ARTÍCULO DE REVISIÓN 
Acta Pediátr Mex ! 
La mitosis y su regulación ůĨƌĞĚŽ ĚĞ :ĞƐƷƐ ZŽĚƌşŐƵĞnjͲ'ſŵĞnj͕1 Sara 
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Acta Pediátrica de México 
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Rodríguez-Gómez AJ y Frias-Vázquez S. La mitosis y su regulación 
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Rodríguez-Gómez AJ y Frias-Vázquez S. La mitosis y su regulación 
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Rodríguez-Gómez AJ y Frias-Vázquez S. La mitosis y su regulación 
REGULACIÓN MOLECULAR DE LA MITOSIS 
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Mitosis

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Mitosis

  1. 1. ARTÍCULO DE REVISIÓN Acta Pediátr Mex ! La mitosis y su regulación ůĨƌĞĚŽ ĚĞ :ĞƐƷƐ ZŽĚƌşŐƵĞnjͲ'ſŵĞnj͕1 Sara ƌŝĂƐͲsĄnjƋƵĞnj1,2 1 ĂďŽƌĂƚŽƌŝŽ ĚĞ ŝƚŽŐĞŶĠƟĐĂ͕ ĞƉĂƌƚĂŵĞŶƚŽ ĚĞ /ŶǀĞƐƟŐĂĐŝſŶĞŶ'ĞŶĠƟĐĂ,ƵŵĂŶĂ͕/ŶƐƟƚƵƚŽEĂĐŝŽŶĂů ĚĞWĞĚŝĂƚƌşĂ͘ 2 ĞƉĂƌƚĂŵĞŶƚŽĚĞDĞĚŝĐŝŶĂ'ĞŶſŵŝĐĂLJdŽdžŝĐŽůŽŐşĂ ŵďŝĞŶƚĂů͕/ŶƐƟƚƵƚŽĚĞ/ŶǀĞƐƟŐĂĐŝŽŶĞƐŝŽŵĠĚŝĐĂƐ͕ UNAM. ZĞĐŝďŝĚŽ͗Ăďƌŝů͕ϮϬϭϯ ĐĞƉƚĂĚŽ͗ƐĞƉƟĞŵďƌĞ͕ϮϬϭϯ Correspondencia Dra. Sara Frias ĂďŽƌĂƚŽƌŝŽĚĞŝƚŽŐĞŶĠƟĐĂ /ŶƐƚŝƚƵƚŽ EĂĐŝŽŶĂů ĚĞ WĞĚŝĂƚƌşĂ /ŶƐƵƌŐĞŶƚĞƐ ^Ƶƌ ϯϳϬϬͲ͕ϲΣƉŝƐŽ ϬϰϱϯϬDĠdžŝĐŽdĞůĠĨŽŶŽLJĨĂdž͗нϱϮϱϱϭϬϴϰϱϱϯϯ͕ ϭϬϴϰϬϵϬϬ͕ĞdžƚĞŶƐŝſŶϭϰϯϲ sarafrias@biomedicas.unam.mx ƐƚĞĂƌơĐƵůŽĚĞďĞĐŝƚĂƌƐĞĐŽŵŽ͗ ZŽĚƌşŐƵĞnjͲ'ſŵĞnj:͕ƌŝĂƐͲsĄnjƋƵĞnj^͘ĂŵŝƚŽƐŝƐLJƐƵ ƌĞŐƵůĂĐŝſŶ͘ĐƚĂWĞĚŝĂƚDĞdžϮϬϭϰ͖ϯϱ͗ϱϱͲϲϴ͘ RESUMEN 3H KP]PZP}U JLSSHY WVY TP[VZPZ LZ KLJPZP]H WHYH LS KLZHYYVSSV KL SVZ VYNHUPZTVZ ` Z YLWYVKJJP}U HUHKV H LSSV LZ ULJLZHYPV XL JHKH UL]H JtSSH ZLH NLUt[PJHTLU[L PKtU[PJH KL SH XL WYV]PLUL ,U SVZ LJHYPVU[LZ LZ[V ZL SVNYH NYHJPHZ H TLJHUPZTVZ JVTWSLQVZXLHZLNYHUSHPU[LNYPKHKKLSTH[LYPHSNL- U}TPJV`ZZLNYLNHJP}UHWYVWPHKHKYHU[LSHTP[VZPZ 3H]PZP}U[YHKPJPVUHSKLSHTP[VZPZSHOHKP]PKPKVLU KPMLYLU[LZL[HWHZXLSVNYHYVUJHYHJ[LYPaHYZLNYHJPHZ HSVZLZ[KPVZTVYMVS}NPJVZLUJtSSHZLUKP]PZP}USVZ H]HUJLZLUIPVSVNxHTVSLJSHYOHUSSL]HKVTmZHSSm LZ[HJHYHJ[LYPaHJP}UKLTHULYHXLHOVYHZLJVUVJL [VKHUHNHTHKLWHY[PJPWHU[LZTVSLJSHYLZ,ULZ[L HY[xJSVZLHIVYKHYmULSWYVJLZVKLSHTP[VZPZJLSSHY `TVSLJSHY`UHIYL]LZxU[LZPZKLSVZHJ[VYLZTVSL- JSHYLZXLYLNSHULZ[LWYVJLZV Palabras clave:KP]PZP}UJLSSHYTP[VZPZJPJSVJLSSHY Mitosis and its regulation ABSTRACT *LSSKP]PZPVUI`TP[VZPZPZLZZLU[PHSMVY[OLKL]LSVWTLU[ VMVYNHUPZTZHUK[OLPYYLWYVKJ[PVUP[PZHSZVULJLZ- ZHY`[OH[LHJOUL^JLSSPZNLUL[PJHSS`PKLU[PJHS[V[OH[ MYVT ^OPJO P[ JVTLZ 0U LRHY`V[LZ [OPZ PZ HJOPL]LK I`[OLWYLZLUJLVMJVTWSL_TLJOHUPZTZ[OH[LUZYL [OL PU[LNYP[` VM NLUVTPJ TH[LYPHS HUK [OLPY WYVWLY ZLNYLNH[PVU KYPUN TP[VZPZ;OL [YHKP[PVUHS ]PL^ VM TP[VZPZOHZILLUKP]PKLKPU[VKPMMLYLU[Z[HNLZ[OH[^LYL JOHYHJ[LYPaLK I` TVYWOVSVNPJHS Z[KPLZ PU KP]PKPUN JLSSZHK]HUJLZPUTVSLJSHYIPVSVN`OH]LSLKIL`VUK [OPZ JOHYHJ[LYPaH[PVU ZV [OH[ ^L UV^ RUV^ H YHUNL VMWHY[PJPWHU[TVSLJSLZ;OPZHY[PJSL^PSSKPZJZZ[OL WYVJLZZVMTP[VZPZIV[OH[[OLJLSSSHYHUKTVSLJSHY SL]LS HUK H IYPLM ZTTHY` VM [OL TVSLJSHY WSH`LYZ [OH[YLNSH[L[OPZWYVJLZZ Key words: +P]PZPVUJLSSHY4P[VZPZ*PJSVJLSSHY www.actapediatricademexico.org 55
  2. 2. Acta Pediátrica de México 56 Volumen 35, Núm. 1, enero-febrero, 2014 ,U SVZ VYNHUPZTVZ WSYPJLSSHYLZ JVTV LS OTHUV XL PUPJPHU Z ]PKH JVU UH JtSSH OL]VVJPNV[VSHKP]PZP}UJLSSHYTP[}[PJHLZ KLJPZP]HWHYHLSKLZHYYVSSV`THU[LUPTPLU[VKL SVZKP]LYZVZ[LQPKVZ}YNHUVZ`ZPZ[LTHZXLSV MVYTHU 3HZ UL]HZ JtSSHZ VYPNPUHKHZ WVY SH TP[VZPZZVUNLUt[PJHTLU[LPKtU[PJHZHSHJtS- SHTHKYLLZ[VZLSVNYHNYHJPHZHTLJHUPZTVZ JVTWSLQVZKLYLNSHJP}UXLHZLNYHUSHPU[L- NYPKHKKLSTH[LYPHSNLU}TPJV`ZZLNYLNHJP}U HWYVWPHKH3HZLYPLKLL]LU[VZXLJVUKJLU H SH KP]PZP}U KL UH JtSSH ZL JVUVJL JVTV JPJSV JLSSHY ` LZ[m JVUZ[P[xKV WVY KVZ MHZLZ WYPUJPWHSLZ! SH PU[LYMHZL ` SH KP]PZPVU JLSSHY XLWLKLZLYTP[VZPZVTLPVZPZ3HTP[VZPZLZ LSWYVJLZVUJSLHYWVYLSJHSSVZJYVTVZVTHZ YLWSPJHKVZ ZL ZLNYLNHU LU KVZ U‚JSLVZ OPQVZ NLULYHSTLU[L]HHJVTWH|HKHKLSHJP[VJPULZPZ XLLZSHKP]PZP}UKLSJP[VWSHZTH`ZLWHYHJP}U MxZPJHKLSHZKVZJtSSHZOPQHZ ,SWYVJLZVTP[}[PJVMLKLZJYP[VWYPTLYHTLU[L WVY -SLTTPUN LU Z KYHJP}U LZ NLUL- YHSTLU[LKLTLUVZKLUHOVYH[PLTWVLULS XLSHJtSSHLZJHWHaKLZLWHYHYZPUMVYTHJP}U NLUt[PJHLUKVZNYWVZPKtU[PJVZXLQU[VJVU LSYLZ[VKLZZJVTWVULU[LZZIJLSSHYLZZLYmU OLYLKHKVZHSHZJtSSHZOPQHZ+YHU[LSHTP[VZPZ SHJtSSHZLVJWHLUUHHJ[P]PKHKWYPUJPWHS XLLZSHZLNYLNHJP}UJYVTVZ}TPJH`WYmJ[PJH- TLU[LKL[PLULLSTL[HIVSPZTVSH[YHUZJYPWJP}U `SH[YHKJJP}U1 (U[LZKLSWYVJLZVKLTP[VZPZKYHU[LSHMHZL: KLSJPJSVJLSSHYSVZJYVTVZVTHZZLYLWSPJHUKL THULYHXLLULSTVTLU[VKLPUPJPHYSHKP]PZP}U JLSSHYLUOTHUVZJHKHUVKLSVZJYVTV- ZVTHZYLWSPJHKVZ[LUKYmKVZJYVTm[PKHZUPKHZ WVYLSJLU[Y}TLYVJHKHUHKLLSSHZYLWYLZLU[H U JYVTVZVTH MUJPVUHS *HKH JYVTm[PKH OLYTHUHKLUJYVTVZVTHZLNYLNHUHJtSSH OPQHHSPNHSXLSHZV[YHZ`QU[HZPU[LNYHYmU LS ¸NYWV¹ KPWSVPKL KL JYVTVZVTHZ KL SH