202 UeIGfo...ll...Two additional points should be stressed. First,the technologies needed to engineer andcontrol apomixis under field conditions are notyet available and must first be developed, apoint that cannot be overstated as theidentification <strong>of</strong> potential regulatory genesprogresses rapidly. Second, this review <strong>of</strong> plantreproduction has focused on a geneticperspective with Mendelian traits controllingsexual and apomictic development. However,gene regulation by genomic imprinting maynot be the only epigenetic mechanism thatshould be considered. Current knowledge <strong>of</strong>the regulation <strong>of</strong> apomixis is fully compatiblewith an epigenetic view <strong>of</strong> this trait and weshould keep our minds open to alternativeexplanations that are epigenetic in nature. Ifapomixis is, for instance, due to epigenetic.in teractions between genomes, theengineering <strong>of</strong> apomixis in sexual species willbe much more complex.Finally, the accessibility <strong>of</strong> apomiXIStechnology to a broad community <strong>of</strong> plantbreeders in the public and private sectorworldwide must be ensured. If universal andequitable access to apomixis technologycannot be achieved through innovativepatenting and licensing, then the e.xcitingscience discussed in this volume will have littlepositive impact (http://billie.btny.purdue.edu/apomixis/).AcknowledgmentsMy special thanks go to Richard Jefferson forhis valuable contributions and comments, andto Gian Nogler for pro<strong>of</strong>reading themanuscript. I would also like to thank JeanPhilippe Vielle-Calzada, Robert Pruitt, andRoss Bicknell for their stimulating discussions,and Alison Coluccio for help with thebibliography. SpeCial thanks go to RobertPruitt, Susan Lolle, Richard Jefferson, WeiYang, Satya Nugroho, lmran Sidiqqi, BrunoTinland, and the members <strong>of</strong> my laboratoryfor allowing me to cite data prior topublication. I am grateful for the supportprovided by the European Molecular BiologyOrganization, the Human Frontiers <strong>of</strong> ScienceProgram, the Janggen-Poehn-Foundation, andthe Demerec-Kaufmann-HollaenderFellowship in Developmental <strong>Genetic</strong>s.Research in my lahoratory was funded by theCold Spring Harbor Laboratory President'sCouncil, Pioneer Hi-Bred International, grantMCB-9723948 <strong>of</strong> the National ScienceFoundation, and grant # 9801207 <strong>of</strong> theNational Research Initiative CompetitiveGrants Program <strong>of</strong> the U.5.D.A.ReferencesAhokos, H. 1977. Amutant <strong>of</strong> barley: triploidinducer. Barley Genet. Newsl. 7: 4-6.Allemon, M., ond, J. Dooor. 2000. Genomicimprinting in planls: observations andevolutionary implications. Plant Mal. Bial43: 147-61.Angenent, G.c., and LColombo. 1996.Molecular control <strong>of</strong> ovule development.Trends Plant Sci 1: 228-32.Antoine, A.F., J·E. Faure, S. Cordeiro, C. Dumas,M. Rougier, and J.A. Feija. 2000. Acalciuminflux is triggered and propagates in thezygote as awavefront during in vitr<strong>of</strong>ertilization <strong>of</strong> flowering ~ants. Proc. Natl.Acad. Sci. USA 97: 10643-48.Aoyama, 1, and N·H. Chua. 1997. Aglucocorticaid·mediated lranscriptionalinduction system in transgenic plants. PlantJ.ll:60S-12.Arabidopsis Genome Initiative 2000. <strong>Analysis</strong> <strong>of</strong>the genome sequence <strong>of</strong> the floweringplant Arabidapsis thaliana. Nature 408:796-8lS.Asker, S.E. 1979. Progress in opomixis research.Herooitas91: 231-40.--.1980. Gametophytic apomixis:Elements ond genetic regulation. Hereditas93: 277-93.Asker, S.E., A. Hagberg, and G. Hagberg. 1983.<strong>Apomixis</strong> in barley? Sver. UtsiidesFiiren.Tidskr. 