欢迎访问作物学报,今天是

作物学报 ›› 2019, Vol. 45 ›› Issue (5): 705-713.doi: 10.3724/SP.J.1006.2019.84121

• 作物遗传育种·种质资源·分子遗传学 • 上一篇    下一篇

芥菜型多室与二室油菜花芽分化过程的比较分析

李新,肖麓,李麟芳,杜德志()   

  1. 青海大学农林科学院 / 青海省春油菜遗传改良重点实验室 / 国家油菜改良中心青海分中心 / 农业部春油菜科学观测实验站 / 青海省春油菜研究开发中心, 青海西宁 810016
  • 收稿日期:2018-09-19 接受日期:2019-01-12 出版日期:2019-05-12 网络出版日期:2019-02-08
  • 通讯作者: 杜德志
  • 基金资助:
    本研究由国家现代农业产业技术体系建设专项(CARS-12);国家自然科学基金项目(31560398);国家重点研发计划项目(2016YFD0101300);青海省春油菜遗传改良重点实验室(2017-ZJ-Y09)

Comparative analysis on flower bud differentiation of multilocular and bilocular traits in Brassica juncea

Xin LI,Lu XIAO,Lin-Fang LI,De-Zhi DU()   

  1. Academy of Agricultural and Forestry Sciences, Qinghai University / Key Laboratory of Qinghai Province for Spring Rapeseed Genetic Improvement / The Qinghai Research Branch of the National Rapeseed Genetic Improvement Center / Spring and Rape Scientific Observation Experimental Station of Ministry of Agriculture and Rural Affair / Qinghai Research and Development Center for Spring Rapeseed, Xining 810016, Qinghai, China
  • Received:2018-09-19 Accepted:2019-01-12 Published:2019-05-12 Published online:2019-02-08
  • Contact: De-Zhi DU
  • Supported by:
    This study was supported by the Earmarked Fund for China Agriculture Research System(CARS-12);the National Nature Science Funds of China(31560398);the National Key Research and Development Plan of China(2016YFD0101300);Key Laboratory of Spring Rape Genetic Improvement of Qinghai Province(2017-ZJ-Y09)

摘要:

芥菜型油菜中存在天然的多室性状。与普通二室油菜相比, 多室油菜因多果室(3~5个腔室)、多粒特征, 具有高产潜力, 为了揭示多室性状的形成过程, 本研究以青海芥菜型多室油菜和塔油2号(芥菜型二室油菜)双亲、BC3F5群体中纯合的二室和多室单株为材料, 通过石蜡切片结合扫描电镜观察, 比较分析表明, 青海芥菜型多室油菜自出苗开始, 约22 d左右, 平均叶片数为5时, 开始花芽分化且经历5个时期; 多室茎顶端分生组织(SAM)和花顶端分生组织(FM)伸长膨大, 直径较二室大; 随心皮分化, 多室花顶端向内凹陷成“X”状的四心皮区域, 而二室花芽中心生长锥向内凹陷成“I”状, 形成二心皮的对半区域, 该凹陷分裂影响心皮数目, 并决定了子房中籽粒的排列数目; 柱头顶端也形成“I”及“X”状凹陷, 可作为一种指标用以早期区分多室及二室性状。

关键词: 多室, 花芽分化, 茎顶端分生组织, 花顶端分生组织, 心皮

Abstract:

Brassica juncea, possessing natural multilocular traits compared with bilocular rapeseed (two-chambered), has high yielding potential due to multiple locules (3-5 locules) and multi-grains. To disclose the formation process of multilocular traits, we constructed a BC3F5 population from a cross between Qinghai multilocular rapeseed and Tayou 2 (two locules) to analyse the whole process of flower bud differentiation of homozygous bilocular plants and multilocular plants through paraffin section along with scanning electron-microscopy observation. The flower bud differentiation in multilocular rapeseed took place at about 22 days after seedling emergence, with an average leaf number of 5, and whole differentiation was divided into five stages. The diameters of shoot apical meristem (SAM) and floral meristem (FM) in multilocular plants were significantly larger than these in bilocular plants. The center of growing cone of the multilocular was deeply recessed into an “X” shaped region with four carpels, while that of the bilocular into an “I” shape, forming half and half regions with two carpels, such differentiation could affect the number of carpels and seeds rows. The tip of the stigma also formed “I” and “X” shapes, which could be used as an indicator, to distinguish between multilocular plants and bilocular plants in the early stage.

