柿性器官败育及相关基因的表达
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摘要
为描述柿Diospyros kaki性器官败育现象,以中国林业科学研究院经济林研究开发中心原阳试验基地10年生的雌雄同株柿品种‘禅寺丸’‘Zenjimaru’为研究对象,通过制作石蜡切片结合外部形态变化,同时利用实时定量聚合酶链式反应(qRT-PCR)技术对6个性别分化相关基因进行表达模式分析,对柿花芽分化过程中的性器官败育的关键时期及分子调控机制进行了研究。结果表明:柿雌花中的雄蕊原基和雄花中的雌蕊原基均在4月中旬发生败育。此时,顶端着生雌花的枝条长为5.0～6.0 cm,着生雄花的枝条长为6.0～8.0 cm,雌花芽长约1.0 cm,雄花芽长为1.0～2.0 cm。除了4月5日和4月8日外, ANT和LYK3基因在其他发育阶段均差异显著(P<0.05)且在雌花芽中的表达量明显高于雄花芽,而AP2和ABCC10基因则是在雄花芽中的表达水平显著高于雌花芽(P<0.05)。在4月5日和4月14日(性器官败育关键时期),基因BI-1在雌花芽中的表达量分别是雄花芽的1.5倍和2.0倍,而在其余阶段,其表达量分别是雄花芽的97.5%, 89.2%和83.9%,推测BI-1作为一种雄性器官细胞凋亡抑制基因,间接促进柿雄花芽的发育。建议对树体进行正常的修剪、施肥等常规处理的同时,在4月中旬花器官原基败育之前对树体进行相应激素的喷施,以调控柿花性别,培育雄性种质。
The abortion of sex organs in Diospyros kaki(persimmon) has not yet been clearly elucidated.To investigate the key period for expression patterns dealing with abortion of sex organs and related-genes in persimmon,paraffin section method and morphology changes,as well as a qRT-PCR analysis of six floral sex differentiation-related genes,were conducted with a 10-year-old monoecious persimmon cultivar 'Zenjimaru' in the Yuanyang Practice Base belonging to the Non-timber Forestry Research & Development Center,Chinese Academy of Forestry.Results showed that stamen primordium in female flowers and pistil primordium in male flowers both aborted in mid-April with the female branch 5.0-6.0 cm in length,the male branch 6.0-8.0 cm in length,the female floral bud 1.0 cm in length,and the male floral bud 1.0-2.0 cm in length.In addition,during differentiation,excluding the April 5 th and April 8 th stages,ANT and LYK3 genes were significantly higher(P <0.05) in female floral buds than male;whereas,AP2 and ABCC10 genes in male floral buds were significantly higher than female(P<0.05).The BI-1 level in female floral buds was 1.5 to 2.0 times greater than those in male buds in April 5 th and April 14 th(the key period for abortion of sex organs),respectively,but it was 97.5%,89.2%,and 83.9% in the other stages.Thus,the BI-1 gene may act as an apoptosis-inhibiter and may be indirectly related to development of stamen primordia.Exogenous phytohormones,that regulate the sex of flower buds and cultivate male persimmon germplasm resources,should be considered along with conventional management before abortion of corresponding sexual organs in mid-April.
The abortion of sex organs in Diospyros kaki(persimmon) has not yet been clearly elucidated.To investigate the key period for expression patterns dealing with abortion of sex organs and related-genes in persimmon,paraffin section method and morphology changes,as well as a qRT-PCR analysis of six floral sex differentiation-related genes,were conducted with a 10-year-old monoecious persimmon cultivar 'Zenjimaru' in the Yuanyang Practice Base belonging to the Non-timber Forestry Research & Development Center,Chinese Academy of Forestry.Results showed that stamen primordium in female flowers and pistil primordium in male flowers both aborted in mid-April with the female branch 5.0-6.0 cm in length,the male branch 6.0-8.0 cm in length,the female floral bud 1.0 cm in length,and the male floral bud 1.0-2.0 cm in length.In addition,during differentiation,excluding the April 5 th and April 8 th stages,ANT and LYK3 genes were significantly higher(P <0.05) in female floral buds than male;whereas,AP2 and ABCC10 genes in male floral buds were significantly higher than female(P<0.05).The BI-1 level in female floral buds was 1.5 to 2.0 times greater than those in male buds in April 5 th and April 14 th(the key period for abortion of sex organs),respectively,but it was 97.5%,89.2%,and 83.9% in the other stages.Thus,the BI-1 gene may act as an apoptosis-inhibiter and may be indirectly related to development of stamen primordia.Exogenous phytohormones,that regulate the sex of flower buds and cultivate male persimmon germplasm resources,should be considered along with conventional management before abortion of corresponding sexual organs in mid-April.
引文
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[4]傅建敏,孙鹏,韩卫娟,等.柿雄性种质资源图志[M].北京:中国林业出版社,2017:4.
[5]苏立娟,靳颖,吕琳,等.植物有性生殖过程中的细胞程序性死亡[J].首都师范大学学报(自然科学版),2005,26(2):69-76,84.SU Lijuan,JIN Ying,L譈Lin,et al.Programmed cell death during sexual reproduction in plants[J].J Capital Norm Univ Nat Sci Ed,2005,26(2):69-76,84.
