!"#"$ %'( ") *++,-"#"./ !"#$%&' ($)*"# +'#,"-."#/ 0121 3#45"--4# 45 0"$)6)7" 89:#)." ; 45 +"#A)7 +"#A)7/ !"#,:7>
C7...
91 downloads
1122 Views
28MB Size
Report
This content was uploaded by our users and we assume good faith they have the permission to share this book. If you own the copyright to this book and it is wrongfully on our website, we offer a simple DMCA procedure to remove your content from our site. Start by pressing the button below!
Report copyright / DMCA form
!"#"$ %'( ") *++,-"#"./ !"#$%&' ($)*"# +'#,"-."#/ 0121 3#45"--4# 45 0"$)6)7" 89:#)." ; 45 +"#A)7 +"#A)7/ !"#,:7>
C7.47)4 3"DD'..4/ 0121 3#45"--4# 45 ?",:.4A4*> :7$ E764A4*> 89:#)." ; 45 +"#A)7 +"#A)7/ !"#,:7>
F).9 647.#)B'.)47- B> G),4 %*B$% E"37,$; ., 8.%02(55< .7, 6%*2(%3.2$,$R- /,%(/,.- %33'68%*&"*+ ,%37 B$2+ %*B .' 37,3?; "( *,3,--%$& "* 3'*-2/.%."'* \".7 % 87&-"3"%* '$ -8,3"%/"-.; \7,.7,$ .7, B'-%+, -37,B2/,- 6,*."'*,B .7,$,"* '$ .7, 3'*.$%"*B"3%0 ."'*- -.%.,B #& .7, 6%*2(%3.2$,$- B"((,$ ($'6 .7, -.%.,6,*.- 6%B, "* .7, 8$,-,*. #''?= J237 ,>%60 "*%."'* "- 8%$."32/%$/& "68'$.%*. \".7 B$2+- .7%. %$, ,".7,$ $%$,/& 2-,B '$ 7%5, #,,* *,\/& $,/,%-,B '* .7, 6%$?,.= c5,$& B'-%+, -37,B2/, '$ ,5,$& ('$6 '( %88/"3%."'* 2-,B "- ,*."$,/& %. .7, 2-,$R'\* $"-? %*B $,-8'*-"#"/".&= 97, %2.7'$- %*B 82#0 /"-7,$- $,d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
"
%0",& &12 %,&1"$(
)*+,-./0,$1*+ 2/+3*4&*+
5(&'($' 6*77/&&'
?0%6@A( B 6&70% C(%,0)$0% \%- #'$* "* _%*'5,$; D,$6%*& "* LMb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d2,-."'*- "* 3/"*"3%/ 6,B"3"*, %*B /%#'$%.'$& 8$%3."3,= 97, 2(#6.,0#$.5 -%&-50) -,3."'* #,+"*- \".7 .7, '$+%*- '( .7, "662*, -&-.,6; ('//'\,B #& % B,0 -3$"8."'* '( .7, $,/,5%*. 3,//- '( .7, "662*, -&-.,6 %*B .7, 6,37%*"-6- #& \7"37 9 %*B A /&687'3&.,%3d2"$, 7"+7 /,5,/- '( -8,3"("3".&= J2$(%3, 6'/,32/,%$, B,-3$"#,B "* B,.%"/ "* B,(,$,*3, .' .7, ,*'$0 6'2- ,687%-"- 8/%3,B '* .7,6 "* 6'-. "662*'0 /'+"3%/ 82#/"3%."'*-= E B,-3$"8."'* '( %33,--'$& 3,//%*B *%.2$%/ ?"//,$ 3,//- ('//'\-= S,>.; .7, 726%* /&687'3&., %*."+,* -&-.,6 "- %*%/&K,B; ('//'\,B #& .7, 8$"*3"8/,- '( %*."+,* 8$'3,--"*+ %*B 7&8,$0 -,*-"."5".& $,%3."'*-= E2.'"662*".& %*B .'/,$%*3, %$, B,-3$"#,B "* .7, /%-. 8%$. '( .7, -,3."'*=
97, 5.'"%.$"%< .--5&8.$&"#) -,3."'* B,-3$"#,.7, 6'-. "68'$.%*. .,-. -&-.,6- "* "662*'/'+&= f)'*5,*."'*%/g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
!7
!"-&2-&( #$%&'()%*'+ ,-.%/.0+)1 D10 +,,(#0 K
49 49_ cEc cE: cAF c) c)1 cDC c!@JE c<E cSE cW cJW CE)J C3`!k"a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`,((,3.a 726%* %*."62$"*, %*."#'B& 7,8%."."- ) 5"$2_'B+?"*R- B"-,%-, 7"+7 ,*B'.7,/"%/ 5,*2/,726%* "662*'B,("3",*3& 5"$2726%* /,2?'3&., %*."+,* 7,%.0-7'3? 8$'.,"* 7,$8,- -"68/,> 5"$2726%* 90/&687'.$'8"3 5"$2"662*, 3'68/,> "*.,$3,//2/%$ %B7,-"'* 6'/,32/, "*.,$/,2?"*0L&03'*5,$."*+ ,*K&6, "*.,$B"+".%."*+ 3,// "*-2/"*0B,8,*B,*. B"%#,.,- 6,//".2"*.,$(,$'* "662*'+/'#2/"*
