禾本科三倍体: 形成、鉴定与利用

doi:10.11983/CBB20166

植物学报 ›› 2021, Vol. 56 ›› Issue (3): 372-387.DOI: 10.11983/CBB20166

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禾本科三倍体: 形成、鉴定与利用

严旭¹^,², 左艳春¹, 王红林¹, 李杨²^,³, 李影正², 寇晶¹, 唐祈林¹, 周晓康²^,^*(), 杜周和¹^,^*()

¹四川省农业科学院蚕业研究所/牧业研究中心, 南充 637000
²四川农业大学玉米研究所, 温江 611130
³绵阳师范学院城乡建设与规划学院, 绵阳 621000

收稿日期:2020-10-09 接受日期:2021-01-21 出版日期:2021-05-01 发布日期:2021-04-30
通讯作者: 周晓康,杜周和
作者简介:duzhouhe@126.com
^*E-mail: tangqilin71@163.com;
基金资助:
四川省重点研发项目(2019YFN0022);现代农业学科建设推进工程No(2021XKJS096);四川省南充市研发项目No(19YFZJ0060);四川省十四五饲草育种攻关No(2021YFYZ0013)

Triploid in Poaceae: Formation, Detection, and Utilization

Xu Yan¹^,², Yanchun Zuo¹, Honglin Wang¹, Yang Li²^,³, Yingzheng Li², Jing Kou¹, Qilin Tang¹, Xiaokang Zhou²^,^*(), Zhouhe Du¹^,^*()

¹Sericulture Research Institute/Animal Husbandry Research Center, Sichuan Academy of Agricultural Sciences, Nanchong 637000, China
²Maize Research Institute, Sichuan Agricultural University, Wenjiang 611130, China
³School of Urban and Rural Planning and Construction, Mianyang Teachers’ College, Mianyang 621000, China

Received:2020-10-09 Accepted:2021-01-21 Online:2021-05-01 Published:2021-04-30
Contact: Xiaokang Zhou,Zhouhe Du

摘要/Abstract

摘要： 禾本科三倍体的形成途径包括2n配子融合、倍性间杂交、多精受精和胚乳培养。其中, 2n配子融合和倍性间杂交分别为自然界和人工合成三倍体的主要途径。该文介绍了形态学观测、染色体分析、流式细胞术和分子标记等倍性鉴定方法在禾本科三倍体中的应用及其优缺点。目前, 三倍体在禾谷类作物中无直接应用价值, 但可作为通往多倍体、非整倍体和转移异源基因的遗传桥梁。多年生禾本科三倍体(特别是异源三倍体)在饲草或能源作物中已得到广泛应用, 在该类型禾本科作物中均可直接尝试三倍体育种。多倍体的三倍体育种和无融合生殖三倍体育种可作为未来禾本科三倍体的研究方向。三倍性胚乳培养可以一步合成三倍体, 多精受精可以实现遗传上3个不同基因组的一步融合, 在三倍体研究中应予以重视。鉴于2n配子融合、多精受精的稀有特性和倍性间杂交、胚乳培养频繁的染色体变异, 高通量三倍体鉴定技术的发展将是三倍体研究实现突破的关键。

关键词: 三倍体, 2n配子, 倍性间杂交, 胚乳培养, 多精受精, 育种利用

Abstract: The formation pathway of triploid in Poaceae includes 2n gamete fusion, interploidy cross, polyspermy, and endosperm culture. The fusion of reduced male (n) and unreduced female gamete (2n) is the main method for forming natural triploids. Interploidy cross is the main method for synthesizing artificial triploids. The application of ploidy identification methods such as morphological observation, chromosome analysis, flow cytometry and molecular markers in gramineous triploids is introduced, and the advantages and disadvantages of different methods are also discussed. At present, triploid has no direct application value in cereal crops, but it can be used as a genetic bridge to synthetize polyploid and aneuploid, as well as to transfer alien genes from wild species to cultivated species. Gramineous triploids (especially allotriploid) are widely cultivated for forage or biofuel production, suggesting that triploidy breeding may be directly performed in this type of grasses. We discuss the future prospect of research on gramineous triploid, e.g., polyploid- and apomixis-triploid breeding. Particularly, endosperm culture can synthesize triploids in one step, and polyspermy can achieve one-step fusion of three genetically different plant genomes, which should be paid attention to in the triploidy research. Due to rare occurrence of 2n gamete fusion and polyspermy, and frequent chromosomal variation in ploidy hybridization and endosperm culture, the development of high-throughput triploid identification technology will become the key for breakthrough in triploidy generation/breeding.

