Desarrollo de Un Protocolo Directo de Regeneración de Plantas in Vitro A Partir de Cannabis Sativa L

 

Agscrrlhhl ag ue prltlmlhl anrgmtl ag rg`gegrcmnýe ag phcetcs ne vntrl c pcrr ag Mceecins scvc H. Gxphcetgs ag phãetuhcs? dlrflhl`äc agh agscrrlhhl ag hc rg`gegrcmnýe ag irltgs y envgh ag phlnaäc ag phcetcs rg`gegrcacs Chigrtl @chãe-Ãvnhc3 *, Ga`cr @crmäc-Flrtgc9, Ocndg [rljges9 y Frcemnsml Ocvngr Jgrrcnz9 3 [hlnay [hlnay cea @geld @geldnm nmss ].H., ].H., Mg Mgetr etrl l Gurlpg Gurlpgl l ag Gdprgs Gdprgscs cs Neelvc Neelvcalr alrcs cs ag Vchge Vchgemnc mnc,, [crm [crm Qg Qgme melh lhù` ù`nm nm,, Vc Vchg hgem emnc nc,, Gspc Gspcóc óc,, 9 Ne Nes stu tutl tl We Wenv nvgr grsn sntc tcrn rnl l ag Ml Mles esgr grvc vcmn mnýe ýe y Dgol Dgolrc rc ag hc C`rlanvgrsnaca Vchgemncec, Wenvgrsntct [lhntêmenmc ag Vchêemnc, Vchgemnc, Gspcóc

Hc rg`gegrcmnýe ag irltgs ne vntrl pugag mletrniunr gfmczdgetg c hc dgolrc ag Mceecins scvc H. rgmc rgmchm hmnt ntrc rcet etg. g. Eugs Eugstr trl l li liog ogv vl l u ugg ag agsc scrr rrlh lhhc hcrr ue prlt prltlm lmlh lhl l ch chtc tcdg dget etgg gf gfmn mnge getg tg pcrc pcrc hc rg`gegrcmnýe anrgmtc ne vntrl ag phcetcs ag M. scvc ag angrgetgs gxphcetgs (mlhgaýe, (mlhgaýe, jnplmlhl y jloc vgracagrc) c pcrr ag phãetuhcs ag dlelnml. M. scvc vcrngacags ag aäc mlrtl Fgrndle, Fghnec=9, Fgalrc30 y W]L=3, ouetl mle hc vcrngaca anlnmc ag aäc egutrl Fnelhc. ]g prlicrle angz dganls ag rg`gegrcmnýe, nemhunals prltlmlhls yc puihnmcals, y mldinecmnlegs ag rg`uhcalrgs ag mrgmndngetl ag phcetcs ansgócacs plr ghhls dnsdls. ]g sn`uný hc dlrlhl`äc agh agscrrlhhl agsag hc `grdnecmnýe ag hcs sgdnhhcs jcstc gh agscrrlhhl ag hcs phãetuhcs gercnzcacs. Cagdãs, sg cechnzý gh envgh ag phlnaäc ag hls gxphcetgs y hls rg`gegrcetgs ne vntrl. Hhg`cdls c hc mlemhusnýe ag qug gh jnplmlhl gs gh dgolr gxphcetg pcrc hc rg`gegrcmnýe anrgmtc ne vntrl ag phcetcs ag M. scvc mle gh 1>.17% ag hls gxphcetgs qug rgspleangrle, dngetrcs qug gh mlhgaýe y hc jloc vgracagrc tuvngrle uec rgspugstc agfmngetg mle, rgspgmvcdgetg, 1.0; y ;.19% ag hls ndphcetgs qug agscrrlhhcrle hcs phcetcs. phcetcs. angrgemncs angrgemncs sn`enfmcvcs sn`enfmcvcs getrg hls dganls ag muhvl muhvl gvchucals gvchucals y hcs vcrngacag vcrngacagss gstuancacs. Ge `gegrch, hls dgolrgs dganls ag rg`gegrcmnýe ugrle _GCYNI 9.; (d` / H) y _GCYNI 3.; (d` / H) + ECC ;.;9 ; .;9 (d` / H) mle 66.60% ag jnplmlhls qug rgspleangrle. ]lrprgeagetgdgetg, ]lrprgeagetgdgetg, hls jnplmlhls muhvcals ge dganl sne rg`uhcalrgs agh mrgmndngetl ag hcs phcetcs dlstrcrle uec gxmghgetg rgspugstc (63.71% ag hls jnplmlhls qug rgspleangrle) y gh gercnzcdngetl gspletãegl ag hls rg`gegrcetgs (30.>1%). Hcs phcetcs rg`gegrcacs ne vntrl sg cmhndctcrle slhl 6 sgdcecs agspuîs agh nenmnl agh muhvl. Gh gstuanl ag hc dlrlhl`äc agh agscrrlhhl su`ngrg qug hls irltgs rg`gegrcals sg lrn`nece ge hcs mîhuhcs agh pgrnmnmhl caycmgetgs c hls plhls ag xnhgdc. ]g agtgmtý plhnsldcc ge jnplmlhls y mlhgalegs ag tlacs hcs vcrngacags gstuancacs, y sg rg`gegrcrle phcetcs anphlnags (5 :;%) y dnxlphlnags (mle mîhuhcs anphlnags y tgtrcphlnags). Eugstrl prltlmlhl pgrdntg uec chtc g f mngemnc ag lr`cel`îegsns ge angrgetgs vcrngacags ag M. scvc. Gh jgmjl ag qug ue plrmgetco plrmgetcogg sn`enfmc sn`enfmcvl vl ag phcetcs phcetcs sgce dnxlphlnags dnxlphlnags pugag prlplrmnle prlplrmnlecr cr uec lrdc chtgre chtgrecv cvcc ag agscrr agscrrlhh lhhcr cr plhnph plhnphlna lnags gs ge M. scvc. scvc. Eugstr Eugstrls ls rgsuht rgsuhtcal calss dugstr dugstrce ce qug hc rg`gegrcmnýe ne vntrl anrgmtc pugag jcmgr uec mletrniumnýe sn`enfmcvc ch agscrrlhhl ag dctgrnchgs Mnvcachgs dgolrcals pcrc cphnmcmnlegs dîanmcs. NEQYLAWMMNÝE Mceecins scvc H. (9e 4 9 Ð 4 9;) gs uec gspgmng anmlhgaýegc pgrtgegmngetg c hc cdnhnc Mceecicmgcg uhnzcac pcrc dÿhphgs prlpýsntls (firc, cmgntg, sgdnhhcs mldgsihgs, dganmnechgs, arl`cs) qug mldprgeag vcrngacags ag aäc mlrtl y egutrl. Getrg sus angrgetgs cphnmcmnlegs, su usl ge dganmnec, agrnvcal ag su mletgenal ge mceecinelnags (Mcsmnl gt ch., 9;30), gstã susmntceal ue netgrîs mrgmngetg. Getrg hls mceecinelnags, gh 3>-tgtrcjnarlmceecinelh (QJM) y gh mceecinanlh (MIA)

 

sle `gegrchdgetg hls dãs ciueacetgs ge hc phcetc (Cearg gt ch., 9;36). Nevgs`cmnlegs rgmngetgs jce nelrdcal ag dumjcs prlpngacags crdcmlaneãdnmcs ag hls mceecinelnags, cdphnceal gh usl pltgemnch ag hls mceecinelnags mceecinelnags ge tgrcpncs dîanmcs (Wrnts gt ch., 9;3>) y prldlvngeal gh agscrrlhhl ag vcrngacags vcrngacags dgolrcacs dgolrcacs ag mceecins mceecins mle pgrfhgs pgrfhgs inlquädnmls inlquädnmls gspgmäfmls. gspgmäfmls. C gstg rgspgmtl, rgspgmtl, gh muhvl ne vntrl gs uec jgrrcdngetc ÿh qug sg jc gdphgcal pcrc mldphgdgetcr gh muhvl mlevgemnlech ag mceecins c trcvîs ag hc dnmrlprlpc`cmnýe c `rce gsmchc ag mhlegs ag îhntg sghgmmnlecals (Hctc gt ch., 9;30), gh agscrrlhhl ag vcrngacags plhnphlnags mle envghgs dgolrcals ag dgtcilhntls sgmueacrnls (Dceslurn y Ic`jgrn , 9;302 [crsles gt ch., 9;3>) l trceslrdcmnýe `geîmc ag tgonals el rg`gegrcetgs (Fggegy y [ueoc, 9;;=, 9;37, 9;302 Pcjiy gt ch., 9;3=, 9;30). ]ne gdicr`l, tlacväc chtc ue prltlmlhl ag rg`gegrcmnýe ne vntrl gfmngetg ge hc cdphnc `cdc ag dctgrnchgs `geîmcdgetg anvgrsls agetrl ag hcs gspgmngs. C gstg rgspgmtl, hc rg`gegrcmnýe ag hcs phcetcs gs ue pcsl gsgemnch pcrc hc dcylräc ag hcs tîmenmcs ag muhvl ne vntrl gdphgcacs ge gh ftldgolrcdngetl. Chtcs tcscs ag rg`gegrcmnýe ag phcetcs ne vntrl c pcrr ag dgrnstgdls cpnmchgs y cxnhcrgs yc agscrrlhhcals ag hc phcetc (Ynmjgz-Audcelns gt ch., 3>:62 Hctc gt ch., 9;;>, 9;36c, i), jlocs oývgegs (Hctc gt ch., 9;3;) y hls mlhgalegs (Mjcljuc gt ch., 9;36) yc jce snal rgplrtcals ge M. scvc. ]ne gdicr`l, vcrnls gstuanls sgóchce ue chtl envgh ag rgmchmntrcemnc ag hc rg`gegrcmnýe ag irltgs ne vntrl c pcrr ag angrgetgs tgonals mldl irãmtgcs dcaurc dca urcs, s, mldphg mldphgols ols ag cetgrc cetgrcss y mãh mãhnmg nmgss y jlocs jlocs vg`gtc vg`gtcvc vcss (Jgdpj (Jgdpjnhh nhh gt ch., ch., 3>0:), 3>0:), jlocs, jlocs, jnplmlhls, mlhgalegs y rcämgs ( Dcealhnel y Ycechhn, 3>>>), jlocs oývgegs, pgmälhls, getrgeuals y irltgs cxnhcrgs (Huscrbngwnmz-Ocrznec gt ch., 9;;7), rcämgs, jlocs y tchhls ([hcwuszgwsbn gt ch., 9;;6), mlhgalegs, tchhls y rcämgs (Pngh`us gt ch., 9;;:), mlhgalegs y gpnmlhls (Dlvcjgan gt ch., 9;37), jlocs g jnplmlhls (Dlvcjgan gt ch., 9;36c, i), rcämgs trceslrdcacs ag mãócdl (Pcjiy gt ch., 9;30) y sg`dgetls ag jnplmlhl (]dôbchlvã gt ch. ch., 9;3>). [lr hl tcetl, hc icoc gfmngemnc ag rg`gegrcmnýe ag hls prltlmlhls ag rg`gegrcmnýe ag phcetcs ne vntrl puihnmcals pcrc gstc gspgmng, y su cdphnc vcrncmnýe getrg hls pls y vcrngacags ag gxphcetgs rgprgsgetce ue ndplrtcetg mughhl ag iltghhc pcrc hc cphnmcmnýe agh muhvl ag tgonals ne vntrl pcrc hc dgolrc ag M. scvc. Cagdãs, ge hc dcylräc ag hcs puihnmcmnlegs cetgs dgemnlecacs, sg gvchuý ue pgqugól eÿdgrl ag vcrngacags, qug el rgprgsgetce tlacs hcs suigspgmngs y snstgdcs rgprlaumvls prgsgetgs ge hc gspgmng. Cagdãs, ge hc `rce dcylräc ag gstls gstuanls, muceal hc rg`gegrcmnýe ag hls irltgs ug gxntlsc, sg agscrrlhhý ag dcegrc neanrgmtc c trcvîs ag uec csg prgvnc ag lrdcmnýe ag mchhls, hl qug pugag mldprldgtgr hc faghnaca `geîmcg ag hls rg`gegrcetgs mle Fnechdgetg, rgspgmtl c hcuec phcetc alecetg (Gvces y Ircvl, 3>:62 Ycdärgz-Dlsqugac N`hgsncs-Ceargu, 9;37). mcrcmtgräsmc mldÿe qug sg pugag negrnr ag hl dgemnlecal cetgrnlrdgetg. Hls gstuanls agdugstrce qug gh agscrrlhhl ag hc lr`cel`îegsns ag irltgs ne vntrl ge gstc gspgmng rgqungrg hc canmnýe ag rg`uhcalrgs agh mrgmndngetl ag hcs phcetcs ch dganl ag muhvl. Qgengeal ge mugetc hls jgmjls dgemnlecals cetgrnlrdgetg, gh liogvl ag gstg gstuanl sg mgetrý ge gh agscrrlhhl ag ue prltlmlhl gspgmäfml ag gspgmngs chtcdgetg gfmngetg pcrc hc rg`gegrcmnýe anr anrgmt gmtcc ne vntrl vntrl ag phcetc phcetcss ag M. scvc. scvc. [crc [crc gst gstl, l, gvchuc gvchucdls dls an angr grget getgs gs gxphce gxphcetgs tgs mldl gh mlhgaýe, gh jnplmlhl y hc jloc vgracagrc prlvgengetgs ag phãetuhcs ag muctrl vcrngacags dlelnmcs ag aäc mlrtl dãs uec vcrngaca ag aäc egutrl ag mãócdl anlnml, qug sg gdphgcrle mldl phcetc phc etcss alecet alecetgs. gs. Hls gxphce gxphcetgs tgs sg muhvc muhvcrle rle ge dganl dganlss mle an angrg grgetg etgss rg`uhc rg`uhcalr alrgs gs agh agh mrgmndngetl ag hcs phcetcs y mlemgetrcmnlegs jlrdlechgs litgenacs ag prltlmlhls yc puihnmcals,  ouetl mle mldinecmnlegs mldinecmnlegs ag ansgól prlpnl ag rg`uhcalrgs agh mrgmndngetl ag hcs phcetcs. Gh

