Honorary President Giorgio Federici Università di Roma “Tor

Honorary President Giorgio Federici Università di Roma “Tor

Honorary
Honorary
Honorary
President
President
President
Giorgio
Giorgio
Federici
Federici
Giorgio
Federici
Università
Università
didi Roma
Roma
“Tor
Vergata”
Vergata”
Università
di Roma
“Tor“Tor
Vergata”
President
President
President
Carmine
Carmine
Carmine
Di Ilio
Di
Di Ilio
Ilio
Università
Università
Università
di Chieti
didi Chieti
Chieti
“G. “G.
d’Annunzio”
“G.d’Annunzio”
d’Annunzio”
Scientific
Scientific
Scientific
Committee
Committee
Committee
Mario
Mario
Mario
Alberghina
Alberghina
Alberghina
- Catania
-- Catania
Catania
Cesare
Cesare
Cesare
Balduini
Balduini
Balduini
- Pavia
-- Pavia
Pavia
Francesco
Francesco
Francesco
Bonomi
Bonomi
Bonomi
- Milano
-- Milano
Milano
Federico
Federico
Federico
Bussolino
Bussolino
Bussolino
- Torino
--Torino
Torino
Filiberto
Filiberto
Filiberto
Cimino
Cimino
Cimino
- Napoli
-- Napoli
Napoli
Chiara
Chiara
Chiara
CiniCini
Cini
- Roma
-- Roma
Roma
Carmine
Carmine
Carmine
Di Ilio
Di
Di Ilio
-Ilio
Chieti
-- Chieti
Chieti
Patrizia
Patrizia
Patrizia
Galletti
Galletti
Galletti
- Napoli
-- Napoli
Napoli
Renato
Renato
Renato
Gennaro
Gennaro
Gennaro
- Trieste
--Trieste
Trieste
Giuseppe
Giuseppe
Giuseppe
Paradies
Paradies
Paradies
- Bari
-- Bari
Bari
Fabiola
Fabiola
Fabiola
Sinigaglia
Sinigaglia
Sinigaglia
- Novara
-- Novara
Novara
Giancarlo
Giancarlo
Giancarlo
Solaini
Solaini
Solaini
- Bologna
-- Bologna
Bologna
Organizing
Committee
Organizing
Organizing
Committee
Committee
Antonio
Aceto
Chieti
Antonio
Aceto
-- Chieti
Antonio
Aceto
- Chieti
Stefania
Angelucci
Chieti
Stefania
Angelucci
-- Chieti
Stefania
Angelucci
- Chieti
Tonino
Bucciarelli
Chieti
Tonino
Bucciarelli
-- Chieti
Tonino
Bucciarelli
- Chieti
Vincenzo
De Laurenzi
Laurenzi
Chieti
Vincenzo
De
-- Chieti
Vincenzo
De Laurenzi
- Chieti
Piero
Del
Boccio
Chieti
Piero
Boccio
-- Chieti
Piero
DelDel
Boccio
- Chieti
Antonio
Di Giulio
Giulio
L’Aquila
Antonio
Di
-- L’Aquila
Antonio
Di Giulio
- L’Aquila
Carmine
Di Ilio
Chieti
Carmine
Di
-- Chieti
Carmine
Di Ilio
-Ilio
Chieti
Luca
Federici
Chieti
Luca
Federici
-- Chieti
Luca
Federici
- Chieti
Mauro
Maccarrone
Teramo
Mauro
Maccarrone
--Teramo
Mauro
Maccarrone
- Teramo
Filippo
Martini
Chieti
Filippo
Martini
-- Chieti
Filippo
Martini
- Chieti
Alfonso
Pennelli
Chieti
Alfonso
Pennelli
-- Chieti
Alfonso
Pennelli
- Chieti
Raffaele
Petruzzelli
Chieti
Raffaele
Petruzzelli
-- Chieti
Raffaele
Petruzzelli
- Chieti
Paolo
Sacchetta
Chieti
Paolo
Sacchetta
-- Chieti
Paolo
Sacchetta
- Chieti
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AMB 1
Ammine Biogene
:WLYTPULHUKZWLYTPKPULL_LY[HZ`ULYNPZ[PJLMMLJ[
VU+5(HNNYLNH[PVU
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AMB 2
Ammine Biogene
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AMB 3
Ammine Biogene
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AMB 4
Ammine Biogene
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AMB 5
Ammine Biogene
05=0;96,--,*;:6--,94,5;,+7(7(@(
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Biochimica Marina e dell’Ambiente
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Biochimica Marina e dell’Ambiente
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Biochimica Cellulare
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BCE 2
Biochimica Cellulare
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JLSS]PHIPSP[`WYLZLY]PUNUL\YP[LZHUKSV^LYPUNJLSSKLH[O
07VZ[LY:LZZPVU
BCE 3
Biochimica Cellulare
(IZJPZPJHJPKLUOHUJLZHKPWVJ`[LKPMMLU[PH[PVUHUKNS\JVZL
\W[HRLPUT\YPUL;3HUKPUO\THU(;4:*
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PUÅHTTH[VY`JLSSZHUKZ[PT\SH[LZZL]LYHSM\UJ[PVUHSHJ[P]P[PLZIV[OH\[VJYPUHSS`HUK
WHYHJYPUHSS`0U[LYLZ[PUNS`()(PZHSZVWYVK\JLKHUKYLSLHZLKI`O\THUHUKT\YPUL
WHUJYLH[PJIL[HJLSSZHUKHKPWVJ`[LZPUYLZWVUZL[VNS\JVZLTVYLV]LYUHUVTVSHY()(
Z[PT\SH[LZNS\JVZLKLWLUKLU[HUKWV[LU[PH[LZNS\JVZLPUKLWLUKLU[PUZ\SPUYLSLHZL
YLSLHZL -PUHSS` ()( JVUJLU[YH[PVU PU O\THU WSHZTH PUJYLHZLZ PU OLHS[O` Z\IQLJ[Z
HM[LY VYHS NS\JVZL SVHK ;V L]HS\H[L [OL LMMLJ[ VM ()( VU HKPWVJ`[L KPMMLYLU[PH[PVU
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SPWPK HJJ\T\SH[PVU KL[LJ[LK I` 6PS 9LK HUHS`ZPZ HUK [YPNS`JLYPKL KLWVZP[PVU (Z H
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HUKP[Z[HYNL[NLULZZ\JOHKPWVULJ[PU(7HUKNS\JVZL[YHUZWVY[LY.3<;^HZ
PUJYLHZLK4VYLV]LYPUTH[\YL;3HK`WVJ`[LZ()(ZPNUPÄJHU[S`Z[PT\SH[LK[OL
IHZHS NS\JVZL \W[HRL TLKPH[PUN I` .3<; [YHZSVJH[PVU [V [OL WSHZTHTLUIYHUL
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L_WLYPTLU[Z()(Z[PT\SH[LZ(R[WOVZWOVY`SH[PVUZPTPSHYS`[VPUZ\SPUHUK[OLLMMLJ[Z
VU NS\JVZL \W[HRL HYL KLJYLHZLK I` JV[YLH[TLU[ ^P[O H 702 PUOPIP[VY Z\NNLZ[PUN
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VMNS\JVZLHUKV[OLYZPUÅHTTH[VY`Z[PT\SPYLSLHZLZ()(PU[OLJ\S[\YLZ\WLYUH[HU[
(Z()(Z[PT\SH[LZPUZ\SPUYLSLHZL[OLLUKVNLUV\ZOVYTVUL()(WYVK\JLKI`[OL
HKPWVZL[PZZ\LJV\SKILPU]VS]LKPUYLN\SH[PVUVMNS\JVZLTL[HIVSPZTPUTHTTHSZ
07VZ[LY:LZZPVU
BCE 4
Biochimica Cellulare
;HYNL[PUN,YI)PTWHPYZ59.KYP]LUTHTTVZWOLYL
MVYTH[PVUVM,YI),YI)L_WYLZZPUN
IYLHZ[JHUJLYZ[LTJLSSZ
.9(A0(56=05*,5A6)(9*(9630+(50,3(./(:,409,A(
:;(::0.069.06
+,3(<9,5A0=05*,5A6 0(*6),330:;,-(56(5+:(3(.0(53<*(
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-VUKHaPVUL¸.+»(UU\UaPV*LU[YV:[\KPZ\SS»0U]LJJOPHTLU[V
*L:0=PH*VSSLKLSS»(YH*OPL[P0;(3@
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7H[OVWO`ZPVSVN`3HIVYH[VY`<UP]LYZP[`VM7HSLYTV 7HSLYTV0[HS`
4LKPH7OHYTHZYS*OPL[P0[HS`
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^OPJOZOV^HNYLH[LYJHWHJP[`[VTHPU[HPU[\TVYPNLULZPZ[OHU[OLYLZ[VM[OLJLSSZHUK
HYLTVYLYLZPZ[HU[[V[YHKP[PVUHSJHUJLY[OLYHWPLZ0[OHZILLUWYVWVZLK[OH[[HYNL[PUN
JHUJLYZ[LTJLSSZPZWVZZPIS`H^H`[VVI[HPUK\YHISLJHUJLY[YLH[TLU[YLZWVUZLZ
/LYL^LPKLU[PÄLK,YI)YLJLW[VYHZHTVSLJ\SHY[HYNL[VMIYLHZ[JHUJLYZ[LTJLSSZ
)*:*Z >L MV\UK [OH[ [OL ,YI)SPNHUK 59. Z[YVUNS` PUK\JLZ JSVUVNLUPJP[`
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MVYTH[PVU HUK (2; HUK ,YR HJ[P]H[PVU 4VYLV]LY [YLH[TLU[ ^P[O ,= H UV]LS
O\THUPaLKHU[PIVK`[HYNL[PUN,YI)PUOPIP[Z59.PUK\JLK,YI)WOVZWOVY`SH[PVU
HUKWYVTV[LZYLJLW[VYKV^UYLN\SH[PVU(SS[VNL[OLY[OLZLKH[HZ[YVUNS`Z\NNLZ[[OH[
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[\TVYZHSSV^PUNLSPTPUH[PVUVMJHUJLYZ[LTJLSSZ^P[OHJ[P]H[LK,YI)
07VZ[LY:LZZPVU
BCE 5
Biochimica Cellulare
(SRHW[VU\YPHPZHUV]LSO\THU
ZLJVUKHY`HT`SVPKVNLUPJKPZLHZL
4033<**03:79,(-0*6(;05;03)9(*650+./,AA03
7(**(.5050,),95(9+050.(4(;633(:*/043<7,;;07
693(5+0504:(5;<**0(
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(SRHW[VU\YPH(2<PZHU\S[YHYHYLKPZLHZLKL]LSVWLKMYVT[OLSHJRVMOVTVNLU[PZPJ
HJPKV_PKHZLHJ[P]P[`JH\ZPUNOVTVNLU[PZPJHJPK/.(HJJ\T\SH[PVU[OH[WYVK\JLZ
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HT`SVPKPZHZZVJPH[LK^P[OHMVYTVMTLSHUPU(ILYYHU[L_WYLZZPVUVMWYV[LPUZPU]VS]LK
PUHT`SVPKVNLULZPZOHZILLUMV\UKPU(2<JLSSZ6\YÄUKPUNZVUHSRHW[VU\YPHHZH
UV]LS[`WL00((HT`SVPKVZPZVWLUUL^PTWVY[HU[WLYZWLJ[P]LZMVYP[Z[OLYHW`ZPUJL
TL[OV[YL_H[L[YLH[TLU[WYV]LK[VZPNUPÄJHU[S`YLK\JLPU]P[YV/.(PUK\JLK(HT`SVPK
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07VZ[LY:LZZPVU
BCE 6
Biochimica Cellulare
-PIYVULJ[PUSHTPUPUHUK[`WL0=JVSSHNLUHYLRL`
JVTWVULU[ZVM[OLZPU\ZVPKZHZZVJPH[LKTLNHRHY`VJ`[L
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4LNHRHY`VJ`[LZ4RZKL]LSVW^P[OPU[OLIVULTHYYV^LU]PYVUTLU[I`PU[LYHJ[PUN
^P[OKPMMLYLU[L_[YHJLSS\SHYTH[YPJLZ,*4ZSVJH[LKH[IVULHUK]HZJ\SHYZ[Y\J[\YLZ
SL]LS 9LJLU[ L]PKLUJLZ KLTVUZ[YH[LK [OH[ 4RZ JVU[YPI\[L [V [OL LZ[HISPZOTLU[ VM
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WYV VY HU[PHUNPVNLUPJ MHJ[VYZ HUK ,*4 JYVZZSPURPUN LUa`TLZ HUK Z\WWVY[PUN
ÄIYVULJ[PU ÄIYPSSVNLULZPZ 0U [OPZ ^VYR ^L OH]L HUHS`aLK [OL ZWH[PHS KPZ[YPI\[PVU VM
4RZHUK,*4ZI`PTT\UVÅ\VYLZJLUJLPUT\YPULMLT\YZLJ[PVUZ>LMV\UK[OH[4RZ
^LYLWYLKVTPUHU[S`SVJH[LKPU[OLMLT\YKPHWO`ZPZ^P[OVUS`VM4RZ^P[OPU›T
MYVT[OLLUKVZ[LHSZ\YMHJLHUKTVYL[OHUVM4RZSVJH[LKSLZZ[OHU›TMYVTH
ZPU\ZVPK*VYYLSH[PVUIL[^LLU4RKPZ[HUJLMYVTZPU\ZVPKZHUKKPTLUZPVUZ\NNLZ[LK
HNYHKPLU[VMTH[\YPUN4RZ[V^HYKZ[OL]HZJ\SHYUPJOL5L_[^LKLJPWOLYLKIVUL
THYYV^ ,*4 JVTWVZP[PVU I` ^LZ[LYU ISV[[PUN HUK THWWLK [OL SVJH[PVU PU ZP[\ VM
KPMMLYLU[JVSSHNLUZ00000==0HUKNS`JVWYV[LPUZÄIYVULJ[PU[OYVTIVZWVUKPUSHTPUPU
]VU>PSSLIYHUKMHJ[VY>LMV\UK[OH[HSS[OLZLWYV[LPUZ^LYLKPMMLYLU[S`SVJH[LKPU[OL
LUKVZ[LHSHY[LYPVSHYHUKZPU\ZVPKHSKPZ[YPJ[ZZ\WWVY[PUN[OLJVUJLW[[OH[YLN\SH[PVU
VM OLTVWVPLZPZ PU [OL IVUL THYYV^ TH` HSZV KLWLUK MYVT TH[YP_ KPZ[YPI\[PVU
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TH[YP_ THPUS` JVTWVZLK VM ÄIYVULJ[PU SHTPUPU HUK [`WL 0= JVSSHNLU 0U[LYLZ[PUNS`
[OLZL[OYLLWYV[LPUZ^LYLHSZVKLTVUZ[YH[LK[VWYVTV[L[OYVTIVWVPL[PUKLWLUKLU[
4R KPMMLYLU[PH[PVU PU PU ]P[YV J\S[\YLZ VM IVUL THYYV^ OLTVWVPL[PJ WYVNLUP[VY JLSSZ
— — — MVSK PUJYLHZL YLZWLJ[P]LS` -PUHSS` ÄIYVULJ[PU
SHTPUPUHUK[`WL0=JVSSHNLU^LYLHSZVKLTVUZ[YH[LK[VILL_WYLZZLKHUKZ`U[OLZPaLK
I`KPMMLYLU[PH[LK4RZPU]P[YVHZKLTVUZ[YH[LKI`7*9HUK^LZ[LYUISV[[PUNHUHS`ZPZ
(SS [VNL[OLY [OLZL YLZ\S[Z Z\NNLZ[LK [OH[ 4RZ HYL PTWVY[HU[ ,*4WYVK\JPUN IVUL
THYYV^JLSSZHUK[OH[YLSLHZLK,*4ZZ\WWVY[TLNHRHY`VWV`LZPZHUKJVUJ\Y[V[OL
NLULYH[PVUVMIVULTHYYV^UPJOLZ
07VZ[LY:LZZPVU
BCE 7
Biochimica Cellulare
=,.-YLJLW[VYPU]VS]LTLU[PU[OLWLYPJ`[LSVZZPUK\JLK
I`,JVSPPUHUPU]P[YVTVKLSVMISVVKIYHPUIHYYPLY
46;;(*(:(34,904)(5-<:6*+((46+,6():*(30(4*
;6:*(564()(3),9./05(4((5+3<76.(
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;OLRL`HZWLJ[VMULVUH[HSTLUPUNP[PZPZYLSH[LK[V[OLHIPSP[`VMWH[OVNLUZ[VPU]HKL
[OLISVVKIYHPUIHYYPLY)))HUK[VWLUL[YH[L[OLJLU[YHSULY]V\ZZ`Z[LTBD
0U HU PU ]P[YV TVKLS VM ))) VU [OL IHZPZ VM JVJ\S[\YPUN WYPTHY` IV]PUL IYHPU
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WYVZ[HNSHUKPU IPVZ`U[OLZPZ ^L TLHZ\YLK 7., JVUJLU[YH[PVU HUK J`[VZVSPJ HUK
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H[[HJR[V)))(IL[[LY\UKLYZ[HUKPUNVM[OLTLJOHUPZTZI`^OPJO,JVSPLU[LY[OL
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07VZ[LY:LZZPVU
BCE 8
Biochimica Cellulare
W2PWPZHULHYS`LMMLJ[VYVMPTH[PUPIHJ[P]P[`PUJOYVUPJ
T`LSVNLUV\ZSL\RLTPH
*(3+(9,3300;9(465;(56(),5*0=,5.(+)69.0((7096AA0(
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;OLPU[YVK\J[PVUPU[OLYHW`VMPTH[PUPITLZ`SH[L:;0HOPNOS`WV[LU[HUKZLSLJ[P]L
PUOPIP[VY VM )*9¶()3 [`YVZPUL RPUHZL HJ[P]P[` SLK [V H [YLTLUKV\Z WVZP[P]L PTWHJ[
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YLJLP]PUNPTH[PUPIHUK[OLYLZWVUZLZPUHJJLSLYH[LKVYISHZ[PJWOHZLZVM[OLKPZLHZL
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^OPSL [OL *20 HJJ\T\SH[PVU ^HZ UV[ L]PKLUJLK PU HU PTH[PUPIYLZPZ[HU[ JSVUL >L
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[OLHU[PWYVSPMLYH[P]LHJ[P]P[`VM)*9¶()3[HYNL[PUNKY\NZ
07VZ[LY:LZZPVU
BCE 9
Biochimica Cellulare
5(+SL]LSZJVU[YVS*HZ[VYLZYLWSLUPZOTLU[HUK
TP[VNLUPUK\JLKPUJYLHZLVMJ`[VZVSPJ*H
I`(+79KLWLUKLU[;974NH[PUNPUO\THU;
S`TWOVJ`[LZ
4(.565,40926 )(<,905.(76..0(3,::(5+96
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+LW[VM,_WLYPTLU[HS4LKPJPUL:LJ[PVUVM)PVJOLTPZ[Y`HUK*LU[LYVM,_JLSSLUJL
MVY)PVTLKPJHS9LZLHYJO*,)9<UP]LYZP[`VM.LUV]H0[HS`"
+LW[VM0U[LYUHS4LKPJPUL<UP]LYZP[`VM.LUV]H0[HS`"
4VSLJ\SHY6UJVSVN`HUK(UNPVNLULZPZ<UP[09**:(6<:HU4HY[PUV0:;5H[PVUHS
0UZ[P[\[LMVY*HUJLY9LZLHYJO.LUV]H0[HS`"
+LW[VM*VTWHYH[P]L)PVTLKPJPULHUK-VVK:JPLUJL<UP]LYZP[`VM7HKV]H0[HS`"
(K]HUJLK)PV[LJOUVSVN`*LU[LY()*.LUV]H0[HS`
0U[YHJLSS\SHY 5(+ SL]LSZ B5(+DP HYL PTWVY[HU[ PU YLN\SH[PUN O\THU ; S`TWOVJ`[LZ
Z\Y]P]HSJ`[VRPULZLJYL[PVUHUK[OLJHWHJP[`[VYLZWVUK[VHU[PNLUPJZ[PT\SP5(+
KLYP]LK*HTVIPSPaPUNZLJVUKTLZZLUNLYZWYVK\JLKI`*+WSH`HWP]V[HSYVSL
PU;JLSSHJ[P]H[PVU
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*H OVTLVZ[HZPZ IV[O PU [LYTZ VM TP[VNLUPUK\JLK B*HDP PUJYLHZL HUK VM
LUKVWSHZTPJYL[PJ\S\T*HZ[VYLYLWSLUPZOTLU[3V^LYPUNB5(+DPI`-2TLKPH[LK
5PJV[PUHTPKL WOVZWOVYPIVZ`S[YHUZMLYHZL PUOPIP[PVU KLJYLHZLK [OL TP[VNLUPUK\JLK
B*HDPYPZLPU1\YRH[JLSSZHUKPUHJ[P]H[LK;S`TWOVJ`[LZ(JJVYKPUNS`[OL*HJVU[LU[
VM;OHWZPNHYNPUZLUZP[P]L*HZ[VYLZ^HZNYLH[S`YLK\JLKPU[OLZLJLSSZPU[OLWYLZLUJL
VM -2 >OLU 5(+ SL]LSZ ^LYL PUJYLHZLK I` Z\WWSLTLU[PUN WLYPWOLYHS ISVVK
S`TWOVJ`[LZ^P[O[OL5(+WYLJ\YZVYZUPJV[PUHTPKLUPJV[PUPJHJPKVYUPJV[PUHTPKL
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M\UJ[PVUZZ\JOHZWYVSPMLYH[PVUHUK03YLSLHZLPUYLZWVUZL[VTP[VNLUZ
07VZ[LY:LZZPVU
BCE 10
Biochimica Cellulare
9LN\SH[PVUVM072NLULL_WYLZZPVU!YVSLVM[OLWYVZZPTHS
WYVTV[LY.*YPJO:WIPUKPUNYLNPVU
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+00306*(9405,(:(0(9+0(+63-6)-(=(3696)(9;636(
+LWHY[TLU[VM:WLYPTL[HSHUK*SPUPJHS:JPLUJLZ
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H
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[YHJ[JHYJPUVTH6UJVNLUL 07VZ[LY:LZZPVU
BCE 11
Biochimica Cellulare
9LKV_YLN\SH[PVUVM[OL[\TVYZ\WWYLZZVY7OVZWOH[HZL
HUK[LUZPUOVTVSVNKLSL[LKVUJOYVTVZVTL[LU7;,5
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M\UKHTLU[HS [V PKLU[PM` [OL \UKLYS`PUN TVSLJ\SHY TLJOHUPZTZ SLHKPUN [V JHUJLY
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OH]L WYL]PV\ZS` ZOV^U [OH[ 4ZY( PZ KV^UYLN\SH[LK PU H U\TILY VM O\THU IYLHZ[
JHUJLYZ SLHKPUN [V [OL KL]LSVWTLU[ VM [^V PTWVY[HU[ OHSSTHYRZ VM [\TVYPNLULZPZ
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RPUHZL702WH[O^H`[OH[PZ\WYLN\SH[LKPUZL]LYHSO\THUJHUJLY6\YO`WV[OLZPZ
PZIHZLKVU[OLMHJ[[OH[4ZY(KV^UYLN\SH[PVUPUJHUJLYJHUWV[LU[PH[L[OL/6
ZPNUHSPUNWH[O^H`I`KPZHJ[P]H[PUN[OL[\TVYZ\WWYLZZVY7;,50UHULMMVY[[VL_WHUK
V\YWYL]PV\Z^VYROLYL^LZOV^[OH[ZPSLUJPUNVM4ZY(PU4+(4)JLSSZPUK\JLZ
WVZ[[YHUZSH[PVUHSKV^UYLN\SH[PVUVM7;,5IV[OPU]P[YVHZ^LSSHZPU]P]VTVKLSZ
9LMLYLUJLZ
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*-H]HSVYV)4L[OPVUPULZ\SMV_PKLYLK\J[HZL(KV^UYLN\SH[PVUPUO\THUIYLHZ[JHUJLYJLSSZYLZ\S[ZPUH
TVYLHNNYLZZP]LWOLUV[`WL7YVJ5H[S(JHK:JP<:(
07VZ[LY:LZZPVU
BCE 12
Biochimica Cellulare
-aKH5V]LS7YVNUVZ[PJ4HYRLYPU)YLHZ[*HUJLY
3(;;(5A0696::(5646;;63,:,4(9*,33(5(;(3070,9.069.06
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3\TPUHS)/,9ULNH[P]LJHUJLYZ;O\ZJ-aKJV\SKILHUL^WYVNUVZ[PJTHYRLY
HUKWV[LU[PHS[HYNL[VM[OLYHW`PUIYLHZ[JHUJLY
9LMLYLUJLZ
9L`H;*SL]LYZ/>U[ZPNUHSSPUNPUZ[LTJLSSZHUKJHUJLY5H[\YL"!
*SL]LYZ/>U[IL[HJH[LUPUZPNUHSPUNPUKL]LSVWTLU[HUKKPZLHZL*LSS"! 4HJ+VUHSK ); L[ HS >U[IL[HJH[LUPU ZPNUHSPUN! JVTWVULU[Z TLJOHUPZTZ HUK KPZLHZLZ +L] *LSS
"! *HU[PSLUH : L[ HS -YPaaSLK YLJLW[VY THYRZ YHYL OPNOS` [\TV\YPNLUPJ Z[LTSPRL JLSSZ PU TV\ZL HUK
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:[.HSSLU0U[LYUH[PVUHS,_WLY[*VUZLUZ\ZVU[OL7YPTHY`;OLYHW`VM,HYS`)YLHZ[*HUJLY(UU
6UJVS"!
07VZ[LY:LZZPVU
BCE 13
Biochimica Cellulare
*LYHTPKLÅV^MYVT[OL,UKVWSHZTPJ9L[PJ\S\T[V[OL
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3,:;<5--/,9=i90)6503(<9(=0(507(63(
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07VZ[LY:LZZPVU
BCE 14
Biochimica Cellulare
*OLTV[OLYHW`HZZVJPH[LKV_PKH[P]LZ[YLZZ!YVSLVM
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PUHTVYLHNNYLZZP]LWOLUV[`WL7YVJ5H[S(JHK:JP<:(
07VZ[LY:LZZPVU
BCE 15
Biochimica Cellulare
5L^WLYZWLJ[P]LPU[OLHZZLZZTLU[
VM[V[HSPU[YHJLSS\SHYTHNULZP\T
:(9.,5;0(AA<99(4,9633,3<*0((5+9,(50.0<30(
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07VZ[LY:LZZPVU
BCE 16
Biochimica Cellulare
0THNPUNÅV^J`[VTL[Y`![OLWYVZWLJ[P]LJLSS\SHYHUHS`ZPZ
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PZ YLK\JLK ^P[O YLZWLJ[ [V [OL JVU[YVS ZHTWSL [OL Å\VYLZJLUJL PU[LUZP[` PU JLSS
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07VZ[LY:LZZPVU
BCE 17
Biochimica Cellulare
4VUP[VYPUNTHNULZP\TLMÅ\_J(47PUK\JLKPU/3JLSSZ
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;OLZL YLZ\S[Z OPNOSPNO[ [OL JHWHIPSP[` VM +*/8 MVY [OL X\HU[PÄJH[PVU VM [V[HS
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4HYYHJJPUP*L[HS*OLT:JP!
9VTHUP(L[HS 1)PVS*OLT!
07VZ[LY:LZZPVU
BCE 18
Biochimica Cellulare
0U[YHJLSS\SHYTHNULZP\TJVU[LU[JOHUNLZK\YPUN
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4,9633,3<*0( *(77(+65,*65*,;;05(-(99<..0(.06=(55(
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BCE 19
Biochimica Cellulare
5-ț)TLKPH[LKKV^UYLN\SH[PVUVMJVSSHNLUZ`U[OLZPZ
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BCE 20
Biochimica Cellulare
;5-ĮTVK\SH[LZ/,4(KLWLUKLU[PUÅHTTH[VY`YLZWVUZL
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BCE 21
Biochimica Cellulare
0UJYLHZLKP56:HJ[P]P[`PU=HZJ\SHY:TVV[O4\ZJSL*LSSZ
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07VZ[LY:LZZPVU
BCE 22
Biochimica Cellulare
ơ*HYV[LULHUKS`JVWLULHMMLJ[LUKV[OLSPHSYLZWVUZL[V
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07VZ[LY:LZZPVU
BCE 23
Biochimica Cellulare
*OHYHJ[LYPaH[PVUVM5P[YPJ6_PKLLUa`TH[PJWYVK\J[PVUPU
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BCE 24
Biochimica Cellulare
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07VZ[LY:LZZPVU
BCE 25
Biochimica Cellulare
AVSLKYVUPJHJPKLUJHWZ\SH[LKPUZLSMHZZLTIS`WLN`SH[LK
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07VZ[LY:LZZPVU
BCE 26
Biochimica Cellulare
7YV[LVTPJWYVÄSLVMJHUJLYZ[LTJLSSZKPMMLYLU[PH[PVU
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07VZ[LY:LZZPVU
BCE 27
Biochimica Cellulare
;YLH[TLU[VM(43JLSSZ^P[OH.X\HKY\WSL_ZLSLJ[P]L
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07VZ[LY:LZZPVU
BCE 28
Biochimica Cellulare
)06*/,40*(3*/(9(*;,90A(;0656--6330*<3(9
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BCE 29
Biochimica Cellulare
,_[YHJLSS\SHYZWOPUNVZPULWOVZWOH[LJVU[YPI\[LZ[V
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BCE 30
Biochimica Cellulare
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BCE 31
Biochimica Cellulare
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BCE 32
Biochimica Cellulare
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BCE 33
Biochimica Cellulare
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BCE 34
Biochimica Cellulare
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BCE 35
Biochimica Cellulare
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BCE 36
Biochimica Cellulare
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BCE 37
Biochimica Cellulare
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BCE 38
Biochimica Cellulare
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07VZ[LY:LZZPVU
BCE 39
Biochimica Cellulare
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07VZ[LY:LZZPVU
BCE 40
Biochimica Cellulare
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07VZ[LY:LZZPVU
BCE 41
Biochimica Cellulare
05=0;96,--,*;:6--,94,5;,+7(7(@(
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07VZ[LY:LZZPVU
BCE 42
Biochimica Cellulare
)PVLULYNL[PJJVUZLX\LUJLZVM7(92T\[H[PVUZPU
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07VZ[LY:LZZPVU
BCE 43
Biochimica Cellulare
-PIYVISHZ[Z[YHUZMLYSPWPKZHUKWYV[LPUZ[V[\TVYJLSSZ
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07VZ[LY:LZZPVU
BCE 44
Biochimica Cellulare
0KLU[PÄJH[PVUHUKJOHYHJ[LYPaH[PVUVMJHUJLYZ[LT
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BCE 45
Biochimica Cellulare
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BCE 46
Biochimica Cellulare
-3(=05(+,505,+05<*3,6;0+,4,;()630:4
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BCE 47
Biochimica Cellulare
0KLU[PÄJH[PVUVMU\JSLHYWYV[LPUZ^OPJOPU[LYHJ[
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BCE 48
Biochimica Cellulare
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07VZ[LY:LZZPVU
BCE 49
Biochimica Cellulare
J4L[[YHUZHJ[P]H[PVUI`-793YLX\PYLZ5(+7/V_PKHZL
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BCE 50
Biochimica Cellulare
5V_PUOPIP[VY`HJ[P]P[`VMUL^Z`U[OLZPaLKV_PUKVSL
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BCE 51
Biochimica Cellulare
)HNWVS`TVYWOPZTZTH`ILHZZVJPH[LK[VOLHY[MHPS\YL
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BCE 52
Biochimica Cellulare
,WPNLUL[PJTVK\SH[PVUVMHUHUKHTPKLO`KYVSHZLI`
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BCE 53
Biochimica Cellulare
;YHUZJYPW[PVUHSHUKLWPNLUL[PJYLN\SH[PVUVM(UHUKHTPKL
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BCE 54
Biochimica Cellulare
,?79,::065(5+7<90-0*(;0656-9,*65)05(5;
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BCE 55
Biochimica Cellulare
0KLU[PÄJH[PVUVMZWLJPÄJWYV[LPUZHMMLJ[LKI`TPJYV95(ZPU
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HUK^LZ[LYUISV[HZZH`Z
9-HYHVUPVL[HS*LSS+LH[O+PMMLY 07VZ[LY:LZZPVU
BCE 56
Biochimica Cellulare
:LYV[VUPUYVSLPUKL]LSVWTLU[M\UJ[PVUHUKKPZLHZLVM
[O`YVPKNSHUK
:03=0(5(9,(:0465(;(-<90(55(*6:;(.3063(
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+LWHY[TLU[VM)PVJOLTPZ[Y`HUK4LKPJHS)PV[LJOUVSVNPLZ
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:LYV[VUPU PZ HU \IPX\P[V\Z TVUVHTPUL ^OPJO L_LY[Z IV[O H TVYWOVNLUL[PJ HUK
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HYL HZZVJPH[LK ^P[O H ]HYPL[` VM KPZVYKLYZ YHUNPUN MYVT KLWYLZZPVU ZJOPaVWOYLUPH
HUK HU_PL[` [V O`WLY[LUZPVU LH[PUN KPZVYKLYZ OLHY[ ]HS]L KPZLHZL PYYP[HISL IV^LS
Z`UKYVTLHUKV[OLYZ
>LMVJ\ZLKV\YPU[LYLZ[VU[OLYVSLVMZLYV[VUPUPU[O`YVPKKL]LSVWTLU[M\UJ[PVUHUK
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07VZ[LY:LZZPVU
BCE 57
Biochimica Cellulare
463,*<3(94,*/(50:46-(*;0656-763@7/,565
,(.9,,5;,(,?;9(*;05),50.5(5+
;<4690.,50*796:;(;,,70;/,30(3*,33:
90AA0-,+,90*(:03=((3,::(5+965(765,330=(3,90(
;(9+0;6:(=,9069(4(AA05(03,(5()65(*0504(9;05((5+
),;;<AA0:(=,906
+LWHY[TLU[VM)PVTLKPJPUL)PV[LJOUVSVN`HUK;YHUZSH[PVUHS9LZLHYJOHUK*LU[YL
MVY4VSLJ\SHYHUK;YHUZSH[PVUHS6UJVSVN`*64;<UP]LYZP[`VM7HYTH7HYTH0[HS`"
5H[PVUHS0UZ[P[\[LVM)PVZ[Y\J[\YLHUK)PVZ`Z[LTZ05))9VTL0[HS`
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7*HI\[[OLTLJOHUPZTVMHJ[PVUPZZ[PSSVIZJ\YL0UJYLHZPUNKVZLZVM7VS`WOLUVU
,7VS`,HZ[HUKHYKPaLK.;,^LYLNP]LU[VPTTVY[HSPaLK75;HTPTPJRPUNPUP[PHS
Z[HNLZ VM 7*H VY TL[HZ[H[PJ HUKYVNLUPUKLWLUKLU[ 7* TPTPJRPUN HK]HUJLK
Z[HNLZ.YV^[OHYYLZ[KLJYLHZLK]PHIPSP[`HUKJLSSKLH[OVJJ\YYLKPUIV[OJLSSSPULZ
I\[75;H^LYLTVYLZLUZPISL[V7VS`,0*$›NT3[OHU7*0*$›N
T3*LSSJ`JSLHYYLZ[VJJ\YYLKH[..WOHZLZMVY75;H^P[O\WYLN\SH[PVUVMW
HUKKV^UYLN\SH[PVUVMJ`JSPU+^OPSLH[.4WOHZLZMVY7*JLSSZPU^OPJOVUS`
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(;-HJ[P]H[PVUVM?)7T95(ZWSPJPUNHUKKYHTH[PJ\WYLN\SH[PVUVM*/67HUK
.(++RL`WYV[LPUZVM,9:TLKPH[LKJLSSKLH[O<UKLY[OLZLJVUKP[PVUZ7*KPLK
]LY`ZSV^S`!JLSSZZOYPURI\[YLTHPUH[[HJOLK[V[OLZ\YMHJL^P[OMVYTH[PVUVMJHSJLPU
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;OLJVTIPUH[PVUVMTVSLJ\SHYTHYRLYZHUKTVYWOVSVNPJHSHUHS`ZPZPZJVTWH[PISL^P[O
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BCE 58
Biochimica Cellulare
*3<:;,9050:,70.,5,;0*(33@9,79,::,+05/<4(5
796:;(;,*(5*,9*,33:
90AA0-,+,90*(*63,;;(4(90(5.,3(9(4(AA05(03,(5(
)65(*0504(9;05(5(765,330=(3,90(:03=((3,::(5+96(5+
),;;<AA0:(=,906
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*LU[YLMVY4VSLJ\SHYHUK;YHUZSH[PVUHS6UJVSVN`*64;<UP]LYZP[`VM7HYTH
7HYTH0[HS`"5H[PVUHS0UZ[P[\[LVM)PVZ[Y\J[\YLHUK)PVZ`Z[LTZ05))9VTL0[HS`
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WYVK\JLK MYVT [OL ZPUNSL JVW` O\THU *3< NLUL I\[ ]LY` SP[[SL PZ RUV^U HIV\[
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OV^L_WYLZZPVUVM*3<PZHMMLJ[LKI`LWPNLUL[PJKY\NZZ\JOHZ(aH»KLV_`J`[PKPUL
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UVYTHSVYPTTVY[HSPaLKJLSSZ(*W.PZSHUKVU*3<WYVTV[LYPZVUS`TL[O`SH[LKPU
JHUJLY JLSSZ ;OL *3< T95( ZPSLUJLK I` JLSS [YHUZMVYTH[PVU PZ YLL_WYLZZLK I`
(A( HUK ;:( 9LL_WYLZZPVU VM ZPSLUJLK *3< T95( ^HZ HZZVJPH[LK [V U\JSLHY
SVJHSPaH[PVU VM *3< HUK PUK\J[PVU VM HWVW[V[PJ JLSS KLH[O ;:( HSVUL VY ^P[O (A(
PUJYLHZLZ / WVZ[[YHUZSH[PVUHS TVKPÄJH[PVU /(JL /3`Z (JL /3`Z 4L[ PU
[OL*3<WYVTV[LYJH\ZPUNHJ[P]L[YHUZJYPW[PVU0UJVUJS\ZPVUHZWLJPÄJ*3<T95(
[YHUZJYPW[PZLWPNLUL[PJHSS`KV^UYLN\SH[LKPUWYVZ[H[LJHUJLYJLSSZ[OYV\NOJOYVTH[PU
YLTVKLSSPUN 9LL_WYLZZPVU VM *3< I` LWPNLUL[PJ KY\NZ \UYH]LSZ P[Z [\TV\Y
Z\WWYLZZVYHJ[P]P[`
07VZ[LY:LZZPVU
BCE 59
Biochimica Cellulare
7YV[LJ[P]LYVSLVMơLZ[YHKPVS[V^HYK03SL\RVJ`[L
L_WYLZZPVU[YPNNLYLKI`PU[LUZL[YHPUPUNPULSP[L`V\UN
MLTHSLH[OSL[LZ
;905.(30*90:;05( :*(3(369,+(5(:03=,:;9003(90(
:*<9(;09(--(,3,40*/0,365.06=(550(3),9;0.0(470,;96(5+
=,5,9(5+6)9<56
+LWHY[TLU[VM4LKPJHS)PV[LJOUVSVN`HUK;YHUZSH[PVUHS4LKPJPUL
<UP]LYZP[`VM4PSHU:JOVVSVM,_LYJPZL:JPLUJLZ30;(
]PH-SSP*LY]P :LNYH[L4PSHU 0[HS`
+LWHY[TLU[VM)PVTLKPJHS:JPLUJLZMVY/LHS[O<UP]LYZP[`VM4PSHU
:JOVVSVM,_LYJPZL:JPLUJLZ
,_LYJPZLWLYMVYTLKH[HJVTWL[P[P]LSL]LSJV\SKKLLWS`TVKPM`[OLPTT\ULZ`Z[LT
HUK[OLJ`[VRPULYLZWVUZLVMH[OSL[LZ0U[OPZYLWVY[^LKLTVUZ[YH[LK[OH[`V\UNLSP[L
MLTHSLH[OSL[LZLUNHNLKPUZ`UJOYVUPaLKZ^PTTPUNHUKHY[PZ[PJN`TUHZ[PJZZOV^LK
HUPUJYLHZLVMPU[LYSL\RPU03HUK[\TV\YULJYVZPZMHJ[VYHSWOH;5-ƠT95(Z
L_WYLZZPVUVMISVVKTVUVU\JSLHYJLSSZPUJVTWHYPZVU[V`V\UNNPYSZWLYMVYTPUNZWVY[
H[HYLJYLH[PVUHSSL]LS;OLPUJYLHZLVM03HUK;5-ƠT95(ZKPYLJ[S`KLWLUKLKVU
[OLPU[LUZP[`VM[OL[YHPUPUN03NLULL_WYLZZPVUHWWLHYLK[VILTVK\SH[LKI`[OL
SL]LSZVMJPYJ\SH[PUNVLZ[YVNLUZHUKWYLW\ILY[HSNPYSZYL]LHSLK[OLOPNOLYPUJYLHZLZVM
030UHKKP[PVU)40IVK`THZZPUKL_WLYJLU[PSL^HZPU]LYZLS`JVYYLSH[LK^P[OIV[O
[OLPUJYLHZLZVM03HUK;5-ƠPUWYLW\ILY[HSH[OSL[LZ;OLJVUZLX\LUJLVM[OLZL
L]LU[Z^HZ[OLZOPM[VM[OLJ`[VRPULWYVÄSL[V^HYKHWYVPUÅHTTH[VY`Z[H[\Z03HUK
;5-ƠHYLHISL[VPTWHPY[OLNYV^[OOVYTVUL./PUZ\SPUNYV^[OMHJ[VY0.-
H_PZ;OLYLMVYL[OLZLTVKPÄJH[PVUZPUK\JLKI`[YHPUPUNWLYMVYTLKH[HULSP[LSL]LSYPZR
[VULNH[P]LS`HMMLJ[[OLNYV^[OVMMLTHSLJOPSKYLUH[OSL[LZ
07VZ[LY:LZZPVU
BCE 60
Biochimica Cellulare
(J[P]H[PVUI`HYHJOPKVUV`SNS`JLYVSVMWSH[LSL[W4(72
J73(WH[O^H`
:0.569,3364(90(.9(A0(.0(*6)),,590*((5+3,65*050.0<30(5(
+LWHY[TLU[VM,_WLYPTLU[HS4LKPJPUL)PVJOLTPZ[Y`ZLJ[PVU
.LUVH<UP]LYZP[`.LUVH0[HS`
;OL LUKVNLUV\Z JHUUHIPUVPK HYHJOPKVUV`SNS`JLYVS (. PZ KLZJYPILK HZ H
WSH[LSL[HNVUPZ[HISL[VPUK\JLHNNYLNH[PVUHUK[VPUJYLHZLPU[YHJLSS\SHYJHSJP\T0U
[OLWYLZLU[YLWVY[^LOH]LJVUÄYTLK[OLZLKH[HHUKKLTVUZ[YH[LK[OH[[OLPUOPIP[VY
VMW4(72:)HUK[OLPUOPIP[VYVMJ73(TL[HIVSPZT,;@(HMMLJ[IV[O[OLZL
WHYHTL[LYZ;O\Z^LHPTLK[VKLÄUL[OLYVSLVMW4(72J`[VZVSPJWOVZWOVSPWHZL
(J73(WH[O^H`PU(.PUK\JLKO\THUWSH[LSL[HJ[P]H[PVUW4(72HJ[P]H[PVU
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HUKHZHYHJOPKVUPJHJPKYLSLHZLHUK[OYVTIV_HUL)MVYTH[PVU0[^HZZOV^U[OH[
(.PUHKVZLHUK[PTLKLWLUKLU[THUULYHJ[P]H[LZW4(72WLHRPUNH[›4
HM[LYTPUVMPUJ\IH[PVU;OL(.LMMLJ[VUW4(72^HZUV[PTWHPYLKI`HW`YHZL
PUKVTL[OHJPUVY9.+:WLW[PKLI\[P[^HZZPNUPÄJHU[S`YLK\JLKI`:9ZWLJPÄJ
PUOPIP[VY VM [`WL JHUUHIPUVPK YLJLW[VY HUK \UHMMLJ[LK I` [OL ZWLJPÄJ PUOPIP[VY VM
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HUK HJ[P]H[PVU 0U HKKP[PVU :9 Z[YVUNS` PTWHPYLK J73( WOVZWOVY`SH[PVU
HYHJOPKVUPJHJPKYLSLHZLHUK[OYVTIV_HUL)MVYTH[PVU^OLYLHZ:9KPKUV[
JOHUNL[OLZLWHYHTL[LYZ-PUHSS`WSH[LSL[Z[PT\SH[PVU^P[O(.SLK[VHUPUJYLHZLPU
MYLLV_`NLUYHKPJHSZWLJPLZ0UJVUJS\ZPVUKH[HWYV]PKLPUZPNO[PU[V[OLTLJOHUPZTZ
PU]VS]LK PU WSH[LSL[ HJ[P]H[PVU I` (. PUKPJH[PUN [OH[ W4(72J73( WH[O^H`
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07VZ[LY:LZZPVU
BCE 61
Biochimica Cellulare
,_WYLZZPVUVM[OLHU[PHWVW[V[PJWYV[LPU)(.PZHMLH[\YL
VMWHUJYLH[PJHKLUVJHYJPUVTHHUKV]LYL_WYLZZPVUPZ
HZZVJPH[LK^P[OWVVYLYZ\Y]P]HS
96:(;0(3,::(5+9(),9:(50:(4(5;/(;(=(56-9(5*,:*(+(33(
76AA(,30:(4(9.6;+,4(9*67(340,904(9;(
+,3(<9,5A0=05*,5A6-9(5*69,5(;6:*6.5(40.306.06:<Ï
7(3(0(9(--(,3,-65;(5((5+9,( +0:,)(:;0(5670,93<0.0
+65(+,3304(::046+(5+603(90(4,+,4(1(57(<3
+012-9,+,902,>,3305.30,2,+0463(-()06-9(5*,:*6
7,AA03309(--(,3,;<9*64(90(*(;,905(:*(97((3+6
+LWHY[TLU[VM7OHYTHJL\[PJHSHUK)PVTLKPJHS:JPLUJLZ<UP]LYZP[`VM:HSLYUV-PZJPHUV:(0[HS`")06<50=,9:(ZYS
<UP]LYZP[`VM:HSLYUV-PZJPHUV:(0[HS`"(9*5,;*LU[LYMVY(WWSPLK9LZLHYJOVU*HUJLYHUK+LWHY[TLU[VM7H[OVSVN`
HUK+PHNUVZ[PJZ/VZWP[HSHUK<UP]LYZP[`;Y\Z[=LYVUH0[HS`"+LWHY[TLU[VM:\YNLY`*HZH:VSSPL]VKLSSH:VMMLYLUaH¹/VZWP[HS
09**::HU.PV]HUUP9V[VUKV0[HS`"+LWHY[TLU[VM3PMLHUK9LWYVK\J[PVU:JPLUJLZ)PVJOLTPZ[Y`:LJ[PVU<UP]LYZP[`VM
=LYVUH=LYVUH0[HS`"+LWHY[TLU[VM)PVTLKPJHS:JPLUJLZ<UP]LYZP[`¸.K»(UU\UaPV¹*OPL[P7LZJHYHHUK-VUKHaPVUL¸.
+»(UU\UaPV*LU[YV:[\KPZ\SS»0U]LJJOPHTLU[V*L:0*OPL[P0[HS`"<6*(UH[VTPH7H[VSVNPJH0Z[P[\[V5HaPVUHSL;\TVYP
-VUKHaPVUL.7HZJHSL5HWSLZ0[HS`"(IKVTPUHS:\YNLY`<UP[0Z[P[\[V5HaPVUHSL;\TVYP-VUKHaPVUL.7HZJHSL5HWSLZ0[HS`"
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BCE 62
Biochimica Cellulare
-HPSPUNJHYKPVT`VJ`[LZYLSLHZL)(.[OH[HJ[P]H[LZ
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07VZ[LY:LZZPVU
BCE 63
Biochimica Cellulare
(J[P]H[PVUVM7SH[LSL[(J[P]H[PUN-HJ[VY9LJLW[VYHUK:09;
ZPNUHSPUNPULUKV[OLSPHSWYVNLUP[VYJLSSZ
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BCE 64
Biochimica Cellulare
7YV[LVTPJHUHS`ZPZVMTVSLJ\SHYYLZWVUZLZ[VHSNHS[V_PUZ
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07VZ[LY:LZZPVU
BCE 65
Biochimica Cellulare
;OLTP[VJOVUKYPHSJP[YH[LJHYYPLY*0*YLN\SH[LZPUZ\SPU
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07VZ[LY:LZZPVU
BCE 66
Biochimica Cellulare
;OL,UKVJHUUHIPUVPK:`Z[LTPU:[LT*LSSZMYVT[OL
(TUPV[PJ-S\PK
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9LMLYLUJLZ
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Z[LTJLSSZ:[LT*LSSZ+L]! 07VZ[LY:LZZPVU
BCE 67
Biochimica Cellulare
;YHUZJYPW[PVUHSYLN\SH[PVUVM[OLO\THU-79(3?NLUL!
