penfigo inmunopatogenia

7/24/2019 penfigo inmunopatogenia

1/15

!"#$%!" '() *+,-. /012304560 *788

272

1 Residente de Dermatologa, Universidad de Antioquia, Medelln-Colombia Correspondencia: [email protected] Mdica Dermatloga, Doctora en Ciencias Bsicas Biomdicas Inmunologa. Jefe Seccin de Dermatologa, Facultad de Medicina, Universidad de Antioquia, Grupo de Investi-

gacin Dermatologica GRID, Grupo de Estudio en Dermatologa

Recibido: agosto 18 de 2010 Aceptado: septiembre 14 de 2010

Inmunopatogenia del pngo vulgar y el pngo foliceo

scar Jairo Valencia Ocampo1, Margarita M. Velsquez - Lopera2

RESUMEN!" $%&'()* +,")-. / 0" $%&'()* '*"(120* 3*& 0&'0.405-503 -4$*""*3-3 -,6*(&4,&03 405(-5-3

$*. -,6*-&6(2,0.$*3 5(.()(5*3 2*&6.- $.*607&-3 50 "*3 5034*3*4-38 "-3 5034*)"07&-3 9 / :;

!361& -3*2(-5-3 2*& 4*"%2,"-3 50" 2*4$"0("(5-5 ?@ABC D,0 $*.

3, 036.,26,.- 6(0&0& "- 2-$-2(5-5 50 $.030&6-. $%$6(5*3 -&6()%&(2*3 50 "-3 5034*)"07&-3;

!& "*3 (&5(+(5,*3 -'026-5*3 30 =-& 5032.(6* "- $.030&2(- 50 "(&'*2(6*3 E / F -,6*..0-26(+*3 /-"60.-2(*&03 0& "- .0),"-2(G& 50" 3(3604- (&4,&0 2*& 5030D,("(>.(* 50 "-3 .03$,036-3 E=9HE=I;

J* 30 2*&*20& 2*& $.02(3(G& "*3 402-&(34*3 50 5-K* $0.* "- (&+036()-2(G& -26,-" (&5(2-

D,0 "*3 -&6(2,0.$*3 6(0&0& ,& $-$0" $-6*)%&(2*8 (&(2(-& 5('0.0&603 2-32-5-3 50 30K-"(L-2(G&

D,0 $.*+*2-& "- -2-&6G"(3(3 / -$*$6*3(3 50 "*3 D,0.-6(&*2(6*3; !" 2*&*2(4(0&6* 50 "-

(&4,&*$-6*)0&(- 50 "-3 0&'0.405-503 -4$*""*3-3 -,6*(&4,&03 =- $0.4(6(5* 0" 503-..*""* /

"- $,036- 0& $.126(2- 50 &,0+-3 -"60.&-6(+-3 60.-$%,6(2-3;

PALABRAS CLAVEB2-&6G"(3(3M N(3(*$-6*"*)7-M O&4,&*"*)7-M P%&'()*

SUMMARY!449:(1;2 1041"(360.(&) 5(30-303 405(-605

>/ -&6(>*5(03 -)-(&36 5034*3*4-" $.*60(&3; E=0/ -.0 36.*&)"/ -33*2(-605 Q(6= 4-("(6/ 2*4$"0R -""0"03 Q(6= 6=0 ->("(6/ 6* $.030&6 -&6()0&(2 $0$6(503 *' 5034*)"0(&3;

O& 6=0 -''02605 (&5(+(5,-"3 6=0 $.030&20 *' -,6*S.0-26(+0 E -&5 F "/4$=*2/6038 -&5 -"60.-6(*&3

!;2603; '()B *+ ,-.C *D* E *FGH /012304560 I :(?304560 *788


2/15

!"#$%!" '() *+,-. /012304560 *788

273

(& 6=0 (44,&0 3/3604 .0),"-6(*& Q(6= (4>-"-&20 *'

6=0 E=9HE=I .03$*&303 =-+0 >00& 5032.(>05; T-4-)0

402=-&(343 -.0 &*6 /06 $.02(30"/ U&*Q& >,6 2,..0&6

(&+036()-6(*& (&5(2-603 6=-6 -&6(>*5(03 $"-/ -&

(4$*.6-&6 $-6=*)0&(2 .*"0V 6=0/ 36-.6 5(''0.0&6 3()&-"(&)2-32-503 6=-6 "0-5 6* -2-&6=*"/3(3 -&5 -$*$6*3(3 *'

U0.-6(&*2/603; F0660. U&*Q"05)0 *' 6=0 $-6=*)0&03(3

*' -,6*(44,&0 >"(360.(&) 5(30-303 =-3 >00& 6=0 >-3(3

'*. 6=0 50+0"*$40&6 -&5 (4$"040&6-6(*& *' &0Q

6=0.-$0,6(2 -$$.*-2=03;

KEY WORDSB2-&6=*"/3(3M O44,&*"*)/M P04$=(),3M P=/3(*S

$-6=*"*)/

INTRODUCCIN!" 6%.4(&*$%&'()*50'(&0 ,& ).,$* 50 0&'0.405-503

-,6*(&4,&03 50 "- $(0" 2-.-260.(L-5-3 $*. "- $.030&2(-

50 -4$*""-3 50>(5-3 - -2-&6G"(3(3 ?30$-.-2(G&

(&60.20","-.C 0 (&4,&*)"*>,"(&- W ?O)WC 5(.()(5- -

"- 3,$0.'(2(0 50 "*3 D,0.-6(&*2(6*3 ?9C; X"13(2-40&60

30 "* =- 5(+(5(5* 0& 2,-6.* 6($*3V $%&'()* +,")-.

?PYC8 $%&'()* '*"(120* ?PNC8 $%&'()* $-.-&0*$"13(2* /

$%&'()* O)B ?IC; Z0 $.030&6- ,&- .0+(3(G& 0&'*2-5- 0&

"- (&4,&*$-6*)0&(- 50" PY / 0" PN;

EPIDEMIOLOGAA- (&2(50&2(- )"*>-" 50 PY 03 50 [89 - [8\ $*.

9[[;[[[H-K*8 2*& +-.(-2(*&03 D,0 50$0&50& 50"

1.0- )0*).1'(2- / 50 "- 06&(-M -'026- - =*4>.03 /

4,(- / P0._ "*3 2,-"03 6(0&0& 2-.-260.736(2-3

2"7&(2-38 =(36*"G)(2-3 0 (&4,&*"G)(2-3 3(4("-.038

$0.* +-.7-& 0& 3, 0$(504(*"*)7-; !" PN! 50" F.-3("

?N*)* 30"+-)04C -'026- - &(K*3 / (6-& 2(0.6-3 4*32-3 &0).-3 ?Z(4,"(,4

$.,(&*3,4CM 413 50" \[` 50 "-3 $0.3*&-3 3-&-3

D,0 .03(50& 0& "-3 L*&-3 5*&50 30 $.030&6- 0" PN!

6(0&0& -,6*-&6(2,0.$*3 O)W -&6(5034*)"07&- 9 / 30

=- 504*36.-5* D,0 "- -$-.(2(G& 50 "- 0&'0.405-5

0361 $.0205(5- $*. ,&- .03$,036- 3*360&(5- 50 5(2=*

-&6(2,0.$*M 30 =-& (&'*.4-5* $*2*3 2-3*3 50 '*)*

30"+-)04 2*& "03(*&03 0& "-3 4,2*3-3 .0"-2(*&-5-32*& "- -$-.(2(G& 50 -,6*-&6(2,0.$*3 -&6(5034*)"07&-

:8 503-..*""-&5* ,&- '*.4- 0&5%4(2- 50 PY ?:C; !"

PN! 50 E_&0L -'026- 413 - 4,.0 0& P0._ 30 =- 504*36.-5* "- $.030&2(- 50

-,6*-&6(2,0.$*3 5(.()(5*3 2*&6.- "- 5034*)"07&-S9

"- 2,-" 30 .0"-2(*&- 2*& "- *2,$-2(G& 0& "--).(2,"6,.- ?\C; !" '*2* 2*"*4>(-&* 30 0&2,0&6.- 0&

0" 4,&(2($(* 50 !" F-).08 B&6(*D,(-8 -'026- -" I8:`

50 3, $*>"-2(G& .,.-"8 0" a\` 50 "*3 2-3*3 *2,..0& 0&

=*4>.03 4-/*.03 50 ][ -K*38 4(&0.*3 / -).(2,"6*.03

/ 03 4-/*. "- $.0+-"0&2(- 0& "*3 (&57)0&-3 50 "-

.0)(G& ?^C;!& X*"*4>(- 30 5032*&*20 "- (&2(50&2(-

50 PY / PN!;

MANIFESTACIONES CLNICAS!" PY 30 2-.-260.(L- 2"7&(2-40&60 $*. -.5*. $.0+(*

- "- -$-.(2(G& 50 -4$*""-3 '"12(5-3 0& "- $(0" / "-34,2*3-38 "-3 D,0 30 $,050& -).,$-. / 50&,5-.

50(0.6-3 $*. 2*36.-3 D,0 -" 2(2-6.(L-.