 

prlmgsl ag dlrlhl`äc agh agscrrlhhl ag hc lr`cel`îegsns ag irltgs ne vntrl ag mlhgalegs, jnplmlhl jnpl mlhlss y jlocs vgracagrcs vgracagrcs sg sn`uný sn`uný y sg rg`nstrý mle ndã`gegs, ndã`gegs, y tcdinîe tcdinîe sg rg`nstrý rg`nstrý hc aurcmnýe ag mcac uec ag hcs gtcpcs ag agscrrlhhl ag hc lr`cel`îegsns. Cagdãs, aginal c hc chtc ag gstuan gst uanls ls slirg slirg hc plhnsl plhnsldc dccc ge gstc gstc gspgm gspgmng ng y su uhnac uhnaca a pltgem pltgemnch nch pcrc pcrc litge litgegr gr ph phcet cetcs cs plhnphlnags mldl jc lmurrnal ge ltrcs gspgmngs mldl gh tldctg (Vce age Iuhb gt ch., 3>>;), gh pgpnel (Mlhnoe-Jllydces gt ch., 3>>1), l dghýe (Yge gt ch., 9;3=), y dãs rgmngetgdgetg ge igrgeogec (@crmäc-Flrtgc gt ch., 9;9;), hls cutlrgs el slhl gstuancrle hc mcpcmnaca rg`gegrcvc ag hls mlhgalegs, jnplmlhls y jlocs vgracagrcs, snel qug tcdinîe cechnzcrle hc phlnaäc. envgh ag gstls gxphcetgs ouetl mle gh ag hls nevntrlrg`gegrcetgs. DCQGYNCHG] ^ DÎQLAL] Dctgrnch ag hc phcetc y mleanmnlegs ag mrgmndngetl  ]gdnhhcs ag vcrngacags dlelnmcs ag M. scvc ag aäc mlrtl Fgrndle, Fghnec=9, Fgalrc30 y W]L=3,  ouetl mle sgdnhhcs ag hc vcrngaca anlnmc ag aäc egutrl Fnelhc sg gstgrnhnzcrle ge supgrfmng ge 07% (v / v) gtcelhaurne`9dnecea=;s, sg`unal plr nedgrsnýe ge =; ` / H ag EcMhL mle ;.3% (v / v) ag Qwgge 9; aurcetg 97 dne, y fechdgetg hcvcal trgs vgmgs ge c`uc agsnlenzcac ge cutlmhcvg. Wec vgz gstgrnhnzcacs, hcs sgdnhhcs `grdnecrle ge phãsmls ag > md ag anãdgtrl qug mletgeäce dganl ag `grdnecmnýe prgvncdgetg cutlmhcvcal muyc mldplsnmnýe grc 3/9 D] schgs icschgs y vntcdnecs (Durcsjn`g y ]bll`, 3>69) + 3.7% (w / v) scmcrlsc + =.7 ` / H @ghrntgY mle ue vchlr ag pJ ag 7.:. Agsp Agspuî uîss ag hc `grd `grdne necm cmný nýe, e, sg gd gdph phgc gcrl rle e ml mlh hga gale legs gs,, jn jnpl plml mlh hls ls y jl jloc ocss vgra vgraca cagr grcs cs ansgmmnlecacs ag phãetuhcs ag 0 aäcs ag gaca mldl gxphcetgs. Ge gstc gspgmng, gstc gtcpc agh agscrrlhhl ag hcs phãetuhcs gs gqunvchgetg ch gstcanl ag mrgmndngetl gelhý`nml mlanfmcal ge Inlhl`nsmjg Iueagscestcht, Iueagsslrtgecdt y MJgdnmch neaustry (gsmchc IIMJ) plr gh eÿdgrl 33 (Dnsjmjgebl gt ch., 9;30). Hls gxphcetgs sg muhvcrle ge hls angrgetgs dganls agsmrntls ge hc Qcihc 3. Hcs phãetuhcs y hls gxphcetgs sg muhvcrle ge mleanmnlegs mletrlhcacs c 99 » M ³ 3 » M y 6;% ³ 3% ag judgaca rghcvc. Gh ltlpgrälal mlesnsý ge 36 j ag huz y : j ag lsmurnaca. Hc huz ug prlplrmnlecac plr tuils ag anlal gdnslr ag huz (HGA) ag 3: P y uec tgdpgrcturc ag mlhlr ag 6;;; B, qug prlplrmnlecrle 6.;3; hux y >;.37 ·dlh d ∙ 9 s ∙ 3. Hls rgmugetls qug prlaumge irltgs irl tgs y rcämgs rcämgs,, y ue eÿdgrl eÿdgrl ag irltgs irltgs agscrrlh agscrrlhhca hcals ls uel plr mcac mcac uel ag hls ndphcet ndphcetgs gs mlrrgspleangetgs sg mletcrle pgrnýanmcdgetg pgrnýanmcdgetg aurcetg 9 sgdcecs ag muhvl. Agspuîs ag gstl Mle gh gdpl, hls rg`gegrcetgs el sg suimuhvcrle neanvnauchdgetg ge tuils ag vnarnl ag 9.7 md ag anãdgtrl y 37 md ag hcr`l, mletgengeal gh dnsdl dganl ge gh much sg `gegrcrle hls irltgs. Muceal hcs rcämgs grce vnsnihgs, hcs phcetcs mle rcämgs gspletãegcs sg muhvcrle ge dcmgtcs (9 H) mle sustrctl mldgrmnch grhnzcal mldpugstl plr uec dgzmhc ag turic eg`rc, turic `rceuhcac y pgrhntc, mle ue vchlr ag pJ ag 6 y uec mleaumvnaca ag 3 d] / md. Cetgrnlrdgetg, gh dganl `ghnfmcal sg hcvcic munacalscdgetg agsag hcs rcämgs. Agspuîs agh trcsphcetg y aurcetg tlal gh prlmgs prl mgsl l ag cmh cmhndc ndctcm tcmnýe nýe,, gh sustrc sustrctl tl sg dcetge dcetgeäc äc hn` hn`grc grcdg dgetg etg judgal judgal y, als vgmgs vgmgs ch aäc (tgdprcel ge hc dcócec y hug`l ge hc tcrag), hls rg`gegrcetgs rgmniäce puhvgrnzcmnýe lhncr mle c`uc. [crc gvntcr hc agsgmcmnýe, hcs phcetcs pgqugócs sg muirngrle mle rgmnpngetgs ag phãsml y sg gxpusngrle prl`rgsnvcdgetg c hc judgaca cdingetch. Jcstc hc cmhndctcmnýe mldphgtc, hcs phcetcs sg muhvcrle ge mleanmnlegs naîemcs ag tgdpgrcturc, ltlpgrälal y huz agsmrntcs cetgrnlrdgetg. Hcs phcetcs gdphgcacs ge gstg gstuanl sg muhvcrle icol hnmgemnc pcrc gh muhvl ag M. scvc mle fegs ag nevgs`cmnýe, gdnal plr gh Dnenstgrnl ag ]chua, ]grvnmnls ]lmnchgs g N`uchaca ag Gspcóc c

 

trcvîs ag hc C`gemnc Gspcólhc ag Dganmcdgetls y [rlaumtls ag schua (C`gemnc Gspcólhc ag Dganmcdgetls y [rlaumtls ]centcrnls l CGD[]) pcrc [hldgräc y @geýdnmc Hta

Gxpgrndgetls ag lr`cel`îegsns ag irltgs ne vntrl  Mle gh fe ag prldlvgr hc lr`cel`îegsns ag irltgs ne vntrl ge M. scvc, sg muhvcrle mlhgalegs, jnplmlhls y jlocs vgracagrcs ansgmmnlecacs ag phãetuhcs ag 0 aäcs ge dganl ag `grdnecmnýe mle hc dnsdc dnsdc mldplsnm mldplsnmnýe nýe agsmrntc agsmrntc cetgrnlrd cetgrnlrdgetg, getg, gxmgptl gxmgptl plr hc canmnýe canmnýe ag phcetcs phcetcs angrgetgs angrgetgs rg`uhc rg` uhcalr alrgs gs ag mrgmnd mrgmndnge ngetl. tl. Ml Mldl dl pcrtg pcrtg ag gstg gstg gstua gstuanl, nl, eugst eugstrl rl liogv liogvl l ug gvchuc gvchucrr mle eugstrls prlpnls `gelpls hc gfmngemnc ag angrgetgs prltlmlhls puihnmcals ag rg`gegrcmnýe ag irltgs ne vntrl agscrrlhhcals pcrc M. scvc. [lr hl tcetl, sghgmmnlecdls gstuanls ge hls qug sg cechnz cec hnzcrl crle e mle îxntl îxntl an angrg grgetg etgss gxphce gxphcetgs tgs,, mntlq mntlqune unenec necss y cuxnec cuxnecss y sus mlemg mlemgetr etrcmn cmnleg legss rgspgmvcs. Ag gstc dcegrc, prlicdls hls dganls uhnzcals ge ue gstuanl slirg hc mcpcmnaca rg`gegrcvc ag hls mlhgalegs dgancetg hc canmnýe ag anczurýe (QA_) y ãmnal γ-ecchgegcmîml (ECC) ch dganl ag muhvl (Mjcljuc gt ch., 9;36), ue trcicol rghcmnlecal ge Yg`gegrcmnýe ne vntrl ag phcetcs c pcrr ag mlhgalegs y gpnmlhls dgancetg 6igemnhcdnelpurnec (IC[) y ãmnal nealh-=iurnml (NIC) (Dlvcjgan gt ch., 9;37), y ltrl ag jlocs g jnplmlhls c trcvîs ag IC[ y ãmnal 9,1anmhlr anm hlrlg lgelx elxncm ncmîm îml l (9,1-A (9,1-A)) (D (Dlvc lvcjg jgang angtch tch., ., 9;36c) 9;36c).. Canmnl Canmnlech echdg dgetg etg,, sg c`rg`ý c`rg`ý c eugstr eugstrl l prl`rcdc ue prltlmlhl ggmvl y rgmngetgdgetg puihnmcal agscrrlhhcal pcrc hcs igrgeogecs ge hcs qug gh usl ag zgcec rniýsnal (_GCYNI), prlplrmnleý iugels rgsuhtcals, el slhl netgrsls ag lr`cel`îegsns agh irltg, plhnphlnanzcmnýe mle iutchslne ag hls rg`gegrcetgs (@crmäc-Flrtgcgtch., 9;9;). Fnechdgetg, mldl sg scig qug gh agscrrlhhl ag rcämgs y irltgs agpgeag ag hc rghcmnýe mntlqunenec? cuxnec y qug chtls envghgs ag mntlqunenec cvlrgmge hc lrdcmnýe ag irltgs (]bll` y Dnhhgr, 3>702 ]u gt ch., 9;33), prlicdls gh ggmtl ag hls dganls mle uec mntlqunenec mlemgetrcmnýe mlemgetrcmnýe 7; vgmgs dcylr qug gh envgh ag cuxnec, ouetl mle angrgetgs agrnvcals ag cagenec y genhurgc mldl IC[, 6igemnhcdnelpurnegrniýsnal (IC[YNI), QA_, lrmhlrgeurle (1M[[W) y _GCYNI dãs ECC, uec cuxnec mldÿedgetg uhnzcac ge prltlmlhls pcrc hc rg`gegrcmnýe ne vntrl ag irltgs ([hcwuszgwsbn gt ch., 9;;62 Pngh`us gt ch., 9;;:2 Hctc gt ch., 9;3;2 Mjcljuc gt ch., 9;36) y gh gercnzcdngetl ne vntrl

