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Go60dbgptl6 (3o68ogbg) gpo@8U6 o68D- receptor- T-ti%6ogolj 6oGo3®co6o), bogco B-go8-
6oob@o tlo8ogpgg:oU6Dg @6b8Dgo gpo@8noooo
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86o68ogob co6bb6oD, 6n8ogtloG B-go88ooo0o
granular lymphocyte), 6co8goboG 86co8b6ocrtobe,b6
@vog38Oab. B-go88cOGOOO b6Oobbo@gob vb6-
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b60oao6oalj B-go@g}noooob o@b6rpa,goa@go606o
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CD19, CD20 gt> CD22.
bDbd6006Oabb b6Oaflgodoab;
T-go@8a]ooO8ao
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at>@O@G6Oao 6OGojoodo - TCR (cD3) vt>68068- 3o3t}g6Goaxp 0666tjso6@6ooob 36ooobo t}t>6at>g
t>6ob @obv68gpogpo.
a,06b. 86b fio8xpo8, 6t>G TCR a60oao6b tlooG-
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d808OO) Ba8OOT88Ogco6a66 06Oi-0607 8ti6aGot>b 8ooo6o - ti86oE86og?0 66rpgo6oaoco Gooodo-
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8U6dooob 608tiRaGo6 t>8 ti*rSoxpoaob g6b&>6o-
at>b i,6 tjti36otm>b 8tigpobb8oatj; at>a6d08Oaob t]0807b808t>HO, cD4 a%6ORO Thi,
- 8o6t)t)ooi o68oGof4oa@g t)%6orgoatj, tiGbrp Th2 t>6 Thi7 gpo8800006R 80806060068Oa6.
dbo8Ogpoab 66 t!081j036uf4 @%6Oxpoab t>668- (bu6DcOO 1.15). @Obo goq}0606006Oaob @O@b6-
07tigoat>b 866dooti6Oaob t>6OEO t>6boa@qpo
au6Oalj, 6t,1jt>G b68Ofo6O u*608Oafoo 3068t>306O
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TCR rp6o bDboljbb. ob Ch, P b6 y, 6 3rpgojo3- 608ogb6o @6d608a8oao t>6 NK-t]*6oRoao
Oogo6o %D83oaobb gb 8bo7or>D6 bbrpGo68atgo 868OO8ut}t>8Oab, 08u6ti6 36tiubb Thl aq]Ooi
CD3 t}083godboco66 V66@og806ogpo. bobbgpob V66866o768ti; IL-4 t}o, 6o@ogbt>G NKT @K6o-
9o88oGo6o)a 90-95%-b Cup TCR, bogrp 5-10%-b fpoao t>6 3Ob006O ti*6O"ao 868OO@uE68Oab,
y8 TCR Dd8b. Th2 8foab 6o)t>8b. t>tif¥t>bt>6b t>g8oBo6oRo Thl7
T-8O880006Oao, cD4 xp6 cD8 8t>6d06Ob qpo88oGo6oaob Ro80606Go6oat>b Go6odo6oao
- IL-6 gb TGFP - oV808b.
Odtj3f4Oijoob 8OboR8OcO, 866tjti868Oaugp 303a-
gt)Goat V66@06gao606. CD4 856do6ob 8do6o Thl ux608Oao, 6O@goboG Ogp,2, f¥088O-
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t}o, Ujo660otjt>g, oo6o6odbot)@6 8@6dGo6b 6tigpoao6 xpo t>Raogrpa6o8 t>6oioaoco 606dGooaEo
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8Ogpodti8Oao]66 d083gpodljtlo 868OOG6Oa06 Ob btia303t]g6006 IL-4, IL-5, IL-6, IL-io, IL-i3-
(tjti6t>coo 2.3). 58go6og, ti*6ogob foog63o6foo b6 ga 8666@gpoGo6-86d6o868ob dogo6oob
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cD8+ Ong6ORoao uot>8806b. cD4 ti*f4O"aob 95% gpo@8cOGOooao, B8OuRoa608, Thl, Th2 R6 Thl7
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8Ob088Oot, 866bb868Oat]gpo 8ti6dGOob tj@a303u- CD8 ax6o"aoG, 868rp8t)tl68oat)go oooodo-
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0708t)t!tlo T gpo88OGO®Oaob a668Ocot>6Oa6
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19

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1995 V. bbjbauB8b (S.Sakaguchi) ortb-
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Vgpobtlo od6b 8rp3o8oatlgo gD j8gpo8D b@ @o8b6- b6o8o @D6oaugo6oaogo T u%do"bo (Treg).
cO@goaoco 86dogpgoat>. "666dgt>tjod@6" T go8-
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q}cOGOooab @Oodtioi86Oab: 06@DaDg 68ab®Ou6O O@ti6t]60 3bbtlbob 8bot68fl68brao @OtD8t>60 6Ogo
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D@ tl08b6Oaoot bbqpbgb880806Og fl%6O"bb
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8bdo®6o T-bet. ob u%6ogoao, IL-12-oortb b6 IL-
a6``6ugobo (G), @o6r.{jo6q?Gob (M), gD ofoogTo6oabgpo 18-oov b6o8ugbGoob @08"b, Thl Goocojo6oab
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a,``gb606aogo, bo@b036fl6O b6 806fltjoot 0680- ILCs-ob Kat]8o 3. IL-17A Rt>^>6 IL-22 oo-
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21

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57

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histocompatibility complex) - Dtj6ug8ab.
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MHC @rr)gojugoao, 6o8ogbbG t>gb@ot>6ob
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CD3

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59

tyrosine-based activation motifs) tjod306bobb nlooGb8b


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d066qiobob 8006 ITAM-Ob OT06Ofo06Oaob 8Oti-
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6t>G o86b 6otl668tj, 6o8 T-go@SoGoOo TCR-Rt>6
808Obt)g b086t>8b t>6 t]36tj@b8ab.
TCR 6oGo30o6ob er fgb P %bB8oaob @bjo-
go6obogo ao6obo o@ti6obgob@go6ob 806oaob
aen8ogoe,ot]6o6 (ob. orit>8o IX). P %683ob 866o6bo-
gfl6 go806t}, o8t6o8gob@go6ob @do@0 %6B8ob
@ti8t)8tJt>R, v, D R6 J a06Obo, togo Ch %6$8Ob
866o6aogB6 go806b, 8btat)do %t>B8ob @t58t>8baR
- V go J ao6obo 6doRo6obb. t>gt}t>6otl6t>8ot>, 6rri8
TCR 6OG03006Ob ]G 806Oao @6OE8608086t>8
@6b3gbgDb VD68rpge,06oqTo.
08fl6Oagobug:p06Obb 8b TCR-Ob @Djog068a-
bu6boio 7.5. T OOOOoodtiotidotiot8Ob b6Ooa06Ob vb6Ir og ao6oab t]o6ob Scot,8t>6o a,66bb868oa6 @5cno
a066. T GOOOooatioj€6O ti%6Ogo 0680006Obti8O 866tib8t>8oat)go oq}oJ0o6t)go 8t6dGooaoRa6 a6-
ti%6ogob q]8963o6foo MHC I+303ooRb tlooG6rnab. CD8 808Ro666ooatj. o8ti6oagobtigo6ob o88doo6tg
dooo30o6o MHC 8cpgpoj@gob or %tB8b t)j68t@o6soa6, 8@6d006b 8Obo j06b0666H6O Ha6Ob Ofo0003O
6Ocot,G T @%6Ogti6 96 t,6Ooa06Ob v6688a06 ti*6Ogb
a,66b6foR3668b. T-t]%6ORob O8Odo06@go 8fl6-
tlo6ob da8tlo6ob 6q)o6obab fo6gob g6, t>@t>t)a)t>6680,
d006 jo t>6 t>6Ob TCR-foo 868Oj08OaH8O. TCR
T t)%6ORrao boa6t>gob b6R6GO@6b tiq]6ti68Ogp808b.
@bog08 66Ooa06Ob a,68OG6Oa6b O@b6bt)6Oa6.
I,8R06t>R, TCR-Ob t306b6666ti6O tia6Ob a06Obo,
a 86 p %6B8Oaob 86606aoRti60 6t>vogoaob
66Oot5boHgob t>6t,goaot]6 t)a66cO66 i)08t,6Oboco,
G8bgoaD9coab T-go@g3cOG0007b b`3OGO8Ojb6Obob
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060 cp a06O bd8b. TCR-Ob jco6boD6Otl6O ub6Ob
66VogEo od8bo 3o306866o6bog@6o @b66o6,
6oon6G TCR a60o806o)66 t|tlt)t)go jo60adob
b88b6Oab.
CD3 do83godtjo raoR8oaa bt>8o o68t>6ot>6o@-
go 3030oRu6o *6B8obaa66 (y, 8 96 €), 6o8ogoG
gobtig808ti6O a8Oaoco 86d68tl06Ob@8 ao8ORO-
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o6co866oo)b o66G8gobt. y, 8 Ra 8 9€t>B8oab o7oorto
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8X6oq?tioq?b
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o6o7@66ocootja66 8a6tib868goa6 (a g6 P gt>goboon, (@b36®q}b8ch ( IgE ti®66®d86b, bngmoao(b6O]08b,


