Fertilization in Higher Plants: Molecular and Cytological Aspects

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Author: Val Raghavan

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1MBUFϒ

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Ϗώϕ

1MBUFϓ

1MBUFɬ 'JHBoF'BUFNBQPGUIFFNCSZPOJDSPPUPG"SBCJ EPQTJTB(MPCVMBSFNCSZPC)FBSUTIBQFEFNCSZPJOXIJDI BMM DFMMT PG UIF JODJQJFOU SPPU NFSJTUFN BSF QSFTFOU 3F QSJOUFE GSPN WBO EFO #FSH FU BM /BUVSF m D$FOUSBMSFHJPOPGUIFSPPUNFSJTUFNPGBNBUVSFFNCSZP TIPXJOHUIFQPTJUJPOPGJOJUJBMTGPSBMMUJTTVFMBZFSTTVSSPVOE JOHUIFDFOUSBMDFMMTE.BHOJêFEWJFXPGUIFDPSFNFSJTUFN TIPXOJOD'JYFEOVNCFSTPGEJĒFSFOUJOJUJBMTGPVOEJOUIF NBUVSFFNCSZPBSFJOEJDBUFEF%JWJTJPOTPGUIFJOJUJBMTUPSF FTUBCMJTIJEFOUJDBMMZHSPVQFEJOJUJBMTBOEEFSJWBUJWFT%BTIFE MJOFêSTUEJWJTJPO EPUTTFDPOEEJWJTJPO TUSJQFEUIJSE SBEJBM EJWJTJPO 3FQSJOUFEGSPN%PMBOFUBM %FWFMPQNFOU m

1MBUFɭ 'JHBoS&YQSFTTJPO PG UIF 4$"3&$308 4$3 HFOF EVSJOH FNCSZPHFOFTJT JO "SBCJEPQTJT BmD &BSMZ HMPCVMBSTUBHF FNCSZPT EmG -BUF HMPCVMBSTUBHF FNCSZPT HmJ 5SJBOHVMBSTUBHF FNCSZPT KmM .JEIFBSU TIBQFETUBHFFNCSZPTNmP5PSQFEPTIBQFETUBHFFNCSZPT QmS )ZQPDPUZM BOE SPPU SFHJPO PG OFBSMZ NBUVSF FNCSZPT B E H K N Q-POHJUVEJOBMTFDUJPOTPGXJMEUZQFFNCSZPT C F I L O R4FDUJPOTPGFNCSZPTGPMMPXJOHJOTJUVIZCSJEJ[B UJPOXJUIBO4$3BOUJTFOTFHFOFQSPCFD G J M P S$POGPDBM JNBHFTPGHSFFOëVPSFTDFOUQSPUFJO ('1 FYQSFTTJPOESJWFO CZUIF4$3QSPNPUFS4DIFNBUJDESBXJOHTPOUIFSJHIUTIPX USBDJOHTNBEFGSPNUIFMFѫNPTUQBOFMTPGUIFQSPHFOJUPSTPG UIFHSPVOEUJTTVF UIFJSEFSJWBUJWFTBOEIZQPQIZTJTDFOUSBM DFMMT4$3HFOFFYQSFTTJPOJTJOEJDBUFECZCMVFTIBEJOHѮF TIJѫPG4$3HFOFFYQSFTTJPOUPUIFJOOFSNPTUDFMMMBZFSBѫFS QFSJDMJOBMEJWJTJPOTJTJOEJDBUFECZBSSPXTѮFCPVOEBSZPG UIF VQQFS BOE MPXFS UJFST PG DFMMT EFMJOFBUFE JO UIF HMPCV MBSTUBHFFNCSZPJTJOEJDBUFECZEBSLMJOFT#BSTEmMʍN NmS ʍN 3FQSJOUFE GSPN 8ZTPDLB%JMMFS FU BM %FWFMPQNFOUm

1MBUFϔ

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1MBUFϕ

1MBUFɮ 'JHBoD1IFOPUZQFT PG UIF NFEFB NFB NVUBOU B 4JMJRVF PG B TFMGFE IFUFSP[ZHPVT NFB.&" QMBOU TIPX JOHBCPSUFETFFETEFSJWFEGSPNGFNBMFHBNFUPQIZUFTDBSSZ JOHUIFNFBBMMFMFC-BUFIFBSUTIBQFEXJMEUZQFFNCSZP D-BUFIFBSUTIBQFEFNCSZPUIBUJOIFSJUFEBNBUFSOBMNFB BMMFMF.PSQIPHFOFTJTJTEFMBZFEBOESFTVMUTJOBMBSHFSUIBO OPSNBM IFBSUTIBQFE FNCSZP UIBU FWFOUVBMMZ BCPSUT #BST B ʍN C D ʍN 3FQSJOUFE GSPN (SPTTOJLMBVT FU BM $VSS0QJO1MBOU#JPMm

