偏孟德尔分离的遗传学实验设计与探讨
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摘要
本文将科研中发现的一个含调控杂种花粉不育基因座(S23)的水稻材料(DSSL)应用于教学,设计了一个基于SSR分子标记对比经典孟德尔分离和偏孟德尔分离的综合性实验。利用位于水稻两条染色体上的4个SSR标记对两亲本及其杂交构建的F_2代群体进行单株基因型检测,用显微图像观察与统计分析亲本及杂种F1的花粉育性,不仅从分子水平上验证了分离定律,更从基因型到表型的观察实验中完整展现了水稻的偏孟德尔分离现象及其原因,加深了学生对植物遗传规律、基因型与表型关系的理解,激发了学生对实验的兴趣与探究动机,增强了学生对实验学习的自觉性和积极性。并在此基础上构思一个科研成果转化至教学应用的可持续发展思路,以推动实验教学的改革与创新。
It has always been a challenge to combine research progress with undergraduate laboratory teaching. Herein we designed a comprehensive experiment to compare classical Mendelian segregation and non-Mendelian distorted segregation by utilizing a rice material(DSSL) containing F1 hybrid male sterility locus S23 constructed previously in our research project. Using the four SSR markers located on two chromosomes of rice, the genotypes of the F2 population and the two parents were analyzed, and the phenotypes of the pollen fertility of the two parents and their F1 plants were investigated. The results not only verified segregation law at the molecular level, but also fully demonstrated the distorted segregation in both genotypes and phenotypes, thus deepening students' understandings of plant genetics and the relationship between genotypes and phenotypes, inspiring students' interests in genetics experiments, and enhancing students' consciousness and enthusiasm for experimental learning. On the basis of this, a sustainable development idea of transforming scientific research progress into teaching applications was conceived to promote the reform and innovation of genetics laboratory teaching.
It has always been a challenge to combine research progress with undergraduate laboratory teaching. Herein we designed a comprehensive experiment to compare classical Mendelian segregation and non-Mendelian distorted segregation by utilizing a rice material(DSSL) containing F1 hybrid male sterility locus S23 constructed previously in our research project. Using the four SSR markers located on two chromosomes of rice, the genotypes of the F2 population and the two parents were analyzed, and the phenotypes of the pollen fertility of the two parents and their F1 plants were investigated. The results not only verified segregation law at the molecular level, but also fully demonstrated the distorted segregation in both genotypes and phenotypes, thus deepening students' understandings of plant genetics and the relationship between genotypes and phenotypes, inspiring students' interests in genetics experiments, and enhancing students' consciousness and enthusiasm for experimental learning. On the basis of this, a sustainable development idea of transforming scientific research progress into teaching applications was conceived to promote the reform and innovation of genetics laboratory teaching.
引文
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[2]Liu XD,Guo HB,Fu XL,Zhang GQ.Genetics teaching reform experiment based on innovative mixed teaching model.High Agric Educ,2008,(8):57-60.刘向东,郭海滨,傅雪琳,张桂权.基于创新型混合教学模式的遗传学教改试验.高等农业教育,2008,(8):57-60.
[3]Liu XD,Guo HB.Experiment course integration for modern agronomic undergraduate program:reform and practice.China Agric Educ,2013,(5):60-63.刘向东,郭海滨.现代农学专业实验课整合改革研究--以华南农业大学为例.中国农业教育,2013,(5):60-63.
[4]Huang XY,Fan K,Ye YF,Wang B,Wu WR,Lan T.Teaching practice and experiences of verifying the three laws of genetics based on the SSLP marker analysis.Hereditas(Beijing),2017,39(9):856-862.黄雪盈,范凯,叶炎芳,汪斌,吴为人,兰涛.基于SSLP分子标记验证遗传学三大定律的教学实践探索与体会.遗传,2017,39(9):856-862.