L]PKLUJLVMHOLYP[HISLNLUL[PJ]HYPHU[[OH[PTWHPYZ
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4(;;6:*06+64,50*6+,3<*((5;65,33(-9(5*/0:(9(
.(;;(=(3,5;05(7(963(90(3,::(5+96>,9)(16:Ï7()36
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+PWHY[PTLU[VKP:JPLUaL6YHSP5HUVL)PV[LJUVSVNPH
*LU[YVKP,JJLSSLUaH:JPLUaLKLSS»0U]LJJOPHTLU[V*OPL[P
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09**:4PSHUV
+PWHY[PTLU[V:JPLUaL-HYTHJVSVNPJOL<UP]LYZP[nKP4PSHUV
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/LYL^LTHWWLKHIWYLNPVU^P[OZ[YVUNWYVTV[LYHJ[P]P[`IW\WZ[YLHT[OL
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LUOHUJLKI`3?([OLSPNHUK^P[O[OLOPNOLZ[HMÄUP[`MVY[OPZYLJLW[VY\UJV]LYPUN
H WVZP[P]L MLLKIHJR SVVW VM YLZVS\[PVU K\YPUN PUÅHTTH[PVU >L HSZV PKLU[PÄLK H
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THKL [OL JVYL WYVTV[LY \UYLZWVUZP]L [V 3?( PU ]P[YV 4VYLV]LY UL\[YVWOPSZ MYVT
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]VS\U[LLYZL_WYLZZPUN[OL^PSK[`WLHSSLSL;VNL[OLY[OLZLYLZ\S[Z\UYH]LS[OL-79
(3?[YHUZJYPW[PVUTHJOPULY`HUKWYV]PKL[OLÄYZ[L]PKLUJLVMNLUL[PJ]HYPHU[Z^P[OPU
[OL-79(3?WYVTV[LY^OPJOPTWHPYNLULL_WYLZZPVUH[[OL[YHUZJYPW[PVUHSSL]LS0U
SPNO[VM[OLL]PKLUJLPU]P]V[OH[-79(3?L_WYLZZPVUSL]LSZYLWYLZLU[WLYZLHRL`
WYVYLZVS\[PVUTLJOHUPZTV\YKH[HTH`VWLU[OL^H`MVY[OLKPZJV]LY`VMHUV]LS
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07VZ[LY:LZZPVU
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BNU 1
Biochimica della Nutrizione
:(--965!(76;,5;0(3*(5+0+(;,-69(56=,3
(5;0*(5*,9+9<.(.(05:;/<4(5796:;(;,
*(5*,9
4(5*050( 30AA0(9+,:0465,(4(996**,33(*,
.9(=05(.3;(;65,*-,:;<**0(*+»(3,::(5+96(4
3PML/LHS[OHUK,U]PYVUTLU[HS:JPLUJLZ<UP]LYZP[`VM3»(X\PSH"
*SPUPJHSHUK)PV[LJOUVSVN`:JPLUJLZ<UP]LYZP[`VM3»(X\PSH"
*OHTILYVM*VTTLYJL(NLUaPHWLYSV:]PS\WWV"
*HUJLYVMWYVZ[H[LPZ[OLTVZ[JVTTVUS`KPHNUVZLKZVSPKTHSPNUHUJ`PUTLUSP]PUNPU
KL]LSVWLKJV\U[YPLZ:HMMYVUJVUZPZ[ZVMKYPLKZ[PNTHZVM*YVJ\ZZH[P]\Z3ÅV^LYHUK
PZRUV^UMVYHU[PJHUJLYWYVWLY[PLZ0U[OPZZ[\K`^LPU]LZ[PNH[LK[OLHU[P[\TVYLMMLJ[Z
HUK\UKLYS`PUNTLJOHUPZTZVM:HMMYVUL_[YHJ[:,HUKP[ZTHQVYJVUZ[P[\LU[JYVJPU
HNHPUZ[WYVZ[H[LJHUJLYPU]P]V(X\LV\ZHUKL[OHUVSPJ:HMMYVUL_[YHJ[:,NTS
^LYLW\YPÄLKHUKJOHYHJ[LYPaLKI`/73*0UVYKLY[VHZZLZ[OLHJ[P]P[`VM:HMMYVU
L_[YHJ[ HUK JYVJPU PU ]P]V YHUKVTPaLK NYV\WZ VM Y]ILHYPUN TPJL ^LYL [YLH[LK
VYHSS`^P[O:,HUKJYVJPUH[TN2NHUKTN2NIVK`^LPNO[YLZWLJ[P]LS`MVY
KH`Z0UWYL]PV\ZPU]P[YVZ[\KPLZ^LKLTVUZ[YH[LK[OH[:,HUKJYVJPUHS[LYLKJLSS
J`JSL WYVNYLZZPVU HUK PUK\JLK HWVW[VZPZ PU H [PTL HUK JVUJLU[YH[PVUKLWLUKLU[
THUULY9LZ\S[ZMYVT[OPZPU]P]VZ[\K`ZOV^LK[OH[:,HUKJYVJPUPUOPIP[LK_LUVNYHM[
NYV^[O VM LZ[HISPZOLK Y] [\TVYZ! H ZPNUPÄJHU[ YLK\J[PVU PU ZVSPK [\TVY ]VS\TL
^HZ MV\UK PU JYVJPU[YLH[LK TPJL ^OLU JVTWHYLK ^P[O JVU[YVS HUPTHSZ ]Z
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THZZ^HZVIZLY]LKI`[YLH[TLU[^P[O:,]ZJVU[YVS TNBTLHU
:+D]ZTNW$0UJYVJPU[YLH[LKTPJLHZPNUPÄJH[P]LYLK\J[PVU
VMHUNPVNLULZPZ^HZHSZVVIZLY]LK-\Y[OLYTVYL:,JV\U[LYHJ[LKV_PKH[P]LZ[YLZZ
I` PUJYLHZPUN [OL HJ[P]P[` VM Z\WLYV_PKL KPZT\[HZL JH[HSHZL HUK NS\[H[OPVUL:
[YHUZMLYHZLSL]LSZ;HRLU[VNL[OLY[OLZLÄUKPUNZZ\NNLZ[[OH[ZHMMYVUHUKJYVJPUL_LY[
H ZPNUPÄJHU[ JOLTPVWYL]LU[P]L LMMLJ[ HNHPUZ[ WYVZ[H[L JHUJLY [OYV\NO PUOPIP[PVU VM
NYV^[OVMWYVZ[H[L[PZZ\LHUKWYV[LJ[WYVZ[H[LMYVTJHUJLY]PHTVK\SH[PUNV_PKH[P]L
Z[YLZZ
07VZ[LY:LZZPVU
BNU 2
Biochimica della Nutrizione
9VSLVMƠ;VJVWOLY`SWOVZWOH[LVU[OLL_WYLZZPVUVM
:LZ[YPUPUWYLHKPWVJ`[LZ50/;3
+<.63(<9(309(5.04,3(50(+(*/±4(905(A05..1,(54(9*
(5+(AA0(5.,36
3HIVYH[VYPVKP)PVJOPTPJH*OPTPJHL5\[YPaPVUL*LU[YVKP9PJLYJH0U[LNYH[V*09
<UP]LYZP[n*HTW\Z)PV4LKPJVKP9VTH9VTH0[HS`
=HZJ\SHY)PVSVN`3HIVYH[VY`14<:+(/59*(H[;\M[Z<UP]LYZP[`)VZ[VU<:(
:LZ[YPUPZHMHTPS`VMWYV[LPUZ^OVZLTHPUYVSLPZ[OLZ\WWYLZZPVUVMSPWPKZHJJ\T\SH[PVU
[OLWYL]LU[PVUVMJHYKPHJTHSM\UJ[PVUZHUK[OLWYV[LJ[PVUVMT\ZJSLMYVTHNLYLSH[LK
KLNLULYH[PVU
:LZ[YPUHJJ\T\SH[LZPUJLSSZPUYLZWVUZL[VZ[YLZZMHJ[VYZWV[LU[PH[L[OLHJ[P]P[`VM
(472HUKPUOPIP[[OLHJ[P]H[PVUVM[HYNL[NLULVMYHWHT`JPUT;69,_WYLZZPVUVM
:LZ[YPUPZPUJYLHZLKI`[OLHJ[P]P[`VMWHUKKLJYLHZLKI`[OLPUZ\SPU(2;ZPNUHSSPUN
WH[O^H`
Ơ;VJVWOLY`SWOVZWOH[LƠ;7PZ[OLWOVZWOVY`SH[LKKLYP]H[LVMƠ[VJVWOLYVSƠ;7
OHZILLUYLWVY[LK[VHJ[HZHYLN\SH[VYVMNLULL_WYLZZPVUHUKJLSSZPNUHS[YHUZK\J[PVU
7YLSPTPUHY`L_WLYPTLU[ZZOV^LK[OH[[OLHKKP[PVUVMƠò;7I\[UV[Ơò[VJVWOLYVS
[V50/;3WYLHKPWVJ`[LZ[YHUZJYPW[PVUHSS`HJ[P]H[LZHZL[VMNLULZ!;9);YPIISLZ
/VTVSVN *-> HU[P.7YV[LPU*V\WSLK 9LJLW[VY 0UK\JLK 7YV[LPU :LZ[YPU
HUK0UZ\SPU0UK\JLK.LUL05:0.WV[LU[PHSS`WYL]LU[PUNMH[HJJ\T\SH[PVUPU[OLZL
JLSSZƠ;7JV\SK[OLYLMVYLOH]LHYVSLPU[OLWYL]LU[PVUVMSPWPKZHJJ\T\SH[PVUK\YPUN
HKPWVJ`[LZKPMMLYLU[PH[PVU
>LYLWVY[OLYL[OLLMMLJ[VMƠ[VJVWOLYVSHUKƠò;7VU[OLL_WYLZZPVUVM:LZ[YPUPU
50/;3 WYLHKPWVJ`[LZ *LSSZ ^LYL [YLH[LK ^P[O Ơ[VJVWOLYVS HUK Ơò;7 H[ [OL
JVUJLU[YH[PVUZVMHUK›4MVYHUKO
:LZ[YPUNLULHUKWYV[LPUL_WYLZZPVU^HZPU]LZ[PNH[LKI`9;7*9HUK^LZ[LYUISV[
6\YYLZ\S[ZZOV^[OH[HM[LYO[YLH[TLU[VMJLSSZ^P[OƠò;7›4[OLL_WYLZZPVU
VMIV[O:LZ[YPUT95(HUK:LZ[YPUWYV[LPU^LYLZPNUPÄJHU[S`PUJYLHZLKJVTWHYLK[V
[OLJLSSZ[YLH[LK^P[OƠ[VJVWOLYVSPU[OLZHTLJVUKP[PVUZ
;OLYLN\SH[PVUVM:LZ[YPUL_WYLZZPVUJV\SKPUKPJH[LHYVSLMVYƠ;7PU[OLTVK\SH[PVU
VMHKPWVZL[PZZ\LTL[HIVSPZTHUKLULYNL[PJOVTLVZ[HZPZ
-\Y[OLYPU]LZ[PNH[PVU^PSSILULJLZZHY`[VJSHYPM`[OLYVSLVMƠ;7VU[OLYLN\SH[PVUVM
NLULL_WYLZZPVUPUHKPWVJ`[LZILMVYLHUKK\YPUNKPMMLYLU[PH[PVU
(aaP(4VSLJ\SHYTLJOHUPZTVMHSWOH[VJVWOLYVSHJ[PVU-YLLYHKPJHSIPVSVN`TLKPJPUL
07VZ[LY:LZZPVU
BNU 3
Biochimica della Nutrizione
9LKV_PTIHSHUJLHJ[P]H[LZ7.*ƠTLKPH[LK
TP[VJOVUKYPHSHU[PV_PKHU[YLZWVUZLK\YPUNTL[HIVSPJZ[YLZZ
(8<03(562(;0()(3+,330:(9(7(.30,0),(;90*,
*0906364(90(96:(
+LW[VM)PVSVN`<UP]LYZP[`VM9VTL¸;VY=LYNH[H¹
=PHKLSSH9PJLYJH:JPLU[PÄJH9VTL0[HS`
09**::HU9HMMHLSL3H7PZHUH=PHKP=HS*HUU\[H 9VTL0[HS`
.S\[H[OPVUL .:/ PZ [OL TVZ[ HI\UKHU[ SV^ TVSLJ\SHY ^LPNO[ [OPVS PU THTTHSPHU
JLSSZHUKHJ[ZHZ[OLTHQVYJLSS\SHYUVULUa`TH[PJHU[PV_PKHU[.:/HSZVOHZHWP]V[HS
M\UJ[PVU PU I\MMLYPUN [OL PU[YHJLSS\SHY WYVK\J[PVU VM UP[YPJ V_PKL 56 WHY[PJ\SHYS` PU
IYHPU^OLYLPU56HJ[ZHZHUPTWVY[HU[UL\YVTVK\SH[VYHUKUL\YV[YHUZTP[[LY>LOH]L
MV\UK[OH[U\[YPLU[SPTP[H[PVUHKPL[HY`YLNPTLU[OH[PZNLULYHSS`SPURLK[VSVUNL]P[`
PZHZZVJPH[LK^P[OHKLJSPULVM.:/SL]LSPUTV\ZLIYHPU;OPZL]LU[PZ[OLWYPT\T
TV]LUZ VM 56TLKPH[LK PUK\J[PVU VM [OL [YHUZJYPW[PVUHS JVHJ[P]H[VY WLYV_PZVTL
WYVSPMLYH[VYHJ[P]H[LKYLJLW[VYƢJVHJ[P]H[VYƠ77(9.*(VY7.*Ơ]PHHUV]LS
WH[O^H`PU]VS]PUNW>LOH]LKLTVUZ[YH[LK[OH[Z\JOWH[O^H`J\STPUH[LZPU[OL
LUOHUJLTLU[VM[OLTP[VJOVUKYPHSHU[PV_PKHU[YLZWVUZLPL:6+PUJYLHZL[OYV\NO
[OLU\JSLHYMHJ[VYLY`[OYVPKKLYP]LKSPRL5-,34VYLV]LY[OLWOHYTHJOVSVNPJHS
KLWSL[PVUVM.:/YLZ\S[ZPU[OLHJ[P]H[PVUVMZ\JOHU[PV_PKHU[WH[O^H`PU[OLIYHPU
6]LYHSSV\YYLZ\S[ZZ\NNLZ[[OH[[OLKLJSPULVM.:/SL]LSVJJ\YYPUN^P[OUVYTHSHNLPUN
HUK\WVUU\[YPLU[SPTP[H[PVUJV\SKILHILULÄJPHSYH[OLY[OHUHKL[YPTLU[HSL]LU[HUK
TH`ZLY]LHZHTVSLJ\SHYZPNUHSMVYHKHW[H[PVU[VLU]PYVUTLU[HSWH[OVSVNPJHSZ[YLZZ
HUKWVZZPIS`PU[OLL_[LUZPVUVMSPMLZWHU
;OPZ^VYR^HZWHY[PHSS`Z\WWVY[LKI`-VUKHaPVUL9VTH.YHU[HUKI`40<9
07VZ[LY:LZZPVU
BNU 4
Biochimica della Nutrizione
+/(46+0-0,:3070+9(-;:;9<*;<9,(5+-<5*;065
054+(4))9,(:;*(5*,9*,33:
*69:,;;67(63(((40+03,;;(465;69-(56.0.3063(
*9,465((5+9,(+6.30(:03=0(490AA6(5.,3(4
+PWHY[PTLU[VKP:JPLUaL-HYTHJVSVNPJOLL)PVTVSLJVSHYP
<UP]LYZP[nKLNSP:[\KPKP4PSHUV
+PWHY[PTLU[VKP)PV[LJUVSVNPLL)PVZJPLUaL<UP]LYZP[nKLNSP:[\KPKP4PSHUV)PJVJJH
0U]P]VHUKPU]P[YVZ[\KPLZZ\NNLZ[[OH[VTLNHWVSP\UZH[\YH[LKMH[[`HJPKZU7<-(
JHUILJHUJLYJOLTVWYL]LU[P]LJOLTVZ\WWYLZZP]LHUKH\_PSPHY`HNLU[ZMVYJHUJLY
[OLYHW`57<-(SPRLKVJVZHL_HLUVPJHJPK+/(JV\SKHS[LY[\TVYNYV^[OI`
PUÅ\LUJPUNJLSSYLWSPJH[PVUPU[LYMLYPUN^P[OJLSSJ`JSLJVTWVULU[ZVYPUJYLHZPUNJLSS
KLH[OLP[OLYI`PUK\JPUNULJYVZPZVYHWVW[VZPZ
>LOH]LVIZLY]LK[OH[+/(PUK\JLZPU4+(4)JLSSZHWVW[VZPZHUKYLK\J[PVU
VMJLSS]PHIPSP[`4VYLV]LY+/(ZPNUPÄJHU[S`YLK\JLZ[OL,.-9SL]LSHUKJVTWSL[LS`
PUOPIP[Z,.-9HJ[P]H[PVU
/V^L]LY[OLTLJOHUPZTI`^OPJOU7<-(PUOPIP[IYLHZ[JHUJLYNYV^[OPZUV[`L[
^LSS\UKLYZ[VVK0[^HZZ\NNLZ[LK[OH[[OLZLMH[[`HJPKZTPNO[JOHUNLJLSSTLTIYHUL
Å\PKP[`HUKZ[Y\J[\YLLZWLJPHSS`VMSPWPKYHM[Z
6\YKH[HVI[HPULKI`/73*.*HUHS`ZPZKLTVUZ[YH[L[OH[+/(PZPUJVYWVYH[LKHUK
TL[HIVSPaLKPU4+(4)TLTIYHUL0UWHY[PJ\SHY+/(PZPUJVYWVYH[LKPUIYLHZ[
JHUJLYSPWPKYHM[Z^P[OKPMMLYLU[ZWLJPÄJP[`MVY[OLWOVZWOVSPWPKTVPL[`HUKPUK\JLZH
YLK\J[PVUVMJOVSLZ[LYVSHUKZWOPUNVT`LSPUJVU[LU[PUKPJH[PUNHWVZZPISLJOHUNLPU
YHM[VYNHUPaH[PVU*OHUNLZPU[OLSPWPKYHM[Z[Y\J[\YL^LYLHSZVHUHS`aLK^P[OH[VTPJ
MVYJLTPJYVZJVW`(-4HUK-V\YPLY[YHUZMVYTPUMYHYLKZWLJ[YVZJVW`-;09;OL(-4
HUHS`ZPZVMIYLHZ[JHUJLYSPWPKYHM[ZPUKPJH[LZHM[LY+/(PUJVYWVYH[PVUHYLK\J[PVUVM
TPJYVKVTHPUZU\TILY4VYLV]LY7<-([YLH[LKYHM[ZHYLKPTLUZPVUHSS`KPMMLYLU[MYVT
JVU[YVSYHM[Z
;OLU^OLU+/(PZPUJVYWVYH[LKPU[VJLSSTLTIYHULP[TH`PUK\JL[OLMVYTH[PVUVM
¸KLJS\Z[LYLKYHM[Z¹^P[OYLVYNHUPaH[PVUVMSPWPKZHUKWYV[LPUZ[OH[TH`NYLH[S`PUÅ\LUJL
ZPNUHS[YHUZK\J[PVU
*HS]PLSSV.L[HS5\[Y*HUJLY *VYZL[[V7(L[HS*LSS)PVJOLT)PVWO`Z
07VZ[LY:LZZPVU
BNU 5
Biochimica della Nutrizione
)Y\[PLYPKPUHUK4LSP[PKPU!UH[\YHSWYVK\J[Z^P[OWV[LU[PHS
O`WVJOVSLZ[LYVSLTPJHJ[P]P[`
4(9;,336,4(5<,3(+0+655(3,65(9+60(*67,;;(+64,50*6
*(77,336(55(9.(33<**0.0:,3+(:05+65(.06=(550(5+
+63*,=05*,5A(
+LWHY[TLU[VM*OLTPZ[Y`HUK+LWHY[TLU[VM7OHYTHJV)PVSVN`
<UP]LYZP[`VM*HSHIYPH(YJH]HJH[HKP9LUKL*:0[HS`
)HJRNYV\UK!:[H[PUZHYL[OLWOHYTHJVSVNPJHSPUOPIP[VYZVMJOVSLZ[LYVSIPVZ`U[OLZPZ
HJ[PUNVURL`LUa`TL/`KYV_`TL[O`SNS\[HY`S*V(YLK\J[HZL/4.9;OL`HYL
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07VZ[LY:LZZPVU
BNU 6
Biochimica della Nutrizione
/LZWLYL[PUHUK]P[HTPU*HU[PPUÅHTTH[VY`WYVWLY[PLZPU
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VYHUNLQ\PJLKLTVUZ[YH[LKP[ZILULÄJPHSLMMLJ[VUJHYKPV]HZJ\SHYYPZRMHJ[VYZZ\JOHZ
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BNU 7
Biochimica della Nutrizione
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07VZ[LY:LZZPVU
BNU 8
Biochimica della Nutrizione
:OVY[[LYTLMMLJ[ZVMVSP]LVPSWVS`WOLUVSZVUSPWPK
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PUJ\S[\YLKYH[OLWH[VJ`[LZ,UKVJYPUVSVN` ! 07VZ[LY:LZZPVU
BNU 9
Biochimica della Nutrizione
7VTLNYHUH[LLSSHNP[HUUPUZPUOPIP[PU]P[YVJHYIVO`KYH[L
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WVS`WOLUVSYPJOL_[YHJ[779,;OL779,VI[HPULK\ZPUN*ZVSPKWOHZLL_[YHJ[PVU
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07VZ[LY:LZZPVU
BNU 10
Biochimica della Nutrizione
(U[PV_PKHU[HJ[P]P[`VMPU]P[YVKPNLZ[LKIV]PULTPSR
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ZOV^LK[OH[[^VWLW[PKLZHYL[OLTHQVYJVU[YPI\[VY[V[OLHU[PV_PKHU[HJ[P]P[`VM[OPZ
MYHJ[PVU;OL[^VWLW[PKLZ^LYLPKLU[PÄLKI`4:4:6\YKH[HZ\NNLZ[[OH[KPNLZ[PVU
VMTPSRPUJYLHZLK[OLHU[PV_PKHU[HJ[P]P[`WV[LU[PHSS`YLK\JPUN[OLV_PKH[P]LZ[YLZZPU
[OLNHZ[YVPU[LZ[PUHS[YHJ[
07VZ[LY:LZZPVU
BNU 11
Biochimica della Nutrizione
)VULTHZZKLUZP[`PZPUKLWLUKLU[S`HZZVJPH[LK^P[O
V_PKH[P]LZ[YLZZPUWVZ[I\[UV[PUWYLTLUVWH\ZHS^VTLU
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:;,-(50(-,99(AA050(3,::(5+9(*(7(;;0,590*(-03(
*(:(30--(5;050.-=05*,5A63(5A(9(3,64(::(90
(5+*(936*,9=,33(;0
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<UP]LYZP[`VM-LYYHYH-LYYHYH0[HS`
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+LWHY[TLU[VM)PVTLKPJHS:JPLUJLZHUK(K]HUJLK;OLYHWPLZ
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7HYHTL[LYZVMIVULTPULYHSKLUZP[`)4+H[MLTVYHSULJRHUKS\TIHYZWPUL^LYL
HSZVHZZLZZLK9LZ\S[Z!7LHYZVU»ZJVYYLSH[PVUHUHS`ZPZ\U]LPSLK[OH[ZWPUHS)4+^HZ
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PU^VTLUPUYLWYVK\J[P]LHNLY$W$5V[L^VY[O`T\S[P]HYPH[LHUHS`ZPZ
ZOV^LK[OH[[OLHZZVJPH[PVUMV\UKPUWVZ[TLUVWH\ZHS^VTLUYL[HPULKZPNUPÄJHUJL
HM[LY[HRPUNPU[VJVUZPKLYH[PVUWV[LU[PHSJVUMV\UKPUNMHJ[VYZW$*VUJS\ZPVU!
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07VZ[LY:LZZPVU
BNU 12
Biochimica della Nutrizione
4LJOHUPZTZILOPUKZ\SMVYHWOHULJHYKPVWYV[LJ[P]LLMMLJ[
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[OYV\NOUVULUa`TH[PJNS`JH[PVUHUKHJJ\T\SH[L\UKLYKPHIL[PJJVUKP[PVU;OLTVZ[
PU]LZ[PNH[LK(.,ZWYLJ\YZVYPZTL[O`SNS`V_HSHTL[HIVSPJJVTWV\UKMVYTLKTHPUS`
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[OYV\NO[OL2LHW¶5YMWH[O^H`
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>LVIZLY]LK[OH[Z\SMVYHWOHUL[YLH[TLU[ZPNUPÄJHU[S`JV\U[LYHJ[LKJLSSKLH[OHUK
HWVW[VZPZPUK\JLKI`TL[O`SNS`V_HSPUOPIP[LKTL[O`SNS`V_HSPUK\JLKHJ[P]H[PVUVM[OL
WYVHWVW[V[PJRPUHZLZ152HUKW4VYLV]LYMVY[OLÄYZ[[PTL^LKLTVUZ[YH[LK[OH[
Z\SMVYHWOHULPZHISL[VZPNUPÄJHU[S`PUJYLHZL.36WYV[LPUL_WYLZZPVUHUKHJ[P]P[`
;OLZL ÄUKPUNZ Z\NNLZ[ [OH[ [HYNL[PUN [OL NS`V_HSHZL Z`Z[LT TPNO[ IL H WYVTPZPUN
[OLYHWL\[PJZ[YH[LN`[VJV\U[LYHJ[JHYKPV]HZJ\SHYKPZLHZLZVMKPHIL[PJWH[PLU[Z
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07VZ[LY:LZZPVU
BNU 13
Biochimica della Nutrizione
,MMLJ[VMWVS`\UZH[\YH[LKMH[[`HJPKZVUSPWVNLUPJMHJ[VYZ
*(7<;64(90,33(+,96:(4(90(*(;,905(9,:*0.56;(50(
A097630/@3+,=(::(336(5;6506+,;644(:05<5A0(;05(
;69056.(,;(56;,**,4(906-,30*,
+LWHY[TLU[VM)PVTLKPJHSHUK7OHYTHJL\[PJHS:JPLUJL
<UP]LYZP[`VM:HSLYUV=PH7VU[L+VU4LSPSSV-PZJPHUV:H0[HS`
7VS`\UZH[\YH[LK MH[[` HJPKZ 7<-( HYL U\[YPLU[Z ^P[O YLSL]HU[ IPVSVNPJHS HJ[P]P[PLZ
HUKWYL]LU[P]LLMMLJ[ZVMJHYKPV]HZJ\SHYHUKULVWSHZ[PJKPZLHZLZHUK[OLYLMVYLP[PZ
PTWVY[HU[ [V JOHYHJ[LYPaL [OL TVSLJ\SHY WYVJLZZLZ PU]VS]LK PU [OLPY HJ[PVU 6U [OPZ
W\YWVZL^LL]HS\H[LK[OLLMMLJ[VM7<-(VU:[LHYV`S*V(+LZH[\YHZL:*+[OL
YH[LSPTP[PUNLUa`TLPU7<-(IPVZ`U[OLZPZHUKVU:[LYVS9LN\SH[VY`,SLTLU[)PUKPUN
7YV[LPU :9,)7J H [YHUZJYPW[PVU MHJ[VY ^OPJO KPYLJ[S` TVK\SH[L P[Z L_WYLZZPVU
:*+PU[YVK\JPUNHKV\ISLIVUKPUMH[[`HJ`S*V(Z\IZ[YH[LZPZVMWHY[PJ\SHYPU[LYLZ[
ZPUJLHS[LYH[PVUZVMP[ZHJ[P]P[`ZLLT[VILPU]VS]LKPUZL]LYHSK`ZSPWPKLTPJJVUKP[PVUZ
*VUZPKLYPUN [OH[ 3P]LY ? 9LJLW[VY 3?9 HJ[P]H[PVU PUK\JLZ SPWVNLUPJ WH[O^H`Z
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YLNPVU^OPSLHZH[\YH[LKZ[LHYPJHJPK:(KPKUV[NP]LHU`LMMLJ[;VM\Y[OLY\UKLYZ[HUK
PM 7<-( WYVK\JL [OLZL LMMLJ[Z [OYV\NO H KPYLJ[ PU[LYHJ[PVU ^P[O 3?9 ^L OH]L HSZV
JHYYPLK V\[ H Z\YMHJL WSHZTVU YLZVUHUJL :79 HUHS`ZPZ ^OPJO ZOV^LK H KPYLJ[
IPUKPUN VM +/( ,7( HUK (( VU 3?9 [YHUZJYPW[PVU MHJ[VY ^OPSL UV PU[LYHJ[PVU ^HZ
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HUHS`aLK7<-(ZLSLJ[P]LS`KV^UYLN\SH[LSPWVNLUPJWH[O^H`Z[OYV\NO3?9PUOPIP[PVU
HUKJVUZLX\LU[:9,)7JHUK:*+L_WYLZZPVUKV^UYLN\SH[PVU
07VZ[LY:LZZPVU
BNU 14
Biochimica della Nutrizione
,MMLJ[VMZ[L]PVSNS`JVZPKLZVUNS\JVZL[YHUZWVY[HJ[P]P[`PU
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<UP]LYZP[nKP)VSVNUH0[HS`
6]LY^LPNO[ VY VILZL WLVWSL PZ JVU[PU\V\ZS` YPZPUN PU PUK\Z[YPHSPaLK UH[PVUZ ;OPZ
WLVWSLPZZ\IQLJ[LK[VHUPUJYLHZLKYPZRMVYHU\TILYVMKLSL[LYPV\ZOLHS[OJVUKP[PVUZ
PUJS\KPUN[`WLKPHIL[LZOLHY[KPZLHZLZHUKJHUJLY;`WLKPHIL[LZPZHZZVJPH[LK
[VPTWHPYLKPUZ\SPUZLJYL[PVUHUKPUZ\SPUYLZPZ[HUJLPUWLYPWOLYHS[PZZ\LZSLHKPUN[V
O`WVPUZ\SPULTPH HUK O`WLYNS`JLTPH ;OLYLMVYL UVU [V_PJ ^LPNO[ JVU[YVSSPUN Z\NHY
Z\IZ[P[\[LZHZ^LSSHZUL^LYHUKZHMLYHNLU[ZMVY[YLH[TLU[VM[`WLKPHIL[LZHYL
ULLKLK :[L]PVS NS`JVZPKLZ UVU JHSVYPJ UH[\YHS Z^LL[LULYZ L_[YHJ[LK MYVT :[L]PH
YLIH\KPHUHOH]LILLU\ZLKHZHU[PO`WLYNS`JLTPJHNLU[ZI\[SP[[SLPZRUV^UHIV\[
[OLPYTLJOHUPZTVMHJ[PVUH[TVSLJ\SHYSL]LSLZWLJPHSS`VU[OLPYWV[LU[PHSPU[LYHJ[PVU
^P[OJLSS\SHYNS\JVZL[YHUZWVY[Z`Z[LT
>LZ[\KPLKNS\JVZL[YHUZWVY[HJ[P]P[`PUKPMMLYLU[JLSS[`WLZL_WYLZZPUN.3<;[OL
IHZHSNS\JVZL[YHUZWVY[LYHUK.3<;[OLPUZ\SPUZLUZP[P]LVUL0[OHZILLURUV^U
[OH[ .3<; SHYNLS` JVUÄULK PU PU[YHJLSS\SHY Z[VYHNL ZP[L ILJVTLZ YLJY\P[LK [V [OL
JLSS Z\YMHJL \UKLY [OL PUÅ\LUJL VM PUZ\SPU HUK V[OLY Z[PT\SP I\[ HSZV [OL PZVMVYT
.3<;[YHUZSVJH[LZ[V[OLWSHZTHTLTIYHULPUYLZWVUZL[VZL]LYHSZ[PT\SP;O\Z
^L PU]LZ[PNH[LK ^OL[OLY YLIH\KPVZPKL ( HUK Z[L]PVZPKL JV\SK IL HISL [V TVK\SH[L
NS\JVZL\W[HRLVM[OLZLJLSSZHUKWVZZPIS`[VPUÅ\LUJL[OLPU[YHJLSS\SHYKPZ[YPI\[PVUVM
.3<;PZVMVYTZ
:\WWVY[LKI`4PUPZ[Y`VM,JVUVTPJ+L]LSVWTLU[7YVQLJ[4HKLPU0[HS`V]LY
5HYPZZHYH3L[HS4L[HIVSPZTWW
/YLSPH:L[HS)PVJOPTPJHL[)PVWO`ZPJH(J[HWW
4HYHSKP;L[HS(U[PV_PKHU[Z9LKV_:PNUHSPUN WW
07VZ[LY:LZZPVU
BNU 15
Biochimica della Nutrizione
4VKPÄJH[PVUZVMSPWV_`NLUHZLMYVTNLUL[PJHSS`TVKPÄLK
ZV`ILHU
(5.,3<**0*36;03+,):()(;<**0(55(3(<9(
+0-9(5*,:*6(5+9,(*0()(;;003(90(+(05,:,,590*6
4(**(9965,4(<96
+LWHY[PTLU[VKP:JPLUaL)PVTLKPJOL<UP]LYZP[nKP;LYHTV;LYHTV0[HSPH
*LU[YVKP9LMLYLUaH5HaPVUHSLWLYSH9PJLYJHKP6.4*96.49VTH0[HSPH
*LU[YV,\YVWLVKP9PJLYJHZ\S*LY]LSSV-VUKHaPVUL¸:HU[H3\JPH¹9VTH
,X\HSS`ZLUPVYH\[OVYZ
0U<:(*HUHKHHUKZL]LYHSKL]LSVWPUNJV\U[YPLZLN*OPUH)YHaPS(YNLU[PUH
[OL \ZL VM NLUL[PJHSS` TVKPÄLK WSHU[Z .47Z PU HNYPJ\S[\YL PZ YHWPKS` NYV^PUN HUK
L_WHUKPUN0U[OPZJV\U[YPLZ.4ZV`ILHUZ465TVKPÄLKMVYYLZPZ[HUJL[V
[OLOLYIPJPKLNS`WOVZH[LPZVULVM[OLTVZ[J\S[P]H[LK.47ZKH[HMYVT0:(((
;OL,\YVWLHU-VVK:HML[`(NLUJ`,-:(PUP[ZN\PKLSPULZLTWOHZPaLZ[OLPTWVY[HUJL
VMPKLU[PM`PUNPU.47Z[OLWYLZLUJLVMM\UJ[PVUHSJOHUNLZPUWYV[LPUZPU]VS]LKPUWSHU[
KLMLUZLTLJOHUPZTZHUKHS[LYH[PVUZPUSPWPKWYVÄSLZ(PTVM[OPZ^VYR^HZ[V]LYPM`
WVZZPISLHS[LYH[PVUZVMHUPTWVY[HU[LUa`TLPU]VS]LKPUWSHU[KLMLUZLTLJOHUPZTZ!
SPWV_`NLUHZL36?>LZOV^THQVYZ[Y\J[\YHSHUKM\UJ[PVUHSJOHUNLZVM36?
PU.4ZV`ILHUZ^P[OYLZWLJ[[V^PSK[`WLJV\U[LYWHY[Z0UWHY[PJ\SHY^LKLTVUZ[YH[L
H Z[YVUNS` YLK\JLK L_WYLZZPVU HUK JH[HS`[PJ LMÄJPLUJ` VM 36? PU 465
ZV`ILHUHUKHYLK\J[PVUVMPYVUPUP[ZHJ[P]LZP[L;OLZL36?HS[LYH[PVUZ^LYL
UV[WHYHSSLSLKI`ZPNUPÄJHU[JOHUNLZPUPYVUIPVH]HPSHIPSP[`IL[^LLU[OL.4ZV`ILHUZ
HUKJVU]LU[PVUHSJV\U[LYWHY[ZHUKSLK[VHZPNUPÄJHU[KLJYLHZLVMWLYV_PKPaLKSPWPKZ
PUZLLKZ0U[LYLZ[PUNS`^LYLJLU[S`KLTVUZ[YH[LK[OH[36?HJ[P]P[`HUKPYVUYLTV]HS
JHUILTVK\SH[LK\WVUL_WVZ\YL[VIHJ[LYPHVM[OL:[YLW[VT`JLZZWLJPLZ6]LYHSS
[OL OLYL YLWVY[LK TVKPÄJH[PVUZ VM 36? JV\SK IL YLSH[LK [V HS[LYH[PVUZ VM OVZ[
WH[OVNLUYLSH[PVUZOPWZPU.4ZV`ILHUZ^P[OYLZWLJ[[V[OLJVU]LU[PVUHSJV\U[LYWHY[Z
+HPULZL,(UNLS\JJP*):HIH[\JJP(+L-PSPWWPZ=4LP.HUK4HJJHYYVUL4
(UV]LSYVSLMVYPYVUPUTVK\SH[PUN[OLHJ[P]P[`HUKTLTIYHULIPUKPUNHIPSP[`VM
H[YPTTLKZV`ILHUSPWV_`NLUHZL;OL-(:,)1V\YUHS
07VZ[LY:LZZPVU
BNU 16
Biochimica della Nutrizione
*HZWHZLTLKPH[LK7(97JSLH]HNLPUS`TWOVJ`[LZMYVT
UL^IVYUIHIPLZVMKPHIL[PJTV[OLYZ
3*90:763;6509;090)<A0
-;(98<050
:769*,33(;0
(+,37056:4(9;056.*6(;(:(9565,,;69365,
)*(77<**050.*+09,5A6(5+(693(**/06
!+LWHY[TLU[VM,_WLYPTLU[HS4LKPJPULHUK)PVJOLTPJHS:JPLUJLZ<UP]LYZP[`VM7LY\NPH"
!+LWHY[TLU[VM6IZ[L[YPJZHUK.`UHLJVSVN`<UP]LYZP[`/VZWP[HS7LY\NPH"+LWHY[TLU[VM
(NYVUVT`HUK,U]PYVUTLU[HS:JPLUJLZ<UP]LYZP[`VM7LY\NPH"!+LWHY[TLU[VM0U[LYUHS4LKPJPUL
,UKVJYPUVSVN`HUK4L[HIVSPZT<UP]LYZP[`VM7LY\NPH"!+LWHY[TLU[VM5LVUH[VSVN`<UP]LYZP[`
/VZWP[HS7LY\NPH!.L)P:H9LZLHYJO-V\UKH[PVU7LY\NPH
:L]LYHSLWPKLTPVSVNPJHSZ[\KPLZZOV^LK[OH[TH[LYUHSKPHIL[LZK\YPUNWYLNUHUJ`PZH
YLJVNUPaLKJH\ZLVMHZLYPLZVMJVUNLUP[HSKLMLJ[ZUL\YHSJHYKPHJYLUHSHUKVMSVUN[LYT
HS[LYH[PVUZPUJHYKPV]HZJ\SHYTL[HIVSPJHUKYLUHSM\UJ[PVUPUVMMZWYPUNK\L[VPUJYLHZLK
HWVW[VZPZHS[LYLKML[HSWYVNYHTTPUNHUKLWPNLUL[PJKHTHNLZ4HU`MHJ[VYZPUJS\KPUN
[OLJ`Z[LPULWYV[LHZLZJHZWHZLOH]LILLUZOV^U[VILPU]VS]LKPU[OLZLL]LU[Z
;OLHPTZVM[OLZ[\K`^HZ[VPU]LZ[PNH[L[OLWYLZLUJLHUKWYVJLZZPUNVMJHZWHZLHUK
7(977VS`(+79PIVZL7VS`TLYHZLPUJVYKISVVKJLSSZPUYLSH[PVU[VNS`JHLTPJJVU[YVS
K\YPUNPU[YH\[LYPULSPML
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9LZ\S[Z! 6\Y YLZ\S[Z ZOV^LK [OL HJ[P]H[PVU VM JHZWHZL VUS` PU JLSSZ MYVT UL^IVYU
IHIPLZVM.+4TV[OLY»Z[OH[KVUV[YLHJ[HUVW[PTHSNS`JLTPJJVU[YVSK\YPUNWYLNUHUJ`
4VYLV]LYPU[OLZHTLZHTWSLZ^LKL[LJ[[OLWYLZLUJLVMPUHJ[P]H[LK7(97LUa`TLZ
^P[OH4>$ R+HPUKPJH[PUNHZWLJPÄJJSLH]HNLTLKPH[LKI`HJ[P]L*HZWHZL
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JSHYPM`[OLTLJOHUPZTI`^P[JO[OLMHPS\YLPUHJOPL]PUNLHYS`NS`JLTPJJVU[YVSPUTH[LYUHS
TL[HIVSPZTJVU[YPI\[LZ[VL_JLZZP]LML[HSHUKULVUH[HSTVYIPKP[`H[[YPI\[HISL[VKPHIL[LZ
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07VZ[LY:LZZPVU
BNU 17
Biochimica della Nutrizione
4VKPÄJH[PVUVMJHZLPUZHZM\UJ[PVUHSIPVTHYRLYZVM
WVS`J`JSPJHYVTH[PJO`KYVJHYIVUZJVU[HTPUH[PVUVMIV]PUL
TPSR
+040*/,3,(5;6506 70,9(.6:;056+(40(5(
+»(3,::(5+9650*63(7(636<9)(50(5+9,(
+,3)6**0670,96
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+LW[VM,_WLYPTLU[HSHUK*SPUPJHS:JPLUJLZ
<UP]LYZP[`¸.K»(UU\UaPV¹VM*OPL[P7LZJHYH0[HS`
9LZLHYJO*LU[YLVU(NPUN*L:0
<UP]LYZP[`¸.K»(UU\UaPV¹VM*OPL[P7LZJHYH0[HS`
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7VS`J`JSPJ HYVTH[PJ O`KYVJHYIVUZ 7(/Z OH]L ILLU PUJS\KLK HZ [V_PJ JVTWV\UKZ
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HUK JHYJPUVNLUPJ WYVWLY[PLZ (M[LY PU[LYUHSPaH[PVU PU IPVSVNPJHS WH[O^H`Z 7(/Z HYL
TL[HIVSPaLKHUKNLULYH[LJVTWSL_TP_[\YLZVMZL]LYHSJVTWV\UKZZ\JOHZO`KYV_`S
TL[HIVSP[LZ6/W`YLULHUK6/WOLUHU[OYLUL^LYLMV\UKPUIV]PULTPSR[O\Z
TL[HIVSP[LZ VM 7(/Z ZOV\SK IL [HRLU PU[V JVUZPKLYH[PVU ^OLU L]HS\H[PUN [OL MVVK
ZHML[` /V^L]LY [OL KL[LJ[PVU VM 7(/ SL]LSZ PU IPVSVNPJHS TH[YPJLZ JV\SK IL UV[
Z\MÄJPLU[[VNP]LHYPNO[X\HSP[H[P]LLZ[PTH[PVUVM[OLÄUHSMVVKWYVK\J[Z;OLW\YWVZL
VM[OPZZ[\K`^HZ[VOPNOSPNO[[OLPU[LYHJ[PVUVMJHZLPUZ7(/ZHM[LYWOV[VPYYHKPH[PVU
PU VYKLY [V PKLU[PM` M\UJ[PVUHS IPVTHYRLYZ VM JVU[HTPUH[PVU (SWOHJHZLPU WYV[LPU
[OH[JVUZ[P[\[LZHVM[OLJHZLPUZWVVSPUIV]PULTPSR^HZ\ZLKHZHTVKLS
VMJVU[HTPUH[PVUPU]P[YVHKKLK^P[O7(/ZHUK[OLPYO`KYV_`STL[HIVSP[LZ:HTWSLZ
^LYL Z\IQLJ[LK [V WOV[VPYYHKPH[PVU [V LTWOHZPaL [OL YLHJ[P]P[` VM JVU[HTPUHU[Z
(M[LY LUa`TH[PJ KPNLZ[PVU VM [OL PUJ\IH[LK WYV[LPU WLW[PKLZ ^LYL L_[YHJ[LK HUK
HUHS`aLK I` THZZ ZWLJ[YVTL[Y` +PMMLYLU[PHS WLW[PKLZ ^LYL OPNOSPNO[LK PUKPJH[PUN
[OL VJJ\YYLUJL VM WYV[LPU TVKPÄJH[PVUZ PUJ\IH[LK ^P[O O`KYV_`STL[HIVSP[LZ
;HUKLTTHZZZWLJ[YVTL[Y`^HZLTWSV`LK[VJOHYHJ[LYPaLHUKPKLU[PM`[OLTVKPÄLK
WLW[PKLZPU]P[YV:\JJLZZP]LS`[OLWV[LU[PHSIPVTHYRLYOPNOSPNO[LK^LYLZLHYJOLK[V
JVU[HTPUH[LKIV]PULTPSR;OLZLWYLSPTPUHY`YLZ\S[ZHSSV^LK\Z[VOPNOSPNO[WYV[LPU
7(/ZHKK\J[ZHZTHYRLYZVMJVU[HTPUH[PVU0MJVUÄYTLK[OLZLL]PKLUJLZJV\SKIL
L_WSVP[LK[VKL]LSVWUL^THYRLYPUKPJH[VYZVMX\HSP[`HNYPMVVKPUK\Z[Y`
9LMLYLUJLZ
3\[a:L[HS1V\YUHSVM(NYPJ\S[\YHSHUK-VVK*OLTPZ[Y`
7OPSPWWL(.\`-YHUJVPZ-LUHPSSL>PSL`7LYPVKPJHSZ0UJ4HZZ:WLJ9L]! 07VZ[LY:LZZPVU
BNU 18
Biochimica della Nutrizione
0KLU[PÄJH[PVUVMWOVZWOVY`SH[LKJVTWVULU[ZPUTPSRVM
KPMMLYLU[ZWLJPLZ
)9<:*/,;;(.-(A06,-9(.(3±:7(90:0456;;0(
-,93(AA6(4
+LWHY[TLU[VM4VYWOVSVN`)PVJOLTPZ[Y`7O`ZPVSVN`HUK(UPTHS7YVK\J[PVU
)PVJOLTPZ[Y`<UP[
7O`ZPVSVN`<UP["
+LWHY[TLU[VM6YNHUPJHUK)PVSVNPJHS*OLTPZ[Y`<UP]LYZP[`VM4LZZPUH
4PSR PZ H IPVSVNPJHS Å\PK ^LSS Z\P[LK MVY U\[YP[PVUHS Z[\KPLZ HUK KPHNUVZ[PJ
W\YWVZLZHZP[JHUILJVSSLJ[LKYV\[PULS`HUKUVUPU]HZP]LS` 7OPNOYLZVS\[PVU
549 ZWLJ[YVZJVW` KVLZ UV[ YLX\PYL HU L_[LUZP]L JOLTPJHS THUPW\SH[PVU VM
ZHTWSLZ HUK JHU LHZPS` OPNOSPNO[ KPMMLYLUJLZ HUKVY ZPTPSHYP[PLZ IL[^LLU ZHTWSLZ
HZ JVTWSL_ HZ TPSR 7 549 HUHS`ZPZ \ZPUN KL[LYNLU[Z H TL[OVK HSYLHK` \ZLK
MVY [OL HUHS`ZPZ VM WSHZTH HUK LY`[OYVJ`[L TLTIYHUL WOVZWOVSPWPK JVTWVZP[PVU
ZOV^LK [OL WYLZLUJL PU HSS TPSR ZHTWSLZ HUHS`aLK VM O`KYVS`ZPZ WYVK\J[Z VM
WOVZWOH[PK`SJOVSPUL HUK WOVZWOH[PK`SL[OHUVSHTPUL! NS`JLYVWOVZWOVY`SJOVSPUL
.7* RUV^U MVY P[Z UL\YVWYV[LJ[PVU LMMLJ[ VU HNLYLSH[LK V_PKH[P]L KHTHNL NS`JLYVWOVZWOVY`SL[OHUVSHTPUL.7,HUKWOVZWOVY`SJOVSPUL7*7OVZWOVJYLH[PUL
PZHSZVWYLZLU[L_JLW[PUO\THUHUKJV^TPSR0UHKKP[PVUTPUVYWOVZWOVY`SH[LKZ\NHYZ
[OH[WYVIHIS`HYLTL[HIVSPJPU[LYTLKPH[LZVMIPVJOLTPJHSWH[O^H`Z^LYLPKLU[PÄLK
NHSHJ[VZLWOVZWOH[L 5HJL[`SNS\JVZHTPULWOVZWOH[L PU NYLH[LY HTV\U[Z PU
KVURL`TPSR[OHUPUO\THUJV^NVH[HUKJHTLSTPSR;OLZLKH[HJVUÄYT[OLUH[\YHS
]HYPHIPSP[`VMTPSRZHTWSLZHTVUNKPMMLYLU[ZWLJPLZ4VYLV]LY[OPZZPTWSPÄLKTL[OVK
JV\SKILWYVWVZLK[VJHYY`VUHJVTWHYH[P]LZ[\K`VMWOVZWOVY`SH[LKJVTWVULU[Z
\ZLKHZTLTIYHULZ[HIPSPaLYZPU[OLHU[PV_PKHU[[OLYHW`
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:\JO`1L[HS 5\[Y9LZ :H]HZ[HUV4L[HS(YJO4LK9LZ
07VZ[LY:LZZPVU
BNU 19
Biochimica della Nutrizione
/<4(5796:;(;,*(5*,979,=,5;065)@.9,,5;,(
*(;,*/05:!