UL]HJtSSH3HTP[VZPZZLJHYHJ[LYPaHWVYKVZ L]LU[VZ PTWVY[HU[LZ XL WLKLU ]PZHSPaHYZL IHQV LS TPJYVZJVWPV KL Sa! SH JVUKLUZHJP}U `SHZLNYLNHJP}UJYVTVZ}TPJH,STV]PTPLU[V JYVTVZ}TPJVLZ[mTLKPHKVWVYUHLZ[YJ[YH JVTWLZ[HWYPTVYKPHSTLU[LWVYTPJYV[‚ISVZ! LS OZV TP[}[PJV XL ZL LUJHYNH KL HSPULHY H SVZ JYVTVZVTHZ YLWSPJHKVZ ` JVUKLUZHKVZ LU LSJLU[YVKLSHJtSSHWHYHHZxWVZPJPVUHYHSHZ JYVTm[PKHZOLYTHUHZJVULSJPUL[VJVYVKLUH HWU[HUKVOHJPHUWVSV`LSKLSHV[YHHWU[HU- KVHSWVSVVWLZ[VLULZLTVTLU[VZLLZJPUKLSH WYV[LxUHXLTHU[LUxHUPKVZSVZJLU[Y}TLYVZ` HSHZJYVTm[PKHZOLYTHUHZ(XxLSOZVTP[}[PJV TLKPHU[LZZTPJYV[‚ISVZJPUL[VJ}YPJVZQHSHSHZ JYVTm[PKHZOHJPHWVSVZVWLZ[VZ,S‚S[PTVWHZV LZSHYLPUZ[HSHJP}UKLUU‚JSLVPU[LYMmZPJV`SH KP]PZP}UKLSJP[VWSHZTHWHYHMVYTHYKVZJtSSHZ OPQHZPKtU[PJHZ;VKVZLZ[VZL]LU[VZZLYLHSPaHU LU JPUJV L[HWHZ UJSLHYLZ! WYVMHZL WYVTL[H- MHZL TL[HMHZL HUHMHZL [LSVMHZL ` ÄUHSTLU[L ZL YLHSPaH SH JP[VJPULZPZ XL LZ SH KP]PZP}U KLS JP[VWSHZTH2-PNYH Figura 1. La mitosis en el contexto de las distintas fases del ciclo celular. ,UUJPJSVJLSSHYTP[}[PJV JVUKYHJP}UHWYV_PTHKHKLOVYHZSHMHZL4ZL KLZHYYVSSHHWYV_PTHKHTLU[LLUUHOVYHKLTHULYH XLJHKHUHKLSHZJPUJVL[HWHZUJSLHYLZ`SHJP[V- JPULZPZ[PLULUSNHYLUTPU[VZ
  3. 3. 57 Rodríguez-Gómez AJ y Frias-Vázquez S. La mitosis y su regulación ETAPAS DE LA MITOSIS Profase 3H[YHUZPJP}UKLPU[LYMHZLHTP[VZPZLZSHWYVMHZL ,ULZ[HL[HWHVJYYLUSVZZPNPLU[LZL]LU[VZ!SH JYVTH[PUH ZL JVUKLUZH WHYH MVYTHY JYVTVZV- THZZLMVYTHLSOZVTP[}[PJV`KLZHWHYLJLSH LU]VS[YHUJSLHY 1) Condensación cromosómica 3HWYPTLYHTHUPMLZ[HJP}U]PZPISLKLSHKP]PZP}U JLSSHY LZ SH JVTWHJ[HJP}U WYVNYLZP]H KL SH JYVTH[PUHUJSLHY-PNYHHWHYHKHYSNHYH OLIYHZJYVTVZ}TPJHZLULZ[LWU[VSVZJYVTV- ZVTHZZLKLÄULUJVTVOLIYHZKVISLZWVYXL`H LZ[mUYLWSPJHKVZLZ[LLTWHXL[HTPLU[VLZPU- KPZWLUZHISLWHYHXLSVZJYVTVZVTHZUVZMYHU HS[LYHJPVULZ NLULYHKHZ WVY LS LZ[YtZ TLJmUPJV HXLZVUZVTL[PKVZKLIPKVHSVZTV]PTPLU[VZ KLS OZV TP[}[PJV KYHU[L SH ZLNYLNHJP}U JYV- TVZ}TPJH2 3H JYVTH[PUH XL LZ[m LU ÄIYHZ KL UT LU PU[LYMHZL JVTPLUaH H JVUKLUZHYZL WVY SH PU[LY- ]LUJP}UKLUJVTWSLQVWYV[LPJVKLJVUKLUZPUH `KL[VWVPZVTLYHZH00XLZWLYLUYVSSHUSHÄIYH KLUTMVYTHUKVHZHZKL(+5ZWLYLUYVSSH- KVHSVSHYNVKLJHKHJYVTm[PKH7HYHTHU[LULY UPKHZHSHZJYVTm[PKHZOLYTHUHZKLSJYVTVZVTH YLWSPJHKV`JVUKLUZHKVPU[LY]PLULV[YVJVTWSLQV WYV[LPJVSSHTHKVJVOLZPUHXLTHU[PLULUPKHZ H SHZ KVZ JYVTm[PKHZ KLZKL XL [LYTPUHYVU Z YLWSPJHJP}ULUMHZL:OHZ[HSHHUHMHZLTP[}[PJH 3HJVUKLUZPUH`SHJVOLZPUHZVULZ[YJ[YHZZP- TPSHYLZXLWLKLUMVYTHYHUPSSVZXLYL[PLULU ZLNTLU[VZKPZ[HU[LZKLJYVTH[PUHUPKVZ-PNYH 3HJVOLZPUHZLLUJLU[YHHSVSHYNVKLSVZ IYHaVZJYVTVZ}TPJVZTHU[LUPLUKVUPKHZSVUNP- [KPUHSTLU[LHSHZJYVTm[PKHZOLYTHUHZ`KHUKV SH HWHYPLUJPH KL UH ZVSH OLIYH HS JYVTVZVTH TP[}[PJV[LTWYHUV;HTIPtUZLLUJLU[YHUPLUKV MLY[LTLU[LHSHZJYVTm[PKHZWVYLSJLU[Y}TLYV SHJVOLZPUHSVJHSPaHKHHSVSHYNVKLSVZIYHaVZZL ZLWHYHKLSHJYVTH[PUHLUWYVMHZL`SHKLSJLU- [Y}TLYVZLYL[PLULOHZ[HHUHMHZL 3HJYVTH[PUHL_[LUKPKHKLSU‚JSLVLUPU[LYMHZL WLYTP[LLSPUNYLZVKLSHTHXPUHYPH[YHUZJYPWJPV- UHSWLYVKYHU[LSHTP[VZPZJHUKVSHJYVTH[PUH [PLULSHTH`VYJVUKLUZHJP}UKLSJPJSVJLSSHY Figura 2. Las distintas fases de la mitosis. *S[P]VKLÄIYVISHZ[VZLUSVZXLZLVIZLY]HU!H7YVMHZLWYVTL[HMHZL! JVUJYVTVZVTHZLUWYVJLZVKLJVUKLUZHJP}U`LU]VS[YHUJSLHYKLZPU[LNYmUKVZLI4L[HMHZL!JYVTVZVTHZ HSPULHKVZLUSHWSHJHLJH[VYPHSJVUTH`VYJVUKLUZHJP}J(UHMHZL!ZLWHYHJP}ULUKVZNYWVZKLSHZJYVTm[PKHZ OLYTHUHZK;LSVMHZL!KP]PZP}UJP[VWSmZTPJH`YLPU[LNYHJP}UKLSHZLU]VS[YHZUJSLHYLZWHYHMVYTHYKVZJtSSHZ OPQHZ-V[VNYHMxHJVY[LZxHKLS+Y1VHXxU*HYYPSSV0UZ[P[[VKL/LTH[VWH[VSVNxH
  4. 4. Acta Pediátrica de México 58 Volumen 35, Núm. 1, enero-febrero, 2014 SH[YHUZJYPWJP}UZLPUOPIL+LIPKVHLZ[VLSU- JSLVSV¶LZ[YJ[YHMVYTHKHWYPTVYKPHSTLU[LWVY WYVKJ[VZKLSH[YHUZJYPWJP}UKLS(+5YPIVZVTHS ` WYV[LxUHZ KLZHWHYLJL ` LU JVUZLJLUJPH [HTIPtUSH[YHKJJP}UZLKL[PLUL 2) Formación del huso mitótico 3VZTPJYV[‚ISVZKLSJP[VLZXLSL[VKLPU[LYMHZL ZL KLZLUZHTISHU KLIPKV H TVKPMPJHJP}U KL WYV[LxUHZHZVJPHKHZJVUTPJYV[‚ISVZV4(7Z SVZTPJYV[‚ISVZZLYLVYNHUPaHUWHYHJVU[YPIPY H SH MVYTHJP}U KLS OZV TP[}[PJV ,Z[H UL]H VYNHUPaHJP}UKLSVZTPJYV[‚ISVZZLPUPJPHWVY SHLZJPZP}UKLUHLZ[YJ[YHKWSPJHKHLUMHZL: KLSJPJSVJLSSHYSSHTHKHJLU[YVZVTHJHKHUV KLSVZKVZJLU[YVZVTHZJVUZ[HKLKVZJLU[YPV- SVZWVZPJPVUHKVZLUmUNSVYLJ[VUVKLSV[YV YVKLHKVZ WVY UH TH[YPa WYV[LPJH +YHU[L SH WYVMHZLLSWYPTLYWHZVWHYHMVYTHYLSOZVTP[}- [PJVLZSHHWHYPJP}UVUJSLHJP}UKLTPJYV[‚ISVZ HSYLKLKVYKLSVZJLU[YVZVTHZXLMVYTHUUH LZWLJPL KL LZ[YLSSH PU[LNYHUKV HZx SVZ mZ[LYLZ J`VZ TPJYV[‚ISVZ [PLULU U L_[YLTV TLUVZ HZVJPHKV HS JLU[YVZVTH ` U L_[YLTV TmZ HSJHSZLHKPJPVUHUKxTLYVZKL[ISPUHH TH`VY]LSVJPKHK*HKHmZ[LYTPNYHHWVZPJPVULZ VWLZ[HZKLU[YVKLSHJtSSHLZ[HISLJPLUKVHZx SVZWVSVZJLSSHYLZHWHY[PYKLKVUKLZLMVYTHYm UOZVTP[}[PJVIPWVSHY-PNYH3VZTPJYV- [‚ISVZKLSVZmZ[LYLZJVU[PU‚HUJYLJPLUKVWVY ZZL_[YLTVZOHZ[HXLHSNUVZKLLSSVZSVZ SSHTHKVZ TPJYV[‚ISVZ JPUL[VJ}YPJVZ LUJLU- [YHULSJPUL[VJVYVKLUHKLSHZKVZJYVTm[PKHZ OLYTHUHZKLUJYVTVZVTHLUKVUKLXLKHU HUJSHKVZ H WYV[LxUHZ KLS JPUL[VJVYV LZWLJx- ÄJHTLU[L KL SH JVYVUH ÄIYVZH JVU H`KH KL V[YHZWYV[LxUHZJVTV5KJ*,57,RPULZPUH `KPULxUH-PNYH3VZTPJYV[‚ISVZKLSWVSV VWLZ[V OHYmU JVU[HJ[V JVU SH V[YH JYVTm[PKH OLYTHUH KLS TPZTV JYVTVZVTH 3VZ TPJYV- [‚ISVZ XL UV LUJVU[YHYVU U JPUL[VJVYV ZL Figura 3. Empaquetamiento del ADN mediado por los complejos de condensina y cohesina. 