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. 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. 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. 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
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SVZTPJYV[‚ISVZZLYLVYNHUPaHUWHYHJVU[YPIPY
H SH MVYTHJP}U KLS OZV TP[}[PJV ,Z[H UL]H
VYNHUPaHJP}UKLSVZTPJYV[‚ISVZZLPUPJPHWVY
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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-
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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
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SSVZLZYLNSHKHWVYWYV[LxUHZHZVJPHKHZKLSHMHTPSPH
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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. 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
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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
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IPLUXLZLPU[LNYLUHMYHNTLU[VZKLYL[xJSV
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.VSNPZLKLZPU[LNYHLUWLXL|HZ]LZxJSHZJ`V
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,U JHSXPLYH KL SVZ KVZ JHZVZ SHZ ]LZxJSHZ
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[LZLNYLNHYZLLUMYHNTLU[VZKLtZ[LOHJPHSHZ
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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
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KLTPJYV[‚ISVZJPUL[VJ}YPJVZXLHWU[HHL_[YLTVZ
VWLZ[VZ3VZTPJYV[‚ISVZPU[LYWVSHYLZJYLJLUWVYZZ
L_[YLTVZWVZP[P]VZZPULUJVU[YHYUJPUL[VJVYVOHZ[H
XLZLZWLYWVULUJVUV[YVTPJYV[‚ISVPU[LYWVSHY