1MBUFɮ 'JHEoH4FDUJPOT PG "SBCJEPQTJT PWVMFT USFBUFE XJUIBOUJTFSVNBHBJOTU"(".064MJLF "(-

TIPXJOH UIF MPDBMJ[BUJPO PG "(- QSPUFJO E F 8JMEUZQF QSPFN CSZPBOEHMPCVMBSFNCSZP SFTQFDUJWFMZG8JMEUZQFFNCSZP BUUIFUSBOTJUJPOTUBHFBSSPXIFBETMBCFMFEFOEPTQFSNOVDMFJ H 8JMEUZQF UPSQFEPTIBQFE FNCSZP & &NCSZP &U FOEP UIFMJVN 4TVTQFOTPS#BSTE G HʍNFʍN 3FQSJOUFE GSPN1FSSZFUBM 1MBOU$FMMm

1MBUFϖ

ϏϏώ

1MBUFɯ 'JHB C1IPUPHSBQIT PG BCOPSNBM FNCSZP QIFOP PWVMFT CPUUPN UXP SPXT B :PVOH BOUJTFOTF PWVMFT BOE UZQFTJOB1&*BOUJTFOTFMJOFPG"SBCJEPQTJT-Fѫ1BSUPGUIF FNCSZPT C -BUFTUBHF BOUJTFOTF PWVMFT BOE FNCSZPT GSPN PQFOFE TJMJRVFT XJUI HSFFO PWVMFT IBSCPSJOH OPSNBM FN UIFTBNFQMBOUBTJOB#BSTʍN 3FQSJOUFEGSPN-JBOE CSZPTBOEXIJUFPWVMFTFODMPTJOHBCOPSNBMFNCSZPT SJHIU ѮPNBT 1MBOU$FMMm FNCSZPT FYDJTFE GSPN HSFFO PWVMFT UPQ SPX BOE XIJUF

1MBUFɯ 'JHDoK$POGPDBM MBTFS TDBOOJOH NJDSPTDPQJD JN BHFTPGQSPQJEJVNJPEJEFTUBJOFEFNCSZPTPGNBJ[FNVUBOUT FNC DmF BOEFNC GmK D&NCSZPEBZTBG UFSQPMMJOBUJPOE&NCSZPEBZTBѫFSQPMMJOBUJPOF&NCSZP EBZTBѫFSQPMMJOBUJPONVUBOUFNCSZPTBSFBSSFTUFEBUBO FBSMZ USBOTJUJPO TUBHF EBZT BѫFS QPMMJOBUJPO G &NCSZP EBZTBѫFSQPMMJOBUJPOH&NCSZPEBZTBѫFSQPMMJOBUJPO I&NCSZPEBZTBѫFSQPMMJOBUJPO4DVUFMMVNCFHJOTUPGPSN

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ϏϏϏ

1MBUFώύ

1MBUFɧɦ 'JHBoI4FFEEFWFMPQNFOUJOUIFXJMEUZQF B D F H BOE BCBJOTFOTJUJWF BCJ NVUBOU C E G I PG "SB CJEPQTJT &NCSZPT EJTTFDUFE GSPN TFFET PG XJMEUZQF BOE NVUBOUIBSWFTUFEBU B C

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1MBUFɧɧ 'JHB" NPEFM GPS EFWFMPQNFOUBM BSSFTU BOE QSF NBUVSFHFSNJOBUJPOPG"SBCJEPQTJTTFFET%VSJOHUIFHSPXUI QIBTF FYQSFTTJPO PG '64$" '64

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1MBUFώώ

ϏϏϐ

BOENVUBUJPOTJOUIFTFHFOFTMFBEUPQSFNBUVSFHFSNJOBUJPO 1SFNBUVSFHFSNJOBUJPOJTSFHVMBUFECZHJCCFSFMMJDBDJE (" EFQFOEFOU BOE ("JOEFQFOEFOU QBUIXBZT 7JWJQBSZ BOE QSFNBUVSF HFSNJOBUJPO JOTJEF UIF TJMJRVF PDDVS JO FNCSZP HSPXUI BSSFTU BOE FNCSZP EPSNBODZ EPVCMF NVUBOUT 3FE BOEHSFFOCPYFTJOEJDBUFHFOFFYQSFTTJPOBOEGVODUJPO 3F QSJOUFEGSPN3B[FUBM %FWFMPQNFOUm

