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Biologia  do  Desenvolvimento	
Fertilização	
Prof.  Pedro  Vale  de  Azevedo  Brito	
No  Momento  da  Fertilização	
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Desafios  para  o  espermatozoide...	
Fecundação  Externa    x    Fecundação  Interna	
-­‐‑Alcançar  o  ovócito	
-­‐‑Ultrapassar  barreiras	
-­‐‑Garantir  fertilização  espécie  -­‐‑  
específica	
-­‐‑Alcançar  o  ovócito	
-­‐‑Ultrapassar  barreiras	
Desafios  para  o  espermatozoide...	
Espermatozoide  precisa  fundir-­‐‑se  à  membrana  celular  para  fertilização  	
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Maioria  dos  Invertebrados	
Envelope  Vitelino	
(membrana  vitelina)	
-­‐‑Glicoproteínas	
-­‐‑Extensões  de  proteínas  da                                  
membrana  plasmática	
Mamíferos	
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Zona  Pelúcida  =  Envelope  
	
 	
 	
            Vitelino	
Corona  Radiata	
Vamos  nos  concentrar  em  dois  modelos	
Ouriço-­‐‑do-­‐‑mar	
Camundongo	
Fertilização  no  Ouriço-­‐‑do-­‐‑Mar	
Como  o  ambiente  marinho  é  compartilhado  por  várias  espécies,  dois  mecanismos  
foram  desenvolvidos:	
Atração  e  Ativação  espécie-­‐‑específica	
Atração  espécie-­‐‑específica  –  Espermatozoides  atraídos  por  quimiotaxia	
-­‐‑Cnidários	
-­‐‑Moluscos	
-­‐‑Equinodermas	
-­‐‑Anfíbios  	
-­‐‑Urocordados	
Fertilização  no  Ouriço-­‐‑do-­‐‑Mar	
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,+0! 0$*'&-! )*! '! -0'(*%$%F0'($! 20+-$)(C! '(/! K#$(! )-! F)(/*! 0$*'&-! +(! -#$! $.-0'&$AABA'0! *)/$C! '!
&+(,+0%'-)+('A!&#'(1$!+(! -#$!&H-+2A'*%)&! *)/$!'&-)L'-$*! -#$! 0$&$2-+0U*! $(5H%'-)&! '&-)L)-H?!"#)*!
'&-)L'-$*! -#$! %)-+&#+(/0)'A! O"VE1$($0'-)(1! '22'0'-B*! '*! K$AA! '*! -#$! /H($)(! O"V'*$! -#'-!
*-)%BA'-$*!,A'1$AA'0!%+L$%$(-!)(!-#$!*2$0%!3S#)%+%B0'!$-!'A?!679R:!W++X!'(/!Q'F&+&X!677=>?!!
The acrosomal reaction in sea urchins!
O!*$&+(/!)(-$0'&-)+(!F$-K$$(!*2$0%!'(/!$11!)*!-#$!acrosomal reaction.!Y(!%+*-!%'0)($!
)(L$0-$F0'-$*C! -#$! '&0+*+%'A! 0$'&-)+(! #'*! -K+! &+%2+($(-*Z! -#$! ,B*)+(! +,! -#$! '&0+*+%'A! L$*)&A$!
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'&0+*+%'A!0$'&-)+(!)(!*$'!B0&#)(*!)*!)()-)'-$/!FH!&+(-'&-!+,!-#$!*2$0%!K)-#!-#$!$11!G$AAH?!W+(-'&-!
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20+-$+AH-)&!$(5H%$*! -#'-!&'(!/)1$*-!'!2'-#! -#0+B1#! -#$! G$AAH!&+'-! -+! -#$!$11!*B0,'&$!3['(!67R8:!
M0'(XA)(!678N:!\$L)($!$-!'A?!6789>?!"#$!*$]B$(&$!+,!-#$*$!$L$(-*!)*!+B-A)($/!)(!M)1B0$!8?6N?!!
