62 Tamara N. Nouma.o and Jeon-Phmppe '(..lIe-Cal.adoThe comparison <strong>of</strong> early fertilization events inPellllisetllnJ ciliare has identified ultrastructuraldifferences related to the control <strong>of</strong> egg cellparthenogenesis, an essential event in theapomictic life cycle that is absent from sexualreprod uction; however, the dynamics <strong>of</strong> pollentube arrival and sperm cell delivery have notbeen investigated. Further ultrastructuralstudies will be necessary to determine if spermcell delivery and movement are equivalent insexual and aposporous female gametophytes.The nature <strong>of</strong> the signal that activates theaposporous egg cell and the fate <strong>of</strong> the spermcells after pollen tube delivery remain to beelucidated. Levels <strong>of</strong> calcium and otherelements have never been measured insynergids <strong>of</strong> apomictic embryo sacs (Chaubaland Reger 1992). Additional studies shouldinclude three-dimensional reconstruction <strong>of</strong>egg cells during the first hours followingpollination and quantitative information onthe pattern <strong>of</strong> synergid degeneration and onthe specific fate <strong>of</strong> sperm cells within themegagametophyte. Attempts to follow themovement <strong>of</strong> fluorochromatically-stainedsperm cells in sexual embryo sacs have beenlimited, and the reliability <strong>of</strong> such studies usingconventional clearing techniques isquestionable; however, in vitro analysis <strong>of</strong>sperm nuclei movement within isolatedembryo sacs has been accomplished (Faure etal. 1994). Isolation and fusion <strong>of</strong> gametes cannow be used to compare the interaction <strong>of</strong>sperm cells with sexual or aposporous eggcells, in order to determine if cell to cellinteractions impede the fusion <strong>of</strong> sperm nucleiwith an ilpomictically-derived egg cell. Finally,some \'ery enlightening findings may bel'btilined in the near future by following onthe pioneering work <strong>of</strong> Olga Erdelska(Skwakian Academy <strong>of</strong> Sciences, Bratislilvil).Recent investigations in Torellia jOllmieri haveshown that, at least in certain species in whichthe megagametophyte can be readily dissectedor is partially uncovered by the integuments,some unique discoveries can be made byobserving the process <strong>of</strong> double fertilizationin vivo (Higashiyama et at. 2000). Microinjectionand cell ablation technology maysoon be combined with microcinerr:atographyto investigate sexual and apomictic megagametophytedevelopment in vivo.AcknowledgmentsWe thank A. Vassilyev and U. Grossniklausfor their critical reading <strong>of</strong> the manuscript andstimulating discussions, and J. Osadtchiy fortechnical assistance. Work by TamaraN'lUmova. was partially supported by a grantfrom the lAC (the Netherlands) and byfinancial support from the InternationalScience Foundation (USA).ReferencesAsker, 5.E., and l. Jerling. 1992. <strong>Apomixis</strong> in Plonts. Boca Rotan, Florida:CRC Press.Bashaw, H., and E.C. Holt. 1958. Megasporogenesis, embryo socdevelopment and embryogenesis in dallisgrass, Paspalum dilatatum.Agran.1. 50: 753-56.Bell, P.R. 1988. Control points in generation cycle <strong>of</strong> ferns. 1. Plant Physiol.132: 378.--. 1992. Apospory and apogamy: implications for understandingthe plant life cycle. Int. 1. Plant Sci. 153: 123-36.---.1995. Incompatibility in flowering plants: adaptation <strong>of</strong> onancient response. The Plont (ell 7: S--16.Burson, B.l., PW. Voigt, B.A. Sherman, and c.L. Dewald. 1990. <strong>Apomixis</strong>and sexuality in eastern gamagrass. (rap. Sci. 30: 81>-89.Campbell, 5.C, WG. Craig, and E.B. ScaN. 1987. <strong>Apomixis</strong> ond sexuality inthree species <strong>of</strong> Amelanchier, shodbush (Rosaceae, Moloideael. Am.1. Bal. 74: 321-28.Corman, J.G., C.F. Crone, and O. Riera-lizarazu. 1991. Comparativehislology <strong>of</strong> cell walls during meiosis and apomicticmegasporogenesis in !wo hexaploid Australasian Elymus species.(rop Sri. 31: '527-32.Corman, J.G. 1997. Asynchronous expression <strong>of</strong> duplicate genes inangiosperms may couse apomixis, bispory, tetraspory, andpolyembryony. Bioi. 1. linn. Soc 61: 51-94.Chapman, G.P, and N. 8usri. 1994. <strong>Apomixis</strong> in Pennisefum: onultrastructural study. International 1. <strong>of</strong> Plonf Sciences 155: 492-97.Chaubal, R., and BJ Reger. 1992. Calcium in the synergid cells and otherregions <strong>of</strong> pearl millet ovaries. Sex. Plant. Reprod. 5: 34-46.Crone, CF., and J.G. Carmon 1987. Mechanisms 01 apomixis in ElymusreC1isefus from eastern Australia and New Zealand. Ame~ 1. Bot. 74:477-96.Diboll, A.G 1968. Fine structural development <strong>of</strong> the megagametophyte01 lea mays follOWing fertilization. Am 1. Bal. 55: 787-806.
