Chapter 5 Genetic Analysis of Apomixis - cimmyt

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234 T1lo.... Dr......... Jo~. G. (..-.. ood Yv•• Sovida.Table 14.2 Examples of patents linked with the engineering of the apomixis trait in sexual crops.Sources: Intellectual Property Network (hflp://www.delphion.com). European Patent Office (http://ep.dips.org/dipsl, and Bicknelland Bicknell (1999).Apomixis technologyPatent number'(Publication date)Breeding strategiesW08900810(Feb. 9. 1989)CN1124564(June 19. 1996)US5710367(Jon. 20.1998)W0971 0704(Sep. 22, 1998)W09833374(Aug. 6, 1998)W0007434(Feb. 17,2000)Stimulation of apomictic reproductionEP0127313(Dec. 5, 1984)SU 1323048(Ju~ 15, 1987)US4818693(April 4. 1989)US5840567(Nov. 24, 1998)Title (and content)De noyo synthesis of apomixis (genes and promoters)W09743427(Nov. 11, 1997)W09808961(March 5, 1998)W09828431(July 2, 1998)US5792929(Aug. 11. 1998)W09836090(Aug. 20, 1998)W09837184(Aug. 27, 1998)US5907082(May 25. 1999)W09935258(July 15, 1999)W09953083(Oct. 21, 1999)W0024914(Moy 4, 2000)Asexual induclion Df heritoble mole sterility and apomixis in plants(use of mole sterility factors).Hybrid vigor fixing breeding process for rice opomixis(breeding ond selection strategy).Apomictic moize (intragression of apomixisfrom Tripsawmlo moize).Apomixis for producing true-breeding plont progenies (introgressionof apomixis from Pennisetum squamulatum to cullivars).Methods for producing apomicitic plants(breeding program).Novel genetic material for transmission intomaize (introgression of apomixis from Tripsawm).The production of haploid seed, of doubled haploids ond ofhomozygous plant lines therefrom (causing apomixis by applyingon apomixic agent).Stimulator of floral apomixis(no file available).Methods and materials for enhanced somalicembryo regeneration in the presence of auxin.Simplified hybrid seed production by latent diploid parthenogenesis andparthenote cleavage (induced by controlled environmental conditions).Production of apomictic seed (using a SERK gene forembryogenic potential).Endosperm and nucellus specific genes, promoters anduses thereof.Transcriptional regulation in planls(using a meiosis specific promoter).Plants with modified flowers (modifying flower cells aftertransformation with foreign DNA).Meonsfor identifying nucleotide sequencesinvolved in apomixis (isolation and modification of sexual genesfor the expression of apomixis in Gramineae).Leafy cotyledon 1genes and their use (using embryo specific genesand their promoters).Ovule-specific gene expression(using ovule·specific genes).Nucleic acid markers for apospory-specificgenomic region (from the genus Paspalum).Seed specific polycomb group gene andmethods of use for same (using repressors of embryo andendosperm development).Apomixis conferred by expression of SERKinteracting proteins (see above W097434271.• WOo US. EP. CN and SU refer 10 World polenls, US-. European. Chinese ond former Sowjet Union polents.Applicant{s)Maxell Hybrids INCChen J.USDAUSDAUniversity of Utah StaleEubanks M.w.Rohm & HaasPollav Selskokhaz IGNikitskijPGSUniversity of CaliforniaNovartis and inventorsDaan. D.N.P., Olsen,O.-A. and Unnestod, CJohn Innes CentreInnov. LTD and inventorsPGSIRD and CIMMYT-ABCUniversity of CaliforniaUniversity of CaliforniaUniversity of GeorgiaResearch Found. INCCold Spring Harbor Lob.Novar1is