;/,*3050*(3;90(3(5+7<;(;0=,463,*<3(9
4,*/(50:4:6-(*;065
90AA0-,+,90*((5+),;;<AA0:(=,906
+LWHY[TLU[VM)PVTLKPJPUL)PV[LJOUVSVN`HUK;YHUZSH[PVUHS9LZLHYJOHUK
*LU[YLMVY4VSLJ\SHYHUK;YHUZSH[PVUHS6UJVSVN`*64;
<UP]LYZP[`VM7HYTH7HYTH0[HS`"5H[PVUHS0UZ[P[\[LVM)PVZ[Y\J[\YLHUK)PVZ`Z[LTZ
05))9VTL0[HS`
7YVZ[H[LJHUJLY7*HPZVULVM[OLTHQVYJHUJLYRPSSLYPUHNLKTLUSP]PUNPU>LZ[LYU
*V\U[YPLZ3P[[SLPZRUV^UHIV\[[OLTVSLJ\SHYTLJOHUPZTZSLHKPUN[VWYVNYLZZPVU
(K]HUJLK 7*H PZ HSTVZ[ PUJ\YHISL ILJH\ZL [OLYHWL\[PJ VW[PVUZ HYL SHYNLS`
\UZH[PZMHJ[VY`;OLYLMVYLP[PZ\YNLU[[VZLHYJOMVYLMMLJ[P]L[OLYHWPLZ^OLUKPZLHZLPZ
Z[PSSPUP[PHSHUKWV[LU[PHSS`J\YHISL;OLSVUNSH[LUJ`THRLZ7*HHUPKLHSJHUKPKH[LMVY
JOLTVWYL]LU[PVUPU[LY]LU[PVUILMVYLPU]HZPVUHUKTL[HZ[H[PJZWYLHKVJJ\YZ
*H[LJOPUZL_[YHJ[LKMVY.YLLU;LH.;,WVZZLZWV[LU[HU[PJHUJLYHJ[P]P[`PU]P[YV
>L ZOV^LK [OH[ .;, HKTPUPZ[YH[PVU ^HZ LMMLJ[P]L H[ PUOPIP[PUN WYVNYLZZPVU VM 7*H
[V JSPUPJHS L]PKLUJL PU WH[PLU[Z ILHYPUN PZVSH[LK WYLULVWSHZ[PJ SLZPVU /.705
0TWYV]LTLU[^HZZ[HISL`LHYZHM[LYZ\ZWLUZPVUVM[YLH[TLU[>LZLHYJOLKMVYNLULZ
^OVZL L_WYLZZPVU ^HZ ZWLJPÄJHSS` JOHUNLK I` .;, HKTPUPZ[YH[PVU PU ]P]V I` +5(
TPJYVHYYH`PU[OLWYVZ[H[LVM.;,[YLH[LK;9(47TPJLH[YHUZNLUPJTVKLSMVY7*H
]LYZ\ZJVU[YVSZSVVRPUNMVYIPVTHYRLYZ[VTVUP[VY.;,HJ[P]P[`HUK[PZZ\LYLZWVUZL
0U[LYLZ[PUNS` ^L MV\UK [OH[ [OL U\TILY VM NLULZ ^OVZL L_WYLZZPVU PZ ZPNUPÄJHU[S`
JOHUNLKI`.;,HYLYLSH[P]LS`ML^(TVUN[OLZLVULVM[OLTVZ[PU[LYLZ[PUNNLUL
^HZ*S\Z[LYPU*3<*3<PZPU]VS]LKPUJLSSZ\Y]P]HSHUKJLSSKLH[OHUKJVUZPKLYLK[V
KYP]L[OLMH[LVM[OLJLSS+\YPUN7*HWYVNYLZZPVUPU;9(47TPJL*3<L_WYLZZPVUPZ
SVZ[H[LHYS`Z[HNLZ>OLUHUPTHSZHYL[YLH[LK^P[O.;,PUKYPURPUN^H[LYHUK
YLZWVUK[VJOLTVWYL]LU[PVU*3<L_WYLZZPVUNL[ZIHJR[VOPNOSL]LSZHUKZLUZP[P]P[`
[V HWVW[V[PJ KLH[O PZ YLZ[VYLK ([ KPMMLYLUJL ^OLU JOLTVWYL]LU[PVU MHPSZ PU .;,
YLZPZ[HU[7*H*3<L_WYLZZPVUPZZ[PSSHIZLU[HUKJLSSZHYLYLZPZ[HU[[VHWVW[VZPZ3VUN
[LYTHKTPUPZ[YH[PVUVM.;,^HZMV\UKZHMLPUO\THUZ;OLYLMVYLJOLTVWYL]LU[PVU
^P[O .;, TH` IL [OL ILZ[ VW[PVU MVY H IL[[LY JSPUPJHS THUHNPUN VM 7*H I\[ TVYL
TVSLJ\SHYKH[HHYLULJLZZHY`[VKPZJYPTPUH[LYLZWVUZP]LMYVTYLZPZ[HU[7*H
07VZ[LY:LZZPVU
BNU 20
Biochimica della Nutrizione
7YVHWVW[V[PJHJ[P]P[`VM[OLWO`[VJOLTPJHS0UKPJH_HU[OPU
VUJVSVYLJ[HSJHYJPUVTHJLSSZ*HJVHUKLWPNLUL[PJ
*W.KLTL[O`SH[PVUVM[OLWYVTV[LYHUKYLHJ[P]H[PVUVM[OL
L_WYLZZPVUVMW
5(:,330-46+0*(4(;;(5A06(*(9(+655(-.,5;03,*
;,:690,9,330=9,(4(
+PWHY[PTLU[VKP:JPLUaLL;LJUVSVNPL4VSLJVSHYPL)PVTVSLJVSHYP
<UP]LYZP[nKP7HSLYTV
7O`[VJOLTPJHSZ WSH` WYVTPULU[ YVSLZ PU O\THU KPL[ HUK U\[YP[PVU HZ WYV[LJ[P]L
YLKVHJ[P]L Z\IZ[HUJLZ PU WYL]LU[PVU VM ZL]LYHS KPZVYKLYZ HUK JOYVUPJ KPZLHZLZ PU
O\THUZ;VKH`[OLPYM\UJ[PVUHZWV[LU[TVK\SH[VYZVM[OLTHTTHSPHULWPNLUVTL
YLN\SH[LKNLULL_WYLZZPVUPZYHWPKS`LTLYNPUN0U[OLWYLZLU[Z[\K`HU[PWYVSPMLYH[P]L
LMMLJ[ZVM0UKPJH_HU[OPU0UKMYVT[OLMY\P[ZVM6W\U[PHÄJ\ZPUKPJHHUKWV[LU[PHS
PUÅ\LUJLVU+5(TL[O`SH[PVUOHZILLUPU]LZ[PNH[LKVU*HJVJLSSZHO\THUJLSS
SPULVMJVSVYLJ[HSJHYJPUVTH0UKJH\ZLKHJSLHYKVZLHUK[PTLKLWLUKLU[KLJYLHZL
VM [OL JLSS WYVSPMLYH[PVU 0* ›4 HZZVJPH[LK [V HWVW[VZPZ HZ KLTVUZ[YH[LK I`
WOVZWOH[PK`SZLYPUL L_[LYUHSPaH[PVU HUK KLWVSHYPaH[PVU VM TP[OVJVUKYPHS TLTIYHUL
0UKKLJYLHZLK[OL.V.WOHZL^OLYLHZPUJYLHZLK:HUK.4WOHZLZVM[OLJLSS
J`JSL;OLWO`[VJOLTPJHSKPKUV[HS[LYLK[OLPU[YHJLSS\SHY96:SL]LSZI\[KLJYLHZLK
[OL B*HDP 0U]LZ[PNH[PVU VU +5( TL[O`SH[PVU \ZPUN 4,:(77*9 4L[O`SH[PVU
:LUZP[P]L(YIP[YHYPS`7YPTLK7VS`TLYHZL*OHPU9LHJ[PVUZOV^LK[OH[›40UK
PUK\JLKHZSPNO[NSVIHSKLTL[O`SH[PVUHM[LYHO[YLH[TLU[(UHS`ZPZVMLWPNLUL[PJ
JOHUNLZ PU [OL +5( TL[O`SH[PVU WH[[LYU H[ *W. WYVTV[LY VM W 052H \ZPUN
4:9,4L[O`SH[PVU:LUZP[P]L9LZ[YPJ[PVU,UKVU\JSLHZLZ4\S[PWSL_7VS`TLYHZL*OHPU
9LHJ[PVU ZOV^LK [OH[ 0UK JH\ZLK *W. KLTL[O`SH[PVU >LZ[LYU ISV[[PUN HUHS`ZPZ
JHYYPLKV\[^P[OWTVUVJSVUHSHU[PIVK`JVUÄYTLK[OLYLHJ[P]H[PVUVM[OLWYV[LPU
L_WYLZZPVU7YLZLU[KH[HZ\NNLZ[PUN[OH[HSVUN[LYTL_WVZ\YL[VPUKPJH_HU[OPUPU
KPL[ TPNO[ WV[LU[PHSS` HMMLJ[ LWPNLUL[PJ THJOPULZ VM [OL PU[LZ[PUHS JLSSZ WYL]LU[PUN
VY YLWHPYPUN PUP[PHS KLYHUNLTLU[ZKPZVYKLYZ LUJV\YHNL Z[\KPLZ VU [OL TLJOHUPZT
PU]VS]LK
3P]YLH4(HUK;LZVYPLYL3PU/LYIHSHUK;YHKP[PVUHS4LKPJPULLKZ7HYRLY36UN*/HSSP^LSS
)4HYJLS+LRRLY
*HYHKVUUH-)HYIH[H.:JPHUKYLSSV.5V]H:JPLUJL7\ISPZOLYZ0UJ5L^@VYR<:([OX\HY[LY
0:)5
07VZ[LY:LZZPVU
BNU 21
Biochimica della Nutrizione
;OLKPL[WO`[VJOLTPJHS0UKPJH_HU[OPUZ\WWYLZZLZ
RL[VJOVSLZ[LYVSPUK\JLK;/7JLSSHWVW[VZPZ]PH
PUOPIP[PVUVMYLKV_\UIHSHUJLJ`[VZVSPJ*HPUJYLHZLHUK
5-R)HJ[P]H[PVU
(;;(5A06(70AA<;6,;,:690,9,3(33,.9(430=9,(4(
+PWHY[PTLU[VKP:JPLUaLL;LJUVSVNPL4VSLJVSHYPL)PVTVSLJVSHYP
<UP]LYZP[nKP7HSLYTV
0UKPJH_HU[OPU0UKPZ[OLIL[HSHPUWPNTLU[JOHYHJ[LYPaPUN[OLLKPISLMY\P[ZVM6W\U[PH
ÄJ\Z PUKPJH (U[PV_PKHU[ HJ[P]P[` VM 0UK PU O\THUZ HUK P[Z IPVHJ[P]P[` OH]L ILLU
YLWVY[LK PU H U\TILY VM Z[\KPLZ PU JLSS J\S[\YLZ HZ ^LSS HZ PU ]P]V ;OPZ Z[\K`
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SPULZ\ITP[[LK[VPUQ\Y`I`RL[VJOVSLZ[LYVS2*HJVTWVULU[VMV_PKPaLK3+3
^OPJO WSH`Z H JLU[YHS YVSL PU H[OLYVZJSLYVZPZ 0U V\Y TVKLS H WH[OVWO`ZPVSVNPJHS
LU]PYVUTLU[ OHZ ILLU ZPT\SH[LK JVPUJ\IH[PUN ›4 0UK ^P[O ›4 2* PL
JVUJLU[YH[PVUZJVTWHYHISL^P[O[OVZLTLHZ\YLKPUO\THUWSHZTHHM[LYJHJ[\ZWLHY
MY\P[PU[HRLHUKRUV^U[VILWYLZLU[H[[OLH[OLYVZJOSLYV[PJWSHX\LYLZWLJ[P]LS`
0UK JV\U[LYHJ[LK [OL 2*¶PUK\JLK JLSS NYV^[O PUOPIP[PVU WYL]LU[PUN [OL HYYLZ[ VM
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JVUZPKLYLK PUJS\KPUN U\JSLHY TVYWOVSVNPJHS JOHUNLZ L_[LYUHSPaH[PVU VM 7: PU
WSHZTHTLTIYHULKLWVSHYPaH[PVUVMTP[VJOVUKYPHSTLTIYHUL7(97JSLH]HNLHUK
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HS[LYH[PVUZ0UKPUOPIP[LK[OLPUJYLHZLVMRL`ZLJVUKTLZZHUNLYZYLN\SH[PUNJLSSKLH[O
Z\JOHZ96:HUK*HPVUZ-PUHSS`0UKWYL]LU[LK[OL2*KLWLUKLU[[OPVSKLWSL[PVU
HUK[OLHJ[P]H[PVUVM[OLJLSS\SHYLMMLJ[VYVMPUÅHTTH[PVU5-R);HRPUNPU[VHJJV\U[
SVJHSPaH[PVU VM 0UK H[ [OL SPWPK IPSH`LYZ YLKV_ HJ[P]P[` VM 0UK ^P[O PU[LYJLW[PVU VM
96:NLULYH[LKI`[OLTHJYVWOHNL5(+7/V_PKHZL56?HUKVYPU[LYMLYLUJL^P[O
TLJOHUPZTZJVU[YVSSPUN*HPUÅ\_TH`ILO`WV[OLZPaLK
3P]YLH4(HUK;LZVYPLYL3PU/LYIHSHUK;YHKP[PVUHS4LKPJPULLKZ7HYRLY36UN*/HSSP^LSS
)4HYJLS+LRRLY
;LZVYPLYL3L[HS(T1*SPU5\[Y 3LTHPYL,^PUN:L[HS*LSS)PVS;V_PJVS ;\YJV3P]LYP43L[HS1(NYPJ-VVK*OLT ¶ 07VZ[LY:LZZPVU
BNU 22
Biochimica della Nutrizione
5\[YHJL\[PJHSHUKHU[PV_PKHU[WV[LU[PHSVM3()!IPVJOLTPJHS
HUKWYV[LVTPJTL[OVKZ[VLS\JPKH[L[OL:LÄ_PUNWH[O^H`
.0<5;(*(9364(5.0(7(5,,902(.(3(56,<.,506
7(34,:,(5.,367,::065,(3,::(5+96(469,:(56(5.,3(
7,::065,,590*(
3PML:JPLUJLZHUK:`Z[LTZ)PVSVN`+LWHY[TLU[<UP]LYZP[`VM;VYPUV0[HS`
+LWHY[TLU[VM6YNHUPJ*OLTPZ[Y`HUK)PVJOLTPZ[Y`<UP]LYZP[`VM5HWVSP0[HS`
7YVIPV[PJSHJ[PJHJPKIHJ[LYPHOH]LM\UKHTLU[HSPTWVY[HUJLMVYHJVYYLJ[M\UJ[PVUPUN
VM [OL O\THU N\[ ;OL` HYL HISL [V WYVK\JL ZL]LYHS TL[HIVSP[LZ Z\JO HZ ZOVY[
JOHPU MH[[` HJPKZ JVUQ\NH[LK SPUVSLPJ HJPKZ IHJ[LYPVJPUZ IPVHJ[P]L TVSLJ\SLZ HUK
L_VWVS`ZHJJOHYPKLZ[OH[PUKPMMLYLU[^H`ZJHUIYPUNHILULÄ[[V[OLOVZ[(TVUN
[OLZL]HYPV\ZWVZZPIPSP[PLZ[OLJHWHIPSP[`[VPU[LYUHSPaLHUKÄ_ZLSLUP\TOHZYLJLU[S`
YLJLP]LKHNYLH[H[[LU[PVU
:LSLUP\T:LPZHULZZLU[PHSKPL[HY`[YHJLLSLTLU[MVYO\THUZHUKTPJYVVYNHUPZTZ
:L KLÄJPLUJ` PZ HZZVJPH[LK ^P[O ]HYPV\Z JOYVUPJ KPZLHZLZ Z\JO HZ JHYKPV]HZJ\SHY
KPZLHZLZ ZVTL RPUKZ VM JHUJLY HUK V_PKH[P]L Z[YLZZ ZPUJL :L PZ WYLZLU[ PU [OL
HJ[P]L ZP[L VM NS\[H[OPVUL WLYV_PKHZL ;OL \ZL VM :LLUYPJOLK SHJ[VIHJPSSP PZ HU
PU[LYLZ[PUN HWWYVHJO [V ZVS]L ZLSLUP\TKLÄJPLUJ` LZWLJPHSS` K\YPUN HNLPUN! [OLZL
TPJYVVYNHUPZTZNYV^UPUWYLZLUJLVMPUVYNHUPJMVYTZVMZLSLUP\THYLHISL[VJVU]LY[
P[PU[VTVYLIPVH]HPSHISLVYNHUPJMVYTZPU[YVK\JPUNP[PU[VWYV[LPUZ;OLHKKP[PVUVM
[OLZLTPJYVVYNHUPZTZ[VHWHY[PJ\SHYMVVKNLULYH[LZHM\UJ[PVUHSMVVK
0UVYKLY[VIL[[LY\UKLYZ[HUK[OLILOH]PVYVM3HJ[VIHJPSS\ZYL\[LYP3I)4+:4
^OLUNYV^UVUZLSLUP\TZL]LYHSZ[YH[LNPLZ^LYLHWWSPLK!HJVTWHYH[P]LWYV[LVTPJ
HWWYVHJO^HZ\ZLK[VOPNOSPNO[JOHUNLZPU[OLWYV[LPUWH[[LYUPUZL]LYHSJLSS\SHYKPZ[YPJ[Z
HUKW0YHUNLZ;OLZLHUHS`ZLZYL]LHSLK[OL\WYLN\SH[PVUVM[OLLULYNL[PJTL[HIVSPZT
NS`JVS`ZPZHUKWLU[VZLWOVZWOH[LWH[O^H`HOPNOL_WYLZZPVUVMHKOLZPVUPU]VS]LK
WYV[LPUZHUKS\Z[I\[UV[SLHZ[[OL\WYLN\SH[PVUVMHZLSLUVJ`Z[LPULS`HZLYLZWVUZPISL
MVY[OLPUZLY[PVUVMZLSLUVJ`Z[LPULPU[VWYV[LPUZ;VZOLKTVYLSPNO[VU[OLHIPSP[`VM[OL
Z[YHPU[VÄ_ZLSLUP\TPU[VWYV[LPUZ:,*HUK/73*JV\WSLK^P[O0*74:L_WLYPTLU[Z
^LYLJHYYPLKV\[;OLVI[HPULKYLZ\S[ZYLWYLZLU[HM\Y[OLYJVUÄYTH[PVUVM[OLHIPSP[`
VM3YL\[LYP3I)4[VNL[TVYLIPVH]HPSHISLMVYTZVMZLSLUP\TZLSLUPaLKWYV[LPUZ
07VZ[LY:LZZPVU
BNU 23
Biochimica della Nutrizione
ơSHJ[VNSVI\SPUZ[Y\J[\YHSYLHYYHUNLTLU[H[PU[LYMHJLZ!
YLSL]HUJL[VP[ZPTT\UVNLUPJILOH]PVY
4090(504(;;,6.,5;030504(;;,64(9,5.64(<96
0(4,;;0:;,-(50(-9•20”9/(55,(5+)65640-9(5*,:*6
:LJ[PVUVM*OLTPZ[Y`HUK)PVTVSLJ\SHY:JPLUJLZ+L-,5:<UP]LYZP[`VM4PSHU
0UZ[P[\[LVM*OLTPZ[Y`HUK)PVJOLTPZ[Y`-HJ\S[`VM3PML:JPLUJLZ
<UP]LYZP[`VM*VWLUOHNLU
;OLWO`ZPVSVNPJHSWYVWLY[PLZVMTHU`WYV[LPUZYLSPLZVU[OLPYILOH]PVYH[[OLTVSLJ\SHY
ZJHSL^OPJOVM[LUPZHMMLJ[LKI`[OLPYZ\YMHJLPU[LYHJ[PVUZ(JLU[YHSPZZ\LMVY[OLPU
]P]VHJ[PVUVMHSSLYNLUZI\[HSZVVMHU[PIHJ[LYPHSVYV[OLY^PZLIPVHJ[P]LWLW[PKLZPZ[OL
Z\YMHJLILOH]PVYPU[LYTZVMPU[LYHJ[PVU^P[OV[OLYTVSLJ\SLZHUKZ\YMHJLZWYLZLU[PU
[OLZHTLLU]PYVUTLU[
ơSHJ[VNSVI\SPU)3.PZ[OLTHQVYIV]PUL^OL`WYV[LPU(R+HKPTLYVMPKLU[PJHS
Z\I\UP[Z)3.PZJVUZPKLYLK[OLTHQVYTPSRHSSLYNLU(NLULYHSYLSH[PVUZOPWIL[^LLU
Z[Y\J[\YLHUKPTT\UVYLHJ[P]P[`VMTPSRWYV[LPUZOHZUV[ILLULZ[HISPZOLK;OLHPTVM
[OPZ^VYRPZ[V\UKLYZ[HUK^OL[OLYHUKOV^[OL)3.JVUMVYTH[PVUHSJOHUNLZKLYP]LK
MYVT [OL PU[LYHJ[PVU ^P[O [^V TVKLS O`KYVWOVIPJ PU[LYMHJLZ H SPX\PK VPSPU^H[LY
UHUVLT\SZPVUHUKZVSPKSH[L_UHUVWHY[PJSLZJHUTVK\SH[LP[ZPTT\UVYLHJ[P]P[`HUK
P[ZHIZVYW[PVUI`O\THUJLSSZPU]VS]LKPU[OLPTT\ULYLZWVUZL
*VUMVYTH[PVUHS JOHUNLZ PU [OL HIZVYILK WYV[LPU ^LYL KLTVUZ[YH[LK [OYV\NO
ZWLJ[YVZJVWPJ HUK SPTP[LK WYV[LVS`ZPZ4: HWWYVHJOLZ ,30:( L_WLYPTLU[Z
KLTVUZ[YH[LK[OH[PTT\UVYLHJ[P]P[`VM)3.PUJYLHZLZ\WVUHKZVYW[PVUVULP[OLYSPX\PK
SPX\PKVYZVSPKSPX\PKO`KYVWOVIPJPU[LYMHJLZSPRLS`HZHJVUZLX\LUJLVMYLJVNUP[PVU
VM JVNUH[L YLNPVUZ VM [OL WYV[LPU I` [OL HU[PIVKPLZ 0UKLLK HM[LY O`KYVS`ZPZ ^P[O
[Y`WZPU[OLV]LYHSSPTT\UVYLHJ[P]P[`VM[OLHIZVYILKWYV[LPUYLTHPUZOPNOJVTWHYLK
^P[O[OH[VM[OLWYV[LPUO`KYVS`aLKPUZVS\[PVU(SZV[OLRPUL[PJZVMPU[LYUHSPaH[PVUI`
TVUVJ`[LZ VM )3. HKZVYILK VU [OL VPS UHUVKYVWSL[Z PZ ^P[O YLZWLJ[ [V MYLL )3.
^OPJOPZHIZVYILKTVYLLMÄJPLU[S`HUKYHWPKS`[OHU)3.HKZVYILKVUHULT\SZPVU
PU[LYMHJL*VTWL[P[PVUL_WLYPTLU[ZJHYYPLKV\[I`HKKPUNMYLL\USHILSLKWYV[LPU[V
IV[OMYLLHUKLT\SZPVU)3.Z`Z[LTZZOV^ZOV^[OLMYLLWYV[LPUZLLTZ[VHJ[ZHZ
HULUOHUJLYMVY[OLPU[LYUHSPaH[PVUVM[OLVPSIV\UKWYV[LPU>OL[OLY[OLZLL]PKLUJLZ
PUKPJH[L PUKLWLUKLU[ WH[[LYUZ MVY PU[YHJLSS\SHY \W[HRL VM MYLL HUK UHUVWHY[PJ\SH[L
IV\UK)3.YLTHPUZ[VIL]LYPÄLK
07VZ[LY:LZZPVU
BNU 24
Biochimica della Nutrizione
5H[\YHSHUKZ`U[OL[PJMVYTZVM]P[HTPU,HZPUOPIP[VYZ
VMZLJYL[VY`WOVZWOVSPWHZL(Z73(PUJ`Z[PJÄIYVZPZ
LWP[OLSPHSJLSSZ
7036330-9(5*,:*(3,.5(0630:03=0()0(5*6504(9*,336
7096++04(9;(.(330-9(5*,:*6(5+.69(**0.0(5-9(5*,:*6
+PWHY[PTLU[VKP4LKPJPUH0U[LYUH<UP]LYZP[nKLNSP:[\KPKP7LY\NPH
:LJYL[VY` WOVZWOVSPWHZL ( Z73( .00( PZ H MVYT VM [OL OL[LYVNLULV\Z MHTPS` VM
73( LUa`TLZ OPNOS` L_WYLZZLK PU PUÅHTTH[VY` [PZZ\LZ HUK PU H ]HYPL[` VM O\THU
[\TVYZ.YHMM91L[HS*SPU*HUJLY9LZ)LZPKL[OLRL`YVSLPUHYHJOPKVUH[L
TL[HIVSPZT HUK [O\Z PU [OL JVU[YVS VM PUÅHTTH[VY` YLHJ[PVUZ [OPZ PZVLUa`TL JHU
WYVK\JL KPMMLYLU[ IPVSVNPJHS YLZWVUZLZ ^OLU YLSLHZLK PU [OL L_[YHJLSS\SHY TLKP\T
Z\JO HZ JLSS RPSSPUN HUK JLSS[VJLSS ZPNUHSPUN HZ ^LSS HZ [PZZ\L KLNLULYH[PVU HUK
YLTVKLSSPUN (JJVYKPUN ^P[O *OHUKYH L[ HS *OHUKYH L[ HS 1 4VS )PVS [OL
MH[ZVS\ISLHU[PV_PKHU[]P[HTPU,PU[OLMVYTVMHSWOH[VJVWOLYVSJV\SKZWLJPÄJHSS`
[HYNL[VUZ73( SLHKPUN[VJVTWL[P[P]LPUOPIP[PVUVM[OLLUa`TLHJ[P]P[`2P$ —
›4/LYL^LWYLSPTPUHYPS`PU]LZ[PNH[LKZ73(.00(L_WYLZZPVUHUKHJ[P]P[`PUS\UN
LWP[OLSPHSJLSSZL_WYLZZPUN[OLJ`Z[PJÄIYVZPZ*-T\[H[PVU¬HUKHZZLZZLKLUa`TL
PUOPIP[PVUHUK[OLJ`[VWYV[LJ[P]LHUKHU[PPUÅHTTH[VY`LMMLJ[ZVMZVTLUH[\YHSHUK
Z`U[OL[PJMVYTZVM]P[HTPU,ILMVYLHUKHM[LY[OLJOHSSLUNLVM*-LWP[OLSPHSJLSSZ^P[O
HJ[P]L7ZL\KVTVUHZ(LY\NPUVZH7(6
<ZPUNHUV]LSHZZH`WYVJLK\YLZL[\WPU[OPZZ[\K`[VZJYLLUZ73(HJ[P]P[`PUSHYNL
ZJHSLWOHYTHJVSVNPJHS[LZ[Z^LJVUÄYTLK[OLPUOPIP[VY`HJ[P]P[`VM]P[HTPU,MVYTZ
^P[ONHTTH[VJVWOLYVSHUKHSWOH;,(HUL[OLYKLYP]H[P]LVMHSWOH[VJVWOLYVSHZ
[^V VM [OL TVZ[ LMMLJ[P]L [LZ[ TVSLJ\SLZ ;OL PUOPIP[VY` HJ[P]P[` ^HZ WYLZLU[ L]LU
HM[LY [OL IHJ[LYPHS JOHSSLUNL ^P[O 7(6 9LHJ[P]L V_`NLU ZWLJPLZ 03 WYVK\J[PVU
HUK*6?HJ[P]P[`^LYLHSZVPUÅ\LUJLKI`[OL[YLH[TLU[Z[OH[^LYL^LSS[VSLYH[LKH[
WOHYTHJVSVNPJHSJVUJLU[YH[PVUZVM#›4;OLÄUKPUNZPU[OPZZ[\K`Z\NNLZ[HYVSL
MVY[OLWOHYTHJVSVNPJHSPUOPIP[PVUVMZ73(HJ[P]P[`PU[OLJVU[YVSVMS\UNPUÅHTTH[PVU
VM*-WH[PLU[Z-\Y[OLYWYLJSPUPJHSPU]LZ[PNH[PVUVM]P[HTPU,KLYP]LKZ73(PUOPIP[VYZ
PU*-TVKLSZ`Z[LTZHYLH^HP[LK
07VZ[LY:LZZPVU
0TT\UVSVNPH
)PVJOPTPJH
04)
IMB 1
Immunologia Biochimica
4,;()63640*:6-4<3;073,4@,364(
-9(5*,:*(-65;(5(16:Ï4(5<,3.(9*0(4(5;,0.(
:03=0(4(90,30:(),;;(4(90(50,590*6*(5,=(
-9(5*,:*6*(45(:064(.+(4(9*(;;0,+6(9+6.(<+,
.06=(55(4<:*696),9;6:0;0(-()06*0*,90
(5+:0465,*,5*0
:HU9HMMHLSL:JPLU[PÄJ0UZ[P[\[L4PSHUV0[HS`
<UP]LYZP[n=P[H:HS\[L:HU9HMMHLSL4PSHUV0[HS`
999LZLHYJOMVY9LU[0[HS`
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4\S[PWSL 4`LSVTH 44 PZ HU PUJ\YHISL ULVWSHZ[PJ KPZVYKLY VM WSHZTH JLSSZ ^OPJO
PU]HKL [OL IVUL THYYV^ ZLJYL[L TVUVJSVUHS PTT\UVNSVI\SPUZ HUK PUK\JL IVUL
SLZPVUZ O`WLYJHSJLTPH HULTPH HUK YLUHS MHPS\YL 44 PZ WYLJLKLK I` H MYLX\LU[
JVUKP[PVU TVUVJSVUHS NHTTVWH[O` VM \UKL[LYTPULK ZPNUPÄJHUJL 4.<: I\[ PZ
KPHNUVZLKVUS`VUJLLUKVYNHUKHTHNLLUZ\LZ
0UZLHYJOMVYLHYS`IPVTHYRLYZVM44^LKLWSV`LK4:IHZLKTL[HIVSPJWYVÄSPUN[V
PU]LZ[PNH[L[OLJVYYLSH[LZVMT`LSVTH[\TVYPNLULZPZHUKWYVNYLZZPVUPUWLYPWOLYHSHUK
IVULTHYYV^WSHZTH(ZH^OVSLTL[HIVSPJMVV[WYPU[PUNZ\JJLZZM\SS`KPZJYPTPUH[LK
HJ[P]L KPZLHZL MYVT JVU[YVSZ JVYYLSH[PUN ^P[O IVUL THYYV^ [\TVY JLSS\SHYP[` 9LUHS
K`ZM\UJ[PVU^HZPUKLWLUKLU[S`HKKYLZZLKSLHKPUN[V[OLPKLU[PÄJH[PVUVMHU\TILY
VM TL[HIVSP[LZ WYL]PV\ZS` HZZVJPH[LK ^P[O JOYVUPJ RPKUL` KPZLHZL HUK WYV]PKPUN
[OL WV^LY [V KPZJYPTPUH[L JHUJLY NYV^[O MYVT LUKVYNHU KHTHNL 44HZZVJPH[LK
MLH[\YLZPUJS\KLKHZPNUPÄJHU[KLJYLHZLPUVULSPWPKJSHZZHUK[OLPUJYLHZLVMHML^YHYL
TL[HIVSP[LZZ\NNLZ[PUN\UL_WLJ[LKWH[OVTLJOHUPZTZ-\Y[OLYTVYLO`WV[OLZPaPUN
L_[YHJLSS\SHYS`ZVSPWPKZ[VILJVUZ\TLKI`44JLSSZ^LVI[HPULKPU]P[YVL]PKLUJL
Z\WWVY[PUNH[YVWOPJYVSLVUT`LSVTHJLSSZ
44 JVUZPZ[Z VM PU[YHTLK\SSHY` KPZZLTPUH[PVU VM THSPNUHU[ JLSSZ JH\ZPUN VZ[LVS`[PJ
SLZPVUZH[OPNOYPZRVMMYHJ[\YL(PTPUN[VHKKYLZZ[OLSVJHSOL[LYVNLULP[`VMT`LSVTH
IVUL KPZLHZL ^L HSZV ZL[ \W [V PU]LZ[PNH[L 44 SLZPVUZ I` PU[HJ[ [PZZ\L /94(:
549 4L[HIVSPJ ÄUNLYWYPU[PUN LMÄJPLU[S` TH[JOLK OPZ[VSVNPJHS ÄUKPUNZ JS\Z[LYPUN
HJJVYKPUN[V[PZZ\LPKLU[P[`4VYLV]LYWLHRHZZPNUTLU[YL]LHSLK[OH[HYLHZVMOPNO
[\TVYJLSS\SHYP[`^LYLHZZVJPH[LK^P[OOPNOJVUJLU[YH[PVUVMSPWPKZ
0U HSS V\Y KH[H Z\NNLZ[ [OH[ TL[HIVSVTPJZ PZ H WV^LYM\S Z[YH[LN` [V HKKYLZZ [OL
JVTWSL_P[`VM44WH[OVWO`ZPVSVN`HUKKL]LSVWUV]LSKPHNUVZ[PJZ[YH[LNPLZ
07VZ[LY:LZZPVU
IMB 2
Immunologia Biochimica
3.(3:)7PUK\JLZ=,-.PUO\THUIYLHZ[JHUJLYJLSSZHUK
WYVTV[LZHUNPVNLULZPZ
;9(050:(9(70**636,5A(;05(9050*63(-0*/,9(044(
+»,.0+064(<90A0(+090:06(55(30:(5(;630*3(9((5+
0(*6),330:;,-(56
4LKPH7OHYTHZYS=PHKLP=LZ[PUP*OPL[P0[HS`"
+LWHY[TLU[VM)PVTLKPJHS:JPLUJLZ.+»(UU\UaPV<UP]LYZP[`HUK-V\UKH[PVU
*OPL[P0[HS`
)HJRNYV\UK! ,SL]H[LK ZLY\T VY [PZZ\L SL]LSZ VM SLJ[PU NHSHJ[VZPKLIPUKPUN ZVS\ISL
IPUKPUN WYV[LPU 3.(3:)7 OH]L ILLU HZZVJPH[LK ^P[O ZOVY[ Z\Y]P]HS HUK
KL]LSVWTLU[ VM TL[HZ[HZPZ PU H ]HYPL[` VM O\THU JHUJLYZ /V^L]LY [OL YVSL VM
3.(3:)7WHY[PJ\SHYS`PU[OLJVU[L_[VM[\TVYOVZ[YLSH[PVUZOPWZPZZ[PSSTPZZPUN
4L[OVKZ!4+(4)O\THUIYLHZ[JHUJLYJLSSZ^LYLZ[HIS`ZPSLUJLKMVYO\THU
3.(3:)7HUKHZZH`LKMVY[OLJVU[LU[VM]HZJ\SHYLUKV[OLSPHSNYV^[OMHJ[VY=,.-
I`,30:(-VYHKOLZPVUHZZH`JLSSZ^LYLWSH[LKVU[V ^LSSJ\S[\YLKPZOLZWYLJVH[LK
^P[O *VSSHNLU 0= *VSSHNLU = -PIYVULJ[PU VY 3HTPUPU (KOLYLU[ JLSSZ ^LYL Z[HPULK
HUK[OLPYHIZVYIHUJLTLHZ\YLKH[UT-VY[YHUZTPNYH[PVUHZZH`JHSJLPUSHILSLK
4+(4)JLSSZ^LYLZ[YH[PÄLKV]LY/<=,*PU[YHUZ^LSSJOHTILYZHUK[OLU\TILY
VM[YHUZTPNYH[PUN[\TVYJLSSZJV\U[LK\ZPUNÅ\VYLZJLU[TPJYVZJVW`;\I\SVNLULZPZ
^HZWLYMVYTLKI`ZLLKPUN/<=,*VUTH[YPNLSHUKJV\U[PUN[OLU\TILYVMIYHUJO
WVPU[ÄLSK \UKLY ? THNUPÄJH[PVU 0U ]P]V HUNPVNLULZPZ ^HZ KVUL \ZPUN TH[YPNLS
WS\NZPTWSHU[LKZ\IJ\[HULV\ZS`PUH[O`TPJU\U\TPJL:L]LUKH`ZSH[LYWS\NZ
^LYL^LYLYLTV]LKHUK[OLOLTVNSVIPUJVU[LU[HZZH`LK
9LZ\S[Z! 3.(3:)7 RUVJRKV^U PU 4+(4) JLSSZ ^HZ HJJVTWHUPLK [V H
KLJYLHZLK JLSS HKOLZPVU [V ÄIYVULJ[PU H YLK\JLK [YHUZLUKV[OLSPHS TPNYH[PVU HUK
TVYL PTWVY[HU[S` H YLK\JLK L_WYLZZPVU VM =,.- 7YVK\J[PVU VM =,.- ^OPJO ^HZ
YLZ[VYLKI`L_WVZ\YLVMZPSLUJLKJLSSZ[VYLJVTIPUHU[3.(3:)7YLX\PYLKHUPU[HJ[
702(R[ ZPNUHSPUN -\Y[OLYTVYL 3.(3:)7 ^HZ HISL [V KPYLJ[S` Z[PT\SH[L PU ]P[YV
/<=,* [\I\SVNLULZPZ HUK PU ]P]V HUNPVNLULZPZ PU H =,.-PUKLWLUKLU[ THUULY
0TT\UVOPZ[VJOLTPJHS HUHS`ZPZ VM O\THU IYLHZ[ JHUJLY [PZZ\LZ ZOV^LK H WVZP[P]L
JVYYLSH[PVU IL[^LLU 3.(3:)7 L_WYLZZPVU =,.- L_WYLZZPVU HUK ISVVK ]LZZLS
KLUZP[`
*VUJS\ZPVUZ! 0U HKKP[PVU [V P[Z WYVTL[HZ[H[PJ YVSL 3.(3:)7 ZLJYL[LK I` O\THU
IYLHZ[JHUJLYJLSSZTH`M\UJ[PVUHZHWYVHUNPVNLUPJMHJ[VY[OYV\NOHK\HSTLJOHUPZT
PLI`PUK\J[PVUVM[\TVY=,.-HUKZ[PT\SH[PVUVMLUKV[OLSPHSJLSSZHUNPVNLULZPZ
07VZ[LY:LZZPVU
IMB 3
Immunologia Biochimica
,=HUV]LSO\THUPaLK,YI)HU[PIVK`Z\WWYLZZLZ
59.ơ,YI)(2;H_PZPU]P[YVHUKIVVZ[[OLHU[P[\TVY
HJ[P]P[`VM[YHZ[\a\THIPU_LUVNYHM[LK)_7*WHUJYLH[PJ
[\TVYPUTPJL
./(:,409,A( 0(*6),330:;,-(56(5+:(3(.0(53<*(
+PWHY[PTLU[VKP:JPLUaL)PVTLKPJOL<UP]LYZP[H»¸.K»(UU\UaPV¹*OPL[P7LZJHYH
-VUKHaPVUL¸.+»(UU\UaPV*LU[YV:[\KPZ\SS»0U]LJJOPHTLU[V
*L:0=PH*VSSLKLSS»(YH*OPL[P0;(3@
4LKPHWOHYTHZYS=PHKLP=LZ[PUP*OPL[P0;(3@
+LYLN\SH[LK ZPNUHSPUN [OYV\NO [OL ,.-9 ,YI) HUK ,YI) TLTILYZ VM [OL
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IMB 4
Immunologia Biochimica
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Immunologia Biochimica
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07VZ[LY:LZZPVU
IMB 6
Immunologia Biochimica
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07VZ[LY:LZZPVU
IMB 7
Immunologia Biochimica
0KLU[PÄJH[PVUJSVUPUNHUKM\UJ[PVUHSJOHYHJ[LYPaH[PVUVM
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Sviluppo Differenziamento e Apoptosi
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SDA 2
Sviluppo Differenziamento e Apoptosi
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9LJLU[ÄUKPUNZZ\NNLZ[HUV]LSWO`ZPVSVNPJHSYVSLMVYLUKVJHUUHIPUVPKZHZZPNUHSPUN
TVSLJ\SLZYLZWVUZPISLMVY[OLJVU[YVSVMWYVSPMLYH[PVUHUKKPMMLYLU[PH[PVUIV[OJLU[YHSS`
HUKWLYPWOLYHSS`/V^L]LYSP[[SLPZRUV^UJVUJLYUPUNOHLTH[VWVPL[PJJLSSKPMMLYLU[PH[PVU
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^HZ UV[ L_WYLZZLK TLNHRHY`VJ`[LZ -\Y[OLYTVYL HKKP[PVU VM HUHUKHTPKL K\YPUN
KPMMLYLU[PH[PVU ZPNUPÄJHU[S` PUJYLHZLK [OL U\TILY VM PU[LYTLKPH[L HUK M\SS` TH[\YL
JLSSZ]PH*)YLJLW[VYZPNUHSSPUN^OPSLZPNUHSPUN[OYV\NO*)YLJLW[VYZYLK\JLK[OL
U\TILYVMM\SS`TH[\YLTLNHRH`VJ`[LZ/V^L]LYUVULLUKVJHUUHIPUVPK^HZHISL[V
HMMLJ[[OLKPMMLYLU[PH[PVUKLNYLLVM[OLJLSSZZPUJL*+L_WYLZZPVUHUKJLSSWSVPK`^HZ
\UHS[LYLKI`LUKVJHUUHIPUVPK[YLH[TLU[
*VUJS\ZPVUZ! 0U SPNO[ VM HSS [OLZL ÄUKPUNZ ^L JHU ZWLJ\SH[L HU PU]VS]LTLU[ VM [OL
LUKVJHUUHIPUVPKZ`Z[LTPU[OLYLN\SH[PVUVM[OLU\TILYVMTH[\YPUNTLNHRHY`VJ`[LZ
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WH[O^H`ZHJ[P]H[LKKV^UZ[YLHTVM*)YLJLW[VYZK\YPUNTLNHRHY`VJ`[VWVPLZPZ
07VZ[LY:LZZPVU
SDA 3
Sviluppo Differenziamento e Apoptosi
Ʒ7<-(9,+<*,*/63,:;,963)06:@5;/,:0:(5+
+6?69<)0*059,:0:;(5*,05*6365*(5*,9*,33:
90.(5;0*/0(9(*69:,;;67(63((465;69-(56.0.3063(
*9,465((5+9,(./0.6+(906)6:0((4(30(90AA6(5.,3(4
+PWHY[PTLU[VKP6UJVSVNPH<UP]LYZP[nKLNSP:[\KPKP;VYPUV
+PWHY[PTLU[VKP:JPLUaL-HYTHJVSVNPJOLL)PVTVSLJVSHYP<UP]LYZP[nKLNSP:[\KPKP4PSHUV
4\S[PKY\N YLZPZ[HUJL 4+9 PUKPJH[LZ H KLJYLHZL PU ZLUZP[P]P[` VM JHUJLY [V H ^PKL
]HYPL[`VMJ`[V[V_PJJVTWV\UKZHZKV_VY\IPJPU0UMHJ[[OLTVZ[MYLX\LU[KYH^IHJR
VMKV_VY\IPJPU[YLH[TLU[PZ[OLVUZL[VMKY\NYLZPZ[HUJLK\L[VP[ZHJ[P]LLMÅ\_MYVT
JHUJLYJLSSZ[OYV\NO7NS`JVWYV[LPU7NW(S[OV\NOHJLU[YHSYVSLOHZILLUHZJYPILK
[V7NWPU49+SPWPKTLTIYHULLU]PYVUTLU[HSZVHWWLHYZ[VILPTWVY[HU["7NWPZ
TLTIYHUL[YHUZWVY[LYHUKPZTHPUS`SVJHSPaLKPUSPWPKYHM[Z0UWHY[PJ\SHYTLTIYHUL
JOVSLZ[LYVSPZLZZLU[PHS[VTHPU[HPUHOPNO7NWHJ[P]P[`JVUZLX\LU[S`[\TVYZ^P[O
H4+9WOLUV[`WLOH]LOPNOLYJVUJLU[YH[PVUZVMWSHZTHTLTIYHULJOVSLZ[LYVS[OHU
JOLTVZLUZP[P]LVULZ
:L]LYHSZ[\KPLZOH]LKLTVUZ[YH[LK[OH[ƷWVS`\UZH[\YH[LKMH[[`HJPKZƷ7<-(
PU[HRLPZJVYYLSH[LK[VSV^PUJPKLUJLVMJVSVYLJ[HSJHUJLY;OLTLJOHUPZTZI`
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O`WV[OLZLZ PUKPJH[LZ [OH[ [OL` TH` HSZV JOHUNL [OL Å\PKP[` VM JLSS TLTIYHUL HUK
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7YLSPTPUHY`YLZ\S[ZPUKPJH[L[OH[+/(HUK,7(HYLPUJVYWVYH[LKPUWSHZTHTLTIYHUL
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ZLUZP[P]LVULZHUKPZKLJYLHZLKKVZLKLWLUKLU[S`I`7<-([YLH[TLU[PUIV[OJLSS
SPULZ 4VYLV]LY Ʒ 7<-( LZWLJPHSS` +/( KL[LYTPUL H YLK\J[PVU VM /4.*V(9
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JVU[LU[
-PUHSS`+/(PUK\JLZKV_VY\IPJPUHJJ\T\SH[PVUPUKY\NYLZPZ[HU[/; K_JLSSZHUK
YLK\JLZ[OL7NWHJ[P]P[`TLHZ\YLKHZ[OLHIPSP[`[VL_[Y\KLYOVKHTPUL
;OLL]HS\H[PVUVMSPWPKYHM[7NWSVJHSPaH[PVUHUKSPWPKJVTWVZP[PVU^PSSNP]L[OLPUKPJH[PVU
VMHKPYLJ[LMMLJ[VMƷ7<-(VUYHM[Z[Y\J[\YLHUKM\UJ[PVU
2VWLJRH11V\YUHSVM*VU[YVSSLK9LSLHZL
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SDA 4
Sviluppo Differenziamento e Apoptosi
5L^HKLUPSPJKPU\JSLV[PKLZ^P[OHU[PSL\RLTPJHJ[P]P[`
:*(9-Ó:650()9<AA65,:(5;05(.06=05,4(9*6
+,-369((5;6506(5+A6**/0,3,5(
+LW[;LYYP[VY`,U]PYVUTLU[HUK3PML<UP]LYZP[`VM.LUV]H"
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9LJLU[S` (+7YPIVZ`S J`JSHZLZ MYVT [OL THYPUL ZWVUNL (_PULSSH WVS`WVPKLZ HUK
MYVTO\THU*+KLTVUZ[YH[LK[VZ`U[OLZPaL[^VKPHKLUVZPULOVTVKPU\JSLV[PKLZ
MYVTJ(+79HUKHKLUPULKLZPNUH[LK7HUK7^OPJOHYLJOHYHJ[LYPaLKI`HU
\U\Z\HS5NS`JVZPKPJIVUKIL[^LLUVULVM[OLHKLUPULZHUKYPIVZLBD
7 HUK 7 HMMLJ[ [OL B*HDP ^OLU HWWSPLK L_[YHJLSS\SHYS` [V PU[HJ[ JLSSZ UV[
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VU[OLPYJHWHJP[`[VTVK\SH[L[OLW\YPULYNPJYLJLW[VY7?HUK7@BD0UWHY[PJ\SHY
7 HJ[P]H[LZ [OL W\YPULYNPJ YLJLW[VYJOHUULS 7? LSPJP[PUN PUÅ\_ VM L_[YHJLSS\SHY
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VIZLY]H[PVUZPUKPJH[L[OH[JLSSZMYVTT`LSVPKHUKS`TWOVJ`[PJSL\RLTPHZZOV\SKIL
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[OH[ 7 PUK\JLZ JLSS KLH[O I` HWVW[VZPZ H[ Z\ITPJYVTVSHY JVUJLU[YH[PVUZ ^OPSL
7PZLMMLJ[P]LH[TPJYVTVSHYJVUJLU[YH[PVUZ0U[OLZHTLJVUKP[PVUZJLSSZPZVSH[LK
MYVTOLHS[O`KVUVYZ^LYLUV[HMMLJ[LKI`7HUK7PUKPJH[PUNHZWLJPÄJJ`[V[V_PJ
LMMLJ[VUSL\RLTPJHUKVUZVSPK[\TVYJLSSZ-\Y[OLYTVYL^LJV\SKJVUÄYT[OH[[OL
J`[V[V_PJHJ[P]P[`VM7^HZTHPUS`H[[YPI\[HISL[V[OLKPZZPWH[PVUVM[OLTP[VJOVUKYPHS
WYV[VUNYHKPLU[[OYV\NO[OLVWLUPUNVM[OLWLYTLHIPSP[`[YHUZP[PVUWVYLYLZ\S[PUNPU
JLSSKLH[O
0UJVUJS\ZPVU^LZ\NNLZ[[OLWVZZPISL\ZLVM7HZHKY\NMVYJHUJLY[OLYHW`PU
OHLTH[VSVNPJHSTHSPNUHUJPLZ^OPSLVUNVPUNZ[\KPLZHYL[Y`PUN[VPKLU[PM`M\Y[OLYJLSS
[`WLZZLUZP[P]L[V[OPZUL^HU[P[\TVYKY\N
)HZPSL.L[HS7YVJ5H[S(JHK:JP<:( )Y\aaVUL:L[HS1)PVS*OLT
)Y\aaVUL:L[HS1)PVS*OLT
07VZ[LY:LZZPVU
SDA 5
Sviluppo Differenziamento e Apoptosi
*LYHTPKLHJ[ZHZHTLKPH[VYVM[OLHU[PWYVSPMLYH[P]LHUK
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JVSVUJHUJLYJLSSZ
()+,3/(+036<)5(.0<::(507(63(=0(507(63(90)6503(<9(
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;OLZWOPUNVSPWPKJLYHTPKLWSH`ZHUPTWVY[HU[YVSLHZHU[PJHUJLYJLSS\SHYTLKPH[VYHUK
PZPTWSPJH[LKPUJOLTV[OLYHW`PUK\JLKHWVW[VZPZI`KPMMLYLU[HNLU[Z/\THUJVSVU
JHUJLYZ [OL ZLJVUK TVZ[ SLHKPUN JH\ZL VM JHUJLYYLSH[LK KLH[OZ PU [OL ^LZ[LYU
JV\U[YPLZ OH]L VUS` OHSM [OL JLYHTPKL JVU[LU[ VM UVYTHS [PZZ\L HUK [OLZL SV^
JLYHTPKL SL]LSZ OH]L ILLU JVYYLSH[LK ^P[O [OLPY PUJYLHZLK YLZPZ[HUJL [V HWVW[VZPZ
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IL[^LLU[OL\URUV^UHU[PJHUJLYTLJOHUPZTVMS\[LVSPUHUKJLYHTPKLPUJVSVUJHUJLY
JLSSZ
<ZPUN[OLO\THUJVSVUJHUJLYJLSSSPUL*HJVHZJLSSTVKLS^LPUP[PHSS`MV\UK[OH[
S\[LVSPUHKTPUPZ[YH[PVU^HZHZZVJPH[LK[V[OLPUOPIP[PVUVMJLSSWYVSPMLYH[PVUHUKH[OPNOLY
KVZLZ[VJLSSKLH[O0UM\Y[OLYL_WLYPTLU[Z^LPU]LZ[PNH[LK[OLJLYHTPKLTL[HIVSPZT
PU*HJVJLSSZHUK[OLWVZZPISLLMMLJ[ZVMS\[LVSPUVUP[\ZPUNYHKPVSHILSLKZWOPUNVZPUL
HZWYLJ\YZVY;OLYLZ\S[ZZOV^LK[OH[*HJVJLSSZLMÄJPLU[S`HUKYHWPKS`NLULYH[L
IV[O JLYHTPKL HUK ZWOPUNVT`LSPU MYVT ZWOPUNVZPUL HUK [OH[ S\[LVSPU [YLH[TLU[
PUK\JLZHKVZLKLWLUKLU[YLK\J[PVUVM[OLZWOPUNVT`LSPUIPVZ`U[OLZPZHZZVJPH[LK
[VHZPNUPÄJHU[PUJYLHZLVM[OLJLSS\SHYJLYHTPKL;OPZL]PKLUJLSL[\ZO`WV[OLZPaL[OH[
JLYHTPKLTH`ILPU]VS]LKPU[OLHU[PWYVSPMLYH[P]LHUK[V_PJLMMLJ[VMS\[LVSPU6U[OLZL
WYVTPZLZ^LL]HS\H[LK[OLLMMLJ[VMJLYHTPKLVUJLSSZ\Y]P]HS>LMV\UK[OH[LP[OLY[OL
HKTPUPZ[YH[PVUVMJLYHTPKLHUHSVN\LZVY[OLPUOPIP[PVUVMZWOPUNVT`LSPUIPVZ`U[OLZPZ
^LYLHISL[VPUK\JL*HJVJLSSKLH[O[O\ZKLTVUZ[YH[PUN[OH[JLYHTPKLPZHISL[V
TPTPJ[OLJ`[V[V_PJLMMLJ[VMS\[LVSPU
6]LYHSS[OLZLYLZ\S[ZKLTVUZ[YH[L[OH[S\[LVSPUJHUHJ[HZHUHU[PWYVSPMLYH[P]LHUKWYV