Espermatozoides  
tornam-­‐‑se   móveis  
quando   em   contato  
com  a  água  do  mar  
(pH),   nadam   em  
círculos	
1   n l   d e   u m a  
solução   10   nM   de  
resact	
Com  a  difusão  do  
f a t o r ,   m a i s  
espermatozoides  
são  atraídos	
Arbacia  punctulata  –  resact  (isolado  a  partir  da  camada  gelatinosa  do  ovo)	
Aumenta  produção  de  ATP  (mitocôndrias)  e  atividade  das  dineínas  =  ativação	
Em  algumas  espécies  a  atração  e  a  ativação  são  realizadas  por  moléculas  diferentes	
Atração  -­‐‑  Quimiotaxia	
-­‐‑  Seletiva  mesmo  entre  espécies  próximas	
-­‐‑  Os  ovócitos  conseguem  regular  o  momento  da  liberação  do  quimiotático	
Mecanismo  de  ação  do  resact	
Reação  Acrossômica	
Nos  invertebrados  marinhos  a  reação  acrossômica  tem  dois  componentes:	
1-­‐‑  Fusão  da  vesícula  acrossômica	
2-­‐‑  Extensão  do  processo  acrossomal	
!
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$%%&!',!(=96!+!-'2=&%!9#,9%,&/+&'#,<!+,-!6#B!9+,!()%/$!7%!)/%0%,&%-!3/#$!&/?',8!&#!3%/&'2'4%!%88(!
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Sperm attraction: Action at a distance!
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(2'-%(<!+,-!()%/$!B%/%!/%2%+(%-!+&!+!9%/&+',!-'(&+,9%!3/#$!&6%!%88(:!@'22%/!3#=,-!&6+&!B6%,!()%/$!
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B%/%!/%+-?!&#!7%!3%/&'2'4%-<!&6%!()%/$!$'8/+&%-!&#B+/-!&6%$:!A6=(<!&6%(%!##9?&%(!9#,&/#2!,#&!#,2?!
&6%!&?)%!#3!()%/$!&6%?!+&&/+9&<!7=&!+2(#!&6%!&'$%!+&!B6'96!&6%?!+&&/+9&!&6%$:!
A  reação  acrossômica  começa  quando  o  
espermatozoide  entra  em  contato  com  a  
camada  gelatinosa  doovo	
As   enzimas   proteolít icas   abrem  
caminho  através  da  camada  gelatinosa	
Uma   vez   que   o   envelope   vitelino   é  
alcançado,  proteossomos  o  digerem	
Reação  Acrossômica	
!
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/.0%)'0%$!+#-.!&)-+#!5+0&/%#-$!:I+0#%4!%-!&0C!=>D?HC!!
 
Species-specific recognition in sea urchins!
Polissacarídeos  sulfatados  na  camada  gelatinosa  do  ovo  são  reconhecidos  pelos  
espermatozoides  para  início  da  reação  acrossômica	
Stongylocentrotus  purpuratus	
Reconhecimento  do  polímero  repetitivo  de  fucose  sulfato  desencadeia:	
1-­‐‑  Entrada  de  Ca+  no  espermatozoide	
2-­‐‑  Bomba  Na-­‐‑H  –  entrada  de  Na  -­‐‑  saída  de  H	
3-­‐‑  Enzima  fosfolipase  produz  inositol  3  fosfato  IP3	
Aumento  de  Ca+  
citoplasmático  e  do  pH  
levam  à  fusão  da  vesícula  
acrossomal	
Aumento  de  Ca+	
Ativação  da  proteína  
RhoB	
Polimerização  de  
actina  no  processo  
acrossomal	
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Species-specific recognition in sea urchins!
Bindin	
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:!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!G%%#!,(52)&%/!7P,*(&,!%&!)2:!;BB9M!V&%)+(!)#/!Q%##)+R!;BB9>:!!
! ! ! ! @',(! HJI?FK)! -+5&%,#! .)3! ')O%! (%O%+)2! +%0,5#(! &')&!! ! ! ! ! :!!!!!!!!!!!!!!!!