6. Acta Pediátrica de México
60
Volumen 35, Núm. 1, enero-febrero, 2014
ZLZLNYLNHUKLTHULYHNLULYHSTLU[LZPTt[YPJH
HSHZJtSSHZOPQHZ
Prometafase
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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
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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
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]PLULUKLWVSVZVWLZ[VZLULZLTVTLU[VSVZ
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TLKPHU[LWVSPTLYPaHJP}UKLZWVSPTLYPaHJP}UKL
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JPULZPUHZ`KPULxUHZ+LIPKVHXLSVZKVZWVSVZ
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JPUL[VJVYVZ·XLWLYTHULJLUUPKVZWVYTL-
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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. 61
Rodríguez-Gómez AJ y Frias-Vázquez S. La mitosis y su regulación
3VZJYVTVZVTHZTV]PKVZWVYLSOZVTP[}[PJV
ZLJVSVJHULULSJLU[YVLU[YLSVZKVZmZ[LYLZ`
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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
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TPJYV[‚ISVZXLLQLYJLUMLYaHKL[YHJJP}U
OHJPHLSWVSVKLSXLWYV]PLULUWVYSVXLSH
WSHJHTL[HMmZPJHZLTHU[PLULWVYLSLXPSPIYPV
LU[YLSHZMLYaHZVWLZ[HZKLSVZWVSVZZVIYLSVZ
JYVTVZVTHZ XL THU[PLULU QU[HZ H ZZ JYV-
Tm[PKHZOLYTHUHZWVYSHJVOLZPUHJLU[YVTtYPJH
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OLYTHUHZHWVSVZVWLZ[VZ
Anafase
UH ]La XL [VKVZ SVZ JYVTVZVTHZ ZL LU-
JLU[YHU LU LS LJHKVY MVYTHUKV SH WSHJH
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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
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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