 

ag M. scvc (Huscrbngwnmz-Ocrznec gt ch., 9;;72 Pce` gt ch., 9;;>2 Dlvcjgan gt ch., 9;36c2 [crsles gt ch., 9;3>). Ge hc Qcihc 3 sg agtchhce hcs angrgetgs mldinecmnlegs jlrdlechgs prgsgetgs ge hls angrgetgs dganls ag neaummnýe ag irltgs gvchucals ge gstg trcicol. Dlrflhl`äc agh agscrrlhhl agh prlmgsl ag rg`gegrcmnýe ne vntrl ]g sn`uný y sg rg`nstrý gh prlmgsl ag agscrrlhhl netg`rch ag hc ne`grl`geîmc ne vntrl, agsag hc `grdnecmnýe ag hcs sgdnhhcs jcstc hc cmhndctcmnýe ag hcs phcetcs y sg rg`nstrý gh gdpl egmgscrnl pcrc mcac uec ag hcs angrgetgs gtcpcs agh agscrrlhhl.Dnmrlsmlpnl ag gstgrglzlld ag hcilrctlrnl ]_ ]_EE-6 6 (L[Q (L[QNB NBC C ].r. ].r.h. h.,, [let [letgr grce cenm nmc, c, Ntch Ntchnc nc)) gq gqun unpc pcal al mle mle ue Lpb LpbcY cY Mã Mãdc dcrc rc an an`n `ntc tchh M-J[ M-J[ (L[QNBC].r.h.). Agtgrdnecmnýe agh envgh ag phlnaäc ag hls gxphcetgs y hls rg`gegrcalrgs ne vntrl ]g gvchuý gh envgh ag phlnaäc ag hls mlhgalegs, jnplmlhls y jlocs ag phãetuhcs ag 0 aäcs ag mrgmndngetl ne vntrl pcrc vgrnfmcr su pctrýe plhnsldãml. Hcs muctrl vcrngacags dlelnmcs ag aäc mlrtl Fgrndle, Fghnec=9, Fgalrc30 y W]L=3, ouetl mle hc vcrngaca anlnmc ag aäc egutrl Fnelhc sg cechnzcrle ge gstg gxpgrndgetl. ]g gdphgcrle trgs phãetuhcs prlmgagetgs ag mcac vcrngaca pcrc gstg gescyl. [lr ltrl hcal, tcdinîe sg gxcdnecrle jlocs oývgegs ag phcetcs rg`gegrcacs ne vntrl. ]g agtgrdneý gh envgh ag phlnaäc ag =7 rg`gegrcetgs ne vntrl (30 ag mlhgalegs, 37 ag jnplmlhls y = ag jlocs). Hls eÿmhgls mghuhcrgs ag hls gxphcetgs ansgmcals sg cnshcrle dgmãenmcdgetg. ]g mlrtcrle sgmmnlegs ag cprlxndcacdgetg ;,7 md9 mle uec mumjnhhc ag cgntcr ge uec phcmc ag [gtrn ag vnarnl ag 6 md ag anãdgtrl qug mletgeäc ;,7 dh ag tcdpýe ag hnsns HI;3 (pJ 0,7) (Apllhgŵgh gt ch., 3>:>), y sg nemuicrle aurcetg 7 dneutls. [lstgrnlrdgetg, hc suspgesnýe qug mletgeäc eÿmhgls y rc`dgetls ag mîhuhcs sg fhtrý uhnzceal ue fhtrl MghhQrnms ag =; ·d (]ysdgx, ]cetOustAgsvgre, Gspcóc). Hls eÿmhgls ge gh fhtrcal sg ógrle mle My]tcne WV [hlnay (]ysdgx) y sg nemuicrle aurcetg 7 dneutls. Hc netgesnaca ag ulrgsmgemnc agh jldl`gegnzcal sg dnaný uhnzceal ue cechnzcalr ag phlnaäc MyFhlw ]ysdgx [crtgm @diJ, cechnzceal ch dgels 1.;;; eÿmhgls pcrc mcac dugstrc. Hcs jlocs oývgegs ag phcetcs anphlnags ag tlacs hcs vcrngacags gstuancacs sg uhnzcrle mldl mletrlh. Gh pnml ag mletrlh anphlnag sg gstcinhnzý ge 7; puetls agh vchlr ag netgesnaca crintrcrnl ag hc ulrgsmgemnc ge gh jnstl`rcdc. Ge mldpcrcmnýe mle gstg pnml, sg mldprliý gh lp agh ltrl tgonal gvchucal. Ceãhnsns ag actls [crc agscrrlhhcr ue prltlmlhl chtcdgetg gfmngetg pcrc hc rg`gegrcmnýe anrgmtc ne vntrl ag phcetcs ag M. scvc, cechnzcdls gstcaäsmcdgetg gh ggmtl ag angrgetgs cmtlrgs mldl gh `gelpl, gh gxphcetg y gh dganl ag muhvl ge hc lr`cel`îegsns ag irltgs ne vntrl. [crc mcac cmtlr, hc dganc ag rgspugstc ag hls gxphcetgs sg rgprgsgetc mldl ue plrmgetcog (³ ]G) ge rghcmnýe mle hc mceaca tltch ag gxphcetgs muhvcals. [crc hcs vcrngacags y hls dganls mle hcs dgolrgs tcscs ag neaummnýe ag irltgs nagefmcacs ge gstg gstuanl, tcdinîe sg gvchuý gstcaäsmcdgetg gh eÿdgrl ag irltgs plr gxphce gxphcetg tg qug rgsplean rgspleaný. ý. Hls actls actls rg`nst rg`nstrca rcals ls jcstc jcstc hc sg`uea sg`ueacc sg sgdce dcecc ag muhvl muhvl sg gdphgcrle pcrc gh ceãhnsns gstcaäsml. Cagdãs, sg agtgrdneý gstcaäsmcdgetg gh ggmtl agh cmtlr gxphcetg ge gh envgh ag phlnaäc ag hls rg`gegrcetgs ne vntrl. [crc mcac gxphcetg, hc dganc ag hls rg`gegrcetgs anphlnags y dnxlphlnags sg gxprgsý mldl ue plrmgetcog (³ ]G) ge rghcmnýe mle hc mceaca tltch ag phcetcs sldgacs ch ceãhnsns ag mntldgträc ag uol. ]n inge gh `gelpl agh cmtlr sg rgprgsgetý c trcvîs ag muctrl vcrngacags dlelnmcs ag aäc mlrtl (Fgrndle, Fghnec=9, Fgalrc30 y

 

W]L=3) dãs uec vcrngaca ag aäc egutrl ag mãócdl anlnml (Fnelhc), hls gxphcetgs gvchucals ge gstg gescyl ugrle mlhgalegs, jnplmlhls y jlocs vgracagrcs ag phãetuhcs ag 0 aäcs ag gaca. Hls 3; dganls angrgetgs agsmrntls ge hc Qcihc 3 mlestuygrle gh cmtlr dganl ag muhvl. Mcac cmtlr sg cechnzý uhnzceal ch dgels mneml rîphnmcs inlhý`nmcs. Mcac rîphnmc inlhý`nmc mlesnsc ge uec phcmc ag [gtrn qug mletgeäc = gxphcetgs prlvgengetgs ag = phãetuhcs angrgetgs ag hc dnsdc vcrngaca ge gh mcsl ag jnplmlhls, y 6 gxphcetgs prlvgengetgs ag = phãetuhcs angrgetgs ag hc dnsdc vcrngaca ge gh mcsl ag mlhgalegs y jlocs vgracagrcs. ]g gdphgcrle ue tltch ag 9,16= gxphcetgs ge gstg trcicol (3,;;; ag mlhgalegs, 907 ag jnplmlhls y 3,3:: ag jlocs vgracagrcs). Hc neagpgeagemnc getrg hcs vcrncihgs (prugic ag anc`rcdc ag anstrniumnýe), hc jldlmgacsmnaca jldlmgacsmnaca (prugic ag Icrthg) y hc elrdch elrdchnac naca a (prugi (prugicc ag ]jcpn ]jcpnrlrl-Pnh Pnhb) b) sg gvchuc gvchucrle rle pcrc pcrc hls actls actls prlvge prlvgeng ngetg etgss ag hls gxpgrn gxp grndge dgetls tls.. Acal Acal qug ene`uel ene`uel ag hls trgs mrn mrntgr tgrnls nls sg mudphn mudphný, ý, sg uhnzý uhnzý hc prugi prugicc el pcrcdîtrnmc ag Brusbch-Pchhns sg`unac ag hc prugic ag Pnhmlxle plr pcrgs (p 8;.;7) pcrc gvchucr hc sn`enfmcmnýe gstcaäsmc ag hcs angrgemncs getrg hls cmtlrgs. Gh ceãhnsns gstcaäsml sg rgchnzý uhnzceal gh slwcrg Y (Njcbccea@gethgdce, 3>>6). YG]WHQCAL] Gh Gfgmtl agh `gelpl, gh gxphcetg y gh dganl slirg hc lr`cel`îegsns ne vntrl agh irltg ag M. scvc ]g lisgrvý rg`gegrcmnýe ne vntrl ag irltgs ge tlacs hcs vcrngacags ag M. scvc, pls ag gxphcetgs y dgancac, rgsuhtceal ge ue tltch ag 977 irltgs rg`gegrcals ne vntrl cuequg hcs angrgemncs sn`enfmcvcs (p 8;.;7) getrg hls angrgetgs envghgs ag hls trgs prnemnpchgs cmtlrgs qug ugrle lisgrvcals. Mle rgspgmtl ch cmtlr gxphcetg y su ggmtl Ge gh plrmgetcog ag gxphcetgs qug agscrrlhhcrle irltgs, sg agtgmtcrl agtgmtcrle e angrgem angrgemncs ncs sn`e sn`enfmc nfmcvcs vcs getrg mlhgalegs, mlhgalegs, jnplmlhl jnplmlhlss y jlocs vgracagrcs. vgracagrcs. Ge prldganl, prldg anl, gh jnplmlhl jnplmlhl dlstrý hc dgolr dgolr rgspugstc rgspugstc ge tîrdnels tîrdnels ag rg`gegrcmn rg`gegrcmnýe ýe anrgmtc anrgmtc ag hc phcetc, chmcezceal 1>.17% ag hls gxphcetgs mle lrdcmnýe ag irltgs, sg`unal ag mlhgaýe mle 1.0;% y jloc vgracagrc mle ;.19% (Qcihc 9). Cagdãs, sg lisgrvcrle angrgemncs sn`enfmcvcs getrg hcs vcrngacags. W]L=3 ug hc dgolr vcrngaca gvchucac ge gstg gxpgrndgetl, mle 39.=9% ag sus gxphcetgs gxjningeal lr`cel`îegsns lr`cel`îegsns ag irltgs, dngetrcs qug hc vcrngaca mle ue dgelr plrmgetcog ag rg`gegrcmnýe ug Fnelhc mle slhl 1.69% (Qcihc 9). Fnechdgetg, slhl gh dganl 3 (QA_ ;.1 d` / H + ECC ;.9 d` / H) dlstrý uec mcpcmnaca sn`enfmcvcdgetg dcylr pcrc prldlvgr hc lr`cel`îegsns agh irltg, hl`rceal ue prldganl ag 37.0:% ag hc tcsc ag neaummnýe (Qcihc 9). Acal qug hcs jlocs vgracagrcs dlstrcrle uec mcpcmnaca aîinh pcrc neaumnr hc lr`cel`îegsns anrgmtc agh irltg, y pcrc ghndnecr gh ggmtl eg`cvl qug gstcice c`rg`ceal c hls cmtlrgs ag vcrngaca y dganl, cechnzcdls plr sgpcrcal hls actls ag hls mlhgalegs (Qcihc =) y hls jnplmlhls (Qcihc (Qc ihc 1). Mle rgspgm rgspgmtl tl c hls mlhg mlhgale alegs, gs, sg lisgrv lisgrvcrl crle e an angrg grgemn emncs cs sn` sn`enf enfmc mcvcs vcs ge getrg trg hcs vcrngacags. W]L=3 chmcezý hc tcsc dãs chtc ag lr`cel`îegsns ag irltgs mle >.9>% (Qcihc =), dngetrcs qug Fghnec=9 gxjniný hc tcsc ag neaummnýe ag irltgs dãs icoc mle 9.3;% (Qcihc =). Mle rgspgmtl ch cmtlr dganl, dganl 3 (QA_ ;.1 d` / H + ECC ;.9 d` / H) ug gh dgolr, hl`rceal hc tcsc ag neaummnýe ag irltg dãs chtc mle ue 99.=9% ag gxphcetgs qug rgspleangrle (Qcihc =). Gh dganl ; (sne rg`uhcalrgs agh mrgmndngetl ag hcs phcetcs) y gh eÿdgrl > (_GCYNI 3 d` / H + ECC ;.;9 d` / H) ugrle hls pglrgs trctcdngetls, sne ene`ÿe gxphcetg qug dugstrg rgspugstc ge tîrdnels ag lr`cel`îegsns agh irltg (Qcihc =).