3HJVUKLUZPUH LZ[m MVYTHKH WVY JH[YV ZIUPKHKLZ KL WYV[LxUHZ :4*`WYV[LxUHZHZVJPHKHZXLZLLUJHYNHUKL JLYYHYVHIYPYLSHUPSSVXLUPYmSVUNP[KPUHSTLU[L HZHZ KL JYVTH[PUH WHYH JVUKLUZHY JHKH UH KL SHZ JYVTm[PKHZOLYTHUHZ3HJVOLZPUHLZ[mMVYTHKHWVY JH[YVZIUPKHKLZKL:4*JVTVZPMLYHUHUP- SSVZLZYLNSHKHWVYWYV[LxUHZHZVJPHKHZKLSHMHTPSPH :**,ZLHUPSSVLUSHaHSHJYVTH[PUHKLSHZKVZJYVTm- [PKHZOLYTHUHZ`SHZULWHYHMVYTHYUJYVTVZVTH TP[}[PJV[xWPJV,Z[HUP}ULZTmZKYHKLYHLU[YLSH JYVTH[PUHKLSJLU[Y}TLYVXLLU[YLSHJYVTH[PUHH SVSHYNVKLSVZIYHaVZJYVTVZ}TPJVZ Figura 4. Papel de los centrosomas en la formación del huso mitótico.3VZJLU[YPVSVZXLZLLUJLU[YHU LUWHYLZZLKWSPJHUKYHU[LSHPU[LYMHZLLUSHMHZL: `ZLPU[LNYHUSVZJLU[YVZVTHZXLKYHU[LSHTP[VZPZ LZ[HISLJLYmUSVZWVSVZKLSOZVTP[}[PJV
  5. 5. 59 Rodríguez-Gómez AJ y Frias-Vázquez S. La mitosis y su regulación SSHTHU TPJYV[‚ISVZ PU[LYWVSHYLZ JVU[PU‚HU JYLJPLUKV WVY Z L_[YLTV OHZ[H XL ZL LU- JLU[YHU`ZLZWLYWVULUJVUSVZL_[YLTVZKL SVZTPJYV[‚ISVZKLSWVSVJVU[YHYPVXLKHUKV HZx U OZV TP[}[PJV MUJPVUHS JVU [YLZ [PWVZ KLTPJYV[‚ISVZ!SVZTPJYV[‚ISVZHZ[YHSLZSVZ TPJYV[‚ISVZJPUL[VJ}YPJVZ`SVZTPJYV[‚ISVZ PU[LYWVSHYLZ-PNYH4 ,_PZ[LU KVZ [PWVZ KL TP[VZPZ! SHZ KLWLUKPLU[LZ ` SHZ PUKLWLUKPLU[LZ KL JLU[YVZVTHZ LU SHZ ZLNUKHZSHUJSLHJP}UKLSVZTPJYV[‚ISVZZL VYPNPUHLUSVZJPUL[VJVYVZ`LZT`WYVIHISLXL HULUSHZJtSSHZJVUJLU[YVZVTHZMUJPVUHSLZ OH`H[HTIPtUUJSLHJP}UKLTPJYV[‚ISVZKLZKL LSJPUL[VJVYVKLSHZJYVTm[PKHZ 3) Desaparición de la envoltura nuclear y fragmentación del aparato de Golgi 7HYHXLSVZTPJYV[‚ISVZJPUL[VJ}YPJVZPU[LYHJ- [‚LUJVUSVZJYVTVZVTHZSHLU]VS[YHUJSLHY KLILKLZHWHYLJLYLZ[VZLYLHSPaHWVYMYHNTLU- [HJP}U KLIPKV H SH PU[LYHJJP}U KLS JVTWSLQV JPJSPUH)*KRJVULSLTLU[VZKLJHKHUVKL SVZ [YLZ JVTWVULU[LZ KL SH LU]VS[YH! a) las TLTIYHUHZUJSLHYLZb)SVZWVYVZUJSLHYLZ `c) SH SmTPUH UJSLHY ,Z WYVIHISL XL SVZ MYHNTLU[VZ YLZS[HU[LZ LU MVYTH KL ]LZxJSHZ ZL KPZWLYZLU H [YH]tZ KL SH JtSSH TP[}[PJH V IPLUXLZLPU[LNYLUHMYHNTLU[VZKLYL[xJSV LUKVWSmZTPJV+LTHULYHZPTPSHYLSHWHYH[VKL .VSNPZLKLZPU[LNYHLUWLXL|HZ]LZxJSHZJ`V KLZ[PUV LZ ZPTPSHY HS KL SH LU]VS[YH UJSLHY ,U JHSXPLYH KL SVZ KVZ JHZVZ SHZ ]LZxJSHZ PUKLWLUKPLU[LZ V IPLU HZVJPHKHZ HS YL[xJSV LUKVWSmZTPJVZLZLNYLNHYmUHSHZJtSSHZOPQHZ `]VS]LYmUHPU[LNYHYZZLZ[YJ[YHZVYPNPUHSLZ KLU[YVKLLSSHZ,SYL[xJSVLUKVWSmZTPJVWVKYxH UVMYHNTLU[HYZLOHZ[H]LZxJSHZ`ZPTWSLTLU- [LZLNYLNHYZLLUMYHNTLU[VZKLtZ[LOHJPHSHZ KVZ JtSSHZ OPQHZ 6[YVZ JVTWVULU[LZ TLT- IYHUVZVZJVTVSHZTP[VJVUKYPHZSPZVZVTHZ` WLYV_PZVTHZ UV ZL KPZNYLNHU ` ZPTWSLTLU[L Figura 5. Estructura del cinetocoro. ,Z[m JVTWLZ[V WVY[YLZSmTPUHZWYV[LPJHZ!UHPU[LYUHUHTLKPH`UH L_[LYUHLUSHXLZLL_[PLUKLUWYV[LxUHZÄIYVZHZXL PU[LNYHUSHJVYVUHÄIYVZH3HWYV[LxUH5KJZLLUJHYNH KLTHU[LULYUPKVZSVZTPJYV[‚ISVZHSHJHWHL_[LYUH KLSJPUL[VJVYVTPLU[YHZZLYLHSPaHSHKLZWVSPTLYPaHJP}U KL[ISPUHWVYRPULZPUHZ Figura 6 Diferentes tipos de microtúbulos en el huso mitótico. 3VZ TPJYV[‚ISVZ HZ[YHSLZ YVKLHU H SVZ JLU[YPVSVZ [PLULU Z L_[YLTV LU SH WLYPMLYPH KLS TH[LYPHS WLYPJLU[YPVSHY ` Z L_[YLTV WVZP[P]V LZ LS XLPYYHKPHOHJPH[VKHZKPYLJJPVULZ3VZTPJYV[‚ISVZ JPUL[VJ}YPJVZZVUSVZTPJYV[‚ISVZXLZLLUJLU[YHU HZVJPHKVZWVYZL_[YLTVWVZP[P]VHSVZJPUL[VJVYVZKL SHZJYVTm[PKHZOLYTHUHZJHKHUHKLLSSHZ[PLULUOHa KLTPJYV[‚ISVZJPUL[VJ}YPJVZXLHWU[HHL_[YLTVZ VWLZ[VZ3VZTPJYV[‚ISVZPU[LYWVSHYLZJYLJLUWVYZZ L_[YLTVZWVZP[P]VZZPULUJVU[YHYUJPUL[VJVYVOHZ[H XLZLZWLYWVULUJVUV[YVTPJYV[‚ISVPU[LYWVSHY
  6. 6. Acta Pediátrica de México 60 Volumen 35, Núm. 1, enero-febrero, 2014 ZLZLNYLNHUKLTHULYHNLULYHSTLU[LZPTt[YPJH HSHZJtSSHZOPQHZ Prometafase ,Z[H ZLNUKH MHZL ZL JHYHJ[LYPaH WVY U TV]P- TPLU[V HJ[P]V XL KPYPNL H SVZ JYVTVZVTHZ HS LJHKVYJLSSHY,SPUPJPVKLSHWYVTL[HMHZLZL YLJVUVJL WVY SH PU[LYHJJP}U KLS OZV TP[}[PJV JVU SVZ JYVTVZVTHZ KWSPJHKVZ KLIPKV H SH KPZVSJP}UKLSHLU]VS[YHUJSLHYKLZWtZKL LZ[VZLPUPJPHUSVZTV]PTPLU[VZJYVTVZ}TPJVZ 3VZL_[YLTVZKLSVZTPJYV[‚ISVZKLSVZmZ[LYLZ ZL TL]LU TLKPHU[L WVSPTLYPaHJP}U OHJPH LS JLU[YV KL SH JtSSH IZJHUKV JYVTVZVTHZ LU UTV]PTPLU[VXLZLWPLUZHXLLZHSHaHY` JHUKV OHJLU JVU[HJ[V JVU U JPUL[VJVYV ZL LZ[HIPSPaHUL]LU[HSTLU[LLSJPUL[VJVYVKLSH V[YHJYVTm[PKHOLYTHUHKLSTPZTVJYVTVZVTH ZLUL[HTIPtUHUNYWVKLTPJYV[‚ISVZXL WYV]PLULU KLS WVSV VWLZ[V :L OH WYVWLZ[V [HTIPtU XL SVZ JPUL[VJVYVZ UV H[YHWHKVZ WLKLUPUPJPHYWVYZxTPZTVZUHUJSLHJP}U