| 研究生: |
方莞婷 Fang, Wan-Ting |
|---|---|
| 論文名稱: |
黍族 ( 禾本科 ) 植物無融合生殖之篩選方法 Screening methods for apomixis in the Paniceae ( Poaceae ) |
| 指導教授: |
郭長生
Kuoh, Chang-Sheng |
| 學位類別: |
碩士 Master |
| 系所名稱: |
生物科學與科技學院 - 生物學系 Department of Biology |
| 論文出版年: | 2003 |
| 畢業學年度: | 91 |
| 語文別: | 中文 |
| 論文頁數: | 81 |
| 中文關鍵詞: | 有性生殖 、禾本科 、黍族 、篩選 、無融合生殖 |
| 外文關鍵詞: | sexual reproduction, Paniceae, apomixis, Poaceae, screen, apospory |
| 相關次數: | 點閱:100 下載:5 |
| 分享至: |
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多數開花植物經由雙重受精行有性生殖,而後產生種子繁衍後代。然而自然界有些物種是以無融合生殖方式繁衍後代,即無性產生種子。無融合生殖能迅速穩定雜種優勢並降低雜交種子的生產成本,可望在21世紀應用無融合生殖於農業和經濟作物改良。在進行無融合生殖之育種及分子層次之各項研究前,篩選無融合生殖物種為首要工作。
本研究利用細胞學透明化觀察法、組織化學切片觀察法與流式細胞種子篩選法篩檢禾本科黍亞科黍族七種不同植物之生殖方式,建立黍族有性生殖與無融合生殖性狀之模式與篩選方法。結果顯示紫果馬唐、圓果雀稗以及百喜草具有蓼型胚囊,經雙重受精形成胚及胚乳,屬有性生殖。大黍、牧地狼尾草、紅毛草以及蒺藜草屬無融合生殖。其中大黍、牧地狼尾草與紅毛草為典型未減數四核黍型胚囊,具有多胚囊現象,胚係由孤雌生殖得來,胚乳則由假受精產生,屬無融合生殖之無孢子生殖。另外,蒺藜草則具有未減數八核山柳菊型胚囊,胚係由孤雌生殖得來,胚乳則自發性產生,屬無融合生殖之無孢子生殖。
由上述可知,利用細胞學觀察法篩檢黍族無融合生殖性狀,以往主要以四核無反足細胞胚囊或多胚囊為主要辨別特徵,但本研究發現蒺藜草為例外之情形,所以在篩選無融合生殖時,除形態特徵外,仍需配合流式細胞種子篩選法。
Most flowering plants reproduce sexually by seeds via double fertilization. However, some plants reproduce asexually by apomixis, a natural way of cloning through seeds. In the crop breeding field, apomixis can fix the genotype of a superior plant and would enable a significant reduction in the cost of hybrid seed production. Therefore, the application of apomixis also holds great promise as an essential tool for sustainable agricultural production and economic crops improvement in the 21 century. In order to research on apomixis at the breeding and molecular levels, screening apomictic plants would be the primary work in advance.
Different reproduction modes of seven species belonging to Paniceae were comparatively studied by histological embryo sac examination and flow cytometry seeds screening. We find that these combined methods are efficient to discriminate apomixis from sexual reproduction. This research indicated that the embryo sacs of Digitaria violascens, Paspalum orbiculare and Paspalum notatum are Polygonum type and reproduce sexually via double fertilization. On the other hand, the embryo sacs of Panicum maximum, Pennisetum polystachion and Rhynchelytrum repens reveal Panicum type. These three latter species show apospory followed by parthenogenesis and pseudogamy. In addition, Cenchrus echinatus is autonomous aposporous apomicts, forming both embryo and endosperm spontaneously inside an unreduced embryo sac of Hieracium type.
Apomixis reproduction modes of Paniceae were screened by general histological embryo sac examination. In which, the 4-nuclear, multiple embryo sacs and absence of antipodal cells are the most obvious characters. However, Cenchrus echinatus does not follow this rule. It is unique in this study. Besides the histological embryo sac examination, FCSS ( flow cytometry seeds screening ) also proved an efficient means to discriminate between the various reproductive routes by reconstruction of the reproductive pathways for seeds.
胡適宜. 1990. 被子植物胚胎學. 曉園出版社.
羅妙琦. 1997. 狼尾草屬無融合生殖性狀之轉移與傳遞. 台灣大學農藝學研究所博士論文.