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SH ZLNYPUH XLKH SPIYL UH WYV[LxUH SSHTHKH
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HSHZJYVTm[PKHZOLYTHUHZ,ULSTVTLU[VLU
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JVSVJHKVZLUSVZWVSVZZVUHSLQHKVZUVKLV[YV
WVYSHPU[LY]LUJP}UKLSVZTPJYV[‚ISVZWVSHYLZ
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L_[YLTVZ ` WVSPTLYPaHU HJ[P]HTLU[L LU ZZ
L_[YLTVZ KPYPNPKVZ HS JLU[YV JLSSHY LZ[VZ
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JPVULZ VWLZ[HZ .YHJPHZ H SH PU[LY]LUJP}U KL
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SVZZLNTLU[VZZWLYWLZ[VZKLSVZTPJYV[‚ISVZ
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¸LU YL]LYZH¹ OHJPH SVZ WVSVZ XL OHJL XL LS
OZVTP[}[PJVLU[V[HSZLOHNHT`SHYNV`ZLWHYL

8. Acta Pediátrica de México
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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
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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
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`SV‚S[PTVKLSHUPSSVKLHJ[PUHTPVZPUH-PUHSTLU[L
ZLZLWHYHUSHZKVZJtSSHZOPQHZ

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
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SHHJ[P]PKHKKLSHZ*+2ZJPUHZHZKLWLUKPLU[LZ
KLJPJSPUHXLHJ[‚HUJVTVJPUHZHZKLV[YHZ
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El punto de control de G2 regula la entrada a
mitosis
+LZWtZKLSHZxU[LZPZKLS(+5`WHYHYLKJPY
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10. Acta Pediátrica de México
64
Volumen 35, Núm. 1, enero-febrero, 2014
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11. 65