:!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!,#&%+)$&!E,&'!G,#/,#:!=&!2%)(&!5#%!56!&'%(%!(,&%(!!!!!!!!!!!!!:!!!!!!!!!!!!!!!!!!!!!!!!
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::!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!5&'%+!(,&%!5+!(,&%(!)--%)+!&5!+%$50#,R%!)!:!!:!::!!!!!!!!!!!!!!!!
[!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!0%#%+)2!G,#/,#!(&+*$&*+%!)#/!$)#!+%$50#,R%!:!!!!
N!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!&'%!G,#/,#!56!.)#3!(-%$,%(:!@'%!G,#/,#!!!!!!!!!!!!!!!!!
L!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!+%$%-&5+(!)+%!&'5*0'&!&5!G%!)00+%0)&%/!,#&5!$5.-2%U%(!5#!&'%!%00!
N!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!$%22!(*+6)$%4!)#/!'*#/+%/(!56!&'%(%!$5.-2%U%(!.)3!G%!#%%/%/!&5!!!!!!!!!!!!!!!!!!!!!!!!!!!
N!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!&!!!!!%&'%+!&'%!(-%+.!&5!&'%!%00!78,0*+%!9:;XL>:!!
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(-%$,%(! 78,0*+%!9:;<LM!C)$D*,%+!)#/!N53!;B99>:!8*+&'%+4! ,&(! ,#&%+)$&,5#!E,&'!%00(! ,(! +%2)&,O%23!
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)$+5(5.%(! 56! S. purpuratus!
G,#/(!&5!,&(!5E#!/%1%22,%/!%00(4!
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punctulata.!! ! !!!
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:!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!G%%#!,(52)&%/!7P,*(&,!%&!)2:!;BB9M!V&%)+(!)#/!Q%##)+R!;BB9>:!!
! ! ! ! @',(! HJI?FK)! -+5&%,#! .)3! ')O%! (%O%+)2! +%0,5#(! &')&! ! ! ! ! ! :!!!!!!!!!!!!!!!!
:!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!,#&%+)$&!E,&'!G,#/,#:!=&!2%)(&!5#%!56!&'%(%!(,&%(!!!!!!!!!!!!!:!!!!!!!!!!!!!!!!!!!!!!!!
Z!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!+%$50#,R%(!5#23!&'%!G,#/,#!56!&'%!().%!(-%$,%(:!@'%!
::!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!5&'%+!(,&%!5+!(,&%(!)--%)+!&5!+%$50#,R%!)!:!!:!::!!!!!!!!!!!!!!!!
[!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!0%#%+)2!G,#/,#!(&+*$&*+%!)#/!$)#!+%$50#,R%!:!!!!
N!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!&'%!G,#/,#!56!.)#3!(-%$,%(:!@'%!G,#/,#!!!!!!!!!!!!!!!!!
L!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!+%$%-&5+(!)+%!&'5*0'&!&5!G%!)00+%0)&%/!,#&5!$5.-2%U%(!5#!&'%!%00!
N!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!$%22!(*+6)$%4!)#/!'*#/+%/(!56!&'%(%!$5.-2%U%(!.)3!G%!#%%/%/!&5!!!!!!!!!!!!!!!!!!!!!!!!!!!
N!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!&!!!!!%&'%+!&'%!(-%+.!&5!&'%!%00!78,0*+%!9:;XL>:!!
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(-%$,%(! 78,0*+%!9:;<LM!C)$D*,%+!)#/!N53!;B99>:!8*+&'%+4! ,&(! ,#&%+)$&,5#!E,&'!%00(! ,(! +%2)&,O%23!
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G,#/(!&5!,&(!5E#!/%1%22,%/!%00(4!
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punctulata.!! ! !!!
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:!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!G%%#!,(52)&%/!7P,*(&,!%&!)2:!;BB9M!V&%)+(!)#/!Q%##)+R!;BB9>:!!