togo y, 8, n 86 8-ti66do8OOTO), 6t>G tj086t>gob 8ssdooa6oaS, Ab o®too6on8oqpohou bb6oo6oqpo806toob)
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GoOo3g6Q]8ti6o 6t>Vogoao. ITAM (immunoreceptor aog306ob do6ocot,go 3o3t)gp6Gooao g`` at>orio 8ti6doooao.
ao6oao @bcognxp o6b6b@o@a6b6ug 3cogpo303- @bco Duocob6ooa06Oa07b6 @rpd8OgoabG tlo@dgobco.
Oogoab bJogo6oat,. bgtjD6otl6b8ob, 6rp8, b60oljboogpotj @1jab8bbg,
bo8botl6o @U0bGoobo 86ofl86ogco8b6 6rpq:pb yoT-t]*6ogobti b60o@06ob ab8rpG6rrtaD gb8nu-
t>ij6t)goatj I,66Otjbo@goT6 @668b8806086Oaob
jopoagpt>g, MHC @nqTojbiTtoaotj Vb6pbo6ob bb-
tlod@6Dtlo. TCR-ob ao6oatlo jo brp@botdo 8tlobGob 6OtloG t]Oadgob. tico8Ogpo8O bd88D6 a,b8cO8806b60,
tjbo6ovrpxp b6 bgoab. b8otjo bbb6b T- gb B-u%6o-
8od6obo6, 6rp@ yoT-t|B€6ogo gbo8ob jo630go
"aob @D6bb8b8Obtgpo g}u6dGOoaoot BOodgoab. bbfob Vb6@obg@06b, D6u o@ti6ud 3Dbtt]Eo olj
8o6t>ogt>6, t)%6og@go R6 3t)8o6t)gpo oat)6ti6o oroT-ux6o"aD8go o6co8obb.
3t>bt)bot5 f)66co8t>ti T-go88rpoooo at>6bt>Cbp86t>8tj,
ob bt>dt)co66 oogpoafoo t>6 I)6xpa 606e,o6oagob.
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quc) @mEpot}uEpoao
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60 8uODGOob tl08Obb8 jo btjocoo gpo88OG06Oaob o68odGot6o t>e,06oo tig36ogolj o6 8t,6bb-
v``6@Od86Ob uobbdqpoaqprpab 86goab. a,b68b b@Otjt>, 8t>8Oat]8 jo@3t>66@06OEO 86t>8gp8Oa6. 806tiboao
tjo&>Oti60 8@06GOob EO8cob8086tlo T-a%6ORb,
8b fooa00607O abd606OD GOOcOQ]rigt)0 D6 a06Ov-
Uoodg¥oarpgb bbjticoD6o MHC @ngpojugob t]oG- 8t}0 86b8g"aD, a0360 3Doirpa,86t6O abdo86Ob
6Oaob u6t>6O gbodt>6ab, 6bG 801jo 8ti6dGOob gb q]naoo6o7o otdb6orpougo 3b6Dfoooo jo
xpDj`>f488bb ab@OOv808gD.
- 8ocbojtgobtlo (06gcobco@oabb gb gpofocobn8oa-
306o806obcbo b61joat)gp T go@BrpGoOcob Eo). t)%6ogoti b8 crt6o ab6bb3b8oaagpo ba6ogb6
Rt,I,bgooaoco 9o% ChpT-tix608Ot>, 1o% -yoT-ti%6O- o680dGoof4o b8060ob ogpo8o6boobb o@@6u6o
9o. cot|8Gt>, dt|B-6t>Vgpt>8ob 06t>doot) go6Vrp8t)61J6 boboo8b bti8Dgbbb8b @fooo} t>b"6b.
gD dt>6tlo yoT-a%6oxpo7b 6brr)xpo6nbb 25-30%-b oo6rpfocogtlo 8Gbco86obo 8o6uboaocob gb
b|rJVo8b. o)o8@bob o60rr)ao6ofoolJ bxp60Ug bob- bbd6Of4Ooaoco 063OG06OaDgT u%6O"alj GO-
8Ooacbo T-go88OGOoot6 t]8Ooottcoat>ti yoT-u%6O- Orr)Oodtjoju6o (CD8+) u%6o"bo b6bgau6oab.
goao tlot)R806t,. o68ooo6oaugo tey}6ogoti 8oj8goo7 ob3rrtaD bb-
OPT-u%6Opoao, a06OotbRb8, cD4+cD8-TH pb I,8O 806utjt]go 6t>vogt>tjoaob6 R6 ti*6ORtloR6
cD4-cD8+ Tc 0%6ORoa06 (bti6t>Oio 7.4 96 7.5). abdoo6ooaob Vb6@cod@6olj V8b6rp, 6oljo ilogoaoG
obo6o bb8bgDtjti8b jgpbtoti MHC 8rpgpojz)g8ab o68odGoobaD6 8bb3o68ogpo n68b6ofo@ob a,D6cob-
EOOG6rpb06 8b, Eobbab@Otjbg, @b6btj8D3Oaog g!u6-
80bo8gpobbb.
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bo6udou6Oao - cD4 8b cD8 @Ogpodugoao - go3gpt>fo@u6o 600ojtgpt)@o, 8ng9€ob jo83god-
MHc I 8D 11 jgbbob 8cogoju8Oaob @D@006cp- tjo, 06gnbrp@oao, gocboljco8oao gD tit.8b u*6ogEo-
abEo @6oT}36ogco8b6 6ngb bb6tlgoatj. CD4 - 8b 3Ofoojugobo) b6boaug 3botco806Oabb gb 8bco
MHCII, CD8 joMHC I jgpbbob 8ngpojugbb fljb8- 36cO8fldooblj T-3Og306tIpO u93608Oao (cD4+)
t]068Oab. t>66O@06Ob i]806cobolj 36OGoljtlo T-goo- bgi@cobf)86tj. Ob fl%6Ogoao, Otb8OD6Oio 8u6dGOob
q3rpGooob Cb8gb3o6foo CD4 gD CD8 @rpgojugpoao 8obog8oco, tjb8 jgpbt)bg odco8b (bu6bo)o 7.6):
TCR-otb6 Dbnoo6"ab. 1. Thl u%6ogoao, 6rp@ogcob 8u6doobb
CD4 6ooo36rp6o oar)bo o8u5me,gcoaugo- fl*60Rt}oR6 8od6coE>otj t}8ga Vt)68co5R806tj,
606o go886ob@b6 T@08g8b6o o6ot%DE8ot>6o 2. Th2 u%6ogoao, drp@goaoG IgE D6oobtio-
@ogojtlsTDb. CD8 @ngoj@gD gobugp80PU60 ugoaocoD gb onfoo6n8ogpoboo7 bb63o6oaoauqgp
a8Oot 8bjb8tl06Oatigpo Ci gb p %bB8Oaobab6 t]08- O8u6tld 6ObdGOoab Ov808b 8b
gat>6o 30006ogo806ot>. 8t>orto Gooo3g6fo@t)6o 3. Thl7 u9€6Ogoao, 6cO8gpoboG, d06OcODgbq?,
6t>Vogpo 85og6o dt>8tlo6tlo6 GoOo3gt>fo8ti6
o7o6rpfoo6do6bfoDbo7b6 (Lck), 6bG D@ @og8juq:poab
boa,6bgob Of4D6bgudGOob bD@ubgoat>1j bdgp031j.
CD4 6oGo30fp6o MHC 11 jgDtjotj 8ogpojugpob
P2 grp806b tjD8tlo6"ab, CD8 joMHC I jgbbob
@°gpoJugob Ci3 go@061j.
MHC 8cogojogpbbcob6 TCR-obb gb CD4 b6
CD8 60GO30rr)6ob O6cog6coUgo gDt}b8Eo602)b
go@8cpOOoob abbDbdoou6OagD8 bb806O b6OO-
@06Ob xpcofobb loo-%06 b@006Oab.
yoT-t)%68goao, do6ootbgbio, n6@b8bg 608t>-
Oot]6o, CD4-CD8- T-u%6ogoaot>. 8t>o7o 8t)6jGoa 6@G8j©ndoo,
a®gpco@8O bb6j8Otgpo b6 b6Ob. 8b607Dgob, foo- @t>6306ul3btlgo
boo6o7 @botbb6b, ChoT-U%688ob 8tlbD8bbg, MHC 8ba®Or,0"bo
I gb 11 jgpbtjob @ngojtgob EOG6rpab t}ot|dgob Li36b070 7.7. Oaboa063do b66Oa0601i 096688o0 8Ob-
(coU@GD ob @Don do6oovbxp 8t6dGoblJ b6 U6gb 8086Ob aq]Oao. Obfoco806t6O i,6008060 u%608t}O
vbd@cob8a06q?Ob). MHc I jgpbljob @bbb8bo cD1 tloodgoab @rpb8gob 8baoooocofoootb gb 3o6ooooncboco,
@Ogpojugpoao gD bocoat6O Bojob Oogb - Hsp b6 8o@a6b6ogT 6ooo30rp6otb6 gbjb8@o6obob i)o@gao8o
(heat schock protein) 8bco go@D6gb Vb68obgao6ti. o60066bgpofoboooon.
o@bTaob, 6rrt8 b@ o6o ab6bb8t>3oaflgo jrp83D60@06-
Oogt>6 808Oat8 3036ORoab MHc 808Ojti8Oaolj
tjtJ8t>gt>tjtJ8t> dgt)tjo Vt>6t>gao6b.
MHC I dgbtjot) 8coqpojtg`rtD Go0oforpgTopb6
@Ogoatg 303008b Ojb8E06Oab, fl%6Ogoij cb08b.
3o6foo 6t>J8b g6 CD8+ T-fl%6ogtj Vt>6@g806tj.
MHC 11 dg6bolJ 8ogojt)ga 8ofood@go6o
d083666806Oogt>6 8Og]Obngp 303008tj @t368i]06-
8Oat> 8t, 8t,tj cD4+ T-t]%6Ogb vt>6ti8806b.
b@806bg, MHc b6t 3Obory7tlocob8boaob 807b-
8bdo t)co@3gpodbolj 8rpq`r`)ojugoao L`60oao6ob
Vb6gao6btlo tl86otl86ogpri3D6ob 6rpgt t>b6tgoab.
6068060 `3Obooi]OotD8boaob t>600806O' -
bu6bcoo 7.8. MHc I 68bbob @®8acti8Ob li66@dot]db. H-b60obo6o - 3o68ogDg %o6% b6og@b ab8no-
MHC I jgbtjob 8rr)gojugb 3rpgo8rl6g}€go er %bB-
t|o6b, 6cog8bbG 06D6b3gb60boob 8riGogoaob
8ObD 8D 8t>tj07t,6 b6bt!cO8bg86Ou6b8 gbjD8tl068aogo
8b@O@v808 b6o0806Oab bt,Dbob07Oa8b. tlo@gbco@EO
f}2-8ot)6rpagprpaogo6otjaD6 i)og8obD. er %b83o u%6ogob
bg@nB6gb, 6o8 b8 b6ooe,06obb 86b8Dgpo a86()b
808a6b6bljonb6bb gbjb8@06Oatgpo.
607ugo j083godbo bj0806bab gb 8bot job-
OrpEoo)b8b8aob @otb8b6o jrp83g8dbolj t>60oe,06oao
ovog`,.
MHc 18t>II dgt>ijot5 8Ogot)tgoao t]%6ORolj
q]ogD3o6tgpo @gojo36rpooo6obob. obo6o tj06Ud-
6t}6Ootb 8D 3u6dGOooi 06Ov8b6Ocoolj @bab8bob,
O)ti@G`>, at>6bti8t>8Oaflgo b06t|jot6tgo tjEa06-
Otot)g8aobbbb6 tl08a,Oat>. ri608O @botab6b @O@a6b-
6bbo)b6 bbgTrpb @g8ab6o rr)o)bo gr.806o bd3b: (r)6o
grp@06o o8t6rpagpnaugtt6u6o grp806ott @1jab8boD,
8o@b6b6obbb6 8rptlo6oaoov 8gobb6o co6o go806o
t)0 a,6d08 8t>61j v668Od86Otj, b6Rt>O @607 303-
0ogoao fljb8Eo698bb. 1Jb8t>g``bb3b Vb68oErr7aob
3030ogoti gbdb3tlo6obt)1Jb gb Vb6@od@6ogpo jco83-
godboaob bb868t>bb86 ti%f6ORobt,R8O vt>698066b
b@ cod jgbbb tlrrt6ob b61joaugo b06tldot6tgo
8t>6bb8t>8oa6 at,6t>3o6rr>aoalj.
MHC I jcpel>ol> 8mcT3jgcpo
1ju6bcr)o 7.9. MHC I 6gbbob @®go8@gob bod@d68db.
MHC I c}qpt>bob 80gpojt|g6 o6o 3rr)go3036ogt|-
tu6bo7Cbo MHC I dgDtjob 8rngpodugob O%fiog8D6oonb
6bvogo gD @Dbortb6 q?bjb8raodobugo 3030ogob 6b- 6o %t>B8obbb6 Eo8gab6o 30006rpgo806ob (bu-
B806oao. 3030ogb6o b66obo6o 8rrtcob8boabgob gD6t)o, 6bor>o 7.8, 7.9). 3cogpo8rT768tgo Ch (@ao8o) %t>E8o
6co@Ogljbo p-EO8Oaobbab6 rao@8aD6 8bj06foo Ch-b3O- b6b3ogo@n6q3ugp P2 @oj6riie,grpaHgo6b b6b-
6DgT88o Vb6@cpd@6otj. P2-@oj6coagcob@go6o Ci %bB8b
jco8bg066t6bg tjL`8i]069Oat>. HLA I jTbbob
b6bjTp8bg06ot6bxp fljD3E068Oab. 8ogojtlgb 3 bbbobDD: HLA-A, HLA-B gb HLA-C.
8bco @o-6 d6o8ribo8t>tlo ab6gpb8oatgpo MHC bo6-
abd606ooaobb gb ljcojnobob bbvo6Dbq@goe,o
oao bjogo6oab (bu6Doio 7.1o). I)2-8oj6oagrpat-
Rt>G8t>ti a,66t>306Oaoat,.
@oj6cobugTo t>60obo6o 8ofoojugu6 jn@3b60- go6ob 8bjrDgo6obogo ao6o @o-15 d6o@cobco@bEOD
8ocot>8boaugo.
@06OT8O 060 bfo007 t,88Oab (bu6t>Oto 7.7):
I forrt8o abdoo6ob, @b8. 8ojmabdoo6oD, 8bj6rp8bb- MHCI jgDljob 8ogpojtgpober %bB8b bb@o U%-
6og8b68o7b grp806o (oil, Ch2 gb Ch3), 06b6b@o@adD6-
t]O 086Oab q:b 0606t>Gogugu6 8Oq]OjugTbtlo
tlgpo 6bvogo gD Ooon3gpDfo@bTao 88Oabd8 ubD6O
Otm8drpblj. bd3b. @t>bortt>6 bbcoGo6oaugpob P2-8oj6coagoatlgo-
I bb8D bbd006Ooao uey:6O"btj 8b6Oot 86b8g-
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3Dcooao6t6o Gogoaolj VD6@rpd86oo7 oV808b. bboo7o
abd086ooao Rb @borto 6fodbojt6o 36ngtldoobo ugpo6Bo a?obugT8og@6 2>8oab 3b8"boot.
Ci,I gb Ci2 grp@86o Ch-b3o6bgobb gb P-Eo8oao-
flB€dogEo o6grpooocofoob afoooi b8"ab, 6ob tlo- b@b6 tlog88ab (btl6boto 7.11). Ci,I 8b ci2 gco8060
98abgbG 8Ofoodugbt]O b88n868Oab.
06q3OdGobfoo b"j8botl60 68b806Oaob078Ob 06o7@b6oonlj obo fljb8raodg8bb, 6rp8 Ch-tijo6Dgoao
p-BO@OaobDaL`6 Tlo@gbb6 8bj06cbo Ovb8b8Obb. tlo-
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3bonrr)ao6o bg8coDBo6rpb gb, t>@bbo)b6b80, tl%6ogr@o 8O8bg, b306Dgoatj Taco6Ob vDf6@rpoj@6Obb gb6O,
608gob a8869Obb b306bgoao vb68Ot>8806b,
8obo bp8ogp898bb6ocoab (Goooforpgtlob ob otu
93bj06b jo - p-BO8Oao. MHc I jgbijob @Ogoju-
80foojt)gbuo) bbb6j8orplj. T-go@3coGo6o b@blJ
gpbtj D60oao6u6o 3830oxpo bvrpdog D@ gb6t]o
MHC @rigojflg8aob q?bti8b6oboot t>tio6boblj. bbd@o
tjb3i]06goab. gib6Ob b06fl5ouf4b agpt8O b6
Od6b gD8bboti6Oa@q?0. Ob
3rpgpo8n6g}ocb8o, u3o6D0obbg,
303Oogotj gb8bjD8u068a8gpo
8t>6Ob 8bj06bb 8b a,306"a-
Cbo @oon8boaflgp b8o6n88b88ab

t>bbbot>07Obb. i}88Oat>g, I jgTb-


bob aD6bb8b8Oau8 b808Oab
ab6t)t]8D8Obbgpo 303Oogob -
tlo@bjDBE06Oaogo bjoGO8O-
dt)6rpbb Dd3b.
HLAI dgDbolj a06Oboti
t)t>8o 866btl8680btgo grr)dUt}o
b6boacoatj gb or>oorioougo @bco-
ab6o gogo 3ogpo@rp68ofo@oo7
bbbob07"bD. D8 gpojtboaob
@obog8oo) jooo6rr)fooarp6U6
bt6boio 7.io. b8b@Ob6ch MHc a06Obch doac8Odlio. MHc d8Ogb 06go8Ogb Od8t!O ab6tjb8b8Oat)go 303Oogob
@06oaotj jr.@3godtio o@ b6ooao6oaob @bjcogodoaogp -EO8bjb8tlofSoaogo gD6O Dd8b. HLAI jgt>boti
ao6obb 8rioob8b, 6n@ogon u@6b8gToboab .3bljobob@ao- t|rr)30g @rlgodt|gbb 86b8bgo 3036ogob gbjb-
bogob bgrr)OribEilj`3g``60b6ob 8coGogo2>bcb8. ob b60oao6-
8t]o6oaD flotdgpoD. drp8ogp o6go8ogtlo bboo7o
obo gb @bcoo ao6obo ljb8 8o6ocrtbq? dgbljbg otinBt>: I, 11
od8tlo @rrtgojflgotj D6boarpab @6ot]86ogpco86DR
8b Ill. I 8b 11 jgTbbob b6Oob06Oao Odlj36Obodoa8gob
q]6gob T-u*608Obot8Ob vb6bb8a060 303Oogobob
@%6ogoaobD gb dbrp3ogobotj foogt,3o6Cbo, Ill jgbtioti
b60o806oao jo tl6bobb gb I)ti3b dbrp8ogTrp8D6 bocoboobEo 6bo"6Oabb.
8®G06jug06O Oogpobob (8ba. c4, c2, 8 g}Ddocodo, TNF). T t*6ogotj 6oGo36o6oMHC I jg6tiob
Ill jgbbob L`66Oe,06Obo 06b6b3gb6Obottb @cOGogobb@O @cogojtgpbbD 8D 8Ob 8b6EO 8cootb8boaug 303-
D6 8ro6bvogonao6. ,`gb@ob6ob MHC I t}gpbbotj bo6oao Oogb o6ot @o7gob6coat>g bgod8b8b gb n6o8ob
bb8() 8o)b8b6o (B,C gb A) gb, bbo80, @o6rn6UgTo gcojtlljo-
o6o796rDuqpbg ujb8t}o6goab. TCR-tj Gbgj8
aolj-8b6 E08aoab. MHc 11 jgbbob a0602>O bflg 8G06O b6(3oao6tdo 3036ogobbg8o Dg3o6rrtab ljDo6oirpg
bt>@ @P, DQ R6 DR) grnt}t]bb EooG68b. MHC bo6oao
b6 abb86ot>, ticogrrt bb8b 3030o"aolj Vb6@gao6o
ab8o66oago t3rpfTo8n68ofo@oco t,bbobot"ab. MHC-tjbg8o dbgoD6 gbabgo b8o6rrtbD bd8b. ob
g306rr)@06o - T-go8g}rpGo0ot) @ood b30Go8ojtl6o
MHc-007 vt>6"06OO lj38GO8Od@6O b6ooa06Ob tloG-
6frtab, `MHC Uofogtig8olj' bbbogocot>b G6coaogo.
D@so6bg, fl%6ogab6oot, o7D8obtl8gbg @drp8o
b60obo6o T-go@q}rnGooob @o86 oa6rp6o6oaugob.
@Obo 8tl6bggoab @07gTob6bg 068OG06Oaugo
tl%6ogoaolj b8rpG6obbfo8b jco6oo666o6oaugo.
G3 grp@06o CD8 8rpgojtlgbb "b3Eo6"aD,
6ob bb8coG MHC I jgblJolJ 8cogodtgob Vb6g-
bo6bono 7.11. MHC I 8gbljob 8ogodt|golj gb6ob 886og 3030ogb 8brr)grog cD8+ 0o0rp0ndbodb6o
b06@do@6b. Chi gb Ci2 80@060 Ci-ij306t,g8aobb gD T-t%dogo G6oab.
I-BO8Oaob@b6 t}Ogbobb. 303Oogotj gD8806O g|b6Ob MHc I jgpbtjob 8cogojugpoao Otootd8Ob d8Ogb
8806goalj a-b306t,goao ,vt>68Od@6Ob, 81jd06ti do aodot8Ob6 u%688foob Odb36Ob06Obflgo. dbco8Og-
b60o3bd``gogur4o 68b f}-Bo8o. q?bljDdD8tlo6obogo 303- @0 @boto Odij36Oboob bb6Obbo ab6bb8b8ObugoD.
Oogob bt,boll 8b6Ob T}1jd06Obb 8D a,306"aob tlob6OcOD 880gpbfoo tb8bg ob gpo@8tl6 dbco8ogfoob Vb6-
Q]og`>`3o6ob 3cogo8rD68ofo@o bb6bDQ)g86D8b.