Ultraslrvdural <strong>Analysis</strong> <strong>of</strong> Apomktit Deveiopmellt 63Dickinson, H.G., ond U. Potter. 1978. Cytoplosmkchanges accompanying the female mei~isin li/ium longiflorum Thunb. 1. (ell Sc;. 29:147-69.Foure, J·E, e. Digonnet, ond e. Dumos. 1994. Anin vitro system for adhesion and fusion <strong>of</strong>maize gometes. 5cience 263: 1598-1600.Gerassimovo·Novoshina, H. 1957. On somecytologicol principles under~ingfertilization. Phytomorphology 7: 1582-98.Golubovskaya, I.N. 1979. Genetk control <strong>of</strong>meiosis. Int. Rev. (yrol. 58: 247-90.Golubovskoya, I.N., N.A. Avalkin, ond W.F.Sheridan. 1992. EIIect <strong>of</strong> several meioticmutants on female meiosis in maize. Dev.Genet. 13: 411-24Gustafsson, A. 1947. <strong>Apomixis</strong> in higher plantsII. lunds Univ. Arsskr NF1I43: 71-179.Huang, B.Q., and S.D. Russell. 1992. The femolegerm unit. In S.D. Russell and e. Dumas(eds.!, Sexual Reproduction in FloweringPlants. Int. Rev. (yrol. 140: 233-93.Higashiyamo 1, H. Kuroiwo, S. Kawano, and 1Kuraiwo. 2000. Explosive discharge afpollen tube contents in Torenio Fournier;.Plant Phys. 122: 11-14.Jensen, W.A. 1965. The ultrastructure andhistochemistry <strong>of</strong> the synergids <strong>of</strong> catton.Amer. 1. 80t. 52: 238-56.Jensen, W.A., and D.B. Rsher. 1968. Coltonembryogenesis: the enlrance and discharge<strong>of</strong> the pollen tube in the embryo sac. Plonto78: 158-83Knox, J.P. 1992. Molecular prabes for the plontcell surface. Protoplosmo 167: 1-9.Knox, R.B., and J. Heslop·Harrisan. 1990. Directdemonslratbn <strong>of</strong> the low permeability <strong>of</strong>the angiosperm meiotk tetrad using aflu orogenic filter. 1. Pflonzenphysiol. 62:451-59.Knox, J.P., P.J.linslead, J. King, e. Cooper, and K.Roberts. 1990. Pectin esterification isspotially reguloted both within cell walls andbetween developing tissues <strong>of</strong> root apices.Plonto 181: 512-21.Koltunow, A.M., K. Soltys, N. Nobumasa, and s.Mcclure. 1995. Anther, ovule, and nucellarembryo development in Gtrus sinensis cvVolencia. (on. 1. 80t. 73: 1567-82.Kronz, E., J. Bautor, and H.larz. 1991.ln vitr<strong>of</strong>ertilizalian <strong>of</strong> single, isolated gametes <strong>of</strong>maize mediated by electro fusion. 5ex. PlantReprod. 4: 12-16.leblanc, 0., M.D. Peel, J.G. Corman, and Y.Savidan. 1995. Megasporogenesis andmegagametogenesis in several Tripsocumspecies (Paoceae) Amer. J. 80t. 82: 57-63ligrane, R., J.G. Duckett, and K.S. Renzaglia.1993. The gametophytk·sporophytkjunction in land plants. Advanced 80tonicolResearch 19: 231-317.Martinez, E.J., F. Espinoza, and e.L Quarin.1994. Bill progeny (2/ltn) from apomicticPospolum nototum obtained through ear~pollination. Journal <strong>of</strong> Heredity. 85: 29>97.Mogensen, H.L 1988. Exclusion <strong>of</strong> malemitochondria and plastids during syngamyin barley as a basis for mote rnaIinheritance. Proe. Notl. Acod. Sri. (USA) 85:2594-97.Naumova, IN. 1978. Peculiarity <strong>of</strong>mauagametogenesis and p~t·fertilizationdevelopment <strong>of</strong> Trillium camS(hatcenseKer.·Gawi. 5oc.80t. Fr. Aduol;tes80toniques 1·2: 183-86.