166 rves Sqyldaounanswered. However, more progress hasbeen achieved toward producing an apomicticgrain during the last ten years than ever before,mostly because <strong>of</strong> the development andapplication <strong>of</strong> new techniques. As moleculardissecting tools continue to improve, we willsee great progress in our understanding <strong>of</strong>how apomixis is controlled and the isolationand manipulation <strong>of</strong> its components. Anotherpromising avenue, approaches based onmutagenesis, is discussed in the following twochapters. These approaches will undoubtedlybetter our understanding <strong>of</strong> the regulation <strong>of</strong>reproduction as a whole. In the end, apomixiscertainly cannot be manipulated without athorough understanding <strong>of</strong> how it is controlledin the wild.ReferencesAsker, S. 1979. Progress in apomixis research.Heredffcs 91: 231-40.Asker, S., ond L. Jerling. 1992. <strong>Apomixis</strong> inPlants. BolO Raton, Florida: CRC Press.Bharathi, M., U.R. Murty, K.B.R.S. Visarada, andA. Annopuma. 1991. Possibility <strong>of</strong>transferring obligate apomixis from(enmrus cilioris L to Sorghum bicolor(LlMoench. <strong>Apomixis</strong> Newsleffer 3: 13-14.Bashaw, E.e. 1975. Problems and possibilities <strong>of</strong>apomixis in the improvemenl <strong>of</strong> tropicalforage grosses. Tropical Forages inUvestock Production Systems. 24: 23-30.Madison, WislOnsin: American Society <strong>of</strong>Agronomy.Boshaw, E.e., AW. Hovin, and E.e. Hoh. 1970.<strong>Apomixis</strong>, its evolutionary significance ondutilization in plant breeding. Proc. XI Int.Grassl. (ongr. Madison, Wisconsin:American Society <strong>of</strong> Agronomy. Pp. 24548.Bergquist, R.R. 1981. Tronsfer from TripsowmdOdyloides to corn <strong>of</strong> 0 mojor gene lorusconditioning resistance to Pu«inio sorghi.Phytopathology 7: 518-20.Bernard, S., and D.C. Jewell. 1985. Crossingmaize with Sorghum, Tripsowm and millet:the products and their level <strong>of</strong>development following pollination. Thear.Appl. Genet. 70: 474--83.Birchler, J.A. 1993. Dosage ana~sis <strong>of</strong> maizeendosperm development. Ann. /lev. Genet.27: 181-204.Brown, W.V., and H.P. Emery. 1958. <strong>Apomixis</strong> inthe Grominelle : Panicoidelle. Amer. 1. Bot.45: 253-63.Burton, GW., !.C. Millot, and W.G. Monson.1973. Breeding procedures for Paniwmmaximum suggested by plant variabilityand made <strong>of</strong> reproduction. (rap Sci. 13:717-20.Corman, 1.G. 1997. Asynchronous expression <strong>of</strong>duplicate genes in angiosperms may lOuseapomixis, bispory, tetraspory, andpo~embryony (review). Bioi. 1. Unn.Soc.61: 51-94.Chu, e.e., e.e. Wong, e.s. Sun, e. Hsu, K.e. Yin,and LV. Chu. 1975. Establishment <strong>of</strong> onefficient medium for onther culture <strong>of</strong> ricethrough lOmparotive experiments on thenitrogen sources. Sci. Sinico 18: 659--il8.de Wet, 1.MJ.de 1979. Tripsowm introgressionand agronomic fitness in maize Ilea maysLl. Proc. (onf. Broadening Genet. Bose inCrops. Wageningen: Pudoc Publish. Pp.203-09.Dujardin, M., and WW. Han no. 19840.Microsporogenesis, reproductive behaviorand fertility in five Pennisetum species.Thear. Appl. Genet. 67: 197-201.--.1984b. Pseudogomausporthenogenesis and fertilizotion <strong>of</strong> apearlmillet xPennisetum orienlole apomicticderivotive.1. Hered. 75: 503-04.--.1986. An apomictic po~hoploidobtoined from 0 peorl millet xPenniselumsquomulatum opomictic interspecifichybrid. Thear. Appl. Genet. 72: 33-36.--.1989. Crassobility <strong>of</strong> Pellrl milletwith wild Pennisetum species. Clop Sci. 29:77-80.Engle, L.M., 1.MJ. de Wet, and J.R. Harlan.1973. Cytology <strong>of</strong> backcross <strong>of</strong>fspringderived from amoize- Tripsowm hybrid.(rap Sci. 13: 690-94.--.1974. Chramosomol variationamang <strong>of</strong>fspring <strong>of</strong> hybrid derivatives with20 lea and 36 Tripsowm chromosOmes.(oryolagio 27: 193-209.Galbraith, D.W., K.R. Harkins, 1.M. Maddox,N.A. Ayres, D.P. Sharma, and E.Firoozaboby. 1983. Arapid flow 1yI0metricana~is <strong>of</strong> cellcyde in intact plant tissues.Science 220: 1049-51.Galinat, W.e. 1971. The origin <strong>of</strong> maize. Ann./lev. Genel. 5: 447-78.Galina!, W.e., P. Chandravadona, and B.GSRoo. 1970. Cytological mop <strong>of</strong> Tripsowmda