 

Mle rgspgmtl rgspgmtl c hls jnplmlh jnplmlhls, ls, sg nagefmcr nagefmcrle le angrgem angrgemncs ncs sn`enfmcv sn`enfmcvcs cs getrg hcs angrgetg angrgetgss vcrngacags y dganls gvchucals ge gstg gxpgrndgetl. Eugvcdgetg, W]L=3 ug hc dgolr vcrngaca gvchucac, mle 03.37% ag sus gxphcetgs agscrrlhhceal irltgs (Qcihc 1), dngetrcs qug Fnelhc y Fgrn Fgrndl dle e u ugr grle le hcs hcs vcrn vcrnga gaca cags gs mle hl hlss plrm plrmge getc tcog ogss ag rg`g rg`geg egrc rcmn mnýe ýe dãs dãs ic icol olss mle, mle, rgspgmvcdgetg, =7.19 y =9.96% ag sus gxphcetgs. irltgs rg`gegrcalrgs (Qcihc 1). Ge rghcmnýe mle gh ggmtl agh dganl ge hc lr`cel`îegsns agh irltg, gh dganl eÿdgrl 1 (_GCYNI 9 d` / H) y gh eÿdgrl > (_GCYNI 3 d` / H + ECC ;.;9 d` / H) rgsuhtcrle ge hc tcsc dãs chtc ag neaummnýe agh irltg mle 66.60% ag rgspugstc gxphcetgs, sg`unals ag dganl ; (sne rg`uhcalrgs agh mrgmndngetl ag hcs phcetcs) y dganl 3 (QA_ ;.1 d` / H + ECC ;.9 d` / H) mle, rgspgmvcdgetg, 63.71 y 71.30% ag lr`cel`îegsns ag irltgs (Qcihc 1). Cagdãs, gh eÿdgrl ag irltgs agscrrlhhcals ge mcac uel ag hls gxphcetgs qug rgspleangrle sg cechnzcrle gstcaäsmcdgetg pcrc vcrngacags y dganls mle hcs tcscs ag neaummnýe ag anspcrls dãs nagefmcacs ge gstg gstuanl. Ge gh mcsl ag hls mlhgalegs, mldl hcs vcrngacags W]L=3, Fgalrc30 y Fgrndle, y hls dganls 3 (QA_ ;.1 d` / H + ECC ;.9 d` / H) y : (1-M[[W 3 d` / H + ECC ;.;9 d` / H) angrle gh dgolr tcscs ag neaummnýe ag irltgs, su eÿdgrl ag irltgs plr gxphcetg ag rgspugstc sg mldpcrcrle gstcaäsmcdgetg (Qcihc =). Cuequg el sg gemletrcrle angrgemncs sn`enfmcvcs getrg hcs vcrngacags y hls dganls ge tîrdnels ag eÿdgrl ag irltgs plr mlhgaýe qug rgspleag, Fgalrc30 dlstrý hls dgolrgs rgsuhtcals mle 3.19 irltgs plr gxphcetg qug rgspleaný, dngetrcs qug gh dganl3 (QA_;.1d` / H + ECC;.9d` / H) chmcezý 3.9: hcezcdngetls pgrmltgamlhgale (Qcihc =). Mle rgspgmtl c hls jnplmlhls, agsag hcs vcrngacags W]L=3 y Fghnec=9, y hls dganls ; (sne rg`uhcalrgs agh mrgmndngetl ag hcs phcetcs), 3 (QA_ ;.1 d` / H + ECC ;.9 d` / H), 1 (_GCYNI 9 d` / H) y > (_GCYNI 3 d` / H + ECC ;.;9 d` / H) chmcezcrle hcs dgolrgs tcscs ag lr`cel`îegsns ag irltgs, su eÿdgrl ag irltgs plr gxphcetg ag rgspugstc tcdinîe sg mldpcrý gstcaäsmcdgetg (Qcihc 1). W]L=3 gxjniný hc dgolr rgspugstc ge tîrdnels ag eÿdgrl ag irltgs plr jnplmlhl ag rgspugstc, chmcezceal 3.09 irltgs plr gxphcetg ag rgspugstc (Qcihc 1). Cagdãs, cuequg el sg gemletrcrle angrgemncs sn`enfmcvcs getrg hls dganls cechnzcals, gh dganl 1 (_GCYNI 9 d` / H), sg`unal ag mgrmc plr gh dganl ; (sne rg`uhcalrgs agh mrgmndngetl ag hcs phcetcs), ug gh dgolr dganl gvchucal ge gstg gxpgrndgetl mle 3.6; y 3.71 irltgs, rgspgmvcdgetg. plr jnplmlhl rgspleagalr (Qcihc 1). Dlrflhl`äc agh agscrrlhhl ag hc lr`cel`îegsns ag irltgs ne vntrl ge M. scvc Ge prn prndgr dgr hu`cr, hu`cr, hcs sgd sgdnhnhhcs hcs ag hcs an angr grget getgs gs vcrnga vcrngacag cagss (Fn (Fn`ur `urcc 3C) sg gstgrn gstgrnhnz hnzcrl crle e ge hc supgrfmng. Getrg 91 y 1: j agspuîs ag sgr muhvcacs ge dganl ag `grdnecmnýe, hcs sgdnhhcs mldgezcrle c `grdnecr y gh dgrnstgdl ag hc rcäz cpnmch sur`ný ag hc tgstc (Fn`urc 3I). Mneml aäcs agspuîs ag hc sngdirc ne vntrl, hcs phãetuhcs hnigrcacs ag hc tgstc grce vnsnihgs dngetrcs gdgr`äce (Fn`urc 3M). Ge gh sîpdl aäc agsag hc sngdirc, gh prndgr pcr ag jlocs vgracagrcs sg gxpceaný mldphgtcdgetg (Fn`urc 3A). Muceal hcs sgdnhhcs hhg`crle c gstc gtcpc ag agscrrlhhl, hls gxphcetgs egmgscrnls pcrc mleeucr hls gxpgrndgetls sg lituvngrle c trcvîs ag ue puetl ag ansgmmnýe mruzcac (gmjc ge hc Fn`urc 3A). ]g agsmcrtý gh ãpnmg agh irltg vg`gtcvl rgstcetg uinmcal ge hc pcrtg supgrnlr ag hc phãetuhc (crrlwneFn`urg3G). Ge gstc gtcpc agh agscrrlhhl ag hcs phãetuhcs, hcs jlocs ag mlhgaýe sg ansgmmnlecrle ãmnhdgetg (Fn`urc 9C). Hls prndgrls pcsls ag hc lr`cel`îegsns agh irltg anrgmtl ugrle rãpnacdgetg vnsnihgs ge hc zlec icsch ag hls mlhgalegs qug rgspleangrle uinmcal ge hc pcrtg prlxndch ag hls

 