! ! ! ! @',(! HJI?FK)! -+5&%,#! .)3! ')O%! (%O%+)2! +%0,5#(! &')&! ! ! ! ! ! :!!!!!!!!!!!!!!!!
:!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!,#&%+)$&!E,&'!G,#/,#:!=&!2%)(&!5#%!56!&'%(%!(,&%(!!!!!!!!!!!!!:!!!!!!!!!!!!!!!!!!!!!!!!
Z!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!+%$50#,R%(!5#23!&'%!G,#/,#!56!&'%!().%!(-%$,%(:!@'%!
::!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!5&'%+!(,&%!5+!(,&%(!)--%)+!&5!+%$50#,R%!)!:!!:!::!!!!!!!!!!!!!!!!
[!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!0%#%+)2!G,#/,#!(&+*$&*+%!)#/!$)#!+%$50#,R%!:!!!!
N!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!&'%!G,#/,#!56!.)#3!(-%$,%(:!@'%!G,#/,#!!!!!!!!!!!!!!!!!
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N!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!$%22!(*+6)$%4!)#/!'*#/+%/(!56!&'%(%!$5.-2%U%(!.)3!G%!#%%/%/!&5!!!!!!!!!!!!!!!!!!!!!!!!!!!
N!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!&!!!!!%&'%+!&'%!(-%+.!&5!&'%!%00!78,0*+%!9:;XL>:!!
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)&&+)$&,5#4!(-%+.!)$&,O)&,5#4!)#/!(-%+.!)/'%(,5#!&5!&'%!%00!(*+6)$%:!
Proteína  insolúvel,  presente  no  processo  acrossomal,  capaz  de  reconhecer  
receptores  no  envelope  vitelino	
O  reconhecimento  espermático  nos  ouriços-­‐‑do-­‐‑mar  ocorrem  na:	
-­‐‑atração  espermática	
-­‐‑ativação  espermática	
-­‐‑reação  acrossômica	
-­‐‑adesão  à  superfície  do  ovo	
Fusão  entre  as  membranas  do  Espermatozoide  e  do  Ovo	
Processo  ativoDependente  de  proteínas  fusogênicas  –  entre  elas  bindin	
Cone  de  fertilização	
Polimerização  de  actina	
Polispermia	
Problemas...	
!
!
The fast block to polyspermy.!
"#$! fast block to poly-spermy! %&! '(#%$)$*! +,!
(#'-.%-.! /#$! $0$(/1%(! 23/$-/%'0! 34! /#$! $..! 20'&5'!
5$5+1'-$6! "#%&!5$5+1'-$! 213)%*$&! '! &$0$(/%)$! +'11%$1!
+$/7$$-!/#$!$..!(,/320'&5!'-*!/#$!38/&%*$!$-)%13-5$-/9!
'-*! /#$! %3-%(! (3-($-/1'/%3-! 34! /#$! $..! *%44$1&! .1$'/0,!
4135! /#'/! 34! %/&! &81138-*%-.&6! "#%&! (3-($-/1'/%3-!
*%44$1$-($! %&! $&2$(%'00,! &%.-%4%('-/! 431! &3*%85! '-*!
23/'&&%85!%3-&6!:$'7'/$1!#'&!'!2'1/%(80'10,!#%.#!&3*%85!
%3-! (3-($-/1'/%3-9! 7#$1$'&! /#$! $..! (,/320'&5! (3-/'%-&!
1$0'/%)$0,! 0%//0$! &3*%856! "#$! 1$)$1&$! %&! /#$! ('&$! 7%/#!
23/'&&%85! %3-&6! "#%&! (3-*%/%3-! %&! 5'%-/'%-$*! +,! /#$!
20'&5'!5$5+1'-$9!7#%(#!&/$'*4'&/0,!%-#%+%/&!/#$!$-/1,!34!
&3*%85!%3-&!%-/3!/#$!33(,/$!'-*!21$)$-/&!23/'&&%85!%3-&!
4135! 0$';%-.! 38/! %-/3! /#$! $-)%13-5$-/6! <4! 7$! %-&$1/! '-!