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Ƶ"0 #1 "(-0 1-&0": .-*' 4'1'*0' )&' 5()6#7/"#0'4 9/ %)7()6'0;
-%&'()(' *++,#& -.$/&+ A(' 5'00 #1 037( )* -%%3*' &'05#*0' 5)2'0 "(' >)/ 1#& "(' 05'7-1-7 -%%3*' &'05#*0'; !* ) 05'7-1-7 7/"#$-*' '*2-*%'*"< "(' 9#4/ 7)* 4': 7-4' >('"('& "# 5''4 "# ) %#&' (3%#&). .-*' #1 4'1'*0' #& ) %#&' 7'..3.)& .-*' #1 4'1'*0'; A(' %-6&)"-#* #1 )*"-6'*:5&'0'*"-*6 7'..0 B@OPC "# "(' ./%5(#-4 #&6)*0 1-&0" "&-66'&0 ) 0/0"'%-7 -%%3*' &'05#*0'< "('* ) 1$1"%2 %$0+"'0$; A(' 05'7-1-7 -%%3*' 0/0"'% 7#*0-0"-*6 #1 A )*4 Q ./%5(#7/"'0 -0 &'05#*0-9.' 1#& "(-0; A('0' 7'.. 0/0"'%0 7)* 5Ƶ' (-6(./ 05'7-1-7 &')7: "-#*0 "# "('-& &'05'7"-2' )*"-6'*0 )*4 3*4'&6# 7.#*). '85)*0-#*< "(30 )7(-'2-*6 ) (-6(./ '11'7: "-2' &'05#*0' "# )*4 %'%#&/ 1#& "(#0' )*"-6'*0;
! 2,#34+/45 67(#'(%5&$
01& *++,#& -.$/&+
"
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
! " 8
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
#?+$?%+@
CD34+
Pluripotent stem cell
Myeloid progenitor
Megakaryocyte
!
Lymphoid progenitor
Myeloblast
T-cell precursor
Monoblast
Thymus
B-cell precursor
Platelets B lymphocytes
P
h
Erythrocytes
a
g
o
c
y
t
e
s T lymphocytes
Basophils Eosinophils
Dendritic Neutrophils Monocytes cells A. Origin of cells of the immune system Innate immunity
Adaptive immunity
Infectious agents
Phagocytes
Natural killer cells
;2' 66 *'0 68=>
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
*'0 9#')+/3&*$)$= *'0 * .*') 2/'# .*0# /A $.*&& &-.%"/9-)#$>
6" 7&')589855)':8$(4 012,+):4 ;:55&( ! &-.%"/9-)#$ *+# .*('&- &//$#&- 0($)+(34)#0 )"+/4,"/4) )"# (')#+A/&&(94&*+ )($$4#B $/.# *+# *&$/ A/4'0 (' )"# (')+*#%()"#&(*& +#,(/'> !"# '4.3#+ /A (')+*#%()"#&(*& &-.%"/9-)#$ *'0 %&*$.* 9#&&$ ('9+#*$#$ 0+*.*)(9*&&- ;"#' ('@ A&*..*)(/' /994+$>
A(%:,+(%8- >%?8.5
Germinal center
White pulp
Sinus
Lymphoid follicle
Arteriole Follicular mantle
Mantle zone
Marginal zone
Marginal zone
B
B Periarterial lymphatic sheath
1. Anatomic structure
B T
B Germinal center
T
T
T
2. Cross-section through arteriole and follicle; lymphocyte circulation
A. Structure of the spleen Capsule
Secondary follicle Paracortex
Marginal sinus Interfollicular sinus
Afferent lymphatic
Medullary sinus Primary follicle
@&.482(.$8- A%:.':,-(5
Red pulp
!
"
Efferent lymphatic
1. Inactive lymph node
2. Active lymph node
B. Structure of the lymph node Follicle center
M cells
Mantle zone
Follicle-associated epithelial cells Dome region
T-cell zone
Postcapillary venule Muscular mucosa
Efferent lymphatic
1. GALT: Gut-associated1. lymphoid tissue; Peyer’s patch C. Mucosa-associated lymphoid tissue
2. BALT: Bronchus-associated 2. lymphoid tissue
! " C
=&*