Key words: multilocular, flower bud differentiation, shoot apical meristem, floral meristem, carpel

图1

芥菜型二室油菜花芽分化进程图 a: 生长锥伸长, 茎顶端分生组织(SAM)、叶原基(L); 图b: 花原基出现, 茎顶端分生组织(SAM)、花原基(f1~f6); 图c~图e: 花原基不断分化膨大, 茎顶端分生组织(SAM)、花原基(f)、花顶端分生组织(FM)、萼片原基(S); 图f: 花萼原基分化, 花顶端分生组织(FM)、萼片原基(S1~S4); 图g: 花萼(S1~S4), 包裹花顶端; 图h: 花瓣原基分化, 花瓣原基(P)、雄蕊原基(St1~St6)和心皮原基(C1, C2)分化; 图i~图k: 子房腔(OV)的形成及花柱伸长; 图l: 柱头(箭头所指)形成。"

图2

芥菜型多室油菜花芽分化进程图 a: 生长锥伸长, 茎顶端分生组织(SAM)、叶原基(L); 图b: 花原基出现, 茎顶端分生组织(SAM)、花原基(F1~F6); 图c~e: 花原基不断分化膨大, 茎顶端分生组织(SAM)、花原基(F)、花顶端分生组织(FM)、萼片原基(S); 图f: 花萼原基分化, 花顶端分生组织(FM)、萼片原基(S1~S4); 图g: 花萼(S1~S4), 包裹花顶端; 图h: 花瓣原基分化, 花瓣原基(P)、雄蕊原基(St1~St6)和心皮原基(C1~C4)分化; 图i~k: 子房腔(OV)的形成及花柱伸长; 图k~l: 柱头(箭头所指)形成。"

表1

大田环境二室和多室油菜花芽组织的分化时间"

类型
Type
生长锥
Growth cone
花原基
Flower
primordium
萼片原基
Sepal
primordium
花瓣原基
Petal
primordium
雄蕊原基
Stamen
primordium
心皮
Carpel
二室Bilocular 2 21 27 33 40 45
多室Multilocular 2 22 31 38 51 52

图3

多室与二室花芽分化比较 图a~b: 二室茎顶端分生组织; 图c: 二室花顶端分生组织; 图d~e: 多室茎顶端分生组织; 图f: 多室花顶端分生组织; 图g: 二室心皮分化(C1, C2); 图h: 二室子房腔(lc); 图i: 二室籽粒着生(O); 图j: 多室心皮分化(C1~C4); 图k: 多室子房腔(lc); 图l: 多室籽粒着生(O); 图m~n: 二室角果内部结构; 图o: 二室柱头; 图p~q: 多室角果内籽粒排列结构; 图r: 多室柱头; P: 花瓣原基; St: 雄蕊原基; C: 心皮; lc: 心室; O: 胚珠; M: 中间区域; L: 横向区域。"

图4

二室与多室SAM及FM的比较 **表示在0.01概率水平上显著相关; *表示在0.05概率水平上显著相关。"