--.1990. Fami~ Trilliaceae. In M.S.Yakolev and lB. Batygina (eds.),(omporotive Embryology <strong>of</strong> FloweringPlants (Butomaceae·lemnaceae).leningrad: Nauka, (in Russian). Pp. 15159.---. 1993 <strong>Apomixis</strong> in Angiosperms:Nucellor and Integumentary Embryony.Boca Raton, Florida: CRC Press.Naumova, IN., A.PM. den Niis, and M.T.M.Willemse. 1992. Quantitative ona~is <strong>of</strong>aposporous parthenogenesis in Pooprotens;s genotypes. Ado 801. Neerlondica43: 1-14.Naumovo, IN., ond F. Matzk. 1998. Differencesin the initiotion <strong>of</strong> the zygotic andparthenogenetic pothwoy in the Salmonlines <strong>of</strong> wheot: ultrostructurolstudies. 5exPlant Reprodudion 11: 121-30.Naumava, IN., J.V. Osadtchiy, V.K. Shorma, PDijkhuis, and K.S. Ramulu. 1999. <strong>Apomixis</strong>in plants: structural and functional ~pects<strong>of</strong> <strong>of</strong> dipl~pry in Poo nemorolis ond P.polustris. Protoplosmo 208: 186-95.Naumova, IN., and M.T.M. Willemse. 1982.Nucellar po~embryany in 5orrococcohumilir. ultrastructural aspects.Phytomorphology 32: 94-108.--.1995. Ultrastructuralchorocterization <strong>of</strong> ap~pory in Ponicummaximum. 5ex. Plant Repr. 8: 197-204.Nogler, G.A. 1984. Gametophytic opomixis. InB.M. Johri (ed.), Embryology <strong>of</strong>Angiosperms. Berlin: Springer·Veriag. Pp.47>-518.Peel, M.D., J.G. Carman, and o. leblonc. 1997.Megasporocyte callose in opomicticbuHelgr~s, Kentucky bluegrass,Pennisetum squomulotum Fresen,Tripsocum Lond weeping lovegrass. (rap5c;. 37: 717-23.Pennell, R.I. 1992. Cell surface orabinogoloctonproteins, arobinogalactons ond plontdevelopment. In J.A. Callow ond J.R. Green(eds.1, Perspedives in Plant (ell Recognition.Society <strong>of</strong> Exp. BioI. Series 48. CambridgeU.K.: Cambridge Univ. Press.Pennell, R.I., l. Jonniche, P. Kjelbom, G.N. Sc<strong>of</strong>ield,J.M. Peart, ond K.A. Roberts. 1991.Developmental regulation <strong>of</strong> a plasmamembrane arabinagalactan protein epitope inoilseed rape flowers. Plant (ell 3: 1317-26.Pennell, R.I., and K.A. Roberts. 1990. Sexualdevelopment in the pea is presaged byaltered exprmion 01 arabinogolactanpratein. Nature 344: 547-49.Philipson, M.N. 1978. <strong>Apomixis</strong> in Cortoderioiuboto (Gramineoel. New Zeal. 1. 801. 16:4>-59.Remy, w., P.G. Gensel, and H. Hoss. 1993. Thegametophyte generation <strong>of</strong> some earlyDevonian land plants. Int. 1. Plont5ci. 154:3>-58.Russell, S.D. 1985. Preferential fertilization inPlumbago; Ultrastructural evidence forgamete·level recognition in an angiosperm.Proe. Notl. Acad. 5ri. (USA) 82: 6129-32.--. 1992. Dauble fertilization. In S.D.Russell and e. Dumas (eds.!, SexualRepoduction in Flowering Plants. Int. Rev.(ylol. 140: 357-88.Savidan, Y. 1982. Nalure et heredite de I'apomixiechez Poniwm maximum Jacq. Trov et DocOR5TOM 153:1-159.Schulz, S.R., and W.A. Jensen. 1968. (opselloembryogenesis: the egg, zygote and yaungembrya. Amer. 1. 80t. 55: 807-19.Vielle, J.p, B.l. Burson, E.e. Bashaw, and M.A.Hussey. 1995. Eorly fertilizotion events in thesexual and aposporous egg apparotus <strong>of</strong>Penn;setum ciliore ILl link. Plant 1. 