50-3-" / "- 50" PN8 "- -2*&6G"(3(3

3,>2G.&0-; A- (&4,&*'",*.0320&2(- 50 -4>*3 6($*3 50

$%&'()* 30 2-.-260.(L- $*. "- $.030&2(- 50 50$G3(6*3 50

O)W8 X: * -4>*3 0& "- 3,$0.'(2(0 50 "*3 D,0.-6(&*2(6*3

0& ,& $-6.G& .06(2,"-.; !36*3 -,6*-&6(2,0.$*3

',0.*& 5032,>(0.6*3 (&(2(-"40&60 0& 0" 3,0.* 50

"*3 $-2(0&603 $*. 405(* 50 (&4,&*'",*.0320&2(-

(&5(.026- / ",0)* *>30.+-5*3 0& "- $(0" $0.("03(*&-"

$*. (&4,&*'",*.0320&2(- 5(.026- ?^8cC;


3/15

!"#$%!" '() *+,-. /012304560 *788

274

ADHESIN EPIDRMICAA- .03(360&2(- 50 "- $(0" 2*&6.- "-3 (&'",0&2(-3

-4>(0&6-"03 / "- (&60).(5-5 .0D,0.(5- $-.- "-

$.*6022(G& 2*&6.- &*R-3 4021&(2-38 D,74(2-3

* 4(2.*>(-&-3 30 50>0& - 5*3 036.,26,.-3 D,0

4-&6(0&0& "- -5=03(G& 0$(5%.4(2-M "- $.(40.-

3*& "*3 5034*3*4-3 D,0 30 0&2-.)-& 50 "- ,&(G&

2%","- - 2%","- / "- 30),&5-8 "*3 =04(5034*3*4-3

D,0 0R$"(2-& "- ,&(G& 50 "-3 2%","-3 - "- 404>.-&-

>-3-"; T('(0.0& 50350 "*3 $,&6*3 50 +(36- 036.,26,.-" /

4*"02,"-. ?IC;

A*3 5034*3*4-3 3*& 0" >"-&2* 50 "*3 -,6*-&6(2,0.$*3

0& 0" PY / 0" PN; !361& 2*4$,036*3 $*. 6.03 '-4("(-3

50 $.*607&-3V 9C "-3 $"-D,(&-3 D,0 2*&36(6,/0& "-

$"-2- (&60.20","-.8 (&2",/0& "- 5034*$"-D,(&-8 "-

$0.($"-D,(&- / "- 0&+*$"-D,(&- / 30 ,&0& - "*3'("-40&6*3 50 D,0.-6(&-M IC "- '-4("(- 50 "-3 $.*607&-3

50" -.4-5(""*V (&2",/0 "- $"-2*'("(&- / "- $"-2*)"*>(&-

D,0 -26_-& 2*4* $,0&603 0&6.0 "- 5034*$"-D,(&-

(&6.-20","-.M :C "- '-4("(- 50 "-3 2-5=0.(&-38 '*.4-5-

$*. 5034*2*"(&-3 / 5034*)"07&-3 D,0 30 ,&0& - 3,3

=*4G"*)-3 0& "-3 2%","-3 +02(&-3 ?'(),.- &;b :C ?dS9IC;

TEORA DE LA COMPENSACIN DE LASDESMOGLENASA-3 5('0.0&2(-3 2"7&(2-3 0 =(36*$-6*"G)(2-3 0&6.0 PY / PN

30 =-& 0R$"(2-5* $*. "- 60*.7- 50 "- 2*4$0&3-2(G& 50

"-3 5034*)"07&-3; T0 "*3 2,-6.* 6($*3 50 5034*)"07&-3

0R(360&6038 0" 9 / 0" : 30 0&2,0&6.-& 5(36.(>,(5*3 0&

5('0.0&603 $.*$*.2(*&03 0& "- $(0" / "-3 4,2*3-3; !&

0" PN 30 $.*5,20& -,6*-&6(2,0.$*3 5(.()(5*3 2*&6.-

"- 5034*)"07&- 9 ?T3)S9C8 0" '0&G40&* 50 -2-&6G"(3(3

.0$.030&6-5* $*. -4$*""-3 3*"* 30 *>30.+- 0& "-

L*&- 3,>2G.&0- 50>(5* - D,0 "- 2-&6(5-5 50 T3)S: &*

-"2-&L- - 2*4$0&3-. 0" >"*D,0* ',&2(*&-" *2-3(*&-5*

$*. "*3 -,6*-&6(2,0.$*3M $*. *6.* "-5*8 "- 2-&6(5-5 50

T3)S: 0& "-3 4,2*3-3 03 3,'(2(0&60 $-.- 4-&60&0. "-

-5=03(G&8 / 036- 03 "- .-LG& $*. "- 2,-" &* 30 $.030&6-&

"03(*&03 -4$*""*3-3 0& "-3 4,2*3-3 50 "*3 $-2(0&6032*& PN; !& "- 4-/*.7- 50 "*3 $-2(0&603 2*& PY 30

0&2,0&6.-& -,6*-&6(2,0.$*3 2*&6.- T3)S9 / T3)S:8

-4>-3 $.*607&-3 3*& >"*D,0-5-3 ',&2(*&-"40&60 /

-$-.020 "- -2-&6G"(3(3 2*& -4$*""-3 6-&6* 0& "- $(0"

2*4* 0& "-3 4,2*3-3 ?'(),.- &;b ]C ?9:C;

Figura n. 2. Pngo foliceo. La piel aparece cubierta por escamas ycostras sobre una base eritematosa, algunas zonas denudadas. Las ampollas ci-das son poco aparentes. Foto Seccin de Dermatologa, Universidad de Antioquia

Figura n. 1. Pngo vulgar. Extensa denudacin de la piel y las mucosas.

Foto Seccin de Dermatologa, Universidad de Antioquia


4/15

!"#$%!" '() *+,-. /012304560 *788

275

Placa Centro Placa Autoantgenos

AUTOANTGENOS!" PY 03 ,&- 0&'0.405-5 405(-5- $*. -,6*-&6(2,0.$*3

5(.()(5*3 2*&6.- "- 5034*)"07&- ?T3)C : / 2*&6.-

*6.*3 -,6*-&67)0&*3 2*4* "- T3)S9 0& 0" \[`Sc\`

50 "*3 2-3*3; !& 0" PN "*3 -,6*-&6(2,0.$*3 0361&

5(.()(5*3 2*&6.- "- T3)S9 ?9]C;Z0 =-& 5032.(6* *6.*3

-,6*-&67)0&*3 0& 0" PY 2*4* "- 5034*$"-D,(&-8 0"

.020$6*. 50 -206("2*"(&- -"'- a / *6.-3 4*"%2,"-3 50

"- 3,$0.'(2(0 50" D,0.-6(&*2(6* 2*4* "- $0&'-R(&-

/ "- -&0R(&- :9 D,0 $*5.7-& ',&2(*&-. 2*4*

.020$6*.03 $-.- -206("2*"(&-; A*3 -,6*-&6(2,0.$*3

D,0 30 $.*5,20& 2*&6.- -,6*-&67)0&*3 5(36(&6*3 - "-3

5034*)"07&-3 6-4>(%& $-.6(2($-& 0& "- $-6*)0&(- 50

"- 0&'0.405-5 ?9\S9aC;

!36,5(*3 3*>.0 0" PN! 2*"*4>(-&* =-& 504*36.-5*8

-50413 50 "*3 -&6(2,0.$*3 2*&6.- T3)S98

-,6*-&6(2,0.$*3 5(.()(5*3 2*&6.- +-.(*3 5*4(&(*3

50 $.*607&-3 50 "- '-4("(- $0.($"-D,(&- ?-&67)0&*3

50" $%&'()* $-.-&0*$"13(2*C8 2*& $03*3 4*"02,"-.03

50 I\[ UT8 I9[ UT / 9a[ UT8 "-3 2,-"03 $*5.7-&

2*..03$*&50. - 5034*$"-D,(&- 98 0&+*$"-D,(&- /

$0.($"-D,(&-8 -50413 50 ,&- $.*607&- 50 I:[ UTM

(&(2(-"40&60 30 $0&3G D,0 036- _"6(4- 2*..03$*&57-

- "- FPI:[8 5032.(6- 2*4* 0" -,6*-&67)0&* 0& 0"

$0&'()*(50 -4$*""*3*8 $0.* -26,-"40&60 30 2.00

D,0 2*..03$*&50 - *6.- $.*607&- 50 (),-" $03*

4*"02,"-. * - ,& 2*4$"0(6-& 0& 0"

'*2* 0&5%4(2* 50 PN! 50 "- 2*4,&(5-5 50 P,0>"*

"(>.0 0& P0._ 30 =-& 504*36.-5* -&6(2,0.$*35(.()(5*3 2*&6.- "- T3)S98 4(0&6.-3 D,0 0&6.0 9c` /

II` 50 "*3 (&5(+(5,*3 3-&*3 50" '*2* 50 E_&0L =-&

504*36.-5* -&6(2,0.$*3 2*&6.- "- T3)S9 ?]C; !& 0" :^`

50 "*3 (&5(+(5,*3 3-&*3 D,0 =->(6-& "- L*&- 5*&50

=-/ PN! 0& F.-3(" 30 (&'*.4-.*& -&6(2,0.$*3 2*&6.-

"- T3)S:M "* 4(34* 30 =-""G 0& 0" ]:` 50 "*3 (&5(+(5,*3

Figura n. 3. Estructura de los desmosomasLas molculas trasmembrana, desmoglenas y desmocolinas, se unen con queratinas del citoesqueleto mediante la interaccincon componentes intracelulares de la placa del desmosoma como desmoplaquina, placo lina, envoplaquina y periplaquina.Modicado de Herti M, Erning R, Veldam C. T cell control in autoimmune skin disorders. J Clin Invest 2006; 116: 115966


