Y,Hou L Q.Studies on floral organ development process of precocious walnut(Juglansregia)by Anatomia[J].ActaHorticulturae Sinica,2011,38(3):434-440(in Chinese)
[20]Srikanth A,Schmid M.Regulation of flowering time:All roads lead to Rome[J].Cellular and Molecular Life Sciences:CMLS,2011,68(12):2013-2037
[21]宋杨,窦连登,张红军.高等植物成花诱导调控的分子和遗传机制[J].植物生理学报,2014,50(10):1459-1468Song Y,Dou L D,Zhang H J.Molecular and genetic mechanisms of control of floral induction in higher plants[J].Plant Physiology Journal,2014,50(10):1459-1468(in Chinese)