Chapter 5 Genetic Analysis of Apomixis - cimmyt

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4 Gary H. Toe"'...breeding: the evolutionary phase, in whichvariable populations are produced, and theevaluation phase, in which desirablegenotypes are selected.Variability, at the heart of the evolutionaryphase, traditionally has been created byhybridization and to a lesser extent bymutations. Wide hybridization throughembryo rescue or somatic hybridization,somaclonal variation, and genetic engineeringare biotechnology tools that can dramaticallyexpand the range of variability available tobreeders. Genetic engineering, especially,should make the process of generatingdesirable variability much more predictableand help obtain other goals that are beyondthe reach of conventional techniques.Meanwhile, the evaluation phase will becomemuch more efficient through the use of thefollowing biotechnology tools: anther cultureto produce doubled haploids and eliminatedominance variance; molecular maps andmarkers of the crop genome to tag and followthe inheritance of genes for important traits,particularly quantitative traits and those thatare difficult to score; and molecular geneticmaps and markers of pests and pathogens thatcan be used to characterize and monitorpopulation structures and dynamiCS, therebypromoting more effective selection anddeployment of resistant plants.The international agricultural research system,which has been so successful at producingimproved varieties for developing countries,is itself evolving and adding new institutions(see Tables 1.1 and 1.2) to take advantage ofthese new tools. It is drawing more on resultsgenerated by fundamental and strategicTable 1.1 International Agricultural Research CentersInternational CenterClAT(IfORClMMYT(IPICARDAIHARM'I(RAFI(RISATIFPRI*lilAIlRIIPGRIIRRIISNAR'IWMI*WARDA(entro Internocional de Agricultura Tropical(enter for Internafional Forestry Research(entro Internacional de Mejoramiento de Maiz yTrigo(entro Internacional de 10 PapaInternafional (enter for AgriculturalResearch in Dry AreasInternational (enter for Uving Aquatic ResourcesInternational (enter for Research in AgrofarestryInternational (rop Research Institutefor the Semi-Arid TropicsInternational Food Policy Reseorch InsfitufeInternational Institute for Tropical AgricultureInternational Uvestock Res. InsfitufeInternational Plant Genefic Resources InstituteInternafional Rice Research InstituteInternational Service for NafionalAgricultural ResearchInternational Water ManagementlnstituleWest Africa Rice Development Association• !CLARM, IFPRI, ISHAR, and IWMI do not directly handle plant research programs.Crops(assava, field beans, riceForestryMaize, wheat, trificalePotatoes, sweet potatoesWheat, barley, lenfils, chickpeaForestry, tree cropsSorghum, pearl millet,groundnut, pigeon pea(assava, yams, cowpea, maizeForagesRiceRiceLocation(olombiaIndonesiaMexicoPeruSyriaPhilippinesKenyaIndiaUSANigeriaKenyaIta~PhilippinesThe NetherlandsSri lanka(6te d'ivoire
Feediog the World ill the 21st Century: Pl••t Breeding. Biotedl.oIogy, .od tbe Pot••tial Role .1 ~pamixls 5research institutions and exploring new waysof gaining access to proprietary technologies.Some of the key institutions that make up thissystem in the era of biotechnology are notedin Figure 1.2. Their work includes• fundamental research conductedprimarily in advanced researchuniversities and institutes that expandsthe knowledge base on plants, insects, andmicrobes, and their interactions with oneanother and with their environment;• strategic research, conducted primarily atagricultural universities, nationalagricultural research institutes, incorporations, and increasingly at theinternational agricultural research centers(IARCs), which generates new andstrengthens existing technologies for cropgenetic improvement;• applied research (including germplasmcollection and evaluation), conductedprimarily at IARCs and national cropbreeding institutions, oftencollaboratively, which generates newbreeding lines;• adaptive research, conducted primarily atthe national level, which combines elitebreeding lines with traditional varieties toproduce improved finished varieties thatare well-suited to local needs andconditions; and• seed multiplication and delivery ofimproved varieties, usually by nationalagencies, local farmers, non-governmentalorganizations (NGOs), and increasinglythrough market mechanisms.The lARCs with plant research programs (seeTable 1.1) have the mandate and primaryTable 1.2 Institutions facilitating the application of biotechnology to international agricultureISAAA - International Service for Acquisition of Agri-Biotech Applications, Ithaca, New Yark, USA, is a not-for-profitinternational organization committed to the acquisition and transfer of proprietary agricultural