KL TPJYV[‚ISVZ +L UH V[YH THULYH SVZ JYVTVZVTHZSSLNHUH[LULYZZKVZJPUL[VJVYVZ JVULJ[HKVZJHKHUVHTPJYV[‚ISVZXLWYV- ]PLULUKLWVSVZVWLZ[VZLULZLTVTLU[VSVZ TPJYV[‚ISVZ QHSHU ` LTWQHU HS JYVTVZVTH TLKPHU[LWVSPTLYPaHJP}UKLZWVSPTLYPaHJP}UKL [ISPUHJVUSHH`KHKLWYV[LxUHZTV[VY[PWV JPULZPUHZ`KPULxUHZ+LIPKVHXLSVZKVZWVSVZ H[YHLUHSTPZTVJYVTVZVTHH[YH]tZKLZZKVZ JPUL[VJVYVZ·XLWLYTHULJLUUPKVZWVYTL- KPVKLSHJVOLZPUH·ZLPUPJPHULZ[HKVLULSXL ZLLTWPLaHUHLXPSPIYHYSHZMLYaHZKL[YHJJP}U KLJHKHWVSVWVSPTLYPaHUKVTmZHJ[P]HTLU[L LSL_[YLTVKLSVZTPJYV[‚ISVZHZVJPHKVZHS JPUL[VJVYVXLZLLUJLU[YHTmZJLYJHUVHZ WVSV ` HJVY[HUKV SVZ TPJYV[‚ISVZ HZVJPHKVZ HS WVSV TmZ SLQHUV -PNYH ,S HJVY[HTPLU[V V LSVUNHJP}U KL SVZ TPJYV[‚ISVZ VILKLJL H SHZKPMLYLUJPHZLUSHZMLYaHZKL[LUZP}ULUSVZ JPUL[VJVYVZ*VULZ[VZTV]PTPLU[VZZLHSJHUaH Figura 7. Polimerización-despolimerización de microtúbulos para formar la placa metafásica. Los JPUL[VJVYVZKLJHKHUHKLSHZKVZJYVTm[PKHZKLSVZ JYVTVZVTHZZLHZVJPHUHTPJYV[‚ISVZXLWYV]PLULU KLWVSVZVWLZ[VZKLSOZVTP[}[PJVNLULYHSTLU[LLU UZP[PVHJtU[YPJVWVYSVXLWHYHSSL]HYSVZHSJLU[YV KLS OZV ` MVYTHY SH WSHJH TL[HMmZPJH SVZ TPJYV[‚- ISVZ TmZ JVY[VZ WVSPTLYPaHU YmWPKHTLU[L OHZ[H LS TVTLU[VLUXLZLHSJHUaHUIHSHUJLKLSVUNP[K JHUKVLZ[VVJYYLSVZJYVTVZVTHZZLLUJLU[YHULU LSLJHKVYJLSSHY`VYPLU[HKVZKL[HSTHULYHXLHS LZJPUKPYZLSVZJLU[Y}TLYVZSHMLYaHKL[YHJJP}USSL]H HUNYWVJYVTVZ}TPJVJVTWSL[VOHJPHUWVSV` HSV[YVNYWVHSWVSVVWLZ[V SHJVUNYLNHJP}UXLLZSHYLUP}UKL[VKVZSVZ JYVTVZVTHZLULSLJHKVYJLSSHYWVZPJPVUHKVZ HOxWVYLSLXPSPIYPVKLSHZMLYaHZKL[YHJJP}U KLSVZWVSVZVWLZ[VZKLSOZV3HJVUNYLNHJP}U JYVTVZ}TPJHTHYJHLSÄUKLSHWYVTL[HMHZL` LSPUPJPVKLSHTL[HMHZL Metafase 3H JVUKLUZHJP}U JYVTVZ}TPJH PUPJPHKH LU SH WYVMHZL JVU[PU‚H WVY SV XL SVZ JYVTVZVTHZ TL[HMmZPJVZ ZL VIZLY]HU JVU UH JYVTH[PUH WLYMLJ[HTLU[LLTWHXL[HKHXLSLWLYTP[LHS TH[LYPHSNLUt[PJVTHU[LULYZPU[LNYPKHKKYHU[L LSLZ[YtZTLJmUPJVKLSVZTV]PTPLU[VZKLHUH- MHZL2,ULZ[HL[HWHLZLUSHXLNLULYHSTLU[L ZLYLHSPaHUSVZLZ[KPVZJYVTVZ}TPJVZKLIPKV HXLZTVYMVSVNxHLZT`JSHYH
  7. 7. 61 Rodríguez-Gómez AJ y Frias-Vázquez S. La mitosis y su regulación 3VZJYVTVZVTHZTV]PKVZWVYLSOZVTP[}[PJV ZLJVSVJHULULSJLU[YVLU[YLSVZKVZmZ[LYLZ` MVYTHUSHSSHTHKHWSHJHTL[HMmZPJH-PNYHI LUSHXLSVZJYVTVZVTHZZLWVZPJPVUHUKL[HS THULYH XL SVZ JPUL[VJVYVZ KL JHKH JYVTm[P- KH OLYTHUH LZ[mU VYPLU[HKVZ OHJPH SVZ WVSVZ VWLZ[VZ2 4HU[LULY H SVZ JYVTVZVTHZ LU LS LJHKVYJLSSHYPTWSPJHULXPSPIYPVLU[YLSHZ MLYaHZKLSVZTPJYV[‚ISVZXL[PLUKLUHTV]LY HSVZJPUL[VJVYVZOHJPHSVZWVSVZVWLZ[VZKL THULYHXLWVZPJPVUHYSVZLULSJLU[YVPTWSPJH UH NYHU JHU[PKHK KL LULYNxH 3H LULYNxH KL SVZ TV]PTPLU[VZ JYVTVZ}TPJVZ WYV]PLUL KL SH WVSPTLYPaHJP}UKLZWVSPTLYPaHJP}U KL SVZ TPJYV[‚ISVZLUSVZXLZL[PSPaHSHJVU]LYZP}U KL .;7 H .+7 ` KL SHZ WYV[LxUHZ TV[VYHZ LU SHZXLZL[PSPaHSHJVU]LYZP}UKL(;7H(+74 ,UJHKHJPUL[VJVYVZLWLKLUHUJSHYLU[YL TPJYV[‚ISVZXLLQLYJLUMLYaHKL[YHJJP}U OHJPHLSWVSVKLSXLWYV]PLULUWVYSVXLSH WSHJHTL[HMmZPJHZLTHU[PLULWVYLSLXPSPIYPV LU[YLSHZMLYaHZVWLZ[HZKLSVZWVSVZZVIYLSVZ JYVTVZVTHZ XL THU[PLULU QU[HZ H ZZ JYV- Tm[PKHZOLYTHUHZWVYSHJVOLZPUHJLU[YVTtYPJH KL THULYH XL JHUKV LZ[H WYV[LxUH ZL YL[PYH KLSJLU[Y}TLYVSHTL[HMHZL[LYTPUH`ZLPUPJPH SHHUHMHZLJVUSHTPNYHJP}UKLSHZJYVTm[PKHZ OLYTHUHZHWVSVZVWLZ[VZ Anafase UH ]La XL [VKVZ SVZ JYVTVZVTHZ ZL LU- JLU[YHU LU LS LJHKVY MVYTHUKV SH WSHJH TL[HMmZPJH SVZ JLU[Y}TLYVZ XL THU[LUxHU UPKHZHSHZJYVTm[PKHZOLYTHUHZZLLZJPUKLU WLYTP[PLUKVXLJHKHJYVTm[PKHTPNYLOHJPHLS WVSVXLLZ[HIHLUJHYHUKVPUPJPmUKVZLHZxSH HUHMHZLXLZLKP]PKLLUHUHMHZL(`HUHMHZL) 3VZTV]PTPLU[VZKLSHHUHMHZL(JVYYLUHJHYNV WYPUJPWHSTLU[LKLSVZTPJYV[‚ISVZJPUL[VJ}- YPJVZXLZLHJVY[HUWVYKLZWVSPTLYPaHJP}ULU HTIVZL_[YLTVZKL[ISPUHJVUPU[LY]LUJP}U KL JPULZPUHZ MHTPSPH LZ[V WLYTP[L QHSHY OHJPH SVZ WVSVZ JLSSHYLZ H SVZ JYVTVZVTHZ ZLUJPSSVZ ·H SVZ XL SSHTHTVZ JYVTm[PKHZ OLYTHUHZJHUKVZLLUJLU[YHUUPKHZWVYLS JLU[Y}TLYV·KP]PKPLUKVLSNLUVTHYLWSPJHKV LUKVZNYWVZJYVTVZ}TPJVZKPWSVPKLZJVU JYVTVZVTHZZLUJPSSVZJHKHUV,USHHUHMHZL) PU[LY]PLULUSVZTPJYV[‚ISVZPU[LYWVSHYLZXLZL LUJHYNHUKLHSLQHYSVZKVZNYWVZJYVTVZ}TPJVZ TLKPHU[LUPUJYLTLU[VLUSHSVUNP[KKLSOZV TP[}[PJV-PNYHJ2 3HHUHMHZL(ZLPUPJPHJHUKVLSJVTWSLQVWYV- TV[VYKLHUHMHZL(7*UPKVH*+*PUKJL SHLZJPZP}UKLSJLU[Y}TLYVKLQHUKVSPIYLZHSHZ JYVTm[PKHZOLYTHUHZWHYHXLJHKHUHTPNYL OHJPHWVSVZVWLZ[VZ,STLJHUPZTVKLHJJP}U KL (7**+* LZ IPXP[PUPaHY UH WYV[LxUH SSHTHKH ZLNYPUH11,12 XL SH LU]xH H KLZ[YJ- JP}UWVYLSWYV[LHZVTH`JVUSHKLZ[YJJP}UKL SH ZLNYPUH XLKH SPIYL UH WYV[LxUH SSHTHKH ZLWHYHZHXLLZUHWYV[LHZHXLHJ[‚HZVIYL SHJVOLZPUHLSJVTWSLQVXLTHU[PLULUPKHZ HSHZJYVTm[PKHZOLYTHUHZ,ULSTVTLU[VLU XLSHJVOLZPUHZLYL[PYHKLSJLU[Y}TLYVL_PZ[L UJHTIPVKL[LUZP}UZVIYLSVZJPUL[VJVYVZSV XLHJ[P]HUHKLZWVSPTLYPaHJP}UKL[ISPUHLU HTIVZL_[YLTVZ`¶KLSVZTPJYV[‚ISVZJPUL- [VJ}YPJVZHSHJVY[HYZLQHSHUHSVZJYVTVZVTHZ UPKVZHLSSVZOHJPHSVZJLU[YVZVTHZIPJHKVZ LUSVZWVSVZJLSSHYLZ-PNYH+YHU[L SHHUHMHZL)SVZKVZNYWVZKLJYVTVZVTHZ`H JVSVJHKVZLUSVZWVSVZZVUHSLQHKVZUVKLV[YV WVYSHPU[LY]LUJP}UKLSVZTPJYV[‚ISVZWVSHYLZ XLKLZWVSPTLYPaHUKxTLYVZKL[ISPUHLUZZ L_[YLTVZ ` WVSPTLYPaHU HJ[P]HTLU[L LU ZZ L_[YLTVZ KPYPNPKVZ HS JLU[YV JLSSHY LZ[VZ TPJYV[ISVZJYLJLU[HU[VXLSVZL_[YLTVZZL JYaHU ` JVU[PU‚HU KLZHYYVSSmUKVZL LU KPYLJ- JPVULZ VWLZ[HZ .YHJPHZ H SH PU[LY]LUJP}U KL WYV[LxUHZTV[VYHZHZVJPHKHZJVUTPJYV[‚ISVZ SVZZLNTLU[VZZWLYWLZ[VZKLSVZTPJYV[‚ISVZ WVSHYLZJVTPLUaHUUWYVJLZVKLKLZSPaHTPLU[V ¸LU YL]LYZH¹ OHJPH SVZ WVSVZ XL OHJL XL LS OZVTP[}[PJVLU[V[HSZLOHNHT`SHYNV`ZLWHYL