[5]Sobrizal,Matsuzaki Y,Sanchez PL,Ikeda K,Yoshimura A.Mapping of F1 pollen semi-sterility gene found in backcross progeny of Oryza sativa L.and Oryza glumaepatula steud.Rice Genet Newsl,2000,17:61-63.
[6]Yu X,Zhao Z,Zheng X,Zhou J,Kong W,Wang P,Bai W,Zheng H,Zhang H,Li J,Liu J,Wang Q,Zhang L,Liu K,Yu Y,Guo X,Wang J,Lin Q,Wu F,Ren Y,Zhu S,Zhang X,Cheng Z,Lei C,Liu S,Liu X,Tian Y,Jiang L,Ge S,Wu C,Tao D,Wang H,Wan J.A selfish genetic element confers non-Mendelian inheritance in rice.Science,2018,360(6393):1130-1132.
[7]Liu HY,Cui JT,Gao YM.Progress of segregation distortion.J Plant Genet Resour,2009,10(4):613-617,622.刘海燕,崔金腾,高用明.遗传群体偏分离研究进展.植物遗传资源学报,2009,10(4):613-617,622.
[8]Cameron DR,Moav RM.Inheritance in Nicotiana tabacum Xxvii.pollen killer,an alien genetic locus inducing abortion of microspores not carrying it.Genetics,1957,42(3):326-335.
[9]Scoles GJ,Kibirge-Sebunya IN.Preferential abortion of gametes in wheat induced by an Agropyron chromosome.Can J Genet Cytol,1983,25(1):1-6.
[10]Rick CM.Abortion of male and female gametes in the tomato determined by allelic interaction.Genetics,1966,53(1):85-96.
[11]Zhang Y,Tan L,Zhu Z,Yuan L,Xie D,Sun C.TOND1confers tolerance to nitrogen deficiency in rice.Plant J,2015,81(3):367-376.
[12]Mangelsdorf PC,Jones DF.The expression of mendelian factors in the gametophyte of maize.Genetics,1926,11(6):423-455.
[13]Wang Z.Distorted segregation in plant hybrids and its implication for evolution.Hereditas(Beijing),2016,38(9):801-810.王哲.植物杂交后代中基因偏分离的产生原因及其进化意义.遗传,2016,38(9):801-810.
[14]Ma SJ,Liu YG,Liu JX.Research Progress of hybrid sterility of rice.J Plant Genet Resour,2014,15(5):1080-1088.马生健,刘耀光,刘金祥.水稻的杂种不育研究进展.植物遗传资源学报,2014,15(5):1080-1088.
[15]Xi ZY,He FH,Zeng RZ,Zhang ZM,Ding XH,Li WT,Zhang GQ.Development of a wide population of chromosome single-segment substitution lines in the genetic background of an elite cultivar of rice(Oryza sativa L.).Genome,2006,49(5):476-484.
[16]He N,Wu R,Pan X,Peng L,Sun K,Zou T,Zhu H,Zeng R,Liu Z,Liu G,Wang S,Zhang G,Fu X.Development and trait evaluation of chromosome single-segment substitution lines of O.meridionalis in the background of O.sativa.Euphytica,2017,213(12):281.
[17]Li DS,Cui DF,Jiang QY,Chen R.Strengthening the practice of basic experiment course teaching in agricultural specialty.Chin Univ Teach,2008,(10):86-88.李大胜,崔大方,江青艳,陈然.加强农科专业基础课实验教学的实践.中国大学教学,2008,(10):86-88.
[2]Liu XD,Guo HB,Fu XL,Zhang GQ.Genetics teaching reform experiment based on innovative mixed teaching model.High Agric Educ,2008,(8):57-60.刘向东,郭海滨,傅雪琳,张桂权.基于创新型混合教学模式的遗传学教改试验.高等农业教育,2008,(8):57-60.
[3]Liu XD,Guo HB.Experiment course integration for modern agronomic undergraduate program:reform and practice.China Agric Educ,2013,(5):60-63.刘向东,郭海滨.现代农学专业实验课整合改革研究--以华南农业大学为例.中国农业教育,2013,(5):60-63.