ϏϏϑ

1MBUFώϏ 1MBUFɧɨ 'JHBoGʃ(MVD VSPOJEBTF (64 FYQSFTTJPO JO UIF FOEPTQFSN OVDMFJ PG USBOTHFOJD PWVMFT PG "SB CJEPQTJT IBSCPSJOH '&3 5*-*; "5*0/*/%&1&/ %&/5 4&&%4 '*4 (64 DPOTUSVDUT B C 6O QPMMJOBUFE PWVMFT TIPXJOH (64 BDUJWJUZ JO UIF DFOUSBM DFMM OVDMFVT DmG (64 BD UJWJUZ JO FOEPTQFSN OVDMFJ GPSNFE JO PWVMFT GPMMPXJOH EPVCMF GFSUJMJ[BUJPO D OV DMFJ E OVDMFJ F OVDMFJ G OVDMFJ #BST NN 3FQSJOUFE GSPN -VP FU BM 1SPD /BUM "DBE 4DJ 64"m

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1MBUFώϐ

ϏϏϒ

1MBUFɧɩ 'JHB C%JBHSBNT TIPXJOH UIF FĒFDU PGHMPCBM%/"IZQPNFUIZMBUJPOPOQBSFOUBMJN QSJOUJOHJO"SBCJEPQTJTB'PSNBUJPOPGFHHBOE TQFSNѮFFOEPTQFSNGPSNFEJOUIFXJMEUZQF BѫFSEPVCMFGFSUJMJ[BUJPODPOUBJOTBSBUJPPGUXP NBUFSOBMHFOPNFTUPPOFQBUFSOBMHFOPNF*OUIF NBUFSOBM HFOPNF NBUFSOBMTQFDJêD JNQSJOUFE HFOFTBSFBDUJWF XIFSFBTQBUFSOBMTQFDJêDHFOFT BSF SFQSFTTFE *NQSJOUFE HFOFT DPOUSJCVUFE CZ UIF QBUFSOBM HFOPNF IBWF B DPNQMFNFOUBSZ FY QSFTTJPO QBUUFSO 8IFO NBUFSOBM HFOPNFT BSF DPOUSJCVUFE CZ B .&5):- 53"/4'&3"4& .&5 BT QBSFOU UIF QBUFSOBMTQFDJê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êD HFOFT SFTVMUJOH JO QIFOPUZQFT TJNJMBS UP UIPTF QSPEVDFE CZ QBSFOUBM JNCBMBODF 3F QSJOUFE GSPN "EBNT FU BM %FWFMPQNFOU m

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ϏϏϓ

1MBUFώϑ

1MBUFɧɪ 'JHB C*O TJUV IZCSJEJ[BUJPO FYQSFTTJPO PG USBO TDSJQUT PG UIF &/%041&3. &/% HFOF PWFS UIF OV DFMMBS QSPKFDUJPO DFMMT PG CBSMFZ FOEPTQFSN B 5SBOTDSJQU FYQSFTTJPOJTJOJUJBUFEJOUIFGSFFOVDMFJBCPWFUIFOVDFMMBS

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1MBUFɧɪ 'JHDoF*O TJUV IZCSJEJ[BUJPO TFDUJPOT PG NBJ[F LFSOFMXJUIBQSPCFTQFDJêDGPSUIF#"4"- &/%041&3. 53"/4'&3 -":&3 #&5- HFOF D 4FDUJPO PG B LFSOFM BU EBZT BѫFS QPMMJOBUJPO ѮF TJHOBM FYUFOET PWFS BU MFBTU UISFFDFMMMBZFSTJOUIFCBTBMFOEPTQFSNE4FDUJPOPGBLFSOFM

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1MBUFώϒ

ϏϏϔ

HSPXUIXJUIFBSMZTUBHFTPNBUJDFNCSZPT TFFOJOTFDUJPOT PODPUZMFEPOTEBZTBѫFSDVMUVSF*OTFU4PNBUJDFNCSZPT BSJTJOH GSPN UIF DBMMVT USBOTGFSSFE BU EBZT BѫFS DVMUVSF JOBOBVYJODPOUBJOJOHNFEJVNUPBNFEJVNMBDLJOHBVYJO GPSEBZT#BSTBʍN CʍN DNN ENN JOTFUNN

ϏϏϕ

1MBUFώϓ

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