Rodríguez-Gómez AJ y Frias-Vázquez S. La mitosis y su regulación
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12. Acta Pediátrica de México
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Volumen 35, Núm. 1, enero-febrero, 2014
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13. 67
Rodríguez-Gómez AJ y Frias-Vázquez S. La mitosis y su regulación
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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

14. Acta Pediátrica de México
68
Volumen 35, Núm. 1, enero-febrero, 2014
20. ^ŚŝďĂƚĂ͕ĂƌƚŽŶ͕EŽŽŶd͕ĂŚŵ͕ĞĐŬďĂƌ͕'ŽŽĚĂƌnjŝ
͕ƂďƌŝĐŚ D͕:ĞŐŐŽ W͘ ZŽůĞ ŽĨ dD ĂŶĚ ƚŚĞ ĂŵĂŐĞ
ZĞƐƉŽŶƐĞ DĞĚŝĂƚŽƌ WƌŽƚĞŝŶƐ ϱϯWϭ ĂŶĚ Dϭ ŝŶ ƚŚĞ
DĂŝŶƚĞŶĂŶĐĞ ŽĨ 'ϮͬD ŚĞĐŬƉŽŝŶƚ ƌƌĞƐƚ͘ Mol Cell Biol
ϮϬϭϬ͖ϯϬ͗ϯϯϳϭʹϴϯ͘
21. ^ƚƵĐŬŝD͕:ĂĐŬƐŽŶ^W͘DϭͬEϭ͗ĂŬĞLJƌĞŐƵůĂƚŽƌŽĨƚŚĞ
DNA damage response in higher eukaryotes. DNA Repair
(Amst)ϮϬϬϰ͖ϯ͗ϵϱϯʹϳ͘
22. ĂƐƚĂŶDE͕ĂƌƚĞŬ:͘ĞůůͲĐLJĐůĞĐŚĞĐŬƉŽŝŶƚƐĂŶĚĐĂŶĐĞƌ͘
NatureϮϬϬϰ͖ϰϯϮ͗ϯϭϲͲϮϯ͘
23. EŝŐŐ͘DŝƚŽƟĐŬŝŶĂƐĞƐĂƐƌĞŐƵůĂƚŽƌƐŽĨĐĞůůĚŝǀŝƐŝſŶĂŶĚŝƚƐ
checkpoints. Nat Rev Mol Cell BiolϮϬϬϭ͖Ϯ͗ϮϭͲϯϮ͘
24. ƂďƌŝĐŚD͕:ĞŐŐŽW͘dŚĞŝŵƉĂĐƚŽĨĂŶĞŐůŝŐĞŶƚ'ϮͬDĐŚĞĐ-
ŬƉŽŝŶƚŽŶŐĞŶŽŵŝĐŝŶƐƚĂďŝůŝƚLJĂŶĚĐĂŶĐĞƌŝŶĚƵĐƟŽŶNat Rev
CancerϮϬϬϳ͖ϳ͗ϴϲϭͲϴϲϵ͘
25. DĂůƵŵďƌĞƐD͕ĂƌďĂĐŝĚD͘DĂŵŵĂůŝĂŶĐLJĐůŝŶͲĚĞƉĞŶĚĞŶƚ
ŬŝŶĂƐĞƐ͘dƌĞŶĚƐŝŽĐŚĞŵ͘^Đŝ͕͘ϮϬϬϱ͖ϯϬ͕ϲϯϬʹϰϭ͘
Ϯϲ͘ ĞůŵŽŶƚ^͘DŝƚŽƟĐĐŚƌŽŵŽƐŽŵĞƐƚƌƵĐƚƵƌĞĂŶĚĐŽŶĚĞŶ-
ƐĂƟŽŶ͘Curr Opin Cell Biol ϮϬϬϲ͖ϭϴ͗ϲϯϮʹϴ͘
27. ,ŝƌĂŶŽ d͘ ŽŶĚĞŶƐŝŶƐ͗ ŽƌŐĂŶŝnjŝŶŐ ĂŶĚ ƐĞŐƌĞŐĂƚŝŶŐ ƚŚĞ
genome. Curr Biol ϮϬϬϱ͖ϭϱ͗Ϯϲϱʹϳϱ͘
28. ,ŝƌŽƚĂd͕'ĞƌůŝĐŚ͕ŽĐŚ͕ůůĞŶďĞƌŐ:͕WĞƚĞƌƐ:D͘ŝƐƟŶĐƚ
ĨƵŶĐƟŽŶƐ ŽĨ ĐŽŶĚĞŶƐŝŶ / ĂŶĚ // ŝŶ ŵŝƚŽƟĐ ĐŚƌŽŵŽƐŽŵĞ
assembly. J Cell Sci ϮϬϬϰ͖ϭϭϳ͗ϲϰϯϱʹϰϱ͘
29. ŚŝŶ͕zĞŽŶŐD͘^ĂĨĞŐƵĂƌĚŝŶŐĞŶƚƌLJŝŶƚŽDŝƚŽƐŝƐ͗ƚŚĞ