5/15

!"#$%!" '() *+,-. /012304560 *788

276

2*& '*)* 30"+-)04M- $03-. 50 "- $.030&2(- 50 036*3

-&6(2,0.$*3 3*"* 0& 4,/ $*2*3 2-3*3 3*& $-6*)%&(2*3

/ 30 $.030&6- ,& +(.--& .0-22(G& 2.,L-5- 2*& "- T3)S]M 30 $"-&60-

D,0 0361& (&+*",2.-5*3 0& "- $-6*)0&(- 50" PY $0.*

30 .0D,(0.0& 036,5(*3 -5(2(*&-"03 $-.- 50'(&(. 0" $-$0"

50 0360 -&67)0&* 0& "- 0&'0.405-5 =,4-&- ?6->"- &;b9C ?IIC;

Figura n. 4. Teora de compensacin de las desmoglenasLas Dsg-1 y Dsg-3 se encuentran distribuidas en diferentes proporciones en la piel y las mucosas. En PF la IgG es contra Dsg-1, el fenmenode acantlisis es subcrneo debido a que la cantidad de Dsg-3 no alcanza a compensar el bloqueo funcional de la Dsg-1; en las mucosas lacantidad de Dsg-3 es adecuada y suciente para mantener la adhesin. En el PV la mayora de los pacientes presentan IgG contra Dsg-1 y Dsg-3, el bloqueo funcional de ambas protenas induce acantlisis en piel y mucosas. Modicado de Stanley JR. Pemphigus. In: Wolff K, GoldsmithLA, Gilchrest BA, Paller AS, Leffell DJ. Fitzpatricks Dermatology in General Medicine. 7th ed. New York. Mc Graw Hill Medical, 2008. p 459-467

MucosaPiel

MucosaPiel

Anticuerpos anti-Dsg1

Solo anticuerpos anti-Dsg3

Anticuerpos anti-Dsg3 y anti-Dsg1

13 3

1

1

3 3

1

13 3

1


6/15

!"#$%!" '() *+,-. /012304560 *788

277

Tabla n. 1. Autoantgenos en pngo vulgar y pngo foliceo

Tipo de pngo Antgenos principales Antgenos accesorios

Pngo vulgar Desmoglena-3Desmoglena-1

Desmoplaquina

Receptor de acetilcolina alfa-9Penfaxina y anexina

Desmoglena-4?

Pngo foliceo espordicoDesmoglena-1

Pngo foliceo endmico

(Colombia) Desmoglena-1

Desmoplaquina-1

Envoplaquina

PeriplaquinaProtena de 230 kD


(Brasil) Desmoglena-1

Desmoglena-3

Desmocolina-1, -3


(Per, Tnez)Desmoglena-1

FACTORES GENTICOS!" PY '*.4- $-.60 50 ,& ).,$* 50 0&'0.405-503

-,6*(&4,&03 ',0.6040&60 -3*2(-5-3 -" 2*4$"0("(5-5 ?Xe@C; !" PY 30 -3*2(- 2*& "*3-"0"*3 @ABSTfF9g[][I 0& $*>"-2(*&03 ("(5-5 -

503-..*""-. PN! 0& F.-3("8 4(0&6.-3 D,0 "- $.030&2(-

50 ,& -"0"* @ABSTh8 0" ThQI8 $-.020 036-. -3*2(-5-

2*& ,& .(03)* 40&*. 50 503-..*""-. "- 0&'0.405-5 0&

036- $*>"-2(G& ?IdS:[C; A*3 -"0"*3 50" @ABSTfF9g9][I8

9][^ / [][] 30 0&2,0&6.-& 2*& 4-/*. '.02,0&2(- 0&

(&5(+(5,*3 3-&*3 2*& -&6(2,0.$*3 -&6(ST3)S9 0& "-

$*>"-2(G& 2*& PN 0&5%4(2* 0& E0.0&-8 F.-3(" ?]C;

FACTORES DESENCADENANTESP0.3*&-3 )0&%6(2-40&60 3,320$6(>"03 6(0&0& 4-/*.

.(03)* 50 503-..*""-. "- 0&'0.405-5 -" 036-. 0R$,036-3

- '1.4-2*3 0& 2,/- 036.,26,.- =-/ ,& ).,$* 6(*"8 2*4*

"- TS$0&(2("-4(&- / 0" 2-$6*$.("M * >(0& 2*& ,& ).,$*

-4(5- 2*4* 0&-"-$.("8 $0&(2("(&-8 20'-"*3$*.(&-3 /

.('-4$(2(&- ?:[C; i&* 50 "*3 $*2*3 036,5(*3 50 ).,$*3

).-&503 ',0 0" 50 @0/4-&& / 2*"->*.-5*.03 D,(0&03

3(),(0.*& - 9\[;[[[ (3.-0"703 $*. \ -K*38 50 "*3 2,-"03:^: $.030&6-.*& PYM 30 =-""G ,&- -3*2(-2(G& ?jfV IC

2*& 0" ,3* 50 $0&(2("(&- 0& "*3 30(3 40303 $.0+(*3 -"

5(-)&G36(2* 50 036- 0&'0.405-5 -4$*""*3- ?:9C;

A- 036.,26,.- 4*"02,"-. 50 -"),&*3 (&).05(0&603 50 "-

5(06- 03 3(4("-. - "- 50 "*3 405(2-40&6*3 (&5,26*.03


7/15

!"#$%!" '() *+,-. /012304560 *788

278

50 $%&'()*; !R(360& (&'*.403 50 2-3*3 D,0 4,036.-& "-

.0"-2(G& 50 -"(40&6*3 2*& 0" '0&G40&* 50 -2-&6G"(3(3

>(*D,74(2-8 2*& 5-K* 5(.026* 50 "*3 5034*3*4-3

* 03674,"* 50 "- -,6*(&4,&(5-5 405(-5- $*.

-&6(2,0.$*3; !&6.0 "*3 -"(40&6*3 .0"-2(*&-5*3 2*&-2-&6G"(3(3 0361& "*3 D,0 2*&6(0&0& ,& ).,$* 6(*"

2*4* 20>*""-38 -*""06-3M 0" '0&*"8

$.030&60 0& "-3 6(&6,.-3 50 >0&(*$3(-3 50 $-2(0&603 2*& PY -26(+* 30

0&2*&6.G OAS] ?,&- 2(6*2(&- E=ICM 0& 2,-&6* - OASI 0

OJNS?2(6*2(&-3 E=9C 30 0&2*&6.-.*& 0& 0" c9` / ]d`

50 "*3 2-3*38 .03$026(+-40&60 ?][C;

A- 30),&5- 5('0.0&2(- 0&6.0 "-3 $0.3*&-3 3-&-3 2*& @AB

50 .(03)* $-.- $%&'()* / "*3 $-2(0&603 D,0 $.030&6-&

"- 0&'0.405-5 3*& "*3 "(&'*2(6*3 E .0),"-5*.03; !&

"*3 (&5(+(5,*3 3-&*38 "- 4-/*.7- 50 "*3 "(&'*2(6*3 E

-,6*..0-26(+*3 2*&6.- T3)S: 3*& .0),"-5*.03 6($*

9 ?E.9C8 D,0 0(5*.- 50 "-

$.*"('0.-2(G& 50 2"*&03 50 AE $-6*)%&(2*38 405(-5-

$*. OAS9[ / EWNS ?]9C; B 5('0.0&2(- 50 "* -&60.(*.80& "*3 $-2(0&603 2*& PY $.05*4(&-& "*3 "(&'*2(6*3

E=I -,6*..0-26(+*3 / 6(0&0& ,&- 40&*. 2-&6(5-5 50

"(&'*2(6*3 E.9M 30 $.*$*&0 D,0 036* $,050 '-+*.020. "-

$%.5(5- 50 6*"0.-&2(- =-2(- "- T3)S: ?]I8]:C;

j6.-3 0+(50&2(-3 (&2",/0& "- $.030&2(- 50 (&'("6.-5*

20","-. 2*& (&2.040&6* 0& 0" &_40.* 50 AE =-36- 0"