b6Ob, 8bb aco608Oao 8b %Obooao bd3b. @cogojti-


gTob b8 bgaogpb T}08bjb8tlo6oaogpo gb6o @inding
groove) ovogoat>.
gib6Ob g}bjodob 8b j88gpobob 0@ fla6Oatlo,
bt>8bG 303008Obob tlojb8tl06Oab 8b vt>69806b
vt>6@Ooalj, b@06cO8ab3boib @O@G8OgToab G8bgo-
bbgoD, 6bG bb6bb8D88buqT 3030ogoacoD6 8Don
c}b8Eodb t}fo6U680gt|rp8b.
MHC 8nqpojugToaob u8cob36ob co8obobbtj
bu6bo7o 7.12. MHC 11 8gbbob 8engodbgotl lj6d@J
b06tldou6t)gpo 3ogo@co6q3ofo@o Vb68rpDgao6b.
2002 Vgob o3gobobco8ob %b686coogoaotj @tm8- oBdb. 11 jgbtjoti @rpgTojtgD E08aoab 06078b6007Otb6
t>6bjo8Dgp86006bg gDjb8Eof4oaflg?o 3rpgpo8ri6g3aj¥o
gpoo 06ab6OfobGOob 60806dgpDot)6Ob jo8OOOoob Ci gb a %DB8oaolJab6. rr)6o8o %b83o Go6®3gpDfo@@6
@oo6 HLA-A gcojt)bob 250, HLA-B gojubob @08a6D6bbotD6t>b got)b8@06Oaugpo. 303Oogotj 8b8bjb8-
488 8b HLA-c godobob 118 Dgogotj D6boaoaD tlodobogTo pb6b Pl gb cil gn@06oao d@6ob.
8cO8a06Ogo, 6Djgoab8 g8Odgt]O, cO06j@Ogtlo,
8Ogo8flo. d8Ogbcbo O@6Otl86Ogo 6bco806coa007DD
Ob 0806tjb XPD I)cO6Bt,Oti ju6cooafoo.
MHC I t}gt>tjob 8ogot}@f¥t) NK-t|g€60Rob
KIR 6oGo36fp6ob gTo8b6got>. 8bbo7b6 KIR 6o-

a
G03006Otj 86j68t]06Oat> NK-t]*6Ogotj d0806fl8
8tl6dGobtj oJ6atl6b3b. orJu bDjuovb6 u%6ogfoo
Odb36Ob06Oatlgo I jgpbbotj @fpgojugb dbo@8
8ofoocboo7 EooG3Dgpb (8o6@bob D6
foo@rpd@o"bob i)ogobDg) b6 bbo6oncog abd6b,
t>b007 tl%608tj NK-a%6Ogo abb6bg8ti6Oblj.
bt58b 6bo8o
___
___
MHC 11 jebbol] 8mc:ojocpb
MHC 11 dgt>bob 8ogodugt>, MHC I dg61JotJ bt6bovo 7.14. MHc I 8b n 3gbbob @08Odti8Oaob
8rr)gojugob @ba,b8bbf8, o6o - Ci @do8o q?a P b66Oa86Ob 8b@bdb33Od8a8go €bdoli o08bdoa0070 8bb-
@1jtatido - 3r.gTo3030ogt6o %bB8obab6 t]ogbo- bbob07Oab. I jgbijob @cDgojtgob gb6O 8bbE6tgob
ab. rp6o8o g3bB8o MHC gojflboo) jrpgo68atgo (gb806Ogo 3030080 gD6Ob jogooab t,6 bGog"aD),
11 jfTbbob @Og8jtlgpob gD6Otj 3060 gob. t8ObbaD@Obbg,
06b6b@O@a6b6ugo GoqTbb (bt6bcoo 7.12, 7.13). 11 jgDbob @ngoj@gDb U86n gogo 3030ogob gt>Bo6D
MHC 11 jgbbob @rpgpojtgbt} oo)tio grp@06o i}oudgot>. I t)gatjoti 8ogpot}tigob 8oo6 Rot)t>8tlo6oa@go
Dd81j: foogb-on, Pi gb d8ogb -Ci2, P2. Cii gD Pi 3030ogo bb€obgcog 8-11 b@o6rp@ab8u6 6Dt]onlJ EooGb8b,
gco@06o, I jgbbob @cogojtgoti Chi gb er2 gn@06ob 11 jgbboij 8006 8bjb8E06Obqjgo 3036080 do floodgoab
8tj8D8bD8, D66Oa06-tlo8bdb8E06Oaog gt>6b d@6Ob 30 b@o6rp@ab8tl6 6bi}cob8go gb 880oG ot|cob.
(I,@6t,o,o 7.14).
806b08b6 HLA-DQ gb HLA-Dp Gogoaob
p2 808060 cD4 8Ogot3t8t,b @jt>8E06"at>, co6080 9€bE8o 3cogo@rr)63tlgoi`, tlco80g t>gb8ob6l}
6olj 8b8rpo MHC 11 jgbbob @oo6 Vt>6gao6ogo
rr)o)b-nor)bo ab6bb8b8obugpo HLA-DQ gD HLA-DP
3830ogob ab@cpG6nbb 8bongog CD4+ T-go8g}co(3-
OogTD tloodgobb 3dco68Ob. bbo80, 6b8at,6 @060-
nob t'ot'dgobo,.
8bgt6bR 3fpgpo@063tlgpo Ch 93bB80, raoodgpoab,
HLAII jgbbob Gog"ao bb@O d06OotbRO
gogobb gb 8b@ob d6o8nbo@oaob HLA-DRBI
bbt)oar)t>b Vt>6@cog806ogo (bo6Do)o 7.10): HLA-
R6 HLA-DR83 %6B8oab Eat)Vtl8ogRolJ, t|rrJ809
DR, HLA-DQ gb HLA-DP. HLA-DR gpcojbtjo o6co
o6go3ogtlo, t]obbdgpnb, rr)or)bo 8b61Jb8D3oaogo
@o6o8Dgtl6bg 3ogo@co63@g or %bB81j (DRA) gD
HLA-DR Gogpb otlcob VD6@rpgao6ogo. otoorto8@g
rr)6 3cogo8o68Ugp P %DB8b (DRBl gb DR83)
t>@ Gogttb (HLA-DR, HLA-DQ, HLA-DP) 86t>8t>g
Djcogo60bt), HLA-DQ gb HLA-DP grr)j@b8bo jo
b600806E6 303Oogcob6 8bjb8tl068aob 3c.006Gob-
- 06o7 3ngo@rp68ug Ch (@01jbab@obbg, 22 a?b 20
go Dd3b. b@o0n@, d6ogo VD6@coljDgao6ob 8oj6o-
bgpogTo) gb odor 3ogo8rp68tlg P %bB8b (t}obD-
atgo b6®0806Ob Otiooto 8uot>GOD, 608Ogoo @bb
ab@ObD8, 53 gb loo bgogo). HLA-DR-Ob bb0607cO
HLA 11 jgL`bob @rpgTojtlgpDbonb6 gbdb8fio6oaobab6
ci 93bB8Ob EOvd8OgoaD DRBl %bE3Otb6 HLA-DRBi
ot)b6olj.
Gogt>b Vt>6@od86ob. HLA-DRB1-olj 260-foo @ooo MHC 11 dgbbob @og8jugToao B-go@8nG-
bgogpob G6cobogo.
o6oaq]8, go696ootigp tl%6ogoafoo, @o6ooooob-
lib 8b 8bj6cO8baoacbob Odb36Obodoatgpo. @Doto
8ti6dGoo CD4+ T-go@8oGo6obco8olj t>66oe,06ob
vb68a06t>b.
MHc 11 dgt>tjob 808Od@8Oao tjb86 ti%6O"a-
fooG (03ocoogu6 96 06fgcocoogfl6 ti%6ogobfoo)
f8Oodgoab Odt,36Ob068Ob, 6t>G b6Otobob j06btlo

a,b@n@t)rab3Oaogo GOood06Oaob (8Da. y-06o06-


806rr)gob) a,b8g86oon t.goad. bbocoo u%6ogo
660oe,06V66@Re,06 8t)6dGoalJ od86tj.
03ogo@orpgcoaot6o d8go3oboo7 100-Cbo @06o
Obocoo gbb8D8Oabb 6D6bt50, 6n8gpoaoG bbdj8Ofl-
q:po HLAI b6II dgbbob 8do60 3o6oab, bbo6otn
3co3UgbooDbcob6 @ogt)68boo7, tl86rp btlo6Dg
tl8ocr)D6gobbco. 8bo)o t)@00obobb btlooo8U6t6o
ao6oaobbt>. @D8t>gocoDg, @bb6jognfoobogpo b3n6-
gogoooot (aob00608Ob 8bb8D"aD) gbD3D8OauqTO
cOOcO6dt>6Ot,60 3t>0006Ooaotj 90-95%-b HLA-827
tj06b070 7.13. MIIc 11 a8bhoti @OgacaBch liodafroadb. t)66Oa86O bd8b,I o03Ob i]bd6Ob6O 8Oba00007 gDt>-
bu6bo)fo8 MHC II jgDbob @rpgpojflgolJ U%6o"D6oonb 3bgobflgo cooot6jb6ob6oaoti 30-50% jooo6oQ]o8n-
6bvogo 8D @btiorib6 gDjb8fl06Oaflgpo 303o08Ob 6D8306Oao. Oob HLA-DQ2/DQ8 @ngot}ugoboo7. I)bo6006otmb
MHC 11 dgbbob 8rpgpojtlgD b6Djo8Dgo60fl6Dp gbjD8- ob 86dooG, 6o8 HLA-DQ6 8ogodt)gt>I oo3olj
t]o6obflgo 3ngo@n6g}ugpo a gD P %bB8obaD6 raoq?aoaD.
goDaoooljab6 8bG8Db 8b6b306coaoab.
HLA 806rpoo3o 8t>6j3oago 33jGo6obobsgao 3t+- T-goa3nco®07b6 6b6bb8806O cO©mtimE06O
btbob bD6ObbbbG @b61jbfop36D3ti. 8be,bgocoDg, DR3 b6®o806oaoi, abcgb@oab8oab (36nooi,o68o)
D60oa86ob @drp60 3o6oao ougbg b6 306 3bbubrpao6 MHC I jgbbotj @rpgojtgbbcot>6 bbcoGo6oau-
8 j03booootj fo08b306@gpo b6Ooa06Ob 8bdG06bb.
go 3ojooq?o Go0rr)forr)gpu6o GogobotJ 806@060t|-
MHc I D6 11 jgbijob 8cogod@goao T-gpo@-
go "2,dbq?boooco @OogoaD. vb6@cod@6Ogpo 30300-
8rr)Go0b Gogprp8b6 b66oao60bb Vb6Ug806b. bg6o xpo o6grp3gTbfo8t6 bb"foo bbgbo6b6obb, bbgbo
@oo8608q?606, 6n@ o8U6flf4o 3bbflbo 3cogobbdb6og- bbgpbgvb68od@6oqp MHC I jgbbolJ 8cogojbgbb
@g gb go3ogu6 b60obo6foo T-gb@rp"ogoaogo bjb8T106"aD.
o8co. o)8gog606, 6rr)8 ob b60oao6oao, 8b68on6o- bo6crt3ob5o flng6ogob Goocoforpgti6o
abgo b06fldofl6oaolj V8Dgpoaooi, B-gpo88rpGoob
Gogobob tl@6b898bcoab 068cO806u6O vb6@cO-
T-fl%6Ogob gbt]t,@b6Oa8D8 Dbdooti6Ob8686. nlrr}aoljbb. ooooforpgflo 8rr)t]8og6ogpo flG15co b6-
ggooljot8Ob Obocoo T-gpo@8cOGOO8aoD 6D6Dbo Oo@06o Eoodgpobt> o@ 8o6@t)olJ b6 b%6oxp@ogb
(@b8. rp6@D8bg 6o@boot6o T g¥o88oGoooao, @oj6obolJ 36coq?fldoo o8rpb, 6rr)@OqTOG @%60gfio
NKT go@8r?oooobo), 6rp@goboG D60oao6b Eoo86b gb bbdtlonb6o bbboGOG15go oodgot}co80b
cDi @Ogojugpoaob q?t>b@b6Oboco G6Oa06. cDi buGogob8g Gogoab vb68Od@6Ob.
8ngojt|gpD MHC t}gbbojt|6 @rr)g8c)tgTbb bd MHc I t38t>tioti 808Oda8Oao vt>6t>Re,0606
Vc>68rrtbge,061j. ob MHC ao6oboot D6 jrr7go6goaD. 6coarp6o tlGbco, btio8o bbdtlcob6 b60obo6oab. 1jo8-
CD1 @ogpojflgb, MHC I jgpDbob @ogpojtgTolt t5o86@6 b%6ogEo MHC I jgpbbob @ogpoj@g=obo
8b8b8bt>g, P2-@oj6n8gnbtgpo6cob6 bbrpGo6oatgo 8U0bGooo7 E0G8gpog, bbj@o)D6o @06oboco D6 rp6j-
bb@O q?O@86Otjab6 i]088aD6O 06b6b8O@a6D6tqpo rrt806oaoco jrpgo6oatgp Gogcp8D6 b6®oao6oalJDG
GogDD (bu6Do)o 7.15). Dg`>8ob6ob CD1-b buono vb6D8a0606.
Oq]080686 I)d81j: CDla-CDle.
Chi gb Ou 9rD806Oaobab6 vbd8fDd@6OgT 8b@DjD8-
tl06Oa8g gib6tio cDl 8cogojugb @Oj6coaob ago-
jrr>go3ogo6 jco83o6060b ojb3nlo6oab. Vb68cod@6og
`CD1-go3ogo' jrp@3godblJ T-go88rr)GoOo tlooG6coalJ.

gb"o6gb, 6n8 CDl-ob fl8G6oab ndo381j: jrnnoobbri@D jo36o"ao


Cup 8b y8 TCR-b tlo@dgpob. b@ afooot t}0006rpab
bo6t>oio 7.16. 36®6obljo@b. Go6rrt3gbq]8btlo bf41joatl-
8Ojtytabj006Oob g0308b6O b6OO@06Obo. b6ooao-
go Gogpoao, @Dco tlco6otj uGbnG, boboo@bofl6bq?
6ob MHC-gD@rpojogoaogTo fioG6rpab y8T-@%6o-
goadbgo6gobb 36noobtjco8btlo.
xpoab bb8b Gogoaob got,8b68aootbG fi8tdgob.
t>8 Oogpoab bbgb8bq80806Ogpo MHc I jgpbt,Otj
8bab81jo %b83oao - MIC (MHC class I-related chains) on©mtmcTo6n oocpoaob
bjcogo6obb. gof6836®OcpO Q?Oa6bpboob
Goocofongudo Gogrp-
8b6o t>60o806obo 36rp6o-
bbo8t>Uo go86Dgo6gobb.
36o®oDbrr)@b 8D6ono bjod-
06ob 36rpooogofob6o Dd-
0oU6rr)botj 8drp6o g}06@060-
or)b tjo@3gpodboD9 60@OgT°G

