青花菜细胞核雄性不育系Y37S的细胞学观察及遗传分析

doi:10.16420/j.issn.0513-353x.2025-0148

摘要/Abstract

摘要：

青花菜细胞核雄性不育材料Y37S是在自交系Y37中发现的一株自然突变不育株，为明确其遗传特性和花药败育原因，从形态学、细胞学、遗传及分子鉴定等方面进行研究。结果表明：Y37S不育系植株雄蕊退化，显著低于柱头，无花粉。扫描电镜观察结果显示，Y37S花药干瘪扭曲，呈现三角形，无花粉粒附着；利用荧光素双醋酸酯（fluorescein diacetate，FDA）对不同大小花蕾的小孢子活力进行检测，不育株花蕾发育至4.5 ~ 5.0 mm时，小孢子活力丧失且无绿色荧光信号。花药细胞学特征观察发现，不育株在单核小孢子期发生小孢子塌陷、绒毡层细胞延迟降解且降解不完全，影响了花药和花粉的正常功能，最终导致败育。用同系内6个可育株分别与Y37S进行杂交，其中Y37-1和Y37-3自交可育株与不育株比例为23︰7和17︰5，Y37-1和Y37-3与Y37S的杂交后代可育株与不育株比例分别为15︰14和28︰30，初步推测Y37S受一对隐性核基因控制。为验证其遗传规律，用Y37S与正常可育自交系B50杂交，F₁代全部可育，F₂群体中可育与不育株比为107︰29，测交群体中可育与不育株比为67︰69，符合孟德尔遗传定律，证明Y37S是由一对隐性基因控制的细胞核雄性不育系。利用已经鉴定的白菜和甘蓝单隐性核不育基因连锁的分子标记对测交群体中的不育株和可育株进行扩增，未扩增出差异条带，推测控制不育系Y37S育性的基因可能为新的隐性核不育基因。

关键词: 青花菜, 细胞核雄性不育, 遗传分析, 分子标记

Abstract:

The nuclear male sterile material Y37S of broccoli is a naturally mutated sterile strain discovered in the inbred line Y37. To clarify its genetic characteristics and the causes of anther failure，studies were conducted from aspects such as morphology，cytology，genetics and molecular identification. The results showed that the stamens of the Y37S sterile line plants were degenerated，significantly lower than the stigma，and there was no pollen. The results of the scanning electron microscope showed that the anthers of Y37S were shriveled and twisted，presenting a triangular shape，with no pollen grains attached. The microspore viability of flower buds of different sizes was detected by fluorescein diacetate（FDA）. When the flower buds of sterile plants developed to 4.5-5.0 mm，the microspore viability was lost and there was no green fluorescence signal. Observation of the cytological characteristics of anthers revealed that in sterile plants，microspore collapse occurred during the mononuclear microspore stage，and the degradation of felt-like layer cells was delayed and incomplete，which affected the normal functions of anthers and pollen，ultimately leading to infertility. Six fertile strains within the same line were crossed with Y37S respectively. Among them，the ratios of self-pollinated fertile strains to sterile strains of Y37-1 and Y37-3 were 23︰7 and 17︰5，respectively，and the ratios of fertile strains to sterile strains of the hybrid offspring of Y37-1 and Y37-3 with Y37S were 15︰14 and 28︰30，respectively. It is preliminarily speculated that Y37S is controlled by a pair of recessive nuclear genes. To verify its genetic pattern，Y37S was crossed with the normal fertile inbred line B50. All F₁ generations were fertile. The ratio of fertile to sterile lines in the F₂ population was 107︰29，and in the test cross population，it was 67︰69. This conforms to Mendel’s law of inheritance and proves that Y37S is a nuclear male sterile line controlled by a pair of recessive genes. The sterile and fertile plants in the test population were amplified using the molecular markers of the single recessive nuclear sterility gene linkage of Chinese cabbage and cabbage that have been identified. No differential bands were amplified，suggesting that the gene controlling the fertility of the sterile line Y37S might be a new recessive nuclear sterility gene.

Key words: broccoli, nuclear male sterility, enetic analysis, molecular marker

姚雪琴, 周维, 刘春晴, 黄雷, 姜晶, 龚静, 李光庆, 谢祝捷. 青花菜细胞核雄性不育系Y37S的细胞学观察及遗传分析[J]. 园艺学报, 2026, 53(1): 99-109.