Key words: triploid, 2n gamete, interploidy cross, endosperm culture, polyspermy, breeding program

严旭, 左艳春, 王红林, 李杨, 李影正, 寇晶, 唐祈林, 周晓康, 杜周和. 禾本科三倍体: 形成、鉴定与利用. 植物学报, 2021, 56(3): 372-387.

Xu Yan, Yanchun Zuo, Honglin Wang, Yang Li, Yingzheng Li, Jing Kou, Qilin Tang, Xiaokang Zhou, Zhouhe Du. Triploid in Poaceae: Formation, Detection, and Utilization. Chinese Bulletin of Botany, 2021, 56(3): 372-387.

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链接本文: https://www.chinbullbotany.com/CN/10.11983/CBB20166

https://www.chinbullbotany.com/CN/Y2021/V56/I3/372

图/表 4

图1 禾本科三倍体的形成途径 (A) 精细胞与卵细胞受精产生二倍性合子后发育成二倍体胚; (B) 精细胞与中央细胞受精产生三倍性初生胚乳细胞后发育成胚乳; (C) 通过离体培养胚或种子萌发发育成二倍体; (D) 通过胚乳组织培养形成三倍体; (E) 通过2n+n生殖方式形成三倍体; (F) 通过二倍体加倍后形成的同源四倍体(2n=4x=AAAA)与二倍体(2n=2x=AA)杂交合成同源三倍体; (G) 通过四倍体(2n=4x=AAAA)与二倍体(2n=2x=BB)杂交合成异源三倍体; (H) 通过多精受精形成三倍体

Figure 1 Pathways to triploidy formation in Poaceae (A) A sperm fertilizes an egg cell to produce diploid zygote which subsequently grows into diploid embryo; (B) A sperm fertilizes a center cell to produce triploid primary endosperm cell which subsequently grows into endosperm; (C) The embryo develops into diploid plant through in vitro culture or seed germination; (D) The endosperm develops into triploid plant via culturing in vitro; (E) Triploid formation by a 2n+n mating; (F) The autotriploid hybrid produced by crossing tetraploid (2n=4x=AAAA) and diploid (2n=2x=AA); (G) The allotriploid hybrid produced by crossing tetraploid (2n=4x=AAAA) and diploid (2n=2x=BB); (H) Formation of triploid by polyspermy

图2 2n配子的形成机制 (A) 与体细胞相比正常配子的染色体数目减半; (B) 通过胞质融合或多核体形成的四倍体花粉母细胞产生PRD型2n配子, 如鸭茅(Falistocco et al., 1995); (C) 在减数分裂I期同源染色体无配对和分裂形成的FDR型2n配子将保留除重组片段外的所有亲本基因, 如雀稗(Filho et al., 2014)和多年生黑麦草(Chen et al., 1997); (D) 在减数分裂II期因分裂缺失形成的SDR型2n配子将保留除重组片段外比正常配子多1套染色体副本的2n配子, 如多年生黑麦草(Chen et al., 1997); (E) 在减数分裂I期部分染色体以姐妹染色单体分离, 其余以二价体分离, 随后在减数分裂II期分裂缺失形成IMR型2n配子, 如百合(Lim et al., 2001); (F) 减数分裂正常而进入配子体发育阶段时基因组加倍形成PMD型2n配子, 如马铃薯(Bastiaanssen et al., 1998)

Figure 2 Mechanisms of 2n gamete formation (A) Chromosome numbers of normal gametes are halved compared with somatic cells; (B) PRD (pre-meiotic doubling) gametes are obtained as a result of the tetraploid pollen mother cells formed by cytomixis or syncytium, e.g., Dactylis glomerata (Falistocco et al., 1995); (C) In the FDR (first division restitution) type, pairing and division of homoeologous chromosomes do not occur during meiosis I, and the FDR 2n gametes maintain all of their parental genes except cross-over fragments, e.g., Paspalum jesuiticum (Filho et al., 2014) and Lolium perenne (Chen et al., 1997); (D) SDR (second division restitution) gametes with two copies of non-recombinant chromosomes are the result of the second division omission or cytokinesis abnormalities after normal anaphase II, e.g., L. perenne (Chen et al., 1997); (E) IMR (indeterminate meiotic restitution) gametes were discovered in Lilium longiflorum × Asiatic hybrid (Lim et al., 2001), in which some chromosomes are separated as univalents during meiosis I whereas the others are separated as bivalents, and subsequently the second division omission occurs during meiosis II; (F) PMD (post-meiotic doubling) gametes occurs due to the genome doubling of haploid spores during microgametogenesis, e.g., Solanum tuberosum (Bastiaanssen et al., 1998)

