Chapter 5 Genetic Analysis of Apomixis - cimmyt

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94 D..iel Grimallelli-, Jo. Tohme, .od Di.go GN,OIt,-d...I.t••--. 19840. Gametophytic apomixis. In8.M. Johri (ed.), Embryology ofAngiosperms. Berlin: Springer-Verlag. Pp_47>-518.--. 1984b. Genetics 01 apospory inapomictic Ranunculus ouricamus V.Conclusions. Botanica Helvetica 94: 41122.Noirot, M. 1993. Allelic ratios and steri~ty in theagamic complex 01 the Maximeoe(Panicoideae): evolutionary role 01 theresidual sexuality.J. Evo/. BioI. 6:9S-101.Nayes, R.O., and R.H. Rieseberg. 2000. Twoindependent loci control agamospermy(apomixis) in the triploid flowering plantfIigeran annuus. Genetics 155: 379-90.Ohad, N., l. Margassian, Y. -C Hsu, CWilliams,p.Repe"i, and R_L Fisher. 1996. Amutationthat allows endosperm developmentdevelopment without fertiliza1ian. Proe.Natl. Acad. Sci. (USA) 93: 5319-24.Ozias-Akins, P. 1998. Tight clustering andhemizygosity of apomixis-linked molecularmarkers in Pennisetum squamulatumimplies genetic control of apospory by adivergent locus that may have no allelic Iform in sexual genotypes. Proe. Natl. Acdd.Sci. (USA) 95: 5127-32.Pessin a, S.C 1997.ldenliflcation of a maizelinkage group related 10 apomixis inBracharia. Theor. Appl. Genet. 94: 43944.Richards, AJ. 1986. Plant breeding systems.london: George Allen and Unwin.Savidan, Y. 1982. Nature et herMite de"apomixie chez Panicum maximum }acq.Ph.D_ dissertation Universite of Paris XI.---. 2000. Apomixis: Genelics and8reeding. Plant Breeding Reviews 18: 1386.Sheridan, W.F., A.A. Nadezhda, 1.1. Shamrov, lB.Batygina, and LN. Golunovskayo. 1996.The mac gene: controlling the commitment10 Ihe meiotic pathway in maize. Genetics142: 1009-20.Shizuya, H., B. 8irren, U.-1. Kim, V. Manano, 1Slepak, Y. Tachiiri, and M. Simon. 1992.Cloning ond stable maintenonce 01300kilobase-poir fragmenls 01 human DNA inEscherichia coli using a Hoctor-basedvector. Pro. Natl. Acod. Sci. (USA) 89:8794-97.van Dijk, PJ. 1999. Crosses between sexual andapomictic dandelions (Taraxacum). II. Thebreakdown of apomixis. Heredity 83:71S-21.Vielle-Calzoda J.P., J. Thomos, L Spillane, A.Coluccio, M.A. Hoeppner, and U.Grossniklaus. 1999. Maintenance ofgenomic imprinting at the Arabidopsismedea locus requires zygotic ODM1activity. Genes Dev. 13(22): 2971-82.Walbot, V. 1992. Strategies lor mutagenesis andgene cloning using transposon lagging andlDNA mutagenesis. Ann Rev. Planf Physiol.Plant Mol. BioI. 43: 49-82.
