240 no- P,ossdlaos, Jo. G.'-.... Y.os ScrrioI..plants, and if so, what would be the impact.This issue is especially important in the centers<strong>of</strong> origin for the crop plants. Furthermore, theissue <strong>of</strong> how apomixis might affect geneticdiversity, and whether it would increase ordecrease monoculture farming needs to beexplored. Based on field studies on herbicideand/or insecticide resistant plants, we canprobably expect engineered apomixis genes tomove through vertical gene transfer (transfer<strong>of</strong> a gene from plant to plant via sexualreproduction/pollen) (Lutman 1999). The rate<strong>of</strong> horizontal gene transfer (asexual gene flowbetween organisms) is relatively low and therisk negligible, however, microbiological riskassessment studies in this area could be useful(Syvanen 1994). Given our current knowledge,it appears unlikely that microorganisms couldgain some advantage over wild relatives afteruptake <strong>of</strong> apomixis genes.Ifapomixis is controlled by multiple genes, theprobability <strong>of</strong> diffusing this trait to wildrelatives is extremely low. The transfer <strong>of</strong>several genes to a wild plant should lower itsfitness to a level unacceptable for survival inthe wild (Berthaud, Chap. 2). If apomixis iscontrolled by a single gene, which would resultin obligate apomictic wild races, these raceswould lose their potential to evolve. Ifdominant, an apomixis gene could rapidlybecome fixed in an outcrossing sexualpopulation. Therefore, in theory, apomixistransgenes could possess advantages thatmight result in the uncontrollable spread <strong>of</strong>the transgenes (van Dijk and van Damme2000). Inducible apomictic systems and malesterility might circumvent these problems.Nevertheless, the described possibilitiesindicate that risk assessment studies andresearch to investigate the ecologicalimplications <strong>of</strong> novel apomictic crops (onceavailable) to the environment are an absolutenecessity. In addition, socioeconomic studieson the positive and negative implications <strong>of</strong>this technology for breeders, seed companies,and farmers-in both developing and developedcountries (see also IPR) will be required, andthe research results should be communicatedto all potential users.SummaryThe extensive introduction <strong>of</strong> apomixis intosexual crops will undoubtedly rely on geneticengineering, as we anticipate that morecandidate genes (especially regulatory genesand tissue/cell-specific promoters) andenabling techniques will be identified anddeveloped in the near future. Transformationtechnology for all major crops is now availableand inducible systems are currently beingdeveloped and optimized, allowing the control<strong>of</strong> transgene expression and activity evenunder field conditions. Adventious apomixisusing already described or novel genes underthe control <strong>of</strong> ovule-, nucellus- or archesporespecificpromoters is probably the easiest wayto engineer the apomixis trait. Plant breedersand seed producers would like to generateinducible obligate mitotic diplospory incombination with autonomous endospermdevelopment. The latter is probably the mostdifficult aspect <strong>of</strong> engineering apomixis,espeCially for cereals such as wheat, rice, andmaize, because <strong>of</strong> dosage and imprintingeffects.Although apomixis is a hot topic in plantresearch, our current understanding <strong>of</strong> bothapomictic and amphimictic reproductionpathways in higher plants is still extremelylimited. The economic potential <strong>of</strong> apomixismight provide the impetus to bring apomicticcrops to the marketplace, and in the process itmay well contribute significantly to our futureunderstanding <strong>of</strong> the molecular regulation <strong>of</strong>the many different sexual and apomictic plantreproduction pathways.International and interdisciplinary approachesand efforts are now needed to study andmanipulate seed reproduction. It will be
Ge..tk Eagloeeri.g .1 Ap.mlxi. io Se..al Ct.,.: A(ritkal A.........' .1 the Apamlli. Tedooology 241necessary (i) to characterize the genetic reproduction through seeds in apomicticregulation <strong>of</strong> apomixis and isolate the systems and sexual crops," coordinated by T.responsible genes, (ii) to analyze the genetic Dresselhaus). In 1999, a transatlanticand molecular bases <strong>of</strong> sexual reproduction consortium was initiated between two publicand to isolate the corresponding genes, and institutions (CIMMYT and IRD) and three(iii) to produce the tissue/cell-specific and private companies (pioneer Hi-Bred, Novartis,inducible/repressible promoters that will be and Group Limagrain). This is just a beginningneeded to control the expression <strong>of</strong> the target and more concerted projects are needed ingenes. Concerted international research efforts order to reach the ambitious aim <strong>of</strong>have been made in Europe aimed at manipulating the apomixis trait in crops.understanding apomictic and sexual<strong>Apomixis</strong> technology will <strong>of</strong>fer many excitingreproduction pathways in order to developopportunities for the agriculture <strong>of</strong> the 21 sftools for the manipulation <strong>of</strong> the apomicticcentury, and indeed many patents alreadytrait (e.g., an E.U. Research Technology andhave been filed with many more yet to come.Development (RTD) project entitled "TheIt is critically important that these patents bemanipulation <strong>of</strong> apomixis for theheld and used for the good <strong>of</strong> all. Publicimprovement <strong>of</strong> tropical forages," coordinatedinstitutions in particular must safeguard theby M. D. Hayward; a RTD project entitledaccess <strong>of</strong> developing countries to these"<strong>Apomixis</strong> in agriculture: a molecularenabling technologies. In all likelihood,approach," coordinated by M. van Lookerenconstraints to the broad and generous use <strong>of</strong>Campagne; and a Concerted Action Projectapomixis technology will be political andentitled "Introducing and controlling asexualeconomic rather than technical in the future.ReferencesBlakey, CA., CL Dewald, and S.L Goldman. Colombo, L, 1. Franken, A.R. Von der Krol, PE.1997. Co-segregation <strong>of</strong> DNA markers with Wittich, HJ. Dons, and G.C Angenent.Adams, S., R. Vinkenoog, M. Spielman, H.G.Tripsacum fertility. Moydica 42: 363-69. 1997. Dawnregulotion <strong>of</strong> ovule-specificDickinson, and RJ. Scott. 2000. Porent-<strong>of</strong>Caddick, M.X., AJ. Greenland, I. Jepson, K.P.MADS box genes from petunia results inorigin effects on seed development inKrouse, N. Qu, K.V. Riddell, M.G. Solter, W. maternally controlled defects in seedArabidapsis thaliana require DNASchuch, U. Sonnewold, and A.8. Tomsett. development. Plant Cell 9: 703-15.methylation. Development 127: 24931998. An ethanol inducible gene switch for Diatchenka, l., Y.·F. Chris Lou, A.P Campbell, A.2502. plonts used to manipulate carbon Chenchik, EMoqodom, B. Huang, S.Alleman, M., and J. Dodar. 2000. Genomicmetabolism. Not. Biotechnal. 16: 177-80. Lukyanav, K. Lukyanaw, N. Gurskaya, E.D.imprinting in plants: observations andChose, S. 1969 Monoploids and monoploid. Sverdloc, ond PD. Siebert. J996.evolutionary implications. Plant Mol. BioI.derivatives <strong>of</strong> maize (leo mays U.Suppression subtractive hybridizotian: A43: 147-61.Botanical Review 35: 117-fJ7.method for generating differentiallyAsker, $.E., and LJerling. 1992. <strong>Apomixis</strong> inChen, E, and M.R. Faolod. 1997. Molecularregulated or tissue-specific eDNA probesPlants. Boca Raton, Florida: CRC Press.organization <strong>of</strong> agene in barley whichand libraries. Proe. Notl. Acad. Sri. (USA)Boi, X., B.N. Peirson, EDong, CXue, ond CA.encodes aprotein similar to aspartic 93: 6025-30.Mokor<strong>of</strong>f. 1999.lsolotion ondpralease and its specific expression in Dresselhaus, 1, S. Cordts, S. Heuer, M. Souter, H.characterization <strong>of</strong> SYNI, a RAD21·likenucellar cells during degenero1;on. Plant liirz, and E. Kronz. J999. Novel ribosomalgene essential for meios~ in Arabidopsis.Mol. BioI. 35: 821-31.genes from maize are differentiallyPlant Cell 11: 417-30.Cheng, M., J.E. Fry,S. Pong, H. Zhau, CM.expressed in the zygotic and somatic cellBoss, B.L 2000. Double-stranded RNA as a Hironaka, D.R. Duncon, 1W. Conner, ond Y. cycles. Mol. Gen. Genet. 261 : 416--27.template for gene silencing. CellI 01:Won. 1997. <strong>Genetic</strong> transformation <strong>of</strong> Dresselhaus, 1, H. Uirz, and E. Kranz. 1994.235-38.wheat medioted by AgrobocteriumRepresenlolive cDNA libraries from fewBicknell, R.A., and K.B. Bicknell. 1999. Who willtumefariens. Plant Physial. 115: 971-80. plant cells. Plant 1. 5: 605-10.benefit from apomixis? Biotechnology andChristou, P1996. Transformation technology. Drews, G.N., D. Lee, and CA. Christensen. 1998.Development Monitor 37: 17-21.Trends Plant Sci. 1: 423-31.<strong>Genetic</strong> analysis <strong>of</strong> female gametophyteBirchler, JA 1993. Dosoge analysis <strong>of</strong> moizede Vries, G.E. 1998. Post, Present And Future development and function. Plant Cell] 0:endosperm development. Annu. Rev.Considerations In Risk Assessment When 5-17.Genet. 27: IBI-204.Using GMOs. Billhaven, The Netherlands: Evans, L.1 199B. Feeding the Ten Billion: PlantsCommission <strong>Genetic</strong> Modificotion.and Population Growth. New York:Cambridge University Press.
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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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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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126 OIlrier Ltblaooc ood AocI...Mo,
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128 or..Ie, leblal( allll Aodrea Ma
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130 Ofjyier LH......AodrH Mom","Mar
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132 Olivier leblaoc and Aldr.. Mall
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134 OIivie' I
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136 or..io. u.~100< awd Aod.... M.n
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138 udda lorge. do Val. aod Joh W.
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140 Ca
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142 C«ida Jorge. do Va" aod Jah W.
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144 (..iIda 80rges cit Va" DOd Ja~.
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146 Cacido ....... Vole..J.b W. MIe
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148 Ca
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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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154 Yve< 5
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156 rn. s.mdaosearch for an opitimu
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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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188 UtI GrOSllilaot.development of
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