表1 禾本科三倍体品种

Table 1 Triploidy cultivars in Poaceae

母本	父本	品种名	用途	参考文献
狗牙根(Cynodon dactylon) 2n=4x=36	非洲狗牙根(C. transvaalensis) 2n=2x=18	Tifway、TifwayII、Midiron、TifEagle、Tifgreen、Tifdwarf、MS-Supreme、MS-Express、TifSport和Champion (2n=3x=27)	草坪草	Kamps et al., 2011
荻(Miscanthus sacchariflorus) 2n=4x=76	芒(M. sinensis) 2n=2x=38	Illinois和Freedom (2n=3x=57)	能源作物	Perera et al., 2015
玉米(Zea mays) 2n=2x=20	四倍体大刍草(Z. perennis) 2n=4x=40	玉草1号(2n=3x=30)	饲用作物	任勇等, 2007
指状摩擦禾(Tripsacum dactyloides) 2n=2x=36	指状摩擦禾(T. dactyloides) 2n=4x=72	Verl (2n=3x=54)	饲用作物	Springer et al., 2006
薏苡(Coix lacryma-jobi) 2n=2x=20	水生薏苡(C. aquatica) 2n=4x=40	丰牧88 (2n=3x=30)	饲用作物	孙福艾, 2017
象草(Pennisetum purpureum) 2n=4x=28	美洲狼尾草(P. americana) 2n=2x=14	热研4号(2n=3x=21)	饲用作物	刘国道等, 2012
美洲狼尾草(P. americana) 2n=2x=14	象草(P. purpureum) 2n=4x=28	闽牧6号和华南1号(2n=3x=21)	饲用作物	陈平等, 2004; 陈钟佃等, 2012

图3 禾本科三倍体的利用 (A) 三倍体作为品种直接利用; (B) 通过辐射、转基因或组织培养等技术改良三倍体品种; (C) 作为通往更高倍性的“三倍体桥”; (D) 转移异源基因; (E) 通过3x/2x合成非整倍体

Figure 3 Application of triploid in Poaceae (A) The direct use of triploid cultivar; (B) The improvement of triploid cultivar through techniques such as radiation, genetic modification, or tissue culture; (C) Use of “triploids bridge” for production of polyploids; (D) Alien genes transfer; (E) Development of aneuploidy by 3x/2x mating