Antephase Checkpoint. Mol Cell BiolϮϬϭϬ͖ϯϬ͗ϮϮʹϯϮ͘
30. ,ĂŐƐƚƌŽŵ ͕ DĞLJĞƌ :͘ ŽŶĚĞŶƐŝŶ ĂŶĚ ĐŽŚĞƐŝŶ͗ ŵŽƌĞ
than chromosome compactor and glue. Nat Rev Genet
ϮϬϬϯ͖ϰ͗ϱϮϬʹϯϰ͘
31. ,ŝƌĂŶŽd͘ƚƚŚĞŚĞĂƌƚŽĨƚŚĞĐŚƌŽŵŽƐŽŵĞ͗^DƉƌŽƚĞŝŶƐŝŶ
ĂĐƟŽŶEĂƚZĞǀDŽůĞůůŝŽůϮϬϬϲ͖ϳ͗ϯϭϭͲϯϮϮ͘
32. KŶŽd͕ĂŶŐz͕^ƉĞĐƚŽƌ͕,ŝƌĂŶŽd͘^ƉĂƟĂůĂŶĚƚĞŵƉŽƌĂů
ƌĞŐƵůĂƟŽŶŽĨĐŽŶĚĞŶƐŝŶƐ/ĂŶĚ//ŝŶŵŝƚŽƟĐĐŚƌŽŵŽƐŽŵĞ
assembly in human cells. Mol Biol CellϮϬϬϰ͖ϭϱ͗ϯϮϵϲʹϯϬϴ͘
33. zĞŽŶŐD͕,ŽŵďĂƵĞƌ,͕tĞŶĚƚ^͕DƵĚƌĂŬ/͕DĞĐŚƚůĞƌ͕
ŽƌĞŐŐĞƌd͕DĂƌĐŚůĞƌͲĂƵĞƌ͕dĂŶĂŬĂ͕WĞƚĞƌƐ:D͕KŐƌŝƐ
͘/ĚĞŶƟĮĐĂƟŽŶŽĨĂƐƵďƵŶŝƚŽĨĂŶŽǀĞůůĞŝƐŝŶͲďĞƚĂͬ^D
ĐŽŵƉůĞdžĂƐĂƉŽƚĞŶƟĂůƐƵďƐƚƌĂƚĞŽĨƉƌŽƚĞŝŶƉŚŽƐƉŚĂƚĂƐĞ
2A. Curr BiolϮϬϬϯ͖ϭϯ͗ϮϬϱϴʹϲϰ͘
34. Mikhailov A, Shinohara M, Rieder CL. The p38-mediated
ƐƚƌĞƐƐͲĂĐƟǀĂƚĞĚĐŚĞĐŬƉŽŝŶƚ͘ƌĂƉŝĚƌĞƐƉŽŶƐĞƐLJƐƚĞŵĨŽƌ
delaying progression through antephase and entry into
mitosis. Cell CycleϮϬϬϱ͖ϰ͗ϱϳʹϲϮ͘
35. ĂŶŐ͕ŚĞŶ:͕tŽŶŐ:͕ĂŶŐ'͘dŚĞĐŚĞĐŬƉŽŝŶƚƉƌŽƚĞŝŶ
,ZŝƐĂůŝŐĂƐĞƚŚĂƚƵďŝƋƵŝƟŶĂƚĞƐWϭĂŶĚŝŶŚŝďŝƚƐϮ
ĂƚƚŚĞ'ϮƚŽDƚƌĂŶƐŝƟŽŶ͘J Cell Biol ϮϬϬϮ͖ϭϱϲ͗ϮϰϵʹϮϱϵ͘
ϯϲ͘ ŝŵ:^͕WĂƌŬzz͕WĂƌŬ^z͕ŚŽ,͕ĂŶŐ͕ŚŽ,͘dŚĞƵƚŽͲ
ƵďŝƋƵŝƚLJůĂƟŽŶŽĨŚĨƌĂƚ'ϮWŚĂƐĞ/ƐZĞƋƵŝƌĞĚĨŽƌĐĐƵ-
ŵƵůĂƟŽŶŽĨWůŬϭĂŶĚDŝƚŽƟĐŶƚƌLJŝŶDĂŵŵĂůŝĂŶĞůůƐ͘
:ŝŽůŚĞŵϮϬϭϭ͘
37. Mikhailov A, Shinohara M, Rieder CL. Topoisomerase II and
ŚŝƐƚŽŶĞĚĞĂĐĞƚLJůĂƐĞŝŶŚŝďŝƚŽƌƐĚĞůĂLJƚŚĞ'ϮͬDƚƌĂŶƐŝƟŽŶ
ďLJƚƌŝŐŐĞƌŝŶŐƚŚĞƉϯϴDWĐŚĞĐŬƉŽŝŶƚƉĂƚŚǁĂLJ͘J Cell Biol
ϮϬϬϰ͖ϭϲϲ͗ϱϭϳʹϮϲ͘
38. Musacchio A, Salmon ED. The spindle-assembly checkpoint
ŝŶƐƉĂĐĞĂŶĚƟŵĞ͘Nat Rev Mol Cell BiolϮϬϬϳ͖ϴ͗ϯϳϵͲϵϯ͘
39. WŝŶĞƐ:͕ZŝĞĚĞƌ͘ZĞͲƐƚĂŐŝŶŐŵŝƚŽƐŝƐ͗ĂĐŽŶƚĞŵƉŽƌĂƌLJǀŝĞǁ
ŽĨŵŝƚŽƟĐƉƌŽŐƌĞƐƐŝŽŶ͘Nat Cell BiolϮϬϬϭ͖ϯ͗ϯͲϲ͘
40. WĠƌĞnjĚĞĂƐƚƌŽ/͕ĚĞĄƌĐĞƌ'͕DĂůƵŵďƌĞƐD͘ĐĞŶ-
ƐƵƐŽĨŵŝƚŽƚŝĐĐĂŶĐĞƌŐĞŶĞƐ͗ŶĞǁŝŶƐŝŐŚƚƐŝŶƚŽƚƵŵŽƌ
cell biology and cancer therapy. CarcinogenesisϮϬϬϳ͖
Ϯϴ͗ϴϵϵͲϵϭϮ͘
41. ŝƚĐŚĮĞůĚ͕:ŽŚŶƐŽŶs͕dŝŐŚĞ͕ůůƐƚŽŶZ͕,ĂǁŽƌƚŚƐ͕