YAO Xueqin, ZHOU Wei, LIU Chunqing, HUANG Lei, JIANG Jing, GONG Jing, LI Guangqing, XIE Zhujie. Cytological Observation and Genetic Analysis of the Nuclear Male Sterile Line Y37S of Broccoli[J]. Acta Horticulturae Sinica, 2026, 53(1): 99-109.

图/表 7

参考文献 40

[1]	Chen Fengxiang, Hu Baocheng, Dai Bingxun, Hou Shumin, Li Qiangsheng, Wu Xinjie, Fei Weixin, Zhang Manlin. 1998. Genetic comparison of recessive superior interaction nuclear sterile lines 9012A with 6AB and S45AB of Rapeseed of the Brassica lanensis type. Anhui Agricultural Sciences,(3):5-7. (in Chinese)
	陈凤祥, 胡宝成, 代秉勋, 候树敏, 李强生, 吴新杰, 费维新, 张曼琳. 1998. 甘蓝型油菜隐性上位互作核不育系9012A与6AB和S45AB遗传对比. 安徽农业科学,(3):5-7.
[2]	Chen Zhuxizhao. 1985. Electron microscopic observation of microspore occurrence and sterile process in nuclear sterile wheat in Taigu. Scientia Agricultura Sinica,(5):16-20,96-98. (in Chinese)
	陈朱希昭. 1985. 太谷核不育小麦小孢子发生和败育过程的电子显微镜观察. 中国农业科学,(5):16-20,96-98.
[3]	Dong Faming. 2012. Genetic pattern correction and physical localization of BnRf loci in the nuclear sterile line 9012AB of rapeseed glauca[Ph. D. Dissertation]. Wuhan: Huazhong Agricultural University. (in Chinese)
	董发明. 2012. 甘蓝型油菜核不育系9012AB的遗传模式修正与BnRf位点的物理定位[博士论文]. 武汉: 华中农业大学.
[4]	Fang Zhiyuan, Liu Yumei, Yang Limei, Wang Xiaowu, Zhuang Mu, Zhang Yangyong, Sun Peitian. 2004. Breeding and seed production technique of dominant genic male sterile(DGMS)line and cytoplasmic male sterile(CMS)line in cabbage. Scientia Agricultura Sinica, 37 (5):717-723. (in Chinese)
	方智远, 刘玉梅, 杨丽梅, 王晓武, 庄木, 张扬勇, 孙培田. 2004. 甘蓝显性核基因雄性不育与胞质雄性不育系的选育及制种. 中国农业科学, 37 (5):717-723.
[5]	Fang Zhiyuan, Sun Peitian, Liu Yumei, Yang Limei, Wang Xiaowu. 1997. Breeding and utilization of dominant male sterile lines in cabbage. Acta Horticulturae Sinica, 24 (3):44-49. (in Chinese)
	方智远, 孙培田, 刘玉梅, 杨丽梅, 王晓武. 1997. 甘蓝显性雄性不育系的选育及其利用. 园艺学报, 24 (3):44-49.
[6]	Gao Rruihong, He Ting, Guo Guimei, Huang Lei, Yao Xxueqin, Lu Ruiju, Xie Zhujie, Liu Chenghong. 2023. Optimization of microspore culture and plant regeneration system of broccoli(Brassica oleracea var. italica). Molecular Plant Breeding, https://link.cnki.net/urlid/46.1068.S.20231130.1544.004. (in Chinese)
	高润红, 何婷, 郭桂梅, 黄雷, 姚雪琴, 陆瑞菊, 谢祝捷, 刘成洪. 2023. 青花菜小孢子培养及再生体系的优化. 分子植物育种, https://link.cnki.net/urlid/46.1068.S.20231130.1544.004.
[7]	Han F Q, Yuan K W, Kong C C, Zhang X L, Yang L M, Zhuang M, Zhang Y Y, Li Z X, Wang Y, Fang Z Y, Lv H H. 2018. Fine mapping and candidate gene identification of the genic male-sterile gene ms3 in cabbage 51S. Theor Appl Genet,131:2651-2661.
[8]	Han F Q, Yuan K W, Sun W R, Zhang X L, Liu X, Zhao X Y, Yang L M, Wang Y, Ji J L, Liu Y M, Li Z S, Zhang J Z, Zhang C Z, Huang S W, Zhang Y Y, Fang Z Y, Lv H H. 2023. A natural mutation in the promoter of Ms-cd 1 causes dominant male sterility in Brassica oleracea. Nat Commun,14:6212.