^[`8 2*& ,&- .0"-2(G& XT]HXTd 50 IV98 / "- .0"-2(G&

5(.026- 0&6.0 "- -26(+(5-5 50 "- 0&'0.405-5 / "*3 &(+0"03

50 XTI\ 0& 0" 3,0.* / 0& 0" "7D,(5* 50 "-3 -4$*""-3 ?:cC;

INMUNIDAD HUMORAL!& "- 3-&).0 $0.('%.(2- 50 $-2(0&603 2*& PY 30 =-&

506026-5* "(&'*2(6*3 F -,6*..0-26(+*3 2*&6.- T3)S

: / 30 =- *>30.+-5* D,0 "*3 -&6(2,0.$*3 50 6($*

O)W D,0 $.*5,20& "*3 "(&'*2(6*3 3*& .03,"6-5* 50 "-

30"022(G& -&6()%&(2-8 /- D,0 6(0&0& ,3* .036.(&)(5*

50 "*3 30)40&6*3 )%&(2*3 Y@ / l@ ?D,0 2*5('(2-&

$-.- "- .0)(G& +-.(->"0 50 "- 2-50&- $03-5-C /

4_"6($"03 4,6-2(*&03 2,/- &-6,.-"0L- / 5(36.(>,2(G&

&* 3*& -"0-6*.(-3 ?]]C; B50413 30 2*4$.*>G D,0 03

(&5(3$0&3->"0 "- 2**$0.-2(G& 0&6.0 "*3 AF / "*3 AE

-,6*..0-26(+*3 $-.- D,0 30 $.*5,L2-& -&6(2,0.$*3-&6(ST3)S:; !+(50&2(-3 (& +(6.* 504*36.-.*& "-

$.030&2(- 50 -,6*-&6(2,0.$*3 -" 036(4,"-. 2*& T3)S

: ,&- 40L2"- 50 "(&'*2(6*3 50 3-&).0 $0.('%.(2-

50 $-2(0&603 2*& PY8 4(0&6.-3 D,0 -" 0"(4(&-. "*3

"(&'*2(6*3 E XT]8 "*3 "(&'*2(6*3 F -,6*..0-26(+*3 &*

$.*5,


8/15

!"#$%!" '() *+,-. /012304560 *788

279

"92(;:23A9061(/

A*3 -,6*-&6(2,0.$*3 0& PY 3*& 50 6($* O)W /

$*"(2"*&-"038 03 502(.8 .02*&*20& 5('0.0&603 0$76*$03

-&6()%&(2*3 $*.D,0 $.*+(0&0& 50 5('0.0&603 2"*&03

50 "(&'*2(6*3 F; B,& 2,-&5* 30 $.*5,20& "*3 2,-6.*3,>6($*3 50 O)W8 0" 3,>6($* O)W] 03 413 '.02,0&60 0&

$-2(0&603 2*& 0&'0.405-5 50 -$-.(2(G& .02(0&60 / 30

0&2,0&6.- 6-4>(%& 0& -D,0""*3 2,/- 0&'0.405-5 =-

3(5* 50 "-.)- 5,.-2(G&M 0& 036*3 _"6(4*3 30 0&2,0&6.-

-50413 O)W9; A*3 -&6(2,0.$*3 0361& 5(.()(5*3 2*&6.-

"- $*.2(G& -4(&*S60.4(&-" 50 "- T3)S: ?:aC;

A*3 =((5* -" $-3* 6.-&3$"-20&6-.(* 50 -,6*-&6(2,0.$*3M

"- 0&'0.405-5 503-$-.020 ",0)* 50" 2-6->*"(34* 50

"*3 4(34*3 ?]^C; !" PN &0*&-6-" 03 ,&- 0&'0.405-5

.-.-M -" .03$026* 30 $.*$*&0 D,0 "- 5(36.(>,2(G&50 "*3 -&67)0&*3 0& "- $(0" 50" &0*&-6* 03 5('0.0&60

2*& $.030&2(- 50 T3)S9 / T3)S: 0& 6*5* 0" 03$03*.

50 "- 0$(50.4(3 / D,0 3*"* 30 $.*5,2(.7-& "03(*&03

2"7&(2-3 0& "*3 2-3*3 2*& -&6(2,0.$*3 5(.()(5*3 2*&6.-

-4>*3 -&67)0&*3; J* 30 =-& (&'*.4-5* 2-3*3 50 PN!

&0*&-6-" / 3*"* 0R(360& 6.03 2-3*3 50 PN &0*&-6-" &*

0&5%4(2* ?]cC; !& 0" 4*50"* 4,.(&*8 "- 6.-&3'0.0&2(-

-5*$6(+- 50 3,0.* 50 $-2(0&603 2*& PY - .-6*&03

&0*&-6*3 FBAFH2 (&5,20 "03(*&03 2"7&(2-3 / 2-4>(*3

=(36*"G)(2*3 67$(2*3 50 PY 2*& -2-&6G"(3(3 / '*.4-2(G&

50 -4$*""-3 3,$.->-3-"03 ?]dC; A- O)W 50 $-2(0&603

2*& $%&'()* $,050 (&5,2(. $%.5(5- 50 "- -5=03(G&20","-. 0& 2,"6(+*3 50 D,0.-6(&*2(6*3 0& -,30&2(- 50

2*4$"040&6* / 50 2%","-3 (&'"-4-6*.(-3 ?]a8\[C;

MECANISMOS DE ACANTLISISZ0 =-& 0+-",-5* 2,-6.* 402-&(34*3 0& "- (&5,22(G&

50 -2-&6G"(3(3V

J0,6.-"(L-2(G&V "*3 .-6*&03 50'(2(0&603 0& T3)S:

60&7-& ,& '0&*6($* 3(4("-. -" 50 "*3 $-2(0&603 2*&

PY ?\9C; i&- 50 "-3 =($G603(3 40"-,2(G& 50 "- $"-2*)"*>(&-; T(36(&6-3 +7-3

/ 4*"%2,"-3 50 30K-"(L-2(G& =-& 3(5* (4$"(2-5-3 0&

036*3 '0&G40&*3 ?\dC;

A- -"60.-2(G& 0& 0" 0&3-4>"-"-


9/15

!"#$%!" '() *+,-. /012304560 *788

280

X ?PkX8 $*. "- 3()"- 0& (&)"%3 50 $.*60(&SU(&-30 XC

D,0 - 3, +0L -26(+- -" 3(3604- $.*60*"76(2* -26(+-5*.

50 $"-34(&G)0&* 6($* ,.*2(&-3- ?,PBCH.020$6*. 50

,PBH$"-34(&-; !" TBW -26(+- 5(.026-40&60 - "- PkX

4(0&6.-3 D,0 0" OP: "* =-20 405(-&60 ,& (&2.040&6*0& "- 2*&20&6.-2(G& (&6.-20","-. 50 2-"2(* D,0 6-4>(%&

""0+- - "- -26(+-2(G& 50 "- 2-"4*5,"(&-; Z0 2.00 D,0

036- +7- -"60.- 0" .02(2"-(. "- -2-&6G"(3(3 0& 0" 4*50"*

4,.(&* 50 6.-&3'0.0&2(- $-3(+- ?\dC;

B" ,&(.30 0" -,6*-&6(2,0.$* - "- T3)S:8 0" 2*4$"0(&- 03 0&5*2(6-5*8 (&60.&-"(L-5*

/ 6.-&3$*.6-5* -" 3(3604- "(3*3*4-"; Z0 2.00 D,0 "-

T3)S: 30 50).-5- 50>(5* - D,0 30 =- 2*4$.*>-5*

D,0 *2,..0 ,&- 5(34(&,2(G& 0& 3, 2-&6(5-5 6-&6*

0& "*3 5034*3*4-3 2*4* 0& *6.*3 2*4$-.6(40&6*320","-.038 ",0)* 50" 6.-&3$*.60 50" 2*4$"0-260.(- D,0 03$027'(2-40&60 -,40&6- "- -26(+(5-5

50 "-3 WEPS-3-3 50 "- '-4("(- f=*; X*& 036- 6*R(&- 30

"*).G 0+(6-. "- .05(36.(>,2(G& 50 "- T3)S: / "- .06.-22(G&

50 '("-40&6*3 (&60.405(*3 50 D,0.-6(&-8 0+(6-&5*

-37 "- 30$-.-2(G& (&60.20","-. ?^9C; A- $:deBPk

6(0&0 *6.-3 ',&2(*&03 0& "- -2-&6G"(3(3V ,&* 50 3,3

3,36.-6*3 03 "- $.*607&-S2(&-3- I -26(+-5- $*. eBPk

?$*. "- 3()"- 0& (&)"%3 50 4(6*)0&S-26(+-605 $.*60(&

U(&-303C ?eBPkBPIC D,0 - 3, +0L '*3'*.("- "- $.*607&-

50 2=*D,0 6%.4(2* Ic ?@ZPIcC ?$*. "- 3()"- 0& (&)"%3

50 =0-6S3=*2U $.*60(&C; B" 30. '*3'*.("-5-8 "- @ZPIc

$,050 .0*.)-&(L-. 5(.026-40&60 0" 2(6*03D,0"06* ?^IC;

Z0 =- $.*$,036* D,0 "- -26(+-2(G& 50 "- $:deBPk

$*5.7- *2,..(. $*. -&6(2,0.$*3 03$027'(2*3 2*&6.- T3)S

: * >(0& $*. 2-4>(*3 0& 0" 2(6*03D,0"06* 2-,3-5*3 $*.-,6*-&6(2,0.$*3 2*&6.- *6.*3 -&67)0&*3 &* 03$027'(2*3

?&* 5034*)"07&-3C; !36*3 -&6(2,0.$*3 (&03$027'(2*3

-26(+-& *6.-3 2(&-3-38 2*4* Z.2 / 2(&-3- 50" .020$6*.