8: 30916.Vielle·Colzada, J.p, C.F. Crane, ond D.M. Stel~.1996a. <strong>Apomixis</strong>: the asexual revolution.5cience 274: 1322-23.Vielle·Calzada, J.p, M.l. Nuccio, M.A. Budimon, 1l.Thomas, B.L Burson, M.A. Hussey, and R.A.Wing. 1996b. Comparative gene expressionin sexual and apamictic ovaries <strong>of</strong>Pennisetum ciliore. Plant Mol. 8iol. 32:1085-92.Voigt, PW., and E.e. 8ashaw. 1972. <strong>Apomixis</strong> andsexuality in &ogr05tis curvulo. (rop Sci. 12:843-47.Wakana, A., ond S. Uemoto. 1987. Advenliveembryogenesis in Glrus (Rutoceae) II. Theoccurrence <strong>of</strong> adventive embryos withoutpollination or fertilizatian Arner. 1. 801. 74:517-30.Wilms, H.J., J.l. van Wenl, M. Cresli, and f.Gampolini. 1983. Advenlive embryagenesisin Citrus. (oryolagio 36: 6>-78.
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(over illustration:Pictured is an i
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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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112 a... A. 8icUeI1993). Before the
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114 ROil A. 8idlooll(Sherwood et al
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116 R... A. IkbeIlinduced mutation
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118 Ron A. Mlltllstudies of apomixi
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120 I... A. BkbollLeblanc, 0., M.D.
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122 Olivier L.bla.,.ncI A.d,ea M.nu
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130 Ofjyier LH......AodrH Mom","Mar
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134 OIivie' I
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136 or..io. u.~100< awd Aod.... M.n
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150 (adlda Borge. da VaNe and Joh.
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158 Yv•• Savid..unreduced polar
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160 Yves SoviclaoBashawet al. (1970
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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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176 U.IGm..lln.apomictic pathway wi
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178 IJeIGr.........development with
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180 Ud Gr....lIoo,(Rhoades and Demp
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182 Ue5Gro........Meiosis is an alm
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184 Uel Gn...lIDo,To date, the phen
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186 Uei Gr....lIa..do not display p
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194 UeI Gm..ikIonsystem developed b
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198 lleIG
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200 UtlG09".ila,ndescribed that are
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202 UeIGfo...ll...Two additional po
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208 Uel GtO,,"ikIa..MIodzik, M., Y.
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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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214 Uta Praebll aod Rod ScottCrosse
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216 Uta Praektlt.. Rod Scana minori
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