G0003qpbcb@Drao @goab6Ooab
(bfl6t>o)o 7.16).
bgbt>6otl6t>8ob, 6rr)a

@b8D-o6086886rr)60t} 8b8-
gpo6ocr) 36rpoobbrr)@bb o@
bua0607Obgcot> (LMp-2 gb
LMP-7-olj) 66t>6bd6oq}oot>
gb tjo6o7ofoo 8t>Otlgcobb,
drp8g8aoo MHC I jgbbob
@ngojbqpoacob6 gbbbjt>8-
tl06Oaog 303008Obb vb6@cO-
d86Ob. b@ 8fooco ab@b-06006-
306rr)6o b60o806ob Vb6ga8-
6Db c,dg006Oab.
303©OQ!Oaob 8bQ3b©b6b

ti06bcoo 7.15. T go@8OOOooao. cD8+ q3b cD4+T t%6Ogoafoo Odb36Oij06Oau- Go®mSmqpoq?o6 06q?m3qpob-
ao6 abgoto
go 6oGo36rp602jo (TCR) MHC I gb 11 jgbbolj acogojugoaoco Vb6gao6og
b6OO@86u6 303Oogoab 80006Oa06. NKT ux6Ogcbo 8db3681j06OaugTO 6bb- Gooco3gbcb@DEo Vb6@nd@-
086bg o68b6ob60tgo TCR 6oGo30ri6o CDld @cogojflgotj @oo6 Vb6gao6og
6ogo 3030ogo o6grr)3gpbq)@tl6
8gojrpgo3ogfl6 D60oe,06tj raooG6rpbti. aD98cbo b3OGODg¥Ofoobago
65

06b6b3cO6006Ob (8Dgb@Ob6Oti) bDT}tbg:poa007 dub, 6co@ogbbG ooonfongt|6o 3030ogo o6go-


b3goab. bd Ob t>t,gb8vbd@Od@6Og MHc I dgpbhob 3gbq]@t6 bbq?8Eo tlobd81J. o8 Uo8ottJ808Di}o, o7fl
8rpgot}ugDb tljb8tlo6"aD. 3030ogo MHC I jgbbob @cogojtgbb @OjoG8g
Ob b3OGobgoq]Oaogo Gogoao D6Oob06Ob 96@t!t>8tl068Oa6 96 `MHc I-3030080' d0838Oj-
36rpGobo68cnb6 bbcoGo6oaugo 06b6b3n6606ob - Ilo VD6@rpod86oaD, MHC I jgt>bob @rrtgoj@qepb
TAP I gb TAP2-ob (transporter associated with antigen 6b3bb06b ljoogp"aD 8b @9€608Ob fo08b306foo
processing -TAP) bDbogootbD G6naogo. bgbb6o- 8bgb06DG8goab. 07t MHc I jgDljob 8coq:poja-
i}6b8ob, 6rp8 TAP I gb TAP2 bo6oao, 6co@goaoo gbt} jo30ogo b6 got)jb8Bo6gb, bt)gpDgvD68co-
LMP-2 gD LMP-7 a86oaob a806goco 8goabdorpbb, d86ogo bf4btjoDbogu6o go@06o (Ch %bB8o gD
ab8b-o6006806on6oco b6o8t}go6goab. f}2-8oj6rpagrpaago6o) 06gco3gpbfo8u6 ab"foo8o
TAP Gogoao do6ocnbgbg o6grp3gDfo8fl6 goa6b806goaD.
bb8Ofoob gcojbgofoobugo. 303608Oaolj 8b- `MHC I-303©nq}o' jm83cpodbob odb360l}nb

gD6t>6b ooonfocogogD6 06gco3gDfo@u6 abgofoo o3Bf6oQ2ob $3pb3o663


ATP-gb8rpjogoaugo 36coGobob. 3030ogcrtb6 gbjb8Eo6oabgo MHC I jgpbbotj
06q!m3cpobao6 atiq!obo `MHC I-303©oq!n' @Ogojtigb bobaogti6 b06@dou6bb vb6@coj8-
jn83cpodliob o6gmab 6Oij 8b t%6Ogotj fo08b306foo 8db36Otj06goab.
06803gbfo@06 aDgofoo 8Ob8086Ogp 303®Ogb ob Ug36ogob foogb3o6foo brpgT%ob b3b6bob
TAp go@86cOD6 8Djt>8@06Oatigo MHc I jgbbob 8bgbbd8b. fl%6opfo8 odb36obo6obugp jo@3gpodbb
8Ogot)t)8t> Odt>8u06Oab (tjt)66cOO 7.17). cD8+T-u%608Obo i)OOG6Oab. cD8 OtD6b6OG03OO-
MHC I jgpDljob 8rpgojugotj bo6o)ofoo 6o MHC I jgpDbob 8cogojugob b6b3ogo@n6-
gD bv8rpbb 86b8bg o)D6D8o@go86Ug 36o- q3tig tiat>6b t)t}t,8E06"at>, 660 T|i%6O"aob
GoblJ @cooob3b. @ngojt|qpob Ch %bB8o gb P2- @ood b60o806olJ tloG6oaDEo 86ofi86ogo8b6 6rr)gpb
8oj6rp8goaugo6o o6g®jgpbfo@u6 bbgofoo Vb6- Dbdugoat,.
@ood86obt> (bb6bo7o 7.18).
Ch %bB8ob ljbarpg®co lj06udou6ob Bb@rptl- 06Qmoo®mbool o3€6oQ2ao @nb8oe6or?o
b6©oao6ob 8bqib@oab8oab (86moobo6ao)
bgoboabEo @6ong86ogo3b6 6ngb bb6tlgoao6
m>6@bgobo (f)bjo6fp6o) Gogpoao - jbg6odbo6o ti%6ogflo o6grpGoococboo) @cob8og6ogo
gb jbg6o®ojUgpo6o. 'Gb6oogo' MHC I jgpbbotj b6ooe,060 068OGOo@6 8Ofoojflgpbtlo 8086D806-
go806o ob3bcbo6oot ojb8rao6goab TAP jrp@3gpo- gobb. 36rpoongofob6o goa6bgbGooov 8og}8atigo
303OO"ao MHc 11 jgDbob 8cogojtlgbtj b@t>80
86g®3gt>b8fl6o bbgnb g6g
8Ofoodti86tlo tidt,8tl06"at>.
b6®o8066o6@q!ao6 0336oq?oaan o3a6oq}-
aof63cnb oopoanb aol}3ee
MHC 11 jgbtjob 8ogojugpobb ob 3030ogobo
ojb3t}o6"bD, 6co8g8boo b3oGobgofoobtg b60o-
-t`'` ,® @06vbd8f9a06 u%608Oat}O 06Ood6D8Ocboat8O
Goqprp8D6o t>60o8068aotjaD6 @oogoab.
u%6Oq?ab6Ocob Gogco3b6O b6Ooa06Ob 36Ofo06-
obGOob (vb6ga06Ob) bbvtiob Oobjlj b6Ooa06vb68-
#b;:d3.:##gb63£b¥3#o#b#Ov:=#L:q°#3: 8a06O t%608Ob 8006 au68a608O b6Ooa06Oti
06b6ti3cof$Oo6oab 36noonbco@bgb6 bt>8nbogpo 3030ogoao uojt>8t]06Oab gb 060066bgofobGob vb6@ODga06tj.
oooocbngogb6 06grp3gDcb@06 bD"foo abgDDd8co. b6Ooa06vb6@q?@06 @%608b D66O@060 1jb8D-

gbtjb8b afooot ujb8806"ab (btl6bcoo 7.7):


I g86g6o0Ugpo tg€6ogobb gb @bj6rpg3bbob
foogb3o6@go 6ooo36co6oao @od6on68b6ocb@o-
bobco8Ob bb0607cO ij06@dofl6Oab tl0006cobij 8b
OJb8t]o6goat>.
I 8bj6co8b8Cbo odb36otjo6oa@go 68oo30rp6oao
b60oljtiougpob Fc-8dbb@060oalj gb jn@3go8066ob
j0830606Ooati (@ba. c3b) OjD8t}06Oati. b6Ooiji5O-
t)gcob6 b6 jn@3go8060otj GogpoboJb6 gbjD8i}odo-
btg b6Ooe,061j 8bj6cO8bao OcogD8 tl07D607db8tj.
I b6Ooe,06vb6@ga06O u%608Oaob b3OGO8Ojt60

bu6boto 7.18. MHc I d8bljob ap]806@8Olj vbd@®d86b. 6ooo30co6ob jb6ao @b8bgocrtob 8 b%6ogob o8tl-
P2 @od6r.8qTcoat}go6otb6 gbjD8t}o6oaD@go Ch %t>B81j
6oagpcobogo6tl6o doGo30co6o. ob 6oGo30n6o,
jbg6odbo6o ujt>8@o6"ab (1). P2 8oj6obgnaugo6o Ch b660806Obbg@0 8b8bgo b806coaob ab80, D66Oao-
%DB8b Ojb8E068Oab (2). 6D3t>b060, jbg6OoodugT06O, 6Ob 03Odo@6 060066bgpofoboobb u93608ab6OoiD
TApl g6 TAP2 3030oRob Rt>8So6 do@3godbb d86ot! (3). I boovbotlo D60oao6ob gbabgo jrrt6oo606bGoob
jgbbob @ngodogotj gbdt}o ooorp3gt>fo@U6o 3030ogob 306Oaoafioo Ov808b,
gbjb8tl06OaD bobbogfld jco@3gTodbb (MHc+383Oogo) 060066bgofooatgo Oogo8b6O b6ooa06O
Vb68rpd86ob (4).
t%6ogtlogb 808a6b6ug 8ofoojugbtlo - 06gotjn-
@btlo b8"aD. 06gotjco8b 3dcooongofot)do 806886- 1jbtlabg8aoot t58Oab. 068rp3gpbfo8u6 bb"foo
ooaotj tlo8o8ogo, Rt>bt>gpo pH-otj 8do60 8ofoodt]- 8Ot,3086Ogpo 303008Oao MHc 11 jgbbob 8coqTojo-
gpbb. 06gr.brp8D gocbcpbco@Db ojb8tlof4goaD. MHC 8t>b 806 t)dt>8E06poa6, 6t>Re,66 8Obo Rt,60
11 jgbbob 8cogojtgoaoot 8gogb6o o6grpbrp@obo o68b6ob60tlgo %bB8ooJ (Ii) b6ob gD8t)6@go. Ii
t>6ooe,06Otj a,6Rt>8tiEt>8Oat>1j6 96 v66R8066Tao 8t>6- Dbo80 8co6bvogorpab MHC 11 jgbbob @coq:pojtgob
bbjuco6obug 6cogpb blJ6bg8bo6 (bU6bono 7.19). bvdrpaDfio 8D tj3Ooobgofoobug 06gcobco8bT}O ab-
06pcobrp@U6 gD goqJobco@o6 8ofoojugpb- gbot>6DEo, bbgbG o60066bgofoobogo Gogobo
tlo 0660666gofooat]go Gogoaob 36OGob06e,0 303Oogoab8 8Ob6D8068Oab.
(8t>R68@E68Oa6). 8ofooj@g6t), 6o8og?bt>o MHC 11 jgt>tiotj
o6oo66bgofooat)go Gogobo o6grpbco8u6 8OgpodtR6 ti*608Ob cb086306Otjd06 a68t>t>d8b,
8b gofoobco@u6 88foodtIpoatlo 806@066tlgp a,fobt]O aD8b@ttlb8Oaugo b6Ooa06Oaob (806@06-
R086t)pt>Got)tj 8t>6008Ob. Gogoti 8Oa6t>86OOOOT 6@gpo xpoe,6t>Rt>ooob tlogo8t>g) t]08o8ogo 8ofoo-
808Oatigpo 303OOR8ao, MHc 11 j8t>tjob 8Ogodo-
jtigoao tidt>8ra068Oa6.
gbbcob6 gDjb8t]o6obob t}08go8, CD4 T q}Po@8oG- @bj608b8OabD 8b Dq:b8OD6Ob 8 u9€d08Obtlo
Oooatj vt>6flxp8Oa6. MHC 11 jgDbolj @cogojtgoboo) @gogD6 86grp-
b6ooa06u6O Gogoao 8bbt>go pH-Ob 306naoa- bco8Oab NIc 11 jgpt>bob jco83b6O@06Otj b6S MIIc
EO 36Ooobfooaoco Poe,f4Dg06q?02>b. 3036Ogoaob (MHC class Il compartment) uvrpgoao6 (cob83olJ a
Vb68rr)d@6bEo 86ot]860grl8b6 6ogpb 8868060o t9€68"at]o o8o8o n6ab6ogpbb Il dgbbob 8ocbojt-
jboo31jo6o btj6ag8ab. ob o6gobrp8bEo t]8ogbcbo gob - CIIV-b Uvogobo6). ob 880gpb o8 DUGogpobogp
gogo 6brpgo6oaoonDb. jrr)@3r')6866b tlooGb81j, 6bG MIIC II jgpDbob 8ogojt-
ti%6oRfoo odb36obo6oat]gpo 8o@a6t>6t]go gpbb07D6 303OORob gDjt>8T]06Oabb uQ]6t68Ogpdco8b.
Gogpb, EobbdgpoaogoD, u%6Ogtlo t,Ogp8OcO608, MHC 11 jcpbl]ol] amcp3joeobo]b6 aoq?a-
o8D8o bfooo) 8cob8golJ. D8go6bg, 8o6UboG jo, 6o- aoae3Oaoeo 303®OQ!Oaol> pbjo8aofi3ab
80goG o6BOGo6oaagpo t%6oq?ob ooorrt3g?bfo8buo MHC 11 jgDbob 8coqTojtgbb MIIC-Eo o68t>6o-
tt8poaD, @08a6b6ugpo Gogoaob Vb6@rpd86ob tlo8go@, D60flgpo %b83o bGogq?oat, 6bG 36ooorpgpofot6o
tlobt>dgcob, oboot 303oogoabg "a6bq?o6"tj, 6rp- 806806OobobD 8D HLA-DM 8Ogod€gob 06Otcoa-
8ogpo)D 36ocbo60boobbbG NIIC 11 jgbbob @cogojo-
go3o foo8od8ogoaob @oxpoaot>.
8Oao t>b060008Obtj. 806t]ti@go 66Ooa06Obt>R@O 36rrtooogofou6o 806806002)o (jD603bo6oao),
b3oGo8odH6o T-3ogp306@go t]%6oRoaot! at>t>doo- 6cO@OgovD ljDt}ubgpoaootbG 060066bgpofooatgo
t)6oab, i]obbdgpnb, t>8 8odb6ofo@oot bgobb. Gogoaogb6 303Oogobo @Oogoab, 063t>6Ob6ougpo
MHc ii jq?bi,oii 8nc:ojoEoi, aonito6o)Obn q2b %t>B8ob go866Rt>oot>b oV808b, 6ob tlo8go86G
83t,OjoEpbao 8bebbQ8O=Oab gb6t}o 8bcogng a.V. II jgpbbcob6 Dbooo6oauqepo
o68b6ob66Ugo %bB8ob 3ojoogo -CLIP (class
MHC 11 dgt>boti 8ogodflgob Ci g6 f} %t>B8o II-associated invariant chain peptide) 6Boab. MHC 11
06R03gt>cb8t}6 a68Ofoo vt>6@Ood@6Oao xp6 jgbbob @rpgojt)gpob gb6tlo CLIP-olj gboco8oab
06o}8b6ootlj bd3o tjb3tlo6q?obb (bu6bo)o 7.19). Eobdgoboglj bgoti 8bbortb6 b66oao6t6o 3030ogolj
bo6@dou6ugbg b6b@8a6D8O 80806Ob tio@8.
8t>tjt,8tl06Oa6b. g66ORt>6 cLlp-Ob @0008Oa6ti
ao@o t>VtioaD Bb3odn6obob (@b8.jbg68dbo6olj) HLA-DM 8cogojugpD Uq)6t|68ogUot}1J.