  8. 8. Acta Pediátrica de México 62 Volumen 35, Núm. 1, enero-febrero, 2014 KLLZ[HTHULYHHSVZKVZNYWVZKLJYVTVZVTHZ XLOHIxHUHYYPIHKVHSVZWVSVZLUHUHMHZL(2,12 Telofase ,U SH [LSVMHZL SVZ TPJYV[‚ISVZ JPUL[VJ}YPJVZ KLZHWHYLJLU`SVZJYVTVZVTHZXLKHUSPIYLZLU HTIVZL_[YLTVZJLSSHYLZ,ULZLTVTLU[VSVZ JYVTVZVTHZJVTPLUaHUHKLZJVUKLUZHYZL`SH LU]VS[YHUJSLHYZLYLPU[LNYH+YHU[LSH[LSVMH- ZL[HYKxHSVZJYVTVZVTHZLTWPLaHUH[YHUZJYPIPY `ZLYLZ[HISLJLLSUJSLVSVTHYJHUKVLS[tYTPUV KLSHTP[VZPZ2 -PNYHK Citocinesis ,ZSH‚S[PTHL[HWHKLSHKP]PZP}UJLSSHYLULSSHLS JP[VWSHZTHZLKP]PKLWHYHKHYSNHYHKVZJtSSHZ OPQHZ JVTWSL[HTLU[L PUKLWLUKPLU[LZ :PU LT- IHYNVSHJP[VJPULZPZLTWPLaHKLZKLSHHUHMHZL JHUKV ZL MVYTH PUTLKPH[HTLU[L WVY KLIHQV KLSHTLTIYHUHWSHZTm[PJHHUP]LSKLSLJHKVY JLSSHYUJPU[Y}UKLWYV[LxUHZÄSHTLU[VZHZ WYPUJPWHSTLU[LHJ[PUH`TPVZPUHXLZLJVU[YHLU ` KHU SNHY H U ZYJV KL KP]PZP}U iZ[L ZL ]H OHJPLUKVWYVNYLZP]HTLU[LTmZWYVMUKVOHZ[H XL SH JPU[YH XL ZL MVYTH SSLNH H [VJHY SVZ TPJYV[‚ISVZXLH‚UXLKHUYLTHULU[LZKLS OZVTP[}[PJV[VKVLZ[LJVUQU[VKLWYV[LxUHZ ZLJVUVJLJVTVJLYWVTLKPV-PUHSTLU[LSH JtSSHZLLZ[YHUNSH`KHSNHYHKVZJtSSHZOPQHZ -PNYH 14 Figura 8. Anafase A.+YHU[LSHHUHMHZL(LSJVT- WSLQV(7*LU]xHWHYHZKLZ[YJJP}UHSHZLNYPUHXL HZ]LaSPILYHHSHWYV[LxUHZLWHYHZH3HZLWHYHZH HJ[‚HZVIYLSHJVOLZPUHXLTHU[PLULUPKHZSHZKVZ JYVTm[PKHZWVYLSJLU[Y}TLYVLULSJYVTVZVTHLU TL[HMHZLJVSVJHKVLULSLJHKVYJLSSHY3PILYHYHSH JVOLZPUHPTWSPJHSHZLWHYHJP}UKLSHZJYVTm[PKHZXL JLKLUHSHZMLYaHZKLSVZTPJYV[‚ISVZJPUL[VJ}YPJVZ WVYKLZWVSPTLYPaHJP}UKLHTIVZL_[YLTVZQHSHUHSVZ JYVTVZVTHZOHJPHSVZWVSVZ Figura 9. Establecimiento del surco de división con la intervención de las proteínas actina y miosina II. La HZVJPHJP}UHJ[PUHTPVZPUHZLPUPJPHPUTLKPH[HTLU[L WVYKLIHQVKLSHTLTIYHUHWSHZTm[PJH`LUKPYLJJP}U WLYWLUKPJSHYHSJLU[YVKLSOZVTP[}[PJV+YHU[LSH JP[VJPULZPZLZ[LHUPSSVJVU[YmJ[PS]HLZ[YHUNSHUKVSH JtSSHXL`HJVU[PLULKVZU‚JSLVZOPQVZWVYLUTLKPV KLtZ[VZOHZ[HXLXLKHZ}SVUJLYWVTLKPVMVY- THKVWVYTPJYV[‚ISVZYLTHULU[LZKLSOZVTP[}[PJV `SV‚S[PTVKLSHUPSSVKLHJ[PUHTPVZPUH-PUHSTLU[L ZLZLWHYHUSHZKVZJtSSHZOPQHZ
  9. 9. 63 Rodríguez-Gómez AJ y Frias-Vázquez S. La mitosis y su regulación REGULACIÓN MOLECULAR DE LA MITOSIS El ciclo celular se regula gracias a un patrón de ciclinas y cinasas dependientes de ciclinas (CDKs) 7HYHZLZ[KPVLSJPJSVJLSSHYWLKLKP]PKPYZL LU KVZ L[HWHZ! SH PU[LYMHZL PU[LNYHKH WVY SHZ MHZLZ . : ` . ` SH MHZL 4 KYHU[L SH XL ZVIYL]PLULU SH ZLNYLNHJP}U JYVTVZ}TPJH ` SH JP[VJPULZPZ 7HYH XL UH JtSSH H]HUJL H SV SHYNVKLSHZMHZLZKLSJPJSVJLSSHY`ÄUHSTLU[L WLKHZLNYLNHYZTH[LYPHSNLUt[PJVZLYLXPLYL UHYLNSHJP}UT`ÄUHXLZLJHYHJ[LYPaHWVY SHHJ[P]PKHKKLSHZ*+2ZJPUHZHZKLWLUKPLU[LZ KLJPJSPUHXLHJ[‚HUJVTVJPUHZHZKLV[YHZ WYV[LxUHZYLNSHKVYHZKLSJPJSVJLSSHY`THYJHU KLLZ[LTVKVSHWYVNYLZP}UKLUHMHZLKLSJPJSV HV[YH,Z[HZJPUHZHZHSJHUaHUZMVYTHHJ[P]HLU U JVTWSLQV OL[LYVKPTtYPJV JVU UH JPJSPUH YLNSHKVYH -PNYH H 3H L_WYLZP}U KL SHZ JPJSPUHZLZUMHJ[VYSPTP[HU[LWHYHSHHJ[P]HJP}U KLSHZ*+2Z,UNLULYHSSVZUP]LSLZKLJPJSPUHZ LZ[mUKL[LYTPUHKVZWVYJVU[YVS[YHUZJYPWJPVUHS` ZWYV[L}SPZPZH[YH]tZKLSZPZ[LTHKLIPJP[PUH WYV[LVZVTH 3H L_WYLZP}U KL SH JPJSPUH + LZULJLZHYPHKYHU[LSHMHZL.WHYHLZ[PTSHY SHLU[YHKHKLSHZJtSSHZHSJPJSVJLSSHY`ZZ JVUJLU[YHJPVULZWLYTHULJLUHS[HZTPLU[YHZOH`H HNLU[LZ TP[VNtUPJVZ 3H JPJSPUH , ZL YLXPLYL LU MHZL . KLZWtZ KL SH PU[LY]LUJP}U KL SH JPJSPUH+`HSJHUaHZTm_PTHL_WYLZP}ULU SH [YHUZPJP}U .: SV XL SH ]LS]L ULJLZHYPH WHYHSHLU[YHKHHSHMHZLKLZxU[LZPZ3HJPJSPUH( ZLYLXPLYLWHYHLSPUPJPVKL:`SH[YHUZPJP}UKL .4TPLU[YHZXLSHJPJSPUH)YLNSHSHLU[YHKH `ZHSPKHKLTP[VZPZ-PNYHI El punto de control de G2 regula la entrada a mitosis +LZWtZKLSHZxU[LZPZKLS(+5`WHYHYLKJPY SHHJTSHJP}UKLLYYVYLZNLUt[PJVZSHZJtSSHZ TVUP[VYLHU XL LS (+5 OH`H ZPKV YLWSPJHKV JVYYLJ[HTLU[LZPKL[LJ[HHSN‚ULYYVYLSJPJSVJL- SSHYZLKL[PLUL`ZPUVKL[LJ[HLYYVYLZSHJtSSH WYVNYLZHYmOHJPHTP[VZPZ,Z[LWU[VKLJVU[YVS ZLKLUVTPUH.V.4*HUKVLULS(+5 L_PZ[LKH|VZLHJ[P]HUSHZJPUHZHZ(;4T[HKV LUH[H_PH[LSHUNPLJ[HZPH`(;9YLSHJPVUHKVJVU (;4`9HK3HWYPTLYHKL[LJ[HYVTWPTPLU[VZ KL JHKLUH KVISL LU LS (+5 TPLU[YHZ XL SH ZLNUKHKL[LJ[HYVTWPTPLU[VZKLJHKLUHZLU- JPSSH3HZJPUHZHZ(;4(;9WYVWHNHUSHZL|HS Figura 10. 7H[Y}UKLHJ[P]PKHKKLSHZ*+2Z`SHZJPJSPUHZHSVSHYNVKLSJPJSVJLSSHY