[1] 刘后利 . 油菜遗传育种学. 北京: 中国农业大学出版社, 2000. pp 215-225.
Liu H L. Rapeseed Genetics and Breeding. Beijing: China Agricultural University Press, 2000. pp 215-225(in Chinese).
[2] 赵洪朝, 杜德志, 刘青元, 李秀萍, 余青兰, 付忠 . 芥菜型多室油菜的主要性状表现. 西北农业学报. 2003,12(3):62-64.
Zhao H C, Du D Z, Liu Q Y, Li X P, Yu Q L, Fu Z . Performances in main characteristics of multilocular B. juncea. Acta Agric Boreali-occident Sin, 2003,12(3):62-64 (in Chinese with English abstract).
[3] 冷锁虎, 唐瑶, 李秋兰, 左青松, 杨萍 . 油菜的源库关系研究: I. 角果大小对油菜后期源库的调节. 中国油料作物学报, 2005,27(3):39-42.
Leng S H, Tang Y, Li Q L, Zuo Q S, Yang P . Studies on source and sink of rapeseed: I. Regulation of pod size on source and sink in rapeseed after flowering. Chin J Oil Crop Sci, 2005,27(3):39-42 (in Chinese with English abstract).
[4] 吕泽文, 徐平, 张向向, 文静, 易斌, 马朝芝, 涂金星, 傅廷栋, 沈金雄 . 芥菜型油菜多室角果的解剖特征及遗传分析. 中国油料作物学报, 2012,34:461-466.
Lyu Z W, Xu P, Zhang X X, Wen J, Yi B, Ma C Z, Tu J X, Fu T D, Shen J X . Primary study on anatomic and genetic characteristics of multi-loculus in Brassica juncea. Chin J Oil Crop Sci, 2012,34:461-466 (in Chinese with English abstract).
[5] Sinhamahapatra S P, Raman R B, Roy S, Roy U S, Raut N M, Ashok K V . Breeding for an ideal plant type in yellow sarson ( Brassica rapa L. yellow sarson). Electronic J Plant Breed, 2010,1:689-694.
[6] Katiyar R K, Chamola R, Chopra V L . Tetralocular mustard, Brassica juncea: new promising variability through interspecific hybridization. Plant Breed, 1998,117:398-399.
[7] 蒙祖庆, 宋丰萍 . 白菜型多室黄籽油菜农艺性状的典型相关分析. 种子, 2012,31(4):96-98.
Meng Z Q, Song F P . Typical correlation analysis on main agronomic and yield characters of yellow-seeded multicapsular silique in Brassica campestris L. Seed, 2012,31(4):96-98 (in Chinese).
[8] 马月萍, 戴思兰 . 植物花芽分化机制研究进展. 分子植物育种, 2003,1:539-545.
Ma Y P, Dai S L . Flower bud differentiation mechanism of anthophyta. Mol Plant Breed, 2003,1:539-545 (in Chinese with English abstract).
[9] 曲波, 张微, 陈旭辉, 李楠, 崔娜, 李天来 . 植物花芽分化研究进展. 中国农学通报, 2010,26(24):109-114.
Qu B, Zhang W, Chen X H, Li N, Cui N, Li T L . Research progress of flower bud differentiation mechanism of plant. Chin Agric Sci Bull, 2010,26(24):109-114 (in Chinese with English abstract).
[10] 官春云 . 油菜花芽分化的研究. 湖南农业科学, 1983, ( 4):20-22.
Guan C Y . Study on flower bud differentiation in rapeseed. Hunan Agric Sci, 1983, ( 4):20-22 (in Chinese).
[11] 张朋飞, 孙万仓, 武军艳, 方彦, 张俊杰, 姜海洋, 陈姣荣, 李学才 . 北方寒旱区冬油菜花芽分化过程观察. 西北农业学报, 2011,20(11):60-65.
Zhang P F, Sun W C, Wu J Y, Fang Y, Zhang J J, Jiang H Y, Chen J R, Li X C . Flower differentiation of winter rapeseed in Northwest cold arid regions. Acta Agric Boreali-occident Sin, 2011,20(11):60-65 (in Chinese with English abstract).
[12] 丁秀琦 . 白菜型春油菜花芽分化研究. 作物学报, 1990,16:83-90.
Ding X Q . Studies on the floral buds differentiation in spring rapes ( Brassica campestris L.). Acta Agron Sin, 1990,16:83-90 (in Chinese with English abstract).
[13] 尹继春, 严敦秀, 张燕, 谢志坚 . 关于甘蓝型油菜的花芽分化研究: 油菜花芽分化的电镜扫描图谱. 作物学报, 1984,10:179-184.
Yin J C, Yan D X, Zhang Y, Xie Z J . On the differentiation of floral buds in rape ( Brassica napus): a sketch of scanning pattern by electron microscope. Acta Agron Sin, 1984,10:179-184 (in Chinese with English abstract).
[14] 何余堂, 龙卫华, 胡进平, 傅廷栋, 李殿荣, 陈宝元, 涂金星 . 白菜型油菜角果多室性状的遗传及解剖学研究. 中国油料作物学报, 2003,25(1):1-4.