[9]	Huang Z, Chen Y, Yi B, Xiao L, Ma C Z,Tu, J X, Fu T D. 2007. Fine mapping of the recessive genic male sterility gene(Bnms3)in Brassica napus L. Theor Appl Genet,115:113-118.
[10]	Kang Jungen. 2006. Cytology and gene expression profiling of four types of male sterility in cabbage(B. oleracea var. capitata)[Ph. D. Dissertation]. Beijing: Chinese Academy of Agricultural Sciences. (in Chinese)
	康俊根. 2006. 四种类型甘蓝雄性不育系花药败育特征及基因表达谱分析[博士论文]. 北京: 中国农业科学院.
[11]	Lei S L, Yao X Q, Yi B, Chen W, Ma C Z, Tu J X, Fu T D. 2007. Towards map-based cloning:fine mapping of a recessive genic male-sterile gene (BnMs2)in Brassica napus L. and syntenic region identification based on the Arabidopsis thaliana genome sequences. Theor Appl Genet,115:643-651.
[12]	Li Shulin, Qian Yuxiu, Wu Zhihua. 1985. Discussion on the genetic law of male sterility in the nuclei of Brassica napus and its application. Shanghai Journal of Agricultural Sciences,(2):1-12. (in Chinese)
	李树林, 钱玉秀, 吴志华. 1985. 甘蓝型油菜细胞核雄性不育性的遗传规律探讨及其应用. 上海农业学报,(2):1-12.
[13]	Li Zhansheng, Liu Yumei, Fang Zhiyuan, Yang Limei, Zhuang Mu, Zhang Yangyong, Lü Honghao, Wang Yong. 2019. Development status,existing problems and countermeasures of China’s broccoli industry. Chinese Vegetables,(4):1-5. (in Chinese)
	李占省, 刘玉梅, 方智远, 杨丽梅, 庄木, 张扬勇, 吕红豪, 王勇. 2019. 我国青花菜产业发展现状、存在问题与应对策略. 中国蔬菜,(4):1-5.
[14]	Liu Yumei. 2003. Research on the cytological characteristics,biochemical basis and molecular markers of dominant nuclear male sterility in cabbage[Ph. D. Dissertation]. Beijing: Chinese Academy of Agricultural Sciences. (in Chinese)
	刘玉梅. 2003. 甘蓝显性细胞核雄性不育的细胞学特征、生化基础及其分子标记的研究[博士论文]. 北京: 中国农业科学院.
[15]	Luo C C, Sun Y Y, Zhang Y X, Guo Y, Klima M, Hu S W. 2018. Genetic investigation and cytological comparison of two genic male sterile lines 9012A and MSL in Brassica napus L. Euphytica,214:124.
[16]	Niu Y, Wu L M, Li Y H, Huang H L, Qian M C, Sun W, Zhu H, Xu Y F, Fan Y H, Mahmood U, Xu B B, Zhang K, Qu C M, Li J N, Lu K. 2020. Deciphering the transcriptional regulatory networks that control size,color,and oil content in Brassica rapa seeds. Biotechnol Biofuels,13:90.
[17]	Pan Tao, Zeng Fanya, Wu Shuhui, Zhao Yun. 1988. Research on the breeding and utilization of male sterile dual-purpose lines of Brassica napus with low erucic acid. Chinese Oil Products,(3):7-10. (in Chinese)
	潘涛, 曾凡亚, 吴书惠, 赵云. 1988. 甘蓝型低芥酸油菜雄性不育两用系的选育与利用研究. 中国油料,(3):7-10.
[18]	Qi Baoying. 2013. Breeding of a new type of Rapeseed recessive nuclear sterile line 72A[M. D. Dissertation]. Changsha: Hunan Agricultural University. (in Chinese)
	祁宝英. 2013. 新型甘蓝型油菜隐性细胞核不育系72A的选育[硕士论文]. 长沙: 湖南农业大学.
[19]	Song Laiqing, Fu Tingdong, Yang Guangsheng, Tu Jingxing, Ma Chaozhi. 2005. Genetic validation of a multiple allele controlled dominant nuclear sterile line 609AB in rape(Brassica napus L.). Acta Agronomica Sinica, 31 (7):869-875. (in Chinese)