参考文献 113

1	陈平, 吴秀峰, 席嘉宾, 梁红 (2004). 华南1号杂交狼尾草选育初报. 仲恺农业技术学院学报 17, 56-59.
2	陈钟佃, 黄勤楼, 黄秀声, 冯德庆, 钟珍梅 (2012). “闽牧6号”狼尾草的选育及田间种植技术. 家畜生态学报 33, 53-55.
3	程祝宽, 李欣, 于恒秀, 顾铭洪 (1996). 一套新的籼稻初级三体的选育和细胞学鉴定. 遗传学报 23, 363-371.
4	党江波, 宋琴, 李彩, 郭启高, 梁国鲁 (2018). 园艺植物中三倍体的应用现状及育种前景分析. 园艺学报 45, 1813-1830.
5	顾洪如, 杨运生, 白淑娟, 陈礼伟 (1992). 牧草新品种“宁牧26-2”狼尾草. 江苏农业科学 ( 4), 61-63.
6	黄群策, 孙敬三 (1999). 通过异倍性水稻间杂交获得同源三倍体植株. 植物学报 41, 741-746.
7	刘国道, 白昌军, 王东劲, 易克贤, 韦家少, 何华玄, 周家锁 (2002). 热研4号王草选育. 草地学报 10, 92-96.
8	刘秀明, 崔腾腾, 葛春霞, 陈翠霞 (2014). 我国野生芒资源的细胞学研究. 草业科学 31, 627-631.
9	罗宗志, 林洁荣, 罗虹建, 林志魁, 陈碧成 (2016). 杂交狼尾草和桂牧1号杂交象草的核型分析. 贵州农业科学 44, 8-12.
10	吕桂华, 唐祈林, 郭国锦, 陈坚剑, 荣廷昭 (2015). 玉米(Zea mays)×四倍体多年生玉米(Zea perennis)可育三倍体形态学和细胞遗传学研究. 植物遗传资源学报 16, 1152-1156.
11	潘志军, 易自力, 杨塞, 肖亮 (2017). 三倍体芒草自然杂交后代数量性状遗传多样性研究. 植物遗传资源学报 18, 984-990.
12	任勇, 陈柔屹, 唐祈林, 荣廷昭 (2007). 新型饲草玉米生长动态及收割期的研究. 作物学报 33, 1360-1365.
13	孙福艾 (2017). 一份薏苡多倍体材料的创制、鉴定及饲用价值初步评价. 硕士论文. 雅安: 四川农业大学. pp. 1-54.
14	孙敬三, 朱至清 (1981). 大麦胚乳植株的诱导及其倍性. 植物学报 23, 262-265.
15	田立忠, 徐爱菊 (2000). 高粱(Sorghum bicolor (L.) Moench)未成熟胚乳培养的研究. 辽宁师范大学学报(自然科学版) 23, 395-402.
16	王敬驹, 陆文梁, 匡柏健 (1982). 小黑麦杂种胚乳的离体培养研究. 植物学报 24, 420-425.
17	王润奇, 高俊华, 王志兴, 王志民 (1994). 谷子三体系列的建立. 植物学报 36, 690-695.
18	汪艳, 肖媛, 刘伟, 李婷婷, 胡锐, 乔志仙 (2015). 流式细胞仪检测高等植物细胞核DNA含量的方法. 植物科学学报 33, 126-131.
19	许蕾, 陈佩琳, 冯光燕, 钟旻依, 景婷婷, 黄琳凯, 张新全 (2019). 利用流式细胞仪鉴定鸭茅倍性. 草业学报 28, 74-84.
20	于卓, 云锦凤, 马有志, 辛志勇 (2004). 加拿大披碱草×野大麦三倍体杂种染色体的分子原位杂交鉴定. 遗传学报 31, 735-739.
21	袁金玲, 顾小平, 岳晋军, 吴晓丽 (2015). 一种毛竹胚乳培养的方法. 中国专利, ZL 201510566151.8. 2015-09-08.
22	张静, 吴先军, 汪旭东, 周开达, 彭海 (2002). 特异同源三倍体水稻材料SAR-3细胞学研究. 作物学报 28, 704-708.
23	张茜, 裘天航, 王安安, 周华健, 袁敏, 李利, 白素兰, 崔素霞 (2020). 北京地区芦苇资源状态及其多样性. 植物学报 55, 693-704.
24	张智奇, 钟维瑾, 唐克轩, 周音, 祝明福 (1994). 异源三倍体水稻原生质体培养及植株再生. 作物学报 20, 578-581.
25	赵世绪, 刘瑞凝, 敖光明, 陈一心 (1984). 小麦、黑麦未成熟胚乳植株的诱导. 北京农业大学学报 10, 129-132.
26	钟声 (2006). 鸭茅不同倍性杂交及后代发育特性的初步研究. 西南农业学报 19, 1034-1038.
27	钟小仙, 刘智微, 刘伟国, 崔莉莉, 吴娟子, 张建丽 (2014). 六倍体杂交狼尾草体细胞突变体特异性分析. 草业学报 23, 107-113.
28	祝剑峰, 刘幼琪, 王爱云, 宋兆建, 陈冬玲, 蔡得田 (2008). 异源六倍体水稻AACCDD和三倍体水稻ACD生殖特性的细胞胚胎学研究. 植物遗传资源学报 9, 350-357.
29	Bajaj YPS, Saini SS, Bidani M (1980). Production of triploid plants from the immature and mature endosperm cultures of rice. Theor Appl Genet 58, 17-18. DOI PMID
30	Bastiaanssen HJM, Van Den Berg PMMM, Lindhout P, Jacobsen E, Ramanna MS (1998). Postmeiotic restitution in 2 n-egg formation of diploid potato. Heredity 81, 20-27. DOI URL
31	Beale KM, Leydon AR, Johnson MA (2012). Gamete fusion is required to block multiple pollen tubes from entering an Arabidopsis ovule. Curr Biol 22, 1090-1094. DOI URL
32	Boller B, Kopeck D (2020). Triploid forage grass hybrids Festuca apennina × F. pratensis display extraordinary heterosis for yield characteristics. Euphytica 216, 143. DOI URL
33	Bretagnolle FO (2001). Pollen production and spontaneous polyploidization in diploid populations of Anthoxanthum alpinum. Biol J Linn Soc 72, 241-247. DOI URL
34	Brownfield L, Köhler C (2011). Unreduced gamete formation in plants: mechanisms and prospects. J Exp Bot 62, 1659-1668. DOI URL
35	Campos JMS, Davide LC, Salgado CC, Santos FC, Costa PN, Silva PS, Alves CCS, Viccini LF, Pereira AV (2009). In vitro induction of hexaploid plants from triploid hybrids of Pennisetum purpureum and Pennisetum glaucum. Plant Breed 128, 101-104. DOI URL
36	Chandra A, Genovesi AD, Wherley BG, Metz SP, Reinert JA, Wu YZ, Skulkaew P, Engelke MC, Hargey D, Nelson LR, Schwartz BM, Raymer PL, Wu YQ, Martin DL, Milla-Lewis SR, Miller G, Kenworthy KE, Munoz P (2015). Registration of ‘DALSA 0605’ St. Augustinegrass. J Plant Regist 9, 27-34. DOI URL