$0$(/13*$! %-/3! '-! $..! '-*! 20'($! '! &$(3-*! $0$(/13*$!
38/&%*$! %/9! 7$! ('-! 5$'&81$! /#$! (3-&/'-/! *%44$1$-($! %-!
(#'1.$! '(13&&! /#$! $..! 20'&5'! 5$5+1'-$6! "#%&! resting 
membrane potential!%&!.$-$1'00,!'+38/!=>!5?9!8&8'00,!
$@21$&&$*! '&! =>!5?! +$('8&$! /#$! %-&%*$! 34! /#$! ($00! %&!
-$.'/%)$0,!(#'1.$*!7%/#!1$&2$(/!/3!/#$!$@/$1%316!
!
!
!
!
!
!
!
!
A%/#%-!B C!&$(3-*&!'4/$1!/#$!+%-*%-.!34!/#$!4%1&/!&2$159!/#$!5$5+1'-$!23/$-/%'0!&#%4/&!/3!
'!23&%/%)$!0$)$09!'+38/!DE>!5?!FG3-.3!$/!'06!BHIJK6!"#%&!(#'-.$!%&!('8&$*!+,!'!&5'00! %-408@!34!
&3*%85! %3-&! %-/3! /#$! $..! FL%.81$! =6EEMK6!M0/#38.#! &2$15! ('-! 48&$!7%/#!5$5+1'-$&! #')%-.! '!
1$&/%-.!23/$-/%'0!34! =>!5?9!/#$,!('--3/!48&$!7%/#!5$5+1'-$&!#')%-.!'!23&%/%)$!1$&/%-.!23/$-/%'09!
&3!-3!531$!&2$15!('-!48&$!/3!/#$!$..6!</!%&!-3/!;-37-!7#$/#$1!/#$!%-(1$'&$*!&3*%85!2$15$'+%0%/,!
%&!*8$!/3!/#$!binding!34!/#$!4%1&/!&2$15!31!/3!/#$!fusion!34!/#$!4%1&/!&2$15!7%/#!/#$!$..!FN380*!'-*!
:/$2#'-3!BHI=9!BHHBO!P(Q8003#!'-*!Q#'5+$1&!BHHEK6!
!
!
The fast block to polyspermy.!
"#$! fast block to poly-spermy! %&! '(#%$)$*! +,!
(#'-.%-.! /#$! $0$(/1%(! 23/$-/%'0! 34! /#$! $..! 20'&5'!
5$5+1'-$6! "#%&!5$5+1'-$! 213)%*$&! '! &$0$(/%)$! +'11%$1!
+$/7$$-!/#$!$..!(,/320'&5!'-*!/#$!38/&%*$!$-)%13-5$-/9!
'-*! /#$! %3-%(! (3-($-/1'/%3-! 34! /#$! $..! *%44$1&! .1$'/0,!
4135! /#'/! 34! %/&! &81138-*%-.&6! "#%&! (3-($-/1'/%3-!
*%44$1$-($! %&! $&2$(%'00,! &%.-%4%('-/! 431! &3*%85! '-*!
23/'&&%85!%3-&6!:$'7'/$1!#'&!'!2'1/%(80'10,!#%.#!&3*%85!
%3-! (3-($-/1'/%3-9! 7#$1$'&! /#$! $..! (,/320'&5! (3-/'%-&!
1$0'/%)$0,! 0%//0$! &3*%856! "#$! 1$)$1&$! %&! /#$! ('&$! 7%/#!
23/'&&%85! %3-&6! "#%&! (3-*%/%3-! %&! 5'%-/'%-$*! +,! /#$!
20'&5'!5$5+1'-$9!7#%(#!&/$'*4'&/0,!%-#%+%/&!/#$!$-/1,!34!
&3*%85!%3-&!%-/3!/#$!33(,/$!'-*!21$)$-/&!23/'&&%85!%3-&!