CJ HLA-DM-I) MHC 11 jgpbljot @rr)qpoj@-


cj u9€6a8Olj q]Oxpb3060 gob 8bat>8bo b66t]dou66 t>d8b. t>@
@jb6bljj68gob8b6 @b61jb8b8Oa007, HLA-
DM 3rpgpo8o63tgo D6 D6otj, o68b6o-
b6Oogp %bB8b b6 Ojb8E068OaD gD
@%6ogob fooR63odcbo 66 odb3601jo6-
goad. 8Obo q3@6dGob MHc 11 dgTDbob
@rpg"jugob g}bdogb6 CLIP-ob @co-
GogToabb. cLlp-Ob @OGogoaob tlo8xpoa
MHC 11 jgt>bob 8ogojtgpb 3830ogb
uJt,8t]o6goat).
o6®o8066bf68q?806o o33f6oq?ol]
boq!b3n6to `MHC II-303©oq}n' jm83-
cpodl]oL} odl]36oboo
303OogoiD6 8bjb8t}06Oaob tlo@"@
bu6D070 7.ig. MHc 11 a8bbo1! @cogacBgch 8b8b8tlodoab 3036OgoDb6. MHC 11 jgDbolj @rr7gojtgolj bobaogu-
06803gt,fo8a6 at>"foo v668Od86OBO R6 068t>6Ot>66Og %698cot>6 6o jco@3godbo u%6ogob foogb3o6objo6
gbdb8Bo60bugo H dgTbtjob @rDgodtlgb o6grpbn8otjjo6 @oo@t>6coobb
8D8bbge,Ogpg8ab. u%609Otj fo08D306fo8
(I). o68b6ob60UgTo %bB8ob "86bgDGoob tlo@gob MHC 8cog8jtgob
gb6o gbBb6ugob CLIP-oov (class II-associated invariant chain peptide) odlj3dot}o6obugp jo83godtjb - `MHC
II-3036ogo' -CD4+ T flg36ogo tlooG-
(2). MHc Il tjRt>boti @cO8Odugotj BO8G8Ogo 8Ocbod@g6 t>66Oa06Ob
t}08G8og 8ocbojtlgbb o6Vtl@oab (3). gb6oq?t>6 CLIP-olj 8rpGogoaolj 6coab, 6bsoG @6o$868grp8b6 6ngplj CD8
@o@"a 3030ogo 11 jgDbob @rDgpojflgbb "b8tlo6"bb. o7b6b68Go30o6o blJ6Ugobb.
67

@836b@8bb: @cO68ugo %bB8Ob007bb vb68cO8a06Ogo. D@


• T a%688Ob abt>doou6Obob bt>vdobo Oob3Ob 8Ogpodt]goalj b06t]joa6ti8O 8tj8t>8tjoa6
b6Oob06vb6@8a86 0%608foo Odb3601j06Obu- bd8b: 3030ogotb6 tlo8bjb8tlo6obogo gb6o

go `MHC-3030ogo' jco83gpodbob lj3ooo8o-


a-ti306bgoaobD gb b6003D6bIpogu6O a

jtl6O EOG6cobb q3D @Ogobrogpo bob6bgolj Bo8oaob8t>6 BORaoat,, Ig-@t>8866o go80602>o


a06Ot801jj06 8bob6Oab. bg6Ong6tigo 36coGobo jo T tl9€6ogob cob6b6oG830o6obb - CD4 gb
T o9€6ogob Cbogb3o6foo odb3dobo6obogo cD8-tj tjb8tl068Oab.
@rpgojflgoaoco bo6oo8ggoaD. • MHC I jgbbob 8r.gpojtlgD ti8ogb bo6o)8ob6
• T a%68q?ob D60oao6rao@o6obo 6ooo3orp6o - og36oq?food odlj3601jo6obtlgpo. MHC 11 jgpbbotj
TCR, 8Otiog83Ogti6O a@8a007 gDjb8tl06Obtigo 8oqpojogpob odlj36oboD, do6ocobgDg, b60o@06-
060 - Ci fpb a - 3cof¥0303Oogfo6O %t>83Obab6 VD6@g806 b%6og8bqJo (go6fp6o0bgo ti%6o-
Eogaoat>. ob *t>B8obo o8@6o8gcoaugo6ob goao, @bj6cO3baobo, 8 gTO@8OOOoobo) ti9Oab.
8rpgojtgob 8do8o gb @btbudo %bB8oaob jco- • CD4 gb CD8 8cogojt)gpo2>o 8VoB0 T gpo@80G-

@Ogcoaotl6Ob. t|n8Ogpo @bot8b60 8b6Obaog¥flf4 Ooobfoo Odb368bodoaugo cob6b6OG03006O-


bob, 608goboo, uoijbab8Otjb8, MHc 11 8b I
96 t}o6ttot>60t)6 tJo88066b i)0oGt>8tj. 8t>60t>-
aogt6 tjoa886Ot]O tjb80 303068b6Obaogb6O jgbljob @rpggojtlgobolJ b6b3cogo@co68Ug
uat>6o aD8coodco8b. 01j doG830rp6ob ob 6DV- t]bt>6b @jt>8E068Ob6. cD4 808Od@86 T aog3-
ogob, 6co@ogoG `MHC-3030ogo' jrp83g8dbb 06foo, CD8 do T Goocoocodbodt)6 t)g€6oRfoo
Ojb8@068OaD. odb36obo6gobt>. TCR-ob 8oo6 MHC-3036ogob
• y8TCR30086cogo806o Tu%6ogo}b @Go60 6D- tloG6rpaob i}o@ovb888bi)o, MHC @ogoju-

rpgo6coabcbob (oap 608D0ofl6 T u%6o"afoo) gt>tjot66 gt,d68E06Oafl8O cD4 96 cD8


@ogodflgo T t]%68REo a,6b6ad6ot6oaog
Odb36Otj06Oaflgo. Ob bb6bb8b88bugpo tjbbob
tjo@6bgb @fob86ob.
t6ooao6ti Eooo6coab. @ob 8oo6 i}oG6coaogo
b6Ooa06O (@bco ti06Ob g03O"ao 8D bb3b 3b-
df¥o6od@do 38@ovt]808obo
Ot>6b 8cogojbgobo) NIlc-01j 81je,D8b, b6t>3cogo- libd8b6eo docob8Oboov
8n68tiqT 8cO8Ojtig?Oa07b6Db 8Djb8tl06Oat}gpo.
• @bgbgo 3®go8co68ofo@oo) bb8rp6Boflgpo job- 2 Vgotl boBIJ ga@g8o696 8do8o drp8bo6o6o-
Orptloo)b8boaob @o)b8b6o jn83godljob (MHC) atigpo O8u6ORoq}OOOOO. d8gpotj 0806tlo fl%6O-

a06Obo, I 8D 11 jgDbob b6Ooa06Obob @b6gb, "aob tlo8G8Ogcoab 606@bgptl6Ob. 6b8Oob-


bt)3b oogobbbG Djcogo6oblj. MHC 8cogot}u- d0006O j890800, 8b98068b, 6cr,a O,08ubt,0
i]08D3Dgo T @x6Ogobot v06D@cO6a08cOD 6b-
q=poao, on8q?D3o68ogDg, dbo8ogo7b 06D6b-
O"6Oa60 606@68ti6Ot>, cO@8G6 1jobbglj6 R6
3gb60booob odb306o@066oaEo bp@rpbBo6ob
306og}06otg go@q!fl6 o6bb6coobflo 8Vo8o T
doa06O ®db6h3gb6Ob®Ob bt>v06bbp@8030
go@g}oGo08bo bbo6onrpg 806 o6bbb. coo@Ubu6
08b6060 3bbubolj bg@d86Ogo D66Oa86Oao. 09€6ogoafoo CD3 t>6 odb368bo6gobb. 6obo
MHc I 8b 11 dgbijob 8Ogojtigoao 303- ti6b6O t>6 bd8b bboco ti3€608Oab7
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3030oq?b
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b60obo6ti ooocoo®dbojt6o T go@8oGoOo
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8t>t!t>8@06Oatigb t!O T aog306O (cD4+). gorp0ogog 66b61j806bfob
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0)b3n VI I I. b6Bna36nl-33i)6ma)@n 1. t>660806OT66 tj3OGO8Otja6O 8t,tj68@06Oa6,
am6b6nEo bBfiodBofioan. 2. t>66Oa06cO66 xp6d68tl06Oaob tl088Oa
bGBnlil]O]E:Oan E® @dmn fioi]OBBmfioan 36Oi0806Ob 868666xpat6Oaogpo t)x608Oaob6 96
8Ogpodugoaob 68Od808Oa6.
b6®Ol,boor?Oao t>8 8@6jGOoab 666Obbot]gpob 060,
b66tid6t)6tg6p 8661)b8a8oat)go, 65Vogo
b3OGO8Odti6O O8ti6ti60 3t>b@t,0 aGbo 6660-
6b06GOogpoab (ba66070 8.2).
a06Ob 68006OaooT Ovdoat>, 66G 8bogpoxp T 86 t>600806cO66 R6b6j68E06Oa08O tiat>6O t,66O-
8 8O88OGOob i]O@d8Ot. 66Ooa06Ob @Otj6G6O-
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a6p O8t]6ti6O tjotj0086 tjt>8O 003Ob 8Ogpod@g6b
V668m6xp806b. ob poxpo @6686g806o86oaoco ti6tjo-
Ot]06Oab. Ob 808Odu8Oa06: 66Oobbougoao,
607Roa6 xp6 8t>606aog@6O (G86goa6xpo) t)a6Ob t>6@
3oboot}oo768boaob 8o7t>8t>6o drp83godt}o g6 T
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b06t]d6t)666 t}otjv68gogo.
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8tl06goa6 608060 O8ti6ti60 3t>bt)t>Ob t!6v8Otj, 660obbougpob 8ogpodt|gob ob 6t>Vogpo, 6n-
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6booo6oa@f¥o 666obbot)go - BCR V668n6gao6b
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606d006 tlood8Oa5 Otiob: d083gp08066Ooi at>6-
8 @K6Osoao 66ootjt,Otig076 tjot36OG06b t>tis0606.
3o6oaoatgo qpofoobo, @%6ogtif¥o Gooooodbo-
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8oo6o7 t]08cob8086Eo) 6gpo68o6.
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666otjbot)gpob 8ogTodUgp6co6 86686g-
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tt8d80or)t>t>, 386o806o6foo Vt>68oJ86ogo Treg
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t,bocoo T-0%68Roaob 8t>8t>6666 866t!t>q]886@8O at)6oa6o8t>g Vt>68od86ogo CD4+CD25+ Treg
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ogo60u6 b3oGoq}ojtl6cobblj t>6 1jbBo6rpoab.
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aoba g6 b6d8oaob t>6oobo6oafoo 6ogpo666ooaob
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u%688Oaob 6bog06Obb 8b6t}gcoab, 6DO b6OO-
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a06-b3OGO8Ojti6O cD4+ 8b cD8+ T t%6O"aob
ti%doRoao 86t>8g8Ob6 80 3t>cooa06@6 T @%6088ab
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jo 6cog86b60rpaob aD68ootD68bbtj oV8081J. q896b@8ab:


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NKT u9€6O"ao @b6ObtlfT068aogo u%6O-
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tfof4a68Oqptlog3b. b8"6b8, O@u6@60 3bbtit,Ob ti6Oaob073Ob buoogobogo 3068Ogpo b086t>gob.
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g}Ddon6oao @bngpcog 6co8ogpo@o d®6j600ugpo OoqTO8b6 b6o0806Oib u@Ooobcoabb 06080 -
b66o@06ob bbvo6bbg8go8rp 3bbflttcbo 8cod88"ab. T- gb B-gpo8g}coGoOo ab8cooG6rr]ab. @oli bb3bbub-
b6bb3OGO8Oju6O do, tl8OIpbb 07u b6b, a036O rp8, tlobbab8Obb8, a%6Ogugo gD 3u8cO6ugpo
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dbgob6 b86Otig bbbjtlo 06908080 a086 b66o@06ob gobs.
b60oao6foo 3bbt)blj 806 D3oonD6oalj. 6bdo8b, %od b60oao6ob rp30o@bgt6o gnfob o30o8bgu6o
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bob @b@co, fonaoodovo b6©obo6olj bbvo6DDg}@po8og o)u b60oao6ob grpcbb dDgob6 gogob, Sob 8o@b6on
@tmgrpg IgM b60obbotlgpob VD68nd86D tlo@dgob. 6coqT06b6Ooab 3Ootb68OaD.
6b8ng3o IgG 3bbobb, do6ocobgbg, 806 b3ocob6oatj. b6Ooa0601j ®6ab6Oq]@BO tlo88b601j aSb.
ao606bgocbobflgo o8tl6ndo8300066b6coab rpdbb6ocb8tlo bo6oao6tlgo, 306n6bgpfl6o
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8Dgoagobb. bvn6og 8b 8bd6co6o ab6D3o6rpaobb tloodgobb Ongpo6b60oaD bb8nov8oob. jb6tlo,
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8OR6Ob tl088Oa 060 ti6bolj 6Ot>dG06 tloodg8a6 (bt)6t>o7o 12.2.).
ab68007D698b: Thl bflb3co3uqp6Gootj @ol68t>6o Gooodo6obo6
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o8@6t6o bobo8@ob 8oo6 bboo7o b6Bo8D6ob acoba t}fo6t)68oguob: IL-12, IFNy, IL-18, IL-23, IL-27.
a,6docooa6ti goRt>8 866u>fo88668ti 08 ti9€6O"aob @obo 8o6oo7bgo q}fl6dGob I)%6ogBogD 3botrpao6otj
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IFN-g
TNF-a

IL-4
IL-5
IL-13

IL-17

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ax6ogflog6 36o7oao6oaolj8a6 o)t>8Rt>Gat>, t>t}Ono8fl6ti6o
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6t)Ocoo8t)6t)6o 36coGoboao, Treg - at)0">a6oboot}8a6
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3 a88boobu8bao dobnb bbvo6bbg@go@m


(bevacizumab) t,6OttD6aocoa06860 cO6ab6b8b VEGF-b
a``b8b@Ob6Obago 8j866bgobb
5 8bogobt}@bbo(daclizumab)bL`bpt>8OL`6Oaogo
oto6j@gob 06b6b3g``60D0oti abt>do006Ob@g T @9€688Obbo agcojb3b
8VBD80 8rr)Gogobb H,-2 doGojon61l

6 ao@outtfl@Dao bb@Ofo6O - gpooc!08Oa6 b%6Ogoafoo


8V8b80 8oognog86og8odo@oob60ottRO80
(gemtuzumab) odb`36obo6oaogo foogb3odogo cD33
abbg.`8Ob6Oatigpo I)60,,ao6o
7 bg086Oqitl@bao(alemtuzumab)abt>xpb8OD6Obogo
b``@Ofo6O - T 8b 8 gpo@q}cOOOO8afoo
8 U%60gugo goojo@ob
odb.3dobo6ob8go CD52 ``6ooao6tt
8 6oO@dbo@t,bo bb8Ofo6O - 8 go@8OOOO8afoo