Arumuganathan K. and Earle E.D. 1991. Estimation of nuclear DNA content of plants by flow cytometry. Plant Mol. Biol. Rptr. 9: 229-233.
Asker S. 1970a. Apomixis and sexuality in the Potentilla argentea complex.Ⅰ. Crosses with other species. Hereditas 66: 127.
Asker S. 1970b. Apomixis and sexuality in the Potentilla argentea complex. Ⅱ. Crosses within the complex. Hereditas 66: 189.
Asker S. 1971. Apomixis and sexuality in the Potentilla argentea complex.Ⅲ. Euploid and aneuploid derivates ( including trisomics ) of some apomictic biotypes. Hereditas 67: 111.
Asker S. 1980. Gametophytic apomixis : elements and genetic regulation. Hereditas 93: 277-293.
Asker S.E. and Jerling L. 1992. Apomixis in Plants. Boca Raton: CRC Press.
Baarlen P. 2001. Apomixis in Taraxacum : an embryological and genetic study. PhD thesis. The Graduate School 'Experimental Plant Sciences' (EPS).
Baarlen P., Jong H. and Dijk P.J. 2002. Comparative cyto-embryological investigations of sexual and apomictic dandelions ( Taraxacum) and their apomictic hybrids. Sex. Plant Reprod. 15: 31–38.
Bantin Jörg., Matzk F. and Dresselhaus T. 2001. Tripsacum dactyloides (Poaceae) : a natural model system to study parthenogenesis. Sex. Plant Reprod. 14: 219–226.
Bashaw E.C. 1962. Apomixis and sexuality in buffelgrass. Crop Sci. 2: 412-415.
Bashaw E.C. 1980a. Apomixis and its application in crop improvement. In Hybridization of Crop Plants. Fehr W.R. and Hadley H.H., eds. Amer. Soc. Agron., Madison, WI. p.45-68.
Bashaw E.C. and Hanna W.W. 1990. Apomictic reproduction. In Reproductive Versatility in the Grasses. G. P. Chapman, ed. Cambridge University Press. P: 100-130.
Bicknell R.A., Borst N.K. and Koltunow A.M. 2000. Monogenic inheritance of apomixis in two Hieracium species with distinct developmental mechanisms. Heredity: 84.
Brown W.V. and Emery W.H.P. 1958. Apomixis in the Graminiae: Panicoideae. Am. J. Bot. 45: 253-263.
Burson B.L. 1975. Cytology of some apomictic Paspalum species. Crop Sci. 15: 229-232.
Burson B.L. 1994. Cytological mechanisms of apospory and diplospory. In Apomixis: Exploiting hybrid vigor in rice. Khush G. S, ed. IRRI Manila. P: 35-38.
Burton G.W. 1999. Efforts to Create Apomictic Diploids of Paspalum notatum var. Parodi. Apomixis Newsl. 11: 14.
Caceres M.E., Matzk F., Busti A., Pupilli F. and Ssergio A. 2001. Apomixis and sexuality in Paspalum simplex : characterization of the mode of reproduction in segregating progenies by different methods. Sex. Plant Reprod. 14: 201-206.
Caceres M.E., Pupilli F., Quarin C.L. and Arcioni S. 1999. Feulgen densitometry of embryo sacs permits discrimination between sexual and apomictic plants in Paspalum simplex. Euphytica 110: 161-167.
Caponio I. and Quarin C.L. 1987. EI sistema genetico de Paspalum simplex de un hibrido interespecifico con Paspalum dilatatum. Kurtziana 19: 35-45.
Carman J.G., Orane C.F. and Riera-Lizarazu O. 1991. Comparative histology of cell walls during meiotic and apomeiotic megasporogenesis in two hexaploid Australasian Elymus species. Crop Sci. 31: 1527-1532.
Casa A.M., Mitchell S.E., Lopes C.R. and Valls J.F.M. 2002. RAPD analysis reveals genetic variability among sexual and apomictic Paspalum dilatatum Poiret biotypes. J. Hered. 93(4): 300-302.
Chao C.Y. 1971. A periodic acid-schiff’s substance related to the directional growth of pollen tube into embryo sac in paspalum ovules. Amer. J. Bot. 58(7): 649-654.
Chapman G.P. and Busri N. 1994. Apomixis in Pennisetum : an ultrastructural study. Int. J. Plant Sci. 155(5) : 492-497.