50" '-26*. 50 2.02(4(0&6* 0$(5%.4(2* ?!WNfk8 $*. "-

3()"- 0& (&)"%3 50 0$(50.4(2 ).*Q6= '-26*. .020$6*.

kC "-3 2,-"03 $-.6(2($-& 0& "- 503$*"(40.(L-2(G&

50 "- -26(&- / 0& "- -).0)-2(G& 50 "*3 '("-40&6*3

(&60.405(*3 50 D,0.-6(&- ?^:C;

A- $%.5(5- 50 -5=03(G& (&60.20","-. 6-4>(%& 30

-'026- $*. "- .05(36.(>,2(G& 50 "- $"-2*)"*>(&- ?PWC;

!36- $.*607&- 6(0&0 +-.(-3 ',&2(*&03 0& "- 2%","-V

-&2"- "- T3) -" 2(6*03D,0"06*8 3(.+0 2*4* 4*"%2,"-

50 30K-"(L-2(G& / .0),"- "- 6.-&32.($2(G& 50 2(0.6*3

)0&03 2*4* 2Se/2 ?^]C; T03$,%3 50 "- ,&(G& 50 "*3

-,6*-&6(2,0.$*3 - "- 3,$0.'(2(0 50 "*3 D,0.-6(&*2(6*3

30 =- .0$*.6-5* ,&- 5(34(&,2(G& 50 "- 2-&6(5-5

50 $"-2*)"*>(&- &,2"0-.; !36* $.*>->"040&60 30

.0"-2(*&0 2*& ,& 6.-&3$*.60 50'(2(0&60 -" &_2"0*

D,0 "0 (4$(50 ""0+-. - 2->* 3, ',&2(G& .0),"-5*.-;

A- $"-2*)"*>(&- &,2"0-. 6(0&0 03$02(-" (4$*.6-&2(-

0& "- 3,$.03(G& 50 "- 6.-&32.($2(G& 50 2Se/28 '-26*.

50 6.-&32.($2(G& .0"-2(*&-5* 2*& "*3 $.*203*3 50

$.*"('0.-2(G& 20","-. / -$*$6*3(3; B" 5(34(&,(. "- PW

0& 0" &_2"0*8 *2,..0 ,& (&2.040&6* 0& "- 2-&6(5-550 2Se/2 / 036* '-+*.020 "- $.*"('0.-2(G& 2*&6(&,-

50 "*3 D,0.-6(&*2(6*3 / 3, $0.4-&0&2(- 0& ,& 036-5*

(&4-5,.*8 0+0&6*3 D,0 30 =-& -3*2(-5* 2*& "-

5(34(&,2(G& 50 "- ',0.L- -5=03(+-; A- (4$*.6-&2(- 50

"- $"-2*)"*>(&- $-.- "- -2-&6G"(3(3 30 =- 0+(50&2(-5*

0& 2,"6(+*3 50 D,0.-6(&*2(6*3 50'(2(0&603 0& 036-

4*"%2,"- "*3 D,08 - 5('0.0&2(- 50 "*3 D,0.-6(&*2(6*3

&*.4-"038 &* $.030&6-& .06.-22(G& 50 '("-40&6*3 50

D,0.-6(&- &( $%.5(5- 50 "- -5=03(G& 20","-. 2,-&5* 30

0R$*&0& - -,6*-&6(2,0.$*3 50 PY ?^\C;

A- -$*$6*3(3 50 "*3 D,0.-6(&*2(6*3 6-4>(%& *2,..0

6.-3 "- ,&(G& 50 "*3 -,6*-&6(2,0.$*3 - "- 3,$0.'(2(0

20","-.; !R(360 0+(50&2(- (&5(2-5*.- 50 D,0 "*3

-,6*-&6(2,0.$*3 (&2.040&6-& "- 37&603(3 50 4*"%2,"-3

.0"-2(*&-5-3 2*& "- +7- 50 N-3 0& "*3 D,0.-6(&*2(6*38

2*4* N-38 N-3A / 2-3$-3- d8 0 (&5,20& -$*$6*3(3 $*.

036- +7-; E-4>(%& 30 =- (&'*.4-5* ,& -,40&6* 0& "-


10/15

!"#$%!" '() *+,-. /012304560 *788

281

0R$.03(G& / 37&603(3 50 4*"%2,"-3 $.*-$*$6G6(2-3 2*4*

GR(5* &76.(2*8 $\:8 F-R8 / 5(34(&,2(G& 50 "- 0R$.03(G&

50 $.*607&-3 -&6(-$*$6G6(2-3 50 "- '-4("(- F2"SI ?^^C;

e05(-&60 -&6(2,0.$*3 2*&6.- N-3A 0 (&=(>(5*.03

50 2-3$-3-3 30 (&=(>0 $-.2(-"40&60 "- -$*$6*3(30+(6-&5* "- -2-&6G"(3(3; J* 0361 2*4$"06-40&60

0&60&5(5- "- .0"-2(G& 0&6.0 "- -$*$6*3(3 / "-3 *6.-3

+7-3 50 30K-"(L-2(G&M 30 3,)(0.0 D,0 "- 30K-"(L-2(G&

405(-&60 N-3 $,050 ""0+-. - -$*$6*3(3 $*. 405(* 50

"- $:deBPk8 * >(0&8 D,0 "- -26(+-2(G& 50 $:deBPk

$*. 0" 5030&3-4>"-"03 -

503-..*""-. "03(*&03 ",0)* 50 "- 6.-&3'0.0&2(- $-3(+-

50 -&6(2,0.$*3 50 PY D,0 "*3 .-6*&03 3("+036.03; J*

30 2*&*20 0" 402-&(34* $*. 0" 2,-" 036-3 2(6*2(&-3

(&'"-4-6*.(-3 (&60.'(0.0& 2*& "- -5=03(G& (&60.20","-.M

$*5.7- 30. D,0 "- $:deBPk 036% (&+*",2.-5- $,036*D,0 2(6*2(&-3 $.*(&'"-4-6*.(-3 $,050& -26(+-."- ?c9C;

INMUNOLOGA APLICADA AL TRATAMIENTO!" $("-. 50" 6.-6-4(0&6* 50 "-3 0&'0.405-503

-4$*""*3-3 -,6*(&4,&03 3*& "*3 2*.6(2*360.*(5038

2*& "*3 D,0 =- 5(34(&,(5* "- 4*.6-"(5-5 50 a[`

- \`; P*. 3, 0'026* -=*..-5*. 50 0360.*(5038 30 =-&

04$"0-5* (&4,&*4*5,"-5*.03 2*4* -L-6(*$.(&-8

4(2*'0&*"-6* 4*'06("8 (&4,&*)"*>,"(&- =,4-&-

(&6.-+0&*3- ?OWOYC8 2(2"*'*3'-4(5-8 406*6.0R-608

2*4$,036*3 50 *.* / $"-34-'%.03(3 ?IC;

X-3( 6*5-3 "-3 2%","-3 6(0&0& .020$6*.03 2(6*$"134(2*3

$-.- "*3 2*.6(2*360.*(503M 2,-&5* 0" 2*.6(2*360.*(50

30 '("-2(*&03 50 "(&'*2(6*3 E - "*3 60


11/15

!"#$%!" '() *+,-. /012304560 *788

282

B,40&6-& 0& 4-/*. $.*$*.2(G& "*3 "(&'*2(6*3 E

2**$0.-5*.03 D,0 "*3 3,$.03*.03 /8 0&6.0 *6.*3

0'026*38 6-4>(%& 30 (&=(>0 "- $.030&6-2(G& -&6()%&(2-

0& "-3 2%","-3 $.030&6-5*.-3 50 -&67)0&*3 ?BPX8 $*.

"- 3()"- 0& (&)"%3 50 -&6()0& $.030&6(&) 20""3C; A-(&=(>(2(G& 50 "*3 "(&'*2(6*3 F .0D,(0.0 5*3(3 4,/ -"6-3

50 2*.6(2*360.*(503; !" 6.-6-4(0&6* $*. $,"3*3 2*&

5*3(3 -"6-3 50 2*.6(2*360.*(503 $0.4(60 5(34(&,(. "-

37&603(3 50 -&6(2,0.$*3 $*. "*3 AF; A*3 2*.6(2*360.*(503

6-4>(%& -'026-& - "*3 0*3(&G'("*3 -" 5(34(&,(. 3,

"(>0.-2(G& 50350 "- 4%5,"- G30- / -,40&6-."03 "-

-$*$6*3(3M *6.-3 2%","-3 -'026-5-3 3*& "*3 4*&*2(6*3

/ "*3 4-2.G'-)*3 ?cIC;

A- -L-6(*$.(&- 03 ,& 50.(+-5* (4(5-LG"(2* 50 "-

^S40.2-$6*$,.(&- ?^SePC 2,/* 402-&(34* $.02(3*

50 -22(G& 30 5032*&*20; Z0 =-& 3,)0.(5* +-.(-3

=($G603(3 2*4* "- $.*5,22(G& 50 ^SeP D,0 -26_-

2*4* ,& -&6(406->*"(6* 50 "-3 $,.(&-38 ,& $*3(>"0

>"*D,0* 50 ).,$*3 oZ@ 405(-&60 -"D,("-2(G&8

"- (&=(>(2(G& 50 4_"6($"03 +7-3 0& "- >(*37&603(3

50 12(5*3 &,2"0(2*3M *6.*3 $*3(>"03 402-&(34*3

3*&V 30 $.0+(0&0 "- $.*"('0.-2(G& 50 "(&'*2(6*3

(&+*",2.-5*3 0& "- 5060.4(&-2(G& / -4$"('(2-2(G&

50 "- .03$,036- (&4,&08 -50413 50" 5-K* -" BTJ -

6.-+%3 50 "- (&2*.$*.-2(G& 50 6(*-&1"*)*3 $,.7&(2*3

?c:C;

j6.* 50 "*3 6.-6-4(0&6*3 D,0 30 =-& 04$"0-5*

03 "- (&4,&*)"*>,"(&- (&6.-+0&*3- ?OWOYCM 0&6.0"*3 402-&(34*3 $.*$,036*3 $-.- 0""- 0361& "*3

3(),(0&603V .0),"-2(G& 50 "-3 ',&2(*&03 50" .020$6*.