[6]杨同文,李潮海.玉米性别决定的激素调控[J].植物学报,2012,47(1):65-73.YANG Tongwen,LI Chaohai.Hormone regulation of sex determination in maize[J].Chin Bull Bot,2012,47(1):65-73.
[7]周劲松,汤泳萍,盛文涛,等.芦笋性别决定与性别分化研究进展[J].植物遗传资源学报,2010,11(5):600-604.ZHOU Jinsong,TANG Yongping,SHENG Wentao,et al.Progress in research on sex determination and differentiation in Asparagus officinalis L.[J].J Plant Genetic Resour,2010,11(5):600-604.
[8]傍島善次,石田雅士,稲葉昭次,他.カキの発芽期以降における花器の発育について[J].京都府立大學學術報告:農學,1974,26:15-20.SOBAJIMA Y,ISHIDA M,INABA A,et al.Studies on the development of floral organ in Japanese persimmon(cv Hiratanenashi)after sprouring(Agriculture)[J].Sci Rep Kyoto Prefe Univ Agric,1974,26:15-20.
[9]YONEMORI K,SUGIURA A,TANAKA K,et al.Floral ontogeny and sex determination in monoecious-type persimmons[J].J Am Soc Hortic Sci,1993,118(2):293-297.
[10]李加茹,孙鹏,韩卫娟,等.不完全甜柿‘禅寺丸’花性别分化形态学关键时期的研究[J].园艺学报,2016,43(3):451-461.LI Jiaru,SUN Peng,HAN Weijuan,et al.Morphological key period study on floral sex differentiation in pollinationconstant and non-astringent persimmon‘Zenjimaru’[J].Acta Hortic Sin,2016,43(3):451-461.
[11]钟云,刘勇,蒋侬辉,等.柿胚胎发生发育的研究[J].园艺学报,2004,31(3):353-356.ZHONG Yun,LIU Yong,JIANG Nonghui,et al.Studies on the embryogenesis of persimmon[J].Acta Hortic Sin2004,31(3):353-356.
[12]SUN Peng,LI Jiaru,DU Gaigai,et al.Endogenous phytohormone profiles in male and female floral buds of the persimmons(Diospyros kaki Thunb.)during development[J].Sci Hortic,2017,218:213-221.
[13]秦俊芬,葛宇,屈淑平,等.南瓜花芽分化解剖结构的研究[J].东北农业大学学报,2011,42(10):66-69.QIN Junfen,GE Yu,QU Shuping,et al.Anatomy of flower bud differentiation in pumpkin[J].J Northeast Agric U-niv,42(10):66-69.
[14]钱桦,刘燕,郑勇平,等.施用6-BA对春石斛花芽分化及内源激素的影响[J].北京林业大学学报,2009,31(6):27-31.QIAN Hua,LIU Yan,ZHENG Yongping,et al.Effects of applying 6-BA on the nobile-type dendrobium flower bud differentiation and changes of hormones[J].J Beijing For Univ,2009,31(6):27-31.
[15]朱伶俐,周兰英.光周期对‘泰山-橙黄’万寿菊花芽分化和开花的影响[J].东北林业大学学报,2017,45(4):33-35,39.ZHU Lingli,ZHOU Lanying.Effect of photoperiod on Tagetes eresta‘Mount Tai Orange’floral bud differentiation and flowering[J].J Northeast For Univ,2017,45(4):33-35,39.
[16]魏永赞,董晨,王弋,等.荔枝花芽分化与花性别分化研究进展[J].广东农业科学,2017,44(7):34-40.WEI Yongzan,DONG Chen,WANG Yi,et al.Advances in floral bud differentiation and floral sex determination in litchi[J].Guangdong Agric Sci,2017,44(7):34-40.
[17]WETZSTEIN H Y,YI Weiguang,PORTER J A,et al.Flower position and size impact ovule number per flower fruitset,and fruit size in pomegranate[J].J Am Soc Hortic Sci,2013,138(3):159-166.
[18]DELONG A,CALDERON-URREA A,DELLAPORTA A S L.Sex determination gene TASSELSEED2 of maize encodes a shortchain alcohol dehydrogenase required for stage-specific floral organ abortion[J].Cell,1993,74(4)757-768.
[19]张敏,王頔,张雷,等.文冠果雌雄花发育过程形态结构比较[J].电子显微学报,2012,31(2):154-162.ZHANG Min,WANG Di,ZHANG Lei,et al.Female and male flower differentiation in Xanthoceras sorbifolia Bunge[J].J Chin Electr Microsc Soc,2012,31(2):154-162.
[20]HARDENACK S,YE De,SAEDLER H,et al.Comparison of MADS-box gene expression in developing male and female flowers of the dioecious pant white campion[J].Plant Cell,1994,6(12):1775-1787.
[21]杨妙贤,肖德兴,梁红,等.中华猕猴桃性别分化的细胞形态学观察[J].园艺学报,2011,38(2):257-264.YANG Miaoxian,XIAO Dexing,LIANG Hong,et al.Cytomorphological observation on sex differentiation of Actinidia chinensis[J].Acta Hortic Sin,2011,38(2):257-264.
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