  10. 10. Acta Pediátrica de México 64 Volumen 35, Núm. 1, enero-febrero, 2014 KLHSLY[HHWV`HKHZWVYSHZWYV[LxUHZTLKPHKVYHZ KLS WU[V KL JVU[YVS XL PUJS`LU H 4+* TLKPHKVY KLS WU[V KL YL]PZP}U KL KH|V HS (+5[HTIPtUJVUVJPKVJVTV5-)+)7 WYV[LxUHKLUP}UHW`)9*(WYV[LxUHKLS JmUJLY KL THTH(KLTmZ ZL YLXPLYL KL SHZ JPUHZHZ[YHUZKJ[VYHZ*/2`*/2(;4 (;9MVZMVYPSHUHSHOPZ[VUH/(?a/(?`LZ[V THYJHSHYLNP}UKLSHJYVTH[PUHXLÅHUXLH JHKHYVTWPTPLU[VHZ]LaSHa/(?WLYTP[L LSHNYWHTPLU[VKLSHZWYV[LxUHZYLWHYHKVYHZLU LSZP[PVKLSKH|V3VZTLKPHKVYLZ)74+* `)9*(MUJPVUHUJVTVUWLU[LTVSLJSHY LU[YLSHOPZ[VUHa/(?`SHZWYV[LxUHZKLSJVT- WSLQVYLWHYHKVY49549,9(+5): ,SJVTWSLQV495`HLULSZP[PVKLSKH|VSSL]H HJHIVSHYLWHYHJP}UKLSYVTWPTPLU[VKLKVISL JHKLUHKLSHOLIYHKLS(+5TLKPHU[LYLJVT- IPUHJP}UOVT}SVNH3HZJtSSHZL]P[HUXLZL SSLNLHSHTP[VZPZJVUKH|VJYVTVZ}TPJVKL THULYH XL TPLU[YHZ ZL LMLJ[‚H SH YLWHYHJP}U KLS(+5 LS WU[V KL JVU[YVS KL . ISVXLH SH HJ[P]PKHK KLS MHJ[VY WYVTV[VY KL SH TP[VZPZ 47-JVTWLZ[VWVYSHJPJSPUH)`SH*+2 3HHJ[P]PKHKKLS47-WLKLZLYPUOPIPKHWVYSH ZL|HSPaHJP}U H [YH]tZ KL (;4(;9 TLKPHU[L KVZHJ[P]PKHKLZWYPUJPWHSLZ! 3HZ JPUHZHZ KL SH MHTPSPH KL W WLKLU PUOPIPYHSHZMVZMH[HZHZ*+*LZ[HZ‚S[PTHZ HJ[P]HUHSJVTWSLQV47-LULSSxTP[LKLSHZ MHZLZ.4WVYSVXLHSZLYPUOPIPKHZUV ZLLU[YHHTP[VZPZ3HZJPUHZHZ*/2 */2WLKLUHJ[P]HYH,,XLLZU YLNSHKVY ULNH[P]V KLS JVTWSLQV 47- V IPLU HJ[P]HY ZLJLUJPHSTLU[L H W ` H .(++UZLJLZ[YHKVYKLJPJSPUH)+L LZ[LTVKVZLL]P[HXLSHJPJSPUH)LU[YLHS U‚JSLV`UVZLMVYTHLSJVTWSLQV47- 2.3. La transición G2/M (entrada a mitosis) 3HWYVNYLZP}UKLSHMHZL.OHJPH4ZLKLILH SHHJ[P]HJP}UKLSJVTWSLQV47-J`HHJ[P]PKHK ZLTHU[PLULKLSHWYVMHZLOHZ[HSHTL[HMHZL,S JVTWSLQV 47- MVZMVYPSH HWYV_PTHKHTLU[L ZZ[YH[VZTP[}[PJVZXLHJ[‚HULUSHZLWHYHJP}U KLSVZJLU[YVZVTHZSHMVZMVYPSHJP}U`HJ[P]HJP}U KLLUaPTHZYLNSHKVYHZKLSHJVUKLUZHJP}UKL SH JYVTH[PUH MVZMVYPSHJP}U KL SH OPZ[VUH / YW[YH KL SH LU]VS[YH UJSLHY ` YLVYNHUPaH- JP}U KLS JP[VLZXLSL[V KL TPJYV[‚ISVZ ` KL HJ[PUHWHYHYLHSPaHYSHTP[VZPZ3HHJ[P]PKHK KLS JVTWSLQV 47- ZL YLNSH H KVZ UP]LSLZ LS WYPTLYVLZ[YHUZJYPWJPVUHSWVYSHWYV[LxUHW XL THU[PLUL SH [YHUZJYPWJP}U ` WVY SV [HU[V SHWYLZLUJPHKL*+2LZ[HISLKYHU[L[VKVLS JPJSVJLSSHY`SHKLJPJSPUH)KLZKLÄUHSLZKL SHMHZL:,SZLNUKVUP]LSKLYLNSHJP}UKLSH HJ[P]PKHKKL47-LZWVYTVKPÄJHJPVULZWVZ[YH- KJJPVUHSLZLZWLJxÄJHTLU[LWVYMVZMVYPSHJP}U PUOPIP[VYPH`HJ[P]HKVYHLUSHMHZL.ZLYLHSPaHU MVZMVYPSHJPVULZ PUOPIP[VYPHZ ZVIYL SH *+2 WVY SHZ JPUHZHZ 4@; `,, ` WVZ[LYPVYTLU[L SH MVZMH[HZH*+*SPILYHSHPUOPIPJP}UH*+2 LZ[VQU[VJVUUHMVZMVYPSHJP}UHJ[P]HKVYHZVIYL SHTPZTH*+2YLHSPaHKHWVY*(2JPUHZHHJ- [P]HKVYHKL*+2UOL[LYVKxTLYVKLJPJSPUH/ `*+2JVUKJLHSHJtSSHHLU[YHYHTP[VZPZ 3H HJ[P]PKHK WYVTV[VYH KL TP[VZPZ KL *+2 WLKL PUOPIPYSH KPYLJ[HTLU[L LS NLU ZWYLZVY KL[TVYW*PWHM,UYLSHJP}UJVUSH JPJSPUH)ZLYLXPLYLUHMVZMVYPSHJP}UWHYHJHT- IPHYZSVJHSPaHJP}UZIJLSSHYKLJP[VWSHZTHH U‚JSLVLUSHZL[HWHZ[LTWYHUHZKLSHTP[VZPZ3H ZHSPKH KL TL[HMHZL ZL JHYHJ[LYPaH WVY SH JHxKH JYx[PJHKLSHZJVUJLU[YHJPVULZKLJVTWSLQV47- KLIPKVHZKLZ[YJJP}U *HUKV SH JtSSH OH WHZHKV LS WU[V KL JVU- [YVSKLSHMHZL.ZZJYVTVZVTHZKWSPJHKVZ PUPJPHU LS WYVJLZV KL JVUKLUZHJP}U NYHJPHZ H SVZ JVTWSLQVZ KL JVUKLUZPUH 0 ` 00 XL MU- JPVUHULTWHXL[HUKVSHJYVTH[PUHKLSLZ[HKV PU[LYMmZPJV H JYVTVZVTH TP[}[PJV WHYH XL WLKHZLYZLNYLNHKHHWYVWPHKHTLU[LLU[YLSHZ JtSSHZ OPQHZ ,U LS JVTWSLQV JVUKLUZPUH
  11. 11. 65 Rodríguez-Gómez AJ y Frias-Vázquez S. La mitosis y su regulación 0 ZL LUJLU[YHU SHZ WYV[LxUHZ :4* Structural maintenance of chromosomeJVUHJ[P]PKHKKL (;7HZH JVTV :4**(7, ` :4**(7* ` WYV[LxUHZUV:4*JVTV*(7.`*(7/ El JVTWSLQVJVUKLUZPUH00JVUZPZ[LKLSHZZIUP- KHKLZ:4*`:4*`SHZWYV[LxUHZO*(7+ O*(7.`O*(7/(SWHYLJLYLSJVT- WSLQV0MUJPVUHLUWYVTL[HMHZL`LSJVTWSLQV 00LUWYVMHZL[LTWYHUH+YHU[LLSLTWHXL- [HTPLU[VSHZJYVTm[PKHZOLYTHUHZWLYTHULJLU UPKHZNYHJPHZHSHZJVOLZPUHZXLZLJHYNHULU SVZJYVTVZVTHZLUSHMHZLZ.[HYKxH:[LTWYHUH `ZLTHU[PLULUOHZ[HXLZLHSJHUaHSHIPVYPLU[H- JP}UKLSHZJYVTm[PKHZLUTL[HMHZL ,SWYVJLZV KLJVUKLUZHJP}UKLSVZJYVTVZVTHZ`SHLU[YH- KHHTP[VZPZWLKLUYL]LY[PYZLLUWYLZLUJPHKL LZ[YtZVJVUKPJPVULZUV}W[PTHZWHYHSHKP]PZP}U JLSSHYNYHJPHZHSHHJ[P]HJP}UKLSHZWYV[LxUHZ KLSWU[VKLJVU[YVSKLHU[LMHZL 3HZWYV- [LxUHZPU]VSJYHKHZLULZLWYVJLZVJVTPLUaHU HWLUHZHZLYKLZJYP[HZWLYV*/-9Checkpoint with FHA and RING domains`SHJPUHZHW`H ZLJVUZPKLYHUWHY[LLZLUJPHSKLSWYVJLZV*/-9 LZUHSPNHZHKLIPJP[PUHZXLZLHJTSHLU YLZWLZ[HHTPJYV[‚ISVZKH|HKVZ`[PLULLU[YL ZZISHUJVZHWYV[LxUHZPTWVY[HU[LZWHYHSHWYV- NYLZP}U TP[}[PJH JVTV(YVYH( ` 732 XL ZVUTHYJHKHZWHYHZKLZ[YJJP}UWVYTLKPVKLS ZPZ[LTHIPJP[PUHWYV[LVZVTHKLLZ[LTVKVZL YL[YHZHSHWYVNYLZP}UTP[}[PJH7VYV[YHWHY[L SHJPUHZHWXLZLHJ[P]HLUJHZVKLKH|VWVY YHKPHJP}US[YH]PVSL[HVHS[LYHJP}UKLSHLZ[YJ- [YHKLSHJYVTH[PUHL]P[HSHLU[YHKHHTP[VZPZ :LOHLZWLJSHKVXLLSISHUJVPUOPIP[VYPVKLS WU[VKLJVU[YVSKLSHHU[LMHZLLZLSJVTWSLQV