(rituximab) b6ulon%do6ob gTo8BO@b odtj36obo6obogo CD20. oV808b


do@06bgo b3co`30rr)fo IJ, CMC, ADCC
9 8bgo8OfoD8bao 6Ob`306bGoag-b06GOOOu60
(palivizumab) 3o68b8go (RSV) o6q}odGo`` cO6a@6b81i Rsv-01i F-Gogbb
abb8b@ttb6Oaago bb8B88atlo
10 Odbtjoacbu@bao ab6t]8OofTO bat,Ob dadub
(trastuzumab) jttanb libvo6bt>g@xp8am bb@ofo6o -HER2/neu (erb82) 6oGo30co6{t
abb8t,@OD6Oaogt, @d866``goab
11
00b60608300(etanercept)t}06V880goGogpb
608@``O®OgUgo b6o7dttoo bb6``p(t TNF 6o(3030m6o

12 bgbgo88@bbo 608@booogugpo t)6o)6o0o,


(adalimumab) j6rp6ob gbD8b"bb, 07da06``8b TNF-ct-b
abb8b@Ob6Oaago VdRt)gpm8t>6o dogooo
13 608Ootlq>O@bao(nimotuzumab)abb8b@Ob6Ob8gpo
bd3D8nfoo6o b%60|otlgpoj``60o6n@b,agorr)@``
on6ao6D8b EGFR-b

15 bgo8©tiq]ti8bbo d6o6odUgo go8g3rpoo0Uf$o


bb8ofo6o CD52, oV808b D3en30rr)fob, CMC,
(alemtuzumab) goodo8ob
abbjpb8OD6Oat,go ADCC
16 b6ulm9€t}o6ob gTo@8co@b
Onljoofl8®8bao(tositumomab)ou`a3,,b
bb@ofo6o CD20, ADCC, oV808b b3®`30cofob

17 0868dbo@bao dogrp60doog(), o}b8ob gb


(cetuximab)do@06ogo jolj6ob bo@bo868 ADcc, Ot6ati6b3b EGFR `ioa6bgTb
17 bo8bGobfl@bbo q}Ogo8ob, coo6d@gob,
(bevacizumab) c}ngrp60doUgo. Crl``8olJ codau6D8b VEGF boa6Dgb
@D``g``@oD6oaogo 08o6ob Rb d@dub dobco.
18 3b6o638tl8bbo®anitumumab)abb8``8Ob6Ob8go

d®gco6odougpo t}oam bocoobodobb EGFR ljoa6bgb

19 Gb688bdljo@bao(catumaxomab)
b8o)8oljobob6o bbGooolj adn60 ADCC, T-o%6og@go gofoolio,
3bG006Ooao EpcAM + g}DanGo©cofoo FeyR-ob 8dm60
do886t)go-o)ba8ot}3oa6,,go tjo@bo86ob g6rplj ag`6oq?obot, @oo6

21 08b©O@38bao d6co6oj@go fETo@8nooo@60 ADCC, CMC


(ofatumumab)@bbgb@ot)6ob@gpo gooj88ob

22 806Oq>3abao at]dt|b 8b 3f40ljoDOob ``Oeeoa06Oab RANKL tjoa6bgb


b,,@bo860
(denosumab)abDgb8Ob6Oaogo

23 o8ogo@8@bbo(ipi]imumab)aD``xp``@Ob6Ob8go @ogpb6on@b t}b@ofo6o CTLA4

24 3060tlq]u8bbo®ertuzumab)@bL`gb8ot>6obogpo dtld8b joarrt


boeeob06Ob`j RER2 boa6bgpb

25 soy.oa6o6ti@Tri8t>aooD"dboob6o(ibritumomabtiuxetan)ont`a3ob ``6tx3n%jo6ot) gpo88n@b

6``gorpofon®co3ttb (9°tt06oo8o) @ovrr)goa``

26 131-I 6obooti@on@bbo(tositumomab)a,ba8ol, D6`ulo%do6ob go@8n@b

6``gtmttfomorp3ttb (]3[-orpgo) 8ovngoaD

27 b606bodlioabbo Err)%jo6ob gpo@8n@b>

30gen©o6o b®o3086o
000onoendtjojflfSo 3603b6boob
Orentuximab vedotin)d,}@06@gTo ao8``6®@6@9€6ogrp8``6o8o88rr)8``
(btl6obob6o6 E) @oVcogobD

ADCC - b60obbo@ggb@rpjogobogo GoOoomdboju6oab

CMC - jco@jgo8060gD@njo"atlgo ooocoondljodo6mab


gb6bdcoo 1
cD 8b6j06Oaolj 8db3doljob 8b 8@6dGob

N CD .1,1b36,]bob qm6`i(]obMHcit]8btjob8bab81io8cogoduc=pb,
12 CDlaCDlb doddolj oio8coooooao,gb6aodib6iiotj

CDlc gt>t}b8@06Oatl8O p2-


I,9€6ogoao, 8ojdmag®a@go6onb6. Vbdbgao6bb6Ooa06b
CDld 88696o®3gou3€6o"ao'ti9€68goao(cDic),6bvgb30b.|3oov.|gofl@o(CDld)

CD2qFA-2) T-Ug€do"bo, bq}3ofooti6o 8ongodogb, 6®@8800


cOO@cOGOooao, NK Odb8E06Oali cD58-ij (LFA-3). uoodgob T-
fl966t|@t|bo tl9€608Oboli abb ®Otl6Oab
34 CD3(T3)CD4 07O@OGO®Oao, T-0966o"ao
8t>t}t>8ta06Oaogob T-t|9€608Og b66Ob06-b30oo8oju66oGo30odcob6.

806bvogo®ali TCR-Oti 8006 boa,6bg=gob


abf9b(i,i@btlo.
ono8®o 6oaob tib3co3tlgbo b,jog3068ao(3060806Oafpo T-@96 8 Oaobgb bgo aocoOdo8ot,b@ogo),@®6OG06Oaogb8D.]6cO9iba,iao MHC 11 t}gbljob 8mgod@gotj
IT4) Ovb6b60003O®60. Odb8u0688ab Ick-b.
HIV-I 8b HIV-2 gpl20-ob 6oGo30co6o.

5 CD5 o7o8moo®oao, T-a96d08Oao,8.ug€6ogoaobbfla3®3flgbt3Ob "b8u068Oab cD72-ij

6 CD7 3g@do3noo60tldoH}06®8b6Otl9€d08Oao,07O@®G06Oao,T-fl966,1gtfoo
OG6®aob. T-u966Ogoaob gb
3gtido3o0066u6o gotSo8b6o
u%608Oaob 8bddot60

7891011 CD8 ovo8eno o aob t}a3m3@gbo b,Go©®©mdbojtl6o T-0%6o"ao(306o806o goT-u%6898aob8b bgco ao no6otoa,1t'b@,leo) MHcl t}gbbol} @ogojogTolJ
ovb6b6oGo36m6o. ot}t>8flo6goab Ick-b

CD16 6oo66oq}ogoao, NK- 8babgb80606O Fc 6OG036Odolj


(FcyRHI)CD|9CD20CD21(CR2) 0%6ogoao'frod6®8t>aoao t}®@30606OO, FeyRIII, ab6b306®aoab
8baoG06®fobb 8b b6©Obt,Otlg.
q?b@oJo"aag U%6ogtlg
t3Ot9®fl®.itjon6Oablj
B-ti966O"ao d86olj don@3godbb CD21-l}b (CR2) gb
CD81-ortb6 (TAPA-1); 8 u%do"aoli
onb6b6,1(3.13(bo6o
8 0%6ogoao 8 u96608Oboti abbd©006Oabtio Bobbd8®6®go

8Vo8o 8 o3€6o"ao, d®83go8066otl dco@3m6060ob C3d- ob


80696o0Ogoti96688Oao 60GO30ondo, CDl9-llb gb CD81-or)b6
fldcobg 8 fl936,|gol} o}b6bd.1(3,13(qodob
12 CD22 8vO8O 8 393608Oao B-ug66o"aob bq?3ocbob 8m6®oo©oaonb6, T
'.
13 CD23 (Fee-RII) @vO8O 8 ti9€do"ao, IgE-8bat>gb806to60 600036cO60.
abbdo006Oau8O CD19.CD21.CD81 otb6b6ooo36®6oll
@bj608baoao,o®fuo6oq}ogobo,8cogot}t}gud-po6€6068go09€6o"ao,botjbgobflodflo(5t|bo 8Oab6F90

14 CD25 (Tac) abbd®03doaogo Ttlg€6o"ao,8t'9€68"ao,806Ot3Od.iao IL-2-olj 6ooo3©codo

15 CD28 T 39€6Opoaoij „@ogub6ti8ogo" T ug€68fgoaob


1jt|b3o3Ugpt>8Gob9 abbd6Ou6Obb, Otb6b8t>1j608ag068bogpo
abbdoo@doau8O bo36bgolj 6oGo36mdo (tioa6t>go2)
3%6ofpoao "b8tl0698ab cD8o-b (87-1) 8b cD86-b (
87-2)
16 CD35 (CR1) 06ootdoooooao, do@3gp8@06®oLJ 6oGo30o6o 1,
u%dogoao, "b8tl068Obb c3b gb c4b, ab6b306maoab
@tr,6®oo©obo,6oo©6®8ogoao,ocofoo6n8ogoao,RO6fg6o68gofl2€6,i@,loo 8bacoGoombb

17 CD40 8 u9€6o"ao, 8 t}96608Oaob cob6b8t>1ioo8t)g068aogo


@m6oooooao, 1joa6bf¥OLi doG03®Odo "b8t)OdR8ab
go6p6o03gofl%d`1@`1ao cD4o goab6gb (T-BAM)

18 cD44 (8®afobtld goojcoooooao, "b8Bodpoat> jobg8606OLi @3b8bb


U%68gcobb6floati6o, Pgp-1) 06oondooo®obo ab6b3o6oboalj goodenoooobob bq}3ofoobb

19 CD45 (bbo6o}m gp80Jooo©8bo otodmfoo6q}mb8b®bfob, bdgoo6oali 8 gb


g8ot}ooobodo T ti9€do"aob t>6Ooa06tido
b66Oa06O (LCA) d0003006Ob libilubgoaoov boa6bgoti
abRbt3ti@t>b
20 CD450 T U96608U8° CD45-ob ofomq}ot6@b, 6en@ogoG b6
1jua3®3ugt>Gob, 839660Rtlgobtla3cO3ugpt>Oot>,8o6noo©oao,@b.idco9}ba.ibo tlooob8b A, B Rb C oaq]en@06obti

21 CD45A 8 u9€6ORoao, T CD45-ob ofomq}®6@b, 6m88goG tlooob81J


09€6ogogobua3cO3Ogpt>Gob(a38ua638Of¥O Ta%6o"ao)8o6m(}ofl,iao 8aSoao6Li

22 CD49a quA-1) abbd®Ou6Oa@f¥O Tu3€do"ao'@on6tr)(}ofl.|bo "b8tl06"bb d®gba06b, gt>@0606b

23 CD54 (ICAM-I) jo8®3®ofoodo gb @9€6ogE®6ot}o bg3ofootl6o 8engpojugb,


b6b3o@o3mo6t|6o (ICAM)-1 "b8806q?Obb cD 1 1 a\cDi8 ajFA-
@9?6oRoao 1) da cDii\cDi8 orAc-1) 06Ooa606Oalj,
d060.qodfltitiaob 6tit3ti3flodo
24 cD55a]AF] 3t|@®3ot|fofl6o gb f9t>tlgob f9b8bfif|b6{iatigo 9!bf|b060
bdulo@co3nofoo6o09€6o"bo a]AF), Odb8u06"abc3b-b, Ov808b c3\c5,}m6.1,|6flbfobb@,|8®6flt>fltj6068080u9€6O"aobb83Oqiotido

25 cD56 OrRE-1)CD580jFA-3)CD59(36onood©o6o) NK a9€68goaojo@®3®oq)t'6o 8bbdulo@m3ooq>U6oa%6o"ao


@mgpojof¥ob oq]®8®6@b orcAM),
bp3.iq]Ofl60 8Ogtitrmgt>
26
98Ot}cO006Oaoti 8ti6dGobbovb6
8btib8806Oaugo b6®Oa06O-3 qjFA-3),
"b88068Oab cD-2-b, bgp3Ob0060
8Ogtit]nebb
27 j88o3ooq]3do pbb6ulo8m3®ocbtl6ou%6o"ao adb8tlo6"ab j®83go@86oob t}m83®606®bc8-tjbfpbcg-tj,ag®db3b8O@a6b6Ob

Efl@t5fl,io ,]®83g[i,]botj bv#Oabb06gcocoogoo8obgoot}oooOU6o


28 CD62E a-1,ogodoo6o) 86Roooogo880
b8jocbou60 8cO8OdtifTb OELAro,
ab6b306®aoab 068cOOv8gou@cbo
6fio6doq)ofptiaob ao6.ibb (rolling)98odoGo0t|6obg3ocbou6o8®gojugpb
29 CD62 aJ-LioF¥od®o6o) 8 u9€608Oao, Ttl%6ogoao,
ajAM), ujb8@06"ab cD34-b, ICAM-tj,
@o6nGoooao, NKfl%6[i@,iao ab6b3o6®aoab o6gonotogou@foo o966ogob
ao6RbLi rolling
30 CD64 (FcgRI) @®6"Go0oao,8b.}6ong]ba.iao IgG-:b @bgbg b8o6U6o 60G830®60

31 CD79cbcD79P (IgobIgp) 8 0%6ogoao 8 t|%6Opob b6Ooa86060 6OG03OcO6Otj


dm83m6066o, CD3-olj b6bg®@o.1)bBo6®bbon6bfpobabgb(3.i8oljt>co.lobcob6b8bij®O@ug06Oaogo,cD28-bD8b

32 CD80 (87-1) 8 8%6898gpobtla303tlgbGob


TLA-4-0 goaD PCDl9gbCD2l-o}b6
0 jb8Bo6tlo 8 O9€609obcob6b6oGo30o61}d86olj
3334 CD81(b6®o3dogo306bGO090b6®obt,o@gob1jb8o$6.1(TAPA-1) go88coGo6obo

CD88 (C5AR) 3ogo8m6q}8gao6on8ugogpootjooo©oao,8bd6mq}baoao,3mtio,itsofl2€6,igtiao


jn83go@06®ob j®@3o6060ob c5a-b
6oGo36o6o

35 CD89 (FcaR) 8on6rpooooao, @bd6co8b@oao'a6b6ug®GOooao,T0%608Ugoboa3®3ugboooao,NK6og11ao IgA-b 6ooo30o6o

3637 CD102 (ICAM-2) 8Ob3068a8gogo8q}rpoo©oao,8co6coooooao,38ogbcbodgoo60boiitigdbd88OaoLi06poo70g8dfla€6,igtiafo,1


"b8tlodgoab CDlla\CD18-s qFA-I)

CD103 HML-1,aE o6 6bo3ooJogU6o aE o6®o@6o6o


o668a6o6o)
go88oGoooao,3860386o3gpo1,out,got,go@T]cot3OO,]aob 24%

38 CD104 (b4, b4 03oovogou@o, fl8b6ob b4 06®Oa6060


o6o8a6o6o) uxdogoao,fooao.i6coo tio8bm6ob
n%6`1g|1ao
39 CD106 ovcAM-1) 06socoof¥oo@oli bq3Ofoou6O @®got}ugb, goab6go
3966osoao VLA4-oliomoli
40 CD132 (ljbo6o}® T gb 8 t]9€6Of9Oao, 8 gb T 896dogoaob fo6gb
ab8t> 96bB8O ) q}Oa6coagbbooao,308o3o86060v06b@06a.ig.ibo