KLH[OHNLU[PU*HJVJLSSZI`\WYLN\SH[PUNJLSS\SHYJLYHTPKL[OYV\NOPUOPIP[PVUVM
P[ZTL[HIVSPZT[VZWOPUNVT`LSPUHUKZ\NNLZ[HWV[LU[PHSYVSLVMS\[LVSPUJLYHTPKLHZ
HU[PJHUJLYHNLU[PUJVSVUTHSPNUHUJPLZ
07VZ[LY:LZZPVU
SDA 6
Sviluppo Differenziamento e Apoptosi
0ZVSH[PVUHUKJOHYHJ[LYPaH[PVUVMO\THUYV[H[VYJ\MM[LUKVU
Z[LTJLSSZ
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*090336-,+,90*(),9.(5;,:650(:*(905.09(--(,33(
./0963+0(5+9,(9(.65,=05*,5A(4(:<AA67(4,3(
*()0;A(7(636;,;;(4(5;0.<0+6.(.30(5650*63,;;(
9(5+,33070,;96(5+(5(:;(:0(3<0.0
09**:7VSPJSPUPJV:HU+VUH[V:HU+VUH[V4PSHULZL0[HS`"
+LWHY[TLU[VM)PVTLKPJHS:JPLUJLZMVY/LHS[O<UP]LYZP[`VM4PSHU4PSHU0[HS`
9V[H[VY J\MM 9* [LUKVUZ HYL VM[LU WYVUL [V SLZPVUZ HUK YHW[\YLZ HZ [V VM [OL WVW\SH[PVU V]LY ÄM[` Z\MMLYZ VM WHY[PHS HUK M\SS[OPJRULZZ 9* [LHYZ :L]LYHS
HWWYVHJOLZOH]LILLUKL]LSVWLKV]LY[OL`LHYZPUJS\KPUN[OL\ZLVMNYV^[OMHJ[VYZ
IVUL TVYWOVNLUL[PJ WYV[LPUZ HUK TVYL YLJLU[S` Z[LT JLSSZ (TVUN HK\S[ Z[LT
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HS[OV\NO[LUKVUKLYP]LKWYVNLUP[VYJLSSZ;:7*ZOH]LILLUMV\UKPUZL]LYHSHUPTHS
ZWLJPLZPUJS\KPUNO\THUZ/V^L]LY[OLPZVSH[PVUVMHJLSSWVW\SH[PVU^P[OZ[LT
JLSSZJOHYHJ[LYPZ[PJZMYVT[OLO\THUYV[H[VYJ\MMOHZ`L[[VILYLWVY[LK
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MYVT [OL SVUN OLHK IPJLW 3/) [LUKVUZ *LSSZ JV\SK IL J\S[\YLK HUK L_WHUKLK
HUK[OL`ZOV^LKHK\S[Z[LTJLSSJOHYHJ[LYPZ[PJZPL[OL`HYLZLSMYLUL^PUNPU]P[YV
JSVUVNLUPJ HUK T\S[PWV[LU[ HZ [OL` JV\SK IL PUK\JLK [V KPMMLYLU[PH[L PU ]P[YV PU[V
KPMMLYLU[ JLSS [`WLZ PUJS\KPUN VZ[LVISHZ[Z HKPWVJ`[LZ HUK ZRLSL[HS T\ZJSL ;OLZL
UL^ JLSS WVW\SH[PVUZ ^LYL WOLUV[`WPJHSS` JOHYHJ[LYPaLK HUK JVTWHYLK [V )4:*Z
HUK[VO\THUKLYTHSÄIYVISHZ[Z+-Z4VYLV]LYHZZWOPUNVSPWPKZHYLLTLYNPUNRL`
YLN\SH[VY` TVSLJ\SLZ VM Z[LT JLSS WYVSPMLYH[PVU HUK KPMMLYLU[PH[PVU [OL ZWOPUNVSPWPK
WH[[LYUVMHSSJLSSZOHZILLUKL[LYTPULK
3\P77HUK*OHU24:[LTJLSSYL]PL^Z )P@,OPYJOPV\+2PS[Z;40URZVU*(,TIYLL4*:VUV`HTH>3P33LL[(0:LV)
4AOHUN3:OP:HUK@V\UN4-5H[\YLTLKPJPUL 07VZ[LY:LZZPVU
SDA 7
Sviluppo Differenziamento e Apoptosi
(JVTWHYPZVUIL[^LLU[OLYVSLVM:7(9*PUVZ[LVNLUPJ
KPMMLYLU[PH[PVUVMTLZLUJO`THSZ[LTJLSSZHUKPU>05
;9(03PUK\JLKHWVW[VZPZPUVZ[LVZHYJVTHJLSSZ
(56;(96::(),33(67,33,90;64.0<30(56(5+.*(3=(9<:6
+PWHY[PTLU[VKP)PVTLKPJPUH:WLYPTLU[HSLL5L\YVZJPLUaL*SPUPJOL
ZLaPVULKP)PVJOPTPJH<UP]LYZP[nKLNSP:[\KPKP7HSLYTV
4LZLUJO`THS Z[LT JLSSZ HYL ^LSS RUV^U [V WVZZLZZ T\S[PWV[LU[PHS KPMMLYLU[PH[P]L
HIPSP[`HUKYLWYLZLU[HNVVK[VVSMVYJSPUPJHSYLZLHYJO
0UV\YZ[\K`HM[LYPUK\J[PVUVMVZ[LVNLUPJKPMMLYLU[PH[PVUVMIVULTHYYV^TLZLUJO`THS
Z[LT JLSSZ )44:* I` \ZPUN JVUKP[PVULK TLKPH ^L MVJ\ZLK V\Y H[[LU[PVU VU
:7(9* H YLN\SH[VY` WYV[LPU ^OPJO HMMLJ[Z JLSS KPMMLYLU[PH[PVU WYVSPMLYH[PVU HUK
Z\Y]P]HS +\YPUN VZ[LVNLUPJ KPMMLYLU[PH[PVU :7(9* SL]LSZ YHPZLK PU [PTLKLWLUKLU[
THUULY HUK ^LYL M\UJ[PVUHS [V [OL PUJYLHZL VM V[OLY VZ[LVNLUPJ THYRLYZ Z\JO HZ
VZ[LYP_ HUK VZ[LVWVU[PU ;OL LMMLJ[Z VM :7(9* KV^UYLN\SH[PVU ^LYL PU]LZ[PNH[LK
[OYV\NO[OL\ZLVMHZWLJPÄJZTHSSPU[LYMLYPUN95(KLTVUZ[YH[PUNHYLK\J[PVUPU[OL
SL]LSZVMVZ[LVNLUPJTHYRLYZPUZP:7(9*JLSSZ
:PUJL:7(9*ILZPKLZ[VP[ZPTWVY[HU[YVSLPUVZ[LVNLUPJKPMMLYLU[PH[PVUJHUTVK\SH[L
JLSSJLSSHUKJLSSTH[YP_PU[LYHJ[PVUZHUKHJ[LP[OLYHZH[\TVYZ\WWYLZZVYVYH[\TVY
WYVTV[LY^L^LYLPU[LYLZ[LK[VPU]LZ[PNH[LP[ZSL]LSZPUVZ[LVZHYJVTH4.JLSSZH
THSPNUHU[VZ[LVISHZ[PJSPRLJLSSSPUL0U[OLZLJLSSZ[OLIHZHSSL]LSZVM:7(9*YLZ\S[LK
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07VZ[LY:LZZPVU
SDA 8
Sviluppo Differenziamento e Apoptosi
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07VZ[LY:LZZPVU
SDA 9
Sviluppo Differenziamento e Apoptosi
.SPVISHZ[VTHT\S[PMVYTL!WOHYTHJVSVNPJHSYLHJ[P]H[PVUVM
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BD4H`V3+L[HS1)PVS*OLT" 07VZ[LY:LZZPVU
SDA 10
Sviluppo Differenziamento e Apoptosi
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SDA 11
Sviluppo Differenziamento e Apoptosi
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07VZ[LY:LZZPVU
SDA 12
Sviluppo Differenziamento e Apoptosi
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07VZ[LY:LZZPVU
SDA 13
Sviluppo Differenziamento e Apoptosi
,MMLJ[ZVM[OLPUQLJ[PVUVMWOHYTHJVSVNPJHSS`HJ[P]L
TPJYVJHYYPLYZYLSLHZPUN/.-0.-HUK[YHUZWVY[PUN
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)PVJOPTPJH<UP]LYZP[nKP)VSVNUH+PWHY[PTLU[VKP7H[VSVNPHLTLKPJPUHKPSHIVYH[VYPV
<UP]LYZP[nKP7HYTH0Z[P[\[V5HaPVUHSLWLYSL9PJLYJOL*HYKPV]HZJVSHYP059*)VSVNUH
05:,94<L<UP]LYZP[nKP(UNLYZ-YHUJPH+PWHY[PTLU[VKP:JPLUaL*OPY\YNPJOL
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KH[HPUKPJH[L[OH[[OLSHYNLZ[PTWYV]LTLU[PUJHYKPHJTLJOHUPJZVJJ\YYLKPU(+:*.-
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07VZ[LY:LZZPVU
SDA 14
Sviluppo Differenziamento e Apoptosi
9VSLVMV_PKH[P]LZ[YLZZPUJOLTPJHSO`WV_PHPUK\JLK
HWVW[VZPZ
*9,465050,3,6569(964(50(90(55(*,9=,33(;0-9(5*6
*,9=,33(;0*(9367(=(5)(9)(9(:;0*6AA0*3(<+0((5+
=(3(**/0.0<:,77,
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+LWHY[TLU[VM)PVSVN`HUK,]VS\[PVU<UP]LYZP[`VM-LYYHYH0[HS`
+LWHY[TLU[VM-VVKHUK5\[YP[PVU2`\UN/LL<UP]LYZP[`:LV\S:V\[O2VYLH
7YVSPMLYH[P]LYL[PUVWH[OPLZHYL[OLTHQVYJH\ZLVMISPUKULZZPUPUK\Z[YPHSPaLKJV\U[YPLZ
HUK[OL`HYLJOHYHJ[LYPaLKI`UL\YVUHSKLWVSHYPaH[PVUJHSJP\TPUÅ\_HUKV_PKH[P]L
Z[YLZZ6:;OPZJVUKP[PVUL]VRLZHU\UIHSHUJLKYLSLHZLVMWYVHUKHU[PHUNPVNLUPJ
MHJ[VYZMYVT[OLYL[PUHSWPNTLU[LWP[OLSP\T97,[V^HYKZ[OLWYVHUNPVNLUPJVULZ^P[O
HJVUZLX\LU[HILYYHU[UL^]LZZLSMVYTH[PVU[OH[[VNL[OLY^P[OUL\YVKLNLULYH[PVU
JH\ZLZPYYL]LYZPISL]PZ\HSSVZZ0[PZ^LSSRUV^U[OH[O`WV_PHHUKPUJYLHZLVMYLHJ[P]L
V_`NLUZWLJPLZ96:^OPJOJVTTVUS`HJJVTWHU`WYVSPMLYH[P]LYL[PUVWH[OPLZHS[LY
[OLIHSHUJLIL[^LLUWYVHUKHU[PHUNPVNLUPJMHJ[VYZ0U[OLWYLZLU[Z[\K`^LOH]L
L]HS\H[LKHWVZZPISLYVSLVM6:PU[OLTVK\SH[PVUVMO`WV_PHHJ[P]H[LK/(MHJ[VYZ
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^P[O[OLPUJYLHZLKL_WYLZZPVUVM=,.-JVUÄYTLK[OH[[OL/97,JLSSZYLZWVUK[V[OL
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JLSSZ[V[OLPUJYLHZLK6:PZ[OLPUK\J[PVUVM5-R)HJ[P]P[`HUK[OPZ^HZJVUÄYTLK
I`PTT\UVO`Z[VJOLTPZ[Y`HUHS`ZPZ0UHKKPJ[PVU[OLPUJYLHZLVM6:PU[OLZLJLSSZPZ
ZOV^UI`ÅV^J`[VTL[Y`
;OLZL YLZ\S[Z WYV]PKL L]PKLUJL VM [OL WVZZPISL TLJOHUPZ[PJ WH[O^H` PU]VS]LK PU
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JLSSZTVY[HSP[`Z\NNLZ[[OH[6:^PSSPUK\JLHWVW[VZPZ
07VZ[LY:LZZPVU
SDA 15
Sviluppo Differenziamento e Apoptosi
9L]LYZPULPUK\JLZLUKVYLWSPJH[PVUVUS`^OLUJLSSJ`JSL
JOLJRWVPU[ZHYLKLYLN\SH[LK
70**6304(9*67(3(AA636.0(*646*65-69;0,902(
*9,67(:8<(3,7(70505(+0(=,5,9(5+6)9<56
;,;;(4(5;0.<0+6(5+(5(:;(:0(3<0.0
3HIVYH[VY`VM:[LT*LSSZMVY;PZZ\L,UNPULLYPUN09**:7VSPJSPUPJV:HU+VUH[V
:HU+VUH[V4PSHULZL0[HS`"
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;OL Z`U[OL[PJ W\YPUL YL]LYZPUL OHZ ILLU ZOV^U [V WVZZLZZ H K\HS HJ[P]P[` HZ P[
WYVTV[LZKLKPMMLYLU[PH[PVUVMHK\S[JLSSZPUJS\KPUNÄIYVISHZ[ZBDI\[P[HSZVPUK\JLZ
JLSS NYV^[OHYYLZ[ HUK \S[PTH[LS` JLSS KLH[O VM JHUJLY JLSSZ 7YL]PV\Z TLJOHUPZ[PJ
Z[\KPLZYL]LHSLK[OH[[OLTVSLJ\SLPZHUPUOPIP[VYVM4,2HUKVMUVUT\ZJSLT`VZPU
OLH]` JOHPU 54400 HUK WVZZLZZ H WV[LU[ PUOPIP[VY` JHWHJP[` VU 47: H JY\JPHS
RPUHZLVM[OLZWPUKSLHZZLTIS`JOLJRWVPU[/V^L]LY[OLHWWHYLU[ZLSLJ[P]P[`
VMYL]LYZPULMVYJHUJLYJLSSZULLKLKM\Y[OLYLS\JPKH[PVULZWLJPHSS`PU[OLWLYZWLJ[P]L
VM H WVZZPISL HWWSPJH[PVU VM [OL TVSLJ\SL HZ HU HU[PJHUJLY KY\N 6U [OLZL IHZLZ
^LWLYMVYTLKHJVTWHYH[P]LHUHS`ZPZVM[OLLMMLJ[ZVMYL]LYZPULVUZL]LYHSO\THU
[\TVYJLSSZHUKUVYTHSO\THUÄIYVISHZ[Z*LSSJ`JSLHUHS`ZPZYL]LHSLK[OH[YL]LYZPUL
[YLH[TLU[JH\ZLK[OLMVYTH[PVUVMTP_LKKPWSVPK[L[YHWSVPKWVW\SH[PVUZPUHSS[YLH[LK
JLSSZ /V^L]LY ^OPSL UVYTHS ÄIYVISHZ[Z ZOV^LK [OL WYLZLUJL VM [^V WVW\SH[PVUZ
KPWSVPK HUK [L[YHWSVPK PU [OLPY YLZWLJ[P]L . WOHZLZ HSS JHUJLY JLSSZ JVU[PU\LK [V
WLYMVYT HILYYHU[ U\JSLP KP]PZPVUZ ^P[OV\[ J`[VRPULZPZ SLHKPUN [V [OL PYYL]LYZPISL
MVYTH[PVUVMT\S[PU\JSLH[LKHUL\WSVPKJLSSZ[OH[KPLKVMTP[V[PJJH[HZ[YVWOL;OLZL
L]PKLUJLZZLLT[VZ\WWVY[[OLO`WV[OLZPZ[OH[YL]LYZPUL[YLH[TLU[JV\SKILSL[OHSMVY
JLSSZ^P[OKLYLN\SH[LKJLSSJ`JSLJOLJRWVPU[ZHZ[OL`JHU»[ISVJRLUKVYLWSPJH[PVU0U
MHJ[JOLTPJHSHUKNLUL[PJHSPUOPIP[PVUVMWPUÄIYVISHZ[ZYLUKLYZ[OLTZLUZP[P]L[V
YL]LYZPUL[YLH[TLU[HUKJLSSZKPLKVMHILYYHU[LUKVYLWSPJH[PVUZHUHSVNV\ZS`[V^OH[
P[^HZVIZLY]LKPUJHUJLYJLSSZ
BD(UHZ[HZPH3L[HS*LSS+LH[OHUK+PMMLYLU[PH[PVU!
BD*OLU:L[HS75(:!
BD:HU[HN\PKH:L[HS1*) !
07VZ[LY:LZZPVU
SDA 16
Sviluppo Differenziamento e Apoptosi
:`ULYNPJLMMLJ[ZVMHSS[YHUZYL[PUVPJHJPKPU
SPTWOVTVUVJ`[LZJVJ\S[\YLZVU[OLNYV^[OVMO\THU
RLYH[PUVJ`[LZHUK.904 L_WYLZZPVU
:(9+(9650*63(7(7(-9(5*,:*6;(;<3364(9*6
50*(:;96(55(90;(+»690(:<:(55(3(0:6.0<30(5()4(99,330
4(::046369<::640*/,3,3077630:96:((5+:*(**6:(3=(;69,
+LWHY[TLU[VM)HZPJ4LKPJHS:JPLUJLZ<UP]LYZP[`VM)HYP¸(4VYV¹"
*HSHIYVKLU[HSZYS*YV[VUL"0)),*59)HYP
/LHKHUKULJRZX\HTV\ZJLSSJHYJPUVTH/5:**YHURZHTVUN[OLTVZ[JVTTVU
JHUJLYZ *`[VRPULZ HUK PUÅHTTH[VY` ZPNUHS WH[O^H`Z HWWLHY [V IL PTWSPJH[LK
PU [OL KL]LSVWTLU[ VM /5:** HZ ^LSS HZ V[OLY VYHS [PZZ\LZ WYLULVWSHZ[PJ SLZPVUZ
VM \URUV^U VYPNPU *`[VRPULZ YLN\SH[L PTT\UP[` PUÅHTTH[PVU HUK JLSS NYV^[O HUK
PUJS\KL PU[LYSL\RPUZ 03Z PU[LYMLYVUZ 0-5Z [\TVY ULJYVZPZ MHJ[VYZ ;5-Z HUK
NYV^[O MHJ[VYZ 6\Y Z[\KPLZ YL]LHS H ZPNUPÄJHU[ PUOPIP[VY` LMMLJ[ VM HSS[YHUZ YL[PUVPJ
HJPKVURLYH[PUVJ`[LZNYV^[O;OPZLMMLJ[HWWLHYZ[VILYLSH[LK[VPUJYLHZLKSL]LSZ
VMWYV[LPU.904 .LULZHZZVJPH[LK^P[O9L[PUVPK0-5PUK\JLK4VY[HSP[`B(UNLSS
1,L[HS1)PVS*OLT!"7HWH-L[HS0U[10TT\U7OHYTHJVS
! D ( UL^ L_WLYPTLU[HS HWWYVHJO I` JVJ\S[\YPUN RLYH[PUVJ`[LZ HUK
WLYPWOLYHSISVVKSPTWOVTVUVJ`[LZZOV^ZHZPUN\SHYZ`ULYNPJLMMLJ[VM(;9([YLH[TLU[
VUPUOPIP[PVUVMJLSSNYV^[O;OL(;9(LMMLJ[ZVU[OL[^VKPMMLYLU[JLSSSPULZTH`[YPNNLY
[OLTP[VJOVUKYPHSWH[O^H`VMHWVW[VZPZ[OYV\NOLUOHUJLTLU[VM.904 HJ[PVUZ
07VZ[LY:LZZPVU
SDA 17
Sviluppo Differenziamento e Apoptosi
5L^PUZPNO[ZPU[V[OLTLJOHUPZTVM152PUOPIP[PVUI`
NS\[H[OPVUL[YHUZMLYHZL7
*(**<90(55(4(90(+,3<*((5(:;(:0(-,+,90*03<*(+,
*(50640*/,3,:;,33(369,5A6
+LWHY[TLU[VM*OLTPJHS:JPLUJLZHUK;LJOUVSVNPLZ
<UP]LYZP[`VM¸;VY=LYNH[H¹9VTL0[HS`"
+LWHY[TLU[VM)PVTLKPJHS:JPLUJLZ<UP]LYZP[`VM*OPL[P.+»(UU\UaPV
*L:0*LU[LYVM,_JLSSLUJLVU(NPUN*OPL[P0[HS`"
+LWHY[TLU[VM0U[LYUHS4LKPJPUL<UP]LYZP[`VM9VTL;VY=LYNH[H
=PH4VU[WLSSPLY9VTL0[HS`
;OLYVSLWSH`LKI`NS\[H[OPVUL[YHUZMLYHZL7.:;7PUTVK\SH[PUN[OLJ1\U
5[LYTPUHSRPUHZL152WH[O^H`OHZILLUL_[LUZP]LS`PU]LZ[PNH[LK\ZPUN152PZVMVYTZ
RUV^U[VL_LY[VWWVZP[LLMMLJ[ZPU[OLJLSSZ>LOH]LL_WYLZZLK[OLPZVMVYT152Ơ
[OH[OHZILLUYLWVY[LK[V[YHUZTP[HWYVHWVW[V[PJZPNUHSHUK^LOH]LHUHS`aLKIV[O
[OLWOVZWOVY`SH[PVUSL]LSHUK[OLHJ[P]P[`VM[OPZRPUHZLPU[OLWYLZLUJLVM.:;7
*VU[YHY`[V^OH[WYL]PV\ZZ[\KPLZZ\NNLZ[^LMV\UK[OH[.:;7PZHISL[VMVYT
H JVTWSL_ ^P[O [OL \UWOVZWOVY`SH[LK HUK PUHJ[P]L 152Ơ PZVMVYT L]LU PU [OL
HIZLUJLVM[OLZ\IZ[YH[L>LHSZVHUHS`aLK[OLJVUZLX\LUJLZVM[OPZPU[LYHJ[PVUVU
[OLHJ[P]P[`VMIV[OLUa`TLZ;OLJVTWSL_Z[YVUNS`YLK\JLK[OLHJ[P]H[PVUVM152Ơ
^OPSLWYLZLY]LK.:;7MYVTPUHJ[P]H[PVU<UL_WLJ[LKS`NS\[H[OPVUL.:/L_LY[LKH
ULNH[P]LLMMLJ[VU[OLHMÄUP[`VM.:;7MVY152ƠZ\NNLZ[PUN[OH[[OLPU[YHJLSS\SHY
SL]LSZVM[OPZ[OPVSTH`HSSV^HÄUL[\UPUNVM[OL4(72ZPNUHSPUNWH[O^H`4VYLV]LY
^LMV\UK[OH[[OLHKK\J[MVYTLKI`.:/HUK[OLZ[YVUN.:;7PUOPIP[VY5)+/,?
HIVSPZOLZ[OLPU[LYHJ[PVUIL[^LLU[OL.:;7HUK152Ơ;OLZLKH[HJVUÄYTHUK
L_[LUKH[[OLTVSLJ\SHYSL]LSWYL]PV\ZL]PKLUJLVI[HPULKPU[\TVYJLSSSPULZ
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BIB 1
Biotecnologie e Biochimica Industriale
7LYV_PYLKV_PUZMVYUHUV[LJOUVSVN`!HIPVJOLTPJHS
HWWYVHJO[VWYVK\JLUHUVJVUK\J[VYZ
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:*6;;0:;,-(56),33,330(5+9,(:(**6**0(-<3=06
077630;096+63-6
+LWHY[TLU[VM3PML/LHS[OHUK,U]PYVUTLU[HS:JPLUJLZ<UP]LYZP[`VM3»(X\PSH
7SL:HS]H[VYL;VTTHZP*VWWP[V(80[HS`
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7SL(SKV4VYV9VTH0[HS`
*VYYLZWVUKPUNH\[OVY!7YVM9VKVSMV0WWVSP[P+LWHY[TLU[VM3PML
/LHS[OHUK,U]PYVUTLU[HS:JPLUJLZ<UP]LYZP[`VM3»(X\PSH7SL:HS]H[VYL;VTTHZP
*VWWP[V(80[HS`7OVUL! -H_! ;`WPJHS *`Z WLYV_PYLKV_PUZ 7Y_Z HYL \IPX\P[V\Z WYV[LPUZ ZOV^PUN [OL ZVJHSSLK
TVVUSPNO[PUN ILOH]PV\Y HZ [OL` HYL PU]VS]LK PU THU` JLSS\SHY HJ[P]P[PLZ! PU UVYTHS
JVUKP[PVUZ[OL`HJ[HZWLYV_PKHZLZHISL[VYLK\JL/[V^H[LY"[OPZM\UJ[PVUJH\ZLZ
HJOHUNLVMZ[Y\J[\YLMYVTYLK\JLKKLJHTLYYPUNZ[VV_PKPaLKKPTLYZ^OPJOPU[\YU
HYLYLK\JLKIHJRI`[OPVYLKV_PU0UZ[LHKPUV_PKH[P]LZ[YLZZJVUKP[PVUZ[OL`Z^P[JO
M\UJ[PVUSVVZPUN[OLWLYV_PKHZLHJ[P]P[`HUKHJX\PYPUNTVSLJ\SHYJOHWLYVULHUKJLSS
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L_JS\ZPVUJOYVTH[VNYHWO`HUK[YHUZTPZZPVULSLJ[YVUTPJYVZJVW`ZOV^LK[OH[[OLZL
ÄSHTLU[ZOH]LH[\I\SHYZ[Y\J[\YL^P[OSLUN[O\W[VUTHUKHYLJVUZ[P[\[LKI`
Z[HJRLKYPUNZWYLZLU[PUNPU[LYUHSHUKL_[LYUHSZ\YMHJLZ^OPJOHYLHTLUHISLVM
THUPW\SH[PVU0U[OPZMYHTL^VYR[OLZLWYV[LPUUHUV[\ILZ75;Z^LYLL_WSVP[LKMVY
HZZLTISPUNTL[HSUHUV^PYLZÄSSPUN[OLPYJH]P[`^P[OHUHYYH`VMNVSKUHUVWHY[PJSLZ
0U[OPZ^VYR^LWYLZLU[[OLZ[YH[LN`MVY[OLWYVK\J[PVUVMWV[LU[PHSOPNOJVUK\JPUN
UHUV^PYLZ WVZZPIS` Z\NNLZ[PUN H ¸NYLLU JOLTPZ[Y`¹ HWWYVHJO [V WYVK\JL TL[HSSPJ
UHUVJVUK\J[VYZ[OH[TH`IL\ZLKPUUHUVJPYJ\P[ZHUKUHUVKL]PJLZ
007VZ[LY:LZZPVU
BIB 2
Biotecnologie e Biochimica Industriale
0KLU[PÄJH[PVUVMUL^[OLYTVWOPSPJLUa`TH[PJHJ[P]P[PLZMVY
OLTPJLSS\SVZLJVU]LYZPVU
*6)<**0765A(56),(;90*,:;9(AA<330(5+9,(.0.30696:(
0(*65696),9;(4(:;<9A6.0<:,77,-(.5(564(::046(5+
469(**04(9*6
0UZ[P[\[LVM7YV[LPU)PVJOLTPZ[Y`¶*VUZPNSPV5HaPVUHSLKLSSL9PJLYJOL
=PH7*HZ[LSSPUV5HWSLZ0[HS`
+PWHY[PTLU[V0UNLNULYPHHNYHYPHL(NYVUVTPH<UP]LYZP[nKP5HWVSP-LKLYPJV00
]PH<UP]LYZP[n7VY[PJP5HWSLZ0[HS`
;OL\ZLVMIPVTHZZMVYLULYN`WYVK\J[PVUPZL_WLJ[LK[VPUJYLHZLPUM\[\YL`LHYZHUK
[OLWYVK\J[PVUVMIPVJOLTPJHSZHUKIPVM\LSZMYVTLULYN`JYVWZHYPZLZMYVT[OLULLK
[VÄUKHS[LYUH[P]LZV\YJLZ[VMVZZPSM\LS.PHU[YLLK(Y\UKVKVUH_3PZHWLYLUUPHS
NYHZZ ILPUN ZLLU HZ VUL VM [OL TVZ[ WYVTPZPUN IPVTHZZLZ MVY [OL WYVK\J[PVU VM
SPNUVJLSS\SVZPJIPVL[OHUVSVMZLJVUKNLULYH[PVU,Ua`TLJVJR[HPSZ[OH[O`KYVS`aLWSHU[
JLSS ^HSS WVS`ZHJJOHYPKLZ HYL H JYP[PJHS JVTWVULU[ VM IPVWYVJLZZPUN JVUÄN\YH[PVUZ
KLZPNULK [V [YHUZMVYT SPNUVJLSS\SVZPJ IPVTHZZ PU[V IPVM\LSZ ;OL SHYNL ]HYPL[` VM
WV[LU[PHSIPVTHZZMLLKZ[VJRZHUKH]HPSHISLWYL[YLH[TLU[ZYLX\PYL[HPSVYLKNS`JVZPKL
O`KYVSHZL WYLWHYH[PVUZ [OH[ M\UJ[PVU VW[PTHSS` \UKLY KP]LYZL JVUKP[PVUZ PUJS\KPUN
OPNO [LTWLYH[\YLZ HUK L_[YLTL W/ ;OPZ Z\NNLZ[Z [OH[ [OLYTVWOPSPJ WYVRHY`V[LZ
TH` IL HU PTWVY[HU[ ZV\YJL VM [OLZL LUa`TLZ 6\Y NYV\W PZ PU]VS]LK ZPUJL SVUN
[PTLPU[OLPKLU[PÄJH[PVUHUKJOHYHJ[LYPaH[PVUVM[OLYTVWOPSPJNS`JVZPKLO`KYVSHZLZ
/LYL^LYLWVY[[^VKPMMLYLU[HWWYVHJOLZHPTLK[VKPZJV]LYYVI\Z[HUKOPNOS`HJ[P]L
OLTPJLSS\SVZL KLNYHKPUN NS`JVZPKL O`KYVSHZLZ 0U [OL ÄYZ[ HWWYVHJO LUYPJOTLU[
J\S[\YLZ VU ( KVUH_ VM TPJYVVYNHUPZTZ SP]PUN PU [OL ZVSMH[HYPJ ÄLSK VM 7PZJPHYLSSP
5HWSLZ ^HZ WLYMVYTLK VU NPHU[ YLLK J\STZ HSSV^PUN [V YLK\JL [OL JVTWSL_P[` VM
[OLTPJYVIPHSJVTT\UP[PLZHUK[VZLSLJ[TPJYVVYNHUPZTHISL[VKLNYHKL[OL[HYNL[
IPVTHZZ0UHZLJVUKHWWYVHJOZ\P[HISL[OLYTVWOPSPJLUa`TH[PJHJ[P]P[`ILSVUNPUN[V
NS`JVZPKLO`KYVSHZLMHTPSPLZ./HUK./OH]LILLUPKLU[PÄLKHUKJOHYHJ[LYPaLK
7YLSPTPUHY`KH[H^PSSILWYLZLU[LKHPTPUN[V[OLKLZPNUVMHJOLTVLUa`TH[PJWYVJLZZ
MVYOLTPJLSS\SVZLJVU]LYZPVU
*VI\JJP7VUaHUV)(\YPSPH=9PJJPV./LUYPZZH[)*V\[PUOV74:[YHaa\SSP(7HK\SH(*VYZHYV
447PLYL[[P.7VJZMHS]P.-P\TL0*HUUPV99VZZP4HUK4VYHJJP41)PVS*OLT 4H\YLSSP3.PV]HUL(,ZWVZP[V(4VYHJJP4-P\TL09VZZP4HUK4VYHUH(,_[YLTVWOPSLZ
007VZ[LY:LZZPVU
BIB 3
Biotecnologie e Biochimica Industriale
0KLU[PÄJH[PVUVMUV]LSHSSVZ[LYPJJOHWLYVULZHZWV[LU[PHS
[OLYHWL\[PJHNLU[ZPU[OL[YLH[TLU[VM7VTWLKPZLHZL
-,99(9(4(90(*(9405(769;6*(;,905(4,304(::04030(56
(*(4769(,44((=6306=(3,90(96:(4(9./,90;(
*6)<**0765A(56),(;90*,*6364)6.069.06(5+90(.,5,96:6
7(9,5;0.0(5*(936469(**04(9*6
0UZ[P[\[LVM7YV[LPU)PVJOLTPZ[Y`*595HWSLZ"
;LSL[OVU0UZ[P[\[LVM.LUL[PJZHUK4LKPJPUL;0.,45HWSLZ"
+LWHY[TLU[VM7LKPH[YPJZ-LKLYPJV00<UP]LYZP[`5HWSLZ"
0Z[P[\[V*OPTPJHKLS9PJVUVZJPTLU[V4VSLJVSHYL*594PSHU
;OL KLÄJPLUJ` VM [OL S`ZVZVTHS LUa`TL ƠNS\JVZPKHZL .(( JH\ZLZ H NLUL[PJ
TL[HIVSPJ T`VWH[O` UHTLK 7VTWL KPZLHZL 7+ ;OPZ S`ZVZVTHS Z[VYHNL KPZLHZL
OHK UV HWWYV]LK [YLH[TLU[ ZPUJL ^OLU ,Ua`TL 9LWSHJLTLU[ ;OLYHW`
,9; L_WSVP[PUN YLJVTIPUHU[ LUa`TL YO.(( OHZ ILLU PU[YVK\JLK ;OPZ [OLYHW`
PTWYV]LZZ\Y]P]HSI\[P[ZLMÄJHJ`PZPTWHPYLKI`ZL]LYHSMHJ[VYZ0U[OLYLJLU[`LHYZ
7OHYTHJVSVNPJHS*OHWLYVUL;OLYHW`7*;LP[OLYHSVULVYPUJVTIPUH[PVU^P[O,9;
OHZILLUWYVWVZLKHZHUHS[LYUH[P]L[OLYHWL\[PJZ[YH[LN`HSZVPU7+7*;^OPJOPZ
IHZLKVU[OL\ZLVMZTHSSTVSLJ\SLZ[OH[HZZPZ[Z[OLJVYYLJ[MVSKPUNVM[OLKLMLJ[P]L
LUa`TLZ PZ HSZV WHY[PHSS` SPTP[LK ILJH\ZL [OLZL TVSLJ\SLZ ILPUN JVTWL[P[P]L
PUOPIP[VYZVM[HYNL[LUa`TLZTPNO[HMMLJ[[OLPYHJ[PVU
/LYL^LYLWVY[[OLPKLU[PÄJH[PVUVMHUV]LSJOHWLYVUL5HJL[`SJ`Z[LPUL5(*HUK
[^VYLSH[LKJVTWV\UKZ5HJL[`SZLYPUL5(:"5HJL[`SNS`JPUL5(.HISL[VZ[HIPSPaL
^PSK[`WL.((H[UL\[YHSW/[VLUOHUJL[OLYLZPK\HSHJ[P]P[`VMT\[H[LK.((HUK
[V PTWYV]L [OL LMÄJHJ` VM YO.(( \ZLK MVY ,9; ;OL Z[\K` VU YO.(( Z\WWVY[LK
HSZVI`HJVTW\[H[PVUHSIPVSVN`HUHS`ZPZVUJLSSZHUKPU]P]VPUKPJH[LK[OH[[OLZL
JOHWLYVULZKPKUV[PU[LYHJ[^P[O[OLJH[HS`[PJKVTHPUVM.((HUKJVUZLX\LU[S`^LYL
UV[JVTWL[P[P]LPUOPIP[VYZVM[OLLUa`TL5(*YLWYLZLU[Z[OLÄYZ[HSSVZ[LYPJJOHWLYVUL
PKLU[PÄLKZVMHYHUKTVYLPUNLULYHSTH`WYV]PKLHUL^[VVSMVY[OL[YLH[TLU[VM
S`ZVZVTHSKPZVYKLYZ
7VY[V*-LYYHYH4*4LSP4(JHTWVYH,(]VSPV=9VZH4*VI\JJP7VUaHUV)*VSVTIV.4VYHJJP
4(UKYPH.7HYLU[P.7OHYTHJVSVNPJHSLUOHUJLTLU[VMHSWOHNS\JVZPKHZLI`[OLHSSVZ[LYPJJOHWLYVUL
5HJL[`SJ`Z[LPUL4VS;OLYHW`PUWYLZZ
007VZ[LY:LZZPVU
BIB 4
Biotecnologie e Biochimica Industriale
.S`JVZ`U[OHZLZ!UV]LSLUNPULLYLKNS`JVZPKHZLZMVY[OL
Z`U[OLZPZVMVSPNVZHJJOHYPKLZMVYIPV[LJOUVSVNPJHSPUK\Z[Y`
*6)<**0765A(56),(;90*,:;9(AA<330(5+9,(),+050,4030(56
*69:(964(90(40*/,3(96::046:Ï(5+469(**04(9*6
0UZ[P[\[LVM7YV[LPU)PVJOLTPZ[Y`¶*VUZPNSPV5HaPVUHSLKLSSL9PJLYJOL
=PH7*HZ[LSSPUV5HWSLZ0[HS`
+PWHY[PTLU[VKP:JPLUaL*OPTPJOL<UP]LYZP[nKP5HWVSP-LKLYPJV00
*VTWSLZZV<UP]LYZP[HYPVKP4VU[L:(UNLSV]PH*PU[OPH5HWSLZ0[HS`
*HYIVO`KYH[LZZ`U[OLZPZPZ\Z\HSS`OHTWLYLKI`SHIVYPV\ZYLNPVHUKZ[LYLVJOLTPJHS
JVU[YVSZSLHKPUN[VWVVY`PLSKZHUKSPTP[PUNP[ZHWWSPJHIPSP[`PUPUK\Z[YPHSIPV[LJOUVSVNPJHS
ZJHSLZ;OPZWYVISLTHUK[OLHIZLUJLVMNLUL[PJJVKLZHZMVYWYV[LPUZHUKU\JSLPJ
HJPKZ OHZ NYLH[S` JVTWSPJH[LK [OL KL]LSVWTLU[ VM H\[VTH[PJ [LJOUPX\LZ MVY [OL
Z`U[OLZPZVMVSPNVHUKWVS`ZHJJOHYPKLZ;OLYLMVYLNS`JHUNS`JVZ`SH[LKJVTWV\UKZ
KLZWP[L [OLPY JY\JPHS YVSL PU ZL]LYHS IPVSVNPJHS WYVJLZZLZ OH]L ILLU L_WSVP[LK PU
IPV[LJOUVSVN`PUPZVSH[LKJHZLZSLH]PUNHO\NLWV[LU[PHSTHYRL[]PY[\HSS`\UL_WSVYLK
*HYIVO`KYH[LHJ[P]LLUa`TLZUHTLS`NS`JVZPKLO`KYVSHZLZHUKNS`JVZ`S[YHUZMLYHZLZ
./ HUK .; YLZWLJ[P]LS` HYL L_JLSSLU[ HS[LYUH[P]L [V JSHZZPJHS TL[OVKZ MVY
JHYIVO`KYH[LZ`U[OLZPZI\[Z\MMLYVMWVVY`PLSKZHUKZLSLJ[P]P[`./VYHYLL_WLUZP]L
HUKKPMÄJ\S[[VPZVSH[L.;.S`JVZ`U[OHZLZHJSHZZVMT\[HU[./LUNPULLYLKPU[OLPY
HJ[P]LZP[LYLWYLZLU[HUPU[LYLZ[PUNHS[LYUH[P]LWYVK\JPUNJHYIVO`KYH[LZ^P[OHSTVZ[
X\HU[P[H[P]L`PLSKZ
6\YNYV\WPZHJ[P]LPU[OPZÄLSKZPUJLTVYL[OHUHKLJHKLHUKOHZUV^HJJ\T\SH[LK
HJVSSLJ[PVUVMƠHUKơNS`JVZ`U[OHZLZ/LYL^L^PSSKLZJYPILOV^NS`JVZ`U[OHZLZ
KLTVUZ[YH[LK[OLPYWSHZ[PJP[`PUKPMMLYLU[WYVJLZZLZHUKZWLJPÄJL_HTWSLZVMơNS\JV
ƠNHSHJ[VHUKƠM\JVZ`U[OHZLZZ`U[OLZLZVMVSPNVZHJJOHYPKLZVMIPV[LJOUVSVNPJHS
PU[LYLZ[ ^OVZL WYVK\J[PVU PZ JOHSSLUNPUN ^P[O JVU]LU[PVUHS TL[OVKZ ^PSS IL
WYLZLU[LK
*VI\JJP7VUaHUV):[YHaa\SSP(9VZZP4HUK4VYHJJP4.S`JVZ`U[OHZLZPU)PVJH[HS`ZPZ(K]:`U[O
*H[HS
*VI\JJP7VUaHUV)4VYHJJP4.S`JVZ`U[OHZLZHZ[VVSZMVY[OLWYVK\J[PVUVMNS`JHUHUHSVNZVMUH[\YHS
WYVK\J[Z5H[7YVK9LW *VI\JJP7VUaHUV)*VU[L-)LKPUP,*VYZHYV447HYYPSSP4:\SaLUIHJOLY.3PWZRP(+HS7PHa-
3LWVYL39VZZP44VYHJJP4*OLT)PVS *VI\JJP7VUaHUV)AVYaL[[P*:[YHaa\SSP(*HYPSSV:)LKPUP,*VYZHYV44*VTMVY[+(2LSS`94
9VZZP44VYHJJP4.S`JVIPVSVN`
007VZ[LY:LZZPVU
BIB 5
Biotecnologie e Biochimica Industriale
;YHUZNS\[HTPUHZL9L[PJ\SH[LK/`KYVJVSSVPK)HZLK-PSTZHZ
,MMLJ[P]L*VH[PUNZVM)HRLKHUK-YPLK-VVKZ
96::04(98<,A.06=(55(+070,996796:7,96,:76:0;64(903,5(
4(9050,336369,+(5((5+769;(9(--(,3,
+LWHY[TLU[VM-VVK:JPLUJL<UP]LYZP[`VM5HWSLZ¸-LKLYPJV00¹7VY[PJP5HWVSP0[HS`
>OL` WYV[LPUWLJ[PU LKPISL ÄSTZ ^LYL WYLWHYLK \UKLY L_WLYPTLU[HS JVUKP[PVUZ MVY
VI[HPUPUN [OL TH_PT\T JVTWSL_H[PVU IL[^LLU [OL [^V O`KYVJVSSVPK JVTWVULU[Z
;OLL_WLYPTLU[Z^LYLJHYYPLKV\[PU[OLHIZLUJLVYWYLZLUJLVM[YHUZNS\[HTPUHZL
HULUa`TLHISL[VJYVZZSPURWYV[LPUZI`MVYTPUNƤƢNS\[HT`SS`ZPULPZVWLW[PKLIVUKZ
6\Y ÄUKPUNZ PUKPJH[LK [OL MVYTH[PVU H[ W/ W/J VM [YHUZNS\[HTPUHZL
JH[HS`aLK JYVZZSPURZ HTVUN ZVS\ISL PVUPJ ^OL` WYV[LPUWLJ[PU JVTWSL_LZ HUK [OH[
[OLVI[HPULKO`KYVJVSSVPKÄSTZNVVKTLJOHUPJHSHZ^LSSHZ^H[LY]HWVYHUKV_`NLU
IHYYPLYWYVWLY[PLZ-PSTHWWSPJH[PVUI`KPWJVH[PUN[LJOUPX\LHSSV^LK[VJYLH[LHSH`LY
VU[OLZ\YMHJLVMIV[OIHRLKIPZJ\P[ZHUKMYPLKKV\NOU\[ZMVVKZHISL[VPUÅ\LUJL
TVPZ[\YL[YHUZMLYHUKVPS\W[HRL0UMHJ[^LVIZLY]LKHZPNUPÄJHU[YLK\J[PVUVM^H[LY
HIZVYW[PVUI`IPZJ\P[Z^OPJOWYL]LU[LK[OLSVZZVM[OLPYJYPZWULZZRLLWPUNUVYTHS[OL
OHYKULZZWHYHTL[LYZHUKPTWYV]PUN[OLZWYPUNPULZZHUKJOL^PULZZVULZ4VYLV]LY
ZPUJL P[ PZ ^LSS RUV^U [OH[ MYPLKMVVKZ JVU[HPU OPNO HTV\U[Z VM MH[ HUK JVU[YPI\[L
PUKL[LYTPUPUNOPNOISVVKJOVSLZ[LYVSSL]LSZ^L\ZLK[OL^OL`WYV[LPUWLJ[PUÄSTZ
VI[HPULKPU[OLWYLZLUJLVM[YHUZNS\[HTPUHZL[VJVH[MYPLKMVVK^P[O[OLHPT[VYLK\JL
[OLPY^H[LYSVZZK\YPUN[OLMY`PUNWYVJLZZ6\YYLZ\S[ZZOV^LK[OH[JVH[LKKV\NOU\[Z
L_OPIP[LKKLJYLHZLKMH[\W[HRLYLK\JLKMH[JVU[LU[HUKPTWYV]LKVPSIHYYPLYPUKL_
[O\Z PUKPJH[PUN [OH[ [OL HWWSPLK JVH[PUN LMMLJ[P]LS` WYL]LU[LK ^H[LY TPNYH[PVU MYVT
[OLJVYL[V[OLJY\Z[K\YPUNMY`PUNI`ÄSSPUN[OL]VPK[OH[UVYTHSS`HSSV^Z[OLMH[[V
LU[LY PUZPKL [OL MVVK 0U JVUJS\ZPVU V\Y KH[H KLTVUZ[YH[L [OH[ [OL PU[YVK\J[PVU
VM [YHUZNS\[HTPUHZLJH[HS`aLK JYVZZSPURZ HTVUN ^OL` WYV[LPUWLJ[PU JVTWSL_LZ
VI[HPULKH[W/JJVUMLYZ[V[OLO`KYVJVSSVPKÄSTH]LY`NVVKWLYMVYTHUJLPUHJ[PUNHZ
^H[LYIHYYPLYMVYIV[OIHRLY`HUKMYPLKMVVKZ
9LMLYLUJLZ
7VY[H9L[HS*YP[9L]-VVK:JP5\[Y
7VY[H9L[HS1)PV[LJOUVS)PVTH[+60 ?L
007VZ[LY:LZZPVU
BIB 6
Biotecnologie e Biochimica Industriale
/(9=)PVYLHJ[VYMVY+J\S[\YL
VMO\THUTLZLUJO`THSZ[LTJLSSZ
7,563(AA03,;0A0(3(4),9;050,30:(),;;(=,**/0(;0509,5(;(
.(4)(9096),9;65(:;9<AA0*3(<+06(5+96),9;(70=(
+LWHY[TLU[VM)PVJOLTPZ[Y`HUK4VSLJ\SHY)PVSVN`HUK
+LWHY[TLU[VM7OHYTHJL\[PJHS:JPLUJLZ<UP]LYZP[`VM-LYYHYH-LYYHYH0;(3@
;OL :@5;/,*65 9V[HY` *LSS *\S[\YL :`Z[LT 9**: ^P[O /(9= /PNO (ZWLJ[
9V[HY` =LZZLSZ [LJOUVSVN` PZ [OL WYLTPLYL KL]PJL [OH[ LMÄJPLU[S` JYLH[LZ HU
LU]PYVUTLU[ [OH[ LUHISLZ JLSS J\S[\YLZ [V NYV^ PU[V JVTWSL_ ZVWOPZ[PJH[LK [OYLL
KPTLUZPVUHS[PZZ\LTVKLSZPU]P[YV;OL9**:PZJ\YYLU[S`ILPUN\ZLKPUH^PKLYHUNL
VMYLZLHYJOWYVQLJ[Z^OLYL[OLWYVK\J[PVUVMOPNOÄKLSP[`[PZZ\LTVKLSZVY[OLSVUN
[LYTTHPU[LUHUJLVM[PZZ\LL_WSHU[ZPZYLX\PYLK
3P[[SLPZZVMHYRUV^UHIV\[[OLJOHUNLZPUWOLUV[`WLHUKNLULL_WYLZZPVUVJJ\YYPUN
^OLUJLSSZHYL[YHUZMLYYLKMYVT+[V+J\S[\YLZ`Z[LTZ;OPZRPUKVMPUMVYTH[PVU
PZ WHY[PJ\SHYS` PTWVY[HU[ PU [PZZ\L LUNPULLYPUN HUK YLNLULYH[P]L TLKPJPUL ILJH\ZL
JLSSZ HYL MYLX\LU[S` WSHJLK PU + ZJHMMVSKZ MVY [OLYHWL\[PJ HWWSPJH[PVUZ (Z ^L HYL
PU[LYLZ[LKPU[OLKL]LSVWTLU[VMPUUV]H[P]LZ`Z[LTZMVY[OLYLWHPYVMIVULKLMLJ[Z
^L ZWLJPÄJHSS` HZRLK ^OL[OLY H YLNPTL VM K`UHTPJ Å\PK ÅV^ YLWYLZLU[LK I` [OL
9**:IPVYLHJ[VYHMMLJ[LK[OLVZ[LVNLUPJWV[LU[PHSVMO\THUTLZLUJO`THSZ[LTJLSSZ
O4:*ZLU[YHWWLKPUZ[H[PJ+Z`Z[LTYLWYLZLU[LKI`HSNPUH[LTPJYVILHKZ
>L \ZLK O4:*Z MYVT KPMMLYLU[ ZV\YJLZ PUJS\KPUN \TIPSPJHS JVYK >OHY[VU»Z QLSS`
IVUL THYYV^ HUK WLYPVKVU[HS SPNHTLU[ HM[LY Z[HUKHYK PTT\UVWOLUV[`WPUN >L
KLTVUZ[YH[LK[OH[:S\N[YHUZJYPW[PVUMHJ[VYL_WYLZZPVUPZLZZLU[PHSMVYVZ[LVNLULZPZ
*VTWHYPUN+]Z+J\S[\YLJVUKP[PVUZO4:*ZVZ[LVNLUPJWV[LU[PHSJLSSPU[LYHJ[PVU
VZ[LVISHZ[PJTHYRLYZL_WYLZZPVUHUKKLWVZP[PVUVMTPULYHSPaLKTH[YP_PTWYV]LK^OLU
HSNPUH[LLU[YHWWLK O4:*Z ^LYL J\S[\YLK PU 9**: IPVYLHJ[VY 0U HKKP[PVU ^OLU
7VS`TLYPJ 4PJLSSLZ ZLSMHZZLTISPUN JVSSVPKHS Z`Z[LTZ ^LYL JOVZLU [V KLSP]LY PU
J\S[\YLTLKP\TVZ[LVNLUPJPUK\JLYZPUZ[LHKVM[YHKP[PVUHSZVS\IPSPaH[PVU^P[O+4:6
^OPJOPZRUV^U[VILJ`[V[V_PJHUPUJYLHZLVMVZ[LVNLUPJWV[LU[PHS^HZVIZLY]LK
6\YKH[HOPNOSPNO[[OLPU[LYLZ[HIV\[[OLJVTIPUH[PVUIL[^LLUO4:*ZHUKZWLJPÄJ
ZJHMMVSKZHUKHYLPTWVY[HU[WYLYLX\PZP[LMVYHULMMLJ[P]LIVUL[PZZ\LLUNPULLYPUN
007VZ[LY:LZZPVU
)PVPUMVYTH[PJH
)5-
BNF 1
Bioinformatica
5V]LS[VVSZMVY[OLKL[LYTPUH[PVUVM
PU[YPUZPJKPZVYKLYPUWYV[LPUZ
+0+64i50*6;64±:>(3:/0(54(9;05(3),9;614
;6:(;;6:03=06*,
+LWHY[TLU[VM)PVSVN`<UP]LYZP[`VM7HKV]H7HKV]H0[HS`
+PZVYKLYLKWYV[LPUYLNPVUZHYLRL`[V[OLM\UJ[PVUVMU\TLYV\ZWYVJLZZLZ^P[OPUHU
VYNHUPZTHUK[V[OLKL[LYTPUH[PVUVMHWYV[LPU»ZIPVSVNPJHSYVSL;OLZJHU[H]HPSHISL
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Enzimiologia e Regolazione Metabolica
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ERM 12
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ERM 13
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GLI 1
Glicobiologia
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GLI 2
Glicobiologia
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GLI 3
Glicobiologia
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GLI 4
Glicobiologia
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007VZ[LY:LZZPVU
GLI 5
Glicobiologia
0:63(;065(5++0--,9,5;0(;0656-:;,4*,33:!