mlhgaleg mlhga legss agspuî agspuîss ag 1 aä aäcs cs ag muhvl muhvl ne vnt vntrl rl (F (Fn`u n`urc rc 9I). 9I). Als sgdcecs sgdcecs agspuî agspuîss ag hc trcesgrgemnc agh gxphcetg ch dganl ag neaummnýe agh irltg, sg lisgrvcrle rg`gegrcetgs vn`lrlsls qug sur`ge agh ilrag prlxndch ag hls mlhgalegs (Fn`urc 9M). Ge gstc gtcpc ag agscrrlhhl, acal qug hls rg`gegrcetgs chmcezcrle cprlxndcacdgetg ue mgedgtrl ge hc chturc y pcrc gvntcr su mletcmtl mle hc tcpc ag hc phcmc ag [gtrn, hls irltgs rg`gegrcals ne vntrl sg suimuhvcrle neanvnauchdgetg ge tuils ag vnarnl qug mletgeäce gh dnsdl dganl ge gh qug sg `gegrcrle (Fn`urc 9A). We tltch ag 10 mlhgalegs rgspleangrle c hls angrgetgs trctcdngetls gvchucals, `gegrceal ue tltch ag 71 irltgs. Chtgrecvcdgetg, sg mlrtcrle jnplmlhls ag phãetuhcs ag 0 aäcs ag gaca. Jnplmlhls gdphgcals ge gstg gxpgrndgetl dganal hle`ntua ag cprlxndcacdgetg ue mgedgtrl (Fn`urc =C). Hc sgmmnýe trcesvgrsch ag jnplmlhls rgmnîe ansgmcals rgvghý su gstrumturc netgrec, mle angrgetgs mcpcs ag tgonal mldl gpnagrdns, mlrtgzc y dîauhc, ouetl mle hc cusgemnc ag trczcs ag dgrnstgdl (Fn`urc =I). Hls gxphcetgs qug rgspleangrle gxjningrle angrgetgs pctrlegs ag lr`cel`îegsns anrgmtc. Ch`uels ag hls jnplmlhls `gegrcrle slhl ue prndlranl ge hc pcrtg supgrnlr agh gxphcetg qug sg lrn`neý ge hc rg`nýe mgetrch ag hc sgmmnýe (Fn`urc =M), y mleeuý su agscrrlhhl jcstc mlevgrrsg ge uec phcetc rliustc agspuîs ag 9 sgdcecs ag muhvl (Fn`urc =A). Ltrls gxphcetgs angrle hu`cr c ue pcr ag prndlranls ge hc pgrngrnc agh ýr`cel y ge hcals lpugstls (gmjcs ge hc Fn`urc =G). Ge gstg ÿhdl mcsl, juil sntucmnlegs ge hcs qug gh agscrrlhhl getrg cdils prndlranls ug csäemrlel, dnge dngetr trcs cs qu qugg ge lt ltrl rlss mcsl mcsls, s, gh mr mrgm gmnd ndng nget etl l y hc ch chtu turc rc ag cdil cdilss rg`g rg`geg egrc rcet etgs gs ugr ugrle le mlnemnagetgs, mldl sg pugag lisgrvcr ge hc Fn`urc =F. Ge gh mcsl nemlrplrcal ge hc Fn`urc =@, cdils rg`gegrcetgs ne vntrl chmcezcrle cprlxndcacdgetg ue mgedgtrl ag chturc 9 sgdcecs agspuîs agh nenmnl agh muhvl, dngetrcs qug tcdinîe sg lisgrvý ue mrgmndngetl gxmgsnvl ag hc dîauhc (uinmcac ge gh dganl agh gxphcetg). Ge gstc gtcpc agh agscrrlhhl lr`cel`îenml, hls irltgs sg sgpcrcrle ag hls jnplmlhls y sg muhvcrle neanvnauchdgetg ge tuils ag vnarnl qug mletgeäce gh dnsdl dganl ge gh qug sg `gegrcrle (Fn`urc =J). We tltch ag 3=6 jnplmlhls rgspleangrle c hls angrgetgs trctcdngetls gvchucals, prlaumngeal uec mceaca tltch ag 3>; irltgs. Fnechdgetg, tcdinîe sg gstuaný hc mcpcmnaca rg`gegrcvc agh prndgr pcr ag jlocs vgracagrcs ag phãetuhcs ag 0 aäcs. [crc gstl, mcac hc jloc sg ansgmmnleý munacalscdgetg (Fn`urc 1C) y sg muhvý ge hls hls an ang grg rget etgs gs dgan dganls ls ge qu qugg u ugg gv gvch chuc ucal al gstg gstg gx gxpg pgrn rndg dget etl. l. Muce Muceal al sg prla prlauo uol l hc rg`gegrcmnýe ne vntrl, hls prndlranls sur`ngrle sngdprg ag hc icsg ag hcs jlocs, gspgmäfmcdgetg agh rc`dgetl agh pgmälhl uenal ch jloc, mldl gs gh mcsl ag hc Fn`urc 1I, aleag sg pugag vgr uec pgqugóc phãetuhc qug sur`g ag hc zlec ag trcesnmnýe jloc-pgmälhl 3 sgdcec agspuîs agh muhvl. Gh agscrrlhhl agh irltg mleeuý jcstc qug hcs phcetcs rg`gegrcacs chmcezcrle cprlxndcacdgetg ue mgedgtrl ag chturc (Fn`urc 1M). Mctlrmg aäcs agspuîs agh nenmnl agh muhvl, hc gsmnsnýe gxntlsc ag hls irltgs sg rgchnzý y hcs phcetcs rg`gegrcacs sg suimuhvcrle neanvnauchdgetg ge tuils ag vnarnl qug mletgeäce gh dnsdl dganl ge gh qug sg `gegrcrle (Fn`urc 1A). ]lhl mneml jlocs rgspleangrle c muchqungrc ag hls angrgetgs trctcdngetls gvchucals, `gegrceal uec mceaca tltch ag 7 irltgs. Gercnzcdngetl ag gxphcetgs y gercnzcdngetl gspletãegl ag phcetcs agrnvcacs ag jnplmlhl  Cuequg gh prgsgetg gstuanl y sus gxpgrndgetls agrnvcals sg mgetrcrle ge hc lr`cel`îegsns ag irltgs irl tgs ne vnt vntrl, rl, ch`uel ch`uelss ag hls gxphc gxphcetg etgss muhv muhvcal calss agscr agscrrlh rlhhcr hcrle le rcämgs rcämgs ge hu`cr hu`cr ag irltgs irltgs.. Gspgmäfmcdgetg, 9 sgdcecs agspuîs agh nenmnl agh muhvl, gh 3,;>% ag hls jnplmlhls muhvcals agscrrlhhcrle rcämgs vn`lrlscs mle pghls rcanmuhcrgs ge hc zlec negrnlr agh gxphcetg (gmjc ge hc

 

Fn`urc 7C). Gstg dnsdl geýdgel, tcdinîe hlmchnzcal ge hc pcrtg prlxndch agh gxphcetg, sg lisgrvý ge gh ;,3% ag hls mlhgalegs muhvcals 9 sgdcecs agspuîs muhvl ag gxphcetgs (gmjc ge hc Fn`urc 7I). Ag ltrc dcegrc, gh gercnzcdngetl gspletãegl ag hls rg`gegrcetgs ne vntrl slhl tuvl hu`cr ge phcetcs agrnvcacs ag jnplmlhl muhvcacs ge dganls sne rg`uhcalrgs agh mrgmndngetl ag hcs phcetcs, aleag gh 30.>1% ag hls jnplmlhls muhvcals agscrrlhhcrle irltgs ge su pcrtg supgrnlr y rcämgs ge su pcrtg negrnlr. gstcice gercnzcacs gspletãegcdgetg y gstcice hnstcs pcrc mldgezcr gh prlmgsl ag cmhndctcmnýe (Fn`urc 7M). Cmhndctcmnýe ag phcetcs rg`gegrcacs ne vntrl  Acal qug slhl hcs phcetcs rg`gegrcacs c pcrr ag jnplmlhls agscrrlhhcrle ue gercnzcdngetl gspletãegl, hcs phcetcs agrnvcacs gxmhusnvcdgetg ag jnplmlhl sg sldggrle ch prlmgsl ag cmhndctcmnýe. Gh prndgr pcsl mlesnsý ge hcvcr munacalscdgetg gh dganl `ghnfmcal rgstcetg ag hcs rcämgs. Agspuîs ag 9: aäcs ag muhvl ne vntrl, hls rg`gegrcetgs dlstrcrle angrgetgs pctrlegs ag dlrl`îegsns ag hc rcäz mldl sg lisgrvc ge hc Fn`urc 6C, aleag sg pugage vnsuchnzcr `rceags y dgancecs mceacags mlrtcs ag mgrl, ouetl mle uec rcäz prnemnpch rliustc mle ue agscrrlhhl prldne prl dneget getgg ag rcämgs rcämgs sgmuea sgmueacrn crncs. cs. Gh agscrr agscrrlhh lhhl l vn`lrl vn`lrlsl sl agh sns snstgd tgdcc rcanmu rcanmuhc hcrr `crce `crcezý zý hc cmhndctcmnýe gxntlsc ag phcetcs agrnvcacs ag jnplmlhl. Ge gstg puetl, hls rg`gegrcetgs gstcice hnstls pcrc gh trcsphcetg ge dcmgtcs pgqugócs (9 h) mle sustrctl mldgrmnch grhnzcal (Fn`urc 6I), cuequg grc egmgscrnc hc mlhlmcmnýe ag rgmnpngetgs ag phãsml trcespcrgetgs pcrc rgtgegr hc judgaca y gvntcr hc agsgmcmnýe ag hcs phcetcs (Fn`urc 6M). ]ne gdicr`l, agspuîs ag 3 sgdcec ag gxplsnmnýe prl`rgsnvc ag hls rg`gegrcetgs c hc judgaca cdingetch, gh prlmgsl ag cmhndctcmnýe fechnzý y hcs phcetcs agrnvcacs ag jnplmlhl dlstrcrle ue mrgmndngetl vn`lrlsl, mldl sg pugag lisgrvcr ge hc Fn`urc 6A, aleag sg agstcmc ue rg`gegrcetg schuacihg 6 sgdcecs agspuîs agh muhvl muh vl ag jn jnplm plmlh lhls. ls. [crc [crc vgrnfm vgrnfmcr cr gh mlrrgm mlrrgmtl tl agscrr agscrrlhh lhhl l ag hls rg`geg rg`gegrce rcetgs tgs ne vnt vntrl, rl, sg muhvcrle phcetcs cmhndctcacs aurcetg als sgdcecs canmnlechgs y sg plhnenzcrle dceuchdgetg. Mldl sg dugstrc ge hc Fn`urc 6G, hcs phcetcs agrnvcacs ag jnplmlhl dlstrcrle uemnlechnaca sgxuch : sgdcecs agspuîs agh gxphcetg ne vntrl nelmuhcmnýe, mldl pugag agaumnrsg agh jgmjl ag qug hcs lrgs gdgenecs agscrrlhhcrle sgdnhhcs vncihgs agspuîs ag hc plhnenzcmnýe dceuch (nesgrtls ge hc Fn`urc 6G). ]n`ungeal gstg prltlmlhl, gh 3;;% ag hcs phcetcs agrnvcacs ag jnplmlhl qug sg gercnzcrle gspletãegcdgetg sg cmhndctcrle mle îxntl. Gvchucmnýe ag phlnaäc ag gxphcetgs y phcetcs rg`gegrcacs ne vntrl ag M. scvc  Gh ceãhnsns agh envgh ag phlnaäc ag mlhgalegs, jnplmlhls y jlocs vgracagrcs ag phãetuhcs ag 0 aäcs ag gaca ag hcs mneml vcrngacags gvchucacs rgvghý qug slhl hcs jlocs vgracagrcs (vgrag) dlstrcrle ue pctrýe anphlnag, dngetrcs qug hls mlhgalegs (czuh) y hls jnplmlhls (rlol) gxjniný ue pctrýe dnxlphlnag (mle mîhuhcs anphlnags y tgtrcphlnags) (Fn`urc 0). Qlacs hcs vcrngacags gvchucacs ge gstg gxpgrndgetl dlstrcrle gh dnsdl pctrýe plhnsldãml pcrc als phcetcs qug sg cechnzcrle. Hls mlhgalegs y hls jnplmlhls prlauogrle, rgspgmvcdgetg, :9,=7 y :6,60% ag rg`gegrcalrgs anphlnags (Qcihc 7). Ge mucetl c hls rg`gegrcalrgs dnxlphlnags, tcetl hls ag hls mlhgalegs mldl hls ag jnplmlhl y hc nejninmnýe ag hc mcpcmnaca ag `gegrcmnýe ag ghhls, rgspgmvcdgetg, mle 30,67 y 3=,==% ag hcs phcetcs rg`gegrcacs ne vntrl dnxlphlnags (Qcihc 7). AN]MW]NÝE

 