37	Chen C, Sleper DA, Chao S, Johal GS, West CP (1997). RFLP detection of 2 n pollen formation by first and second division restitution in perennial ryegrass. Crop Sci 37, 76-80. DOI URL
38	Costich DE, Friebe B, Sheehan MJ, Casler MD, Buckler ES (2010). Genome-size variation in switchgrass ( Panicum virgatum): flow cytometry and cytology reveal rampant aneuploidy. Plant Genome 3, 130-141.
39	De Storme N, Zamariola L, Mau M, Sharbel TF, Geelen D (2013). Volume-based pollen size analysis: an advanced method to assess somatic and gametophytic ploidy in flowering plants. Plant Reprod 26, 65-81. DOI PMID
40	Delomas TA (2019). Differentiating diploid and triploid individuals using single nucleotide polymorphisms genotyped by amplicon sequencing. Mol Ecol Resour 19, 1545-1551. DOI PMID
41	Dewald CL, Taliaferro CM, Dunfield PC (1992). Registration of four fertile triploid germplasm lines of eastern gamagrass. Crop Sci 32, 504. DOI URL
42	Duan QH, Liu MCJ, Kita D, Jordan SS, Yeh FLJ, Yvon R, Carpenter H, Federico AN, Garcia-Valencia LE, Eyles SJ, Wang CS, Wu HM, Cheung AY (2020). FERONIA controls pectin- and nitric oxide-mediated male-female interaction. Nature 579, 561-566. DOI URL
43	Dvorak J, Harvey BL, Coulman BE (1973). The use of nitrous oxide for producing eupolyploids and aneuploids in wheat and barley. Can J Genet Cytol 15, 205-214. DOI URL
44	Erichsen AW, Ross JG (1963). A triploid derived from a selfed haploid Sorghum plant. Crop Sci 3, 99-100. DOI URL
45	Falistocco E, Tosti N, Falcinelli M (1995). Cytomixis in pollen mother cells of diploid Dactylis, one of the origins of 2n gametes. J Hered 86, 448-453. DOI URL
46	Filho RAB, Santos ACC, Souza FHD, Valls JFM, Pagliarini MS (2014). Complete asynapsis resulting in 2n pollen formation in Paspalum jesuiticum Parodi (Poaceae). Genet Mol Res 13, 255-261. DOI URL
47	Gao LH, Diarso M, Zhang A, Zhang HK, Dong YZ, Liu LX, Lv ZL, Liu B (2016). Heritable alteration of DNA methylation induced by whole-chromosome aneuploidy in wheat. New Phytol 209, 364-375. DOI URL
48	Ghimire BK, Seong ES, Nguyen TX, Yoo JH, Yu CY, Kim SH, Chung I (2016). Assessment of morphological and phytochemical attributes in triploid and hexaploid plants of the bioenergy crop Miscanthus × giganteus. Ind Crop Prod 89, 231-243. DOI URL
49	Hagerup O (1947). The spontaneous formation of haploid, polyploid, and aneuploid embryos in some orchids. Biol Meddel Kongol Danske Vidensk Selsk 20, 1-22.
50	Hanna WW, Schertz KF (1971). Trisome identification in Sorghum bicolor( L.) Moench by observing progeny of triploid × translocation stocks. Can J Genet Cytol 13, 105-109. DOI URL
51	Harlan JR, DeWet JMJ (1975). On Ö. Winge and a prayer: the origins of polyploidy. Bot Rev 41, 361-390. DOI URL
52	Henry IM, Dilkes BP, Miller ES, Burkart-Waco D, Comai L (2010). Phenotypic consequences of aneuploidy in Arabidopsis thaliana. Genetics 186, 1231-1245. DOI
53	Hoshino Y, Miyashita T, Thomas TD (2011). In vitro culture of endosperm and its application in plant breeding: approaches to polyploidy breeding. Sci Hortic 130, 1-8. DOI URL
54	Hu FR, Zhang L, Wang XY, Ding J, Wu DX (2005). Agrobacterium-mediated transformed transgenic triploid bermudagrass ( Cynodon dactylon × C. transvaalensis) plants are highly resistant to the glufosinate herbicide liberty. Plant Cell Tissue Organ Cult 83, 13-19. DOI URL
55	Iqbal MZ, Cheng MJ, Zhao YL, Wen XD, Zhang P, Zhang L, Ali A, Rong TZ, Tang QL (2018). Mysterious meiotic behavior of autopolyploid and allopolyploid maize. Comp Cytogenet 12, 247-265. DOI URL