Chapter 7The Gene Effect:Genome Collisions and ApomixisJOHN G. CARMANIntroductionIn the vast majority of angiosperms, femalemeiosis results in the formation of a tetrad ofmononucleate megaspores, of which threedegenerate and one forms the geneticallyreduced8-nucleate female gametophyte(Polygonum-type embryo sac). Consistentlyexpressedcytological deviations from thisnorm occur in certain species in 506 of the13,479 genera of angiosperms recognized bythe Kew Botanical Gardens (Carman 1997).However, most species in most of these 506genera reproduce normally. Thus, thepercentage of species consistently expressingreproductive anomalies (probably < 0.3 %) isfar less than the percentage of genera (3.8 %)currently known to contain anomalous species.Reproductively-anomalous species occur in atleast 184 families, which is 53 % of those inwhich some embryological analyses have beenreported, and are much more abundant insome families than in others (Carman 1997).The reproductive anomalies considered in thischapter generally belong to three categories:gametophytic apomixis, polyspory, andpolyembryony. Gametophytic apomictsproduce unreduced embryo sacs that containparthenogenetic eggs, are generally polyploid,and occur in at least 33 of 460 families ofangiosperms. Diplospory and apospory aretwo major subdivisions of gametophyticapomixis (referred to as apomixis hereafter)and occur when unreduced embryo sacs formprecociously from ameiotic megaspore mothercells (MMCs) or nearby somatic cells,respectively (Asker and Jerling 1992; Carman1997; Peel et al. 1997a, b; Crane, Chap. 3).Polysporic species (bisporic or tetrasporic) aresexual and occur in at least 88 families. As indiplospory, embryo sacs of polysporic speciesform precociously from MMCs, but onlyportions of meiosis not critical to geneticreduction are affected (Battaglia 1989; Johri etal. 1992; Carman 1997). Polyembryonyinvolves the formation of embryos from cellsof other embryos, synergids, antipodals,nucelli, integuments, and even leaves (Tisseratet al. 1979; Johri et al. 1992). Like parthenogenesisin apomicts, polyembryony oftenbegins before pollination (Naumova 1993).This chapter summarizes these anomalies interms of developmental similarities,phylogenetic associations, and gene effecthypotheses, and it discusses implications of thegene effect hypotheses for future research andplant improvement. It concludes that a majorpurging of some widely accepted dogmaconcerning the evol ution and geneticregulation of apomixis will probably occur asthe mechanisms involved give way to moreaccurate evolutionary, developmental, andmolecular characterizations.Developmental Biology andPhylogeny of ReproductivelyAnomalous SpeciesAt the developmental level, some apomicticmechanisms resemble sexual polysporicmechanisms more than other apomictic
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Institut de Recherche pour Ie Devel
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iv33 Outlook33 References35 Appendi
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vi131 Screening Procedures: Advanta
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VIIITables4 Table 1.15 Table 1.29 T
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xAcknowledgmentsWe are grateful to
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xiifood crops such as maize, wheat,
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2 Gary H. T....i.....much food as i
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4 Gary H. Toe"'...breeding: the evo
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6 Gary H. T....it....The potential
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Chapter 2Apomixis and the Managemen
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10M.. Be,th.dcategories n + nand 2n
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12 J.&eo BertIoaodtwo tetraploid sp
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14 JoG.. B.rth""dTable 2.8 Distribu
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16 M......thodhexaploidy through 2n
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18 JI&.. lertIoaodheterozygous cond
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20 JM Iertltaodmarkers to retain th
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22 JoA•• BerthudFurthermore, if
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Chapter 3Classification of Apomicti
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26 (llarIesf.(r••3) The Ixeris-
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simultaneous division of the proxim
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30 (\aries f. Crao.differentiating
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32 CWIts F. C,..haploids from norma
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34 cales F. (,...Campbell, C. S., C
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36 GaOO F. er...media. There may al
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42 C"Ie
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Page 80 and 81: 66 Robe" T. SherwoodIn mitotic dipl
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150 (adlda Borge. da VaNe and Joh.
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152 Ccdda Borge. do v..... J.h W. M
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156 rn. s.mdaosearch for an opitimu
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158 Yv•• Savid..unreduced polar
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162 hOI Scrvidao211 = 56 chromosome
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164 r.., SaYid..Transfer of Gene(s)
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166 rves Sqyldaounanswered. However
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Chapter 12From Sexuality to Apomixi
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170 lie. Gros..iklalSgametes and em
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172 Uti Gro....1aosGunning 1990). T
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174 UeIGr.........embryogenesis, wh
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206 UeRG'....ild...--.1974. Reprodu
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21 0 Ud Gromild.1S--.1982. Nature e
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Chapter 13Induction of Apomixis inS
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progression or inhibition of develo
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