	宋来强, 傅廷栋, 杨光圣, 涂金星, 马朝芝. 2005. 1对复等位基因控制的油菜(Brassica napus L.) 显性核不育系609AB的遗传验证. 作物学报, 31 (7):869-875.
[20]	Song Renjing, Zhang Qingqin, Liu Linong, Li Huaqin. 1988. Cytological abservations on anther and pollen developments of qianxing wale sterile and sterility maintainer lines of wheat. Guizhou Agricultural Sciences,(4):7-13. (in Chinese)
	宋仁敬, 张庆勤, 刘丽农, 李华琴. 1988. 黔型小麦雄性不育系及其保持系花药和花粉发育的细胞形态学观察. 贵州农业科学,(4):7-13.
[21]	Tan C, Liu Z Y, Huang S N, Feng H. 2019. Mapping of the male sterile mutant gene ftms in Brassica rapa L. ssp. pekinensis via BSR-Seq combined with whole-genome resequencing. Theor Appl Genet,132:355-370.
[22]	Tang Xin, Li Yuanyuan, Lu Junxing, Zhang Tao. 2021. Morphological and cytological study on anther abortion of thermosensitive nuclear male sterile line 160S in Brza napus. Acta Agronomica Sinica, 47 (5):983-990. (in Chinese) doi: 10.3724/SP.J.1006.2021.04207 URL
	唐鑫, 李圆圆, 陆俊杏, 张涛. 2021. 甘蓝型油菜温敏细胞核雄性不育系160S花药败育的形态学特征和细胞学研究. 作物学报, 47 (5):983-990. doi: 10.3724/SP.J.1006.2021.04207
[23]	Wan L L, Xia X Y, Hong D F, Li J, Yang G S. 2010. Abnormal vacuolization of the tapetum during the tetrad stage is associated with male sterility in the recessive genic male sterile Brassica napus L. line 9012A. J Plant Biol,53:121-133.
[24]	Wan Lili. 2010. Cytology studies with two types of genic male sterility and characterization of fertility related genes in rapeseed(Brassica napus. L).[Ph. D. Dissertation]. Wuhan: Huazhong Agricultural University. (in Chinese)
	万丽丽. 2010. 油菜细胞核雄性不育的细胞学研究以及育性相关基因的克隆与功能分析[博士论文]. 武汉: 华中农业大学.
[25]	Wang Fuqing, Wang Cuilan, Song Zaihua. 2001. Morphological observation on anther and pollen development of male sterile line 88-3 of Chinese cabbage. Acta Botanica Sinica, 21 (3):570-574,621. (in Chinese)
	王福青, 王翠兰, 宋再华. 2001. 大白菜雄性不育系88-3花药和花粉发育的细胞形态学观察. 西北植物学报, 21 (3):570-574,621.
[26]	Wang Guohuai, Guan Chunyun, Chen Sheyuan, Liu Benkun, Liu Zhong. 2003. Study on the utilization of nutritional advantages of interspecific hybrids of Brassicae in Cruciferae. Acta Cropologica Sinica, 29 (1):54-58. (in Chinese)
	王国槐, 官春云, 陈社员, 刘本坤, 刘忠. 2003. 十字花科芸薹属种间杂种营养优势的利用研究. 作物学报, 29 (1):54-58.
[27]	Wang Jiansheng, Shen Yusen, Yu Huifang, Sheng Xiaoguang, Song Mengfei, Gu Honghui. 2024. Research progress of broccoli breeding in China. Journal of Zhejiang Agricultural Sciences, 36 (8):1934-1944. (in Chinese)
	王建升, 沈钰森, 虞慧芳, 盛小光, 宋蒙飞, 顾宏辉. 2024. 中国西兰花育种研究进展. 浙江农业学报, 36 (8):1934-1944. doi: 10.3969/j.issn.1004-1524.20240449
[28]	Wang Xinzhe. 2017. Development of molecular markers for nuclear male sterility genes in Chinese kale[M. D. Dissertation]. Shenyang: Shenyang Agricultural University. (in Chinese)
	王馨哲. 2017. 白菜薹细胞核雄性不育基因分子标记的开发[硕士论文]. 沈阳: 沈阳农业大学.
[29]	Xiong X P, Li X Y, Zhang S T, Hu Zn, Liu T T, Qiu Z M, Cao J S, Huang L, Yan C H. 2024. Identification and fine mapping of Brmmd1 gene controlling recessive genic male sterility in Brassica rapa L. Gene,924:148558.