He Y T, Long W H, Hu J P, Fu T D, Li D R, Chen B Y, Tu J X . Anatomic and genetic studies on multicapsular character in Brassica campestris L. Chin J Oil Crop Sci, 2003,25(1):1-4 (in Chinese with English abstract).
[15] 赵洪朝 . 芥菜型油菜多室性状的遗传表现及其与甘蓝型油菜的种间远缘杂交 . 西北农林科技大学硕士学位论文, 陕西西安, 2003.
Zhao H C . Study on Multilocular Heredity of B. juncea and Interpecific Hybridization between B. juncea and B. napus. MS Thesis of Northwest A&F University, Xi’an, Shaanxi, China, 2003 (in Chinese with English abstract).
[16] 赵洪朝, 杜德志, 刘青元, 余青兰, 王瑞生 . 芥菜型油菜多室性状的遗传研究. 西北农林科技大学学报(自然科学版), 2003,31(6):90-92.
Zhao H C, Du D Z, Liu Q Y, Yu Q L, Wang R S . Study on multilocular heredity of B. juncea. J Northwest A&F Univ ( Nat Sci Edn), 2003,31(6):90-92 (in Chinese with English abstract).
[17] 朱彦涛, 胡选萍, 刘湛, 李殿荣, 郭蔼光 . 油菜角果多室性状及其研究进展. 中国油料作物学报, 2012,34:321-325.
Zhu Y T, Hu X P, Liu Z, Li D R, Guo A G . Research progress on multi-loculus silique of rapeseed. Chin J Oil Crop Sci, 2012,34:321-325 (in Chinese with English abstract).
[18] 赵会彦 . 芥菜型油菜多室基因Bjln1定位的研究 . 青海大学硕士学位论文, 青海西宁, 2014.
Zhao H Y . Fine Mapping of Multilocular Gene Bjln1 in Brassica juncea L. MS Thesis of Qinghai University, Xining, Qinghai, China, 2014 (in Chinese with English abstract).
[19] 赵文路 . 种间杂交创建的甘蓝型多室油菜的多室等位性研究及其不育材料JN55A不育类型的鉴别. 青海大学硕士学位论文, 青海西宁, 2007.
Zhao W L . Study on Multilocular Alleles and Identified Sterile Style of JN55A CMS in Brassica napus with Multiloculus Created by Interspecific Crossing. MS Thesis of Qinghai University, Xining, Qinghai, China, 2007 (in Chinese with English abstract).
[20] 吕泽文 . 芥菜型油菜多室性状的遗传分析及其基因的分子标记. 华中农业大学硕士学位论文, 湖北武汉, 2011
Lyu Z W . Study on Inheritance and Molecular Markers for Multilocular Trait in Brassica juncea. MS Thesis of Huazhong Agricultural University, Wuhan, Hubei, China, 2011 (in Chinese with English abstract).
[21] 王刚, 张向向, 徐平, 吕泽文, 文静, 易斌, 马朝芝, 涂金星, 傅廷栋, 沈金雄 . 芥菜型油菜多室基因Bjmc2的精细定位. 作物学报, 2016,42:1735-1742.
Wang G, Zhang X X, Xu P, Lyu Z W, Wen J, Yi B, Ma C Z, Tu J X, Fu T D, Shen J X . Fine mapping of polycyetic gene ( Bjmc2) in Brassica juncea L. Acta Agron Sin, 2016,42:1735-1742 (in Chinese with English abstract).
[22] Choudhary B R, Solanki Z S . Inheritance of siliqua locule number and seed coat colour in Brassica juncea. Plant Breed, 2007,126:104-106.
[23] Xu P, Lyu Z W, Zhang X X, Wang X H, Pu Y Y, Wang H M, Yi B, Wen J, Ma C Z, Tu J X, Fu T D, Shen J X . Identification of molecular markers linked to trilocular gene ( mc1) in Brassica juncea L. Mol Breed, 2014,33:425-434.
[24] Coen E S, Meyerowitz E M . The war of the whorls: genetic interactions controlling flower development. Nature, 1991,353:31-37.
[25] Xiao L, Zhao H Y, Zhao Z, Du D Z, Xu L, Yao Y M, Zhao Z G, Xing X R, Shang G X, Zhao H C . Genetic and physical fine mapping of a multilocular gene Bjln1 in Brassica juncea to a 208-kb region. Mol Breed, 2013,32:373-383.
[26] Fan C C, Wu Y D, Yang Q Y, Meng Q W, Zhang K Q, Li J G, Wang J F, Zhou Y M . A novel single-nucleotide mutation in a CLAVATA3 gene homolog controls a multilocular silique trait in Brassica rapa L. Mol Plant, 2014,7:1788-1792.
[27] Xiao L, Li X, Liu F, Zhao Z, Xu L, Chen C P, Wang Y H, Shang G X, Du D Z . Mutations in the CDS and promoter of BjuA07.CLV1 cause a multilocular trait in Brassica juncea. Sci Rep, 2018,8:5339.
[28] 张玉英 . 制作石蜡切片易出现的问题及改进措施. 中国科技信息, 2007, ( 6):91-92.