KLJVUKLUZPUH00 2.4. El punto de control de metafase ,U[YL SVZ WU[VZ KL JVU[YVS LZWLJPHSPaHKVZ LU KL[LULYSHWYVNYLZP}UTP[}[PJHLSTmZLZ[KPH- KV ` XPam LS WYPUJPWHS LZ LS JVUVJPKV JVTV :(*:WPUKSL(ZZLTIS`*OLJRWVPU[VWU[VKL JVU[YVSTP[}[PJV,S:(*ZLHJ[P]HLUJHKHJPJSV JLSSHY PUTLKPH[HTLU[L KLZWtZ KL SH LU[YHKH H TP[VZPZ ` MUJPVUH YL[YHZHUKV SH LU[YHKH H HUHMHZLOHZ[HXL[VKVZSVZJYVTVZVTHZLZ[mU JVYYLJ[HTLU[LHUJSHKVZLUSHWSHJHTL[HMmZPJH IPVYPLU[HKVZ`JVU[LUZP}UHKLJHKH *HUKV LS HSPULHTPLU[V KL SVZ JYVTVZVTHZ ZH[PZMHJLHS:(*ZLWYVJLKLHSHZLNYLNHJP}UKL SVZJYVTVZVTHZ,Z[LWYVJLZVZLPUPJPHJHUKV LSJVMHJ[VYWYV[LPJV*+*ZLUL`HJ[P]HHS JVTWSLQV WYVTV[VY KL SH HUHMHZLJPJSVZVTH (7** XL [PLUL MUJP}U KL SPNHZH KL IP- JP[PUHZ*HUKVLS(7**ZLHJ[P]H[YHUZÄLYL [YLZIPJP[PUHZHSHJPJSPUH)`HSHWYV[LxUH ZLNYPUH THYJmUKVSHZ KL LZ[L TVKV WHYH Z KLNYHKHJP}U ]xH WYV[LVZVTH : 3H ZLNYPUH LZ PTWVY[HU[L WVYXL ZL LUJHYNH KL THU[LULY ZLJLZ[YHKH H SH WYV[LHZH ZLWHYHZH JHUKV tZ[H LZ SPILYHKH NYHJPHZ H SH KLNYHKHJP}U KL SH ZLNYPUH [PLUL SNHY SH KLNYHKHJP}U KL SH JVOLZPUH XL THU[PLUL UPKHZ H SH JYVTm[P- KHZ OLYTHUHZ 3H KLNYHKHJP}U KL SH JVOLZPUH WLYTP[LSHZLNYLNHJP}UJYVTVZ}TPJHTPLU[YHZ XLSHKLNYHKHJP}UKLSHJPJSPUH)[PLULJVTV YLZS[HKVSHPUHJ[P]HJP}UKLSH*+2NYHJPHZH LZ[VZLJVUZPNLZHSPYKLTP[VZPZ *HUKVLS HSPULHTPLU[V KL SVZ JYVTVZVTHZ UV ZH[PZMHJL HS:(*SHZL|HSWYV]LUPLU[LKLJPUL[VJVYVZUV HUJSHKVZPUKJLLSYLJS[HTPLU[VKLWYV[LxUHZ KLSWU[VKLJVU[YVSJVTV4(+KLÄJPLU[LLU SHKL[LUJP}UTP[}[PJH))NLTHJP}UUVPU- OPIPKHWVYILUaPTPKHaVS`))9YLSHJPVUHKV JVU)),Z[HZL|HSJVUKJLHSHMVYTHJP}UKL KVZJVTWSLQVZPUKLWLUKPLU[LZUVJVTWLZ[V WVY))))9`LSV[YVWVY*4(+*+* 3HUP}UKLLZ[VZKVZJVTWSLQVZKHYmSNHYHS 4**JVTWSLQVTP[}[PJVKLJVU[YVSLULSXL `HOHXLKHKVZLJLZ[YHKV*+*PUOPIPLUKV HZxSHHJ[P]HJP}UKLSJVTWSLQV(7**+LLZ[L TVKV SH ZLNYPUH UV ZLYm THYJHKH WHYH Z KLNYHKHJP}U SH ZLWHYHZH UV ZL SPILYHYm ` SVZ JYVTVZVTHZ UV ZLNYLNHYmU OHZ[H XL SVZ YL- XLYPTPLU[VZKLS:(*ZLZH[PZMHNHU (WLZHY KLSHS[VNYHKVKLLZWLJPHSPaHJP}UKLS:(*tZ[L
  12. 12. Acta Pediátrica de México 66 Volumen 35, Núm. 1, enero-febrero, 2014 UVLZJHWHaKLYLJVUVJLYSHZUPVULZLYY}ULHZ LU[YLLSJPUL[VJVYV`SVZTPJYV[‚ISVZJVUVJPKHZ JVTVUPVULZTLYV[tSPJHZZPULTIHYNVLZ[HZ ‚S[PTHZWLKLUYLWHYHYZLNYHJPHZHSHHJ[P]PKHK KLS JVTWSLQV *7* *OYVTVZVTL 7HZZLUNLY *VTWSL_ ,S JVTWSLQV *7* LZ[m MVYTHKV WVY SHZWYV[LxUHZ:9ZY]P]PUH)69IVYLHSPUH 05*,57WYV[LPUHJLU[YVTtYPJH`(Y)HYVYH JPUHZH ) ` OHZ[H HOVYH LZ LS TLQVY JHUKPKH[V WHYH ZLY LS JVTWSLQV ZLUZVY KL [LUZP}U KL SHZ UPVULZHSJPUL[VJVYV3HHJ[P]PKHKKL(Y)LZ WYPTVYKPHS WVYXL KL[LJ[H ` KLZLZ[HIPSPaH SHZ UPVULZ LYY}ULHZ LU[YL SVZ TPJYV[‚ISVZ ` SVZ JPUL[VJVYVZ [HSLZ JVTV UPVULZ ZPU[tSPJHZ ` TLYV[tSPJHZ KL LZ[L TVKV NLULYH JPUL[VJVYVZ UV HUJSHKVZ XL ZVU KL[LJ[HKVZ WVY LS :(* ` ZLJVUKJLHSHKL[LUJP}UKLSHTP[VZPZ(SSSL- NHYHHUHMHZLSH(Y)ZLKLZWSHaHKLSHYLNP}U JLU[YVTtYPJH+YHU[LSHWYVTL[HMHZL*+* ` [VKHZ SHZ WYV[LxUHZ KLS :(* ZL JVUJLU[YHU LU SVZ JPUL[VJVYVZ NYHJPHZ HS YLJS[HTPLU[V LMLJ[HKVWVYSHZJPUHZHZ))47:`(Y) WLYVZLHJ[P]HUHU[LSHZZL|HSLZKLJPUL[VJVYVZ UVHUJSHKVZ7HYHLSZPSLUJPHTPLU[VKLS:(* ZLOHUWYVWLZ[V]HYPVZTLJHUPZTVZ,SWYPTL- YVJVUZPZ[LLUYL[PYHYWVYTLKPVKLSHKPULxUH H SHZ WYV[LxUHZ KLS :(* LU SVZ TPJYV[‚ISVZ JVYYLJ[HTLU[L HUJSHKVZ LS ZLNUKV LZ LS KL ZPSLUJPHTPLU[VXLVJYYLJHUKVSHWYV[LxUH *,57, ZL UL H SVZ TPJYV[‚ISVZ L PUOPIL SH HJ[P]PKHK JPUHZH KL ))9 LS [LYJLY WYVJLZV KL ZPSLUJPHTPLU[V LZ LS TLKPHKV WVY SH UP}U PUOPIP[VYPHKLW*64,;H4(+*+*L PUJSZVSHKLNYHKHJP}UKLSVZJVTWVULU[LZKLS :(* WVY LS ZPZ[LTH KL IPJP[PUHWYV[LVZVTH TLKPHKHWVYLSJVTWSLQV(7** 3H WYVNYLZP}U H [YH]tZ KL TP[VZPZ UV Z}SV LZ[m YLNSHKH WVY SVZ WU[VZ KL JVU[YVS ZPUV XL [HTIPtU LZ[m xU[PTHTLU[L YLSHJPVUHKH JVU SH HJ[P]PKHKKLS(7**XLLUSHTP[VZPZ[HYKxHZL HZVJPHJVU*+/`WLYTHULJLHJ[P]HH[YH]tZ KLSHMHZL.ZPNPLU[L(7***+*KLNYHKH HSHJPJSPUH)`HSHJVOLZPUHJHUKV[VKVZSVZ JYVTVZVTHZ OHU OLJOV JVU[HJ[V JVU LS OZV IPWVSHY ` ZL TL]LU H [YH]tZ KL SH WSHJH TL- [HMmZPJHSVXLJVUKJLHSHZLWHYHJP}UKLSHZ JYVTm[PKHZOLYTHUHZ7VYZWHY[LLSJVTWSLQV (7**+/THYJHWHYHKLNYHKHJP}UHSHZJPUHZHZ [PWVWVSV732HYVYH(ZY]P]PUH5,2 *+*`:27KYHU[LSHHUHMHZL`KYHU[LSH MHZL.ZPNPLU[L3VZUP]LSLZKL*+/WLYTH- ULJLUYLSH[P]HTLU[LJVUZ[HU[LZKYHU[L[VKVLS JPJSV JLSSHY ` Z HJ[P]PKHK LZ YLNSHKH ZVIYL [VKVH[YH]tZKLSHZMVZMVYPSHJPVULZKLWLUKPLU[LZ KLSJPJSVJLSSHY3HMVZMVYPSHJP}UKL*+/WVY SHZ*+2ZKYHU[L:.`4[LTWYHUHPUOPILZ UP}UH(7**TPLU[YHZXLZKLZMVZMVYPSHJP}U LUSHMHZL4[HYKxHWLYTP[LZUP}UH(7**` SHHJ[P]HJP}UKLSJVTWSLQV-PNYH 2.5. Regulación de la citocinesis 3HJP[VJPULZPZUVKLILVJYYPYZPUVOHZ[HXLSH ZLWHYHJP}UKLSVZJYVTVZVTHZZLOHJVTWSL[H- KV ,U NLULYHS ZL JVUZPKLYH XL SH LQLJJP}U HWYVWPHKH KL SH JP[VJPULZPZ YLJHL LU SH VYNH- UPaHJP}ULZ[YJ[YHSJLSSHY`UVLUTVStJSHZ KPMZPISLZZPULTIHYNVLZ[KPVZYLJPLU[LZOHU LUJVU[YHKV XL SHZ KLÄJPLUJPHZ LU SH JPUHZH */2WLKLU[LULYJVTVJVUZLJLUJPHLYYVYLZ LU SH ZLNYLNHJP}U JYVTVZ}TPJH YLNYLZP}U KL SH JP[VJPULZPZ ` IPUJSLHJP}U (ZPTPZTV LZ[H KLÄJPLUJPH JVYYLSHJPVUH JVU UH SVJHSPaHJP}U LYY}ULHKLSHZWYV[LxUHZ(Y)`*/2XLZL HJTSHU LU LS ZYJV KL ZLNTLU[HJP}U WHYH MHJPSP[HYSHWYVKJJP}UKLKVZJtSSHZOPQHZ,Z[V WVULKLTHUPÄLZ[VXLSHZWYV[LxUHZXLYLNSHU SVZWU[VZKLJVU[YVSKLSJPJSVJLSSHY[HTIPtU WLKLU[LULYMUJPVULZPTWVY[HU[LZKYHU[LSH ZLNYLNHJP}UJYVTVZ}TPJH`SHJP[VJPULZPZ 2.6. Modificaciones covalentes de las histonas durante la mitosis 3HZ OPZ[VUHZ ZVU U NYWV KL WYV[LxUHZ JVU- ZLY]HKHZ XL [PLULU U WHWLS JYx[PJV LU LS LTWHXL[HTPLU[V KLS (+5 KL SHZ JtSSHZ