N28 5(34(&,2(G& 50" 5-K* 6(3,"-. 405(-5* $*.

2*4$"040&6*8 &0,6.-"(L-2(G& 50 -,6*-&6(2,0.$*3

$*. -&6(2,0.$*3 -&6(5(*6($*8 (&60.'0.0&2(- 50

2(6*2(&-38 4*5,"-2(G& 50 "-3 ',&2(*&03 0'026*.-3 50

"*3 AE8 AF / 50" 3(3604- .06(2,"*0&5*60"(-" / -,40&6*

50" -2"-.-4(0&6* 50 "*3 -&6(2,0.$*3; !36,5(*3 0&

4*50"*3 4,.(&*3 =-& 504*36.-5* D,0 "- 0'(2-2(-

60.-$%,6(2- 50" 6.-6-4(0&6* 2*& OWOY 0& PY

50$0&50 50" .020$6*. &0*&-6-" N2f&M "- 0R$"(2-2(G&

- 036* &* 03 2"-.- $0.* $.*>->"040&60 30 50>- - ,&

402-&(34* D,0 (&+*",2.- -&6(2,0.$*3 -&6(5(*6($*;

B504138 "*3 .-6*&03 50'(2(0&603 0& N2f& .03(360&

"- (&5,22(G& 0R$0.(40&6-" 50 PY / 4,036.-& ,&-

5(34(&,2(G& 50 O)W $-6*)%&(2-; !" N2f& $,050 30.

,& >"-&2* 60.-$%,6(2* $.*40605*. 0& PY ?c]C;

!" 4(2*'0&*"-6* 4*'06(" 03 ,& (&=(>(5*. 50 "- 0&L(4-

(&*3(&S4*&*'*3'-6*S503=(5.*)0&-3- .0D,0.(5- $-.-

"- 37&603(3 50 &*+* 50 "-3 $,.(&-3 0 (&5(3$0&3->"0

$-.- "- $.*"('0.-2(G& 50 "(&'*2(6*3M 30 "* ,3- 2*4*

2*-5/,+-&60 0& "- 60.-$(- 50" PY / 0" PN 50 5('72("4-&0*.-5*.03 036,5(-.*& - ]I $-2(0&603 2*& PN /

PY 50 6.-6-4(0&6* 5('72(" - D,(0&03 "03 -54(&(36.-.*&

4(2*'0&*"-6* 0& 5*3(3 50 :\ - ][ 4)HU)M (&'*.4-.*&

.04(3(G& 0& 0" c\` 50 "*3 2-3*38 *>60&(5- (&2",3* 2*&

"- 40.4- $.*).03(+- 50 "*3 2*.6(2*360.*(503 ?c\C;

Z0 =- $.*$,036* D,0 "*3 -)*&(36-3 2*"(&%.)(2*3

$,050& 30. _6("03 0& 0" 6.-6-4(0&6* 50" PY; J),/0&

/ 2*"->*.-5*.03 036,5(-.*& ,& 4*50"* 4,.(&*

503-..*""-5* 405(-&60 "- 6.-&3'0.0&2(- $-3(+- 50

-&6(2,0.$*3M - 03*3 .-6*&03 30 "03 -54(&(36.G 2-.>-2*"8,& (&=(>(5*. 50 "- -206("2*"(&0360.-3-8 / 30 0+(50&2(G

40"-2(*&03 50 "(&'*2(6*3 -,6*..0-26(+*3 0'026*.03

/ "(&'*2(6*3 E .0),"-5*.03; A- 2*4$.0&3(G& 50 "-

(&4,&*$-6*)%&03(3 $0.4(6(.1 0" 503-..*""* 50 &,0+*3

6.-6-4(0&6*38 413 03$027'(2*3 / 30),.*3;


12/15

!"#$%!" '() *+,-. /012304560 *788

283

REFERENCIAS BIBLIOGRFICAS9; i50/ eX8 Z6-&"0/ lf; P04$=(),3SS5(30-303 *' -&S

6(5034*3*4-" -,6*(44,&(6/; lBeB; 9aaa B,)

99MIdI?^CV\cIS^;

I; p*"'' k8 W*"534(6= A8 k-6L Z8 W("2=.036 F8 P-""0. B8

A0''0"" T; N(6L$-6.(2U,3 T0.4-6*"*)/ (& W0&0.-" e05(S

2(&0; J0Q n*.UV e2W.-QS@(""M I[[d;

:; f*2=-SB"+-.0L f8 j.60)-SA*-/L- BW8 N.(054-& @8

X-4$>0"" O8 B*U( Y8 f(+(66( !B8 06 -"; !&504(2 $04$=(S

),3 +,")-.(3; B.2= T0.4-6*"; I[[c l,"M9]:?cCVda\Sa;

]; B>(5- j8 k-""0"SZ0""-4( e8 l*"/ P8 F0& B/05 e8 q(6*,&(

e8 e-34*,5( B8 06 -"; B&6(S5034*)"0(& 9 -&6(>*S

5(03 (& =0-"6=/ .0"-605 -&5 ,&.0"-605 3,>.r,SY0"0L Be8 @-3=(4*6* E8 F*""-) pF8 E*>G&

B..*/-+0 Z8 B>.r,SY0"0L X!8 A*&5*K* eA8 06 -"; B

,&(D,0 '*.4 *' 0&504(2 $04$=(),3 (& &*.6=0.&

X*"*4>(-; l B4 B2-5 T0.4-6*"; I[[: j26M]a?]CV\aaS

^[d;

d; l,55 kP8 A0+0. pN; X*..0"-6(*& *' -&6(>*5(03 (& 3U(&

-&5 30.,4 Q(6= 5(30-30 30+0.(6/ (& $04$=(),3; B.2=

T0.4-6*"; 9aca B$.M99\?]CV]IdS:I;

a; W*.>3U/ W8 Z60(&>0.) eZ; O3*"-6(*& *' 6=0 (&60.20"",S

"-. )"/2*$.*60(&3 *' 5034*3*403; l X0"" F(*"; 9ad9

l,"Ma[?9CVI]:Sd;

9[; W.00& kl8 P-../ TB8 Z60(&0.6 Pe8 Y(.-6- eA8 p-)&0.fe8 B&)36 FT8 06 -"; Z6.,26,.0 *' 6=0 =,4-& 5034*S

$"-U(&3; O4$"(2-6(*&3 '*. ',&26(*& (& 6=0 5034*3*S

4-" $"-D,0; l F(*" X=04; 9aa[ l," \MI^\?9aCV99][^Sc;

99; X*Q(& P8 k-$$.0"" @P8 N.-&U0 pp8 E-4U,& l8 @/&03

fj; P"-U*)"*>(&V - $.*60(& 2*44*& 6* 5(''0.0&6

U(&53 *' (&60.20"","-. -5=0.(&) (&0. F8 06 -"; T034*3*4-" 2-5=0S

.(& >(&5(&) 5*4-(&3 *' $"-U*)"*>(&; l F(*" X=04;9aa^ e-/ :MIc9?9dCV9[a[]Sa;

9:; k*2= Pl8 p-"3= el8 Z2=40"L e8 W*"532=4(56 eT8

q(4>0"4-&& f8 N.-&U0 pp; O50&6('(2-6(*& *' 503S

4*)"0(&8 - 2*&36(6,6(+0 5034*3*4-" )"/2*$.*60(&8

-3 - 404>0. *' 6=0 2-5=0.(& '-4("/ *' 20"" -5=03(*&

4*"02,"03; !,. l X0"" F(*"; 9aa[ j26M\:?9CV9S9I;

9]; W-..*5 Tf8 e0..(66 Bl8 J(0 q; T034*3*4-"

2-5=0.(&3; X,.. j$(& X0"" F(*"; I[[I j26M9]?\CV\:cS

]\;

9\; e-=*&0/ eW8 p-&) q8 f*6=0&>0.)0. k8 k*2= Pl8

B4-)-( e8 Z6-&"0/ lf; !R$"-&-6(*&3 '*. 6=0 2"(&(2-"-&5 4(2.*32*$(2 "*2-"(L-6(*& *' "03(*&3 (& $04S

$=(),3 '*"(-20,3 -&5 +,")-.(3; l X"(& O&+036; 9aaa

N0>M9[:?]CV]^9Sd;

9^; B4-)-( e8 k"-,3Sk*+6,& Y8 Z6-&"0/ lf; B,6*-&6(>*S

5(03 -)-(&36 - &*+0" 0$(6=0"(-" 2-5=0.(& (& $04$=(S

),3 +,")-.(38 - 5(30-30 *' 20"" -5=03(*&; X0""; 9aa9

J*+ IaM^c?\CVd^aScc;

9c; J),/0& YE8 J5*/0 B8 W.-&5* ZB; J*+0" =,4-&

-"$=-a -206/"2=*"(&0 .020$6*. .0),"-6(&) U0S

.-6(&*2/60 -5=03(*& (3 6-.)0605 >/ P04$=(),3

+,")-.(3 -,6*(44,&(6/; B4 l P-6=*"; I[[[ j26M

9\c?]CV9:ccSa9;