Gh jnplmlhl prgsgetc ue chtl pltgemnch pcrc hc rg`gegrcmnýe anrgmtc ne vntrl ag phcetcs ag M. scvc sne canmnýe canmnýe ag rg`uhc rg`uhcalr alrgs gs agh mrgmndng mrgmndngetl etl ag hcs phcetcs phcetcs ch dganl dganl ag muhv muhvl. l. Cuequg Cuequg sg gemletrcrl geml etrcrle e angrgem angrgemncs ncs sn`enfmcv sn`enfmcvcs cs pcrc tlals hls cmtlrgs cmtlrgs gvchucals gvchucals,, hcs angrgem angrgemncs ncs dãs ndplrt ndp lrtcet cetgs gs ugrle ugrle lisgrv lisgrvcac cacss ge gh pl ag gxphce gxphcetg. tg. Gspgm Gspgmäfm äfmcdg cdgetg etg,, gh jnplml jnplmlhl hl ug sn`enfmcvcdgetg dgolr qug hls mlhgalegs y hcs jlocs ag mletcmtl ag pltgemnch lr`cel`îenml ag irltg, mldl sg pugag mlemhunr agh jgmjl ag qug gh jnplmlhl rgsuhtý ge ue cudgetl ag dãs ag 3; vgmgs ge hc tcsc ag neaummnýe ag irltgs ge mldpcrcmnýe mle gh mlhgaýe. Ge mletrcstg, hcs jlocs dlstrcrle uec plirg mcpcmnaca pcrc prldlvgr hc lr`cel`îegsns agh irltg. [lr ltrl hcal, qug agdugstrc mýdl gh `gelpl cgmtc hc rg`gegrcmnýe ag phcetcs agrnvcacs ag jnplmýhl ge M. scvc, cuequg nemhusl hc vcrngaca ag pglr rgeandngetl gvchucac ge gxpgrndgetls ag jnplmýhl chmcezý ue chtl `rcal ag lr`cel`îegsnshc vcrncmnýe getrg hc tcsc ag lr`cel`îegsns ne vntrl ag hls jnplmlhls prlvgengetgs ag neanvnauls ag hc dnsdc vcrngaca ug rghcvcdgetg icoc ge eugstrls gxpgrndgetls. Gh jgmjl ag qug sg lisgrvcrle vchlrgs icols ag gstcaäsmcs ag vcrncmnýe agetrl ag mcacc vcrng mca vcrngaca aca su`ngr su`ngrgg qug hc vcrncm vcrncmnýe nýe net netrcv rcvcrn crngtc gtchh pcrc pcrc hc mcpcmn mcpcmnaca aca ag rg`geg rg`gegrcm rcmnýe nýe ag jnplmlhl gstã hndntcac ge M. scvc, hl qug mleslhnac gh chtl pltgemnch qug gstg gxphcetg prgsgetc pcrc hc rg`gegrcmnýe ne vntrl ag phcetcs ge gstc gspgmng. Mldpcrceal eugstrls rgsuhtcals mle hc inihnl`rcc rghcmnlecac, cuequg Pcjiy gt ch. (9;30) el puangrle rg`gegrcr phcetcs c pcrr ag mchhls agrnvcals ag jnplmlhl ag mãócdl trceslrdcal, y ]dôbchlvã gt ch. (9;3>) su`nrngrle qug hls jnplmlhls jnplmlhls el sle gxphcetgs gxphcetgs cagmucals cagmucals pcrc hc duhphnm duhphnmcmnýe cmnýe agh mãócdl, qug mcrgmge mcrgmge ag pltgemnch lr`cel`îenml prlicihgdgetg aginal c su icoc mlemgetrcmnýe ag mntlqunenecs cmvcs, vchg hc pgec sgóchcr qug hc rg`gegrcmnýe ne vntrl ag phcetcs c pcrr ag mchhls agrnvcals ag jnplmlhl yc sg jc nelrdcal ge M. scvc (Dcealhnel y Ycechhn, 3>>>2 Dlvcjgangtch ., 9;36c, i)]ne gdicr`l, ge hcs prndgrcs ãirnmcs, hc tcsc ag rg`gegrcmnýe el sg mucefmc y, prlicihgdgetg aginal c hc canmnýe ag rg`uhcalrgs agh mrgmndngetl ag hcs phcetcs ge gh dganl ag muhvl, tlacs hcs csgs ag lrdcmnýe tldce hu`cr ge hc lr`cel`îegsns, hl qug pugag mldprldgtgr hc faghnaca `geîmc ag hls `gegrcalrgs mle rgspgmtl c hls ndphcetgs (Gvcescea Ircvl, 3>:62 Ycdärgz-DlsuDlsu-Ceargu-Dlsmu-Ceargu-Dlsmu Dlsu-Ceargu-Dls mu-Ceargu-Dlsmu-Cearî-Dlsmu-Cea -Cearî-Dlsmu-Ceargu-Dlsmu-Cearî) rgu-Dlsmu-Cearî).. Ge gstg mletgxtl, agig tgegrsg ge mugetc qug, jcstc aleag scigdls, gstg gs gh prndgr nelrdg ag rg`gegrcmnýe ne vntrl anrgmtc ag phcetcs ag M. scvc c pcrr ag jnplmlhls, y gh prndgr trcicol ge gh qug sg prldugvg hc gfmngetg lr`cel`îegsns anrgmtc ag irltgs ne vntrl mle gxphcetgs qug mcrgmge ag dgrnstgdls yc agscrrlhhcals y muhvcals ge dganl sne rg`uhcalrgs agh mrgmndngetl ag hcs phcetcs. Acal qug hc gaca oug`c ue pcpgh mhcvg ge gh pltgemnch lr`cel`îenml agh irltg ag hls gxphcetgs, mldl sg jc agsmrntl ge hls mlhgalegs ag M. scvc (Mjcljuc gt ch., 9;36), prlicihgdgetg ge eugstrl mcsl, gh usl ag jnplmlhls ag phãetuhcs ag 0 aäcs ag gaca mle phng`ugs ag prndgrc dcel mldphgtcdgetg gxpceanals , ouetl mle hc suprgsnýe ag hls rg`uhcalrgs agh mrgmndngetl ag hcs phcetcs ge gh dganl ag neaummnýe ag irltgs, pugag `gegrcr angrgemncs mle hls gstuanls prgvncdgetg puihnmcals slirg hc rg`gegrcmnýe ag phcetcs agrnvcacs ag jnplmlhl ge hc scturcmnýe ag M. C hc nevgrsc ag hl qug lmurrg mle hls mlhgalegs, qug el pugage rg`gegrcr hcs phcetcs ge ue dganl hnirg ag jlrdlecs, eugstrl prltlmlhl pgrdntg uec chtc tcsc ag lr`cel`îegsns ag irltgs ag jnplmlhls ag tlacs hcs vcrngacags prlicacs sne hc canmnýe ag rg`uhcalrgs agh mrgmndngetl ag hcs phcetcs ch dganl ag neaummnýe ag irltgs. Hc rg`gegrcmnýe ne vntrl ag phcetcs c pcrr ag jnplmlhls sne uscr rg`uhcalrgs ag mrgmndngetl ag phcetcs, sg jc nelrdcal ge gspgmngs mldl Mcpsnmud ceeuud H. (Gzurc gt ch., 3>>=) l [cssn  lrc sgtcmgc ag A. M. M. scvc (Vngnrcplaräce gt ch., 9;31). Hls envghgs prgsgetgs ge hls jnplmlhls gstcr neuygeal geag su jlrdlecs mcpcmnaca geaý`gecs ag `gegrcr

 

phcetcs c trcvîs agh muhvl ne vntrl, mldl pugag agaumnrsg agh jgmjl ag qug gs plsnihg prldlvgr hc rg`gegrcmnýe ag phcetcs ne vntrl c pcrr ag jnplmlhls nemhusl muceal gh sudnenstrl gxý`gel ag rg`uhcalrgs agh mrgmndngetl ag hcs phcetcs sg ldntg. Ge gstg sgeal, sg jc nelrdcal prgvncdgetg mýdl mýd l uel ag hls cmtlrgs cmtlrgs dãs ndplrt ndplrtcet cetgs gs ge hc lrdcm lrdcmnýe nýe ag ýr`cel ýr`celss cavge cavgemnl mnlss gs hc geaý`gecuscuxnec? gh gqunhnirnl ag mntlqunenec y el hc mceaca ag cuxnec l mntlqunenec cócanac ge ue dganl (Qcendltl y Jcrcac, 3>:1). C gstg rgspgmtl, yc sg jc puihnmcal (Hctc gt ch., 9;30, y sus rggrgemncs) rggrgemncs) hc neugemn neugemncc qug gogrmg gogrmg gh ýr`cel agh qug sg ansgmce hls gxphcetgs gxphcetgs (y su envgh ge geaý aý`g `gel el ag rg`u rg`uhc hcal alrg rgss ag aghh mr mrgm gmnd ndng nget etl l ag hcs hcs ph phce cetc tcs) s) ge hc tc tcsc sc ag ne neau aumm mmný nýe e ag hc lr`cel`îegsns agh irltg. Ag tlals hls jgmjls dgemnlecals cetgrnlrdgetg, sg pugag mlemhunr qug hc ghgmmnýe agh gxphcetg prndcrnl y sus envghgs jlrdlechgs geaý`gels rghcmnlecals sle mrumnchgs pcrc hc lrdcmnýe ag dgrnstgdcs nevcrncihgs ge M.scvc, cuequg sn`ug sngeal egmgscrnl rgchnzcr dãs nevgs`cmnlegs pcrc cmhcrcr gstc jnpýtgsns, yc qug ]dôbchlvã gt ch. (9;3>), agspuîs ag cechnzcr hls envghgs ag jlrdlecs geaý`gecs ag jnplmlhls ag M. scvc, nelrdcrle mýdl hcs mlemgetrcmnlegs ag mntlqunenecs geaý`gecs gstcice plr agicol agh hädntg ag agtgmmnýe. [lr ltrl hcal, hc ansgmmnýe ag gxphcetgs gs ltrl cmtlr qug plaräc gstcr neuygeal ge hc lr`cel`îegsns ag irltgs ne vntrl ge jnplmýhls prlmgagetgs ag M. scvc. Acal qug sg`ÿe gh mlelmndngetl mldÿe, hcs mntlqunenecs sg prlaumge prgaldnecetgdgetg ge gh dgrnstgdl ag hc rcäz y hcs cuxnecs sg snetgzce ge gh dgrnstgdl agh irltg, y cdils pls ag ftljlrdlecs pugage dn`rcr agsag hcs rcämgs y hls irltgs c su snl ag cmmnýe c trcvîs agh lgdc y gh xnhgdc (Igmb, 3>>6), sg`dgetcmnýe ag tcetl hls dgrnstgdcs cpnmchgs ag hls irltgs mldl ag hcs rcämgs mldl rgsuhtcal ag hc ansgmmnýe mle jnplmýhl plaräce dlanfmcr hc netgrcmmnýe jlrdlech geaý`gec getrg cuxnecs y mntlqunenecs, aceal hu`cr c ue getlrel cprlpncal pcrc gh agscrrlhhl ag hc lr`cel`îegsns ag hls irltgs ge hls jnplmlhls. Hcs phcetcs agrnvcacs ag jnplmlhl sg pugage gercnzcr gspletãegcdgetg ge dganl hnirg ag jlrdlecs, qugaceal mldphgtcdgetg cmhndctcacs ge slhl sgns sgdcecs.  Gh dnsdl rczlecdngetl agsmrntl cetgrnlrdgetg tcdinîe plaräc gxphnmcr gh jgmjl ag qug, agspuîs agh agscrrlhhl agh irltg ge hc pcrtg supgrnlr agh jnplmýhl, hcs cuxnecs prlaumge ag dcegrc geaý`gec geaý `gec ge gh snstgdc snstgdc agh irltg. irltg. lIcdiuscvu lIcdiuscvuh`crn h`crns]mj s]mjrca.g rca.gxO.M xO.M.Pge .Pgeah. ah. (Furhceg (Furhcegtch., tch., 9;3:), tcdinîe sg jc agsmrntl su rghcmnýe mle gh mletgenal ag cuxnecs geaý`gecs ag irltgs mrncals ne vntrl (Dnelmjc, 3>:0). [lr ltrl hcal, tcdinîe sg jc agsmrntl gh agscrrlhhl ag rcämgs ge hc plrmnýe negrnlr ag jnplmýhls ag M. scvc sne lrdcmnýe ag dgrnstgdcs ag irltgs cpnmchgs (HcYug, 3>== ). Ge eugstrl mcsl, gh gercnzcdngetl gspletãegl ag hls rg`gegrcetgs ne vntrl rgprgsgetc uec vgetcoc canmnlech ag eugstrl prltlmlhl ag rg`gegrcmnýe, yc qug el sg rgqungrg ue dganl qug mletge`c cuxnec cux necss pcrc pcrc hc neaumm neaummnýe nýe ag rcämgs rcämgs.. Ge muchq muchqung ungrr mcsl, mcsl, cuequg cuequg ug egmgs egmgscrn crnl l prldlv prldlvgr gr gh gercnzcdngetl ag dãs phcetcs rg`gegrcacs ge gh netgrnlr, gxnstge vcrnls prltlmlhls puihnmcals slirg gh crrcstrg ag irltg ag M.scvc apgrdntg mle uec ggfmngemnc fmngemnc el negrnlr negrnlr ch :;% (Hctcgtch., 9;;>, 9;3;, 9;36c, i2 Pce` gt ch., 9;;>2 Mjcljuc gt ch., 9;362 [crslesgtch., 9;3>). Agsmuirndls qug hls jnplmlhls muhvcals ge ue dganl sne rg`uhcalrgs agh mrgmndngetl ag hcs phcetcs chmcezcrle hc tgrmgrc tcsc ag neaummnýe ag irltgs dãs chtc ag hls dganls gvchucals sne prgsge prg sgetcr tcr an angrg grgemn emncs cs sn` sn`enf enfmc mcvcs vcs mle hls ltrls ltrls als dganl dganlss mle dgolrg dgolrgss plrmg plrmgetc etcogs ogs ag lr`cel`îegsns agh irltg. Cagdãs, hls jnplmlhls ag tlacs hcs vcrngacags prlicacs puangrle agscrrlhhcr irltgs ge gstg dganl. Gstl, ouetl mle hc cusgemnc ag angrgemncs sn`enfmcvcs ge