  13. 13. 67 Rodríguez-Gómez AJ y Frias-Vázquez S. La mitosis y su regulación LJHYPVU[LZ /H` JH[YV [PWVZ KL OPZ[VUHZ U- JSLVZVTHSLZ/(/)/`/`SHOPZ[VUH /KLUP}UHS(+5LZWHJPHKVY3HZOPZ[VUHZ WVZLLU MUJPVULZ YLNSH[VYPHZ XL ZL SSL]HU H JHIVNYHJPHZHSHZTVKPÄJHJPVULZJV]HSLU[LZKL SVZYLZPKVZKLHTPUVmJPKVZKLZZJVSHZ:L OHUKLZJYP[VTJOVZ[PWVZKLTVKPÄJHJPVULZLU SHZOPZ[VUHZXLPUJS`LUMVZMVYPSHJP}UHJL[PSH- JP}UTL[PSHJP}UIPJP[PUHJP}U`ZTVPSHJP}U 3HZKPMLYLU[LZJVTIPUHJPVULZKLTVKPÄJHJPVULZ OPZ[}UPJHZ ZL OHU WYVWLZ[V JVTV U J}KPNV XL PKLU[PÄJH KPMLYLU[LZ LZ[HKVZ MUJPVUHSLZ KL SH JYVTH[PUH LZ[VZ J}KPNVZ WVZLLU U WH- WLS YLSL]HU[L LU SH YLNSHJP}U LWPNLUt[PJH KL SH L_WYLZP}U KL SVZ NLULZ ,Z[HZ TVKPÄJHJPV- ULZ [HTIPtU ZVU YLSL]HU[LZ WHYH LS WYVJLZV TP[}[PJV 3H MVZMVYPSHJP}U KL /: ZLYPUH KL SH OPZ[VUH ` /: ZLYPUH KL SH PZ[VUH ZL YLXPLYL WHYH SH JVUKLUZHJP}U KL SVZJYVTVZVTHZ`ZZLNYLNHJP}U3HTL[PSHJP}U KL/2 SPZPUH KLSHOPZ[VUHWYVTL]LSH MVYTHJP}UKLOL[LYVJYVTH[PUHLPUOPILSHMVZ- MVYPSHJP}UKL/:`]PJL]LYZHSVXLPUKPJH UHPU[LYHJJP}ULU[YLLZ[HZKVZTVKPÄJHJPVULZ 3HMVZMVYPSHJP}UKL/:L]P[HSHUP}UKLSH WYV[LxUH/7WYV[LxUHMVYTHKVYHKLOL[LYVJYV- TH[PUHXLZLKPZVJPHKLSHJYVTH[PUHKYHU[L SH TP[VZPZ JHUKV VJYYL SH KLZMVZMVYPSHJP}U KL/:SHWYV[LxUH/7WLKLUPYZLHJYV- TH[PUH (SWHYLJLYSHIPJP[PUHJP}UKLSH OPZ[VUH/(PUOPILSHHJ[P]PKHKKLSHHYVYH( LUL[HWHZXLUVZLHUSHTP[VZPZTPLU[YHZXLZ KLZIPJP[PUHJP}UZLYLXPLYLWHYHXLSH/: ZLMVZMVYPSL`VJYYHSHZLNYLNHJP}UJYVTVZ}TPJH +YHU[L SH KP]PZP}U JLSSHY SH HJL[PSHJP}U ` SH TL[PSHJP}UKLOPZ[VUHZZVUL]LU[VZHU[HN}UPJVZ JVUKPZTPUJP}UKLSHHJL[PSHJP}ULPUJYLTLU[V KLSHTL[PSHJP}UKYHU[LLSWYVJLZV REFERENCIAS 1. ^ĂůĂƺŶ W͕ ZĂŶŶŽƵ z͕ WƌŝŐĞŶƚ ͘ ĚŬϭ͕ WƐ͕ ƵƌŽƌĂƐ͕ ĂŶĚ EƐ͗ ƚŚĞ ŵŝƚŽƟĐ ďŽĚLJŐƵĂƌĚƐ͘ Adv Exp Med Biol ϮϬϬϴ͖ ϲϭϳ͗ϰϭʹϱϲ͘ 2. tĂůĐnjĂŬ ͕ Ăŝ ^ LJ ŚŽĚũĂŬŽǀ ͘ DĞĐŚĂŶŝƐŵƐ ŽĨ ĐŚƌŽ- mosome behaviour during mitosis. Nat Rev Mol Cell Biol ϮϬϭϬ͖ϭϭ͗ϵϭͲϭϬϮ͘ 3. ,ŝƌĂŶŽd͘ƚƚŚĞŚĞĂƌƚŽĨƚŚĞĐŚƌŽŵŽƐŽŵĞ͗^DƉƌŽƚĞŝŶƐŝŶ ĂĐƟŽŶ͘Nat Rev Mol Cell Biol ϮϬϬϲ͖ϳ͗ϯϭϭͲϯϮϮ͘ 4. Sharp DJ, Rogers GC, Scholey JM. Microtubule motors in mitosis. Nature ϮϬϬϬ͖ϰϬϳ͗ϰϭͲϰϳ͘ 5. KĞŐĞŵĂ ͕ ,LJŵĂŶ ͘ Ğůů ĚŝǀŝƐŝŽŶ ;:ĂŶƵĂƌLJϭϵ͕ ϮϬϬϲͿ͘ tŽƌŵŽŽŬ͕ ĞĚ͘ dŚĞ C. elegans Research Community, tŽƌŵŽŽŬ͕ĚŽŝͬϭϬ͘ϭϴϵϱͬǁŽƌŵŬ͘ϭ͘ϳϮ͘ϭ͕ŚƩƉ͗ͬͬǁǁǁ͘ wormbook.org. ϲ͘ 'ƺƫŶŐĞƌ ^͕ ĂƵƌĞůů ͕ ƵƚĂLJ h͘ KƌĐŚĞƐƚƌĂƟŶŐ ŶƵĐůĞĂƌ envelope disassembly and reassembly during mitosis. Nat Rev Mol Cell Biol ϮϬϬϵ͖ϭϬ͗ϭϳϴͲϭϵϭ͘ 7. WĞĐŽƚDz͕DĂůŚŽƚƌĂs͘'ŽůŐŝŵĞŵďƌĂŶĞƐƌĞŵĂŝŶƐĞŐƌĞ- ŐĂƚĞĚĨƌŽŵƚŚĞĞŶĚŽƉůĂƐŵŝĐƌĞƟĐƵůƵŵĚƵƌŝŶŐŵŝƚŽƐŝƐŝŶ mammalian cells. Cell ϮϬϬϰ͖ϭϭϲ͗ϵϵʹϭϬϳ͘ 8. tŝƩŵĂŶŶd͕,LJŵĂŶ͕ĞƐĂŝ͘dŚĞƐƉŝŶĚůĞ͗ĂĚLJŶĂŵŝĐ assembly of microtubules and Motors. Nat Cell BiolϮϬϬϭ͖ ϯ͗ϮϴͲϯϰ͘ 9. ĂŵƉĞƌƚ͕tĞƐƚĞƌŵĂŶŶ^͘ďůƵĞƉƌŝŶƚĨŽƌŬŝŶĞƚŽĐŚŽƌĞƐͶ new insights into the molecular mechanics of cell division Nat Rev Mol Cell Biol ϮϬϭϭ͖ϭϮ͗ϰϬϳͲϰϭϮ͘ 10. ŬŝLJŽƐŚŝ͕^ĂƌĂŶŐĂƉĂŶŝ͕WŽǁĞƌƐ͕EĞůƐŽŶZ͕ZĞŝĐŚŽǁ SL, Arellano-Santoyo H, Gonen T, et al . Tension directly sta- ďŝůŝnjĞƐƌĞĐŽŶƐƟƚƵƚĞĚŬŝŶĞƚŽĐŚŽƌĞͲŵŝĐƌŽƚƵďƵůĞĂƩĂĐŚŵĞŶƚƐ EĂƚƵƌĞϮϬϭϬ͖ϰϲϴ͗ϱϳϲͲϱϴϭ͘ 11. ĐƋƵĂǀŝǀĂ͕WŝŶĞƐ:͘dŚĞĂŶĂƉŚĂƐĞͲƉƌŽŵŽƟŶŐĐŽŵƉůĞdžͬ ĐLJĐůŽƐŽŵĞ͗ Wͬ͘ ϮϬϬϲ͘ :ŽƵƌŶĂů ŽĨ Ğůů ^ĐŝĞŶĐĞ ϮϬϬϲ͖ ϭϭϵ͗ϮϰϬϭͲϰ͘ 12. Sullivan M, Morgan DO. Finishing mitosis, one step at a ƟŵĞ͘EĂƚZĞǀDŽůĞůůŝŽůϮϬϬϳ͖ϴ͗ϴϵϰͲϵϬϯ͘ 13. WŝŶĞƐ:͘DŝƚŽƐŝƐ͗ĂŵĂƩĞƌŽĨŐĞƫŶŐƌŝĚŽĨƚŚĞƌŝŐŚƚƉƌŽƚĞŝŶ ĂƚƚŚĞƌŝŐŚƚƟŵĞ͘Trends Cell Biol ϮϬϬϲ͖ϭϲ͗ϱϱͲϲϯ͘ 14. EŽƌĚĞŶ͕DĞŶĚŽnjĂD͕ŽďďĞůĂĞƌĞ:͘dŚĞEŽƵƚWĂƚŚǁĂLJ ŝŶŬƐŽŵƉůĞƟŽŶŽĨLJƚŽŬŝŶĞƐŝƐƚŽ^ƉŝŶĚůĞDŝĚnjŽŶĞƵŶĐƟŽŶ ƚŽWƌĞǀĞŶƚŚƌŽŵŽƐŽŵĞƌĞĂŬĂŐĞ͘ϮϬϬϲCellϭϮϱ͗ϴϱʹϵϴ͘ 15. ǀĂŶ ĚĞŶ ,ĞƵǀĞů͕ ^͘ ĞůůͲĐLJĐůĞ ƌĞŐƵůĂƟŽŶ ;^ĞƉƚĞŵďĞƌ Ϯϭ͕ ϮϬϬϱͿ͕tŽƌŵŽŽŬ͕ĞĚ͘dŚĞC. elegans Research Commu- ŶŝƚLJ͕ tŽƌŵŽŽŬ͕ ĚŽŝͬϭϬ͘ϭϴϵϱͬǁŽƌŵŬ͘ϭ͘Ϯϴ͘ϭ͕ ŚƩƉ͗ͬͬ www.wormbook.org. ϭϲ͘ DĂůƵŵďƌĞƐD͕ĂƌďĂĐŝĚD͘ĞůůĐLJĐůĞ͕ƐĂŶĚĐĂŶĐĞƌ͗ ĂĐŚĂŶŐŝŶŐƉĂƌĂĚŝŐŵ͘EĂƚZĞǀĂŶĐĞƌϮϬϬϵ͖ϵ͗ϭϱϯͲϭϲϲ͘ 17. ƌĂŶnjĞŝ͕ŽŝĂŶŝD͘ZĞŐƵůĂƟŽŶŽĨEƌĞƉĂŝƌƚŚƌŽƵŐŚŽƵƚ the cell cycle Nat Rev Mol Cell Biol ϮϬϬϴ͖ϯ͗ϮϵϳͲϯϬϴ͘ 18. ZĞŝŶŚĂƌĚƚ,͕zĂīĞD͘ŝŶĂƐĞƐƚŚĂƚĐŽŶƚƌŽůƚŚĞĐĞůůĐLJĐůĞ ŝŶƌĞƐƉŽŶƐĞƚŽEĚĂŵĂŐĞ͗ŚŬϭ͕ŚŬϮ͕ĂŶĚDϮ͘Ƶƌƌ KƉŝŶĞůůŝŽůϮϬϬϵ͖Ϯϭ͗ϮϰϱʹϮϱϱ͘ 19. ŝƚĂŐĂǁĂZ͕ĂŬŬĞŶŝƐƚ:͕DĐŝŶŶŽŶW:͕ĂƐƚĂŶD͘WŚŽƐ- ƉŚŽƌLJůĂƟŽŶŽĨ^DϭŝƐĂĐƌŝƟĐĂůĚŽǁŶƐƚƌĞĂŵĞǀĞŶƚŝŶƚŚĞ dDʹE^ϭʹZϭƉĂƚŚǁĂLJ͘Genes DevϮϬϬϰ͖ϭϴ͗ϭϰϮϯʹϯϴ͘
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