Go..'k 1091....i.9 .1 Aporo" io S....ICr.ps: A(,ilkal A..........'.I ... Apomlxi. Tech.logy 235gametophyte (Grossniklaus, Chap. 12; Cordtsand Dresselhaus, unpublished results).Through the use of mutant approaches(Vollbrecht and Hake 1995; Drews et a!. 1998;Yang and Sundaresan 2000; Grossniklaus,Chap. 12; Praekelt and Scott, Chap. 13), we cananticipate that many more genes involved infemale gametophyte development will soon beisolated. Gene trap screens such as T-DNAinsertional mutagenesis, transposonmutagenesis, and enhancer detection(Grossniklaus, Chap. 12) are very powerfulmolecular tools for isolating the correspondinggenes and!or their promoters from sexualmodel plants like maize and Arabidopsis.Further tissue!cell-specific genes and theirpromoters will be isolated by transcriptprofiling methods (e.g., Liang and Pardee 1992;Welford et al. 1998; Matsumura et a!. 1999) andfrom tissue!cell-specific cDNA libraries (e.g.,Dresselhaus et a!. 1994; Diatchenkoetal. 1996).Initial attempts have been made to comparegene expression profiles between sexual andapomictic lines within the same species. A fewgenes that are specifically expressed in theovules of either sexual or apomictic lines wereisolated (Vielle-Calzada et al. 1996b). Thesegenes may eventually be useful tools forinducing apomictic development in sexuallines or sexual development in apomictic lines.Parthenogenetic embryogenesis fromunreduced eggs is the next required step forsuccessfully engineering the apomixis trait.Whether this will occur spontaneously oncethe egg is diploid has yet to be shown. Quarinand Hanna (1980) found that doubling a sexualdiploid Paspalum line generated a tetraploidthat was facultative aposporous, thusunreduced egg cells developed parthenogeneticallyinto embryos. Spontaneousparthenogenetic development was observed ata low frequency in maize (Chase 1969; Bantinand Dresselhaus, unpublished results). Wheatlines have been described that produced up to90% parthenogenetic haploids (Matzk et al.1995). Very little molecular data concerningparthenogenesis are available for higherplants. One protein (a-tubulin) was identifiedwhose expression is associated with theinitiation of parthenogenesis in wheat (Matzket a!. 1997). And auxin (2,4 D) treated sexualeggs from maize can be triggered to initiateembryo development at a low frequency'r.(Kranz et al. 1995), however, the molecularmechanism is not understood. Three geneswere used to successfully initiate the formationof embryo-like structures on vegetative tissue(lec1: leafy cotyledon 1, Lotan et a!. 1998; andbbm1: baby boom1, Boutilier et a!., unpublishedresults) or to enhance the rate of somaticembryos in culture (SERK1: somaticembryogenesis receptor-like kinase 1, Hecht et a!.,unpublished results), respectively. It remainsto be demonstrated whether these genes arealso useful for inducing embryo developmentin reproductive cells.Parthenogenesis may also arise as a functionof timing, taking into account thatparthenogenetic embryogenesis is usuallyinitiated before anthesis. In contrast to sexualeggs, parthenogenetic eggs (e.g., Pennisetumciliare and wheat) contain ample amounts ofribosomes and polysomes and a large numberof cristae in mitochondria, thus suggesting ahighly active metabolic status prior topollination (Naumova and Vielle-Calzada,Chap"4; Naumova and Matzk 1998). Incontrast to sexual eggs, degeneration ofsynergids in aposporous Pennisetum ciliarefemale gametophyte was precocious andrapid. In addition, a complete cell wall aroundthe eggs was already generated before thearrival of the pollen tube (Vielle et al. 1995). Inmaize, zygotic gene activation (ZGA), theswitch from maternal to embryonic control ofdevelopment, occurs soon after fertilization(Sauter et al. 1998; Dresselhaus et al. 1999;Bantin and Dresselhaus, unpublished).Precocious expression of zygotic genes beforepollination!fertilization could thus eventually
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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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Page 226 and 227: Chapter 13Induction of Apomixis inS
Page 228 and 229: 214 Uta Praebll aod Rod ScottCrosse
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Chapter 5 Genetic Analysis of Apomixis - cimmyt

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