9d; J),/0& YE8 J5*/0 B8 W.-&5* ZB; P04$=(),3 +,")-S

.(3 -&6(>*5/ (50&6('(03 $04$=-R(&; B &*+0" U0.-6(&*S

2/60 -&&0R(&S"(U0 4*"02,"0 >(&5(&) -206/"2=*"(&0; l

F(*" X=04; I[[[ Z0$ IIMIc\?:dCVIa]^^Sc ;

9a; X*LL-&( !8 T-" F0""* eW8 e-36.*)(-2*4* B8 T.*30.-

e8 P-.*5( B; B&6(5034*$"-U(& -&6(>*5(03 (& $04$=(S

),3 +,")-.(3; F. l T0.4-6*"; I[[^ B$.M9\]?]CV^I]Sd;

I[; B>.r,SY0"0L Be8 F0,6&0. !@8 e*&6*/- N8 F*""-) pF8

@-3=(4*6* E; B&-"/303 *' -,6*-&6()0&3 (& - &0Q

'*.4 *' 0&504(2 $04$=(),3 '*"(-20,3 (& X*"*4>(-;

l B4 B2-5 T0.4-6*"; I[[: j26M]a?]CV^[aS9];

I9; @(3-4-63, n8 B>.0, Y0"0L Be8 B4-)-( e8 j)-Q- ee8

k-&L-U( E8 @-3=(4*6* E; X*4$-.-6(+0 36,5/ *' -,S

6*-&6()0& $.*'("0 >06Q00& X*"*4>(-& -&5 F.-L("(-&

6/$03 *' 0&504(2 $04$=(),3 '*"(-20,3 >/ +-.(*,3

>(*2=04(2-" -&5 4*"02,"-. >(*"*)(2-" 602=&(D,03; l

T0.4-6*" Z2(; I[[: l,&M:I?9CV::S]9;


13/15

!"#$%!" '() *+,-. /012304560 *788

284

II; J-)-3-U- E8 J(3=(',=-.5 kA8 i60. p8 p-334,6= f8 W.sS

6L(&)0. l8 Z2=,"6L !8 06 -"; E 20"" .02*)&(6(*& *' 503S

4*)"0(& : $0$6(503 (& $-6(0&63 Q(6= $04$=(),3 +,"S)-.(3 -&5 =0-"6=/ (&5(+(5,-"3; l O44,&*"; I[[] e-.

9\M9cI?^CV:dd:SaI;

I^; k"0(& l8 Z-6* B; E=0 @AB 3/3604; N(.36 *' 6Q* $-.63; J

!&)" l e05; I[[[ Z0$ cM:]:?9[CVc[ISa;

Ic; Y0)-Se04(0=-+(*.-"8 405(2-"8 -&5

D,-"(6-6(+0 '**5 '.0D,0&2/ D,036(*&&-(.0; O&6 l T0.S

4-6*"; I[[9 Z0$M][?aCV\^ISa;

:\; B>.%, Y%"0L Be8 p-.'+(&)0 W8 @0..0.- pA8 B>.%, Y%S

"0L X!8 e*&6*/- e N8 @-.5/ Te8 06 -"; T06026(*& *'

40.2,./ -&5 *6=0. ,&5060.4(&05 4-60.(-"3 (& 3U(&

>(*$3(03 *' 0&504(2 $04$=(),3 '*"(-20,3; O&6 l T0.S

4-6*"; I[[: j26MI\?\CV:d]Sa9;

:^; p-&) WSh8 v, @8 p-&) nSk8 W-* vS@8 q=-* n8 @0 X8 06

-"; @()=0. $.0+-"0&20 *' =,4-& =0.$03+(.,3 d TJB30D,0&20 -&5 3$02('(2 O)W -&6(>*5(03 (& $-6(0&63

Q(6= $04$=(),3 (& X=(&-; l B4 B2-5 T0.4-6*"; I[[\

e-.M\I?: P6 9CV]^[Sc;

:c; q(""(U0&3 T8 B4>-2= B8 q0&6&0. B8 T,440.

f8 Z2=w33"0. e8 F,.) W8 06 -"; !+(50&20 '*.

20""405(-605 (44,&0 402=-&(343 (& 6=0

$-6=*"*)/ *' $04$=(),3; F. l T0.4-6*"; 9aa: l,&M

9Id?^CV^:^S]:;

:d; Y0"54-& X8 Z6-,>0. B8 p-334,6= f8 i60. p8 Z2=,S

"0. W8 @0.6" e; T(2=*6*4/ *' -,6*.0-26(+0 E=9 -&5

E=I 20"" .03$*&303 6* 5034*)"0(& : (& $-6(0&63 Q(6=

$04$=(),3 +,")-.(3 ?PYC -&5 =0-"6=/ 2-..(0.3 *'PYS-33*2(-605 @AB 2"-33 OO -""0"03; l O44,&*"; I[[:

l-&M9M9c[?9CV^:\S]I;

:a; k.(2=0"( T8 T-+(5 e8 N.,3(2Sq"*6U(& e8 W*"534(6= T8

f->(&*+ e8 Z,"U03 l8 06 -"; E=0 5(36.(>,6(*& *' $04S

$=(),3 +,")-.(3SO)W 3,>2"-3303 -&5 6=0(. .0-26(+(S

6/ Q(6= 5034*)"0(& : -&5 9 (& $04$=(),3 $-6(0&63

-&5 6=0(. '(.36S50).00 .0"-6(+03; F. l T0.4-6*"; I[[[

B,)M9]:?ICV::cS]I;

][; f(2* el8 F0&&(&) X8 p0(&)-.6 !Z8 Z6.0("0(& fT8 @-""

fP; X=-.-260.(L-6(*& *' 3U(& 2/6*U(&03 (& >,""*,3

$04$=()*(5 -&5 $04$=(),3 +,")-.(3; F. l T0.4-6*";

9aaa l,&M9][?^CV9[caSd^;

]9; Y0"54-& X8 @s=&0 B8 T(02U4-&& T8 Z2=,"0. W8 @0.6"

e; E/$0 O .0),"-6*./ E 20""3 3$02('(2 '*. 5034*)"0(& :

-.0 4*.0 '.0D,0&6"/ 50602605 (& =0-"6=/ (&5(+(5,-"3

6=-& (& $-6(0&63 Q(6= $04$=(),3 +,")-.(3; l O44,S

&*"; I[[] e-/ 9\M9cI?9[CV^]^dSc\;


14/15

!"#$%!" '() *+,-. /012304560 *788

285

]I; Y0"54-& X8 P-=" B8 F0(330.6 Z8 @-&30& p8 F,0. l8

T(02U4-&& T8 06 -"; O&=(>(6(*& *' 6=0 6.-&32.($6(*&

'-26*. N*R$: 2*&+0.63 5034*)"0(& :S3$02('(2 6/$0

9 .0),"-6*./ E 20""3 (&6* E=IS"(U0 20""3; l O44,&*";

I[[^ e-. 9M9c^?\CV:I9\SII;]:; @0.6" e8 !4(&) f8 Y0"54-& X; E 20"" 2*&6.*" (&

-,6*(44,&0 >,""*,3 3U(& 5(3*.50.3; l X"(& O&+036;

I[[^ e-/M99^?\CV99\aS^^;

]]; h(-& n8 T(-L AB8 n0 l8 X"-.U0 Z@; T(33026(&) 6=0

-&6(S5034*)"0(& -,6*.0-26(+0 F 20"" .0$0.6*(.0 (&

$04$=(),3 +,")-.(3 $-6(0&63; l O44,&*"; I[[c e-/

9M9cd?aCV\adISa[;

]\; J(3=(',*5/ $.*5,26(*&; l O&+036

T0.4-6*"; I[[[ l-&M99]?9CVddSa];

]^; e0."*> P8 e06LU0. B8 @-L-L F8 f*)*+(& @8 f0(3&0.

Z@; J0*&-6-" $04$=(),3 +,")-.(3; P05(-6.(23; 9ad^

T02Mcd?^CV99[IS\;

]c; W-"-.L- X8 W,6(%..0L !A8 f-4*3 p8 E0""* e8 f*&20.*3

W8 B"+(L,.( Z8 06 -"; t!&504(2 $04$=(),3 '*"(-20,3

(& - $.0)&-&6 Q*4-&; f0$*.6 *' *&0 2-30u; f0+ e05

X=("; I[[a Z0$M9:c?aCV9I[\Sd;

]d; B&=-"6 Wl8 A->(> fZ8 Y**.=003 ll8 F0-"3 EN8 T(-L AB;

O&5,26(*& *' $04$=(),3 (& &0*&-6-" 4(20 >/ $-33(+0

6.-&3'0. *' O)W '.*4 $-6(0&63 Q(6= 6=0 5(30-30; J !&)"l e05; 9adI e-/ I[M:[^?I[CV99daSa^;

]a; Z2=("6L lf8 e(2=0" F; P.*5,26(*& *' 0$(50.4-"

-2-&6=*"/3(3 (& &*.4-" =,4-& 3U(& (& +(6.* >/

6=0 O)W '.-26(*& '.*4 $04$=(),3 30.,4; l O&+036

T0.4-6*"; 9ac^ B,)M^c?ICVI\]S^[;

\[; p-32=U0 l8 F.,))04-& P8 F-,4)-.6&0. p8 q(""(U0&3

T8 T.0&2U=-=& T; P04$=(),3 '*"(-20,3 O)W 2-,303

5(33*2(-6(*& *' 5034*)"0(& 9S2*&6-(&(&) "*2U(&) 5034*)"0(& 9 6.-&3(&60.-26(*&;

l X"(& O&+036 I[[\ J*+M99\?99CV:9\cS^\;