:,(9*/05.-695,>4(92,9:
),9.(5;,:650(7(70505(+0(*9,67(:8<(3,;699,;;(,590*(
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)9<56;,;;(4(5;0.<0+6(5+(5(:;(:0(3<0.0
3HIVYH[VY`VM:[LT*LSSZMVY;PZZ\L,UNPULLYPUN09**:7VSPJSPUPJV:HU+VUH[V
:HU+VUH[V4PSHULZL0[HS`"
+LWHY[LTLU[VM)PVTLKPJHS:JPLUJLZVM/LHS[O<UP]LYZP[`VM4PSHU0[HS`
0Z[P[\[V*SPUPJV/\THUP[HZ09**:9VaaHUV0[HS`
:[LTJLSSZHYLWV[LU[PHS[OLYHWL\[PJHNLU[Z[OH[VMMLYNYLH[WYVTPZLZPUYLNLULYH[P]L
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HZ [OL` JHU IL PZVSH[LK MYVT HSTVZ[ HU` [PZZ\L [OL` ZOV^ H NVVK ZLSMYLUL^HS
JHWHJP[`PU]P[YVHUK[OL`HSZVWVZZLZZNVVK¸WSHZ[PJP[`¹6KKS`4:*ZHYLPKLU[PÄLK
HUKKLÄULKI`HJVTIPUH[PVUVMTHYRLYZ[OH[HYLUV[KPZ[PUJ[P]LHZ[OL`HYLZOHYLK
I` V[OLY JLSSZ PUJS\KPUN ÄIYVISHZ[Z ;OLYLMVYL W\YL WVW\SH[PVUZ VM 4:*Z JHUUV[
ILPZVSH[LKHZ[OL`HYLHS^H`ZJVU[HTPUH[LKI`V[OLYHK\S[JLSSZ[OH[VM[LUKVUV[
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SPWPKZJVTWVZLKVMHJLYHTPKLIHJRIVULHUKHZ\NHYOLHKNYV\WMV\UKPU[OLV\[LY
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)44:*ZZPNUPÄJHU[S`JOHUNLZK\YPUNJLSSKPMMLYLU[PH[PVU[V^HYKVZ[LVISHZ[ZHUKP[
JV\SKIL\ZLK[VPKLU[PM`HUKJOHYHJ[LYPaL[OLKPMMLYLU[PH[PVUZ[H[\ZVM[OLJLSS
1PHUN@L[HS5H[\YL!
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GLI 6
Glicobiologia
*(5;T\[H[PVUZPU+LZI\X\VPZK`ZWSHZPHJH\ZLHKLMLJ[
PUWYV[LVNS`JHUZ`U[OLZPZ
465;03<*(/<),9*i305,+,3,65(9+0:-()06;,5509<..,96
-693056(5;65,33(*,;;(.0<:,77,),9;6304(9;(
-9(+054i3(50,3,4,99,94(9;05,3,.6--*(905,
*6940,9+(09,=(3i90,(5+96::0(5;6506
<UP]LYZP[`VM7H]PH+LW[4VSLJ\SHY4LKPJPUL7H]PH0[HS`"
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/WP[HS5LJRLY,UMHU[Z4HSHKLZ7HYPZ-YHUJL
+LZI\X\VPZK`ZWSHZPH++PZHUH\[VZVTHSYLJLZZP]LJOVUKYVK`ZWSHZPHJOHYHJ[LYPaLK
I` HU[LUH[HS HUK WVZ[UH[HS NYV^[O YL[HYKH[PVU T\S[PWSL KPZSVJH[PVUZ HUK HK]HUJLK
JHYWHSVZZPÄJH[PVU;^VMVYTZVM++OH]LILLUKLZJYPILKVU[OLIHZPZVM[OLWYLZLUJL
[`WLVYHIZLUJL[`WLVMJOHYHJ[LYPZ[PJOHUKKLMVYTP[PLZ:[\K`PUN++[`WL
MHTPSPLZT\[H[PVUZPU[OL*HSJP\T(J[P]H[LK5\JSLV[PKHZLNLUL*(5;OH]LILLU
PKLU[PÄLK/\ILY*L[HS (T14LK.LUL[HUK5PaVU4L[HS
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UVYTHS\UKLYIHZHSJVUKP[PVUZI\[ZPNUPÄJHU[YLK\JLK.(.Z`U[OLZPZ^HZVIZLY]LK
PUWYLZLUJLVMơ+_`SVZPKLHJVTWV\UK^OPJOLUOHUJLZZ`U[OLZPZHUKZLJYL[PVUVM
JOVUKYVP[PUHUKKLYTH[HUZ\SMH[LJOHPUZHJ[PUNHZHJOHPUPUP[PH[VY-\Y[OLYTVYLNLS
ÄS[YH[PVUJOYVTH[VNYHWO`VU:\WLYVZLVM.(.ZYLSLHZLKMYVTUL^S`Z`U[OLZPaLK
WYV[LVNS`JHUZ HM[LY ơLSPTPUH[PVU KLTVUZ[YH[LK [OH[ .(. JOHPUZ ^LYL ZOVY[LY
JVTWHYLK [V [OL JVU[YVSZ /`HS\YVUPJ HJPK Z`U[OLZPZ ^OPJO VJJ\YZ PU [OL WSHZTH
TLTIYHUL H KPMMLYLU[ JVTWHY[TLU[ MYVT WYV[LVNS`JHUZ ^HZ ^P[OPU UVYTHS ]HS\LZ
Z\WWVY[PUN[OLPU]VS]LTLU[VM*(5;PU[OL,9.VSNPJVTWHY[TLU[
;OLZL KH[H Z\NNLZ[ [OH[ *(5; WSH`Z H YVSL PU WYV[LVNS`JHU HUK .(. TL[HIVSPZT HUK Z\WWVY[ P[Z
PU]VS]LTLU[PU[OLLUKVJOVUKYHSVZZPÄJH[PVUWYVJLZZ
>VYRZ\WWVY[LKI`;LSL[OVU0[HS`NYHU[UV..7 007VZ[LY:LZZPVU
GLI 7
Glicobiologia
:6+4040*4U;47`79,+<*,:05-3(44(;065
05+<*,+)@+,.9(+,+/@(3<965(505*/65+96*@;,:
:;04<3(;,+>0;/-L0073<:(:*69)(;,
+»(:*63((5.,3((=,56:6(5.,3(:*<9<*/040*/,3,
5(:;(:0.0(5*(93640*(30(5;65067<AA636+64,50*6
*(3(;9650(3),9;6*(476:(3=(;69,*(476.0<:,77,4(<90A06
+LWHY[TLU[VM)PVTLKPJHS:JPLUJLZHUK4VYWOVSVNPJHSHUK-\UJ[PVUHS0THNLZ
:JOVVSVM4LKPJPUL<UP]LYZP[`VM4LZZPUH
7VSPJSPUPJV<UP]LYZP[HYPV ¶4LZZPUH0[HS`
+\YPUN WH[OVSVNPJHS JVUKP[PVUZ V_PKH[P]L I\YZ[ NLULYH[LZ O`HS\YVUHU /(
MYHNTLU[H[PVU ^P[O JVUZLX\LU[ PUJYLHZL PU ZTHSS /( VSPNVZHJJOHYPKL SL]LSZ ;OLZL
MYHNTLU[Z HYL HISL [V Z[PT\SH[L PUÅHTTH[VY` YLZWVUZL PU KPMMLYLU[ JLSS [`WLZ I`
HJ[P]H[PUN[OL*+HUKIV[O[OL[VSSSPRLYLJLW[VYZ;39HUK;39*+HUK
;39ZZ[PT\SH[PVUPU[\YUHJ[P]H[LZ[OL5-R)[OH[PUK\JLZ[OLWYVK\J[PVUVMZL]LYHS
WYVPUÅHTTH[VY` TLKPH[VYZ [OH[ HTWSPM` HUK WLYWL[\H[L PUÅHTTH[PVU >L HPTLK
[V Z[\K` [OL LMMLJ[ VM [OL HU[PV_PKHU[ :6+ TPTPJ Z`U[OL[PJ THUNHULZL WVYWO`YPU
4U000 [L[YHRPZ5TL[O`SW`YPKPUP\T`SWVYWO`YPU 4U;47`7 PU
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O`KYV_`SYHKPJHSHUKWLYV_`UP[YP[LWYVK\J[PVUHUKPUJYLHZLPUIV[OSPWPKWLYV_PKH[PVU
HUK/(KLNYHKH[PVU/(MYHNTLU[ZWYVK\JLKOPNOT95(L_WYLZZPVUHUK[OLYLSH[LK
WYV[LPU WYVK\J[PVU VM *+ ;39 HUK ;39 5-R) HJ[P]H[PVU HUK H ZPNUPÄJHU[
\WYLN\SH[PVUVM[OLPUÅHTTH[VY`J`[VRPULZ[\TVYULJYVZPZMHJ[VYHSWOH;5-HSWOH
PU[LYSL\RPUIL[H 03IL[H HUK V[OLY WYVPUÅHTTH[VY` TLKPH[VYZ Z\JO HZ TH[YP_
TL[HSSVWYV[LHZL447HUKPUK\JPISLUP[YPJV_PKLZ`U[OHZLP56:;YLH[TLU[VM
JLSSZ^P[O4U;47`7^HZHISL[VH[[LU\H[L[OLV_PKH[P]LI\YZ[/(KLNYHKH[PVU
HUK5-R)HJ[P]H[PVUTHYRLKS`KLJYLHZLK*+HUK;39ZT95(L_WYLZZPVUHUK
[OL YLSH[LK WYV[LPU Z`U[OLZPZ HZ ^LSS HZ [OL \WYLN\SH[LK PUÅHTTH[VY` TLKPH[VYZ
WYVK\J[PVU (KKP[PVU [V JLSSZ VM /( ISVJRPUN WLW[PKL 7,7 /()7 ZPNUPÄJHU[S`
YLK\JLKHSS[OLPUÅHTTH[VY`WHYHTL[LYZ\WYLN\SH[LKI`-L00WS\ZHZJVYIH[LHUK
Z[YLUN[OLUZ4U;47`7HJ[P]P[`;OLZLKH[HZ\NNLZ[[OH[/(KLNYHKH[PVUWSH`Z
HRL`YVSLPU[OLLHYSPLZ[PUÅHTTH[VY`YLZWVUZLVMJHY[PSHNLHUKHU[PV_PKHU[ZJV\SKIL
H\ZLM\S[VVS[VWYL]LU[[OLHTWSPÄJH[PVUVM[OLTLJOHUPZT
007VZ[LY:LZZPVU
GLI 8
Glicobiologia
*64)05,+;9,(;4,5;>0;//(05/0)0;697,7(5+
(:,3,*;0=,(+6(9,*,7;69(.650:;9,+<*,:
05-3(44(;06505,?7,904,5;(3(9;/90;0:
*(3(;9650(3),9;6(=,56:6(5.,3(+»(:*63((5.,3(
5(:;(:0.0(5*(93640*(30(5;65067<AA636+64,50*6
79,:;07056=,9(:*<9<*/040*/,3,*(476:(3=(;69,*(476.0<:,77,4
+LWHY[TLU[VM)PVTLKPJHS:JPLUJLZHUK4VYWOVSVNPJHSHUK-\UJ[PVUHS0THNLZ:JOVVS
VM4LKPJPUL<UP]LYZP[`VM4LZZPUH7VSPJSPUPJV<UP]LYZP[HYPV ¶4LZZPUH0[HS`
(IZ[YHJ[
9OL\TH[VPK HY[OYP[PZ 9( PZ H JOYVUPJ HUK Z`Z[LTPJ KPZVYKLY JOHYHJ[LYPZLK I` [OL
WYVNYLZZP]L KLZ[Y\J[PVU VM HY[PJ\SHY JHY[PSHNL HUK IVUL 7YL]PV\Z PU]LZ[PNH[PVUZ
YLWVY[LK [OH[ [OL KLNYHKH[PVU VM UH[P]L OPNOS` WVS`TLYPaLK O`HS\YVUHU /( PU[V
ZTHSSVSPNVZHJJOHYPKLZTH`WSH`HYVSLPU[OLKL]LSVWTLU[HUKWYVNYLZZPVUVM9(
0UÅHTTH[VY`YLZWVUZLZVJJ\YI`TVK\SH[PUN[OL[VSSSPRLYLJLW[VY;39HUK[OL
;39 HUK *+ H UH[\YHS /( YLJLW[VY 0[ ^HZ HSZV ZOV^U [OH[ [OL HKLUVZPUL (
YLJLW[VY((9WSH`ZYVSLPUHY[OYP[PZL_LY[PUNHU[PPUÅHTTH[VY`HJ[P]P[`;OLZ[PT\SH[PVU
VM ;39 ;39 HUK *+ JVU]LYNL [V HJ[P]H[L 5-R) ^OPJO PU [\YU Z[PT\SH[LZ
[OL WYVK\J[PVU VM WYVPUÅHTTH[VY` J`[VRPULZ HUK V[OLY KL[YPTLU[HS TLKPH[VYZ 0U
JVU[YHZ[[OLZ[PT\SH[PVUVM[OL((9PUOPIP[Z5-R)HJ[P]H[PVU;OLHPTVM[OPZZ[\K`
^HZ[VPU]LZ[PNH[L[OLLMMLJ[VMHJVTIPULK[YLH[TLU[\ZPUN[OL/(PUOPIP[VY7LW
HUKHZLSLJ[P]L((9HNVUPZ[*=PUJVSSHNLUPUK\JLKHY[OYP[PZ*0(PUTPJL
(Y[OYP[PZ^HZPUK\JLK]PHPU[YHKLYTHSPUQLJ[PVUVMHULT\SZPVUJVU[HPUPUNIV]PUL[`WL00
JVSSHNLUPUJVTWSL[L-YL\UK»ZHKQ\]HU[4PJL^LYL[YLH[LK^P[O7LWWS\Z*=
PU[YHWLYP[VULHSS`KHPS`MVYKH`Z
*0( PUJYLHZLK ;39 ;39 *+ HUK ((9 T95( L_WYLZZPVU HUK [OL YLSH[LK
WYV[LPUZPU[OLQVPU[JHY[PSHNLVMHY[OYP[PJTPJL:PNUPÄJHU[S`PUJYLHZLKSL]LSZVMIV[O
T95( HUK YLSH[LK WYV[LPU ^LYL VIZLY]LK MVY [\TVY ULJYVZPZ MHJ[VY HSWOH ;5-Ơ
PU[LYSL\RPUIL[H03ơPU[LYSL\RPU03TH[YP_TL[HSSVWYV[LHZL447
HUKPUK\JPISLUP[YPJV_PKLZ`U[OHZLP56:7LW[VNL[OLY^P[O*=[YLH[TLU[
ZPNUPÄJHU[S` YLK\JLK *0( KHTHNL HUK KLJYLHZLK HSS [OL \WYLN\SH[LK IPVJOLTPJHS
WHYHTL[LYZ 9LK\J[PVU PU IPVJOLTPJHS WHYHTL[LYZ ^HZ Z\WWVY[LK I` OPZ[VSVNPJHS
HUHS`ZPZHUKZ`UV]PHSÅ\PK/(SL]LSZ
;OLZLYLZ\S[ZZ\NNLZ[[OH[I`HJ[PUN[OYV\NOHWWVZP[L[VVSZVU[OL/(KLNYHKH[PVU
WH[O^H`ZHUKH[[OLZHTL[PTL^P[OV[OLYVWWVY[\ULZWLJPÄJ[VVSZVUHKLUVZPUL
WH[O^H`ZP[^V\SKILWVZZPISL[VKL]LSVWHU[PPUÅHTTH[VY`Z[YH[LNPLZ[OH[HTLSPVYH[L
HY[OYP[PZZ`TW[VTZTVYLLMMLJ[P]LS`
007VZ[LY:LZZPVU
GLI 9
Glicobiologia
(*;0=(;0656-(+,56:05,(9,*,7;699,+<*,:
05-3(44(;69@9,:765:,=0(,7(*(5+7(9;3@=0(
72(0546<:,:@56=0(3-0)96)3(:;::;04<3(;,+
>0;//@(3<965(5630.6:(**/(90+,:
.0(5*(9365(:;(:0(5.,3((=,56:6(5.,3(+»(:*63(
40*/,3,:*<9<*/0(5;650640*(30+64,50*67<AA636
(3),9;6*(3(;9650:(3=(;69,*(476.0<:,77,4(<90A06*(476
+LWHY[TLU[VM)PVTLKPJHS:JPLUJLZHUK4VYWOVSVNPJHSHUK-\UJ[PVUHS0THNLZ
:JOVVSVM4LKPJPUL<UP]LYZP[`VM4LZZPUH7VSPJSPUPJV<UP]LYZP[HYPV
¶4LZZPUH0[HS`
(IZ[YHJ[
:THSSO`HS\YVUHU/(MYHNTLU[ZPUK\JLPUÅHTTH[VY`YLZWVUZLPUKPMMLYLU[JLSS[`WLZ
I` WYPTHYPS` Z[PT\SH[PUN [OL [VSSSPRL YLJLW[VY ;39 ;OPZ PU[LYHJ[PVU HJ[P]H[LZ H
ZLYPLZVMPU[LYTLKPH[LZ[OH[H[[OLLUKHJ[P]H[L[OLU\JSLHYMHJ[VYRHWWH)5-R)5-
R)PU[\YUPUK\JLZ[OL[YHUZJYPW[PVUVMPUÅHTTH[PVUJ`[VRPULZZ\JOHZ[\TVYULJYVZPZ
MHJ[VYHSWOH ;5-HSWOH HUK PU[LYSL\RPUIL[H 03IL[H [OH[ WYPTL PUÅHTTH[PVU
6U [OL JVU[YHY` [OL W\YPUL U\JSLVZPKL HKLUVZPUL (+6 PU[LYHJ[PUN ^P[O [OL OPNO
HMÄUP[`((YLJLW[VYHJ[P]H[LZHWH[O^H`SLHKPUN[VPUJYLHZLKJ(47SL]LSZ;OLJ(47
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L_JOHUNLWYV[LPUZHJ[P]H[LKI`J(47,7(*TH`HSZVHJ[VU5-R)PUOPIP[PVU^L
HPTLK[VPU]LZ[PNH[L[OLPU]VS]LTLU[VM,7(*PUTLY/(VSPNVZHJJOHYPKLPUK\JLK
PUÅHTTH[VY`YLZWVUZLPUTV\ZLZ`UV]PHSÄIYVISHZ[Z5:-
;YLH[TLU[VM5:-^P[OTLY/(PUJYLHZLK;39;5-HSWOHHUK03IL[HT95(
L_WYLZZPVUHUK[OLYLSH[LKWYV[LPUWYVK\J[PVUHZ^LSSHZ5-R)HJ[P]H[PVU(KKP[PVU
[V5:-WYL]PV\ZS`Z[PT\SH[LK^P[OTLY/(VSPNVZHJJOHYPKLZVM(+6ZPNUPÄJHU[S`
YLK\JLK 5-R) HJ[P]H[PVU ;5-HSWOH HUK 03IL[H L_WYLZZPVU ;OL WYL[YLH[TLU[
VM5:-^P[OJ(47HUKVY72(HUKVY,7(*ZWLJPÄJPUOPIP[VYZZPNUPÄJHU[S`PUOPIP[LK
[OLHU[PPUÅHTTH[VY`LMMLJ[L_LY[LKI`(+60UWHY[PJ\SHY[OL,7(*PUOPIP[VYYLK\JLK
(+6LMMLJ[[VHTHQVYL_[LU[[OHU72(PUOPIP[VY;OLZLYLZ\S[ZTLHU[OH[IV[O72(
HUK,7(*WH[O^H`ZHYLPU]VS]LKPU(+6PUK\JLK5-R)PUOPIP[PVUHS[OV\NO,7(*
ZLLTZ[VILTVYLPU]VS]LK[OHU72(
007VZ[LY:LZZPVU
GLI 10
Glicobiologia
(.,ơNS`JVWYV[LPU0HJ[P]H[LZKLUKYP[PJJLSSZHUKPUK\JLZ
HKHW[P]LPTT\UP[`PUWH[PLU[Z^P[OHU[PWOVZWOVSPWPK
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GLI 11
Glicobiologia
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Glicobiologia
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GLI 13
Glicobiologia
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GLI 14
Glicobiologia
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V_PKH[P]L Z[H[\Z ([ JLSS\SHY SL]LS [OL PTWYV]LTLU[ VM *V8 V_PKH[P]L Z[H[\Z ^HZ
HZZVJPH[LK ^P[O LUOHUJLK TP[VJOVUKYPHS M\UJ[PVUHSP[` HUK KLJYLHZLK Z\ZJLW[PIPSP[`
[V V_PKH[P]L PUZ\S[ TLHZ\YLK PU [LYTZ VM TP[VJOVUKYPHS TLTIYHUL WV[LU[PHS \ZPUN
[OL ULYUZ[PHU WYVIL 1* /V^L]LY TP[VJOVUKYPHS HJ[P]P[` ^OLU Z[PT\SH[LK I`
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V\YL_WLYPTLU[HSZL[\W^HZHSZVHZZVJPH[LK^P[OPUJYLHZLKPU[YHJLSS\SHY96:SL]LSZ
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YLTHYRHIS`V_PKPZPUNJVUKP[PVUZ-\Y[OLYZ[\KPLZHYLYLX\PYLK[VIL[[LYJOHYHJ[LYPaL
PU]P]VK\YPUNL_OH\Z[P]LL_LYJPZL[OLWYVLULYNL[PJHUKW\aaSPUNHU[PWYVV_PKH[P]L
LMMLJ[VM*V8HUKSPWVPJHJPK
007VZ[LY:LZZPVU
MEB 2
Membrane e Bioenergetica
4LSH[VUPUTVK\SH[LZTP[VJOVUKYPHSM\UJ[PVUPU/H*H;JLSSZ
]PHUP[YPJV_PKLWYVK\J[PVU
(9,:,4(9A0(4(.50-0*64(90(*/0(9(4(:;9650*63(+(50,3(
-69;,,3,5(.0<--9Ï(3,::(5+96.90336*(;,905(
(3;0,90-()06)3(5*2;/64(:11(5+:(9;07(636
+LWHY[TLU[VM)PVJOLTPJHS:JPLUJLZ:HWPLUaH<UP]LYZP[`VM9VTL
*590UZ[P[\[LVM4VSLJ\SHY)PVSVN`HUK7H[OVSVN`9VTL
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4LSH[VUPUPZHOVYTVULWYVK\JLKI`[OLWPULHSNSHUKMVSSV^PUNJPYJHKPHUYO`[OTZ
HS[OV\NOTHU`V[OLYJLSS[`WLZHYLHISL[VWYVK\JLP[;OPZTVSLJ\SLPZPU]VS]LKPU
THU` JLSS M\UJ[PVUZ PUJS\KPUN YLN\SH[PVU VM ZWLJPÄJ LUa`TLZ PU]VS]LK PU JLSS YLKV_
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YLZWVUZLZ"[OLZLPUJS\KLZTVV[OT\ZJSLYLSH_H[PVUUL\YV[YHUZTPZZPVUPUOPIP[PVUVM
WSH[LSL[ HNNYLNH[PVU JLSS TPNYH[PVU HUK TP[VJOVUKYPHS YLZWPYH[PVU 0U THTTHSZ 56
PZ Z`U[OLZPaLK I` [OYLL KPMMLYLU[ NLULLUJVKLK 56 Z`U[OHZL 56: [OL UL\YVUHS
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VMZPNUHSSPUNJVTWV\UKZ6\Y^VYR^HZHPTLKH[Z[\K`PUN[OLLMMLJ[ZVMTLSH[VUPU
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SL]LS[OLIPVLULYNL[PJWHYHTL[LYZZ\JOHZSHJ[H[LHJJ\T\SH[PVU(;7WYVK\J[PVUHUK
TP[VJOVUKYPHS TLTIYHUL WV[LU[PHS MVSSV^LK Z`UJOYVUV\Z JOHUNLZ JVTWH[PISL ^P[O
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007VZ[LY:LZZPVU
MEB 3
Membrane e Bioenergetica
9HWHT`JPUYLK\JLZV_PKH[P]LZ[YLZZ
PUMYH[H_PUKLÄJPLU[`LHZ[JLSSZ
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MEB 4
Membrane e Bioenergetica
4P[VJOVUKYPHSZVS\ISLHKLU`S`SJ`JSHZLZ(*JVVWLYH[LZ
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J(47JVVWLYH[LPU[OLYLN\SH[PVUVMJVTWSL_0HJ[P]P[`I`TVK\SH[PUN[OLPTWVY[HUK
K`UHTPJHZZLTIS`L_JOHUNLVMWLYPWOLYHSZ\I\UP[Z
007VZ[LY:LZZPVU
MEB 5
Membrane e Bioenergetica
6]LYL_WYLZZPVUPU,JVSPHUKYLJVUZ[P[\[PVUPUSPWVZVTLZ
VM[OL6*;5:3*([YHUZWVY[LY
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L_WYLZZLK PU OL[LYVSVNV\Z VYNHUPZTZ ;OL V]LYL_WYLZZPVU PU , JVSP VM O\THU
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H2TMVYJHYUP[PULVM\4ZPTPSHY[V[OH[MV\UKPUJLSSZ6UHWO`ZPVSVNPJHSWVPU[
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BD;HTHP0L[HS1)PVS*OLT
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MEB 6
Membrane e Bioenergetica
(\[VUVTV\Z\S[YHJPYJHKPHUYO`[OTZVMTP[VJOVUKYPHS
YLZWPYH[VY`HJ[P]P[`
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KLSSH:VMMLYLUaH6WLYHKP7HKYL7PVKH7PL[YLSJPUH:HU.PV]HUUP9V[VUKV-.0[HS`
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MEB 7
Membrane e Bioenergetica
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MEB 8
Membrane e Bioenergetica
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MEB 9
Membrane e Bioenergetica
;OL5)+KVTHPUZVM[OL497()**
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MEB 10
Membrane e Bioenergetica
/@76?0((5+(4@360+ơ7,7;0+,40./;
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MEB 11
Membrane e Bioenergetica
=+(*[VWVSVN`PU[OLV\[LYTP[VJOVUKYPHSTLTIYHUL
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MEB 12
Membrane e Bioenergetica
:WLJPÄJȕZ[YHUKZJVU[HJ[PUN[OL5[LYTPUHSKVTHPUVM
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MEB 13
Membrane e Bioenergetica
(YL=+(*SL]LSZPU+YVZVWOPSHTLSHUVNHZ[LYTP[VJOVUKYPH
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MEB 14
Membrane e Bioenergetica
:OVY[[LYTLMMLJ[VMKPPVKV3[O`YVUPULVUMH[[`HJPK
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MEB 15
Membrane e Bioenergetica
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MEB 16
Membrane e Bioenergetica
*VTWHYH[P]LHUHS`ZPZVMTP[VJOVUKYPHSK`ZM\UJ[PVUZPU
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MEB 17
Membrane e Bioenergetica
9L[PUVPKZHUK72(ZPNUHS[YHUZK\J[PVUPU[LYHJ[PVUYLN\SH[L
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MEB 18
Membrane e Bioenergetica
7YV[LVTPJL_WYLZZPVUWYVÄSLPUTP[VJOVUKYPHRLYH[PUVJ`[LZ
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3077630:96:(7(7(-9(5*,:*650*(:;96(55(90;(:(9+(96
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VMTP[VJOVUKYPHS6?7/6:JVTWSL_LZ
007VZ[LY:LZZPVU
MEB 19
Membrane e Bioenergetica
,?;9(40;6*/65+90(36?0+(;0=,7/6:7/69@3(;065!
(5,>46+,3-69*6<7305.),;>,,59,:709(;69@
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9(=,9(:03=0(7(5-6300:(),33()(9;63<**04(9;05(*(3A0(+(50,3(
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9LJLU[WHWLYZYLWVY[HTVYLWLY]HZP]LL_WYLZZPVUVM(;7Z`U[OHZL[OHUL_WLJ[LK
B9L]PL^LKPUD0ZVSH[LKT`LSPU]LZPJSLZBDHUKYL[PUHSYVKV\[LYZLNTLU[KPZRZBD
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[OL6?7/6:WYV[LPUZHUKÅ\VYLZJLU[WYVILZHZ4P[V;YHJRLYHUK1*BD
6_`NLUJVUZ\TW[PVUI`7SHZTHISHZ[Z7)JLSSZLUYPJOLKPULUKVWSHZTPJYL[PJ\S\T
^HZ ZPNUPÄJHU[S` OPNOLY ^OLU Z[PT\SH[LK ^P[O 5(+/ [OHU PU X\PLZJLU[ ISHZ[Z )
9H]LYHL[HS<UW\ISPZOLKKH[H4VYLV]LY7)KPZWSH`LKHZPNUPÄJHU[S`KPMMLYLU[>)
ZPNUHSYH[PVVM(;7Z`U[OHZL]Z(5;^P[OYLZWLJ[[V)
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VM [OL J\YYLU[ TVKLS MVY WYV[VU / [YHUZMLY MYVT YLZWPYH[VY` JVTWSL_LZ [V -V PZ
WYVWVZLK0USPUL^P[OL]PKLUJLZ\NNLZ[PUN[OH[/TV]L^P[OPU[OLIPSH`LYYH[OLY[OHU
L_JOHUNPUN ^P[O [OL HX\LV\Z WOHZL BD ^L WVZ[\SH[L H WOHZL ZLWHYH[PVU HTVUN
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TLTIYHUL ^OPSL (;7 Z`U[OLZPZ [HRLZ WSHJL PU [OL HX\LV\Z WOHZL ;OPZ TVKLS PZ
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BD >*3H\139\IPUZ[LPU5H[\YL
007VZ[LY:LZZPVU
5\JSLV[PKP
(JPKP5\JSLPJP
.LUVTP
5(.
NAG 1
Nucleotidi, Acidi Nucleici, Genomi
9LJ\YYLU[JVW`U\TILYHIUVYTHSP[PLZHUKJVW`UL\[YHS
SVZZVMOL[LYVa`NVZP[`PUJVSVYLJ[HSJHUJLYYL]LHSLKI`
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3HIVYH[VY`VU*VTWSL_:`Z[LTZ:J\VSH:\WLYPVYLKP*H[HUPH<UP]LYZP[`VM*H[HUPH0[HS`"
-VUKHaPVUL4LKP[LYYHULH¸.)4VYNHNUP¹*H[HUPH0[HS`"
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;^V MVYTZ VM NLUL[PJ PUZ[HIPSP[` OH]L ILLU VIZLY]LK PU JVSVYLJ[HS JHUJLY *9*!
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H ZLYPLZ VM *9* ZHTWSLZ HUK [OLPY YLSH[LK UVYTHS T\JVZHL I` OPNO YLZVS\[PVU
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VM )*5(Z U\TILY JVUÄYTZ [OH[ PU [OL THQVYP[` VM JHZLZ JOYVTVZVTHS PUZ[HIPSP[`
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HZTHSS[\TVYZ\INYV\W^P[OKV\ISLNLUVTPJPUZ[HIPSP[`4:0*055VJVYYLSH[PVU
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ZVTH[PJ JVW` UL\[YHS36/ YLNPVUZ Z\NNLZ[PUN [OH[ KPMMLYLU[ TLJOHUPZTZ \UKLYSPL
Z\JO JOYVTVZVTHS HIUVYTHSP[PLZ (UHS`ZPZ VM YLJ\YYLU[ *5(Z PKLU[PÄLK ZVTH[PJ
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7VZ[LY7YLZLU[H[PVU
007VZ[LY:LZZPVU
NAG 2
Nucleotidi, Acidi Nucleici, Genomi
(ZP95(HWWYVHJOZWLJPÄJHSS`[HYNL[Z[OLT\[HU[HSSLSLHUK
YLK\JLZT\[HU[JVSSHNLUPU)Y[STPJLHT\YPULTVKLSMVY
VZ[LVNLULZPZPTWLYMLJ[H
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/,@4(55+64058<,),:0696),9;(96::0(5;6506
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;OtYHWPLKLZ;\TL\YZ6ZZL\ZLZ7YPTP[P]LZ-HJ\S[tKL4tKLJPUL5HU[LZ-YHUJL
65090:5HU[LZ([SHU[PJ*VSSLNLVM=L[LYPUHY`4LKPJPUL-VVK:JPLUJLHUK,UNPULLYPUN-
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THPUS`I`T\[H[PVUZPU[`WL0JVSSHNLUNLULZ:L]LYLWOLUV[`WLPUWH[PLU[ZJV\SKIL
JVU]LY[LK[VHTPSKLYV\[JVTLI`HZWLJPÄJZ\WWYLZZPVUVM[OLT\[H[LKHSSLSL(PTVM
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60[YLH[TLU[;OL60)Y[STV\ZLJHYY`PUNH[`WPJHSNS`JPULZ\IZ[P[\[PVU.S` *`ZPU
HJVSHHSSLSL^HZ\ZLK
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TPZTH[JOLZ /,2 JLSSZ HUK WYPTHY` )Y[S ÄIYVISHZ[Z ^LYL \ZLK MVY [YHUZMLJ[PVU
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T\[HU[HUK>;HSSLSLZPU)Y[SÄIYVISHZ[Z^LYLL]HS\H[LKI`TLHUZVMX7*9*VSSHNLU
MYVT)Y[SJLSSZ^HZHUHS`aLKI`:+:7(.,
0U[YHUZMLJ[LK/,2JLSSZVUS`[^VZP95(ZKPKUV[JOHUNLS\JPMLYHZLHJ[P]P[`^OLU[OL
S\JPMLYHZLNLULJVU[HPULK[OL^PSK[`WLJVSHL_VUZLX\LUJL^P[OPUP[Z»<;9
^OLYLHZ[OL`LUHISLKHMVSKKLJYLHZLVM[OLS\JPMLYHZLHJ[P]P[`^OLUS\JPMLYHZLNLUL
JVU[HPULK[OLT\[H[LKJVSHL_VUZLX\LUJL;OLZL[^VZP95(Z[YHUZMLJ[LKPU
)Y[SÄIYVISHZ[ZLUHISLKHYLK\J[PVUVM[OLT\[H[LKJVSH[YHUZJYPW[]LYZ\ZH
YLK\J[PVUVM[OL^PSK[`WLVULHZTLHZ\YLKI`HSSLSLZWLJPÄJX7*9(KKP[PVUHSS`
ZO95(Z JVYYLZWVUKPUN [V [OLZL [^V LMMLJ[P]L HUK ZLSLJ[P]L ZP95( ^LYL JSVULK PU
H SLU[P]PYHS ]LJ[VY HUK VUL VM [OLT LUHISLK H Z[HISL ZWLJPÄJ Z\WWYLZZPVU VM
[OL T\[H[LK JVSH ^P[OV\[ LMMLJ[PUN [OL >; HSSLSL PU )Y[S ÄIYVISHZ[Z HUK JH\ZLK
YLK\J[PVU VM [OL T\[HU[ JVSSHNLU 6\Y KH[H Z\WWVY[ [OL \ZL VM ZO95(Z HZ H
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007VZ[LY:LZZPVU
NAG 3
Nucleotidi, Acidi Nucleici, Genomi
<UJV]LYPUN[OLWO`ZPVSVNPJHSYVSLVMO\THUJ`[PKPUL
KLHTPUHZL[OYV\NO[OLJOHYHJ[LYPaH[PVUVMP[ZM\UJ[PVUHS
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7`YPTPKPULWYVKY\NZT\Z[ILJVU]LY[LKPU[VHJ[P]LJVTWV\UKZ[OYV\NOHZLYPLZVM
YLHJ[PVUZPU]VS]PUN[OLLUa`TLZVM[OLW`YPTPKPULZHS]HNLWH[O^H`Z0U[LYPUKP]PK\HS
]HYPH[PVUPU[OLHJ[P]P[`VM[OLZLTL[HIVSPaPUNLUa`TLZJHUHMMLJ[[OLL_[LU[VMWYVKY\N
HJ[P]H[PVUHUKHZHYLZ\S[HJ[VU[OLLMÄJHJ`VMJOLTV[OLYHW`[YLH[TLU[;^V:PUNSL
5\JSLV[PKL7VS`TVYWOPZTZ:57ZPU[OLJ`[PKPULKLHTPUHZLNLUL^LYLKL[LJ[LK!VUL
[OL (%*:57YLZ\S[PUNPUH28JOHUNLHUK[OL.%(:57WYVK\JPUNHU
HSHUPUL[V[OYLVUPULZ\IZ[P[\[PVU(;^P[OPU[OLJVUZLY]LKJH[HS`[PJKVTHPU0U
[OPZ^VYR^LMVJ\ZLKVU[OL (%*:57HUKP[ZWYVK\J[ZUHTLK8HUK2*+(
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WVZP[PVULZ[HISPZOLZHUPVUPJPU[LYHJ[PVU^P[OHNS\[HTTPJHJPKYLZPK\LH[WVZP[PVU
[OH[PZYLZWVUZPISLMVYHNYLH[LYZ[Y\J[\YHSZ[HIPSP[`VM[OL]HYPHU[JHYY`PUN[OL2^P[O
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WVZP[PVU PUZ[LHK VM S`ZPUL PZ YLZWVUZPISL MVY [OL THQVY JH[HS`[PJ LMÄJPLUJ` [V^HYKZ
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HZZLZZLK[OLRPUL[PJWYVWLY[PLZVM[OLYLJVTIPUHU[OL[LYVa`NV\ZO\THU*+(82
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WSHZTPKL_WYLZZPUNZPT\S[HULV\ZS`IV[O*+(NLULZ;OL82*+(VI[HPULK^HZHISL
[VMVYTH[L[YHTLYNP]PUNYPZL[VÄ]LPZVMVYTZVMJ`[PKPULKLHTPUHZLMVYTLKI`[OL
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2…OU2L[HS )PVJOLT)PVWO`Z9LZ*VTT\U ¶
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NAG 4
Nucleotidi, Acidi Nucleici, Genomi
9LN\SH[PVUVM;OL7@3PRL9LJLW[VY.79I`
(NVUPZ[0UK\JLK+LZLUZP[PaH[PVUPU[YHUZMLJ[LK
5JLSSZ
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Z\NNLZ[PUN[OH[U\JSLV[PKLZOH]LHOPLYHYJO`PUWYVK\JPUNKLZLUZP[PaPUNZPNUHSZ
0U [OPZ ^VYR ^L PU]LZ[PNH[LK [OL YVSL VM .92 HUK IL[HHYYLZ[PUZ PU [OL YLN\SH[PVU
VM .79 YLZWVUZLZ PU 5 JLSSZ [YHUZMLJ[LK ^P[O /([HNNLK .79 )V[O
W\YPULYNPJHUKJ`Z3;HNVUPZ[ZPUK\JLH[PTLKLWLUKLU[YLJLW[VYWOVZOVY`SH[PVUVU
;OYLVUPUL HUK :LYPUL YLZPK\LZ )V[O .79 HNVUPZ[Z PUK\JL .92 [YHUZSVJH[PVU
MYVTJ`[VWSHZT[VWSHZTHTLTIYHULHUK[OLWO`ZPJHSHZZVJPH[PVUIL[^LLU.92
HUK.79I\[[OLZLLMMLJ[Z^LYLTVYLTHYRLK^P[O[OLJ`Z3;SPNHUK^P[OYLZWLJ[
[V [OL W\YPULYNPJ HNVUPZ[ .92 HWWLHYZ [V IL LZZLU[PHS PU 3;+PUK\JLK .79
OVTVSVNV\ZKLZLUZP[PaH[PVU^OPSLP[PZUV[WYPTHYPS`PU]VS]LKPU.79OVTVSVNV\Z
HUK OL[LYVSVNV\Z KLZLUZP[PaH[PVU PUK\JLK I` <+7NS\JVZL )V[O SPNHUKZ HJ[P]H[L
PU[YHJLSS\SHY,92WOVZWOVY`SH[PVU^P[OHKPMMLYLU[RPUL[PJZHUK[OYV\NOH.WYV[LPU
KLWLUKLU[ HUK . WYV[LPUPUKLWLUKLU[ TLJOHUPZT MVY 3;+ HUK <+7NS\JVZL
YLZWLJ[P]LS`
([ [OL SPNO[ VM [OLZL YLZ\S[Z ^L ZWLJ\SH[L [OH[ .79 HJ[P]H[PVU I` J`Z3;Z HUK
W\YPULYNPJSPNHUKZKPYLJ[Z[OLYLJLW[VY[V^HYKZKPMMLYLU[PU[YHJLSS\SHYYV\[L
*PHUH7L[HS,4)61!
+HUPLSL:L[HS17OHYTHJVS,_W;OLY! 007VZ[LY:LZZPVU
NAG 5
Nucleotidi, Acidi Nucleici, Genomi
5(+)06:@5;/,:0:,5A@4,:-964;/,)(*;,90(3
>693+(:=(3<()3,;663:-69=0;(405)
46565<*3,6;0+,:8<(5;0-0*(;065054(44(3:
*(99(+69090;(4690=(3,906(40*0(+63-6
69:64(5+6.0<:,77,:69*03,65(9+64(AA63(-9(5*,:*(
)6**07(63(9<..0,90:03=,9069(--(,3305(+0(
+LWHY[TLU[VM*SPUPJHS:JPLUJLZ+0:*6:LJ[PVUVM)PVJOLTPZ[Y`HUK
+LWHY[TLU[VM5\[YP[PVUHS,U]PYVUTLU[HSHUK(NYPJ\S[\YHS:JPLUJLZ+(
7VS`[LJOUPJ<UP]LYZP[`VM4HYJOL=PH9HUPLYP(UJVUH0[HS`
;OLW`YPKPULTVUVU\JSLV[PKLZ545HUK5(45OH]LILLUMVYHSVUN[PTLJVUZPKLYLK
HZ]P[HTPU)KLYP]LK5(+WYLJ\YZVYZ^P[OUVV[OLYIPVSVNPJHSM\UJ[PVUPUTHTTHSZ
/V^L]LY YLJLU[ TV\U[PUN L]PKLUJL Z\NNLZ[Z H KPYLJ[ YVSL VM 545 HZ H RL`
PU[LYTLKPH[L PU ]HYPV\Z TVSLJ\SHY WYVJLZZLZ SPRL [OVZL HMMLJ[PUN H_VU ]PHIPSP[` VY
NS\JVZLYLSH[LKTL[HIVSPJKPZVYKLYZ\UKLYZJVYPUN[OLULLKMVYZ\P[HISLX\HU[P[H[P]L
KL[LJ[PVU TL[OVKZ ;OL HUHS`[PJHS KL[LYTPUH[PVU VM [OLZL TVUVU\JSLV[PKLZ PU
IPVSVNPJHS ZHTWSLZ OHZ ILLU ZV MHY OHTWLYLK I` [OLPY SV^ HTV\U[Z HUK JOLTPJHS
WYVWLY[PLZ;VKH[LVUS`HML^545HUK5H45KL[LJ[PVUTL[OVKZTHPUS`YLS`PUN
VUYHKPVHJ[P]P[`Å\VYPTL[Y`VYZVWOPZ[PJH[LK3*4:HUHS`ZLZOH]LILLUYLWVY[LK
ZVTL[PTLZ ^P[O JVU[YHZ[PUN YLZ\S[Z ;OLYLMVYL [OL WH[OVWO`ZPVSVNPJHS YLSL]HUJL VM
[OLZLTVSLJ\SLZPZZ[PSS\UJSLHY/LYL^LYLWVY[VU[^VUV]LSLUa`TLJV\WSLKHZZH`Z
VM^PKLHWWSPJHIPSP[`MVY[OLKL[LYTPUH[PVUVM545HUK5H45PUTHTTHSPHUZ`Z[LTZ
;OLZLTL[OVKZHYLIHZLKVU[OLÅ\VYPTL[YPJKL[LJ[PVUVM[OLTVUVU\JSLV[PKLZLP[OLY
HM[LYKPYLJ[JOLTPJHSKLYP]H[PaH[PVUVYHM[LY[OLPYLUa`TH[PJJVU]LYZPVU[V5(+HUKP[Z
Z\IZLX\LU[X\HU[P[H[PVUI`HJ`JSPUNHZZH`>LL_WSVP[5(+IPVZ`U[OL[PJLUa`TLZ
MYVT IHJ[LYPHS ZV\YJLZ ^P[O WLJ\SPHY JH[HS`[PJ WYVWLY[PLZ [OH[ V\Y SHIVYH[VY` ÄYZ[
JVU[YPI\[LK [V PKLU[PM` HUK JOHYHJ[LYPaL PL P 5H45 HKLU`S`S[YHUZMLYHZL VM 5HK+
MHTPS` ZWLJPÄJHSS` JVU]LY[PUN 5H45 PU[V KLHTPKV5(+ PP 545 KLHTPKHZL HUK
545Z`U[OL[HZLYLZWVUZPISLMVY[OLZWLJPÄJPU[LYJVU]LYZPVUVM545PU[V5H45HUK
]PJL]LYZHYLZWLJ[P]LS`)V[OTL[OVKZYLZ\S[LK[VILHZZLUZP[P]LHZ[OL3*4:IHZLK
HZZH`Z^P[O[OLHK]HU[HNLVMILPUNT\JOMHZ[LYHUKSLZZL_WLUZP]L;OL`HSSV^LK
\Z[VWLYMVYTHWYLSPTPUHY`TL[HIVSPJWYVÄSPUNVMW`YPKPULTVUVU\JSLV[PKLZPUTV\ZL
[PZZ\LZHUKZLYH9LTHYRHIS`[OL`JHUILM\Y[OLYLTWSV`LKMVYTLHZ\YPUNHSSRUV^U
5455(45 Z`U[OLZPaPUN LUa`TL HJ[P]P[PLZ ^OVZL KPMMLYLU[PHS JVU[YPI\[PVU [V 5(+
Z`U[OLZPZPUWO`ZPVWH[OVSVNPJHSJVUKP[PVUZPZHTH[[LYVMPUJYLHZPUNPU[LYLZ[
007VZ[LY:LZZPVU
NAG 6
Nucleotidi, Acidi Nucleici, Genomi
0U]VS]LTLU[VMJ500PUWYVKY\NTL[HIVSPZT
+(50,3(50*63,-0365050*63,;;(:*/0),*05(;630:*0(330
96::(5(7,:0:0465,(33,.90504(90(.9(A0(;6AA0
+PWHY[PTLU[VKP*OPTPJHL-HYTHJPH<UP]LYZP[nKP:HZZHYP"
+PWHY[PTLU[VKP)PVSVNPH<UP]LYZP[nKP7PZH
*`[VZVSPJ »U\JSLV[PKHZLWOVZWOV[YHUZMLYHZL 00 J500 ZWLJPÄJ MVY V_`W\YPUL
TVUVWOVZWOH[LZ PZ H ^PKLZWYLHK LUa`TL ^P[O [OL OPNOLZ[ HJ[P]P[` PU [PZZ\LZ
JOHYHJ[LYPaLKI`HOPNO[\YUV]LYVMU\JSLPJHJPKZHZPUO\THUJVSVUJHYJPUVTHHUK
THSPNUHU[S`TWOVJ`[LZ(ZHTLTILYVM[OLV_`W\YPULJ`JSLJ500PZPU]VS]LKPU[OL
YLN\SH[PVUVMPU[YHJLSS\SHYJVUJLU[YH[PVUVM047(47.47HUKHSZV7977[OLYLMVYLPU
[OLYLN\SH[PVUVMW\YPULHUKW`YPTPKPULKLUV]VHUKZHS]HNLZ`U[OLZPZ
*SPUPJHS PU[LYLZ[ PU J`[VZVSPJ »U\JSLV[PKHZL 00 OHZ ILLU PUJYLHZPUN LUVYTV\ZS` PU
[OLZLSHZ[`LHYZUV[VUS`ILJH\ZLP[ZT95(SL]LSZLLTZ[VILHWYVNUVZ[PJMHJ[VY
PUHK\S[HJ\[LT`LSVPKSL\RLTPHI\[HSZVMVYP[ZWVZZPISLPU]VS]LTLU[PUHU[PJHUJLY
HUK HU[P]PYHS WYVKY\N TL[HIVSPZT HZ \UKLYSPULK I` PU ]P[YV Z[\KPLZ *500 PU
MHJ[ UV[ VUS` JV\SK IL YLZWVUZPISL MVY YLZPZ[HUJL [V U\JSLVZPKL HUHSVN\LZ ^P[O
P[Z JH[HIVSPJ M\UJ[PVU I\[ P[ TPNO[ HSZV [HRL WHY[ PU [OLPY HJ[P]H[PVU WH[O^H` I` P[Z
WOVZWOV[YHUZMLYHZL HJ[P]P[` 5V[^P[OZ[HUKPUN THU` YLZLHYJOLZ OH]L ILLU JHYYPLK
V\[PUVYKLY[V\UYH]LS[OPZWVPU[ZL]LYHSJVU[YHZ[PUNHZWLJ[ZYLTHPU[VILZVS]LK
-VY[OPZYLHZVU^LWLYMVYTLKL_]P]VL_WLYPTLU[Z\ZPUNO\THULTIY`VUPJRPKUL`
JLSSZ V]LYL_WYLZZPUN VY UV[ J500 [YLH[LK ^P[O [OYLL KPMMLYLU[ U\JSLVZPKL HUHSVNZ!