41 CD152 (CTLA-4) T b986O"ao T @9€d08Oaoti O8a6udo 3btit]hoii


Ub68®q}oono doatigboob: CD80/86mb
goab6go
42 CD154 (CD40L, T- T u9€6Ogoao Otb6b8bboo8tigodoaogo @®gotitlgb;
BAM) CD40-olj gop,b6@o
Qb6b6O.0 2. Ooono06Oach atxDquono6O O8a3boho

00®qxp6o q}a do6oouq?o OOcxpe06ch dofroouq?oba@rfe69g}afyoq?oq?6


to@ofroaegiv ftyrfu 6qu•'I,LEI. ckro"ogivO80to°
I I ootob aodrREofty o"fro
o6®06qpoo3o6-2 T ox6oq?oTox6oq?oCD4+Tox6oq?o CD25 (IL-2Rci) T "6oqr>: 36onqpo806boob q?b
(IL-2)o6®06qpongo6-3 CD122 (IL-2RP) 'CD132(yc) q?o30606oo6oab o8od®on6oqp a?a
@obboo6obch ox6oq?oDbq? ;
ocb6t]53oqpg on3b T
@36Oaoqp06Oaoqpo io}c6Oc9Obob
ab63OcDb6Oabb, abe?b6805bb q?b
sO6doocofro6ODbb.
N K "6oq}o= 36onq{lo806boob,
abb.1oon6.lob
CD123 (IL-3R) toubapbq]O:mc6Oq]Obch3afrob60
(IL-3)o5®06cpoo3o5-4 CD131 (Pc)
fty6bfro6Doq2O@O= Oife03b g8Oq::}b
liotibqp@abc9O Ox6Oq?OQ::io bbfoob
@on@6o3oabb
CD124 (IL-4R) a ex6oqu: ocbonoo3ob abq?b6o]3o
(IL-4) (Th2), 3onboo6o CD132 (yc) IgE-too
Ox6oq?o T a)c6Ocgrt: Th2 q7080605OOs006,
3froqp0306boob
aa8froBbao: bq=i®065b®Ooq::to
abbd®006Ob3 q7b IFN-y-O]O
ab63 06coaoBoq::io 3q::>bb03060
8bbd0006Oaob q23cO6a063bg{.Ifro6ngoqu:abbd®006Obb,
o5oo6qpoo3o5-5 CD4+ T "6oq?o CD125 (IL-5R)
(IL-5) (Th2), ILCs -Xa39o2 CD131 (Pc)
abdqpoo6o@oqpo Vb6@onj@5b

o6006quoo3o5-6 ab36®8bao, CD126 (IL-6R) eeodqu: ap3b30 8rfeoii OoqpoDoij


(IL-6) 06q?OcDoqro60 CD130 (gpl30) bo5o]ocbo
ex6oq?o, T 8 exfuqu:
Ox6ocgo b60obboocp6b6@on@J@5oqpo
ox6oq?oaob 3 6onqpo806boob
T "6oqxp: Th 17 a?o80606oo6oosrfurfucacoqofo
o6®06qpoo3o5-7 8o06o@qpbb®o, CD127 (IL-7R)
(IL-7) d3qpob ®8o5ch CD132 (yc)
"6Oq}Ofrots 606b8nd@Oqgobo: T a?a
b®6on@oqpo 8 Ox6Oq?Oaob bq?6Ooqpo
Ox6oq?o fro 6o@on6boq?ob oli 3 6onqg}o306bo ob
T qua8Ooobo: ace:i:Ob698O¢::)a a?6
a.ibuoLi6ODch fx6Oqpobob abq2b6806b3obqofro:p)c6oq?o,8{g)c6oq?o,T
o6®06qpoo3o6-9 CD4+ T ox6oq?od3qull®3o6ob CD129 (IL-9R)
(IL-9)o6006qpoo3o6-11 CD132 (yc)
a?Cfoqur q?a givngoqu8a6o
"froqroho: abqpb6806b q,a
abbd®006Obb
I L-1 1 Rcl o]6on@aveoooaob 6b6@ond@5b
(IL-11) b®6on@oqpoflx6oq?o CD130 (gpl30)

o6®06qpoo3o5-12 @b36on8bao, CD212 (IL- T "6Oqr>= Th 1 q2030606Oodoob


(IL-12): a?06q?6o®oqpo 12RP1) NK qtb T "froqu: IFN-y
IL-12A (p35)IL-12B(p40)o6oo6qpoo3o5-13 IL-12RP2 bo6o)ofoo, oo®moondbo306o
Ox6Oq?o
bdoo86onoob abfo6q?b
llll||-±¥|-Ill-
CD4+ T #x6o@o CD213al (IL- 8 "6Oq?O= Ofrob03Ob a6qp®6O]3b
(IL-13) (Th2), NKT 13Ral) IgE-fog
"6oq?o, ILCs CD213a2 (IL- 03oapoqp:givo :mc6oq?o= qp®6Sonb
-Xas8o 2, 13Ru2)
abdqpo6oboqpo Sb6@ onda 6b
3enboo6o CD132 (yc) fro3fro8bap : bqp0066boooqpo
3X6Oq?O abb,ib006OBb
o6oo6qpoo3o5-15 @b36033ao, bb3b IL-15RG N K ox6oq}o= 36cngpo806boob
(IL15) Oo3ob CD122 (IL-2RP) T "6oqu: @obboo6oach
Ox6Oq?oao CD132 (yc) cD8+io}c6Oq?O@Oij abq?b6806b q?a
3froqp0806boob
o6oo6qpoo3o6- CD4+ T ox6oq?o CD217 (IL-17RA) 0 "6Oqxp:
17A (IL-17A) (Thl7), ILCs- IL-17RC de@On305Ooch abdqp06ODoqpo
o6®06qpoo3o6- Xa38o 3 6b6aond@6b
17F (IL-17F) fro36ngb8o : do@on3o6o@chb q?b
oooon3o6oach ab6qpo6oboq{.o
5b6@ond@60
o "6oqu: GM-CSF a?b
G-CSF-b36®q?odoob
o6006qpoo3o6-21 Th2 ex6OQO, CD360 (IL-21R) 8 "6Oq}O: abbd®006Oab,
(IL-21) Thl7ox6oq?o, CD132 (yc) 3froqp0306boob, q?080606Ooboob
Tfh ox6oq?o Tfh "6oqu= 6b6@ond@6b
Th 17 :mc6oq?o: andqpoo6oDoqpo
Sb6@®d@6b
Nlt :mc6Oc9O: 836:le0060
@o@6o8ob3
o5®06qpoo3o6-23 @b36cO8bao, IL-23R T :mc6oq?o= Th 17 ox6oq?oaoii
(IL-23): a?86q?6o0ocpo CD212 (IL- q?o306050oboob q?a bb@6b3qpo8b
IL-23A (pl9)IL-12B(p40) Ox6oq?o 12RP1)

o5®06qpoo3o6-25 T ox6oQo, IL-17RB T :g}c6oq?o q?a H.ts -xao8o 2: IL-


(IL-25; IL-17E) 3oboo6oOx6Ocgo,o®foofro8oqpo,@b36n8bao,qpon6Son3b5o03ocoo"6o3X6oq?o 5, IL-13-b 5b6@onj@6b

o5®06qpoo3o6-27 @b36®8bao@O, IL-27Ro T :mc6Oqro= Th 1 c9080606Oobooch


(IL-27): q?06Q6oooqpo CD130 (gpl30) abdqp006Oab q?a Th 1 7-Ob
IL-27 (p28)E813(IL-27B) Ox6oq?o q2bo]6a063S q2080606Ooboob

a?06co3b5o d3cpob o3ofrob CD117 (KIT) 3qpigivo3cOo6®givohot}bqp@bxpo


Ox6oq?ch b®6on@oqpo sO608b6O :mc®Oq]O: Oifeo8b seocI7b
8bdoon6o (C-Kitqpoab6q?o) Ox6Oq?obo 306m3onocbo6o brfech @on@So8oabb

b6b6oqpq}ooO- T ox6oQo' CD116 (GM- :papnga6o 6o5a@o6boq}o : o63ogiv


@05veooch @b36on8bao, CSFRo) a6b5Oqpcooo®Oaobb q?b
30qpon6ob@bboo@o 06q?®cooqpo6o CD131 (Pc) @05tyioooob ch @Onapo8Oobu
qp06Oaoqpo Ox6oq?o, @b3608baob abb3®006Obb
8bdoon6o (GM-CSF) 8oD6m@qpbboo

@on6onoooob @b36m8baoao, CD115 (CSFIR) toUbququ "6ogivLe


3onqp®6ob@bboo@o 05q?®o)oqpo6o So6oan6Doq?o: o$303li @on6onooooach
qpo6oboqpo 0%6oq?o, d3qpch @on@6o3obbb
8bd®on6o (M-CSF, 03o6ob
CSF1) Ox6oQo,9}oa6onaqpbbbo

a6b6oquonoo®oli @b36®3bcoao, CD114 (CSF3R) toubququ "6ogivLe


3onqu6ob@bboo@o 8Oa6Oaqpbb®O, ysbasfty: Oife03li
qp06oaoqpo 06q?oncooue6o a6b5Ocponoooobob acoap03Oabb
8bdoon6o (G-CSF,CSF3) 3X6oq?o

11 oo3ch oobmjo6obch {v]aabho


IFN-c[ 3qubcb@boo®onoq} I FNAR 1 {raoqpb {g)c6Oq?O: b6®0306Oboqpo
06o CD118 (IFNAR2) bob®ol)o, fo6q?ob MHC I 3qpbbch
q?06q?6o®oqpo Odue6Obobb
Ox6Oq?O'@b.36mg}bao NK Ox6Oq?O@O : abbd0006Oab

IFN-P 3oa6onoqubu®o, IFNAR1 {B0qpb :g)C60q?o= b6®o3o6oboqpo


3qpbcb@booocooq? CD118 (IFNAR2) bob®obo, fo6q}ol} M HC I 3cpbbob
vurfur.
edue6Obobb
a?05q?6oooqpo"6oq?o N K "6Oqu= abbd®006O@b

o5®06806on5-y T ox6oQo CD119 (IFNGR1) @b3ch8®ap : 3qpbb03060


(IFN-y) (Thl, CD8+ Tox6oq?o),NK3X6Oq)Oq?bILcsuxac80I IFNGR2
abbd®006Obb (bdqp006Obb
@oj6®aonooq?06 306doobb)
a "6Oqu: Ocbcot)03Ob abq2b6O]8b
@bon8ho6ofoo6oaoqp q?b
3on@3qpo@06och ab8odbo6oboq::i Ig G
de03qpbbo8foo tqpbq7cO5O qpob
cobasoaao)
T "6Oqrt: Th lq?no0605Ooboob
bb8bq}bbb8b :g}C6oq?o: fo6@ob I qib I
I 3q:lbbob M H C @onqpo3oq::lo8ch
Odue6Obobb, bdqp006ODb b6®Oa05Ob
bbq?b@Oab3Oabb q}b T
ox6oqrooobo73 ob ffb6q? an6bb8b3ch8bac,a?06givooap
o5®06qpoo3o5-10 6b36on8bao, T CD210 (IL-10ROL)
(IL-10)o6®06qpoo3o6-22 Ox6Oq?o IL-10RP
"6Oqap: IL-12-ch,
(do6ocobq?bq? cob5b@bboo@Oqlt06Oaoqpo q?a M Hc 11
@b6Oasqp06Obo
3qpbbch 6onqpo3oqpoDob edb36oboob
qpo T ox6oq?o) q2bo]6a#53b
Thl7 ox6oq?o IL-22Rol 03Ooragiv ex6rm :
(IL-22) IL-10RP2 or IL-22oL2IL-10RP2
q?0806bo5o@ob Sb6@on5@5b, Cb6q?ob
bb6oo6oqp 306doobb
303unooo: 8xpb6806b
: qpo3onqpocoo

o6®06306®5-^ q?06q?6oooqu IFNLR1 (IL-28R) 03Ooragiv ex6Oqgiv :


(type Ill Ox6oq?o CD210B (IL- b5®o3o6oboqpo I)Oboobo
o6t)06g}.|6on6o)qp0030@ool, 10RP2)
O@D6o®6oqpo CD118 (LIFR) Q06q8fro ex6Oqu:
@bof3cfoo6oboqpo8b3®®6o(LIE) ®60305®®q?06@0'd8qub03o6oul,®6on@oqpo CD130 (gpl30) q?030606Oobooob a qp®306OBb
ox6oq?o
o53®b®boo5 M d3qpob ®3o6ob OSMR if eqrraprygiv "6grEN
b06on@oqpo CD130 (gpl30) 306 utcr}ocbo6o oo®on3o6o@ob
9X,6.1q?o 6b6@®]@6Ob 6Oanqpb Ob
TN F-ch onx6hob aodmuo6tgiv CINFSF) . °
uo@bo36ob ab36on3bao, NK CD120a
6ce6onfoob
deqprranggiv "f rgrun
Ox6oq?o' T (TNFRSF1) jGCD abba®006ODb (b6cooab,
8bdoon6o(TNF,TNFSF1)quo@8onoondbo5-a 3X6Oq?oTox6oQo, 8 120b (TNFRSF2)
3Ouaoqpboob)
6Ooofro8Oqpo : abbd®006Obb
3oB©qpbftyho : oboqpoDb
306cro, obo@o= 3b®ba®qpofo@o
(3Sbodbob)
CD120a Oa030, 6bo TNF
(LTh, TNFSF1) Ox6oq?o (TNFRSF1) bGCD120b(TNFRSF2)

qpo@8onoondbo6- T ox6oQo, NK LTPR qungigivodho8Oq]chto6O@:pqpr>


ap(LThp) Ox6oq?o, "froqu Qa qxp8rm6
8mqpo3oqpo6o qra6givoD:rm. imc6Ogo: de@0306ch
8 ox6oQo, Odue6Ouob q}b qpo@8060
qu@3®oq?06o06q?od®®6oOx6oq?o on6ab5onao6ofoo

BAFF -TNF-ob a?06q?6o®oqpo BAFF-R 8 "froqu: b6q7b6806b,


onxbbob a Ox6Oq?o' (TNFRSF13C) c>G 36onqpo806boob
Ox6Oq?ob @®6onoo®o, TACI
8babbj®006Oooqpo 80qpo3€qp36 (TNFRSF13B)
8bd®co6o a?06q?6oooqpo bGBCMA -8
(CD257, Ox6oq?o,B Ox6OQch
TNFSF13B) @Onaposobob Ooqpb(TNFRSF17)
ex6OQo
APRIL - T ox6oq?o' TACI 8 "froqu: 8bqpb6806b,
3froqu806SOOOu 905q?6oooqpo (TNFRSF13B) t>G 3froqpog!06boob
@bo5q?ooo6oboqpo Ox6oq?o' BCMA - a
qpoab5q?o - @ofrooooo, Ox6OQob
(CD256, 8onqpo3oqpo6 @®apo8OBob Ooqpb
TNFSF13) a?06@6oooqpo3X6oq?o (TNFRSF17)

onboout6on®oa6o5 onbooon@qpbb®o RANK froongqpaltoch6o6


o (OPG, (bo6cosoqpo "6Oqu: cO6a05b8b
TNFRSFllB) 8btoon6o nB-bqpoac6q7Obab@bbd0006Oaoqpo60co3®ql6o)qpoa66@o chooon3qpbbo ol} q?o80606ooboobb

IL|1-aliapadbchaobqjo6abo
o6®06qpoo3o6-1c[ @b36cO8bao, CD121a (IL-1R1) grnyprrtyrygivyuRTrfufjsfi.
(IL-lot) a?06q?6o®oqpoOx6OQo, IL-1RAP c>GCD121b
abbdo006Oab