56	Iwata N, Omura T (1975). Studies on the trisomics in rice plants (Oryza sativa L.): III. Relation between trisomics and genetic linkage groups. Jpn J Breed 25, 363-368. DOI URL
57	Jansen RC, Den Nijs APM (1993). A statistical mixture model for estimating the proportion of unreduced pollen grains in perennial ryegrass (Lolium perenne L.) via the size of pollen grains. Euphytica 70, 205-215. DOI URL
58	Johansen B, Von Bothmer R (1994). Pollen size in Hordeum L.: correlation between size, ploidy level, and breeding system. Sex Plant Reprod 7, 259-263. DOI URL
59	Johri BM, Bhojwani SS (1965). Growth responses of mature endosperm in cultures. Nature 208, 1345-1347. DOI URL
60	Kamps TL, Williams NR, Ortega VM, Chamusco KC, Harris-Shultz K, Scully BT, Chase CD (2011). DNA polymorphisms at bermudagrass microsatellite loci and their use in genotype fingerprinting. Crop Sci 51, 1122-1131. DOI URL
61	Kapadia ZJ, Gould FW (1964). Biosystematic studies in the Bouteloua curtipendula complex. III. Pollen size as related to chromosome numbers. Am J Bot 51, 166-172. DOI URL
62	Kato A (1999). Induction of bicellular pollen by trifluralin treatment and occurrence of triploids and aneuploids after fertilization in maize. Genome 42, 154-157. DOI URL
63	Kato A, Birchler JA (2006). Induction of tetraploid derivatives of maize inbred lines by nitrous oxide gas treatment. J Hered 97, 39-44. DOI URL
64	Katsiotis A, Forsberg RA (1995). Pollen grain size in four ploidy levels of genus Avena. Euphytica 83, 103-108. DOI URL
65	Keller B, Feuillet C (2000). Colinearity and gene density in grass genomes. Trends Plant Sci 5, 246-251. PMID
66	Kindiger B, Dewald C (1994). Genome accumulation in eastern gamagrass, Tripsacum dactyloides(L.) L. 92, 197-201.
67	King IP, Morgan WG, Harper JA, Thomas HM (1999). Introgression mapping in the grasses. II. Meiotic analysis of the Lolium perenne/Festuca pratensis triploid hybrid. Heredity 82, 107-112. DOI URL
68	Kirov I, Divashuk M, Van Laere K, Soloviev A, Khrustaleva L (2014). An easy "SteamDrop" method for high quality plant chromosome preparation. Mol Cytogenet 7, 21. DOI PMID
69	Krans JV, Philley HW, Goatley JM Jr, Maddox VL (1999). Registration of ‘MS-Supreme’ bermudagrass. Crop Sci 39, 287.
70	Kreiner JM, Kron P, Husband BC (2017). Frequency and maintenance of unreduced gametes in natural plant populations: associations with reproductive mode, life history and genome size. New Phytol 214, 879-889. DOI URL
71	Kron P, Husband BC (2015). Distinguishing 2N gamete nuclei from doublets in pollen using flow cytometry and pulse analysis. Cytometry Part A 87, 943-957. DOI URL
72	La Rue CD (1949). Cultures of the endosperm of maize. Am J Bot 36, 798.
73	Lamote V, Baert J, Roldán-Ruiz I, De Loose M, Van Bockstaele E (2002). Tracing of 2n egg occurrence in perennial ryegrass (Lolium perenne L.) using interploidy crosses. Euphytica 123, 159-164. DOI URL
74	Lim KB, Ramanna MS, De Jong JH, Jacobsen E, Van Tuyl JM (2001). Indeterminate meiotic restitution (IMR): a novel type of meiotic nuclear restitution mechanism detected in interspecific lily hybrids by GISH. Theor Appl Genet 103, 219-230. DOI URL
75	Loginova DB, Silkova OG (2017). Mechanisms of unreduced gamete formation in flowering plants. Russ J Genet 53, 741-756. DOI URL