Chatterji H.K. and Timothy D.H. 1969. Apomixis and tetraploidy in Pennisetum orientale Rich. Crop Sci. 9: 796.
Chaudhury A.M., Ming L., Miller C., Craig S., Dennis E.S. and Peacock W.J. 1997. Fertilization-independent seed development in Arabidopsis thaliana. Proc. Natl. Acad. Sci. USA 94: 4223-4228.
Chen L.Z. and Kozono T. 1994. Cytology and quantitative analysis of aposporous embryo development in guineagrass (Panicum maximum Jacq.) Cytologia 59: 253-260.
Christoff M. and Christoff M.A. 1948. Meiosis in the somatic tissue responsible for the reduction of chromosome number in the progeny of Hieracium hoppeanum Schult. Genetics 33: 36.
Crane C. 2001. Classification of Apomictic Mechanisms. In: Savidan Y., Carman J.G., Dresselhaus T., eds. The Flowering of Apomixis: from Mechanisms to Genetic Engineering. Mexico, DF: CIMMYT, IRD, European Commission DG VI (FAIR).
Darlington C.D. 1939. The Evolution of Genetic Systems. Cambridge University Press, London.
Dijk P.J. and Damme J. 2000. Apomixis technology and the paradox of sex. Trends Plant Sci. Vol. 5, No. 2.
Dijk P.J., Tas I.C.Q., Falque M. and Bakx-Schotman J.M.T. 1999. Crosses between sexual and apomictic dandelions (Taraxacum) II. The breakdown of apomixis. Heredity 83:715–721.
Dresselhaus T. and Colombo L. 2001. 2nd International Apomixis Conference (APO2001) at Como, Italy, 24–28 April 2001. Sex. Plant Reprod. 14:245–251.
Dujardin M. and Hanna W.W. 1984b. Microsporogenesis, reproductive behavior, and fertility in five Pennisetum species. Theor. Appl. Genet. 67: 197-201.
Dujardin M. and Hanna W.W. 1989a. Crossability of pearl millet with wild Pennisetum species. Crop Sci. 29: 77-80.
Elizangela R. Alves, Vera T.C. Carneiro and Ana C.G. Araujo. 2001. Direct evidence of pseudogamy in apomictic Brachiaria brizantha (Poaceae). Sex. Plant Reprod. 14: 207–212.
Emidio A., Andrea P., Francesco F., Lara R., Gianni B., Bruno R. and Mario F. 2001. Apospory and parthenogenesis may be uncoupled in Poa pratensis: a cytological investigation. Sex. Plant Reprod. 14: 213–217.
García R., Asíns M.J., Forner J. and Carbonell E.A. 1999. Genetic analysis of apomixis in Citrus and Poncirus by molecular markers. Theor. Appl. Genet. 99: 511–518.
Grimanelli D., Leblanc O., Espinosa E., Perotti E., Leon D.G. and Savidan Y. 1998. Mapping diplosporus apomixis in tetraploid Tripsacum: one gene or several genes? Heredity 80: 33–39.
Grimanelli D., Leblanc O., Perotti E. and Grossniklaus, U. 2001. Developmental genetics of gametophytic apomixis. Trends Genet. 17:597–604.
Grossniklaus U., Koltunow A. and Lookeren C.M. 1998a. A bright future for apomixis. Trends Plant Sci. 3: 415–416.
Grossniklaus U., Nogler G. and Dijk P. 2001b. How to avoid sex: the genetic control of gametophytic apomixis. Plant Cell 13: 1491–1498.
Gupta F., Shivanna K.R. and Mohan R.H.Y. 1996. Apomixis and polyembrony in Guggul Plant, Commiphora wightii. Annals of Bot. 78: 67-72.
Guptafsson A. 1943. The genesis of the European blackberry flora, Lunds Unvi. Arsskr. N. F. Avd. 2. 39: 1.
Hanna W.W. 1995. Use of apomixis in cultivar development. Adv. Agron. 54: 333-350.
Hanna W.W. and Bashaw E.C. 1987. Apomixis: its identification and use in plant breeding. Crop Sci. 27: 1136-1139.
Hanna W.W., Dujardin M., Ozias-Akins P., Lubbers E. and Arthur L. 1993. Reproduction, cytology and fertility of pearl millet ╳ Pennisetum squamulatum BC4 plants. J. Hered. 84 (3): 213-216.