J`[HYHIPULÅ\KHYHIPULHUKNLTJP[HIPUL0UMHJ[L]LUPM[OLYVSLVMJ500PUU\JSLVZPKL
HUHSVN\L YLZPZ[HUJL OHZ ILLU L_[LUZP]LS` Z[\KPLK M\Y[OLY PU]LZ[PNH[PVU PZ YLX\PYLK
[V[OYV^SPNO[VUZL]LYHSHZWLJ[Z^OPJOZ[PSSYLTHPU\UJSLHY"PUHKKP[PVU[OLYLPZSP[[SL
L]PKLUJLMYVTL_]P]VZ[\KPLZ[OH[·U\JSLV[PKHZLZHJ[\HSS`WOVZWOVY`SH[LKLV_`
U\JSLVZPKLHUHSVN\LZ
=LY` WYLSPTPUHY` YLZ\S[Z ZLLT [V Z\NNLZ[ UV[ VUS` [OH[ J500 PZ KPYLJ[S` PU]VS]LK PU
WYVKY\NTL[HIVSPZTI\[HSZVUL^J`[V[V_PJZ[YH[LNPLZMVY[OVZLWYVKY\NZJVUZPKLYLK
JOHPU[LYTPUH[VYZ
;VaaPL[HS (YJO)PVJOLT)PVWO`Z
0WH[HHUK;VaaP7\YPU:PNUHSS
.HSTHYPUPL[HS3L\RLTPH
1VYKOLPTHUK+\TVU[L[))(
007VZ[LY:LZZPVU
NAG 7
Nucleotidi, Acidi Nucleici, Genomi
+PMMLYLU[PHS9VSLZVM:5(0:3<.;YHUZJYPW[PVU-HJ[VYPU
[OL,WP[OLSPHSHUK:[YVTHS*VTWHY[TLU[ZVM[OL/\THU
7YVZ[H[L*HUJLY
:699,5;056*(936 ;<765,4(90(.9(A0(+04,6:,9,5(
4<:0(5070,96+00306*(9405,+0*(936,44(
:LJ[PVUVM(UH[VTPJ7H[OVSVN`HUK4VSLJ\SHY4LKPJPUL+LWHY[TLU[VM4LKPJPUL
HUK:JPLUJLZVM(NPUNHUK+LWHY[TLU[VM)PVTLKPJHS:JPLUJLZ¸.K»(UU\UaPV¹
<UP]LYZP[`VM*OPL[P7LZJHYH*OPL[P0[HS`"
*L:0(NPUN9LZLHYJO*LU[LY¸.K»(UU\UaPV¹<UP]LYZP[`-V\UKH[PVU*OPL[P0[HS`
0U[YVK\J[PVU:5(0:3<.PZHaPUJÄUNLY[YHUZJYPW[PVUMHJ[VY[OH[HJ[ZHZHTHZ[LY
YLN\SH[VY VM JLSS TPNYH[PVU I` [YPNNLYPUN LWP[OLSPHSTLZLUJO`THS [YHUZP[PVU ,4;
K\YPUNLTIY`VUPJKL]LSVWTLU[HUK[\TVYTL[HZ[HZPaH[PVUHUKHSZVYLN\SH[LZJLSS
J`JSLWYVNYLZZPVUHUKZ\Y]P]HS0U[OPZZ[\K`^LPU]LZ[PNH[LKP[ZYVSLPUWYVZ[H[LJHUJLY
7*H\UKLYUVYTHSHUKHUKYVNLUKLWYP]H[PVU[OLYHW`(+;JVUKP[PVUZ
4L[OVKZ3HZLYJHW[\YLTPJYVKPZZLJ[PVUHUKYLHS[PTL9;7*9^LYL\ZLK[VKL[LYTPUL
:3<. NLUL L_WYLZZPVU SL]LSZ PU UVYTHS HUK JHUJLYV\Z WYVZ[H[PJ LWP[OLSP\T HUK
Z[YVTHKPMMLYLU[PH[PUNULVWSHZ[PJMVJP^P[OSV^]LYZ\ZOPNO.SLHZVUNYHKL,WP[OLSPHS
HUKZ[YVTHSJLSSZ^LYLPZVSH[LKMYVTJHUJLYV\ZHUKUVYTHSWYVZ[H[LZWLJPTLUZMYVT
\U[YLH[LK HUK (+;[YLH[LK WH[PLU[Z ^OV \UKLY^LU[ YHKPJHS WYVZ[H[LJ[VT`
MVY 7*H 0TT\UVZ[HPUPUNZ HUK JVUMVJHS TPJYVZJVW` ^LYL \ZLK [V ]PZ\HSPaL :3<.
L_WYLZZPVUH[[OLWYV[LPUSL]LS
9LZ\S[Z :3<. NLUL L_WYLZZPVU SL]LSZ ^LYL ZPNUPÄJHU[S` 7# SV^LY PU [OL
ULVWSHZ[PJ^P[OUVHWWYLJPHISLKPMMLYLUJLIL[^LLUSV^HUKOPNONYHKL7*H]LYZ\Z
UVYTHS WYVZ[H[PJ LWP[OLSPH HUK PU ULVWSHZ[PJ ]LYZ\Z UVYTHS Z[YVTH :3<. ^HZ
WYLKVTPUHU[S`L_WYLZZLKPU[OLZ[YVTHSYH[OLY[OHU[OLLWP[OLSPHSJVTWHY[TLU[^P[O
OPNOLYSL]LSZPU[OLULVWSHZ[PJZ[YVTH
0TT\UVZ[HPUPUNZZOV^LK[OH[PU[OLUVYTHSWYVZ[H[LNSHUKZIV[OIHZHSHUKS\TPUHS
ZLJYL[VY` JLSSZ Z[YVUNS` L_WYLZZLK :3<. PU [OLPY U\JSLP ^OLYLHZ :3<. Z[HPUPUN
YHUNLKMYVT]LY`^LHR[VTVKLYH[LPU[OLULVWSHZ[PJNSHUKZ:3<.L_WYLZZPVU^HZ
PU]LYZLS`YLSH[LK[V[OH[VM[OLJLSSJ`JSLYLN\SH[VYJ`JSPU+HUK[OLWYVSPMLYH[PVU
THYRLY2P
3HZ[S`(+;NLULYHSS`LUOHUJLK:3<.L_WYLZZPVUPU[OL7*HZ[YVTHSJVTWHY[TLU[
*VUJS\ZPVUZ ;OLZL WYLSPTPUHY` ÄUKPUNZ Z\NNLZ[ [OH[ PU [OL WYVZ[H[L :3<. TH`
PUOPIP[ ULVWSHZ[PJ LWP[OLSPHS JLSS WYVSPMLYH[PVU YH[OLY [OHU HJ[ HZ HU ,4; WYVTV[LY 0[
TH`HSZVOH]LHKPMMLYLU[YVSLPU[OL7*HHZZVJPH[LKZ[YVTH^OVZLYLZWVUZL[V(+;
TH` OH]L PTWVY[HU[ JSPUPJHS HUK WH[OVSVNPJHS PTWSPJH[PVUZ (SS [OLZL PZZ\LZ HYL [OL
Z\IQLJ[VMVUNVPUNPU]LZ[PNH[PVUPUV\YSHIVYH[VYPLZ
007VZ[LY:LZZPVU
NAG 8
Nucleotidi, Acidi Nucleici, Genomi
;OL,TIY`VUPJ:[LT*LSS4HYRLY:6?PZ+V^U9LN\SH[LK
PU7YVZ[H[L*HUJLYHUK<W9LN\SH[LKI`(UKYVNLU
>P[OKYH^HS
+04,6:,9,5( :699,5;056*(936;<765,4(90(.9(A0(
4<:0(5070,96+00306*(9405,+0*(936,44(
:LJ[PVUVM(UH[VTPJ7H[OVSVN`HUK4VSLJ\SHY4LKPJPUL+LWHY[TLU[VM4LKPJPUL
HUK:JPLUJLZVM(NPUNHUK+LWHY[TLU[VM)PVTLKPJHS:JPLUJLZ¸.K»(UU\UaPV¹
<UP]LYZP[`VM*OPL[P7LZJHYH*OPL[P0[HS`"
*L:0(NPUN9LZLHYJO*LU[LY¸.K»(UU\UaPV¹<UP]LYZP[`-V\UKH[PVU*OPL[P0[HS`
0U[YVK\J[PVU :6? [YHUZJYPW[PVU MHJ[VY PZ LZZLU[PHS [V THPU[HPU ZLSMYLUL^HS VM
\UKPMMLYLU[PH[LK LTIY`VUPJ Z[LT JLSSZ ,:* HUK WSH`Z H YVSL PU KPMMLYLU[PH[PVU
WYVSPMLYH[PVU HUK HWVW[VZPZ *VUZPZ[LU[ ^P[O [OL O`WV[OLZPZ [OH[ Z[LTULZZ HUK
LTIY`VUPJWH[O^H`ZTH`YLHJ[P]H[LK\YPUNVUJVNLULZPZ:6?PZV]LYL_WYLZZLKPUH
]HYPL[`VM[\TVYZHUKHSZVHZZVJPH[LK^P[OJHUJLYHNNYLZZP]LULZZ0UWYVZ[H[LJHUJLY
7*HOV^L]LYP[ZYVSLPZZ[PSS\UJSLHYZPUJLJVU[YHZ[PUNKH[HHYLYLWVY[LKHIV\[P[Z
L_WYLZZPVUPUJLSSSPULZHUKO\THUZHTWSLZ
4L[OVKZ >L \ZLK SHZLY JHW[\YL TPJYVKPZZLJ[PVU MVSSV^LK I` YLHS[PTL 9;7*9
[V KL[LYTPUL :6? NLUL L_WYLZZPVU SL]LSZ PU [OL UVYTHS HUK JHUJLYV\Z WYVZ[H[PJ
LWP[OLSP\THUKZ[YVTHHUKKPZJYPTPUH[LULVWSHZ[PJMVJP^P[OSV^]LYZ\ZOPNO.SLHZVU
NYHKL,WP[OLSPHSHUKZ[YVTHSJLSSZ^LYLPZVSH[LKMYVTJHUJLYV\ZHUKUVYTHSWYVZ[H[L
ZWLJPTLUZ MYVT \U[YLH[LK HUK HUKYVNLU KLWYP]H[PVU [OLYHW` (+;[YLH[LK
7*H WH[PLU[Z ^OV \UKLY^LU[ YHKPJHS WYVZ[H[LJ[VT` MVY 7*H HUK UVYTHS WYVZ[H[L
ZWLJPTLUZ MYVT WH[PLU[Z ^OV OHK \UKLYNVUL J`Z[VWYVZ[H[LJ[VT` MVY ISHKKLY
JHUJLYJVU[YVSWH[PLU[Z
9LZ\S[Z:6?NLULL_WYLZZPVUSL]LS^HZZPNUPÄJHU[S`7#OPNOLYPU[OLUVYTHS
WYVZ[H[PJLWP[OLSP\T[OHUPU[OLZ\YYV\UKPUNZ[YVTHHUKVMNYLH[LYPTWVY[HUJL^HZ
ZPNUPÄJHU[S`YLK\JLKPU[OLULVWSHZ[PJLWP[OLSP\TMYVTIV[OSV^HUKOPNONYHKL7*H
HUKZSPNO[S`YLK\JLKPU[OLULVWSHZ[PJHZVWWVZLK[V[OLUVYTHSZ[YVTH/VYTVUL
[OLYHW`PUJYLHZLK:6?L_WYLZZPVUPUULVWSHZ[PJHUKWHY[PJ\SHYS`PUUVYTHSWYVZ[H[PJ
LWP[OLSP\T;OLTLHUL_WYLZZPVUSL]LSVM:6?PUUVYTHSZHTWSLZMYVT[OL\U[YLH[LK
WH[PLU[Z^HZZPTPSHY[V[OH[VI[HPULKPUUVYTHSZHTWSLZMYVTJVU[YVSZ
*VUJS\ZPVUZ6\YWYLSPTPUHY`KH[HPUKPJH[L[OH[:6?L_WYLZZPVUPZKV^UYLN\SH[LK
K\YPUN WYVZ[H[L JHYJPUVNLULZPZ HUK [OH[ HUKYVNLU ^P[OKYH^HS SLHKZ [V P[Z \W
YLN\SH[PVU0[^V\SKZLLT[OH[PU[OLWYVZ[H[L¸Z[LTULZZ¹HUK¸THSPNUHUJ`¹KVUV[
^HSRHYTPUHYT,:*YLSH[LKNLULZZ\JOHZ:6?TH`ILKPMMLYLU[PHSS`YLN\SH[LKPU
[OLKPMMLYLU[HK\S[[PZZ\LZHUKJHYJPUVNLUPJWYVJLZZLZ;OPZPZZ\LHUK:6?M\UJ[PVUZ
PU7*HHYLZ\IQLJ[ZVMPU]LZ[PNH[PVUPUV\YSHIVYH[VYPLZ
007VZ[LY:LZZPVU
NAG 9
Nucleotidi, Acidi Nucleici, Genomi
*VYYLSH[PVUIL[^LLU:S\N[YHUZJYPW[PVUMHJ[VYHUKTP9
PUIYLHZ[JHUJLYJLSSZ
3(4),9;050,30:(),;;(36330(5+9,(=,AA(30-,+,90*(
7,563(AA03,;0A0(.(4)(9096),9;6(5+96),9;(70=(
+LWHY[TLU[VM)PVJOLTPZ[Y`HUK4VSLJ\SHY)PVSVN`
<UP]LYZP[`VM-LYYHYH-LYYHYH0;(3@
)YLHZ[JHUJLYZHUK[OLPYTL[HZ[H[PJWYVNYLZZPVUHYLTHPUS`KPYLJ[LKI`LWP[OLSPHS[V
TLZLUJO`THS [YHUZP[PVU ,4; H WOLUVTLUVU Z\WWVY[LK I` ZWLJPÄJ [YHUZJYPW[PVU
MHJ[VYZHUKTP95(ZVUS`PUWHY[RUV^U
0UVYKLY[VPU]LZ[PNH[LHWVZZPISLJVYYLSH[PVUIL[^LLU:S\N[YHUZJYPW[PVUMHJ[VYHUK
TP9^LWLYMVYTLKNLULZPSLUJPUNHNHPUZ[:S\NHUKTP9PUHNNYLZZP]LIYLHZ[
JHUJLYJLSSSPULZHUKL]HS\H[LKI`X9;7*9HUK>LZ[LYUISV[[OLL_WYLZZPVUVMNLULZ
PU]VS]LKPUZ\WWVY[PUN[OLIYLHZ[JHUJLYWOLUV[`WL
>LZOV^LK[OH[:S\NZPSLUJPUNZPNUPÄJHU[S`KLJYLHZLK[OLSL]LSVMTP9WVZP[P]LS`
HMMLJ[LK [OL L_WYLZZPVU VM LWP[OLSPHS THYRLYZ PUJS\KPUN ,Z[YVNLU 9LJLW[VY Ơ
,JHKOLYPUHUK;97:HUKJH\ZLKHKLJYLHZLVM]PTLU[PUHUKHZ[YVUNPUOPIP[PVUVM
JLSSTPNYH[PVUL]HS\H[LKI`^V\UKOLHSPUNHZZH`Z>LKLTVUZ[YH[LK[OH[TP9PZ
H:S\N[HYNL[NLULHUKPKLU[PÄLKI`JOYVTH[PUPTT\UVWYLJPWP[H[PVUHZWLJPÄJYLNPVU
VM TP9 WYVTV[LY [OH[ PZ [YHUZJYPW[PVUHSS` HJ[P]L HUK PZ PU]VS]LK PU [OL IPUKPUN
VM:S\N¸PU]P]V¹*VTWSL[LRUVJRKV^UVMTP9L_WYLZZPVUI`[YHUZMLJ[PVU^P[O
HU[HNVTP9JH\ZLZHZPNUPÄJH[P]LKLJYLHZLVM:S\NL_WYLZZPVUZ\NNLZ[PUN[OH[
:S\N PZ UV[ H KPYLJ[ [HYNL[ VM TP9 I\[ YH[OLY [OH[ H ULNH[P]L YLN\SH[VY VM :S\N
JV\SK IL H TP9 [HYNL[ /V^L]LY TP9 RUVJRKV^U ZPNUPÄJHU[S` H[[LU\H[LK
[OL NHW JSVZPUN PU [OL JLSSZ I\[ UV[ HZ T\JO HZ [OH[ VIZLY]LK PU :S\NYLWYLZZLK
JLSSZ 0U HKKP[PVU YLZJ\L L_WLYPTLU[ ^P[O WYLTP9 KPK UV[ IHSHUJL ZP:S\N
TLKPH[LKLMMLJ[ZZ\NNLZ[PUN[OH[:S\NZPSLUJPUNJV\SKZPNUPÄJHU[S`WYV[LJ[JLSSZMYVT
WYVNYLZZPVU[V^HYKZHUHNNYLZZP]LWOLUV[`WLVYTL[HZ[H[PJZ[PT\SP[OH[PU[OPZJHZL
HYLYLWYLZLU[LKI`TP9V]LYL_WYLZZPVU
-VY[OLÄYZ[[PTL[OLL]PKLUJLVMHM\UJ[PVUHSSPURIL[^LLU:S\NHUKTP9PUIYLHZ[
JHUJLY JLSSZ PZ YLWVY[LK ;OLYLMVYL PU [OL JVU[L_[ OLYL HUHS`aLK ^L TH` JVUJS\KL
[OH[TP9L_WYLZZPVUPZPUWHY[KLWLUKLU[VU:S\NHUK[OH[:S\NWSH`ZHTVYL
PTWVY[HU[YVSL[OHUTP9PUJLSSTPNYH[PVUHUKPU]HZPVU
007VZ[LY:LZZPVU
NAG 10
Nucleotidi, Acidi Nucleici, Genomi
7YLJSPUPJHS[LZ[PUNVM757PUOPIP[VYZHUHSVN\LZVM
0TT\JPSSPU.MVY[OL[OLYHW`VM3LZJO5`OHUKPZLHZL!
WYLSPTPUHY`PU]P[YVZ[\KPLZ
)(3+050,=( 1(*64,330.()90,33(*69,330-,+,90*6
40*/,30=(55(
+PWHY[PTLU[VKP)PV[LJUVSVNPL
+PWHY[PTLU[V-HYTHJV*OPTPJV;LJUVSVNPJV<UP]LYZP[nKP:PLUH
3LZJO5`OHU +PZLHZL 35+ PZ H YHYL ?SPURLK NLUL[PJ KPZLHZL JOHYHJ[LYPaLK I`
O`WLY\YPJHLTPH NV\[ HUK ZL]LYL UL\YVSVNPJHS Z`UKYVTL /`WV_HU[OPULN\HUPUL
WOVZWOVYPIVZ`S[YHUZMLYHZL/79;KLÄJPLUJ`VJJ\YZJH\ZPUN\YPJHJPKO`WV_HU[OPUL
HUK _HU[OPUL HJJ\T\SH[PVU <YPJ HJPK L_JLZZ PZ JVTTVUS` THUHNLK I` _HU[OPUL
V_PKHZLPUOPIP[VYZ`PLSKPUNPU_HU[OPULHUKO`WV_HU[OPULWYVK\J[PVU;OLIPVJOLTPJHS
IHZPZVM[OLUL\YVSVNPJHSWH[OVSVN`PZZ[PSS\UJSLHYHUKUVLMMLJ[P]L[OLYHW`PZH]HPSHISL
+LHaHN\HUPUL KLYP]H[P]LZ PUOPIP[PUN O\THU W\YPUL U\JSLVZPKL WOVZWOVY`SHZL 757
^LYL\ZLKPUJSPUPJHS[YPHSZ[VSV^LY\YPJHJPKPUNV\[`WH[PLU[Z;OPZZ[\K`PZHPTLKH[
[LZ[PUN[OLYLSPHIPSP[`VM757PUOPIP[VYZHZH[OLYHW`MVY35+\YH[LHUKO`WV_HU[OPUL
L_JLZZ
( KLHaHN\HUVZPUL HUHSVN\L VM 0T\JPSSPU. :LTLYHYV L[ HS 1 4LK *OLT "
JVTWV\UK H ^HZ \ZLK HZ 757 PUOPIP[VY 757 PUOPIP[PVU HUK HWWHYLU[ RPUL[PJ
JVUZ[HU[Z^LYLTLHZ\YLKPUJY\KLS`ZH[LZMYVTUVYTHSLY`[OYVJ`[LZHUKÄIYVISHZ[Z
I` /73*SPURLK TL[OVKZ 4PJOLSP L[ HS ))( 5HUVTVSHY H JVUJLU[YH[PVUZ
THYRLKS`KLJYLHZLK=TH_UV[HMMLJ[PUN2TPUVZ\NNLZ[PUNUVUJVTWL[P[P]LPUOPIP[PVU
7YPTHY` J\S[\YLZ VM JVU[YVS Z\IQLJ[Z ZRPU ÄIYVISHZ[Z ^LYL NYV^U ^P[O +4,4 WS\Z
OP-*:WLUPJPSSPUZ[YLW[VT`JPUH[‡*HUK*6PU[OLWYLZLUJLVY
HIZLUJLVMHU4HUK›4HUKVM›4PUVZPULMVYHUKO7LYJOSVYPJ
L_[YHJ[ZVI[HPULKMYVTTLKP\THUKJLSSZ^LYL[OLUHUHS`ZLKI`/73*5VZPNUPÄJHU[
YLK\J[PVUVM]PHISLJLSSZ^HZKLTVUZ[YH[LKI`4;;[LZ[HUKUVYLTHYRHISLHS[LYH[PVU
PU JLSS U\JSLV[PKL WH[[LYU ^HZ MV\UK HM[LY HU` PUJ\IH[PVU JVUKP[PVU ^OPSL ›4 H
JH\ZLKO`WV_HU[OPULHUK_HU[OPULKLJYLHZL[VNL[OLY^P[OPUVZPULHUKN\HUVZPUL
PUJYLHZLPU[OLJ\S[\YLTLKP\T
7YLZLU[KH[HKLTVUZ[YH[LLMMLJ[P]L757PUOPIP[PVUI`SV^HJVUJLU[YH[PVU^P[OUV
HWWYLJPHISL[V_PJP[`:[\KPLZHYLPUWYVNYLZZPU/79;KLÄJPLU[JLSSZPU]PL^VMHUL^
[OLYHWL\[PJZ[YH[LN`HNHPUZ[V_PW\YPULHJJ\T\SH[PVU
(JRUV^SLKNLTLU[Z![OPZZ[\K`PZM\UKLKI`35+-HTPNSPL0[HSPHUL653<:
007VZ[LY:LZZPVU
NAG 11
Nucleotidi, Acidi Nucleici, Genomi
;VIYHT`JPUPUOPIP[ZPU[LYSL\RPUNLULL_WYLZZPVUHUK
HJ[ZHZZ\WWYLZZVYVMWYLTH[\YL[LYTPUH[PVUJVKVUZPU[OL
HIZLUJLVM54+
(3;(4<9(50*63( .(4)(9096),9;6)69.(;;04650*(
0UZ[P[\[LVM)PVTLTIYHULHUK)PVLULYNL[PJZ*59
=PH(TLUKVSH()HYP0[HS`"
+LWHY[TLU[VM)PVJOLTPZ[Y`HUK4VSLJ\SHY)PVSVN`
<UP]LYZP[`VM-LYYHYH=PH-VZZH[VKP4VY[HYH-LYYHYH0[HS`"
(TPUVNS`JVZPKLZ HYL ^PKLS` \ZLK MVY [OL [YLH[TLU[ VM J`Z[PJ ÄIYVZPZ *- H MH[HS
H\[VZVTHS YLJLZZP]L NLUL[PJ KPZLHZL JOHYHJ[LYPaLK I` WLYZPZ[LU[ W\STVUHY`
PUMLJ[PVU HUK L_[LUZP]L S\UN PUÅHTTH[PVU 0U HKKP[PVU [V [OLPY YVSL HZ HU[PIPV[PJZ
HTPUVNS`JVZPKLZOH]LILLUZOV^U[VHJ[HZZ\WWYLZZVYZVMUVUZLUZLT\[H[PVUZ[OH[
NLULYH[L WYLTH[\YL [YHUZSH[PVU [LYTPUH[PVU 7;*Z 7;*Z JVUZ[P[\[L [OL TVSLJ\SHY
IHZPZVMTHU`NLUL[PJKPZLHZLZPUJS\KPUN*-VMJ`Z[PJÄIYVZPZHSSLSLZJVU[HPU
7;*Z HZ SLHK [V [OL Z`U[OLZPZ VM [Y\UJH[LK UVUM\UJ[PVUHS VY WHY[PHSS` M\UJ[PVUHS
WYV[LPU :\WWYLZZPVU VM [YHUZSH[PVU [LYTPUH[PVUZ H[ 7;*Z YLHK[OYV\NO OHZ ILLU
KL]LSVWLKHZH[OLYHWL\[PJZ[YH[LN`[VYLZ[VYLM\SSSLUNO[WYV[LPUPUZL]LYHSNLUL[PJ
KPZLHZLZ7OLUV[`WPJJVUZLX\LUJLZVM7;*ZJHUILL_HJLYIH[LKI`[OLUVUZLUZL
TLKPH[LK T95( KLJH` 54+ WH[O^H` [OH[ KL[LJ[ HUK KLNYHKL 7;* JVU[HPUPUN
T95(4VK\SH[PVUVM54+[OLYLMVYLPZHSZVVMPU[LYLZ[HZHWV[LU[PHS[HYNL[MVY[OL
Z\WWYLZZPVU [OLYHW` ;VIYHT`JPU PZ HU HTPUVNS`JVZPKL HU[PIPV[PJ UVYTHSS` \ZLK [V
[YLH[ 7ZL\KVTVUHZ HLY\NPUVZH W\STVUHY` PUMLJ[PVU PU *- WH[PLU[Z 0U [OL WYLZLU[
Z[\K`^LOH]LPU]LZ[PNH[LKVUVULOHUK[OLWYVWLY[PLZVM;VIYHT`JPUHZHTVK\SH[VY
VM[OLL_WYLZZPVUVM03HZZVJPH[LK^P[OPUÅHTTH[PVUVM[OL*-HPY^H`WH[OVSVN`PU
*-0)JLSSZPUK\JLK^P[OWYVPUÅHTTH[VY`J`[VRPUL;5-Į6U[OLV[OLYOHUKI`
\ZPUN[OL`LHZ[HZHNLUL[PJZ`Z[LT^LOH]LL_HTPULK[OLHIPSP[`VM;VIYHT`JPU[V
Z\WWYLZZ7;*ZHZHM\UJ[PVUVM[OLWYLZLUJLVYHIZLUJLVM54+9LZ\S[ZKLTVUZ[YH[L
[OH[;VIYHT`JPUPZHNVVKPUOPIP[VYVM03L_WYLZZPVUHUKL_OPIP[ZYLHK[OYV\NOHIPSP[`
VU7;*ZPUJVUKP[PVUZPU^OPJO54+PZUV[VWLYH[PUNPU[OLJLSS
(JRUV^SLKNLTLU[Z;OPZ^VYR^HZZ\WWVY[LKI`JVU[YPI\[PVUMYVT--*0[HSPHU*`Z[PJ-PIYVZPZ9LZLHYJO
-V\UKH[PVU[V4)--*
40<90[HSPHU4PUPZ[Y`VM<UP]LYZP[`HUK9LZLHYJO-VUKHaPVUL*HYPWHYV
*HZZHKP9PZWHYTPVKP7HKV]HL9V]PNV*0)HUK;LSL[OVU..7
007VZ[LY:LZZPVU
NAG 12
Nucleotidi, Acidi Nucleici, Genomi
7LW[PKLU\JSLPJHJPKZ[HYNL[PUNTP9TVK\SH[LW2PW
L_WYLZZPVUPUIYLHZ[JHUJLY4+(4)JLSSZ
)96.5(9(,3,6569(-())90,590*((040-()06(3,?
)0(5*/050*63,;;(-056;;0(3,::0()9,=,.30,90.0<30(
)69.(;;04650*(*699(+05096),9;64(9*/,33096:(5.,3(
.(4)(9096),9;6
)PV7OHYTH5L[+LWHY[TLU[VM)PVJOLTPZ[Y`HUK4VSLJ\SHY)PVSVN`
-LYYHYH<UP]LYZP[`-LYYHYH0[HS`"
+LWHY[TLU[VM6YNHUPJHUK0UK\Z[YPHS*OLTPZ[Y`<UP]LYZP[`VM7HYTH0[HS`"
3HIVYH[VY`MVY[OL+L]LSVWTLU[VM7OHYTHJVSVNPJHSHUK7OHYTHJVNLUVTPJ;OLYHW`
VM;OHSHZZHLTPH)PV[LJOUVSVN`*LU[LY-LYYHYH<UP]LYZP[`-LYYHYH0[HS`"
7LW[PKL 5\JSLPJ (JPKZ 75(Z HYL +5( TPTPJZ ^OLYL WZL\KVWLW[PKL IHJRIVUL
PZ JVTWVZLK VM 5HTPUVL[O`SNS`JPUL \UP[Z 75(Z ^LYL MV\UK [V IL L_JLSSLU[
JHUKPKH[LZMVYHU[PZLUZLHUKHU[PNLUL[OLYHWPLZ4PJYV95(ZTP95(ZTP9ZHYLH
MHTPS`VMZTHSSUVUJVKPUN95(Z[OH[YLN\SH[LNLULL_WYLZZPVUI`ZLX\LUJLZLSLJ[P]L
[HYNL[PUN VM T95(Z SLHKPUN [V H [YHUZSH[PVUHS YLWYLZZPVU VY T95( KLNYHKH[PVU
4PJYV95(ZWSH`PUNHJY\JPHSYVSLPU[OLPUP[PH[PVUHUKWYVNYLZZPVUVMO\THUJHUJLYHYL
KLÄULKHZVUJVNLUPJTP95(ZVUJVTP9Z0UTHU`[`WLZVM[\TVYZOPNOSL]LSZVM
TP9PZMV\UKHZZVJPH[LK^P[OPUOPIP[PVUVML_WYLZZPVUVMP[Z[HYNL[W2PWT95(
^OPJOLUJVKLZH]LY`PTWVY[HU[VUJVZ\WWYLZZVYWYV[LPU0U[OPZZ[\K`^LKLZJYPIL
[OLHJ[P]P[`VMH75([HYNL[PUNTPJYV95(HZZVJPH[LK[VIYLHZ[JHUJLY>LÄYZ[
HUHS`aLK[OLL_WYLZZPVUVMTP9HUKW2PWT95(PUKPMMLYLU[IYLHZ[JHUJLYJLSS
SPULZ HUK MV\UK [OH[ OPNO L_WYLZZPVU VM TP9 HUK SV^ WYVK\J[PVU VM W2PW HYL
WYLZLU[PU4+(4)JLSSZ75(HNHPUZ[TP9^LYLKLZPNULKPUVYKLY[VIPUK
]LY`LMÄJPLU[S`[V[OL[HYNL[95(Z[YHUKHUK[V\UKLYNVLMÄJPLU[\W[HRLPU[HYNL[JLSSZ
(WVS`HYNPUPUL75(JVUQ\NH[L[HYNL[LKHNHPUZ[TP99WLW75(HZOV^LK
IV[O ]LY` OPNO HMÄUP[` MVY JVTWSLTLU[HY` U\JSLPJ HJPKZ HUK LMÄJPLU[ \W[HRL ^P[OPU
[HYNL[JLSSZ^P[OV\[[OLULLKVM[YHUZMLJ[PVUYLHNLU[Z<UTVKPÄLK75(^P[O[OLZHTL
ZLX\LUJL KPZWSH`LK ]LY` WVVY JLSS\SHY \W[HRL *VUZPZ[LU[S` VUS` 9WLW75(H
Z[YVUNS`PUOPIP[LKTP9;HYNL[PUNTP9I`75(YLZ\S[LKPUHSV^LYPUNVM[OL
O`IYPKPaH[PVUSL]LSZVMTP9TLHZ\YLKI`9;X7*9I\WYLN\SH[PVUVMW2PW
T95( HUK WYV[LPU TLHZ\YLK I` 9;X7*9 HUK >LZ[LYU )SV[ HUHS`ZPZ ;OL THQVY
JVUJS\ZPVUVM[OPZZ[\K`PZ[OH[LMÄJPLU[KLSP]LY`VMHU[PTP975([OYV\NOHZ\P[HISL
WLW[PKLJHYYPLY9WLW75(HSLHKZ[VPUOPIP[PVUVMTP9HJ[P]P[`HS[LYPUN[OL
L_WYLZZPVUVMTP9YLN\SH[LKM\UJ[PVUZPUIYLHZ[JHUJLYJLSSZ
007VZ[LY:LZZPVU
NAG 13
Nucleotidi, Acidi Nucleici, Genomi
;YHUZJYPW[PVUMHJ[VYZ;-ZULNH[P]LS`YLN\SH[PUNƢNSVIPU
NLUL[YHUZJYPW[PVU!)*3(PZKV^UYLN\SH[LKK\YPUN
TP[OYHT`JPUPUK\J[PVUVMLY`[OYVPKJLSSZMYVTȕ[OHSHZZLTPH
WH[PLU[Z
-056;;0(3,::0( )0(5*/050*63,;;(A<**(;6*90:;05()9,=,.30,90
.0<30(3(47965;003(90()96.5(9(,3,6569(.(4),90504(90(
90;()69.(;;04650*(.(4)(9096),9;6
3HIVYH[VY`MVY[OL+L]LSVWTLU[VM7OHYTHJVSVNPJHSHUK7OHYTHJVNLUVTPJ;OLYHW`
VM;OHSHZZLTPH)PV[LJOUVSVN`*LU[LY-LYYHYH<UP]LYZP[`-LYYHYH0[HS`"
+LWHY[TLU[VM9LWYVK\J[PVUHUK.YV^[O7HLKPH[YPJHUK(KVSLZJLU[<UP[
:(UUH/VZWP[HS-LYYHYH0[HS`
0UK\J[PVUVMML[HSOLTVNSVIPU/I-PZJVUZPKLYLKHUL^WYVTPZPUNZ[YH[LN`[V[YLH[
ȕ[OHSHZZLTPH ^OLYL [OL WYVK\J[PVU VM HK\S[ OLTVNSVIPU /I( PZ PTWHPYLK I`
T\[H[PVUZHMMLJ[PUN[OLȕNSVIPUNLUL;OYLLTHQVYSVJP?TU/).ZPUNSLU\JSLV[PKL
WVS`TVYWOPZT /):34@) PU[LYNLUPJ YLNPVU VU JOYVTVZVTL X HUK )*3(
JVU[YPI\[L[VOPNO/I-WYVK\J[PVU;OLZLKH[HHYLPUSPUL^P[OHNLULYHSJVUJLW[[OH[^HZ
MVSSV^LK\ZPUN+5(IPUKPUNKY\NZ+)+VYVSPNVU\JSLV[PKLKLJV`ZMVY[YHUZJYPW[PVU
MHJ[VYZKLJV`6+5HISL[VPUOPIP[;-+5(PU[LYHJ[PVUZ[OL[YHUZJYPW[PVUMHJ[VYKLJV`
Z[YH[LN`;-+"PM[OLO\THUȕNSVIPUNLULZHYL\UKLYULNH[P]L[YHUZJYPW[PVUHSJVU[YVS
JH\ZLKI`[YHUZJYPW[PVUMHJ[VYZL_LY[PUNYLWYLZZVYHJ[P]P[PLZ[OLU[HYNL[PUN[OLZL;-Z
TPNO[IL[OLYHWL\[PJHSS`YLSL]HU[7\[H[P]LYLWYLZZVYZVMȕNSVIPUNLUL[YHUZJYPW[PVU
HYL6J[4@)HUK)*3(
>L OH]L Z[\KPLK [OL L_WYLZZPVU VM )*3( K\YPUN /I- PUK\J[PVU VM LY`[OYVPK
WYLJ\YZVY JLSSZ ,Y7* MYVT ȕ[OHSHZZLTPH WH[PLU[Z [YLH[LK ^P[O TP[OYHT`JPU ;OL
SL]LSZ VM ȕNSVIPU HUK )*3( T95( ^LYL HZZH`LK I` X\HU[P[H[P]L YLHS[PTL 9;
7*9"[OLWYVK\J[PVUVMOLTVNSVIPUI`/73*
;OL YLZ\S[Z VI[HPULK Z\NNLZ[ [OH[ PUJYLHZL VM ȕNSVIPU NLUL L_WYLZZPVU HUK /I-
WYVK\J[PVU PZ WYLJLKLK I` H ZOHYW KLJYLHZL VM )*3( SL]LSZ PU ,Y7* MYVT
ȕ[OHSHZZLTPHWH[PLU[Z[YLH[LK^P[OU44;/
;OLZL YLZ\S[Z Z\NNLZ[ [OH[ )*3( SL]LSZ HYL HZZVJPH[LK ^P[O ȕNSVIPU NLUL
L_WYLZZPVU;OLYLMVYLKPZY\W[PUN[OLIPUKPUNZVM[OL)*3([YHUZJYPW[PVUHSJVTWSL_
[V[OLȕNSVIPUNLULWYVTV[LYWYV]PKLZHUV]LSHWWYVHJOMVYPUK\JPUNL_WYLZZPVUVM
[OLȕNSVIPUNLULZ
(JRUV^SLKNLTLU[Z ;OPZ ^VYR ^HZ Z\WWVY[LK I` H NYHU[ I` 40<9 0[HSPHU 4PUPZ[Y` VM <UP]LYZP[` HUK
9LZLHYJO-VUKHaPVUL*HYPWHYV*HZZHKP9PZWHYTPVKP7HKV]HL9V]PNV*0)I`;LSL[OVU..7
;OPZYLZLHYJOPZHSZVZ\WWVY[LKI`(ZZVJPHaPVUL=LUL[HWLYSH3V[[HHSSH;HSHZZLTPH(=3;9V]PNV
007VZ[LY:LZZPVU
5L\YVJOPTPJH
5,<
NEU 1
Neurochimica
;YPTL[O`S[PU;4;[YLH[TLU[VM7*JLSSZPUK\JLZ[OL
YLSLHZLVMNYV\W00(ZLJYL[VY`WOVZWOVSPWHZL(.00(73(
)0(.0650(5.,30,*69=056=4(9*/,:,,40*/,;;0-
5(9+0**/0=)0(5*6504(5+.69(**0.
+LWHY[TLU[VM0U[LYUHS4LKPJPUL:LJ[PVUVM)PVJOLTPZ[Y`<UP]LYZP[`VM7LY\NPHHUK
0UZ[P[\[LVM(UH[VT`HUK*LSS)PVSVN`*H[OVSPJ<UP]LYZP[`9VTL
;4;PZHUL\YV[V_PJVYNHU[PUJVTWV\UK[OH[PUK\JLZUL\YVKLNLULYH[PVUWHY[PJ\SHYS`PU
OPWWVJHTW\Z.LSVZVL[HS;YLH[TLU[VM\UKPMMLYLU[PH[LK7*JLSSZ^P[O;4;
JH\ZLZK`ZYLN\SH[PVUVMPU[YHJLSS\SHY*HHUKHWVW[V[PJJLSSKLH[O4PZP[PL[HS"
7PHJLU[PUPL[HS;OLZHTLL_WLYPTLU[HSTVKLSOHZILLU\ZLK[VPU]LZ[PNH[L
VU[OLLMMLJ[VM;4;VU73(HJ[P]P[`7*JLSSZL_WYLZZ]HYPV\Z73([`WLZHUK
HTVUN [OLT *HKLWLUKLU[ NYV\W 00( PZVMVYT .00( PZ THPUS` SVJHSPaLK PU 7*
TP[VJOVUKYPHHUKP[ZYLSLHZLMYVTYH[IYHPUTP[VJOVUKYPH^OLUTLTIYHULWV[LU[PHS
PZYLK\JLK^HZKLTVUZ[YH[LK4HJJOPVUPL[HSZ\NNLZ[PUNP[ZPU]VS]LTLU[PU
UL\YVUHSJLSSKHTHNL.VYHJJPL[HS0U[OLZLZLYPLZVML_WLYPTLU[ZJLSSZ^LYL
[YLH[LK ^P[O ;4; ¶ 4 MVY VY OV\YZ 73( HJ[P]P[` ^HZ HZZH`LK PU
7*S`ZH[LZHUKPUJ\S[\YLTLKP\T\ZPUNHÅ\VYVNLUPJZ\IZ[YH[L7,+I`HUL^
KL]LSVWLKWYVJLK\YL;OL[YLH[TLU[^P[O;4;JH\ZLKZPNUPÄJHU[KLJYLHZLVMJLSS\SHY
73( HJ[P]P[` ^OPJO ^HZ KVZL HUK [PTL KLWLUKLU[ HUK P[ ^HZ HJJVTWHUPLK I` H
WHYHSSLSPUJYLHZLVMLUa`TLHJ[P]P[`PU[OLJ\S[\YLTLKPH^P[OV\[ZPNUPÄJHU[JOHUNLZVM
3+/HJ[P]P[`;OLYLSLHZLK73(HJ[P]P[`^HZHSTVZ[JVTWSL[LS`ISVJRLKI`HZWLJPÄJ
PUOPIP[VYVM.00(*OPYPJVaaPL[HS0[JHUILJVUJS\KLK[OH[.00(WHY[PJPWH[LZ[V
[OL TVSLJ\SHY L]LU[Z PU]VS]LK PU UL\YVUHS JLSS KLH[O PUK\JLK I` ;4; PU[V_PJH[PVU
7HY[PJ\SHYS`[OLYLSLHZLKLUa`TLTH`JH\ZL[OLWYVWHNH[PVUVM[OL[V_PJLMMLJ[VM;4;
[V[OLV[OLYJLSSZVMIYHPU[PZZ\LJVU[YPI\[PUN[V[OLHNNYH]H[PVUVMUL\YVKLNLULYH[PVU
*OPYPJVaaP,L[HS15L\YVJOLT
.LSVZV4*L[HS5L\YVJOLT0U[
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NEU 2
Neurochimica
7`Y\]H[LHKTPUPZ[YH[PVUWYL]LU[ZJVNUP[P]LKLÄJP[ZPUH;N
HUPTHSTVKLSVM(SaOLPTLY»ZKPZLHZL
0:670,30:(:,5:03:;,-(56
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007VZ[LY:LZZPVU
NEU 3
Neurochimica
4PJYVHYYH`HUHS`ZPZVMO\THUUL\YVISHZ[VTHJLSSZL_WVZLK
[VPYVUơHT`SVPKVY[OLơPYVUJVTWSL_
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007VZ[LY:LZZPVU
NEU 4
Neurochimica
73.(57ZMVY,Ua`TL9LWSHJLTLU[;OLYHW`VM
3`ZVZVTHS:[VYHNL+PZLHZLZ
763*/0(;(5*050);6:0.:,=,9050.44(.050(
<9)(5,3303)69;6;)+63*,;;(+=(5+,3304(-6950-
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+LW[VM7OHYTHJL\[PJHS:JPLUJLZ+LW[VM)PVTLKPJHS:JPLUJLZ<UP]LYZP[`VM
4VKLUHHUK9LNNPV,TPSPH0[HS`
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.HYVMVSV;YPLZ[L0[HS`
¸4H\YV)HZJOPYV[[V¹0UZ[P[\[LMVY9HYL+PZLHZLZ)09+-V\UKH[PVU=PJLUaH0[HS`
3`ZVZVTHS Z[VYHNL KPZLHZLZ 3:+ HYL H NYV\W VM PUOLYP[LK KPZVYKLYZ JH\ZLK I`
[OL KLÄJPLUJ` VM S`ZVZVTHS LUa`TLZ ;OLYL HYL UV J\YLZ MVY 3:+ HUK [YLH[TLU[
PZ TVZ[S` Z`TW[VTH[PJ ,Ua`TL 9LWSHJLTLU[ ;OLYHW` ,9; PZ ZHML HUK LMMLJ[P]L
I\[PULMÄJPLU[PU[OL[YLH[TLU[VMUL\YVSVNPJHSTHUPMLZ[H[PVUHZL_VNLUV\ZLUa`TLZ
JHUUV[JYVZZ)))7VS`TLYPJUHUVWHY[PJSLZ57ZYLWYLZLU[VULVM[OLTVZ[PUUV]H[P]L
UVUPU]HZP]LHWWYVHJOLZMVYKY\NZKLSP]LY`0UWHY[PJ\SHY[OL\ZLVMUHUVJHYYPLYZMVY
THJYVTVSLJ\SL KLSP]LY` TH` HSSV^ [V V]LYJVTL ZVTL SPTP[Z VM [OL THJYVTVSLJ\SL
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TLTIYHUL IHYYPLYZ PL ))) +\L [V [OLPY IPVKLNYHKHIPSP[` HUK IPVJVTWH[PIPSP[`
WVS`SHJ[PKLJVNS`JVSPKL 73.( 57Z HYL VUL VM [OL TVZ[ ^PKLS` HWWSPLK Z`Z[LTZ
MVYKY\NKLSP]LY`/LYL^LL_WSVYLK[OL\ZLVMHJPKHSWOHNS\JVZPKHZL.((SVHKLK
73.(57ZPU7VTWL+PZLHZL7+,9;7+PZH3:+JOHYHJ[LYPaLKI`[OLHJJ\T\SH[PVU
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MHY7+[YLH[TLU[JVUZPZ[LKVM[OLHKTPUPZ[YH[PVUVM4`Va`TLŽHYLJVTIPUHU[O\THU
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007VZ[LY:LZZPVU
NEU 5
Neurochimica
4HZZZWLJ[YVTL[Y`IHZLKTL[HIVUVTPJZVM[OL
JLYLIYVZWPUHSÅ\PKPU4\S[PWSL:JSLYVZPZWH[PLU[Z
70,9(.6:;056+(40(5(+0060(4(90(96::0*3(<+0(A<**/,330409*6
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:(**/,;;(7(636<9)(50(5+9,(+,3)6**0670,96
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JVUKP[PVU0UHYLJLU[^VYR^LYLWVY[LKHU3*4:IHZLKSPWPKVTPJZHWWSPJH[PVUPU
4: YL]LHSPUN HS[LYLK ZLY\T WOVZWOVJVSPULZ HUK S`ZVWOVZWOVJVSPULZ SL]LSZ PU 4:
Z\IQLJ[Z KLTVUZ[YH[PUN H TL[HIVSPJ PTIHSHUJL HS[OV\NO [OL ZHTWSL PZ [HRLU MHY
MYVT[OLZP[LVMPUÅHTTH[PVU/LYL^LHPTLK[VPKLU[PM`HTL[HIVSPJHS[LYH[PVUPU[OL
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IHZLKTL[HIVUVTPJZWSH[MVYT0UWHY[PJ\SHY^LMVJ\ZLKV\YH[[LU[PVUVUPU]LZ[PNH[PUN
*:-WOVZWOVJVSPULZWH[[LYUZI`4(3+0;6-4:[LJOUVSVN`3PWPKVTPJZKH[H^LYL
JVTIPULK[VHTPUVHJPKZHUKJHYUP[PULZSL]LSZPU*:-PUHWYLSPTPUHY`JYVZZZLJ[PVUHS
PU]LZ[PNH[PVU VM 4: HMMLJ[LK WH[PLU[Z ]Z WH[PLU[Z ^P[O V[OLY UL\YVSVNPJHS KPZLHZLZ
-PUHSS`H[LU[H[P]LVMJVYYLSH[PVUIL[^LLUL_WLYPTLU[HSYLZ\S[ZHUKJSPUPJHSKH[H^HZ
THKLZOV^PUNWYLSPTPUHY`L]PKLUJLMVY[OLKPZJV]LY`VMJOHYHJ[LYPZ[PJ*:-TL[HIVSPJ
¸ÄUNLYWYPU[WH[[LYU¹PU4:KPZLHZL
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007VZ[LY:LZZPVU
¶
NEU 6
Neurochimica
0UJYLHZLVMOPWWVJHTWHSHS\TPU\TPZSPURLK[V[OL
WYVNYLZZPVUVMHT`SVPKWH[OVSVN`PUH[YPWSL[YHUZNLUPJ
TV\ZLTVKLSVM(SaOLPTLY»ZKPZLHZL
*0(=(9+,330+64,50*6*65:(3=6(+(.(;;(=(3,5;05(
96::0*3(<+0(369,+(5(9,55(:(**/,;;(7(636
+00306*(9405,:,5:0:;,-(56
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,_WLYPTLU[HSHUK*SPUPJHS:JPLUJL+LWHY[TLU[¸.K»(UU\UaPV¹<UP]LYZP[`
*OPL[P7LZJHYH"-\UJ[PVUHS.LUVTPJ<UP[*[YVM,_JLSSLUJLVU(NPUN*OPL[P0[HS`"
4VS5L\YVS+LW[VM5L\YVZJPHUK0THNPUN*OPL[P7LZJHYH0[HS`"
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JVUJLU[YH[PVUZZOV^LKHZPNUPÄJHU[WVZP[P]LJVYYLSH[PVU^P[O[OLPUJYLHZLKSL]LSZVM
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NLULZRUV^U[VILPU]VS]LKPUIYHPU(SPUÅ\_Z\JOHZ[OL[YHUZMLYYPUYLJLW[VY;-9
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SL]LSZ PUJYLHZLK ^P[O HNPUN TH[JOPUN [OL HNLYLSH[LK PUJYLHZL PU OPWWVJHTWHS (S
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NEU 7
Neurochimica
+PHIL[LZPUK\JLKT`LSPUHIUVYTHSP[PLZHYLHZZVJPH[LK^P[O
HUHS[LYLKSPWPKWH[[LYU!WYV[LJ[P]LLMMLJ[ZVM3?9HJ[P]H[PVU
40;9650*6*,94,5(;0.(0(())0(;0-,+,90*6
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ILJH\ZL P[Z TVSLJ\SHY TLJOHUPZT PZ Z[PSS VIZJ\YL ;OPZ WH[OVSVN` PUK\JLZ
UL\YVSVNPJHS JVTWSPJH[PVUZ VU IV[O ULY]L M\UJ[PVU HUK Z[Y\J[\YL Z\JO HZ T`LSPU
4`LSPUPZHIPVSVNPJHSTLTIYHULJOHYHJ[LYPaLKI`OPNOSPWPKJVU[LU[HUKJVU[YPI\[LZ
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:[LYVS9LN\SH[VY`,SLTLU[)PUKPUN-HJ[VYJ:9,)-JHNLULPU]VS]LKPUMH[[`HJPK
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NEU 8
Neurochimica
+PZ[PUJ[TVK\SH[PVUVMLUKVJHUUHIPUVPKZ\WVURHPUPJHJPK
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NEU 9
Neurochimica
(NVUPZ[PUK\JLKKLZLUZP[PaH[PVUVM.79PU
VSPNVKLUKYVJ`[LWYLJ\YZVYJLSSZ!HYVSLPU[OLTH[\YH[PVU[V
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NEU 10
Neurochimica
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NEU 11
Neurochimica
447 PZVMVYTZPUZLY\TVM4\S[PWSL:JSLYVZPZWH[PLU[Z
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NEU 12
Neurochimica
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NEU 13
Neurochimica
+7(.,KL[LJ[PVUVM5L\YVÄSHTLU[3PNO[*OHPU5-
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NEU 14
Neurochimica
TPJYV95(TLKPH[LKS`ZVZVTHSLUa`TLZKV^UYLN\SH[PVU
PUJLSSVMWLYPWOLYHSZ`Z[LTMYVT(SaOLPTLY»ZKPZLHZL
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693(**/06(3+6
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+PWHY[PTLU[VKP4LKPJPUHKLP:PZ[LTP<UP]LYZP[n¸;VY=LYNH[H¹9VTH5L\YVWZPJVSVNPH*SPUPJH
6ZWLKHSL:.PV]HUUP)H[[PZ[H-VSPNUV.L)P:H9LZLHYJO-V\UKH[PVU7LY\NPH
)HJRNYV\UK! (SaOLPTLY»Z KPZLHZL (+ [OL TVZ[ JVTTVU MVYT VM KLTLU[PH PU [OL
LSKLYS`PZJOHYHJ[LYPaLKI`UL\YVÄIYPSSHY`[HUNSLZL_[YHJLSS\SHYHT`SVPKơWSHX\LZHUK
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JYVZZYVHKPU^OPJOL[PVSVNPJHSMHJ[VYZPU(+WH[OVNLULZPZJVU]LYNL;VIL[[LY\UKLYZ[HUK
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NEU 15
Neurochimica
/73*(::(@6-46<:,/@+96?@05+63,6
4,;/@3;9(5:-,9(:,/064;059,;05(3
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3TL[OPVUPUL :(4 HUK 5HJL[`S ZLYV[VUPU 5(: HZ [OL Z\IZ[YH[LZ ;OPZ WYV[LPU
PZ WYLKVTPUHU[S` L_WYLZZLK PU [OL ]LY[LIYH[L WPULHS NSHUK ^OLYL P[ WYVK\JLZ
TLSH[VUPUMYVT5(:I`[OLHJ[P]H[PVUVM[OLYH[LSPTP[PUNLUa`TLHY`SHSR`SHTPUL5
HJL[`S[YHUZMLYHZL((5(;^OPJO\UKLYNVLZ[OLJPYJHKPHUYLN\SH[PVU)LZPKL[OLTHPU
WPULHSVYPNPUVMTLSH[VUPU[OLYLPZHSZVL]PKLUJLJVUJLYUPUNP[ZZ`U[OLZPZPU[OLYVKLU[
HUKO\THUYL[PUH6U[OLV[OLYOHUK[OLTLHZ\YLTLU[VMYL[PUHS/064;HJ[P]P[`PZ
JYP[PJHS ZPUJL [OL LUa`TL PZ H SV^HI\UKHU[ WYV[LPU PU [OPZ [PZZ\L (Z ^LSS /064;
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OHZILLUTHWWLKPUZPKLHOPNOMYLX\LUJ`TLPV[PJYLJVTIPUH[PVUJOYVTVZVTLZP[L
[OLWZL\KVH\[VZVTHSYLNPVU7(9HZOVY[YLNPVUVMOVTVSVN`IL[^LLU[OL?HUK@
JOYVTVZVTLZ
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^LYLPUJ\IH[LK^P[O›45(:›4 /:(4¶*PTTVS4LSH[VUPU
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Z\ZWLUKLK PU TVIPSL WOHZL HUK PUQLJ[LK PU[V [OL JOYVTH[VNYHWO MVY [OL PZVJYH[PJ
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NEU 16
Neurochimica
973*<=7+(4,(:<9,4,5;6-;/,
(5;0+,79,::(5;;9(A6+65,(5+0;:4(169
4,;()630;,T*7705/<4(5:,9<4
7(3,.63),;;03*90:;6-(9644(:(3(0A(56;;0:7(**0(9+0
)),330:3<*/050-4(<9043<*(**/050(.0(55(**050.