76	Lu SY, Peng XX, Guo ZF, Zhang GY, Wang ZC, Wang CY, Pang CS, Fan Z, Wang JH (2007). In vitro selection of salinity tolerant variants from triploid bermudagrass ( Cynodon transvaalensis × C. dactylon) and their physiological responses to salt and drought stress. Plant Cell Rep 26, 1413-1420. DOI URL
77	Maceira NO, De Haan AA, Lumaret R, Billon M, Delay J (1992). Production of 2 n gametes in diploid subspecies of Dactylis glomerata L. 1. Occurrence and frequency of 2n pollen. Ann Bot 69, 335-343. DOI URL
78	Mason AS, Nelson MN, Yan GJ, Cowling WA (2011). Production of viable male unreduced gametes in Brassica interspecific hybrids is genotype specific and stimulated by cold temperatures. BMC Plant Biol 11, 103. DOI URL
79	Multani DS, Jena KK, Brar DS, De Los Reyes BG, Angeles ER, Khush GS (1994). Development of monosomic alien addition lines and introgression of genes from Oryza australiensis Domin. to cultivated rice O. sativa L. Theor Appl Genet 88, 102-109.
80	Mutlu SS, Mutlu N, Tokgöz S, Çakır M, Selim C (2020). Development of vegetative triploid turf-type bermudagrass [ Cynodon dactylon × C. transvaalensis (C. × mangennisii Hurcombe)]. Genet Resour Crop Evol 67, 177-189. DOI URL
81	Naganowska B, Zwierzykowski Z, Zwierzykowska E (2001). Meiosis and fertility of reciprocal triploid hybrids of Lolium multiflorum with Festuca pratensis. J Appl Genet 42, 247-255. PMID
82	Nakano H, Tashiro T, Maeda E (1975). Plant differentiation in callus tissue induced from immature endosperm of Oryza sauva L. Z Pflanzenphysiol 76, 444-449. DOI URL
83	Nakel T, Tekleyohans DG, Mao YB, Fuchert G, Vo D, Groß-Hardt R (2017). Triparental plants provide direct evidence for polyspermy induced polyploidy. Nat Commun 8, 1033. DOI URL
84	Norstog K, Wall WE, Howland GP (1969). Cytological characteristics of ten-year-old rye-grass endosperm tissue cultures. Bot Gaz 130, 83-86. DOI URL
85	Norstog KJ (1956). Growth of rye-grass endosperm in vitro. Bot Gaz 117, 253-259. DOI URL
86	Nunes JD, Azevedo ALS, Pereira AV, Paula CMP, Campos JMS, Lédo FJS, Santos VB (2013). DNA elimination in embryogenic development of Pennisetum glaucum × Pennisetum purpureum(Poaceae) hybrids. Genet Mol Res 12, 4817-4826. DOI PMID
87	Pagliarini MS, Valle CB, Santos EM, Mendes DV, Bernardo ZH, Mendes-Bonato AB, Silva N, Calisto V (2012). Microsporogenesis in Brachiaria brizantha(Poaceae) as a selection tool for breeding. Genet Mol Res 11, 1309-1318. DOI PMID
88	Pécrix Y, Rallo G, Folzer H, Cigna M, Gudin S, Le Bris M (2011). Polyploidization mechanisms: temperature environment can induce diploid gamete formation in Rosa sp. J Exp Bot 62, 3587-3597. DOI URL
89	Pepin GW, Funk CR (1971). Intraspecific hybridization as a method of breeding Kentucky bluegrass ( Poa pratemis L.) for turf. Crop Sci 11, 445-448. DOI URL
90	Perera D, Barnes DJ, Baldwin BS, Reichert NA (2015). Direct and indirect in vitro regeneration of Miscanthus × giganteus cultivar freedom: effects of explant type and medium on regeneration efficiency. In Vitro Cell Dev Biol Plant 51, 294-302. DOI URL
91	Ramsey J, Schemske DW (1998). Pathways, mechanisms, and rates of polyploid formation in flowering plants. Annu Rev Ecol Syst 29, 467-501. DOI URL
92	Rhoades MM (1936). Note on the origin of triploidy in maize. J Genet 33, 355-357. DOI URL
93	Sandfaer J (1975). The occurrence of spontaneous triploids in different barley varieties. Hereditas 80, 149-153. DOI URL
94	Schwartz BM, Harris-Shultz KR, Contreras RN, Hans CS, Hanna WW, Milla-Lewis SR (2013). Creation of artificial triploid and tetraploid centipedegrass using colchicine and breeding. Int Turfgrass Soc Res J 12, 327-334.