Johnson T, Mortlock A, Keen A, Keen N, Taylor SS. Aurora
ĐŽƵƉůĞƐĐŚƌŽŵŽƐŽŵĞĂůŝŐŶŵĞŶƚǁŝƚŚĂŶĂƉŚĂƐĞďLJƚĂƌ-
ŐĞƟŶŐƵďZϭ͕DĂĚϮ͕ĂŶĚĞŶƉͲƚŽŬŝŶĞƚŽĐŚŽƌĞƐ͘:Ğůů
ŝŽůϮϬϬϯ͖ϭϲϭ͗ϮϲϳͲϴϬ͘
42. ŽƉƐ':͕tĞĂǀĞƌ͕ůĞǀĞůĂŶĚt͘KŶƚŚĞƌŽĂĚƚŽĐĂŶĐĞƌ͗
ĂŶĞƵƉůŽŝĚLJ ĂŶĚ ƚŚĞ ŵŝƚŽƟĐ ĐŚĞĐŬƉŽŝŶƚ͘ Nat Rev Cancer
ϮϬϬϱ͖ϱ͗ϳϳϯͲϴϱ͘
43. ,ŽǁĞůů :͕ DĐǁĞŶ ͕ ĂŶŵĂŶ :͕ ,ŽīŵĂŶ ͕ ĂƌƌĂƌ
D͕Z/ĞĚĞƌ͕^ĂůŵŽŶ͘LJƚŽƉůĂƐŵŝĐĚLJŶĞŝŶͬĚLJŶĂĐƟŶ
drives kinetochore protein transport to the spindle poles
ĂŶĚŚĂƐĂƌŽůĞŝŶŵŝƚŽƟĐƐƉŝŶĚůĞĐŚĞĐŬƉŽŝŶƚŝŶĂĐƟǀĂƟŽŶ͘J
Cell BiolϮϬϬϭ͖ϭϱϱ͗ϭϭϱϵʹϳϮ͘
44. DĂŽz͕ĞƐĂŝ͕ůĞǀĞůĂŶĚt͘DŝĐƌŽƚƵďƵůĞĐĂƉƚƵƌĞďLJ
EWͲ ƐŝůĞŶĐĞƐ ƵďZϭͲĚĞƉĞŶĚĞŶƚ ŵŝƚŽƚŝĐ ĐŚĞĐŬƉŽŝŶƚ
signaling. J Cell BiolϮϬϬϱ͖ϭϳϬ͗ϴϳϯʹϴϬ͘
45. yŝĂ'͕ƵŽy͕,ĂďƵd͕ZŝnjŽ:͕DĂƚƐƵŵŽƚŽd͕zƵ,͘ŽŶĨŽƌ-
ŵĂƟŽŶͲƐƉĞĐŝĮĐ ďŝŶĚŝŶŐ ŽĨ Ɖϯϭ;ĐŽŵĞƚͿ ĂŶƚĂŐŽŶŝnjĞƐ ƚŚĞ
ĨƵŶĐƟŽŶŽĨDĂĚϮŝŶƚŚĞƐƉŝŶĚůĞĐŚĞĐŬƉŽŝŶƚ͘EMBO JϮϬϬϰ͖
Ϯϯ͗ϯϭϯϯʹϰϯ͘
ϰϲ͘ WĂůĨƌĂŵĂŶt:͕DĞĞŚů:͕:ĂƐƉĞƌƐĞŶ^͕tŝŶĞLJD͕DƵƌƌĂLJ
t͘ ŶĂƉŚĂƐĞ ŝŶĂĐƚŝǀĂƚŝŽŶ ŽĨ ƚŚĞ ƐƉŝŶĚůĞ ĐŚĞĐŬƉŽŝŶƚ͘
ScienceϮϬϬϲ͖ϯϭϯ͗ϲϴϬʹϲϴϰ͘
47. WĞĚĚŝƚůĂ ^͕ ZŽƐĞŶ :͘ ŚĞĐŬŝŶŐ ĂŶĚ ĞdžĞĐƵƚŝŶŐ ĐĞůů
division to prevent genomic instability. Cell Cycle 2009
ϴ͗ϭϱ͕ϮϯϯϵͲϰϮ͘
48. Luger K, Richmond TJ. The histone tails of the nucleosome.
Curr Opin Genet Dev ϭϵϵϴ͖ϴ͗ϭϰϬͲϲ͘
49. yƵ͕Ăŝ:͕ƵĂŶY͕ŽƐƚĂD͕Ăŝt͘ŽǀĂůĞŶƚŵŽĚŝĮĐĂƟŽŶƐ
of histones during mitosis and meiosis. Cell CycleϮϬϬϵ͖
ϴ͗ϯϲϴϴͲϵϰ͘
50. ŝƐƐĞŶďĞƌŐ:͕ůŐŝŶ^͘dŚĞ,WϭƉƌŽƚĞŝŶĨĂŵŝůLJ͗ŐĞƫŶŐĂ
ŐƌŝƉŽŶĐŚƌŽŵĂƟŶ͘Curr Opin Genet DevϮϬϬϬ͖ϭϬ͗ϮϬϰͲϭϬ͘
51. ŝƐĐŚůĞt͕dƐĞŶŐ^͕ŽƌŵĂŶŶ,͕hĞďĞƌŚĞŝĚĞD͕'ĂƌĐŝĂ
͕^ŚĂďĂŶŽǁŝƚnj:͕,ƵŶƚ͕ƵŶĂďŝŬŝ,͕ůůŝƐ͘ZĞŐƵůĂƟŽŶ
ŽĨ,WϭͲĐŚƌŽŵĂƟŶďŝŶĚŝŶŐďLJŚŝƐƚŽŶĞ,ϯŵĞƚŚLJůĂƟŽŶĂŶĚ
ƉŚŽƐƉŚŽƌLJůĂƟŽŶ͘NatureϮϬϬϱ͖ϰϯϴ͗ϭϭϭϲͲϮϮ͘
52. :ŽŽ,z͕ŚĂŝ͕zĂŶŐ͕^ŚƵLJŝEŝĞ͕ƌĚũƵŵĞŶƚͲƌŽŵĂŐĞ,͕
dĞŵƉƐƚW͕ŚĂŶŐ͕tĂŶŐ,͘ZĞŐƵůĂƟŽŶŽĨĐĞůůĐLJĐůĞƉƌŽ-
ŐƌĞƐƐŝŽŶ ĂŶĚ ŐĞŶĞ ĞdžƉƌĞƐƐŝŽŶ ďLJ ,Ϯ ĚĞƵďŝƋƵŝƟŶĂƟŽŶ͘
Nature ϮϬϬϳ͖ϰϰϵ͗ϭϬϲϴͲϳϮ͘