biotechnologies fromthe industrial countries for the benefit of the developing world. It assists in identifying biotechnology needs andopportuni~es, evaluates the availability of proprietary lechnologies, serves as an "honest broker" that malches needswith available technology, and when necessary mobilizes the financial resources required to implement brokeredproposals.ILTAB . International Laboratory for Tropical Agricultural Biotechnology, SI. Louis, Missouri, USA, is a unit of theDan forth Plant Science (enter. Technology is transferred first from the center, where scientists are engaged inpioneering work on the development of disease resistant plants, to five ILTAB scientists and fellows from developingcountries. These scientists use the technology to produce new sources of disease resistance in tropical crops includingcassava, rice, sweet potato, and yam.CAMBIA - (enter for the Application of Molecular Biology to International Agriculture, (anberra, Australia is aresearch and technology transfer organization committed to the applica~on of biotechnology to internationalagriculture. It specializes in producing inexpensive biotechnology tools that can be effernve~ u~lized in developingcountries.ICGEB - International (enter for Genetic Engineering and Biotechnology was established by the United Na~onsIndustrial Development Organization. It has headquarters and information gathering and dissemina~on facili~es inTrieste, Italy, and agricultural biotechnology research and training facilities in New Delhi, India.ABSP . Agricultural Biotechnology for Sustainable Productivity, East Lansing, Michigan, USA, is a projectheadquartered at Michigan Slate University and funded by USAID. It is a unique bilateral program in that it supportsresearch at and technology transfer from public and corporate sector crop research insntu~ons to developing countries.IBS . Intermediary Biotechnology Service, The Hague, Netherlands, is a unit of the International Service for Na~onalAgricultural Research (ISNAR). It provides national agricultural research agencies with informa~on, advice, andassistance to help strengthen their agricultural biotechnology capacities and to enable them to establish collaborativearrangements with international biotechnology programs.BA( . Biotechnology Advisory (enter, Stockholm, Sweden, a unit of the Stockholm Environmentlns~tuIe, is anindependent resource for impartial biosafety advice. It was established to help developing countries assess the possibleenvironmental, health, and socioeconomic impacts of proposed biotechnology introductions.
Page 2 and 3: (over illustration:Pictured is an i
Page 4 and 5: Institut de Recherche pour Ie Devel
Page 6 and 7: iv33 Outlook33 References35 Appendi
Page 8 and 9: vi131 Screening Procedures: Advanta
Page 10 and 11: VIIITables4 Table 1.15 Table 1.29 T
Page 12 and 13: xAcknowledgmentsWe are grateful to
Page 14 and 15: xiifood crops such as maize, wheat,
Page 16 and 17: 2 Gary H. T....i.....much food as i
Page 20 and 21: 6 Gary H. T....it....The potential
Page 22 and 23: Chapter 2Apomixis and the Managemen
Page 24 and 25: 10M.. Be,th.dcategories n + nand 2n
Page 26 and 27: 12 J.&eo BertIoaodtwo tetraploid sp
Page 28 and 29: 14 JoG.. B.rth""dTable 2.8 Distribu
Page 30 and 31: 16 M......thodhexaploidy through 2n
Page 32 and 33: 18 JI&.. lertIoaodheterozygous cond
Page 34 and 35: 20 JM Iertltaodmarkers to retain th
Page 36 and 37: 22 JoA•• BerthudFurthermore, if
Page 38 and 39: Chapter 3Classification of Apomicti
Page 40 and 41: 26 (llarIesf.(r••3) The Ixeris-
Page 42 and 43: simultaneous division of the proxim
Page 44 and 45: 30 (\aries f. Crao.differentiating
Page 46 and 47: 32 CWIts F. C,..haploids from norma
Page 48 and 49: 34 cales F. (,...Campbell, C. S., C
Page 50 and 51: 36 GaOO F. er...media. There may al
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Page 54 and 55: 40 CWIes F.
Page 56 and 57: 42 C"Ie
Page 58 and 59: Chapter 4Ultrastructural Analysis o
Page 60 and 61: 46 T.""". N. Naomo•• ..dJ...·P
Page 62 and 63: (Figure 4.2a,b,c). Their cytoplasm
Page 64 and 65: 50 Tomaro N. Naurnovo ond Jeon-Phil
Page 66 and 67: 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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190 Ud e;,....1Ia..Arabidopsis, unp
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192 Ueli Gro".llae,system to identi
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194 UeI Gm..ikIonsystem developed b
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196 Ud Gro...ild...concentrate our
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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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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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