Hanna W.W. and Powell J.B. 1973. Stubby head, and induced facultative apomict in pearl millet. Crop Sci. 13: 726-728.
Harlan J.R., Brooks H., Borgaonkar D.S. and Wet J.M.J. 1964. Nature and inheritance of apomixis in Bothriochloa and Dichanthium. Bot. Gaz. ( Chicago ) 125: 41-46.
Herr J.M. 1971. A new clearing-squash technique for the study of ovule development in angiosperms. Am. J. Bot. 58: 785-790.
Iwamasa M., Ueno I. and Nishiura M. 1967. Inheritance of nucellar embryony in Citrus. Bull Hortic Res Stn Jpn, Ser B 7:1–8.
Jongedijk E. 1985. The pattern of megasporogenesis and megagametogenesis in diploid Solanum species hybrids; its relevance to the origin of 2n-eggs and the induction of apomixis. Euphytica 34: 599.
Khush G.S., Brar D.S., Bennett J. and Virmani S.S. 1994. Apomixis for rice improvement in Apomixis : Exploiting Hybrid Vigor in Rice, Khush G.S., ed. IRRI Manila. P: 1-2.
Kindinger B., Bai D. and Sokolov V. 1996. Assignment of a gene(s) conferring apomixis in Tripsacum to a chromosome arm: cytological and molecular evidence. Genome 39: 1133-1141.
Kitano H., Tamura Y., Satoh H. and Nagato Y. 1993. Hierarchical regulation of organ differentiation during embryogenesis in rice. Plant J. 3: 607-610.
Knox R.B. 1967. Apomixis : Seasonal and population difference in a grass. Science 157: 325-326.
Koltunow A.M. 1993. Apomixis: Embryo sacs and embryos formed without meiosis or fertilization in ovules. Plant Cell 5: 1425-1437.
Koltunow A.M., Bicknell R.A. and Chaudhury A.M. 1995. Apomixis : molecular strategies for the generation of genetically identical seeds without fertilization. Plant Physiol. 108: 1345-1352.
Koltunow A.M. and Grossniklaus U. 2003. APOMIXIS: A developmental perspective. Annu. Rev. Plant Biol. 54: 547–74.
Koltunow A.M., Hidaka T. and Robinson S.P. 1996. Polyembryony in Citrus. Accumulation of seed storage proteins in seeds and in embryos cultured in vitro. Plant Physiol. 110: 599–609.
Koltunow A.M., Johason S.D. and Bicknell R.A. 2000. Apomixis is not developmentally conserved in related, genetically characterized Hieracium plants of varying ploidy. Sex. Plant Reprod. 12: 253–266.
Koltunow A.M., Johason S.D. and Bicknell R.A. 1998. Sexual and apomictic development in Hieracium. Sex. Plant Reprod. 11: 213-230.
Leblanc O., Grimanelli D., Islam-Faridi N., Berthaud J. and Savidan Y. 1996. Reproductive behavior in maize-Tripsacum polyhaplid plants : implication for the transfer of apomixis into maize. J. Hered. 87 : 108-111.
Leblanc O., Grimanelli D., Leon D.G. and Savidan Y. 1995a. Detection of apomictic mode of reproduction in maize-Tripsacum hybrids using maize RFLP markers. Theor. Appl. Genet. 90: 1198-1203.
Leblanc O., Peel M.D., Carman J.G. and Savidan Y. 1995b. Magasporogenesis and megagametogenesis in several Tripsacum species ( Poaceae ). Am. J. Bot. 82(1): 57-63.
Luo M., Bilodeau P., Koltunow A., Dennis E.S., Peacock W.J. and Chaudhury A.M. 1999. Genes controlling fertilization-independent seed development in Arabidopsis thaliana. Proc. Natl. Acad. Sci. USA 98: 296–301.
Lutts S., Ndikumana J. and Louant B.P. 1994. Male and female sporogenesis and gametogenesis in apomictic Brachiaria brizantha, Brachiaria decumbens and F1 hybrids with sexual colchicines induced tetraploid Brachiaria ruziziensis. Euphytica 78: 19–25.
Matzk F., Armin M. and Ingo S. 2000. An efficient screen for reproductive pathways using mature seeds of monocots and dicots. The Plant Journal 21 (1): 97-108.