95	Sehgal CB (1974). Growth of barley and wheat endosperm in cultures. Curr Sci 43, 38-40.
96	Sheidai M, Jafari S, Taleban P, Keshavarzi M (2009). Cytomixis and unreduced pollen grain formation in Alopecurus L. and Catbrosa Beauv.(Poaceae). Cytologia 74, 31-41. DOI URL
97	Sieber VK, Murray BG (1979). The cytology of the genus Alopecurus(Gramineae). Bot J Linn Soc 79, 343-355. DOI URL
98	Springer TL, Dewald CL, Sims PL, Gillen RL, Louthan VH, Cooper WJ, Taliaferro CM, Maura C, Pfaff S, Wynia RL, Douglas JL, Henry J, Bruckerhoff SB, Grinten M, Salon PR, Houck MJ, Esquivel RG (2006). Registration of 'Verl' eastern gamagrass. Crop Sci 46, 477. DOI URL
99	Straus J, LaRue CD (1954). Maize endosperm tissue grown in vitro I. Culture requirements. Am J Bot 41, 687-694. DOI URL
100	Suarez EY, Lopez AG, Naranjo CA (1992). Polyspermy versus unreduced male gametes as the origin of nonaploids (9x) common wheat plants. Caryologia 45, 21-28. DOI URL
101	Sun S, Wu Y, Lin XY, Wang J, Yu JM, Sun Y, Miao YL, Li QP, Sanguinet KA, Liu B (2017). Hybrid weakness in a rice interspecific hybrid is nitrogen-dependent, and accompanied by changes in gene expression at both total transcript level and parental allele partitioning. PLoS One 12, e0172919. DOI URL
102	Tamaoki T, Ullstrup JA (1958). Cultivation in vitro of excised endosperm and meristem tissues of corn. Bull Torrey Bot Club 85, 260-272. DOI URL
103	Tan GY, Dunn GM (1973). Relationship of stomatal length and frequency and pollen-grain diameter to ploidy level in Bromus inermis Leyss. Crop Sci 13, 332-334. DOI URL
104	Thomas H, Morgan WG, Humphreys MW (1988). The use of a triploid hybrid for introgression in Lolium species. Theor Appl Genet 76, 299-304. DOI PMID
105	Thomas TD, Chaturvedi R (2008). Endosperm culture: a novel method for triploid plant production. Plant Cell Tissue Organ Cult 93, 1-14. DOI URL
106	Toda E, Ohnishi Y, Okamoto T (2016). Development of polyspermic rice zygotes. Plant Physiol 171, 206-214. DOI URL
107	Toda E, Okamoto T (2016). Formation of triploid plants via possible polyspermy. Plant Signal Behav 11, e1218107. DOI URL
108	Van Santen E, Hugessen PM, Casler MD (1991). Identification and frequency of tetraploid progeny from 2 x-4x and 4x-2x crosses in Dactylis. Genome 34, 273-278. DOI URL
109	Wang C, Liu Q, Shen Y, Hua YF, Wang JJ, Lin JR, Wu MG, Sun TT, Cheng ZK, Mercier R, Wang KJ (2019). Clonal seeds from hybrid rice by simultaneous genome engineering of meiosis and fertilization genes. Nat Biotechnol 37, 283-286. DOI URL
110	Wang X, Yamada T, Kong FJ, Abe Y, Hoshino Y, Sato H, Takamizo T, Kanazawa A, Yamada T (2011). Establishment of an efficient in vitro culture and particle bombardment-mediated transformation systems in Miscanthus sinensis Anderss, a potential bioenergy crop. GCB Bioenergy 3, 322-332. DOI URL
111	Younis A, Hwang YJ, Lim KB (2014). Exploitation of induced 2 n-gametes for plant breeding. Plant Cell Rep 33, 215-223. DOI URL
112	Yu CY, Kim HS, Rayburn AL, Widholm JM, Juvik JA (2009). Chromosome doubling of the bioenergy crop, Miscanthus × giganteus. GCB Bioenergy 1, 404-412. DOI URL
113	Zeng RZ, Zhu J, Xu SY, Du GH, Guo HR, Chen JJ, Zhang SZ, Xie L (2020). Unreduced male gamete formation in Cymbidium and its use for developing sexual polyploid cultivars. Front Plant Sci 11, 558. DOI URL

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禾本科三倍体: 形成、鉴定与利用

Triploid in Poaceae: Formation, Detection, and Utilization

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