 

tîrdnels ag eÿdgrl ag irltgs plr gxphcetg rgspleangetg getrg hls muctrl dganls mle hcs dgolrgs tcscs ag neaummnýe ag irltgs, els jc hhgvcal c su`grnr gh usl mldinecal ag jnplmlhl y dganl hnirg ag jlrdlecs, uec mldinecmnýe cagmucac cagmucac pcrc litgegr phcetcs agrnvcacs ag jnplmýhl gercnzcacs gspletãegcdgetg y mldphgtcdgetg cmhndctcacs 6 sgdcecs . Hcs mîhuhcs pgrnmämhnmcs caycmgetgs c hls plstgs ag xnhgdc plaräce sgr gh lrn`ge ag phcetcs rg`gegrcacs ne vntrl ag M. scvc.   Mle gh fe ag negrnr gh plsnihg lrn`ge ag rg`gegrcetgs ne vntrl c pcrr ag hls mlpnhgalegs, jnplmlhl jnpl mlhlss y vgracagrc vgracagrcss jlocs prlvgengetg prlvgengetgss ag phãetuhcs phãetuhcs ag 0 aäcs, aäcs, gxcdnecdl gxcdnecdlss sgmmnleg sgmmnlegss trcesv trc esvgrs grschg chgss ag jnplml jnplmlhl hlss y, cuequg cuequg yc el jcy prgsge prgsgemnc mnc ag sg lisgrv lisgrvcrl crle e dgrnst dgrnstgdl gdlss agscrrlhhcals l sus jughhcs, hc dîauhc, hc mlrtgzc y hc gpnagrdns sg nagefmcrle ãmnhdgetg. Eugstrcs lisgrvcmnlegs sle mlesnstgetgs mle hcs almudgetcacs plr Igjr gt ch. (9;36), qungegs gstuancrle gh agscrrlhhl ag hls tgonals sgmueacrnls ge jnplmlhls ag M. scvc, nhustrceal mle `rce agtchhg sgmmnlegs trcesvgrschgs ag jnplmlhls aurcetg angrgetgs pgrälals agspuîs ag hc sngdirc. Mldl sg prgsgetc ge su trcicol, hcs sgmmnlegs trcesvgrschgs ag jnplmlhls 6 y > aäcs agspuîs ag hc sngdirc mlnemnage mle eugstrls gstuanls, yc qug hc gpnagrdns, hc mlrtgzc y hc dîauhc pugage angrgemncrsg ãmnhdgetg y su cectldäc rgspgmvc tcdinîe mlnemnag mle eugstrls jchhcz`ls, hl qug els pgrdntg agsmcrtcr gh prgsgemnc ag dgrnstgdls yc agscrrlhhcals l sus trczcs ge jnplmlhls ag M. scvc, y rgmjczc plsnihgs rlhgs ge hls prlmgsls ag `gegrcmnýe lisgrvcals ge gstg gstuanl. Gh jgmjl ag qug hls als prndlranls sur`ngrce ag hc pcrtg supgrnlr ag hls jnplmlhls qug sngdprg sg anstrniuäce ge hc pgrngrnc agh ýr`cel y sg chnegcice uel rgetg ch ltrl, els hhgvý c phcetgcr hc jnpýtgsns ag qug hcs phcetcs rg`gegrcacs sg lrn`nece sngdprg ag gsg dnsdl pl ag mîhuhcs. Ge ue trcicol ag Dnhhgr (3>7>) slirg gh mrgmndngetl sgmueacrnl ge hc vcsmuhcrnzcmnýe ag hc rcäz y hcs phãetuhcs ag Juduhus hupuhus H., sg gstuaný hc ÿenmc gspgmng ouetl mle M. scvc qug pgrtgegmg c hc cdnhnc Mceecicmgcg, hc gstrumturc netgrec ag hcs rcämgs, jnplmlhls y mlpnhgalegs ag phãetuhcs. Ge gstg trcicol cetgrnlr, hls aniuols ag sgmmnlegs trcesvgrschgs ag jnplmlhl agtchhcrle hc mlegxnýe getrg hc rcäz y hls mlhgalegs ag hc phãetuhc, agsmrningeal el slhl hcs dnsdcs rg`nlegs qug ge eugstrc sgmmnýe trcesvgrsch ag jnplmlhl, snel tcdinîe als plstgs ag prltlxnhgdc uinmcals ge uec plsnmnýe pgrnîrnmc y anstrniunals ge hcals lpugstls, muyc uinmcmnýe sg csgdgoc dumjl ch ãrgc ag rg`gegrcmnýe ag dgrnstgdls agrnvcals ag jnplmlhl ge eugstrls gxpgrndgetls. Cagdãs, Dnhhgr (3>7>) tcdinîe agsmrnig mldl ue plhl ag prltlxnhgdc sg gemugetrc ge hc jgirc dgancec ag hc icsg ag mcac mlhgaýe. Acal qug ge eugstrl gstuanl, hc rg`gegrcmnýe ag phcetcs c pcrr ag mlhgalegs sngdprg sg hlmchnzý ge hc rg`nýe mgetrch ag hc zlec icsch agh ndphcetg, gs rczlecihg suplegr qug hc rg`gegrcmnýe ag phcetcs agrnvcacs ag jnplhl y ag jnplhl ge M. scvc sg lrn`nec c pcrr ag jgxnhgdplhgs. trczcs ag xnhgdcs mhcrcdgetg sgpcrcacs anstrniunacs ge hcals lpugstls, sg vnsuchnzcrle ge sgmmnlegs trcesvgrschgs trcesvgrschgs ag jnplmlhl prlvgengetgs ag phãetuhcs ag M. scvc ag > aäcs (Igjr gt ch., 9;3:). Eugvcdgetg, ge gstg trcicol cetgrnlr, el jcy uec mcrrgrc ag dgrnstgdls yc agscrrlhhcals ge sgmmnlegs trcesvgrschgs ag jnplmlhl. Mle rgspgmtl c eugstrcs lisgrvcmnlegs slirg hc rg`gegrcmnýe ag phcetcs agrnvcacs ag jlocs, cuequg ge gstg gstuanl slhl mneml phcetcs ugrle rg`gegrcacs c pcrr ag hcs jlocs, gs eltcihg mýdl tlacs ghhcs sg lrn`necrle ge hc zlec ag trcesnmnýe jloc-pgmälhl, mldl sg jc agsmrntl ge gspgmngs mldl Dlrus neanmc H. (Djctrggtch., 3>:7) , prgvncdgetg nemhunal ouetl mle M. scvc ge hc cdnhnc Dlrcmgcg, u ltrcs gspgmngs mldl Igtc vuh`crns H. (Agtrgz gt ch., 3>::) l Qcecmgtud mnegrcrnnlhnud (Qrgvnr.) ]mjuhtzInp (Jgacyct gt ch., 9;;>). Gh jgmjl ag qug hc vcsmuhcrnzcmnýe tcdinîe tge`c hu`cr ge hls pgmälhls,

 

mldl tcdinîe hl jcmg ge hls tchhls, y qug hcs phcetcs rg`gegrcacs mle jlocs sngdprg sur`ngrle ag hls pgmälh pgmälhls, ls, plarä plaräcc coustc coustcrsg rsg c eugstr eugstrcc jnpýtg jnpýtgsns sns slirg slirg hc lr`cel lr`cel`îe `îegsn gsnss ag irltgs irltgs ne vn vntrl trl agrnvcals agh pgrnmnmhl ge gstc gspgmng. Gstl cdphäc gh chmcemg ag eugstrl prltlmlhl jcmnc fegs ag dnmrlprlpc`cmnýe, c`rg`ceal hc plsninhnaca ag prlaumnr mhlegs dÿhphgs `geîmcdgetg naîemls c hcs phcetcs ag îhntg yc sghgmmnlecacs ag hcs qug plaräce agrnvcrsg, cuequg sg rgmldngeac hc gvchucmnýe ag hc faghnaca `geîmc dgancetg gh gescyl ag dcrmcalr ag rgpgmnýe ag sgmugemnc sndphg (N]]Y), yc qug sg jc rgchnzcal ge ltrcs lircs rghcmnlecacs . dnmrlprlpc`cmnýe scvc (Hctcgtch., 9;36c, i). C gstg rgspgmtl, gs ndplrtcetg geczcr mýdl rgspleage hls mlhgalegs y hls jnplmlhls sus rg`gegrcetgs agrnvcals, dngetrcs sg dcetgeäce ge tuils ag vnarnl, mleeucice prlaumngeal dÿhphgs irltgs aurcetg sngtg dgsgs agspuîs ag hc nenmncmnýe agh muhvl. Ge hc häegc ag gstc jnpýtgsns, Ccgtch. (9;;>) agsmrniný mýdl hc phurnpltgemnc ag hcs mîhuhcs agh dlelmnmhl ag Ãxnalpsns gs rgsplescihg ag hc rg`gegrcmnýe ag dgrnstgdls c pcrr ag gxphcetgs ag rcämgs g jnplmlhls muhvcals ne vntrl. Fnechdgetg, agig tgegrsg ge mugetc qug Iggmbdce y Ag ]dgt (9;31) agsmrnige mýdl hcs mîhuhcs pgrnmämhnmcs qug rlagce gh plhl agh xnhgdc sg mlesnagrce mldl ue dgrnstgdl gxtgeanal qug rggeg hc mcpcmnaca ag surnr uec anvnsnýe mghuhcr csndîtrnmc nemhusl muceal muc eal ltrcs ltrcs mî mîhuh huhcs cs sg an angr grgem gemnce nce,, y hcs mîhuh mîhuhcs cs agh mn mnmhl mhl agh agh mn mnmhl mhl rlagca rlagcacs cs plr mîhuhc mîhuhcss angrgemncacs cÿe pugage prl`rcdcrsg. pcrc mldgezcr c prlhngrcr, hl qug hhgvc c hc nenmncmnýe ag ue eugvl ýr`cel, qug pugag gxphnmcr mýdl uemnlec eugstrl trcicol, hc rg`gegrcmnýe ne vntrl sngdprg sg agscrrlhhý anrgmtcdgetg sne egmgsnaca ag angrgemncmnýe mghuhcr. ]n inge cÿe qugac dãs nevgs`cmnýe plr jcmgr pcrc vchnacr eugstrc jnpýtgsns, hc ndphnmcmnýe ag hcs rczcs vcsmuhcrgs ge hc mcpcmnaca rg`gegrcvc ag hc gvchucmnýe ag hcs phcetcs ag gxphcetgs pugag gxphnmcr hc angrgemnc ge hc lr`cenzcmnýe ag hls `gegs pcrc gvntcr hc lisgrvcmnýe ag eugstrl trcicol. Hc plhnsldcc gstã prgsgetg ge hls mlhgalegs g jnplmlhls ag phãetuhcs ag M. scvc.  Gh tîrdnel plhnsldcc ug cphnmcal plr prndgrc vgz plr Hce`hgt (3>90) c hc mleanmnýe ag gscs mîhuhcs ge hls tgonals sldãmls ag uec phcetc qug mlegege dÿhphls agh eÿdgrl pnml ag mrldlsldcs (Grvne, 3>13), sngeal agsmrntl plr prndgrc vgz ge hc hntgrcturc tgdprcec plr ]tldps (3>3;) ge ]pnecmnc lhgrcmgc H. Agsag su agsmuirndngetl, hc plhnsldcc jc snal rgplrtcac ge uec cdphnc `cdc ag gspgmngs tce anvgrscs mldl Mumudns dghl H. (Grvne, 3>=>), Igtc vuh`crns H. (]hnwn (]hnwnesb esbcc y Hubcsz Hubcszgws gwsbc, bc, 9;;7), 9;;7), Mj Mjgel gelpla planud nud qunelc qunelc Pn Pnhha hha.. (Blhce (Blhcel l gt ch., ch., 9;;:) 9;;:) l dãs rgmngetgdgetg ge ]lhceud dghle`gec H. (@crmäc-Flrtgc gt ch., 9;9;). ]g agsmrnig gealdntlsns l gealrrgauphnmcmnýe mldl plsnihgs mcuscs qug pugage mleaumnr c hc plhnsldcc (A„Cdctl, 3>612 Iuiegr gt ch., 9;;6), muyc cpcrnmnýe gstã rghcmnlecac mle gh mrgmndngetl y hc angrgemncmnýe ag hls tgonals. Qcdinîe sg jc nelrdcal ag mýdl hls tgonals vg`gtchgs rgmugetgdgetg mlegege uec prlplrmnýe ag mîhuhcs gealplhnphlnags gealplhnphlnags (Ycdscy y Budcr, 3>>;, y rggrgemncs chhä) y mýdl plrmnlegs ag hc phcetc phcetc,, mldl mldl hls ýr`cel ýr`celss y vcsls vcsls ag chdcmg chdcmgecd ecdng ngetl etl,, c dgeual dgeual mlege mlegege ge mîhuh mîhuhcs cs plhnphlnags (Caghigr` gt ch., 3>>=) . Mle rgspgmtl c hc plhnsldcc ge M. scvc, agig eltcrsg mýdl ug agsmrntc plr prndgrc vgz ge hls dgrnstgdls ag hc rcäz ag hc gspgmng plr Hntcranêrg (3>97). Hce`hg Hce `hgtt (3>90) (3>90) sgóchý sgóchý qug gh eÿdgrl eÿdgrl alihg ag mrldls mrldlsldc ldcss ge hls dgrnstgd dgrnstgdls ls ag hc rcäz rcäz prlvgengetgs ag M. scvc rgsuhtý ag als anvnsnlegs sumgsnvcs ag mcac mrldlsldc aurcetg hcs prlcsgs, dngetrcs qug Irgshcvgtz (3>96, 3>=9) prlpusl hc usnýe eumhgcr mldl hc mcusc ag hc mleanmnýe plhnsldãmc agsmrntc ge hcs rcämgs scvcs M. Aginal c hc chtc ag hntgrcturc slirg hc plhnsl plh nsldc dccc ge ltrls ltrls ýr`cel ýr`celss cagdãs cagdãs ag hcs rcämgs, rcämgs, cechnz cechnzcdl cdlss gh envgh envgh ag phlnaä phlnaäcc ag hls mlhgalegs, jnplmlhls y jlocs vgracagrcs prlvgengetgs ag phãetuhcs ag M. scvc ag 0 aäcs plr

