152 Ccdda Borge. do v..... J.h W. MilesSherwood, R.T., CC Berg, ond B.A. Young.1994.lnheritonce <strong>of</strong> opospory inbuffelgross. Clop Sci. 34: 149G-94.Swenne, A., B-P louont, ond M. Dujordin. 1981.Induction por 10 cokhicine de formesoutotetro~oides chez Brochiorio ruziziensisGermain el Evrard (Grominlies). Agron.Trap. 36: 134-41.lorlOfeHo, CM., and E.C Bushaw. 1966.Inheritonce ond conlrol <strong>of</strong> ob~goleapomixis in breeding buffe~ross,Pennisetum aliarB ((enehM ciliorisl. ClopSci. 6: 473-76.Toledo, J.M. (ed.). 19B2. Manual para 10evo/uaaiin ogroniimico: Red Internocionolde Evo/uadiin de Postos Tropicoles. Coli,Colambio: CIAT.Volle, CB. do. 1986. Cytology, mode <strong>of</strong>reproduction ond forage quolily <strong>of</strong> selectedspecies <strong>of</strong> Brochiorio Griseb. Ph.D.dissertation. University <strong>of</strong> lIfinois-Urbono.Urbano, Illinois. USA.--.1990. (oI~iio de germoplasmo dBespeeies de Brochioria no ClAT: estudosb6si!os visondo 00 melhoromento geootico.EMBRAPA·CHPGC, Documentos, 46, CompoGrande, Brazil: EMBRAPA-CHPGCVolle, CB. do, S. Calixta, and M.C Amezquha.19930. Agronomic evaluation <strong>of</strong> Brochioriogermplasm in Brozil. PrO(, XVlllnt, Grassl,Congr., HZGA, TGSA, HZSAp, ASAP·Old, ondHZIAS, Polmer>ton North, New Zealand.1993. pp. 511-12.Volle, CB, do, and CGlienke. 1991. New sexualaccessions in Brochiorio. <strong>Apomixis</strong>NBlVSietter 3: 11-13.Volle, CB. do, CGlienke, and G.O.CLeguizaman 1993b. Breeding <strong>of</strong> opomicticBroehiorio through inlerespecifichybridisotion. Prac. XVII Int. Grassl, (ongr.,NZGA, TGSA, NZSAp, ASAP-Old, ond NZIAS,Palmerllon North, New Zeolond. 1993, Pp.427-28.Volle, CB. do, G. leguizomon, and N.R. Guedes,1991.lnter>pecific hybrids <strong>of</strong> Brochior;o(Gramineoe). <strong>Apomixis</strong> Newsletter 3: 1GIl.Volle, CB. do, B.L Moass, CB. de Almeido, andJ.CG. (0110. 1993c. Morpho/ogicolmaracterizolion <strong>of</strong> Broch;orio germplosm.Proc. XVII Int. Grassl. Congr., NZGA, TGSA,NZSAp, ASAP-Old, ond NZIAS, PalmerstonNorth, New Zealand. 1993. pp. 20~9.Volle, CB. do, Y. Sovidon, and LJonk. 19B9.<strong>Apomixis</strong> and sexuolity in Brachioriodecumbens Stapf. Proc XVI Int.Gross/.(ongr., Nice, Fronce. 1989. Assoc.Fr. Prod. Fourragilre. v.l. pp. 407~8.Volle, CB., ond Y. Sovidon. 1996. <strong>Genetic</strong>s,cytogenetics and reproductive biology <strong>of</strong>Brochiaria.. ln J.W. Miles, B.L Maoss, andCB. do VoDe (eds.), Brachiaria: Biology,Agronomy, and Improvement. (oli,Co/ombia and (ampa Grande, Brazil: ClATand (NPG(-EMBRAPA. Pp.147-163.Volle, CB., J R. Valerio, S. Colixlo, A. O.Barcellos. 1997. Characteristics <strong>of</strong> selectedgenotypes <strong>of</strong> Brachiar;a for Brazilianposlures. In Proc. XVIII Int. Grassl. (ongr.,Conadian Forage (ouncil, 1999. Winnipeg,(anada. pp.l/81-l/82. CD-ROM.v.l,section 1. 10 1358.Voigt, P.W., and E.C Bashaw. 1972. <strong>Apomixis</strong>and sexualily in fragrostis wrvu/o. (ropSci. 12: 843-47.Voigt, P.W., and B.L Burson. 1992. <strong>Apomixis</strong> infrogrostis. In J.H. 8gin Jr. and J.P. Miksme{edl.}. PrO(. <strong>Apomixis</strong> Workshop, Atlonta,Georgia, USA. February 11·12, 1992.USDA/ARS. ARS-l04. Pp. S-11.Young, B.A., R.T. Sherwood, and H. Boshaw.1979. CIeored-pistyl ond thick-sectioningtechniques for detecting aposporousapomixis in grosses. (an. 1. Bot, S7:166S-72.