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TLJOHUPZTVMHJ[PVUVM;9AJVUZPZ[ZPUP[ZHU[HNVUPZTVU/;(*YLJLW[VYZ[OLKY\N
ILPUNH[[OLZHTL[PTLHSV^HMÄUP[`ISVJRLYVM[OL/;YL\W[HRLZP[L:,9;;9A
OHZILLUMV\UK[VPU[LYHJ[HSZV^P[OOPZ[HTPULHUKƠHKYLULYNPJYLJLW[VYZ\I[`WLZ
HSILP[[OLKY\NWOHYTHJVSVN`OHZUV[ILLUM\SS`LS\JPKH[LKLZWLJPHSS`P[ZLMMLJ[ZVU
ZSLLW M\UJ[PVU ,]LU PM ;9A PZ H ^LSS[VSLYH[LK WZ`JOV[YVWPJ HNLU[ [OL TVUP[VYPUN
VM[OPZJVTWV\UKHUKTL[HJOSVYVWOLU`SWPWLYHaPULT*77TL[HIVSP[LPU[YLH[LK
WH[PLU[Z JHU IL YLSL]HU[ MVY KVZL HKQ\Z[TLU[ [OLYHW` WLYZVUHSPZH[PVU LZWLJPHSS`
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SPX\PKWYVJLK\YL\ZPUNUL_HULHUKI\ZWPYVULHZ[OLPU[LYUHSZ[HUKHYK0:"ZWLJPÄJP[`
HUKHJJ\YHJ`HYLTVUP[VYLKI`<=7+(KL[LJ[PVU"MVYWLHRPU[LNYH[PVUHUKHUHS`ZPZ
[OL7+(PZZL[H[[OL;9AHUKT*77ƪTH_ $UTHUKI\ZWPYVULƪTH_ $UT
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YLJV]LYPLZ SPULHYP[` Y JVLMÄJPLU[Z % HUK PU[YHKH`PU[LYKH` YLSH[P]L
Z[HUKHYKKL]PH[PVU9:+^P[OPU;9AHUKT*77368]HS\LZHJJV\U[MVYI`
›N 3 HUK ›N 3 YLZWLJ[P]LS` 0U HKKP[PVU [OL WYVWVZLK TL[OVK ZOV^Z UV
ZPNUPÄJHU[LS\[PVUPU[LYMLYLUJLZ^P[OHZLYPLZVMWZ`JOVHJ[P]LHU[PKLWYLZZHU[KY\NZ
;OL\ZLVM7+(ZWLJ[YHSSPIYHY`HKKP[PVUHSS`TVUP[VYZWV[LU[PHSPU[LYMLYLUJLZTHRPUN
[OL WYVJLK\YL WHY[PJ\SHYS` \ZLM\S MVY ;9A HUK T*77 TLHZ\YLTLU[ PU ZLY\T VM
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007VZ[LY:LZZPVU
NEU 17
Neurochimica
-HZ[VULZ[LWSPX\PKJOYVTH[VNYHWO`KL[LYTPUH[PVUVM
W\YPULJVTWV\UKZPUISVVK^P[OWOV[VKPVKLHYYH`KL[LJ[VY
.<,304(90(*65*,;;(:(3,40.0<:,77,
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<UP]LYZP[nKLNSP:[\KPKP7HSLYTV
<YPJ HJPK <( H WVS`LKYPJ M\UJ[PVUHS JVTWV\UK PZ H Z[YVUN UH[\YHS HU[PV_PKHU[ PU
O\THUZ^OLYLP[PZ[OLLUKWYVK\J[VMW\YPULTL[HIVSPZT;OLYH[PVUHSWYVJLZZMVY[OPZ
Z[\K`^HZ[OLJSPUPJHSPU[LYLZ[VM[OLTL[HIVSP[LZHZTHYRLYZMVYLULYN`KPZ[\YIHUJL
PUPZJOLTPHO`WV_PHHU[PV_PKHU[JHWHJP[`HUKKPZLHZLHJ[P]P[`PU]P]V<UMVY[\UH[LS`
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WYV]PKLZ H OPNOLY ZLUZP[P]P[` ^P[O H JSLHU JOYVTH[VNYHT ;OL YLWVY[LK YLMLYLUJL
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VMH\[OLU[PJZ[HUKHYKZHUKI`ZWPRPUNZ[HUKHYKZVS\[PVUZ[V\S[YHÄS[YH[LKWSHZTH;OPZ
/73*7+(Z`Z[LTPZ]LY`\ZLM\S[VVSMVY[OLHZZH`VM<(HUKW\YPULKLYP]H[P]LZIV[O
PUYLZLHYJOHZZH`ZHUKPUJSPUPJHSSHIVYH[VYPLZ
007VZ[LY:LZZPVU
NEU 18
Neurochimica
:,9<4:,96;6505(5+4,3(;6505057(;0,5;:>0;/
466+(5?0,;@(5+:3,,7+0:69+,9:;9,(;,+>0;/
;/,(5;0+,79,::(5;;9(A6+65,
*90:;6-(964),;;037(3,.634(:(3(0A(56;;0:7(**0(9+0
)),330:3<*/050-4(<9043<*(**/050(.0(55(**050.
+LWHY[TLU[VM7Z`JOPH[Y`5L\YVIPVSVN`7OHYTHJVSVN`HUK)PV[LJOUVSVN`<UP]LYZP[`VM7PZH
:SLLW KPZVYKLYZ OH]L ]LY` OPNO PUJPKLUJL PU WZ`JOPH[YPJ KPZLHZLZ ILPUN OV^L]LY
VM[LUV]LYSVVRLKPU[OLJSPUPJHSWYHJ[PJL(S[LYH[PVUZPU[OLK\YH[PVUWHJLHUKX\HSP[`
VMZSLLWHJX\PYLKPHNUVZ[PJHUKWYVNUVZ[PJZPNUPÄJHUJLPUYLSH[PVU[V[OLWZ`JOPH[YPJ
KPZVYKLYHZZVJPH[LK*VYVSSHY`[V[OH[PZ[OLUV[L^VY[O`YLSL]HUJLVMZSLLWKPZVYKLYZ»
THUHNLTLU[LP[OLYPUYLK\JPUN[OLZ\MMLYPUNVMWH[PLU[ZVYPUPTWYV]PUN[OLPYX\HSP[`VM
SPMLI`HMMLJ[PUN[OLJV\YZLVM[OLYLSH[LKWZ`JOPH[YPJKPZLHZLHZ^LSSHZI`WYL]LU[PUN
P[Z YLSHWZL 5L]LY[OLSLZZ HU PTWLKPTLU[ [V [OL NVVK [YLH[TLU[ VM ZSLLW KPZVYKLYZ
JVUULJ[LK[VWZ`JOPH[YPJKPZLHZLZPZYLWYLZLU[LKI`[OLT\S[PWSL]HYPHISLZPU]VS]LK
(PT VM [OPZ YLZLHYJO ^HZ [V L]HS\H[L [OL JSPUPJHSIPVJOLTPJHS LMMLJ[Z VM [YHaVKVUL
HT\S[PM\UJ[PVUHSHU[PKLWYLZZHU[KY\N^OPJOHSZVKPZWSH`ZNVVKO`WUV[PJWYVWLY[PLZ
![OLZLYV[VULYNPJHUKTLSH[VULYNPJTL[HIVSPZT^HZKL[LYTPULKPUWH[PLU[Z^P[O
TVVKHU_PL[`KPZVYKLYZ\UKLY[YHaVKVUL[OLYHW`MVYZSLLWKPZ[\YIHUJLZ[OYV\NO[OL
TLHZ\YLTLU[VMZLYV[VUPU/;HUKTLSH[VUPUZLY\TSL]LSZILMVYL;HUKHM[LY;H
[YLH[TLU[^P[OSV^KY\NKVZLZTNKH`MVY[OLO`WUV[PJHJ[PVU:LY\TVI[HPULK
MYVTWLYPWOLYHS]LUV\ZISVVK^P[OKYH^UIL[^LLU HT^HZL_[YHJ[LK^P[O
L[OHUVS/;HUKKPJOSVYVTL[OHULTLSH[VUPUKYPLK\UKLYOPNO]HJ\\THUKZ[VYLK
H[ ‡* \U[PS HZZH` (U PZVJYH[PJ YL]LYZLKWOHZL SPX\PK JOYVTH[VNYHWOPJ 973*
TL[OVK^HZHWWSPLK\ZPUN*WYLHUKHUHS`[PJHSJVS\TUZLX\PSPIYH[LKPUHTVIPSL
WOHZL JVTWVZLK I` H TP_[\YL TL[OHUVSWOVZWOH[L I\MMLY W/ ! ]] H[ [OL
JVUZ[HU[ ÅV^ YH[L VM TS TPU (UHS`[L WLHRZ ^LYL YL]LHSLK I` TLHUZ VM H 7,+
LSLJ[YVJOLTPJHSKL[LJ[VYHUKHTWLYVTL[YPJTLHZ\YLTLU[
7YLSPTPUHY`YLZ\S[ZVI[HPULKPUZL]LUZ\IQLJ[ZZOV^LKHZL_WLJ[LKPU[LYPUKP]PK\HS
]HYPHUJL JVUJLYUPUN ; HUK ; ZLY\T TLSH[VUPU HUK /; ( [YLUK [V^HYKZ [OL
YLK\J[PVU VM IV[O HUHS`[LZ TLHU JVUJLU[YH[PVUZ H[ ; ^HZ VIZLY]LK ULLKPUN
OV^L]LYJVUÄYTH[PVU[OYV\NO[OLL_[LUZPVUVML_HTPULKJHZLZ
:[HOS: *5::WLJ[Y!
007VZ[LY:LZZPVU
7YV[LPUL
796
PRO 1
Proteine
,MMLJ[VM4\J\UH7Y\YPLUZSLH]LZL_[YHJ[VUO\THU
RLYH[PUVJ`[LZ7YV[LVTPJWYVÄSPUN
*69;,3(AA6(3,::063(47(90,3363<*0(:;0*6AA0*3(<+0(
.<,99(5;096),9;6=(3(**/0.0<:,77,
+LWHY[TLU[VM0U[LYUHS4LKPJPUL,UKVJYPUL4L[HIVSPJ:JPLUJLZHUK)PVJOLTPZ[Y`
<UP]LYZP[`VM:PLUH:PLUH0[HS`"
+LWHY[TLU[VM*OLTPZ[Y`<UP]LYZP[`VM:PLUH:PLUH0[HS`"
+LWHY[TLU[VM,]VS\[PVUHY`)PVSVN`<UP]LYZP[`VM-LYYHYH-LYYHYH0[HS`"
+LWHY[TLU[VM-VVKHUK5\[YP[PVU2`\UN/LL<UP]LYZP[`:LV\S:V\[O2VYLH
;OLWSHU[KLYP]LKJVTWV\UKZHYLHUPTWVY[HU[ZV\YJLVMKY\NZMVY]HYPV\ZKPZLHZLZ
;OL TLKPJPUHS WSHU[ 4\J\UH WY\YPLUZ 4W PZ [OL TVZ[ WVW\SHY KY\N PU (`\Y]LKPJ
Z`Z[LT VM TLKPJPUL 0[Z KPMMLYLU[ WYLWHYH[PVUZ MYVT ZLLKZ OH]L ILLU \ZLK HZ
[OLYHWL\[PJ TLKPJPUL MVY ZL]LYHS KPZLHZLZ Z\JO HZ YOL\TH[VPK HY[OYP[PZ KPHIL[LZ
H[OLYVZJSLYVZPZ ULY]V\Z KPZVYKLYZ HUK 7HYRPUZVU KPZLHZL BD 7SHU[Z MYVT 5PNLYPH
ZLLKZ^LYLNYV^UPU)V[HUPJHS.HYKLU<UP]LYZP[`VM:PLUH[OLSLH]LZ^LYLJVSSLJ[LK
KYPLKW\S]LYPaLKHUKL_[YHJ[LK^P[OTL[OHUVS;OLL_[YHJ[Z^LYLL_HTPULKPU]P[YV
MVY [OLPY HU[PV_PKHU[ HJ[P]P[` )LPUN [OL ZRPU VUL VM [OL THPU [HYNL[ VM L_VNLUV\Z
V_PKH[P]LZ[YLZZ<=6ZTVRLL[JBDHUKILPUNZL]LYHSZRPUWH[OVSVNPLZYLSH[LK[V
PUJYLHZLKV_PKH[P]LZ[YLZZBD0U[OPZZ[\K`RLYH[PUVJ`[LZ/H*H;]PHIPSP[`I`TLHUZ
VMKPMMLYLU[S`KPS\[LKZHTWSLZOHZILLUTLHZ\YLKHUKJOHUNLZPUWYV[LPUZL_WYLZZPVU
OH]LILLUL]HS\H[LKHM[LY4WSLH]LZTL[OHUVSPJL_[YHJ[4W3,[YLH[TLU[7YV[LPUZ
MYVTJLSSZS`ZH[LZ^OLYLHUHS`aLKI`:+:7(.,HUK+,7YV[LPUIHUKZHUKZWV[Z
^LYL L]PKLUJLK I` ZPS]LY Z[HPUPUN HUK KP]PKLK PU NYV\WZ IHZLK VU [OL TVSLJ\SHY
^LPNO[ HZ MVSSV^Z! 2+ 2+ HUK 2+ ;OL HUHS`ZPZ
YL]LHSLK[OH[HM[LY4W3,[YLH[TLU[[OLYL^LYL HUKUL^ZWV[ZPUNYV\WZ
HUKYLZWLJ[P]LS`0UHKKP[PVU[OL[YLH[TLU[ZOV^LK[OLSVZZVM HUK
VYPNPUHSPUNYV\WZHUKYLZWLJ[P]LS`6MUV[L[OL4W3,[YLH[TLU[ZPNUPÄJHU[S`
KLJYLHZLK[OLSL]LSZVM/5,WYV[LPUHKK\J[ZLZWLJPHSS`MVYNYV\W;OLZLYLZ\S[Z
Z\NNLZ[[OH[[OL\ZLVM4W3,TPNO[ILOLSWM\SPU[OL[YLH[TLU[VMV_PKH[P]LZ[YLZZ
YLSH[LKZRPUKPZLHZLZ
9LMLYLUJLZ!
4PZYH3L[HS0UKPHU1V\YUHSVM)PVJOLTPZ[Y`HUK)PVWO`ZPJZ
=HSHJJOP.L[HS-,):3L[[
)YPNHU[P-HUK7PJHYKV41,\Y(JHK+LYTH[VS=LULYLVS 007VZ[LY:LZZPVU
PRO 2
Proteine
7YV[LVTPJHUHS`ZPZVMO\THUWSHZTHWYV[LVTLJOHUNLZ
KLYP]LKMYVTPU]P[YVWYV[LVS`[PJHJ[P]P[`VM,JOPZJHYPUH[\Z
]LUVT
*69;,3(AA6(3,::06.<,99(5;096),9;6
/67,65@,2>,9,55(+6A0,
-<93(50,4030(56)0503<*(3,65*05096),9;6
+LWHY[TLU[VM0U[LYUHSTLKPJPUL,UKVJYPUL4L[HIVSPJ:JPLUJLZHUK)PVJOLTPZ[Y`
<UP]LYZP[`VM:PLUH:PLUH0[HS`
+LWHY[TLU[VM4VSLJ\SHY)PVSVN`<UP]LYZP[`VM:PLUH:PLUH0[HS`
0U [OPZ Z[\K` ^L PU]LZ[PNH[LK [OL PU ]P[YV LMMLJ[Z VM ZUHRL ]LUVT WLW[PKHZLZ MYVT
[OL =PWLYPKHL MHTPS` VU O\THU WSHZTH WYV[LPUZ ;V JHYY` V\[ V\Y L_WLYPTLU[Z ^L
\ZLK ,JOPZ JHYPUH[\Z ]LUVT ,= 0[ PZ H JVTWSL_ TP_[\YL VM WYV[LHZLZ ILZPKLZ
WLW[PKLZ[V_PUZHUKWSH[LSL[HNNYLNH[PVUPUOPIP[VYZBD,=JVU[HPUZ[^V^LSSRUV^U
TL[HSSVWYV[LPUHZLZ ^OPJO HYL WYV[OYVTIPU HJ[P]H[VY! LJHYPU HUK JHYPUHJ[P]HZL ;OL
LMMLJ[ZVM,=VU[OLJVHN\SH[PVUWYVJLZZHYLHSZVYLWYLZLU[LKI`[OLIYLHRKV^UVM
[OLÄIYPUVNLUBD0U[OPZ^VYR^LWYLWHYLK[OLWSHZTHMYVTHUOLHS[O`]VS\U[LLYHUK
[OLU[OLWSHZTH^HZKLWSL[LK\ZPUN7YV[LV7YLWŽ)S\L(SI\TPU0N.+LWSL[PVU2P[
+LWSL[LKWSHZTH^HZPUJ\IH[LK^P[O,=VY^P[OZHSPULZVS\[PVUH[TPUH[‡*
+,^HZWLYMVYTLK[VHUHS`aL[OLJVTWSL_TP_[\YLVMLU[PYLWYV[LPUZHUKMYHNTLU[Z
WYVK\JLK I` [OL WYV[LVS`[PJ HJ[P]P[` /\THU WSHZTH ZOV^Z ZPNUPÄJHU[ JOHUNLZ PU
WYV[LPUZPU]VS]LKPUISVVKJVHN\SH[PVUI\[HSZVPUKPMMLYLU[WO`ZPVSVNPJHSTLJOHUPZTZ
^OLU P[ ^HZ [YLH[LK ^P[O [OL ZUHRL ]LUVT ,HJO WYV[LPU ZWV[ ^HZ HUHS`aLK \ZPUN
,;;(54(3+0;6-HUKTHZZÄUNLYWYPU[PUNZLHYJOPUN^HZJHYYPLKV\[\ZPUN4(:*6;
(M[LY WYV[LPU PKLU[PÄJH[PVU ^L Z[\KPLK [OLPY ZPTPSHYP[` \ZPUN )3(:; HUK 7YVZP[L
4VYLV]LY^LPU]LZ[PNH[LK[OLWYV[LHZLZHJ[P]P[PLZ\ZPUN*\[+)HUK4,967:;OL
PKLU[PÄLKWSHZTHWYV[LPUZJHUILSPURLK[VHZWLJPÄJWH[OVSVNPJHSWYVJLZZLZ;OLPY
K`ZM\UJ[PVUJH\ZLKI`WYV[LHZLHJ[P]P[`SLHKZ[VKPZLHZLHZZVJPH[LKWYV[LPU;OLZL
YLZ\S[ZNP]L\ZUL^PUMVYTH[PVUHIV\[O\THUWSHZTHWYV[LVTLWYV[LHZLZHUKZUHRL
]LUVT WVPZVU ;OPZ PUMVYTH[PVU JHU OLSW \Z [V \UKLYZ[HUK [OL YVSL VM [OLZL [OYLL
TVPL[PLZPUTLKPJHSHWWSPJH[PVU
9LMLYLUJLZ!
*HS]L[L11:HUa3.\[PtYYLa14=LUVTZ]LUVTPJZHU[P]LUVTPJZ-,):3L[[ .\LYYHU[P96N\LSP0.)LY[VJJP,7YV[LVTPJHUHS`ZPZVM[OLWH[OVWO`ZPVSVNPJHSWYVJLZZPU]VS]LKPU
[OLHU[PZUHRL]LUVTLMMLJ[VM4\J\UHWY\YPLUZL_[YHJ[7YV[LVTPJZ
007VZ[LY:LZZPVU
PRO 3
Proteine
),;(.3@*6796;,050:,3,*;0=,3@05/0)0;:;/,
796*6(.<3(5;-<5*;065:6-;/964)05!(56=,3
7/@:0636.0*(3(5;0*6(.<3(5;4,*/(50:405
/(,46:;(:0:
+,-030770:=05*,5A6 76AA050*63((*8<(:(30,5;,3(<9(
-9(::6596),9;()(5A(;6(3,::(5+9((9*6=0;6(3,::(5+96
:*(.3065,.06=(5503<*(+,*90:;6-(969(0465+67,5.6=0;;6906
3HIVYH[VY`VM7YV[LPU*OLTPZ[Y`+LWHY[TLU[VM7OHYTHJL\[PJHSHUK
7OHYTHJVSVNPJHS:JPLUJLZHUK
+LWHY[TLU[VM*HYKPHJ;OVYHJPJHUK=HZJ\SHY:JPLUJLZ
<UP]LYZP[`VM7HK\H7HK\H0[HS`
0UZ[P[\[LVM)PVJOLTPZ[Y`HUK*SPUPJHS)PVJOLTPZ[Y`HUK
0UZ[P[\[LVM0U[LYUHS4LKPJPUL/HLTVZ[HZPZ9LZLHYJO*LU[YL*H[OVSPJ<UP]LYZP[`
:JOVVSVM4LKPJPUL9VTL0[HS`
)HJRNYV\UK! ơNS`JVWYV[LPU 0 ơ.W0 PZ HU HI\UKHU[ WSHZTH WYV[LPU TN
TS PU]VS]LK PU [OL WH[OVNLULZPZ VM [OL HU[PWOVZWOVSPWPK Z`UKYVTL I\[ ^OVZL
WO`ZPVSVNPJHSM\UJ[PVUPZZ[PSSLS\ZP]L4L[OVKZ!;OLLMMLJ[VMơ.W0VU[OLWYVJVHN\SHU[
ÄIYPU NLULYH[PVU HUK WSH[LSL[ HNNYLNH[PVU HUK HU[PJVHN\SHU[ WYV[LPU * HJ[P]H[PVU
M\UJ[PVUZ VM [OYVTIPU ^HZ PU]LZ[PNH[LK I` LUa`TL PUOPIP[PVU HZZH`Z Z\YMHJL
WSHZTVUYLZVUHUJL:79[\YIPKPTL[YPJHUKPTT\UVJ`Å\VYPTL[YPJHUHS`ZPZ9LZ\S[Z!
ơ.W0KVLZUV[HMMLJ[[OLHMÄUP[`VM[OLLUa`TLMVYZVTLHJ[P]LZP[LPUOPIP[VYZSPRL
WHTPUVILUaHTPKPUL HUK [OL 5[LYTPUHS KVTHPU VM OPY\KPU +PYLJ[ HUHS`ZPZ VM
ơ.W0[OYVTIPU PU[LYHJ[PVU I` :79 `PLSKZ H 2K$U4 +PZWSHJLTLU[ L_WLYPTLU[Z
JHYYPLK V\[ ^P[O ZWLJPÄJ IPUKLYZ H[ [OYVTIPU L_VZP[L OPY\NLU HUK /+ HW[HTLY
VYL_VZP[LÄIYPUVNLUƢ»WLW[PKLHUK/+HUK^P[O[OYVTIPUT\[HU[Z(YN(SH
HUK (YN(SH OH]PUN VUL VM [OL [^V L_VZP[LZ ZLSLJ[P]LS` JVTWYVTPZLK PUKPJH[L
[OH[IV[OL_VZP[LZVM[OLWYV[LHZLHYLPU]VS]LKPUơ.W0IPUKPUNơ.W0ZPNUPÄJHU[S`
WYVSVUNZ[OLJSV[[PUN[PTLPUÄIYPUNLULYH[PVUHZZH`ZHUKOPUKLYZWSH[LSL[ZHNNYLNH[PVU
0*$ƫ4I`PUOPIP[PUNJSLH]HNLVM7(9VUPU[HJ[WSH[LSL[Z0*$ƫ4HUKPU
ZVS\[PVUơ.W0KVLZUV[HS[LY[OLHIPSP[`VM[OYVTIPU[VNLULYH[L[OLHU[PJVHN\SHU[
WYV[LPU * ^P[O VY ^P[OV\[ [OYVTIVTVK\SPU HKKLK *VUJS\ZPVUZ! ơ.W0 IPUKZ [V
[OYVTIPUH[IV[OL_VZP[LZ^OPSLSLH]PUN[OLWYV[LHZLHJ[P]LZP[LHJJLZZPISLơ.W0
PUOPIP[Z [OL RL` WYVJVHN\SHU[ WYVWLY[PLZ VM [OYVTIPU ^P[OV\[ HMMLJ[PUN P[Z \UPX\L
HU[PJVHN\SHU[ M\UJ[PVU ;OLZL YLZ\S[Z HYL \UWYLJLKLU[LK HUK WH]L [OL ^H` [V [OL
KPZJV]LY`VMHUV]LSWO`ZPVSVNPJHSHU[PJVHN\SHU[WH[O^H`PUOHLTVZ[HZPZ
007VZ[LY:LZZPVU
PRO 4
Proteine
6Z[LVWVU[PUHUK:`Z[LTPJ:JSLYVZPZ!WYLSPTPUHY`YLZ\S[ZVM
6Z[LVWVU[PUZLJYL[PVUI`ZJSLYVKLYTPJÄIYVISHZ[Z
3(5+0.*69(336*=(55650+.069+(5653,65*0509
+LWHY[TLU[VM0U[LYUHS4LKPJPUL,UKVJYPUL4L[HIVSPJ:JPLUJLZHUK)PVJOLTPZ[Y`
<UP]LYZP[`VM:PLUH0[HS`
6Z[LVWVU[PU 675 PZ H NS`JVWYV[LPU L_WYLZZLK I` H SHYNL ]HYPL[` VM [PZZ\LZ HUK
WSH`Z HU PTWVY[HU[ YVSL PU L_[YHJLSS\SHY TH[YP_ YLTVKLSPUN 9LJLU[ Z[\KPLZ YLWVY[LK
OPNOSL]LSZVM675PUZLY\THUKKLYTPZVMWH[PLU[ZHMMLJ[LKI`:`Z[LTPJ:JSLYVZPZ
::JHUKZ\NNLZ[[OH[675JV\SKWSH`HUPTWVY[HU[YVSLPU[OLKL]LSVWTLU[VM[OL
KPZLHZL ;OL HPT VM [OPZ Z[\K` PZ [V L]HS\H[L [OL JVU[YPI\[PVU VM ÄIYVISHZ[ PU [OL
L_WYLZZPVU HUK ZLJYL[PVU VM 675 PU ::J -VY [OPZ Z[\K` ^L PZVSH[LK ZJSLYVKLYTPJ
ÄIYVISHZ[Z MYVT SLZPVUHS ::J HUK UVU SLZPVUHS 5( ZRPU VM ZJSLYVKLYTPJ WH[PLU[Z
*LSSZ^LYLJ\S[\YLKPUHZLY\TMYLLTLKP\THUK[YLH[LK^P[O)SLVT`JPU)3,6T<
TS HUK LUKV[OLSPU ,; U4 (M[LY O J\S[\YL TLKPH Z\WLYUH[HU[Z ^LYL
JLU[YPM\NLK HUK JVSSLJ[LK MVY HUHS`ZPZ 675 ZLJYL[PVU ^HZ L]HS\H[LK ^P[O /\THU
6Z[LVWVU[PU,30:(RP[()*(4*HTIYPNL;OPZHZZH`LTWSV`ZHUHU[PIVK`ZWLJPÄJ
MVYO\THU6Z[LVWVU[PUJVH[LKVUH ^LSSWSH[L7YLSPTPUHY`YLZ\S[ZZOV^LKZPTPSHY
675SL]LSZVU\U[YLH[LKÄIYVISHZ[Z:[H[PZ[PJHSS`YLSL]HU[KPMMLYLUJLZPUJ\S[\YLTLKPH
675 JVUJLU[YH[PVU ^LYL MV\UK ^OLU [OL JLSSZ ^LYL [YLH[LK ^P[O )SLVT`JPU TVYL
[OHU,;;OLZLWYLSPTPUHY`YLZ\S[ZZ\NNLZ[[OH[OPNO675SL]LSZHSYLHK`V\[SPULKPU
YLJLU[Z[\KPLZJV\SKILYLSH[LK^P[O[OLWH[OVNLULZPZVMZJSLYVKLYTHHUKHSZV[OLPU
]P[YVTVKLSVMPUK\JLKKLYTHSÄIYVZPZI`ISLVT`JPUIL[[LYZPT\SH[L[OLILOH]PVYVM
ZJSLYVKLYTPJÄIYVISHZ[ZPU]P]V
>\4L[HS6Z[LVWVU[PUPUZ`Z[LTPJZJSLYVZPZHUKP[ZYVSLPUKLYTHSÄIYVZPZ10U]LZ[+LYTH[VS
1\U"!
2L`^VYKZ!:`Z[LTPJ:JSLYVZPZ6Z[LVWVU[PU+LYTHS-PIYVZPZ)SLVT`JPU,UKV[OLSPU
007VZ[LY:LZZPVU
PRO 5
Proteine
(U[PTPJYVIPHSWLW[PKLZKLYP]LKMYVT[YPWZPUKPNLZ[PVUVM
IV]PULSHJ[VMLYYPU
(55(90;(30AA0=,9650*(*(950*,330-9(5*,:*/05050*63(
70,9(.6:;056+(40(5((5+(5;6506+0.0<306
+LW[VM)PV[LJOUVSVNPJHSHUK(WWSPLK*SPUPJHS:JPLUJLZ<UP]LYZP[`VM3»(X\PSH
+LW[VM,_WLYPTLU[HSHUK*SPUPJHS:JPLUJLZ<UP]LYZP[`VM*OPL[P7LZJHYH0[HS`
)V]PULSHJ[VMLYYPUI3MPZHUPYVUIPUKPUNNS`JVWYV[LPUJVUZPZ[PUNVM HTPUVHJPKZ
MV\UKPUKPMMLYLU[IPVSVNPJHSÅ\PKZVMTHTTHSZP[L_LY[ZZL]LYHSIPVSVNPJHSM\UJ[PVUZ
HTVUN^OPJOHU[PTPJYVIPHSHU[P]PYHSHUKHU[PPUÅHTTH[VY`HJ[P]P[PLZ3HJ[VMLYYPUPZH
KVTPUHU[^OL`WYV[LPUPUTPSR[OYV\NOV\[SHJ[H[PVUHUKPZHISL[VIPUK[^VMLYYPJPVUZ
BD >OLU [OPZ WYV[LPU PZ KPNLZ[LK H[ HJPKPJ W/ I` WYV[LHZLZ P[ `PLSKZ OPNOS` HJ[P]L
HU[PTPJYVIPHS WLW[PKLZ [OH[ PUOPIP[ [OL NYV^[O VM H U\TILY VM .YHTULNH[P]L HUK
.YHTWVZP[P]LIHJ[LYPHPUJS\KPUNZ[YHPUZ[OH[^LYLYLZPZ[HU[[VUH[P]L3MBD
;OLHPTVM[OPZZ[\K`^HZ[VPU]LZ[PNH[LPM[OL[YPWZPUO`KYVS`ZPZVM^OVSLI3MHZ^LSS
HZ[OLPYVUKLWSL[LKWYV[LPUHWVI3MYLSLHZLZV[OLYUV[`L[^LSSRUV^UHU[PTPJYVIPHS
WLW[PKLZ(47Z(M[LYO`KYVS`ZPZ[OLWLW[PKLWVVSZ^HZKP]PKLKPU[^VMYHJ[PVUZI`
*LU[YPJVUÄS[LYZJ\[VMMR+H;OLMYHJ[PVUZ[VNL[OLY^P[O[OLWHYLU[WYV[LPUZ^LYL
[LZ[LKMVY!P[OLHU[PTPJYVIPHSHJ[P]P[`]LYZ\ZZL]LYHS,JVSPHUK7HLY\NPUVZHIHJ[LYPHS
Z[YHPUZ" PP [OL WLY[\YIPUN LMMLJ[ VU IHJ[LYPHS I` ;,4 TPJYVZJVW` HUK PPP [OL HIPSP[`
PU PUK\JPUN Å\VYLZJLU[ WYVILZ YLSLHZL VM MYVT Z`U[OL[PJ ]LZPJSLZ SPWVZVTLZ ;OL
WLW[PKLWVVS4>#R+HKLYP]LKMYVTHWVI3-O`KYVS`ZPZ^OPJOYLZ\S[LK[OLTVYL
HJ[P]LZHTWSLZLWHYH[LKI`/73*HUKHUHS`ZLKI`THZZZWLJ[YVZJVW`YL]LHSLK[OL
WYLZLUJLVMZL]LYHSZTHSSWLW[PKLZ0U[OPZTVTLU[^LMVJ\ZV\YH[[LU[PVUVUH
HTPUVHJPKZWLW[PKL[OH[ZLLTZ[VIL[OLYLZWVUZPISLMVY[OLOPNOHU[PTPJYVIPHSHIPSP[`
VM[OLWVVS
3PaaPL[HS4PUP9L]4LK*OLT ! ;VTP[HL[HS1+HPY`:JP! 007VZ[LY:LZZPVU
PRO 6
Proteine
;OLHSSLYNLU4\ZT!L]PKLUJLVMHNNYLNH[PVU
-,99(90,3,5(*(:(30,4(5<,3((5+(3),9;6:70:50
+PWHY[PTLU[VKP:JPLUaL)PVTLKPJOL)PV[LJUVSVNPJOLL;YHZSHaPVUHSP:)P)P;
<UP]LYZP[nKLNSP:[\KPKP7HYTH
7YV[LPUHNNYLNH[PVUPZYLSL]HU[[VH]HYPL[`VMIPV[LJOUVSVNPJHSWYVJLZZLZ!MYVT[OL
MVYT\SH[PVUVMWYV[LPUKY\NZSPRL]HJJPULZ[V[OLWYVK\J[PVUHUKZ[VYHNLVMOPNOX\HSP[`
WYV[LPUZ/LYL^LWYLZLU[HUPU]LZ[PNH[PVUVU[OLWV[LU[PHSHNNYLNH[PVUVM[OLHSSLYNLU
4\ZTHUKP[ZT\[HU[*:;OLL_WLYPTLU[HSJVUKP[PVUZHSSV^LK[VPKLU[PM`
[OL MVYTH[PVU VM JV]HSLU[ T\S[PTLYZ HZ ^LSS HZ VM ZVS\ISL HNNYLNH[LZ! [OL MVYTLY
PUK\JLKI`[OLYTHSKLUH[\YH[PVUVM4\ZT^OPSL[OLSH[[LYHYLVIZLY]LKPUUVU
KLUH[\YH[PUNJVUKP[PVUZMVYIV[OWYV[LPUZ
/LH[KLUH[\YLKWYV[LPUZ^LYLHUHS`aLKI`:+:7(.,HUKYL]LHSLK[OH[PU[OLJHZL
VM4\ZT*`ZPZPU]VS]LKPUHUPU[LYJOHUNLWYVJLZZ^P[O[OLVUS`KPZ\SÄKL
IYPKNLWYLZLU[PU[OLWYV[LPUHUPU[LYTVSLJ\SHYKPZ\SWOPKLIYPKNLMVYTH[PVUWSH`PUNH
YVSLPUJV]HSLU[HNNYLNH[PVU
+`UHTPJ 3PNO[ :JH[[LYPUN +3: ^HZ \ZLK [V Z[\K` [OL HNNYLNH[PVU Z[H[L VM 4\Z
THUKP[ZT\[HU[PUUVUKLUH[\YH[PUNJVUKP[PVUZ;OL[^VWYV[LPUZHWWLHYLK
PU [^V KPZ[PUJ[ ZPaL WVW\SH[PVUZ! H[ WYV[LPUZ JVUJLU[YH[PVU %$ TNYTS [OL ZTHSSLY
ZPaL MYHJ[PVU ZOV^LK H O`KYVK`UHTPJ KPHTL[LY JVYYLZWVUKPUN [V [OL LZ[PTH[LK
TVUVTLYPJ THZZ ^OPSL [OL SHYNLY ZPaL MYHJ[PVU JVYYLZWVUKLK [V H OPNOLY VYKLY
HNNYLNH[L-VSSV^PUNKPS\[PVUZ\W[VTNYTS[OLPU[LUZP[`VM[OLWLHRZVM[OLUH[P]L
HSSLYNLUYLZ\S[LKL_[YLTLS`TVUVKPZWLYZL^OPSL[OLT\[HU[SV^ZPaLPU[LUZP[`WLHR
ILJHTLWYVNYLZZP]LS`WVS`KPZWLYZLILPUN[OLYLZ\S[VMHTP_[\YLVMTVUVKPHUK
[YPTLYPJWYV[LPUMVYT
0UJVUJS\ZPVU[OLWYVJLZZVMJV]HSLU[HNNYLNH[PVU^HZHJRUV^SLKNLKVUS`HM[LY[OLYTHS
KLUH[\YH[PVUVM[OLUH[P]LWYV[LPU^OPSL[OLLX\PSPIYP\TIL[^LLU[OLTVUVTLYZHUK
HTPUVYHSPX\V[VMZVS\ISLHNNYLNH[LZ^HZHWWYLJPH[LKPUUH[P]LJVUKP[PVUZMVYIV[O
WYV[LPUZ >VY[O JVUZPKLYPUN PZ [OH[! [OL KPS\[PVU VM [OL T\[HU[ ZHTWSL PUJYLTLU[LK
[OLVSPNVTLYZWVW\SH[PVUHUPU[LYLZ[PUNÄUKPUNHZ[YHUZPLU[KPTLYZHYLL_WLJ[LK[V
Z[PT\SH[L[OLHSSLYNLUPJYLHJ[PVU
:9VLMZHUK2+L2Y\PMM ,\Y1)PVJOLT 43PL[HS1)PVS*OLT
007VZ[LY:LZZPVU
PRO 7
Proteine
-\UJ[PVUHSJOHYHJ[LYPaH[PVUI`ZP[LKPYLJ[LKT\[HNLULZPZ
VMO\THUV_PKVZX\HSLULJ`JSHZL6:*L_WYLZZLKPU7PJOPH
WHZ[VYPZ
=063(-9(5*( )(330(56.0(5509<-(9405;(9(4056:03=0(
+0)(9;636.0<:,77,630(96)6::6:0465,;;(
+PWHY[PTLU[VKP:JPLUaHL;LJUVSVNPHKLS-HYTHJV¶<UP]LYZP[nKP;VYPUV
-OVMMTHU3H9VJOL(.7OHYTHYLZLHYJO+PZJV]LY`*OLTPZ[Y`
)HZLS:^P[aLYSHUK
0UZ[LYVSIPVZ`U[OLZPZV_PKVZX\HSLULJ`JSHZL6:*JH[HS`aLZ[OLTVZ[V\[Z[HUKPUN
Z[LW VM [OL WH[O^H`! [OL ZOHWPUN VM [OL [V[HSS` VWLU [YP[LYWLUL PU[LYTLKPH[L
V_PKVZX\HSLUL NLULYH[LK I` [OL TL]HSVUH[L WH[O^H` PU[V SHUVZ[LYVS [OL ÄYZ[
WVSPJ`JSPJWYLJ\YZVYVMJOVSLZ[LYVS9LJLU[S`HUL^PU[LYLZ[MVY[OPZLUa`TLOHZILLU
YHPZLKI`HUPU]LYZLKVJRPUNZ[\K`[OH[PKLU[PÄLZ6:*HZ[OLOPNOLZ[YHURWV[LU[PHS
[HYNL[VM7904(HZTHSSTVSLJ\SLHJ[P]H[PUNT\[HU[WWYV[LPUYLZ[VYPUNP[Z[\TVY
Z\WWYLZZVYM\UJ[PVUBD4VYLV]LYHU[PWYVSPMLYH[P]LHUKHU[PHUNPVNLUPJHJ[P]P[`VM9V
VULVM[OLTVZ[WV[LU[PUOPIP[VYZVM6:*OHZILLUYLJLU[S`ZOV^UBD
7YL]PV\Z T\[HNLULZPZ Z[\KPLZ VU [OL `LHZ[ 6:* HUK VU [OL OVTVSVN\L IHJ[LYPHS
LUa`TL :/* OH]L ZOV^U [OH[ ZVTL HTPUVHJPKPJ YLZPK\LZ UV[ ILSVUNPUN [V [OL
HJ[P]LZP[LHUKSVJHSPaLKPUHUO`KYVMVIPJJOHUULSO`WV[PaLK[VHSSV^[OLHJJLZZ[V
HJ[P]LZP[LHYLLZZLU[PHSMVYTHPU[HPUPUN[OLJH[HS`[PJHJ[P]P[`VM[OLLUa`TLBD;OL
JY`Z[HSZ[Y\J[\YLVMO\THU6:*ZOV^Z[OLWYLZLUJLIV[OVM[OLO`KYVMVIPJJOHUULS
HUKVMHTVYLWVSHYJOHUULSHSSV^PUNHJOHPUVM^H[LYTVSLJ\SLZ[VYLHJO[OL, YLZPK\LH[HJ[P]LZP[L!HWVZZPISL^H`VMYLWYV[VUH[PVUVMJH[HS`[PJ+
0U[OPZZ[\K`^LWYLZLU[[OLLMMLJ[ZVM[OLT\[HNLULZPZVMKPMMLYLU[HTPUVHJPKZVM
O\THU6:*ILSVUNPUN[V[OLHJ[P]LZP[LVY[V[OLO`KYVWOVIPJJOHUULSVY[V[OLWVSHY
VUL6\YYLZ\S[ZJVUÄYT[OLYVSLVM[OLWVSHYJOHUULSZPUJL[OLTVKPÄJH[PVUZVM, HUKVM[OLUL[^VYRVM/IVUKJVUULJ[PUNP[[V2HUK8 JVTWSL[LS`Z\WWYLZZ
[OLJH[HS`[PJHJ[P]P[`VM6:*
.YPU[LY:AL[HS14VS.YHWO4VK ¶
4LKH*L[HS-,):177 :[HLKSLY+L[HS14LK*OLT ò
6SPHYV)VZZVL[HS73V:65,!LKVP!