4135! 0$';%-.! 38/! %-/3! /#$! $-)%13-5$-/6! <4! 7$! %-&$1/! '-!
$0$(/13*$! %-/3! '-! $..! '-*! 20'($! '! &$(3-*! $0$(/13*$!
38/&%*$! %/9! 7$! ('-! 5$'&81$! /#$! (3-&/'-/! *%44$1$-($! %-!
(#'1.$! '(13&&! /#$! $..! 20'&5'! 5$5+1'-$6! "#%&! resting 
membrane potential!%&!.$-$1'00,!'+38/!=>!5?9!8&8'00,!
$@21$&&$*! '&! =>!5?! +$('8&$! /#$! %-&%*$! 34! /#$! ($00! %&!
-$.'/%)$0,!(#'1.$*!7%/#!1$&2$(/!/3!/#$!$@/$1%316!
!
!
!
!
!
!
!
!
A%/#%-!B C!&$(3-*&!'4/$1!/#$!+%-*%-.!34!/#$!4%1&/!&2$159!/#$!5$5+1'-$!23/$-/%'0!&#%4/&!/3!
'!23&%/%)$!0$)$09!'+38/!DE>!5?!FG3-.3!$/!'06!BHIJK6!"#%&!(#'-.$!%&!('8&$*!+,!'!&5'00! %-408@!34!
&3*%85! %3-&! %-/3! /#$! $..! FL%.81$! =6EEMK6!M0/#38.#! &2$15! ('-! 48&$!7%/#!5$5+1'-$&! #')%-.! '!
1$&/%-.!23/$-/%'0!34! =>!5?9!/#$,!('--3/!48&$!7%/#!5$5+1'-$&!#')%-.!'!23&%/%)$!1$&/%-.!23/$-/%'09!
&3!-3!531$!&2$15!('-!48&$!/3!/#$!$..6!</!%&!-3/!;-37-!7#$/#$1!/#$!%-(1$'&$*!&3*%85!2$15$'+%0%/,!
%&!*8$!/3!/#$!binding!34!/#$!4%1&/!&2$15!31!/3!/#$!fusion!34!/#$!4%1&/!&2$15!7%/#!/#$!$..!FN380*!'-*!
:/$2#'-3!BHI=9!BHHBO!P(Q8003#!'-*!Q#'5+$1&!BHHEK6!
Bloqueio  rápido  à  Polispermia	
!
"#$!%&'()*+,-$!(.!/(0%1&!%(,/!+,0!*#$!-#+,2$!%,!)$/*%,2!'(*$,*%+3!4+/!0$&(,/*)+*$0!56!
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Abertura  de  canais  de  Na+  na  membrana  do  ovo  quando  o  primeiro  
espermatozoide  o  fecunda	
Ocorre  em  ouriços,  sapos,  mas  provavelmente  não  em  mamíferos	
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Bloqueio  Lento  à  Polispermia	
Liberação  de  serina  protease	
	
-­‐‑  Clivagem  dos  receptores  de  bindin	
	
-­‐‑  Clivagem  das  ligações  proteicas  entre  
membrana  e  envelope	
A  formação  do  envelope  de  
fertilização  começa  na  região  de  
fusão  do  espermatozoide	
O  Cálcio  é  o  iniciador  da  reação  cortical	
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Uma  onda  de  cálcio  se  
propaga  a  partir  do  ponto  
de  entrada  do  
espermatozoide	
Proximidade  entre  o  RE  e  os  
grânulos  corticais	
Ativação  do  metabolismo  do  ovo	
-­‐‑Retirada  dos  inibidores  dos  RNAm  do  ovo	
-­‐‑Retirada  dos  inibidores  de  divisão  celular	
O  Influxo  de  Ca+  no  citoplasma  do  ovo  também  atua  na  ativação  do  metabolismo  	
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Duas  hipóteses  para  o  mecanismo  de  
liberação  do  Ca+	
Ativação  do  metabolismo  do  ovo	
!
The Activation of Gamete Metabolism!