8o06onbqpbb®o,05q?oncooqro6oOx6oq?o,306bbo5veooo (IL-1R2)
LI ) fi R 3 3 n

me3n I. naD6gfin Linb833nLI tinaef=n b6©n806o


f=)ti)I.ne013ao... 5 b6®o806ol} do6ocr)Dq?a jcTDl}06n ..................................... 40

oat)6t)6o I,oi>©08nit 8oo6 3aoim806oi, b6©o806-b6©nbboocpnl> jb8aof6n ................................... 40

8bi,b6bc9eiuf6Oapbq?8bamt]06Oaucp003Od©m6t)CPO bq?nu8D6©o
8odb60Q]6oao in)3n v. jn33E=33368nb LinLib33o 44
au6oa6o8o (oib6q2bt]mcpocpo) cgb bcgb3©bGou6o jma3cpoao6©ob 8bbd©oU6oab 44
(8Od06nqpo) 08u6t)6n 3bijubo .................................... 7 L}m83TO806©oi, 8bbd©nu6oaoi, bcp©066b©nucTo
au6oa6o8 o8o6u6 3bl>obE}n 6rn6b6ocTo u386oq?06o a?a
@rytcpojucpoao jm83cp0806©Oi, 8t>Od©Ot)f6Oaoi, jcpt>i,oju60 8Q36 .. 46
bq?D3©®ooo6 o8€606 3bl>uban 8rr)6b6ocpo o3€6oq?obn c9b jm83Cpo806©oi, 8bbd©ou6oaoi, qpoci©o6ni, 8cbb46
@ncpoJUCPO8o jma3cp0806©Oi, 8bbd©Ou6Oaoi, amcpm e3bq,b .... 47
oau6u6o 3bijuboi, aoc9oeioao 14 jm@3cp0806©Oi, 6OG03©mf6Oao .............................. 48

0183n ii. n3363fi 3ebgi]3n @n6e6nE=3 mf68b6nQDaoi, Oib8QbG8b jma3cpoa06©Oi,


333fi3q!3an 16 Qb@bfobo606ocpo 8md8ocgoaoi,8b6 ......................... 48
B-cpn@e3mGo©Oao jm83cpo806©oi, aomTmetnu6o o30d©oao .......... 5o
T-cpO@e3mGo©oao 8ou8b6n l]bl}D68oacpm bd©no6m6oao .......................... 50
Tqu6q}rnoo(So6ob"bf6bjcpDhodo6n"ha63m3.opboooan.........19 jm83ro806©oli a?b8bttnD606cTo bj©nu6rnaoao ........... 50
m6@b8Dq? 608D®no6o T o366ocgoao ....................... „ ..... 19 jrn83cro806©ol} l)ol)©08nl} 8oof6 8oj6m6cnb j8cTol>
y6T u3€6ocgoan 0$6U680CP8rnq}D
au6oa6n3n 8b608ocTo6obocpo T o366oc908o ............... 19 bDd©06ooanb co®8q2bo8b jrn83cpo606©ol> q!babtob606ocro
NKT o3a6oeobo So@md8ocgoaol>D8b6

NK-U3€f60C9obn jrn83cTo806©nl) C908oon©o


onb6cgbt]mcpoqpo cpo8e3nocgu6o u3€fioc9oao ....... 2o o»3n vl. gg3ri3f=3anLI @n3fieune Eo )6013aB ..... 52
8Ooqpmoq2u6Ot)336Oc9Oao 20 b6oioaoi> f9f6mi, n8u6t)6 fiobdGn06ao 8m6b6OcpO
8bj6rnq}b8oao/6m6moo©oan 2o i)3€6ocgoao
C906C96n©UCPo O3€60Qo6o 21 b6010aoli 8b63oolb6oab
cpD68063D6l)nb o3a6oq?08o 22 Cpoojcnoo©c»b@o86bGnb

q}mcpojucpo6-cgo6q?6n©ocro u8a6oq?08n .................... 22 u336oq?am6olJn Dcg3obnol> @nqpojucpo8n ...... „ ............. 54


8bDd©ou608ucpn B-cpo8q3moo©o ................................. 22 u336oq}oaob 6n86Donob 8odD6o%8o ............................... 55
86b6ucpmGn©oao 23 md3n VIl. J6Bnao6nli aou6ma.an @mG®6iiE@
6on©f6mq3ocro6n 23 @nT3JgE=3an. T g3efiaEnLi fi3u3BBnfin fee
omSn6mq3oCTn 23 3nLIBn330l)3Li3anLi am)3ofin jn3BE=3dLinii
3rtiboo6o o386oq?oao 24 3nE=3jgE=3an 58
aDsmq)ocpo6o 24 TU3€6oq3oi> 60oo3©m6o (TCR) .............................. 58
crt6m6anGo©oao 25 3olt©m8oolb8ijoaoi> 8Oib8b6o Jm63qpodboi, (MHc)
me3nlll. naB6gfin iinl-03@nL. nfiae6n3bn...26 8mcp0l)OCP0ao
o8u6uf6O I,Oi,©08Oi, G06©6Dcpu6O (3n63OcTbc9O) MHC I jcpDL,oL, 6cr,qpojupD
mfa8b6moao MHC 11 jcpol)ol} @roqpojucpD
cl8cpol> ©8o6o T-cpO8e3nGO©Oib6 6bfai,bc9806O GO©mq,mcrt)6n
a,o8ubo b6©oao6oaoi, 8bcgb6uab8oab (36mGoi,n68o) ..... 64
o8u6t)6o ijob©o8ob 306oe306oucpo (@om6bcgo) on©rntJmcpu6o Gncpoartll q)06@06©ocTn cgo86DcgbGnb ... 64
m68b6moao 303©oq?obol} 8bq?b©b6D on©msmcpoq?D6 06q?rn3cTbs@06

CTo8q3o6o j8®6clo abQoto


opo6ou 06em3cTbt)@t)6 aDq?0%o `MHC I-303©oq3o' jm83cpodl}nl)
8ujn%bl}ou6 bl)mon608ocpo cpn88U6o dl)m8oC:Pn b6tl"6b
`MHC I-303©oq?n' jcn83cpodbnl} odl}360l}nD o3€f6oq?ol}

c?na3rr)Go©Oaoi, 60G06jucpboob 34 S`oQo3n6So


ole3n lv. 9I)bnLi 33i)6na) n336gfin 06QmGn©ncbooiu3a6oQaoamb3oQ6ocpob6©o806ob
LlnL1333ni. 3n3ri 37 8aq?t>8t)at>3oat> (36mGoi,o68o) ................................. 65

oib6Q®dmcpncrn o8u6n©o©ob bb@n$60 8mcpojuc=poan D6©n8066b6@e806 u336ocgoaao u336ocg8Df6oou oncro8nb

(PAMP, DAMP) a?a @bcr)a ao@06mbo 6oG83©m6obn ...... 37 @ob8Cpb

uGbmit aoG6mab bQb3©bGou6o oat)6U6o boi,©08oii MHc ii jcpDi,Oi, amcpojucpoi, anmi,06Oiocbo c9b
8006 8ocbojucTb8o 8bcgbbc98ocToab ................................. 66
MHC 11 jc¥til>ob @mcpojt)cpoliou6 8oq?oaoaa8oaocpo
ao608ocpo6oaocpo T o3€6ocgo8ob (Go66f6ocpu6n Treg)
303©ocpoanb cgbjb8an6oab
g®68nd86b
66©n8066668Q806o o3€6oc9ob eiopo3o6e]o `MHC 11-
306oe306nt)cpo ©mTo6b6©mab 96
383©oq?o' jm83TOJbob odb36obnb .... 68
T-u3a6oQoaoi, bd©ou6maoij Qbo}68U68b ........... 96
mb3n vlll. d6Bnao6nii aoi]6ma®@n @m6b6neo B-t}3€6OQ?Oaoi, bd©Ou6maoi, a?boi68u68b .......... 97
Liefi3dBBfi3an. o6@nbt]3Bf=3an fee aeoin
l,U360l)Ob
fi3i)33bmfi3an...................68
306Oq30606e]O 06q2OG0606OcpO cD4+
b6©ni,boucp060
@D608ucpo608ocpo u336oq?06o ............................. 98
b6©nbboocpnb l}©6od©U6b
bb8a CD4+ T 8o608ocpo6oaocpo I)3€6oq?oart ....... 99
o6©obboocpcoo l>®6od©u6ocpo 8o6oo6©oao .............. 70
CD8+ 8b608ucpn606ocpo o366oq?08o ................ 100
b6©obboocpou jcpttl)obn
NKT o3€6oa?Can
b6©Oi,dot)p06Oi, 03Od©m6ucpO e3u6dGOoao ..... 73
y6T o%6oq?08o
b6@r)I,6oocTo)b 6oGo3©m6oao 73
oia3n xl I. n@g63fin 8OI.3Ifnti ri333f=3une ioi
oie3n lx. o6Bna36nb ae3n@i]6nan liBfiDd83ri3anu bl)bJO
@fio3oE=g3fin363anti @nt3t3an ..................... 76 6bj8oamab
b6©Oi,boucpoib 60306©ub6Oi, 6b6@md@6b ........ 76
Oau6u60 3bi,uboi, 608ucpbGob b6©0806Oon .... 1 0 1
806oanb 6ojrtt6ao6boobao @rn6b6rtpo q306@06©obn .... 78 b6©0806Ob a06Oao
brn@b®u6o 3o306@u©bonb
b6©o806ol} a?nQ]b.
oq)n©n3ob 8bq?b6Cn8b
o6©o806obrnf68b6o$8anaoti8D6ob8Sb......................101
T-u336Oenb 6oGo3©m6ou (TCR) 86b8bq- b6©o806-6b6@Q806o u3€6oc9oaoi, 6mcpo o@u6u6o
e306m86oaoi, 8ottoboao 3buubob 608ucpbGobao
in.3n x. T- pb 8 "gfioEoanL aJ]dbiiofi@ab ....... 81 oat)6t)60 3t>i,i)boi, ©03u Th ti336Oc9Oao
l)Dgtjobo q}D8b
8b6ltbq]CP86b8lt
oo6©6Dcpu6n e3Dtlb NK I)3e60Qoao Oat)6m8o608t)cpo6oaoap
l)b6mcpmm q3bbb
oo©mjo6oai,tt Qtt do8mjo6oab 8o8mo8t)at>8oab ... 1 o3
cD4+ T I)3€60q306oi> 8bbd©ou6oab 81
NKTu336Oc9Oao
o]b6D6ooo3©rn6nbb a?b l>b8D bcg3oSoo6o 8nqpojupoaob
t>6©oiibot)poaoi, 6mpo oat)6t}6o adituboit
cgD6n86opoab 608ucpbGnbao
cno6a8ob©o8t)cpo606ocpo lto866cpnb ao6066 ................ 82
OQom©03ti6o di,Oqpo
cpn@e3mGo©oi,8bbd©ot)6oaoi,ao@q?eim8U3a6oqaao
o8U6U6o 3bituboii 806o©oju6o jm6©6mcpo ... 1 o5
8o68ooiti6oat)cpri 36mGoi,oao 83
GO©mi}06Oao a?b do8mj06Oao io5
T-cpn8q3noo6nb 86.d®oo606ou ao8Q8o6 096f6oq?ao
b3m3©mQ>n
8D68oow606ocpo 36mool!oao oau6u6bcg 36o8ocpo8o6oaucpo I,m6oao .......... 1 o6
cD4+ T-u336Oq?Oaob i,ua3m3ucpbGooao ............... 84
Oat)6U60 3bliubol> 600f6m06Qrrlj606UcPo
T-B I)3€6oq?oaoi, jmm306bGob. a U3€6oq?Oaoit
8mqiupbGob
8bbcl©ot)60ab 608om6ucmo@u6n©o©o:b3OGobcpncboaucTOO8u6u60
t,oQt>3o6t}cpo oat)6m8cpmatiqo6ooi 8 I)3e6oQoit
3bi,ubo o3noioqpu6 dbm3ocpi,b a?b jb6ao ................... 1 o7
8bbd©ou6oaob u336oq?aoQb 8ttoao ..................... 87
03ncoocTu6naD6oo6oanbo8u6n®o©nbon8nbobt)6oab6n107
BD6bi,bbmob6 oo6©6ao ao8en6b60 36mGoi,Oao .......... 87
jb6ano806u63bl]ubou6gDjD8an606upoq?Db8Dq208obot08
cD8+ Go©m©mdi,OUT)6o T-u336Oq?Oaoi>
Oie3n xlll. t33aril)6na3E=nanL. fi3Ogt]n3an.
8bbd©ou6Oab I Bn3nL. t]3aafitl6na3E?nanli fie)3un3an ..1io
Go©m©mdi>odt)6o T-I)336oc906oi, aoo6 I,t>8ocb60
I ©03Oi, t,Oa86c)6maoTmaoi, 6ObdGOoao ........... 1 1 o
u336Oc9Oaoi, 8b6be8u6Oab
b06i,06Ocpost>Oooi,e3tit,t>
8ucpua6ti8oc?in T-I)3a6oC9oaoi, 8bbci©Ou60aoi, TH2 u3€6Oeoaob 8bDd©006Obb ................................ „ llo
ijb3b aQ]oao 8 o866ocgo6ob 8ttod©ot)6oao. IgE-tio oe>n®o3nb
bo306D6©n806n
8Dq3b6o)8b
a00606OOcpO OOcToao c9O T-0366Oq3Ob 8Oq30306OcpO
8bbd©ou6oaoi, e3bcbb
8mqpojupo6obl>bgo6Dbc?8cgo8no6©obboucpoao.......91
oe3Oci©m6uc]O q3bcbb
T-gt>8mt)Jog?oaocpo oat)6t)6o 3tti,I)bo: 3rnbno6ou336ocgoaob,abeme3ncpoanbDq?borne)o6mq}ocpo8nb
B-t}336oc906oi, 8bbd©ot)6oab T-u3a60q?Oaob 6mcpn CgbU8rn8608ocpo %o@86cl6m6ocpm6nb 6obdooobnb
qib68b6Oaoi, 8b6Oao
8b68nco®606Dan
m®3ii xi. I]06Bfi.E]fin q!d B]filigofilioen 3mboo6n U3€6oq?Oaoi, aoo6 6b68md86ocpo
bnE=3fie6BnaJ 80q?Ob©m60ao
G06©6bcpu6O ©mcp06b6©mab 94 bq?606b68md86ocpo 8ocgob©m6oao ................. „. 1 14
608b©nt)6n lJoe?odGnb
8omcpm8nu6n D@n6oao 114
86o6t)cpoaobGr)cpoaoq?o36fn®om8cpoja606o..........114
IV BnBnu b3a3fid6na3=nanb (T 33efi3f=3ano)
de novo gb68md860PO 80Q?Ob©m60ao ................. 1 t4
ab6Bnfimaoa]En) fiobdunoan ....................... 1 33
CToojm&6no6obo
ao6ocpoat)cpo ©riaou t,oa86d6maocpmar)I,
36mb©b8pD6q?n6oan
60bclG00ao
o]6rn86rnGn®06ob 8d@6od®no6oaocpn q3od®rn6o a}AF).1 15
i}m6©bd©u6o t>oa86ci6maoqmab ........................ 133
on©rnjo6oan
bo6boaocpn%booob q}DSD
Omcb06me3OEpOu6mcpoI©03OuQ>Oa86d6m6Ocpn6ou
8D8rn8po6ob q}b%b
6obciGooaao
Qtit)tjmo6oaoqo a?I 88m>6o 8t>t>ob 606dGooao 1 1 6 jm6©bd©06o a?068b©o©o
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