[30]	Xin Q, Wang X, Gao Y P, Xu D D, Xie Z Q, Dong F M, Wan L L, Yang L Y, Yang G S, Hong D F. 2020. Molecular mechanisms underpinning the multiallelic inheritance of MS 5 in Brassica napus. Plant J,103:1723-1734.
[31]	Xu Daixiang. 2018. Cytological observation and omics analysis of homozygous two-type dominant sterility line D3AB in Brassica napus[M. D. Dissertation]. Chongqing: Southwest University. (in Chinese)
	许代香. 2018. 甘蓝型油菜显性核不育纯合两型系D3AB的细胞学观察与组学分析[硕士论文]. 重庆: 西南大学.
[32]	Xu Junjie. 2022. Fine localization and candidate gene identification of male sterility mutant gene BrABCG26 in Chinese cabbage[Ph. D. Dissertation]. Shenyang: Shenyang Agricultural University. (in Chinese)
	徐均杰. 2022. 大白菜雄性不育突变基因BrABCG26的精细定位与候选基因鉴定[博士论文]. 沈阳: 沈阳农业大学.
[33]	Xue Meihui. 2023. Clone and identification of male-sterile mutant gene BrRNR1 in Chinese cabbage(Brassica rapa L. ssp.)[Ph. D. Dissertation]. Shenyang: Shenyang Agricultural University. (in Chinese)
	薛美惠. 2023. 大白菜雄性不育突变基因BrRNR1的克隆与鉴定[博士论文]. 沈阳: 沈阳农业大学
[34]	Yan X H, Zeng X H, Wang S S, Li K Q, Yuan R, Gao H F, Luo J L, Liu F, Wu Y H, Li Y J, Zhu L, Wu G. 2016. Aberrant meiotic prophase I leads to genic male sterility in the novel TE5A mutant of Brassica napus. Sci Rep,6:33955.
[35]	Yang Guangsheng, Fu Tingdong. 1991. A prelominary study on the restoring-maintaining relationship in rapeseed(Brassica napus and Brassica campestris). Acta Agronomica Sinica, 17 (2):151-156. (in Chinese)
	杨光圣, 傅廷栋. 1991. 油菜细胞质雄性不育恢保关系的研究. 作物学报, 17 (2):151-156.
[36]	Yang Guangsheng, Qu Bo, Fu Tingdong. 1999. Cytological study on microspore formation in three recessive nuclear male sterile lines of rapeseed of Brachycephalus orientalis. Journal of Huazhong Agricultural University, 18 (6):520-523. (in Chinese)
	杨光圣, 瞿波, 傅廷栋. 1999. 三个甘蓝型油菜隐性细胞核雄性不育系小孢子发生的细胞学研究. 华中农业大学学报, 18 (6):520-523.
[37]	Yi B, Chen Y N, Lei S L, Tu J X, Fu T D. 2006. Fine mapping of the recessive genic male-sterile gene(Bnms1)in Brassica napus L. Theor Appl Genet,113:643-650.
[38]	Yi Bin, Tu Jinxing, Fu Tingdong. 2014. Study and utilization of recessive nuclear male sterility in Brassica napus. Science China:Life Sciences, 44 (8):752-757. (in Chinese)
	易斌, 涂金星, 傅廷栋. 2014. 甘蓝型油菜隐性细胞核雄性不育的研究及利用. 中国科学:生命科学, 44 (8):752-757.
[39]	Zhao Ying. 2022. Cloning and identification of male sterile genes BrGGL7 and BrRBL3 in Chinese cabbage(Brassica rapa ssp. pekinensis)[Ph. D. Dissertation]. Shenyang: Shenyang Agricultural University. (in Chinese)
	赵莹. 2022. 大白菜雄性不育基因BrGGL7和BrRBL3的克隆与鉴定[博士论文]. 沈阳: 沈阳农业大学.
[40]	Zu Feng. 2009. Molecular markers of male sterility genes in recessive cell nuclei of rapeseed brassicolor[M. D. Dissertation]. Wuhan: Huazhong Agricultural University. (in Chinese)
	俎峰. 2009. 甘蓝型油菜隐性细胞核雄性不育基因的分子标记[硕士论文]. 武汉: 华中农业大学.