93: 7s-76.Asker, S.E., and LJerling. 1992. <strong>Apomixis</strong> inPlants. Boca Raton, Florida: CRC Press.Baker, S.c., K. Robinson·Beers, J.M. Villanueva,J.c. Gaiser, and C.S. Gasser. 1997.Interactions among genes regulating ovuledevelopment in Arabidopsis tholiono.<strong>Genetic</strong>s 14S: 1109-24.Ballinger, D.G., and S. Benzer. 1989. Targetedgene mutations in Drosophila. Proc. Natl.Acad. Sci. (USA) 86: 9402-ll6.Barcacoa, G., A. Manucato, E. Albertini, J.lethaf, A. Gerats, M. Penotti, M. Falcinelli.1998. Inheritance <strong>of</strong> parthenogenesis inPoa pratensis L: auxin test and AFLPlinkage ana~ses support monogeniccontrol. Theoretical and Applied <strong>Genetic</strong>s97: 74-82.8ashaw, E.C., and W.W. Hanna. 1990. Apomicticreproduction.. In G.P. Chapman led.},Reproductive Versatility in the Grosses.Cambridge, U.K.: Cambridge UniversityPress. Pp. 10~30.8echtold, N., J. Ellis, and G. Pelletier. 1993. Inplanta Agrobaeterium gene transfer byinfiltration 01 adult Arabidapsis thalianaplants. CR. Acad. Sci. Paris. 316: 1194-99.
F",," s.....Uy I. <strong>Apomixis</strong>: MoIeuIor ..dGHelk Ap,.-M, 203Bedinger, P., and S.D. Russell. 1994.Gametogenesis in maize. In M. Freeling andV. Walbot (eds.), The Maize Handbook. NewYork: Springer-Verlag. Pp. 48-61.Belanger K.D., and R.s. Quatrano. 2000.Polarity: the role <strong>of</strong> localized secretion. Curr.Gp. Plant Bioi. 3: 67-72.Bellen, HJ., CJ. O'Kane, C. Wibon, U.Grossniklaus, R_K_ Pearson, and WJ.Gehring. 1989. P-element-mediatedenhancer deteelion: aversatile method 10study development in Drosophila. Genes andDev. 3: 1288-1300.Bellen, HJ., C. Wilson, and WJ. Gehring. 1990.Dissecting the complexity <strong>of</strong> the nervouss~tem by enhancer detection. Bioessays 10:199-204.Belostotsky, D.A., and R.B. Meagher. 1996. Apollen-, ovule-, and ear~ embryo-specificpo~ (A) binding pratein from Arabidopsiscomplements essential functions in yeast.Plant Ce1/8: 1261-75.Bennelzen, lL, and M. Freeling. 1993. Grassesas asingle genetic system: genomecomposition, colinearity ond compatibility.Trends in <strong>Genetic</strong>s 9: 259-61.Bensen, RJ., G.S. Jahal, V.c. Crane, J.T. Tossberg,P.S. Schnable, R.B. Meeley, and S.P. Briggs.1995. Ooning and characterization <strong>of</strong> themaize AnI gene. Plant Ce1/7: 75-84.Berger, F. 1999. Endosperm development. Curr.Op. Plant Bioi. 2: 28-32.Bicknell, R.A. 1997. Isolation <strong>of</strong> adiploid,apomictic plant <strong>of</strong> Hierodum aurontiacum.Sex. Plant Reproo. 10: 168-72.Bicknell, R.A., N.K. Barsl, and A.M. Kahunow.2000. Monogenic inheritalKe <strong>of</strong> opomixis inIwo Hieradum species wilh dislinctdevelopmental me
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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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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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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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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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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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124 Olivier lt~1ao< aod Aock.. Mau"
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132 Olivier leblaoc and Aldr.. Mall
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134 OIivie' I
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142 C«ida Jorge. do Va" aod Jah W.
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148 Ca
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150 (adlda Borge. da VaNe and Joh.
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