\9; k*2= Pl8 e-=*&0/ eW8 O3=(U-Q- @8 P,"UU(&0& A8i(66* l8 Z=,"6L A8 06 -"; E-.)0605 5(3.,$6(*& *' 6=0

$04$=(),3 +,")-.(3 -&6()0& ?5034*)"0(& :C )0&0 (&

4(20 2-,303 "*33 *' U0.-6(&*2/60 20"" -5=03(*& Q(6= -

$=0&*6/$0 3(4("-. 6* $04$=(),3 +,")-.(3; l X0"" F(*";

9aac l,& IM9:c?\CV9[a9S 9[I;

\I; B4-)-( e; B,6*(44,&(6/ -)-(&36 5034*3*4-"

2-5=0.(&3 (& $04$=(),3; l T0.4-6*" Z2(; 9aaa

l,&MI[?ICVaIS9[I;

\:; A-$(0.0 lX8 W,(6-.6 l8 !66"(& TB8 X=0& Te8 B4-)-( e8

X=-& AZ; P.0'0.0&6(-" -26(+-6(*& *' 6=0 2*4$"040&63/3604 (& 6=0 "*Q0. 0$(50.4(3 *' $-6(0&63

Q(6= $04$=(),3 +,")-.(3; F. l T0.4-6*"; 9aad

J*+M9:a?\CVd\9S];

\]; F.w))04-&& e8 p(""(-43 WE8 F(&5*& XO8 X"-.U ef8

p-"U0. ef8 l0''0.(3 f8 06 -"; X*4$-.(3*& *' 6=0

0''026*. ',&26(*&3 *' =,4-& (44,&*)"*>,"(&3 ,3(&)

- 4-62=05 306 *' 2=(40.(2 -&6(>*5(03; l !R$ e05;

9adc J*+ 9M9^^?\CV9:\9S^9;

\\; e-32-.G le8 !3$-K- B8 A(, q8 T(&) v8 ZQ-.6L Zl8

N-(."0/ lB8 06 -"; e02=-&(343 *' -2-&6=*"/3(3 (&

$04$=(),3 +,")-.(3V .*"0 *' O)W +-"0&20; X"(&

O44,&*" O44,&*$-6=*"; 9aac j26Md\?9CVa[S^;

\^; B&=-"6 Wl8 E("" Wj8 T(-L AB8 A->(> fZ8 P-60" @P8

!-)"360(& JN; T0'(&(&) 6=0 .*"0 *' 2*4$"040&6

(& 0R$0.(40&6-" $04$=(),3 +,")-.(3 (& 4(20; l

O44,&*"; 9ad^ J*+ 9M9:c?aCVId:\S][;

\c; A(, q8 W(,5(20 Wl8 ZQ-.6L Zl8 N-(."0/ lB8 E(""

Wj8 E.*/ lA8 06 -"; E=0 .*"0 *' 2*4$"040&6 (&

0R$0.(40&6-" >,""*,3 $04$=()*(5; l X"(& O&+036;

9aa\ B$.Ma\?]CV9\:aS]];

\d; Z1&2=0LSX-.$(&60.* O8 !3$-K- B8 P0"-2=* F8 AG$0L

e*.-6-""- J8 f,>0&360(& TZ8 T(-L AB8 06 -"; O& +(+*

>"*2U-50 *' $04$=(),3 +,")-.(3 -2-&6=*"/3(3 >/

(&=(>(6(*& *' (&6.-20"","-. 3()&-" 6.-&35,26(*&

2-32-503; F. l T0.4-6*"; I[[] Z0$M9\9?:CV\^\Sc[;

\a; n-4-4*6* n8 B*/-4- n8 Z=, !8 E3,&*5- k8

B4-)-( e8 k(6-"(360.(&); l X0"" F(*";

I[[^ T02 ]M9c\?\CVcI9Sc;

^9; @*''4-&& X8 P*$ e8 A004=,(3 l8 Z2=(.40. l8 BU6*.(03

k8 Z2=4(56 W; E=0 n0.3(&(- $30,5*6,>0.2,"*3(3

2/6*6*R(2 &02.*6(L(&) '-26*. ?XJNnC 30"026(+0"/

-26(+-603 f=*B; l F(*" X=04; I[[] B$.

9^MIca?9^CV9^[I^S:I;


15/15

!"#$%!" '() *+,-. /012304560 *788

286

Î; F0.U*Q(6L P8 @, P8 A(, q8 T(-L AB8 !&)=("5 ll8 X=,-

eP8 06 -"; T034*3*40 3()&-"(&); O&=(>(6(*& *'

$:deBPk $.0+0&63 $04$=(),3 +,")-.(3 O)WS(&5,205

2/6*3U0"06*& .0*.)-&(L-6(*&; l F(*" X=04; I[[\ l,&

I]MId[?I\CVI:ccdSd];

^:; X=0.&/-+3U/ BO8 B..05*&5* l8 k(6-(& -3 U0/ 3,$$.033*. *' 2Se/2 (& 6=0 3U(&;

!eFj l; I[[^ l," I^MI\?9]CV:IadS:[a;

66. Wang X, Brggre F, Frusi-Zlotkin M, Feinmesser M,

e(2=0" F8 e("&0. n; P*33(>"0 -$*$6*6(2 402=-&(34 (&

0$(50.4-" 20"" -2-&6=*"/3(3 (&5,205 >/ $04$=(),3

+,")-.(3 -,6*(44,&*)"*>,"(&3; B$*$6*3(3; I[[]

e-.Ma?ICV9:9S]:;

^c; P,+(-&( e8 e-.2*&( B8 X*LL-&( !8 P(&20""( X; N-3

"()-&5 (& $04$=(),3 30.- (&5,203 U0.-6(&*2/60

-$*$6*3(3 6=.*,)= 6=0 -26(+-6(*& *' 2-3$-30Sd; l

O&+036 T0.4-6*"; I[[: l-&M9I[?9CV9^]Sc;

^d; X(.(""* J8 A-&L- e8 N04(-&* N8 W-06- We8 T0 f*3-

B8 W*4>*3 N8 06 -"; O' $04$=(),3 +,")-.(3 O)W -.0

6=0 2-,30 *' -2-&6=*"/3(38 &0Q O)WS(&50$0&50&6

402=-&(343 -.0 6=0 2*&2-,30; l X0"" P=/3(*"; I[[c

Z0$MI9I?:CV\^:Sc;

â; F=*" kX8 T03-( B8 k,4-.( Z8 X*"*& l!8 B=405

Bf; P04$=(),3 +,")-.(3V 6=0 .*"0 *' OAS9 -&5 OAS9

.020$6*. -&6-)*&(36 (& $-6=*)0&03(3 -&5 0''0263 *'

(&6.-+0&*,3 (44,&*)"*>,"(& *& 6=0(. $.*5,26(*&;

X"(& O44,&*"; I[[9 B,)M9[[?ICV9cISd[;

c[; j."*+ eT8 X=0.&/-+3U/ BO8 B..05*&5* l8 W.-&5*

ZB; Z/&0.)(36(2 -26(*&3 *' $04$=(),3 +,")-.(3 O)W8N-3S"()-&5 -&5 6,4*. &02.*3(3 '-26*.S-"$=- 5,.(&)

(&5,26(*& *' >-3-" 20"" 3=.(&U-)0 -&5 -2-&6=*"/3(3;

B,6*(44,&(6/; I[[^ J*+M:a?cCV\\cSÎ;

c9; j&* k8 @-& l; E=0 $:d 3()&-" 6.-&35,26(*& $-6=Q-/V

-26(+-6(*& -&5 ',&26(*&; X0"" Z()&-"; I[[[;

l-&M9I?9CV9S9:;

cI; k-3$0.U(0Q(2L e8 Z2=4(56 !; X,..0&6 6.0-640&6 *'

-,6*(44,&0 >"(360.(&) 5(30-303; X,.. T.,) T(32*+

E02=&*"; I[[a T02M^?]CVIc[Sd[;

73. Bardek I, Milavec-Pureti V, Lipozenci J.

BL-6=(*$.(&0 (& 50.4-6*"*)/; B26- T0.4-6*+0&0.*"X.*-6; I[[c l-&M9\?]CVI^]Sd;

c]; A( J8 q=-* e8 @("-.(*SY-.)-3 l8 P.(3-/-&= P8 p-..0&

Z8 T(-L AB8 06 -"; X*4$"060 N2f& 50$0&50&20 '*.

(&6.-+0&*,3 O) 6=0.-$/ (& -,6*(44,&0 3U(& >"(360.(&)

5(30-303; l X"(& O&+036; I[[\ T02M 99\?9ICV:]][S\[;

c\; e(4*,&( T8 B&=-"6 Wl8 X,44(&3 TA8 k*,>-

Tl8 E=*.&0 l!8 J*,3-.( @X; E.0-640&6 *'

$04$=(),3 +,")-.(3 -&5 $04$=(),3 '*"(-20,3 Q(6=

4/2*$=0&*"-60 4*'06("; B.2= T0.4-6*"; I[[: l,&M

9:a?^CVc:aS]I;

c^; J),/0& YE8 B..05*&5* l8 X=0.&/-+3U/ BO8 P(660"U*Q

ef8 k(6-

penfigo inmunopatogenia

Documents