不育基因 Sterile genes	引物名称 Primer name	正向引物（5′-3′） Forward primer sequence	反向引物（5′-3′） Reverse primer sequence
Brnnd1	HABS378	CCGAAGACTAGAACGTCCAA	ACAATGAACACACCGGTCTA
Brnnd1	HABS386	GAATTCTAGCAGGTAGCTGACG	GCGGAAACCTTTGTGAGTTT
BoTPD1	YIndel0116	TGCAGAGTTGAGTCACGATG	TGTTGGCTGTTCTGCGATAC
	YIndel012	CGCCTTGCTTCTATTCCAAG	TGTTGGCTGTTCTGCGATAC
	Ms3-dec	GAATGTATCAGGCGTGGAGG	TCACCGATTTGGTATGAGTTT

不育基因 Sterile genes	引物名称 Primer name	正向引物（5′-3′） Forward primer sequence	反向引物（5′-3′） Reverse primer sequence
Brnnd1	HABS378	CCGAAGACTAGAACGTCCAA	ACAATGAACACACCGGTCTA
Brnnd1	HABS386	GAATTCTAGCAGGTAGCTGACG	GCGGAAACCTTTGTGAGTTT
BoTPD1	YIndel0116	TGCAGAGTTGAGTCACGATG	TGTTGGCTGTTCTGCGATAC
	YIndel012	CGCCTTGCTTCTATTCCAAG	TGTTGGCTGTTCTGCGATAC
	Ms3-dec	GAATGTATCAGGCGTGGAGG	TCACCGATTTGGTATGAGTTT

群体 Group	可育株数 Number of fertile plants		分离比 Separation ratio	卡方值 Value	χ²_0.05,1
Y37S × Y37-1	15	14	1.00︰0.93	0.034	3.84
Y37S × Y37-2	18	0	全部可育All fertile	0	3.84
Y37S × Y37-3	28	30	1.00︰1.07	0.069	3.84
Y37S × Y37-4	52	0	全部可育All fertile	0	3.84
Y37S × Y37-5	47	0	全部可育All fertile	0	3.84
Y37S × Y37-6	28	0	全部可育All fertile	0	3.84
Y37-1 ⊗	23	7	3.28︰1.00	0.044	3.84
Y37-2 ⊗	31	9	3.44︰1.00	0.133	3.84
Y37-3 ⊗	17	5	3.40︰1.00	0.060	3.84
Y37-4 ⊗	37	0	全部可育All fertile	0	3.84
Y37-5 ⊗	22	0	全部可育All fertile	0	3.84
Y37-6 ⊗	26	0	全部可育All fertile	0	3.84
F₁（Y37S × B50）	34	34	全部可育All fertile	0	3.84
F₂（Y37S × B50）	107	29	3.68︰1.00	0.980	3.84
B₁（Y37S × F₁）	67	69	1.00︰1.02	0.029	3.84

群体 Group	可育株数 Number of fertile plants		分离比 Separation ratio	卡方值 Value	χ²_0.05,1
Y37S × Y37-1	15	14	1.00︰0.93	0.034	3.84
Y37S × Y37-2	18	0	全部可育All fertile	0	3.84
Y37S × Y37-3	28	30	1.00︰1.07	0.069	3.84
Y37S × Y37-4	52	0	全部可育All fertile	0	3.84
Y37S × Y37-5	47	0	全部可育All fertile	0	3.84
Y37S × Y37-6	28	0	全部可育All fertile	0	3.84
Y37-1 ⊗	23	7	3.28︰1.00	0.044	3.84
Y37-2 ⊗	31	9	3.44︰1.00	0.133	3.84
Y37-3 ⊗	17	5	3.40︰1.00	0.060	3.84
Y37-4 ⊗	37	0	全部可育All fertile	0	3.84
Y37-5 ⊗	22	0	全部可育All fertile	0	3.84
Y37-6 ⊗	26	0	全部可育All fertile	0	3.84
F₁（Y37S × B50）	34	34	全部可育All fertile	0	3.84
F₂（Y37S × B50）	107	29	3.68︰1.00	0.980	3.84
B₁（Y37S × F₁）	67	69	1.00︰1.02	0.029	3.84