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phosphatidylinositol 4,5-bisphosphate!=PIPKB!'.!.6&'0!M/!01-!-+N/)-!phospholipase C!=PLCB!0*!
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Atuação  do  Ca+  no  citoplasma	
Ativação  do  metabolismo  do  ovo	
!
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The Activation of Egg Metabolism!
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Early responses!
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Ou  seja...	
Fusão  do  Material  Genético	
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Fusion of genetic material in mammals!
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Após  a  Fecundação:	
-­‐‑Mitocôndrias  originadas  do  espermatozoide  se  degeneram	
-­‐‑Centrossomo  originado  do  ovo  se  degenera	
Fecundação  em  Mamíferos	
Dificuldades:	
-­‐‑  Fecundação  interna	
-­‐‑  População  heterogênea  de  espermatozoides  no  ejaculado	
-­‐‑  Poucos  espermatozoides  chegam  ao  ovócito  1  em  10.000	
-­‐‑  Mais  de  um  mecanismo  para  reação  acrossômica  e  reconhecimento  da  ZP	
Como  os  gametas  chegam  ao  local  da  fecundação?	
Translocação  do  ovócito	
-­‐‑É  essencial  a  presença  das  células  do  cumulus	
-­‐‑Batimentos  ciliares	
-­‐‑Contrações  de  musculatura  lisa	
Como  os  gametas  chegam  ao  local  da  fecundação?	
Translocação  do  espermatozoide	
Os  batimentos  flagelares  parecem  ter  uma  importância  
menor  na  translocação  dos  espermatozoides	
1-­‐‑  As  contrações  uterinas  são  essenciais  para  
o  espermatozóide  chegar  ao  oviducto	
2-­‐‑  A  região  do  oviducto  antes  da  ampola  
diminui  a  velocidade  dos  espermatozoides	
3-­‐‑  A  motilidade  dos  flagelos  é  importante  
quando  os  espermatozoides  alcançam  o  
ovócito	
4-­‐‑  Os  espermatozoides  seguem  um  gradiente  
de  temperatura  e  concentração  de  
quimiotaticos  liberados  pelo  cumulus	
5-­‐‑  O  espermatozoide  sofre  maturação  ao  
longo  do  trajeto	
Capacitação	
O  Primeiro  espermatozoide  a  chegar  na  ampola  será  aquele  que  irá  fertilizar?	
Eventos  importantes  da  capacitação:	
1)  Retirada  de  colesterol  da  membrana  do  
espermatozoide	
2)  Retirada  dos  carboidratos  de  glicoproteínas	
3)  Mudança  no  potencial  de  membrana  	
4)  Fosforilação  de  proteínas	
5)  Aproximação  entre  membrana  acrossômica  e  
membrana  do  espermatozoide	
Capacitação	
Reação  Acrossômica  em  Mamíferos	
Zona  Pelúcida  composta  por  ZP1,  ZP2  e  ZP3	
Até  então,  acreditava-­‐‑se  que:	
-­‐‑Reação  acrossômica  na  Zona  Pelúcida	
-­‐‑Espermatozoide  se  ligando  a  ZP3	
Reação  Acrossômica  em  Mamíferos	
Estudos  mais  recentes  mostram  que:	
-­‐‑Reação  acrossômica  ocorre  no  cumulus	
-­‐‑Espermatozoide  se  liga  a  ZP2	
Fusão  dos  Gametas	
-­‐‑  Contato  lateral  dos  espermatozoides  (região  equatorial)	
-­‐‑  Polimerização  de  
actina  do  ovócito	
Bloqueio  a  Polispermia	
Apenas  bloqueio  lento  à  polispermia	
-­‐‑Reação  cortical	
-­‐‑Clivagem  de  proteínas  ZP2	
Fusão  de  Material  Genético	
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Demora  12  h	
Entrada  do  pronucleo  
masculino,  ondas  de  
Ca+  estimulam  
termino  da  Meiose  II	
Apos  15h  os  
pronucleos  
(duplicados)  se  
aproximam