Zhang Y Y . Problems and improvement measures of making paraffin sections. Chin Sci Technol Inf, 2007, ( 6):91-92 (in Chinese).
[29] 安彩泰, 陈军 . 甘蓝型油菜自交不亲和系柱头和花粉粒不亲和互作的扫描电镜研究. 甘肃农业大学学报, 1992, ( 2):159-162.
An C T, Chen J . The interactions of stima papills and pollen grains in self-incompatible line of rape under scanning electron microscope. J Gansu Agric Univ, 1992, ( 2):159-162 (in Chinese with English abstract).
[30] 侯燕鸣, 胡剑江, 方蔷, 张倩, 雷洪涛, 王毅 . 扫描电镜的不同含水量植物叶片样品的处理及观察方法研究. 分析仪器, 2011, ( 5):45-48.
Hou Y M, Hu J J, Fang Q, Zhang Q, Lei H T, Wang Y . Study on sample treatment methods of plant leaves with different water contents for SEM. Analy Instrument, 2011, ( 5):45-48 (in Chinese with English abstract).
[31] 梅德圣, 李云昌 . 中国油菜高产育种研究进展. 湖北农业科学, 2003, ( 4):35-39.
Mei D S, Li Y C . Advances in high yield breeding of rapeseed in China. Hubei Agric Sci, 2003, ( 4):35-39 (in Chinese).
[32] 张锦芳, 周贤琼, 蒲晓斌, 李浩杰, 蒋梁材 . 高产、双低杂交油菜产量构成因素与产量的相关分析. 西南农业学报, 2008,21:939-941.
Zhang J F, Zhou X Q, Pu X B, Li H J, Jiang L C . Correlation analysis between yield components and yield per plant of high yield rapeseed hybrid. Southwest Chin J Agric Sci, 2008,21:939-941 (in Chinese with English abstract).
[33] 胡虹文 . 甘蓝型油菜12种主要性状与产量的关系. 中国油料, 1997,19(3):10-11.
Hu H W . Relationship between 12 main characters and yield of rapeseed ( Brassica napus L.). Chin J Oil Crop Sci, 1997,19(3):10-11 (in Chinese).
[34] Schoof H, Lenhard M, Haecker A , Mayer K F X, Jürgens G, Laux T . The stem cell population of Arabidopsis shoot meristems is maintained by a regulatory loop between the CLAVATA and WUSCHEL genes. Cell, 2000,100:635-644.
[35] Jiang Y, Zu W, Wu C X . Maintenance, transition and reversion of the shoot apical meristem during plant development. Chin J Cell Biol, 2008,30:147-152.
[36] Stahl Y, Grabowski S, Bleckmann A, Kühnemuth R, Weidtkamp- Peters S, Pinto K G, Kirschner G K, Schmid J B, Wink R H, Hülsewede A, Felekyan S , Seidel C A M, Simon R . Moderation of Arabidopsis root stemness by CLAVATA1 and ARABIDOPSIS CRINKLY4 receptor kinase complexes. Curr Biol, 2013,23:362-371.
[37] Ogawa M, Shinohara H, Sakagami Y, Matsubayashi Y . Arabidopsis CLV3 peptide directly binds CLV1 ectodomain. Science, 2008,319:294-294.
[38] Durbak A R, Tax F E . CLAVATA signaling pathway receptors of Arabidopsis regulate cell proliferation in fruit organ formation as well as in meristems. Genetics, 2011,189:177-194.
[39] Song S K, Lee M M, Clark S E . POL and PLL1 phosphatases are CLAVATA1 signaling intermediates required for Arabidopsis shoot and floral stem cells. Development, 2006,133:4691-4698.
[40] Muños S, Ranc N, Botton E, Bérard A, Rolland S, Duffé P, Carretero Y, Paslier M L, Delalande C, Bouzayen M, Brunel D, Causse M . Increase in tomato locule number is controlled by two single-nucleotide polymorphisms located near WUSCHEL. Plant Physiol, 2011,156:2244-2254.
[1] 赵改会, 李书宇, 詹杰鹏, 李晏斌, 师家勤, 王新发, 王汉中. 甘蓝型油菜角果数突变体基因的定位及候选基因分析[J]. 作物学报, 2022, 48(1): 27-39.
[2] 杨阳, 李淮琳, 胡利民, 范楚川, 周永明. 白菜型油菜srb多室性状的遗传分析与分子鉴定[J]. 作物学报, 2021, 47(3): 385-393.
[3] 王刚,张向向,徐平,吕泽文,文静,易斌,马朝芝,涂金星,傅廷栋,沈金雄*. 芥菜型油菜多室基因Bjmc2的精细定位[J]. 作物学报, 2016, 42(12): 1735-1742.
[4] 张焱;官春云. 内源赤霉素与油菜不同种性品种花芽分化的关系的研究[J]. 作物学报, 1993, 19(04): 365-371.
[5] 丁秀琦. 白菜型春油菜花芽分化研究[J]. 作物学报, 1990, 16(01): 83-90.
Viewed
Full text


Abstract

Cited

  Shared   
  Discussed   
No Suggested Reading articles found!