Yr."" .f Apomi.i. I.rotgir Wid. Cr.ue. 167leblonc, 0., o. Grimonelli, N. Islom-Foridi, J.8erthaud, and Y. Sovidan. 1996.Repraductive behavior in moize- Tripsacumpa~haploid plants: Implications far thetransfer <strong>of</strong> apomixis into maize. 1. Hered.87: 108--11.leblonc, 0., M.D. Peel. J.G. (arman, and Y.Savidon. 19950. Megosporogenesis andmegagometogenesis in several Tripsacumspecies (Pooceoe). Amer. 1. Bot. 82: 5763.leblanc, 0., and Y. Sovidan. 1994. Timing <strong>of</strong>megasporagenesis in Tripsacum species(Pooceoe) as related to the control <strong>of</strong>apomixis and sexuolity. Polish. Bot. Stud.8: 75-8l.Mangelsdorf, P.c., and R.G. Reeves. 1931.Hybridizalion <strong>of</strong> maize, Tripsacum, andEuchlaena. 1. Hered. 22: 339--43.Marsholl, R.D., and A.H.D. Brown. 1981.Estimotion <strong>of</strong> the level <strong>of</strong> apamixis in plontpopulations. Heredity 32: 321-33.Martinez, EJ., EEspinoza, and C.L Quarin.1994. Bill progeny (2ntnl from apomicticPaspalum nolatum obtained through eor~pollination. 1. Hered. 85: 295-97.Mogie, M_ 1988 Amadel for the evolution andcontrol <strong>of</strong> generative apomixis. Bioi. 1.Unn. Soc. 35: 127-53.Murashige, 1, ond ESkoog. 1962. Arevisedmedium for rapid growth and bioassoyswith tobocco tissue cultures. Physiol. Plant.15,413-97.Ndikumona, J. 1985. Etude de rhybridationentre especes opomitiques et sexuees damIe genre Brachiaria. Ph.D. dissertation.Univ. <strong>of</strong> lauvoin, louvoin-lo-Neuve,8elgium.Nogler, G.A. 1984. Gametophytic apomixis. In8.M. Jahri (ed.), Embryology <strong>of</strong>Angiosperms. 8erlin: Springer-Verlag.Pp.475-518.Noirot, M. 1993. Allelic rotios and sterility in lheogamic complex <strong>of</strong> the Maximae(Panicoideael: evolutionary role <strong>of</strong> theresidual sexuolity. 1. Evol. 8io. 6: 95-101.Ozias-Akim, P., E.l. lubbers, W.W. Hanna, ondJ.W. McNay. 1993. Transmission <strong>of</strong> theapomictic mode <strong>of</strong> reproduction inPennisetunr. co-inheritalKe <strong>of</strong> the trait andmoleculor markers. Theor. Appl. Genet. 85:632-38.Ozios-Akins, P., O.Rache, and w.w. Honna.1998. Tight clustering and hemizygosity <strong>of</strong>opomixis-linked molecular morke~ inPennisetum squomulatum implies geneticcontrol <strong>of</strong> apospory by adivergent locusthot may have no allelic form in sexuolgenotypes. Proc. Nal. Acad. Sci. (USA) 95:5127-32.Pernes, J. 1972. Organisotion evalutive d'ungroupe agomique: la section des Maximaedu genre Panicum (GramineesJ. Ph.D.dissertotion, University <strong>of</strong> Poris.Petrov, OJ., N.I. 8elausava, and E.S. Fokino.1979. Inheritance <strong>of</strong> apomixis and itselements in carn-Tripsocum hybrids.Genetika 15:1827-36.Pelrov, DJ., N.I. 8elousovo, E.5. Fokina, R.M.Yalsenka, L1. laikova, and 1P. Sorokina.1984. Tramfer <strong>of</strong> some elements <strong>of</strong>apomixis from Tripsocum to maize. In OJ.Petrov led.), <strong>Apomixis</strong> and Its Kole inEvolution and Plant Breeding. RussianTranslotion Series 22. Rotterdam: AABalkena. Pp. 9-78.Purseglove, J.W. 1972. Tropical Crops,Monocotyledons. New York: langman.Ramulu, KS., P. Dijkuis, E. Rutgers, J. 8laas, EA.Krens, JJ.M. Dons, eM. (alijn-Hoojmans,ond H.A. Verhoeven. 1996.Microprataplost-medioted transfer <strong>of</strong> singlespecific chromosomes between sexual~incampotible plants. Genome 39: 921-33.Sovidan, Y_ 19820. Nature et heredite deropomixie chez Panicum maximum Jacq.Travaux & Documents ORSTOM 153: 1159.--.1982b. Embryologkal ana~sis <strong>of</strong>facultative apomixis in Panicum maximumJacq. [rap Sci. 22: 467-69.---.2000. <strong>Apomixis</strong>: Genetia and8reeding. Plant Breeding Keviews 18: 1386.Simone, G.W., and A.L Hooker. 1976.Monogenic resistance in corn toHelminthosporium turcicum derived fromTripsacum floridanum. Proc. Am.PhytopathoJ. Soc. 3: 207.
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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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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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38 (IIarIe,F.e-.The following recip
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40 CWIes F.
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42 C"Ie