Transfer <strong>of</strong> <strong>Apomixis</strong> throughWide Crosses<strong>Chapter</strong> 1 1YVES SAVIDANIntroductionInterspecific hybridization has been usedextensively to transfer agronomicallyimportant genes that control resistances todiseases and insect pests. Recent advances intissue culture, especially in molecular biology,have further widened the scope <strong>of</strong> alien genetransfer-and the outlook for widehybridization in crop improvement seemsmore promising than ever. But does thatoutlook also apply to the transfer <strong>of</strong> sequencesinvolved in plant reproduction, especiallythose involved in apomixis?The interface between conventionalcytogenetical approaches and new moleculartechniques makes the "conventional" widecross approach very competitive when the traitis simply controlled and the gene(s) to betransferred is (are) available in a species thatbelongs to the secondary gene pool. Thegenetic analyses reviewed by Savidan (2000)and Sherwood (Chap. 5) suggest that apomixisa good candidate and <strong>of</strong>fer support for theongoing wide cross projects. Such projects haveencountered unexpected difficulties, andseveral papers have questioned the ultimatelikelihood <strong>of</strong> transferring apomixis to any crop.Nevertheless, knowledge gathered through thequest for wide crossing apomixis into usefulcrop species, which relates to the geneticcontrol, transmission, and expression <strong>of</strong> thetrait (Grimanelli et aI., Chap. 6), has provenextremely valuable for those investigatingother approaches. Accordingly, three paths arenow being pursued in the effort to introduceapomixis into major crops: (i) the widehybridization, (ii) the identification, isolation,and manipulation <strong>of</strong> sequences from wildapomicts, and (iii) the creation <strong>of</strong> an apomicticreproduction de novo, from individualmutations (Grossniklaus, Chap. 12; Praekeltand Scott, Chap.13). In this chapter, I reviewprogress to date and the problems or questionsthat have emerged from work aimed at widecrossing <strong>of</strong> the apomictic trait.Scientists have tried for decades to use widecrosses to transfer the apomixis trait intovaluable food crops, including wheat, maize,and pearl millet. The first attempt involvedmaize and was initiated approximately fortyyears ago (Petrov et al. 1979, 1984). Crossing atetraploid maize (2n =4x =40) with a tetraploidTripsacum dacty/aides (2n =4x =72), the Russianscientists successfully produced maizeTripsacum Fls and BC lhybrid derivatives that,according to progeny tests, reproducedapomictically. The BC lplants combined 20maize chromosomes with one complete set (18)<strong>of</strong> Tripsacum dacty/aides chromosomes. Efficienttechniques for evaluating chromosomenumbers, embryo-sac analysis, etc., were notavailable, making screening <strong>of</strong> large numbers<strong>of</strong> progenies for apomixis difficult.Consequently, little progress was made in thistransfer effort. Recently, the Russian materialswere transferred to the United States, andintrogression efforts were reinitiated. Animportant piece <strong>of</strong> information generated by
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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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202 UeIGfo...ll...Two additional po
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204 Uel G.....ilcmDiboll, A.G., and
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206 UeRG'....ild...--.1974. Reprodu
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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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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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