[4]Huang XY,Fan K,Ye YF,Wang B,Wu WR,Lan T.Teaching practice and experiences of verifying the three laws of genetics based on the SSLP marker analysis.Hereditas(Beijing),2017,39(9):856-862.黄雪盈,范凯,叶炎芳,汪斌,吴为人,兰涛.基于SSLP分子标记验证遗传学三大定律的教学实践探索与体会.遗传,2017,39(9):856-862.
[5]Sobrizal,Matsuzaki Y,Sanchez PL,Ikeda K,Yoshimura A.Mapping of F1 pollen semi-sterility gene found in backcross progeny of Oryza sativa L.and Oryza glumaepatula steud.Rice Genet Newsl,2000,17:61-63.
[6]Yu X,Zhao Z,Zheng X,Zhou J,Kong W,Wang P,Bai W,Zheng H,Zhang H,Li J,Liu J,Wang Q,Zhang L,Liu K,Yu Y,Guo X,Wang J,Lin Q,Wu F,Ren Y,Zhu S,Zhang X,Cheng Z,Lei C,Liu S,Liu X,Tian Y,Jiang L,Ge S,Wu C,Tao D,Wang H,Wan J.A selfish genetic element confers non-Mendelian inheritance in rice.Science,2018,360(6393):1130-1132.
[7]Liu HY,Cui JT,Gao YM.Progress of segregation distortion.J Plant Genet Resour,2009,10(4):613-617,622.刘海燕,崔金腾,高用明.遗传群体偏分离研究进展.植物遗传资源学报,2009,10(4):613-617,622.
[8]Cameron DR,Moav RM.Inheritance in Nicotiana tabacum Xxvii.pollen killer,an alien genetic locus inducing abortion of microspores not carrying it.Genetics,1957,42(3):326-335.
[9]Scoles GJ,Kibirge-Sebunya IN.Preferential abortion of gametes in wheat induced by an Agropyron chromosome.Can J Genet Cytol,1983,25(1):1-6.
[10]Rick CM.Abortion of male and female gametes in the tomato determined by allelic interaction.Genetics,1966,53(1):85-96.
[11]Zhang Y,Tan L,Zhu Z,Yuan L,Xie D,Sun C.TOND1confers tolerance to nitrogen deficiency in rice.Plant J,2015,81(3):367-376.
[12]Mangelsdorf PC,Jones DF.The expression of mendelian factors in the gametophyte of maize.Genetics,1926,11(6):423-455.
[13]Wang Z.Distorted segregation in plant hybrids and its implication for evolution.Hereditas(Beijing),2016,38(9):801-810.王哲.植物杂交后代中基因偏分离的产生原因及其进化意义.遗传,2016,38(9):801-810.
[14]Ma SJ,Liu YG,Liu JX.Research Progress of hybrid sterility of rice.J Plant Genet Resour,2014,15(5):1080-1088.马生健,刘耀光,刘金祥.水稻的杂种不育研究进展.植物遗传资源学报,2014,15(5):1080-1088.
[15]Xi ZY,He FH,Zeng RZ,Zhang ZM,Ding XH,Li WT,Zhang GQ.Development of a wide population of chromosome single-segment substitution lines in the genetic background of an elite cultivar of rice(Oryza sativa L.).Genome,2006,49(5):476-484.
[16]He N,Wu R,Pan X,Peng L,Sun K,Zou T,Zhu H,Zeng R,Liu Z,Liu G,Wang S,Zhang G,Fu X.Development and trait evaluation of chromosome single-segment substitution lines of O.meridionalis in the background of O.sativa.Euphytica,2017,213(12):281.
[17]Li DS,Cui DF,Jiang QY,Chen R.Strengthening the practice of basic experiment course teaching in agricultural specialty.Chin Univ Teach,2008,(10):86-88.李大胜,崔大方,江青艳,陈然.加强农科专业基础课实验教学的实践.中国大学教学,2008,(10):86-88.