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Chapter 4Ultrastructural Analysis o
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46 T.""". N. Naomo•• ..dJ...·P
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(Figure 4.2a,b,c). Their cytoplasm
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50 Tomaro N. Naurnovo ond Jeon-Phil
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Figure 4.2 (cont'd)
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54 Tamara N. Nauma.a and J...-Phaip
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56 Tamara H. Haumaya ...d J...-Phmp
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58 Tamara N. N....... Old JOGO-PUIp
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60 T.mar. N. N••may•••dJ
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62 Tamara N. Nouma.o and Jeon-Phmpp
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Chapter 5Genetic Analysis of Apomix
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66 Robe" T. SherwoodIn mitotic dipl
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68 ••It T. Sllorwoodmegasporoge
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70 Robert T. SherwoodIdentification
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72 R....11 T. SlMrwoodwe postulate
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74 Rob.rt T. Sherwoodin the gametop
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76 Rokrt T. SHtwoodPerhaps the most
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78 Robe" T. S~erwoodEnvironment pla
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80 Robert T. SherwoodBanaglio, E. 1
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82 Robe" 1. Sherwood---. 1989. Apom
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84 Dcaitl ~11i-, lot To...., .od Di
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86 Do.lel Griome/I-, Joe To~ ....,
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88 Oaoitl G
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90 o.lel ~II-, Jo. lob.., aod Diego
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and meiotic or developmental mutant
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94 D..iel Grimallelli-, Jo. Tohme,
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96 10k. G.(o,.,..mechanisms (Figure
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98 Jolo.G.(_explain the existence o
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100 JolI.G.C....parameters affectin
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102 1010. G.(arma.was developed in
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104 J... G.(.....effects), which co
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106 J.hG.eforseveral thousand to a
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108 Jelo.G.(_large linkage group in
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11 0 Joh. G.(orma.---. 1997. Asynch
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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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216 Uta Praektlt.. Rod Scana minori
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21 8 Uta P...ke~ .d Rod S
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220 Ura P"",k.h .d Rod ScottStubby
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222 Uta Praekelt Old Rod S,ollelong
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224 Uta P...kek .d Rad Scaliresulti
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226 Uta P,..kelt ..dRod S
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228 Uta P..utt aod Rod 5
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230 1\omas Dresselhaus, Joh. G. (IJ
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progression or inhibition of develo
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234 T1lo.... Dr......... Jo~. G. (.
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236 TIoo.... D........., J.~. G. (a
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238 T1tonoas Dr......... Joino G. '
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240 no- P,ossdlaos, Jo. G.'-.... Y.
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242 1110""" 0,........, M. G. c,.._
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