Matzk F., Meyer H.M., Horstmann C., Balzer H.J., Bäumlein H. and Schubert I. 1997. A specific alpha-tubulin is associated with the initiation of parthenogenesis in ‘Salmon’ wheat lines. Hereditas 126: 219-224.
Mogie M. 1988. A model for the evolution and control of generative apomixis. Biol. J. Linn. Soc. 35: 127-153.
Mogie M. 1992. The Evolution of Asexual Reproduction in Plants. Chapman and Hall. London.
Muntzing A. 1940. Further studies on apomixis and sexuality in Poa. Hereditas 26: 115.
Muntzing A. 1958. Hereploidy and polymorphism in some apomict species of Potentilla. Hereditas 44: 280.
Muntzing A. and Muntzing G. 1943. Spontaneous changes in chromosome number in apomictic Potentilla collina. Hereditas 29: 451.
Naumova T.N. 1993. Apomixis in Angiosperm : Nucellar and Integumentary Ebmryony. CRC . Floria, U.S.A.
Naumova T.N., Hayward M.D. and Wagenvoort M. 1999. Apomixis and sexuality in diploid and tetraploid accessions of Brachiaria decumbens. Sex. Plant Reprod. 12: 43-52.
Naumova T.N., Laak J., Osadtchiy J., Matzk F., Kravtchenko A., Bergervoet J., Ramulu K.S. and Boutilier K. 2001. Reproductive development in apomictic populations of Arabis holboellii (Brassicaceae). Sex. Plant Reprod. 14:195–200.
Nijs A.P.M. and Dijk G.E. 1994. In Plant Breeding: Principles and Prospects. M. Hayward, N. Bosemark, I. Romagosa,ed. Chapman & Hall. London. P: 229-242.
Nogler G.A. 1984. Gametophytic apomixis. In: Johri BM, ed. Embryology of Angiosperms, Springer, Berlin, Heidelberg, New York.
Nogler G.A. 1995. Genetics of apomixis in Ranunculus aurcomus. VI. Epilogue. Bot. Helv. 105:111–115.
Norrmann G.A., Quarin C.L. and Burson B.L. 1989. Cytogenetics and reproductive behavior of different chromosome races in six Paspalum species. J. Hered. 80: 24-28.
Noyes R.D. and Rieseberg L.H. 2000. Two independent loci control agamospermy ( apomixis ) in the triploid flowering plant Erigeron annuus. Genetics 155: 379-390.
Ohad N., Margossian L., Hsu Y.C., Williams C., Repetti P. and Fischer R.L. 1996. A mutation that allows endosperm development without fertilization. Proc. Natl. Acad. Sci. USA 93: 5319-5324.
Ohad N., Yadegari R., Margossian L., Hannon M., Michael D., Harada J.J., Goldberg R.B. and Fischer R.L. 1999. Mutations in FIE a WD polycomb group gene, allow endosperm development without fertilization. Plant Cell 11: 407–415.
Ozias-Akins P., Lubbers E.L., Hanna W.W. and Mcnay J.W. 1993. Transmission of the trait and molecular makers. Theor. Appl. Genet. 85: 632-638.
Ozias-Akins P., Roche D. and Hanna W.W. 1998. Tight clustering and hemizygosity of apomixis-linked markers in Pennisetum squamulatum implies genetic control of apospory by a divergent locus that may have no allelic form in sexual genotypes. Proc. Natl. Acad. Sci. 95: 5127-5132.
Parlevliet J.E. and Cameron J.W. 1959. Evidence on the inheritance of nucellar embryony in Citrus. Proc. Am. Soc. Hortic. Sci. 74: 252–260.
Powers L. 1945. Fertilization without reduction in guayule ( Parthetium argentatum Gray ) and a hypothesis as to the evolution of apomixis and polyploidy. Genetics 30: 323.
Pullaiah T. and Febulaus G.N.V. 2000. Embryology and Apomixis in Grasses. Regency Pubilications. Old Market, West Patel Nagar, New Delhi.
Pupilli F., Caceres M.E., Quarin C.L. and Arcioni S. 1997. Segregation analysis of RFLP markers reveals a tetrasomic inheritance in apomictic Paspalum simplex. Genome 40: 822–828.
Pupilli F., Labombarda P., Cáceres M.E., Quarín C.L. and Arcioni S. 2001. The chromosome segment related to apomixis in Paspalum simplex is homoeologous to the telomeric region of the long arm of rice chromosome 12. Mol. Breed. 8: 53-61.