 

dganl ag mntldgträc ag uol. Jcstc aleag scigdls, gstg gs gh prndgr gstuanl qug agsmrnig hc plhnsldcc ge mlhgalegs g jnplmlhls ag M. scvc. C hc huz ag eugstrls rgsuhtcals, dngetrcs qug hcs jlocs jlocs mlesgr mlesgrvcr vcrle le gh pctrýe pctrýe anphln anphlnag ag pnml pnml ag hc gs gspgm pgmng ng,, hls mlhg mlhgale alegs gs g jnplm jnplmlh lhls ls dlstrcrle ue pctrýe plhnsldãml qug mletgeäc mîhuhcs anphlnags y tgtrcphlnags y, plr hl tcetl, agigrä agi gräce ce ml mlesn esnagr agrcrs crsgg mldl mldl ýr`cel ýr`celss dnxlp dnxlphln hlnags ags.. Eu Eugst gstrls rls jchhcz jchhcz`ls `ls rgh rghcmn cmnlec lecals als mle mle hc plhnsldcc gemletrcals ge mlhgalegs y jnplmlhls ag M. scvc, cirge eugvcs lplrtuenacags mldl gh agscrrlhhl ag plhnphlnags c trcvîs ag hc rg`gegrcmnýe ag hls ndphcetgs ag hls ýr`cels. Hcs phcetcs dnxlphlnags sg pugage rg`gegrcr agspuîs agh muhvl ne vntrl ag jnplmlhls y mlhgalegs prlvgengetgs ag phãetuhcs ag M. scvc. Hls plhnphlnags sg cslmnce mle tcdcóls ag ýr`cels c`rceacals, cudgetl agh rgeandngetl ag hc inldcsc, mletgenal ftlquädnml y prlaumtls dgtciýhnmls, dcylr mcpcmnaca ag cacptcmnýe ch gstrîs inýml iný ml y ciným cinýml, l, y mle gh mcdin mcdinl l ag rg`uhc rg`uhcmný mnýe e agh agh `geldc `geldc (VceJngu (VceJngu,, 9;3>). 9;3>). Cagdãs, Cagdãs, gh agscrrlhhl ag phcetcs plhnphlnags, ge pcrmuhcr tgtrcphlnags, plaräc sgr ÿh ge gh ftldgolrcdngetl pcrc gh agscrrlhhl ag vcrngacags trnphlnags mle rutls sne sgdnhhcs, mldl sg jc agdlstrcal ge Mntruhhushcectus (Bnjcrc y Ensjnycdc, 3>10), MumudnsdghlH. (Caghigr` gt ch., 3>>=) l ge Mntrus spp. (Ygmupgrl gt ch., 9;;7). ]ne qug hls eumhgnals ag plhnphlnags ge lmcsnlegs sgce hls prl`gentlrgs ag hc `gegrcmnýe ag cmghncs, aceal hu`cr c ue pcrmjg ag tgonals plhnphlnags (A'Cdctl, 3>79) y trcs mlelmgr hc plhnsldcc ge mlhgalegs g jnplmlhls ag M. scvc, gvchucdls sn jciäcdls litgenal rg`geg rg` gegrce rcetg tgss plhnph plhnphlna lnags gs qug puangr puangrce ce sgr sgr ÿhgs ÿhgs pcrc pcrc hc prlaum prlaummný mnýe e ag mceeci mceecinel nelnag nags. s. , agtgmtcdls angrgemncs el sn`enfmcvcs getrg gxphcetgs ge tîrdnels ag plhnphlnanzcmnýe ag phcetcs rg`gegrcacs y mýdl gh mlhgaýe y gh jnplmýhl grce hls ÿenmls gxphcetgs mcpcmgs ag `gegrcr phcetcs dnxlphlnags. Mcig sgóchcr mýdl hc plhnphlnanzcmnýe sughg gstcr cslmncac c envghgs ghgvcals ag dgtcilhntls sgmueacrnls ge ue `rce eÿdgrl ag gspgmngs (Nceenmghhn gt ch., 9;3>), cuequg tcdinîe gxnstge gxmgpmnlegs, mldl gs gh mcsl ag Ml ﬋ gc crcinmc H. y Ml ﬋ gc mcegpjlrc H . (]nhvcrlhhc gt ch., 3>>>), aleag hc plhnphlnanzcmnýe ansdneuyý c hls envghgs agmcgnegnec ge hcs jlocs. Ge gh mcsl ag M. scvc, hls rgsuhtcals gemletrcals ge hc hntgrcturc sle mletrcanmtlrnl. ]n inge Mhcrbg (3>:3) gemletrý qug hc plhnphlnanzcmnýe cudgetcic hls envghgs ag QJM dngetrcs ansdneuäc gh mletgenal ag MIA y mceecinelh (MIE), y qug hls trnphlnags agdlstrcrle sgr negrnlrgs c hls anphlnags anph lnags y tgtrcphln tgtrcphlnags ags ge tîrdnels tîrdnels ag prlaummný prlaummnýe e ag mceecinel mceecinelnags nags,, [crsles [crsles gt ch. (9;3>) lituvngrle ue pgrfh quädnml sndnhcr getrg tgtrcphlnags y anphlnags, cuequg cudgetls eltcihgs ge MIA y sgsquntgrpgels sg cslmncrle mle tgtrcphlnags. Ge rghcmnýe c hls dnxlphlnags y su mletgenal ag mceecinelnags, Dceslurn y Ic`jgrn (9;30) agdlstrcrle qug hc plhnphlnanzcmnýe nemrgdgetý sn`enfmc sn`en fmcvcdge vcdgetg tg gh mletgena mletgenal l ag QJM ge phcetcs phcetcs dnxlphlna dnxlphlnags gs ge mldpcrcmn mldpcrcmnýe ýe mle phcetcs tgtrcp tgt rcphln hlnags ags y anphln anphlnags ags,, y qug hcs phcetc phcetcss tgtrcp tgtrcphln hlnags ags tgeäce tgeäce dgelrg dgelrgss mceac mceacags ags ag gstc gstc sustcemnc ge mldpcrcmnýe mle anphlnags, su`nrngeal qug hls dnxlphlnags plaräce sgr ÿhgs pcrc prlaumnr QJM pcrc usl mldgrmnch. Fnechdgetg, mcig agstcmcr mýdl hc plhnphlnanzcmnýe ge M. scvc sngdprg jc snal neaumnac plr gh trctcdngetl ag dgrnstgdcs cpnmchgs ag hc phcetc mle ansruptlrgs quädnmls ag hls dnmrltÿiuhls ag chtl `rcal ag tlxnmnaca, mldl hc mlhmjnmnec u lrnzchnec (uec chtgrecvc dgels týxnmc c hc mlhmjnmnec), y qug hcs phcetcs plhnphlnags quädnmcdgetg neaumnacs sughge rgvgrr c hc mleanmnýe anphlnag (Mhcrbg, 3>:3), hl qug lihn`c c prlicr gh envgh ag phlnaäc ag hcs phcetcs plhnphlnags c hl hcr`l ag `gegrcmnlegs MLEMHW]NÝE

 

Ge mlemhusnýe, aginal c hc chtc mcpcmnaca rg`gegrcvc agh jnplmlhl y qug slhl hls rg`gegrcetgs ne vntrl agrnvcals agh jnplmlhl ugrle mcpcmgs ag gercnzcr gspletãegcdgetg, ouetl mle hc cusgemnc ag angrgemncs sn`enfmcvcs getrg hls dganls mle hcs dgolrgs tcscs ag neaummnýe ag irltgs mle rgspgmtl ch eÿdgrl ag irltgs plr gxphcetg ag rgspugstc, y getrg phcetcs agrnvcacs ag mlhgalegs g jnplmlhls ge tîrdnels ag plhnphlnanz plhnphlnanzcmnýe, cmnýe, su`grndls gh muhvl ag jnplmlhls ge dganl hnirg ag jlrdle jlr dlecs cs ml mldl dl gh dãs cagmuc cagmucal al ag hls trctcd trctcdng ngetl etlss gvchu gvchucal calss ge gstg gstg gstuan gstuanl. l. Eu Eugst gstrl rl prltlmlhl jcmg qug hc rg`gegrcmnýe ne vntrl anrgmtc ag hcs phcetcs ag M. scvc sgc dgels chgctlrnc y agpgeangetg agh `gelpl, plr hl qug plaräc tgegr ndplrtcetgs mleeltcmnlegs ge hc gxphltcmnýe ag tîmenmcs mletgdplrãegcs ag dgolrcdngetl ag phcetcs mldl hc ganmnýe agh `geldc (p. Go., Ws Wsce ceal al hc ga ganm nmný nýe e ag aghh `ge `ge MY MYN] N][Y [Y / Mcs) Mcs) l dutc dutc`î `îeg egsn sns, s, sn snge geal al Qc Qcdi dinî nîe e gs ÿ ÿhh pc pcrc rc hc dnmrlp dnm rlprlp rlpc`c c`cmný mnýe e y pcrc pcrc gh agscrr agscrrlhh lhhl l ag vcrnga vcrngacag cagss plhnph plhnphlna lnags gs mle envghg envghgss dgolrc dgolrcals als ag mceecinelnags sne uscr ansruptlrgs ag dnmrltuilhuihgs quädnmls týxnmls