Quarin C.L., Espinoza F., Martinez E.J., Pessino S.C. and Bovo O.A. 2001. A rise of ploidy level induces the expression of apomixes in Paspalum notatum. Sex. Plant Reprod. 13: 243-249.
Ramulu K.S., Dijkhuis P., Rutgers E., Blaas J., Krens F.A., Verbeek W.H.J., Colijn-Hooymans and Verhoeven H.A. 1996. Intergeneric transfer of a partial genome and direct production of monosomic addition plants by microprotoplast fusion. Theor. Appl. Genet. 92: 316-325.
Reiser L. and Fischer R.L. 1993. The ovule and the embryo sac. The Plant Cell 5: 1291-1301.
Richards A.J. 1986. Agamospermy in Plant Breeding Systems. Allen and Unvwin, London. P: 403-456.
Savidan Y. 1980. Chromosomal and embryological analysis in sexual ╳ apomictic hybrids of Panicum maximum Jacp. Theor. Appl. Genet. 57: 153.
Savidan Y. 1982. Nature et heredite de I ‘ apomixis chez Panicum maximum Jacq. PhD thesis Universite of Paris.
Savidan Y. 1990. The genetic control of apomixis. Apomixis Newsl 2: 24.
Savidan Y. 2000. Apomixis: genetics and breeding. Plant Breed. Rev. 18: 13–86.
Savidan Y. and Pernès J. 1982. Diploid-tetraploid-diploid cycles and the evolution of Panicum maximum Jacq. Evolution 36: 596–600.
Schmelzer G. 1998. Pennisetum section Brevivalvula: morphological and genetic variation in an agamic species complex. WAU dissertation no. 2488.
Sherwood R.T., Berg C.C. and Young B.A. 1994. Inheritance of apospory in buffelgrass. Crop Sci. 34: 1490-1494.
Sherwood R.T., Young B.A. and Bashaw E.C. 1980. Facultative apomixis in buffelgrass. Crop Sci. 20: 375-379.
Sorensen T. 1958. Sexual chromosome-aberrants in triploid apomictic Taraxaca. Bot. Tidsskr. 54: 1.
Sorensen T. and Gudjonsson G. 1946. Spontaneous chromosome-aberrants in apomictic Taraxaca. K. Dan. Vidensk. Selskab. Biol. Skr. 4: 1.
Spillane C., Vielle-Calzada J.P. and Grossniklaus U. 2001. APO2001 : A sexy apomixer in Como. The Plant Cell 13 : 1480-1491.
Spillane C., Steimer A. and Grossniklaus U. 2001. Apomixis in agriculture: the quest for clonal seeds. Sex. Plant Reprod. 14: 179–187.
Taliaferro C.M. and Bashaw E.C. 1966. Inheritance and control of obligate apomixis in breeding buffelgrass, Pennisetum cilliare. Crop Sci. 6: 473-476.
Tiantang N., Yi S. and Zhiliang B. 1987. Stimulating Sorghum apomixis through distant pollination and hormone treatment. Acta Agric. Boreali. Sinica. 2: 1.
Tucker M.R., Paech N.A., Willemse M.T.M. and Koltunow A.M.G. 2001. Dynamics of callose deposition and β-1,3-glucanase expression during reproductive events in sexual and apomictic Hieracium. Planta Vol. 212: 487-498.
Tzvelev N.N. 1989. The systems of Grasses ( Poaceae ) and their evolution. Bot. Rev. 55: 141-203.
Vielle-Calzada J.P., Crane C.F. and Stelly D.M. 1996a. Apomixis: The asexual revolution. Science 274: 1322-1323.
Warmke H.E. 1954. Apomixis in Panicum maximum. Am. J. Bot. 41: 5–11.
Young B.A., Shewood R.T. and Bashaw E.C. 1979. Cleared-pistil and thick – sectioning techniques for detecting aposporous apomixes in grasses. Can. J. Bot. 57: 1668-1672.
Zhao Z.Y. and Gu M.G. 1984. Production of diploid pure lines of maize by chemically induced parthenogenesis. Acta Genet. Sinica. 11: 39.
Zhao Z.Y. and Gu M.G. 1988. Cytogenetic studies of chemically-induced parthenogenetic maize plants. Acta Genet. Sinica. 15: 89.