007VZ[LY:LZZPVU
PRO 8
Proteine
;OLWVS`8WYV[LPUH[H_PUWYV[LJ[ZHNHPUZ[Z[YLZZ
JVUKP[PVUZ
)65(56404(9*,33("7(:;690=(3,5;05("05=,950AA0.(,;(56
9,.65,:04(90(,3,5(";69;69(7(636
+PWHY[PTLU[VKP)PV[LJUVSVNPLL)PVZJPLUaL<UP]LYZP[nKP4PSHUV)PJVJJH4PSHUV0[HS`
:WPUVJLYLILSSHYH[H_PH[`WL:*(PZHUL\YVKLNLULYH[P]LKPZVYKLYJH\ZLKI`[OL
L_WHUZPVUIL`VUKHJLY[HPU[OYLZOVSKVMHWVS`NS\[HTPULWVS`8[YHJ[PU[OLWYV[LPU
H[H_PU (; (; JVUZPZ[Z VM H NSVI\SHY 5[LYTPUHS 1VZLWOPU KVTHPU HUK VM H
ÅL_PISL*[LYTPUHS[HPSJVU[HPUPUN[OLWVS`8[YHJ[;OLWH[OVSVN`YLZ\S[ZMYVTWYV[LPU
TPZMVSKPUNHUKPU[YHJLSS\SHYHJJ\T\SH[PVUVMÄIYPSSHYHT`SVPKSPRLHNNYLNH[LZ;OLSVZZ
VM M\UJ[PVU YLZ\S[PUN MYVT TPZMVSKPUN TPNO[ HSZV IL PU]VS]LK PU [OL TLJOHUPZTZ VM
WH[OVNLULZPZ(;PZHJVUZLY]LKHUK\IPX\P[V\ZWYV[LPURUV^U[VIPUKWVS`\IPX\P[PU
JOHPUZ HUK [V M\UJ[PVU HZ H KL\IPX\P[PUH[PUN LUa`TL 0[ ZLLTZ [V IL PU]VS]LK PU
KPMMLYLU[ JLSS\SHY WH[O^H`Z PL HNNYLZVTL MVYTH[PVU HUK \IPX\P[PUWYV[LHZVTL
WH[O^H`I\[[OLWO`ZPVSVNPJHSYVSLPZZ[PSSWVVYS`\UKLYZ[VVK;VPU]LZ[PNH[LWVZZPISL
M\UJ[PVUZ VM (; PU JLSS\SHY WH[O^H`Z [OH[ YLZWVUK [V HS[LYLK WYV[LPU OVTLVZ[HZPZ
^LJVUZ[P[\[P]LS`L_WYLZZLKZL]LYHS(;]HYPHU[ZPU7PJOPHWHZ[VYPZHTL[O`SV[YVWOPJ
`LHZ[SHJRPUNWYV[LPUZOVTVSVNV\Z[V(;.YV^[OHZZH`ZZOV^LK[OH[L_WYLZZPVU
VM ^PSK [`WL (; VY [OL ZVSL 1VZLWOPU KVTHPU HSSV^Z `LHZ[ NYV^[O L]LU \UKLY
Z[YLZZ JVUKP[PVUZ Z\JO HZ OLH[ZOVJR VY ,9 Z[YLZZLZ KP[OPV[OYLP[VS [\UPJHT`JPU
0UJVU[YHZ[[OLL_WYLZZPVUVMHJH[HS`[PJHSS`PUHJ[P]L]HYPHU[JV\SKUV[YLZ[VYL`LHZ[
]PHIPSP[`^OPJOZ\NNLZ[Z[OH[[OLKL\IPX\P[PUH[PUNHJ[P]P[`VM(;PZYLX\PYLKMVY[OL
WYV[LJ[P]LLMMLJ[>LHSZVJYLH[LKH[YHUZNLUPJ*LSLNHUZZ[YHPUV]LYL_WYLZZPUN^PSK
[`WL(;PUUL\YVUZ\UKLY[OLJVU[YVSVMHWHUUL\YVUHSWYVTV[LY\UJ )HZLK
VUIVK`ILUKZJV\U[PUNWYLSPTPUHY`Z[\KPLZZ\NNLZ[[OH[\UKLYZ[YLZZJVUKP[PVU*
LSLNHUZV]LYL_WYLZZPUN^PSK[`WL(;ZOV^ZHUPUJYLHZLKTV[PSP[`JVTWHYLK^P[O
^PSK[`WL6\YKH[HZ\NNLZ[[OH[(;[OHURZ[VP[ZKL\IPX\P[PUH[PUNHJ[P]P[`KPZWSH`ZH
WYV[LJ[P]LLMMLJ[HNHPUZ[Z[YLZZJVUKP[PVUZ
007VZ[LY:LZZPVU
PRO 9
Proteine
:[Y\J[\YHSJOHYHJ[LYPaH[PVUVMKPVPJPUUV]LS[`WL
YPIVZVTLPUHJ[P]H[PUNWYV[LPUMYVTSLH]LZVM7O`[VSHJJH
KPVPJH3
9<::696:0;(+04(96(5;046*/(4),9@(5.,3((5+
7(9,5;,(<.<:;6
+PWHY[PTLU[VKP:JPLUaLL;LJUVSVNPL(TIPLU[HSP)PVSVNPJOLLK-HYTHJL\[PJOL
:LJVUKH<UP]LYZP[nKP5HWVSP=PH=P]HSKP0*HZLY[H0[HS`
9PIVZVTL PUHJ[P]H[PUNWYV[LPUZ 907Z" ,* HYL ZPUNSLJOHPU [`WL VY
OL[LYVKPTLYPJJ`[V[V_PJ[`WLZHUKLUa`TLZLUKV^LK^P[OY95(5ơNS`JVZPKHZL
HJ[P]P[`BD907ZYLTV]LHZP[LZWLJPÄJZPUNSLHKLUPULYLZPK\L(PU[OLJHZLVMYH[
SP]LY YPIVZVTLZ MYVT [OL OPNOS` JVUZLY]LK ZHYJPUYPJPU SVVW VM [OL : Y95( [O\Z
HYYLZ[PUNWYV[LPUZ`U[OLZPZ BD;OLNLU\Z7O`[VSHJJHPZRUV^U[VILHYPJOZV\YJLVM
ZL]LYHS[`WL907BD!P7(7000HUK000MYVTSLH]LZVM7O`[VSHJJHHTLYPJHUH3"PP7+
:MYVTZLLKZ"7+3MYVTSLH]LZVMHK\S[WSHU[ZVM7KPVPJH3"KPVPJPULKPVPJPU
MYVTSLH]LZVM`V\UNWSHU[Z\W[V[OYLL`LHYZHUKMYVTKL]LSVWPUNSLH]LZVMHK\S[
WSHU[ZBD0U[OLSHZ[L_WLYPTLU[HSZ`Z[LTKPVPJPUPZHS^H`ZWYLZLU[^OPSLKPVPJPUPZ
L_WYLZZLK\W[VKH`ZL]LU[LLU;OLMHJ[[OH[7KPVPJHZ`U[OLZPaLZHUKHJJ\T\SH[LZ
907ZPUKPYLJ[S`PUKPJH[LZ[OH[[OLPYWYLZLUJLTPNO[ILILULÄJPHS[VWSHU[Ä[ULZZ
/LYL^LYLWVY[[OLKL[LYTPUH[PVUVMZ[Y\J[\YHSHUKM\UJ[PVUHSWYVWLY[PLZVMKPVPJPU
PZVSH[LKHJJVYKPUN[V7HYLU[LL[HSBD;OLW\YP[`VM[OLWYV[LPU^HZ]LYPÄLKI`:+:
7(.,HUK,:08;6-4:HUHS`ZPZ;OLU\ZPUNHZ[YH[LN`^OPJOJVTIPULZ,KTHU
KLNYHKH[PVU HUK THZZ ZWLJ[YVTL[Y` ,:08;6- [OL VM KPVPJPU WYPTHY`
Z[Y\J[\YL^HZKL[LYTPULK
;OLH]HPSHISLWYPTHY`Z[Y\J[\YLVMKPVPJPUPZOPNOS`PKLU[PJHS[V7(700MYVT7O`[VSHJJH
PJVZHUKYH 3 -\Y[OLYTVYL [OL JVTWSL[PVU VM P[Z WYPTHY` Z[Y\J[\YL ^PSS HSSV^ \Z [V
LZ[HISPZOWO`SVNLUL[PJYLSH[PVUZOPWZ^P[OHSSRUV^U907ZMYVT7O`[VSHJJHJLHL
9LMLYLUJLZ!
BD.PYILZL[HS4PUP9L]4LK*OLT
BD,UKV@HUK;Z\Y\NP2 1)PVS*OLT BD7L\THUZ>1L[HS-(:,)1 BD7HYLU[LL[HS;V_PJWSHU[WYV[LPUZ3VYK14/HY[SL`49,KZ:WYPUNLY/LPKLSILYNWW 007VZ[LY:LZZPVU
PRO 10
Proteine
7+A+VTHPUZHUK4LTIYHUL5\JSLVSPUPU[OL,UNPULLYPUN
HUK7YVK\J[PVUVM*OPTLYPJ;V_PUZ(NHPUZ[*HUJLY*LSSZ
-9(5*,:*6.0(5:(5;03<(5(+03,(5+96:()905(4,0
(55(4(90(*0405096+63-6077630;0
+LWHY[TLU[VM/LHS[O3PMLHUK,U]PYVUTLU[HS:JPLUJLZ<UP]LYZP[`VM3»(X\PSH
03»(X\PSH0[HS`
0U[OLYLZLHYJOVMUL^LUNPULLYLK[V_PUZ[V[HYNL[JHUJLYJLSSZ^LOH]LZ[\KPLK7+A
WYV[LPUKVTHPUHZWVZZPISL[VVSZJLSS\SHY[HYNL[PUNVM[OLYPIVZVTLPUHJ[P]H[PUNWYV[LPU
ZHWVYPU7+AKVTHPUZHYLMV\UKPUWYV[LPUZPU]VS]LKPUJLSS\SHYWYVJLZZLZMYVT[OL
ZPNUHS[YHUZK\J[PVU[V[OL[YHUZWVY[HUKWHY[PJ\SHYS`PUZ`UHWZLMVYTH[PVUPUTHTTHSZ
;OL`YLJVNUPaLHUKIPUKZOVY[WLW[PKLZLX\LUJLZSVJH[LKH[[OL*[LYTPU\ZVMHWYV[LPU
9LJVNUP[PVUVM[OL*[LYTPUPYLWYLZLU[ZHºUVUPU]HZP]L»PU[LYHJ[PVU]LY`^LSSZ\P[LK
[VTLKPH[LVYNHUPaH[PVUVM[YHUZWVY[SVJHSPaH[PVUZVY[PUNHUKZWH[PHSHYYHUNLTLU[VM
WYV[LPUZL_WVZPUN[OLPYPUKP]PK\HS*[LYTPUHS[HPSZ[OH[JHUILYLJVNUPaLKHUKOHUKSLK
I`]HYPV\Z7+AKVTHPUZ>LOH]LMVJ\ZZLKV\YH[[LU[PVU[V[OL7+AKVTHPUMYVT
O*(:2/\THUJHSJP\TJHSTVK\SPUKLWLUKLU[ZLYPULWYV[LPURPUHZL[OH[OHZILLU
KLTVUZ[YH[LK[VIPUKL_[YHJLSS\SHY*+ PULWP[OLSPHSJLSSZYLJVNUPaLKHZHTHYRLY
MVY ZL]LYHS O\THU [\TVYZ >L WYVK\JLK M\ZPVUZ VM O*(:27+A ^P[O ZHWVYPU HUK
[LZ[LK P[ VU O\THU NSPVISHZ[VTH JLSSZ ;OLZL JVUZ[Y\J[Z WYV]LK [V IL [V_PJ ^P[O
PUJYLHZPUNHJ[P]P[`HZHM\UJ[PVUVM[OLU\TILYVM7+AKVTHPUZHUKPUK\JLJLSSKLH[O
I`HWVW[V[PJTLJOHUPZTZ-\Y[OLYTVYL^LHYLL_WSVYPUNU\JSLVSPUL_WYLZZLKVU[OL
JLSSTLTIYHULVMJHUJLYJLSSZHZHZWLJPÄJ[HYNL[VMO\THUNSPVISHZ[VTHZPUJLP[Z
TLTIYHULL_WYLZZPVUPUNSPVTHZPZZ[YPJS[S`JVYYLSH[LK^P[O[OLPYTHSPNUHUJ`HUKJHU
ILL_WSVP[LK[VKLSP]LY[V_PJZ\IZ[HUJLZ>LZOV^PU[OPZ^VYRWYLSPTPUHY`KH[HVUH
+5(JVUZ[Y\J[JHYY`PUN[OLNLULVM[OLZHWVYPU[V_PUSPURLK[V[OL(:HW[HTLY
ZWLJPÄJMVYU\JSLVSPU;V_PJP[`VUNSPVISHZ[VTHJLSSZPZYLWVY[LK7VZZPISLHWWSPJH[PVUZ
VM[OLZL[HYNL[PUNZ[YH[LNPLZ^PSSILKPZJ\ZZLK
007VZ[LY:LZZPVU
PRO 11
Proteine
(UHSPZPWYV[LVTPJHKLSSHMYHTTLU[HaPVULKLSSLWYV[LPUL
WSHZTH[PJOLLK\YPUHYPL
PUWHaPLU[PZVNNL[[PH[YHWPHU[VYLUHSL
-<93(50,4030(563,65*05096),9;6:*(7,33(;6*(936
*,5:<9(;6*6:;(5;05(.<,99(5;096),9;6
+PWHY[PTLU[V4LKPJPUH0U[LYUH:JPLUaL,UKVJYPUV4L[HIVSPJOLL)PVJOPTPJH
<UP]LYZP[nKP:PLUH
<6*3HIVYH[VYPV7H[VSVNPH*SPUPJH(aPLUKH6ZWLKHSPLYH<UP]LYZP[HYPH:LULZL
<6*5LMYVSVNPH+PHSPZPL;YHWPHU[V(aPLUKH6ZWLKHSPLYH<UP]LYZP[HYPH:LULZL
5LSWSHZTHLULSSL\YPULZVUVWYLZLU[PWYV[LPULPU[LYLLYLSH[P]PMYHTTLU[PJYLH[PKHSSL
WYV[LHZPÄZPVSVNPJOLWV[LUaPHSTLU[LPTWVY[HU[PULSSHKPHNUVZ[PJHKPSHIVYH[VYPVWVPJOt
SHWYV[LVSPZPW\~LZZLYLHS[LYH[HKH]HYPLJVUKPaPVUPWH[VSVNPJOL
0U \U YLJLU[L Z[\KPV Z\ WHaPLU[P JVU PUZ\MÄJPLUaH YLUHSL JYVUPJH 09* ZVUV Z[H[P
PKLU[PÄJH[P KLP WLW[PKP [YH P X\HSP MYHTTLU[P KP HSI\TPUH 3H KL[LYTPUHaPVUL
KLSS»HSI\TPUHULSSL\YPULuMVUKHTLU[HSLULPWHaPLU[PJVUWH[VSVNPLULMYVSVNPJOLMYH
J\P S»(J\[L 2PKUL` 0UQ\Y` (20 9PN\HYKV HP WHaPLU[P ZV[[VWVZ[P H [YHWPHU[V YLUHSL
TVS[PKPLZZP]HUUVPUJVU[YVHK\UHMVYTHKP(20JOLJVUZPZ[LPU\UHYP[HYKH[HYPWYLZH
M\UaPVUHSLKLSS»VYNHUVV+LSH`LK.YHM[-\UJ[PVU+.-0UVS[YLULNSP\S[PTPHUUPuÄVYP[H
SHYPJLYJHKPU\V]PIPVTHYRLYKP(20L+.-WPƒWYLJVJPLKLMÄJPLU[PKPX\LSSP[YHKPaPVUHSP
3V ZJVWV KP X\LZ[H PUKHNPUL WYLSPTPUHYL u K\WSPJL! PUKP]PK\HYL P MYHTTLU[P
KLSS»HSI\TPUH \YPUHYPH TVKPÄJH[P YPZWL[[V HS WSHZTH WLY NL[[HYL S\JL Z\SS»H[[P]P[n
WYV[LHZPJHULSJVU[LZ[VKLS[YHWPHU[VYLUHSL"JVUMYVU[HYLSVZJOLTHKLPMYHTTLU[PKP
HSI\TPUHWSHZTH[PJHL\YPUHYPH[YHSLK\LJH[LNVYPLKPWHaPLU[PJOL]HUUVPUJVU[YV
VUVU]HUUVPUJVU[YVH+.-WLY]HS\[HYUL\UWVZZPIPSL\[PSPaaVJVTLTHYJH[VYLKP
YLJ\WLYVKLSS»VYNHUV
7LY X\LZ[V HIIPHTV LMML[[\H[V S»LSL[[YVMVYLZP IPKPTLUZPVUHSL Z\ NLS KLSSL WYV[LPUL
WSHZTH[PJOL L \YPUHYPL KP ZVNNL[[P [YHWPHU[H[P :\ [\[[P P JHTWPVUP PUVS[YL u
Z[H[V LZLN\P[V \U PTT\UVISV[ JVU SV Z[LZZV (I HU[PHSI\TPUH PTWPLNH[V ULSSL
KL[LYTPUHaPVUP ULMLSVTL[YPJOL WLY L]PKLUaPHYL L]LU[\HSP KPMMLYLUaL ULS WH[[LYU KLP
MYHTTLU[PWYV[LVSP[PJP
0UVZ[YPYPZ\S[H[PL]PKLUaPHUVHSJ\ULKPMMLYLUaLMYHPMYHTTLU[PWSHZTH[PJPLX\LSSP\YPUHYP
JVZyJVTL[YHPWHaPLU[PJVU+.-LX\LSSPZLUaH+.-8\LZ[PKH[PWYLSPTPUHYPWLYLZZLYL
JVUMLYTH[POHUUVIPZVNUVKP\S[LYPVYPZ[\KPJVU\UTHNNPVYU\TLYVKPJHTWPVUP
4L[aNLY1L[HS<YPUHY`L_JYL[PVUVM[^LU[`WLW[PKLZMVYTZHULHYS`HUKHJJ\YH[LKPHNUVZ[PJWH[[LYUVM
HJ\[LRPKUL`PUQ\Y`2PKUL`0U[¶
007VZ[LY:LZZPVU
PRO 12
Proteine
0TWSPJH[PVUZVM3HTPUHUK,TLYPUT\[H[PVUZPU+PSH[LK
*HYKPVT`VWH[OPLZ
)633(;040*/,3(-(=(330=(3,5;05((9)<:;050,360:(
)636.5,:04(9;056
+LWHY[TLU[VM)PV:JPLUJLZ<UP]LYZP[`VM4PSHUV4PSHUV0[HS`
*LU[YLMVY0UOLYP[LK*HYKPV]HZJ\SHY+PZLHZLZ
-VUKHaPVUL09**:7VSPJSPUPJV:HU4H[[LV7H]PH0[HS`
3HTPU(*PZHU\JSLHYZWLJPÄJ[`WL=PU[LYTLKPH[LÄSHTLU[WYV[LPUI\PSKPUN\W[OL
THQVYZ[Y\J[\YHSJVTWVULU[ZVM[OLU\JSLHYSHTPUH+LMLJ[ZPU[OL3HTPUNLULOH]L
ILLUSPURLK[VTVYL[OHU[LUKPMMLYLU[KPZLHZLZRUV^UHZSHTPUVWH[OPLZ"[OLZLHYL
HZZVJPH[LK [V JHYKPHJ JVTWSPJH[PVUZ [OH[ VM[LU [\YU PU[V KPSH[LK JHYKPVT`VWH[O`
+*4HTHQVYJH\ZLVMJVUNLZ[P]LOLHY[MHPS\YLBD>P[OPU[OLU\JSLHYSHTPUH3HTPU
PU[LYHJ[Z^P[OZL]LYHSU\JSLHYTLTIYHULWYV[LPUZ[O\ZWYV]PKPUNHUL^RL`[HYNL[MVY
[OL Z[\K` VM [OL LMMLJ[Z VM +*4YLSH[LK T\[H[PVUZ (TVUN PU[LYHJ[VYZ ,TLYPU OHZ
ILLUWYV]LK[VIPUKKPYLJ[S`[VSHTPUHUKT\[H[PVUZPU[OL,TLYPUNLULJH\ZL+*4
ZPTPSHYS` [V 3HTPU T\[H[PVUZ :V MHY UV Z[Y\J[\YLZ VM M\SS SLUN[O 3HTPU HUK ,TLYPU
OH]L ILLU YLWVY[LK OHTWLYPUN WHY[ VM [OL Z[Y\J[\YLIHZLK KY\N YLZLHYJO 0U [OPZ
JVU[L_[^LKLZPNULKUV]LSJVUZ[Y\J[ZVMM\SSSLUN[O3HTPUHUKVM[OL,TLYPU3HTPU
IPUKPUNKVTHPU[OH[OH]LILLUW\YPÄLK"JY`Z[HSSPaH[PVUHUKJVJY`Z[HSSPaH[PVU[YPHSZHYL
J\YYLU[S`VUNVPUN0UWHYHSSLS^LZVS]LK[OLJY`Z[HSZ[Y\J[\YLZVM[^V345(*VPS)
KVTHPUT\[HU[Z,2HUK9>[OH[HSSV^JOLJRPUN[OLLMMLJ[ZVMT\[H[PVUZPU
[LYTZVMZ[Y\J[\YL]HYPHIPSP[`VM[OLHMMLJ[LKWYV[LPUHUKOLSWZ[\K`PUNV]LYHSSWYV[LPU
Z[HIPSP[`HUK[OLPTWSPJH[PVUZVMT\[HU[ZPU[OLJVU[L_[VM+*4BD
(YI\Z[PUP,L[HS(\[VZVTHSKVTPUHU[KPSH[LKJHYKPVT`VWH[O`^P[OH[YPV]LU[YPJ\SHYISVJR!HSHTPU(*
KLMLJ[YLSH[LKKPZLHZL1(T*VSS*HYKPVS ¶ )VSSH[P 4 L[ HS :[Y\J[\YLZ VM [OL SHTPU (* 9> HUK ,2 T\[HU[Z! PTWSPJH[PVUZ MVY KPSH[LK
JHYKPVSHTPUVWH[OPLZ)PVJOLT)PVWO`Z9LZ*VTT\U
007VZ[LY:LZZPVU
PRO 13
Proteine
-YVT.LUVTL[V(U[PNLU!H:[Y\J[\YHS9L]LYZL=HJJPUVSVN`
HWWYVHJO[VWV[LU[PHS)\YROVSKLYPHWZL\KVTHSSLPHU[PNLUZ
3(::(<?7(;90*0(.6<93(@36<0:,7,90*3(<+06-,99,95(=(96
4(906*65*/0336:63i6:*(9+(<9(?(=0,9365./09,5(;6
*6364)6.069.06(5+)636.5,:04(9;056
+LWHY[TLU[VM)PV:JPLUJLZ<UP]LYZP[`VM4PSHU4PSHU0[HS`
+LWHY[TLU[VM*VTW\[H[PVUHS)PVSVN`0Z[P[\[VKP*OPTPJHKLS9PJVUVZJPTLU[V
4VSLJVSHYL*VUZPNSPV5HaPVUHSLKLSSL9PJLYJOL4PSHUV0[HS`
0UZ[P[\[LVM)PV[LJOUVSVN`HUK)PVTLKPJPUL<UP]LYZP[H[(\[~UVTHKL)HYJLSVUH:WHPU
=HJJPUL KL]LSVWTLU[ OHZ YLJLU[S` ILLU JOHUNLK I` [OL H]HPSHIPSP[` VM NLUVTLZ
MYVTTHU`WH[OVNLUZ"HKKP[PVUHSS`UL^WLYZWLJ[P]LZPU]HJJPUL\ZLOH]LLTLYNLK
Z\JOHZ[HYNL[PUNHK\S[ZJVTIPUPUNHU[PNLUZ\ZPUNUL^TL[OVKVMHKTPUPZ[YH[PVU
Z[PT\SH[PUNPUUH[LPTT\ULYLZWVUZLOLSWPUNJ\YLVMJOYVUPJPUMLJ[PVUZVYKLMLUKPUN
[OLWVW\SH[PVUMYVTIPV[LYYVYPZT
0U [OPZ JVU[L_[ ^L MVJ\ZLK VU )\YROVSKLYPH WZL\KVTHSSLP HU HU[PIPV[PJ YLZPZ[HU[
WH[OVNLUSPZ[LKHZHIPVSVNPJHS[OYLH[HNLU[VMJSHZZYLZWVUZPISLVM4LSPVPKVZPZ
H KPZLHZL HMMLJ[PUN IV[O O\THUZ HUK HUPTHSZ ÄYZ[ KLZJYPILK PU 0U [OL SHZ[
[LU`LHYZ4LSPVPKVZPZOHZILJVTLLUKLTPJPU[YVWPJHSHYLHZVM:V\[OLHZ[(ZPHHUK
5VY[OLYU (\Z[YHSPH ^P[O OPNO TVY[HSP[` YH[LZ P[Z KPHNUVZPZ ILPUN KPMÄJ\S[ ZPUJL [OL
Z`TW[VTZTPTPJV[OLYKPMMLYLU[PUMLJ[PVUZ
)HZLK VU ) WZL\KVTHSSLP NLUVTPJ PUMVYTH[PVU ^L HWWSPLK HU PTWYV]LK 9L]LYZL
=HJJPUVSVN` HWWYVHJO [V ZLSLJ[ WV[LU[PHS WYV[LPU HU[PNLUZ [V IL [LZ[LK HZ ]HJJPUL
JHUKPKH[LZ 6\Y HWWYVHJO MVYLZLLZ [OL HWWSPJH[PVU VM M\UJ[PVUHS NLUVTPJZ WYV[LPU
TPJYVHYYH`Z IPVPUMVYTH[PJZJVTW\[H[PVUHS IPVSVN` JVTWSLTLU[LK I` Z[Y\J[\YHS
IPVSVN`Z[\KPLZ0UKLLKZPUJLPTT\UP[`PZLUOHUJLKI`YLJVNUP[PVUVMHULWP[VWL
WHY[ VM H WYV[LPU PU V\Y JHZL P[Z [OYLLKPTLUZPVUHS Z[Y\J[\YL Z[HIPSP[` HUK YLSH[LK
K`UHTPJZHYLRL`MHJ[VYZN\PKPUNHU[PNLUZLSLJ[PVU
;VKH[LJVUZ[Y\J[ZOH]LILLUKLZPNULKPUV\YSHIMVYOL[LYVSVNV\ZL_WYLZZPVUPU
,JVSPHZ.:;VY/PZ[HNNLKM\ZPVUZWYV[LPUZ"WYV[LPUW\[H[P]LHU[PNLUZOH]LILLU
JY`Z[HSSPaLKHUK+Z[Y\J[\YLZZVS]LK;OLWYVNYLZZTHKL[VKH[LPZOLYLYLWVY[LK"[OL
)WZL\KVTHSSLP6WW([HYNL[^PSSILWYLZLU[LK[VKLZJYPIL[OL^OVSLUL^TL[OVKVSVN`
HUKP[ZPTWSPJH[PVUZ
9HWW\VSP99L]LYZL=HJJPUVSVN`*\YYLU[6WPUPVUPU4PJYVIPVSVN`!¶
.V\YSH`31:HU[P07LaaPJVSP(.YHUKP.:VYPHUP4HUK)VSVNULZP4 .YV\W):[YLW[VJVJJ\Z
W\SS\SHUHZLJY`Z[HSZ[Y\J[\YLZPU[OLJVU[L_[VMHUV]LSZ[YH[LN`MVY]HJJPULKL]LSVWTLU[1)HJ[LYPVS
007VZ[LY:LZZPVU
PRO 14
Proteine
;OLYVSLVM[OLTVS[LUNSVI\SLZ[H[LPU[OLMVYTH[PVUVM
HT`SVPKSPRLZ[Y\J[\YLZI`O\THUƠHJPKNS`JVWYV[LPU
)(3+(::(99,4(<90A06:*09Ï(5+9,()(9;/(5+9,(:
.(3,(AA096),9;((5+;(5-(50-()06
+PWHY[PTLU[VKP:JPLUaLKLSSH=P[HLKLSS»(TIPLU[L
<UP]LYZP[n7VSP[LJUPJHKLSSL4HYJOL(UJVUH0[HS`
+LWHY[TLU[VM)PVJOLTPZ[Y`HUK)PVWO`ZPJZ
:[VJROVST<UP]LYZP[`:[VJROVST:^LKLU
Ơ(JPKNS`JVWYV[LPU(.7PZHOLH]PS`NS`JVZPSH[LKWSHZTH[PJWYV[LPUILSVUNPUN[V[OL
PTT\UVJHSPUMHTPS`HZ\INYV\WVM[OLSPWVJHSPUMHTPS`PU]VS]LKPUPTT\ULYLZWVUZL
;OLWYV[LPUZ[Y\J[\YLMLH[\YLZHJLU[YHSơIHYYLSMVYTPUNHO`KYVWOVIPJIPUKPUNZP[LMVY
H^PKLU\TILYVMIPVSVNPJHSS`HJ[P]LL_VNLUV\ZHUKLUKVNLUV\ZJVTWV\UKZ^OPSL
[OLVSPNVZHJJOHYPKLZHSSV^MVY[OLPU[LYHJ[PVU^P[OHUKYLJVNUP[PVUI`ZWLJPÄJJLSS
[`WLZVM[OLPTT\ULZ`Z[LT-V\YPLY;YHUZMVYTPUMYHYLKZWLJ[YVZJVW`VM[LTWLYH[\YL
PUK\JLKKLUH[\YH[PVUVM(.7YL]LHSLK[OLWYLZLUJLVMHTVS[LUNSVI\SLSPRLMVSKPUN
PU[LYTLKPH[L4.H[UL\[YHSHUKHJPKPJW/]HS\LZ^P[OUVZ\JOPU[LYTLKPH[LILPUN
KL[LJ[LK PU HSRHSPUL VY Z[YVUNS` HJPKPJ LU]PYVUTLU[Z 0U[LYLZ[PUNS` M\Y[OLY OLH[
KLUH[\YH[PVUVM[OL4.Z[H[LVI[HPULKH[HJPKPJW/]HS\LZPUP[PH[LKHNNYLNH[PVUVM[OL
WVS`WLW[PKLJOHPUZPU[VPU[LYTVSLJ\SHYHU[PWHYHSSLSơZOLL[Z;OPZPZHJJVTWHUPLKI`
[OL[`WPJHSZWLJ[YHSJOHUNLZVJJ\YYPUNPU[OLHIZVYW[PVUHUKÅ\VYLZJLUJLLTPZZPVU
ZWLJ[YH VM [OL HT`SVPKIPUKPUN K`LZ *VUNV 9LK HUK ;OPVÅH]PU ; YLZWLJ[P]LS`
Z\NNLZ[PUNMVYTH[PVUVMÄIYPSZMLH[\YPUNHJYVZZơHYJOP[LJ[\YL7VZP[P]LYLZ\S[Z^LYL
HSZVVI[HPULK\WVUHUHS`ZLZVM[OLHNNYLNH[LKZHTWSLZ^P[O[YHUZTPZZPVULSLJ[YVU
HUKWVSHYPaLKSPNO[TPJYVZJVWPLZ
0U VYKLY [V WYVIL PU NYLH[LY KL[HPS [OL JVUULJ[PVU VM 4. HUK [OL HNNYLNH[PVU VM
(.7[PTLYLZVS]LKPUMYHYLKZWLJ[YH^LYLJVSSLJ[LK^P[OTPSSPZLJVUKYLZVS\[PVUHM[LY
PUZ[HU[YLSLHZLVMWYV[VUZI`[OLWOV[VYLHJ[P]L¸JHNLK¹JVTWV\UKUP[YVWOLU`S
L[O`SZ\SWOH[L(JOHUNLPU[OL[PTLJVUZ[HU[VMHU[PWHYHSSLSơZOLL[MVYTH[PVUJV\SK
IL TLHZ\YLK HSSV^PUN \Z [V HZZLZZ [OL LMMLJ[ VM [LTWLYH[\YL HUK W/ VU [OL 4.
KYP]LUHNNYLNH[PVUVM(.7;OLKH[HZ\NNLZ[[OH[WHY[PHSVYJVTWSL[LL_WVZ\YLVM[OL
HNNYLNH[PVUWYVULYLNPVUZJV\SK[YPNNLY(.7HZZLTIS`PU[VHT`SVPKZ[Y\J[\YLZ
007VZ[LY:LZZPVU
PRO 15
Proteine
(UHS`ZPZVMW2PWU\JSLHYTL[HIVSPZTPUYL[PUVPJ[YLH[LK
UL\YVISHZ[VTHJLSSZ
),5*0=,5.(+)69.0((;9(465;(56(*(3+(9,33007096AA0(
630=((+,33(9(.065,-(5+)6990,336(
+LWHY[TLU[VM)PVJOLTPZ[Y`)PVWO`ZPJZHUK.LULYHS7H[OVSVN`
:LJVUK<UP]LYZP[`VM5HWSLZ=PH+L*YLJJOPV5HWSLZ"
;OLZLH\[OVYZLX\HSS`JVU[YPI\[LK[V[OLZ[\K`
W2PWPZHJ`JSPUKLWLUKLU[RPUHZL*+2PUOPIP[VY*20ILSVUNPUN[V[OL*PW2PW
MHTPS`[OH[HSZVPUJS\KLZW*PWHUKW2PW;OLWYV[LPUYLN\SH[LZJLSSWYVSPMLYH[PVU
I` TVK\SH[PUN [OL HJ[P]P[` VM *+2Z \UKLY ZL]LYHS JVUKP[PVUZ PUJS\KPUN JLSS[VJLSS
JVU[HJ[ PUOPIP[PVU HUK [YLH[TLU[ ^P[O H SHYNL U\TILY VM HU[PWYVSPMLYH[P]L KY\NZ 0U
HKKP[PVU[V[OLTVK\SH[PVUVMJLSSWYVSPMLYH[PVUW2PWJVU[YVSZJLSSKPMMLYLU[PH[PVU
JLSSTV[PSP[`HUKPU[LYHJ[PVU^P[O[OLZ\IZ[YH[LNLUL[YHUZJYPW[PVUHUK+5(K\WSPJH[PVU
W2PW HJ[P]P[` PZ JVU[YVSSLK I` P[Z JVUJLU[YH[PVU P[Z KPZ[YPI\[PVU HTVUN KPMMLYLU[
JLSS\SHYJVTWSL_LZHUKP[ZJLSS\SHYSVJHSPaH[PVU(U\TILYVML]PKLUJLKLTVUZ[YH[LK
[OH[[OLTL[HIVSPZTHUKPU[\YU[OLPU[LYHJ[VYZVM[OL*20HYLTVK\SH[LKI`P[ZWVZ[
[YHUZSH[PVUHSTVKPÄJH[PVUZHUKTHPUS`I`WOVZWOVY`SH[PVU
:L]LYHS KPMMLYLU[ YLZPK\LZ PU O\THU W2PW OH]L ILLU KLTVUZ[YH[LK [V IL
WOVZWOVY`SH[LK UHTLS` :LY ;`Y ;`Y ;`Y ;OY ;OY HUK ;OY ;OY WOVZWOVY`SH[PVU I` HJ[P]L *+2 PZ YLX\PYLK [V JYLH[L H WOVZWOVKLNYVU
ULJLZZHY` MVY :RW IPUKPUN HUK [OL Z\IZLX\LU[ *20 \IPX\P[PUH[PVUKLNYHKH[PVU
*VU]LYZLS`:LYWOVZWOVY`SH[PVUOHZILLUYLWVY[LK[VPUJYLHZLW2PWZ[HIPSP[`I`
LSVUNH[PUNP[ZOHSMSPML6\YWYL]PV\ZZ[\KPLZKLTVUZ[YH[LK[OH[UL\YVISHZ[VTHJLSSZ
[YLH[TLU[^P[OHSS[YHUZYL[PUVPJHJPKPUK\JLZHYHWPKUL\YVUHSSPRLJLSSKPMMLYLU[PH[PVU
HZZVJPH[LK[VHUPUJYLHZLVMWOVZWOV:LYW2PWPU[OLU\JSLHYJVTWHY[TLU[
;OL NVHS VM [OL WYLZLU[ Z[\K` OHZ ILLU [V PKLU[PM` [OL TVSLJ\SHY TLJOHUPZTZ
YLZWVUZPISLVMWOVZWOV:LYW2PWPUJYLHZLKZ[HIPSP[`;V[OPZHPT^LWYLSPTPUHY`
KL]LSVWLK H TL[OVKVSVN` MVY [OL JOHYHJ[LYPaH[PVU VM W2PW WOVZWOVPZVMVYTZ
IHZLKVU+WYV[LPUZLWHYH[PVUHZZVJPH[LK[V[OL\ZLVM^LSSJOHYHJ[LYPaLKZWLJPÄJ
HU[PIVKPLZ HNHPUZ[ [OL *20 WOVZWOVPZVMVYTZ 6\Y ÄUKPUNZ KLTVUZ[YH[L [OH[ [OL
PUJYLHZLVMWOVZWOV:LYW2PWZ[HIPSP[`PZK\L[V[OLZJHYJLWOVZWOVY`SH[PVUVM
[OLPZVMVYTVU;OY([WYLZLU[^LHYLPU]LZ[PNH[PUN^OL[OLY[OPZLMMLJ[PZK\L[VH
KPTPUPZOLKPU[LYHJ[PVU^P[O*+2VY[VHUHS[LYLKIPUKPUNVM[OLWYV[LPU[V[OLRPUHZL
007VZ[LY:LZZPVU
PRO 16
Proteine
(UHS`ZPZVMLY`[OYVJ`[LTLTIYHULZI`KPMMLYLU[WYV[LVTPJ
TL[OVKVSVNPLZPU9L[[:`UKYVTL
*69;,3(AA6(3,::06.<,99(5;096),9;63,65*050:03=0(
:0.569050*05A0(3(5+0.0(*6467,*69,330(3,::(5+9(
=(3(**/0.0<:,77,+,-,30*,*3(<+06*0**6303<*0(
/(@,216<::,-
+LWHY[TLU[VM0U[LYUHS4LKPJPUL,UKVJYPUL4L[HIVSPJ:JPLUJLZHUK)PVJOLTPZ[Y`
<UP]LYZP[`VM:PLUH:PLUH0[HS`"+LWHY[TLU[VM7H[OVWO`ZPVSVN`,_WLYPTLU[HS
4LKPJPULHUK7\ISPJ/LHS[O<UP]LYZP[`VM:PLUH:PLUH0[HS`"+LWHY[TLU[VM
,]VS\[PVUHY`)PVSVN`<UP]LYZP[`VM-LYYHYH-LYYHYH0[HS`"5LVUH[HS0U[LUZP]L*HYL
<UP[<UP]LYZP[`/VZWP[HS(6<::PLUH0[HS`"*OPSK5L\YVWZ`JOPH[Y`<UP[
<UP]LYZP[`/VZWP[HS(6<::PLUH0[HS`
9L[[Z`UKYVTL9;;^P[OHMYLX\LUJ`VMe!¶!MLTHSLZPZHYHYLHUK
JVTWSL_ UL\YVKL]LSVWTLU[HS KPZVYKLY BD ^OPJO WYLZLU[Z HZ H UL\YVYLNYLZZPVU
MVSSV^PUNHUVYTHSKL]LSVWTLU[MVY¶TVU[OZ^P[OSVZZVMJVNUP[P]LZVJPHSHUK
TV[VYZRPSSZPUH[`WPJHSZ[HNLYLNYLZZPVU;VKH[LT\[H[PVUZPUTHPUNLULZ4L*7
®*+23%-6?.OH]LILLUYLSH[LK[V9;;JSPUPJHSMLH[\YLZ(^PKLWOLUV[`WPJHS
OL[LYVNLULP[`PZ^LSSRUV^UPU[OLKPZLHZL^OPJOPUJS\KLZH[SLHZ[THQVYKPMMLYLU[
JSPUPJHS WYLZLU[H[PVUZ PL [`WPJHS WYLZLY]LK ZWLLJO 7:= LHYS` ZLPa\YL ,:= HUK
JVUNLUP[HS]HYPHU[ZBD9LJLU[S`V\YNYV\WOHZZOV^ULY`[OYVJ`[LZOHWLJOHUNLZPU
[`WPJHS9;;WH[PLU[ZHUKÄUK[VILYLSH[LK[VTLTIYHULV_PKH[P]LZ[YLZZBDIL[[LY
JOHYHJ[LYPaL [OL LY`[OYVJ`[LZ PU 9;; H JSHZZPJHS WYV[LVTPJ HUHS`ZPZ ^HZ \ZLK [V
LZ[HISPZOHYLMLYLUJLTHWVMTLTIYHULWYV[LPUZPU[OLKPZLHZLHUKJVTWHYLK^P[O
[OLWYV[LPUZWH[[LYUPUOLHS[O`JVU[YVSZ7YLHUHS`[PJHSZ[\K`PUKPJH[LK[OH[[OLZ[VYHNL
I` MYLLaPUN H[ ‡* JHU JH\ZL WYV[LPU KLNYHKH[PVU ^P[O KPMMLYLU[ WYV[LPU WH[[LYU
WYVÄSL JVTWHYLK [V [OL YLMLYLUJL THW +PMMLYLU[PHS L_[YHJ[PVU WYVJLK\YLZ ^LYL ZL[
\W HSSV^PUN ZLWHYH[PVU VM PU[LNYHS TLTIYHUL WYV[LPUZ MYVT WLYPWOLYHS ZWLJPLZ HUK
[V KL[LYTPUL [OL TVZ[ HWWYVWYPH[L L_[YHJ[PVU TL[OVK [OVYV\NO HUHS`ZPZ VM WYV[LPU
WH[[LYUZPU9;;WH[PLU[Z^PSSILVMTHQVYOLSWPUKLÄUPUNVM[OLTVZ[LMÄJPLU[Z[YH[LN`
MVYPKLU[PM`PUNPUKP]PK\HSWYV[LPUZHUK[VPU]LZ[PNH[LVUWVZZPISLYLSH[PVUZOPWZIL[^LLU
LY`[OYVJ`[LWYV[LPUWH[[LYUZHUKJSPUPJHSWOLUV[`WLHUKVYKPZLHZLZL]LYP[`
9LMLYLUJLZ!
BD5L\S132H\MTHUU>,.SHaL+.L[HS(UU5L\YVS"! BD*PJJVSP3+L-LSPJL*7HJJHNUPUP,L[HS)PVJOPTPJHL[)PVWO`ZPJH(J[H"
;OLYLZLHYJOOHZILLUM\UKLKI`;\ZJHU`9LNPVUB¸(U[PV_PKHU[Ʒ7VS`\UZH[\YLK-H[[`(JPKSPWVPJHJPK
Z\WWSLTLU[H[PVUPU9L[[Z`UKYVTL!HUV]LSHWWYVHJO[V[OLYHW`¹D
007VZ[LY:LZZPVU
PRO 17
Proteine
:PUNSLYLZPK\LJVU[YPI\[PVU[V[OLZLSMHNNYLNH[PVUHUK
HJ[P]P[`VMHUHU[PM\UNHSPTT\UVNSVI\SPUKLYP]LKWLW[PKL
7,9;05/,A;/,34(*06*063(;,*3(*65;0:;,-(50(
76365,3303<*0(56:70:50(3),9;6
3HIVYH[VYPV*04;LJUVWVSV7HYTH"+PWHY[PTLU[VKP:JPLUaL)PVTLKPJOL
)PV[LJUVSVNPJOLL;YHZSHaPVUHSP¶:)P)P;<UP]LYZP[nKP7HYTH0[HSPH
(U[PTPJYVIPHSWLW[PKLZHYLPTWVY[HU[LMMLJ[VYTVSLJ\SLZVM[OLPUUH[LPTT\ULZ`Z[LT
7LW[PKLZ KLYP]LK MYVT [OL O\THU PTT\UVNSVI\SPU 0N WYV[LVS`ZPZ L_LY[ M\UNPJPKHS
HJ[P]P[` HNHPUZ[ *HUKPKH HSIPJHUZ PU ]P[YV HUK PU ]P]V ;OL KLJHWLW[PKL 52
58=:3;*3=2 KLYP]LK MYVT 0N. L_OPIP[LK ZPNUPÄJHU[ HJ[P]P[` ,* VM ¶ ›4HNHPUZ[KPMMLYLU[M\UNHSZ[YHPUZHUKZOV^LKUVOLTVS`[PJVYJ`[V[V_PJLMMLJ[ZVU
O\THUYLKISVVKJLSSZHUKTHTTHSPHUJLSSZ
52 ZLX\LUJL PZ JOHYHJ[LYPaLK I` [OL HS[LYUH[PVU VM O`KYVWOVIPJO`KYVWOPSPJ
YLZPK\LZHUKKPZWSH`LK[OLHIPSP[`[VZWVU[HULV\ZS`ZLSMHZZLTIS`PU[VYPJOơZOLL[
Z[Y\J[\YLPUHX\LV\ZZVS\[PVUVYPU[OLWYLZLUJLVM*HSIPJHUZ,SLJ[YVUTPJYVNYHWOZ
VM ULNH[P]LS` Z[HPULK 52 KLTVUZ[YH[LK [OL WYLZLUJL VM H UL[^VYR VM ÄIYPSSPRL
Z[Y\J[\YLZ 0U VYKLY [V JYP[PJHSS` LZ[HISPZO [OL YLSL]HUJL VM LHJO YLZPK\L PU [OL ZLSM
HNNYLNH[PVUHUKÄIYPSMVYTH[PVUWYVJLZZLZ^LHUHS`aLK52\ZPUNHSHUPULZJHUUPUN
I` *PYJ\SHY +PJOYVPZT -S\VYLZJLUJL :WLJ[YVZJVW` HUK ,SLJ[YVU HUK ;YHUZTPZZPVU
4PJYVZJVW`
;OL HSHUPUL Z\IZ[P[\[PVU ZOV^LK PUJYLHZLK \UHS[LYLK VY KLJYLHZLK JHUKPKHJPKHS
HJ[P]P[`PUJVTWHYPZVU[VWHYLU[HSWLW[PKL-VYPUZ[HUJLYLWSHJLTLU[VMJ`Z[LPUL[OH[
OHTWLYZ KPZ\SWOPKL IVUK MVYTH[PVU KL[LYTPULK [OL TVZ[ ZPNUPÄJHU[ YLK\J[PVU VM
JHUKPKHJPKHSHJ[P]P[`!PUMHJ[[OLHIZLUJLVMWYLHSPNUTLU[VM[OLTVUVTLYZYLK\JLZ
[OL LU[YVWPJ JVZ[ VM HZZLTIS` HUK JVUZLX\LU[S` [OL MVYTH[PVU VM PU[LYTVSLJ\SHY
/IVUKZ :\IZ[P[\[PVU VM [OL WVZP[P]LS` JOHYNLK YLZPK\L THRLZ [OL WLW[PKL OPNOS`
O`KYVWOVIPJSV^LYPUNP[ZZVS\IPSP[`HUKHJ[P]P[`
6]LYHSS ZLX\LUJL HUHS`ZPZ VM [OL WYVWLUZP[` VM JYVZZ ơHNNYLNH[PVU HTPUV HJPK
]VS\TL /IVUK WV[LU[PHS O`KYVWOVIPJP[` ^HZ L_WSVP[LK [V KL]PZL HU L_WSHUH[PVU
VM [OL JVU[YPI\[PVU VM PUKP]PK\HS YLZPK\L PU IV[O ZLSMHNNYLNH[PVU HUK JHUKPKHJPKHS
HJ[P]P[`
37VSVULSSP;*PVJPVSH>4HNSPHUP77AHULSSV;+»(KKH-+L)LYUHYKPZ:(YHUJPH,.HIYPLSSP,
7LYPJVSPUP(=LJJOPHYLSSP;(7LY[PUOLa(:WPZUP:*VU[P¸7LW[PKLZVM[OL*VUZ[HU[9LNPVUVM
(U[PIVKPLZ+PZWSH`-\UNPJPKHS(J[P]P[`¹7SVZ6ULL
007VZ[LY:LZZPVU
PRO 18
Proteine
5\JSLVWOVZTPUIPUKZ.X\HKY\WSL_YLNPVUZ
H[YPIVZVTHS+5(
*/0(9,33(::*(.3065,.336:;,9A6*+04(;;,6(
(9*6=0;6(-,+,90*03
+PWHY[PTLU[VKP:JPLUaL)PVJOPTPJOL¸:HWPLUaH¹<UP]LYZP[nKP9VTH9VTH
0Z[P[\[VKP)PVJOPTPJHL)PVJOPTPJH*SPUPJH
<UP]LYZP[n*H[[VSPJHKLS:HJYV*\VYLKP9VTH9VTH
*L:0*LU[YV:JPLUaLKLSS»0U]LJJOPHTLU[VL+PWHY[PTLU[VKP:JPLUaL:WLYPTLU[HSP
L*SPUPJOL<UP]LYZP[nKP*OPL[P¸.+»(UU\UaPV¹*OPL[P
5\JSLVWOVZTPU 574 PZ HU HI\UKHU[ WOVZWOVWYV[LPU [OH[ WSH`Z H RL` YVSL
PU YPIVZVTL IPVNLULZPZ 574 IPUKZ U\JSLPJ HJPKZ HUK OHZ PU[YPUZPJ 95(HZL HUK
JOHWLYVUL HJ[P]P[PLZ (S[OV\NO THPUS` SVJHSPaLK H[ U\JSLVSP 574 JVU[PU\V\ZS`
ZO\[[SLZIL[^LLU[OLU\JSL\ZHUK[OLJ`[VWSHZT[VM\SSÄSSP[ZM\UJ[PVUZ
;OLNLULLUJVKPUN574PZT\[H[LKPU¶VMUVYTHSRHYPV[`WLHJ\[LT`LSVPK
SL\RLTPH (43 WH[PLU[Z ( KPZ[PUJ[P]L MLH[\YL VM 574 T\[HU[Z PZ [OLPY Z[HISL HUK
HILYYHU[SVJHSPaH[PVUPU[OLJ`[VWSHZTVMSL\RLTPJJLSSZ(SST\[H[PVUZSLHK[VJYP[PJHS
JOHUNLZH[[OL574*[LYTPU\Z!PSVZZVM[Y`W[VWOHUYLZPK\LZHUK VY HSVUL^OPJOKLZ[HIPSPaLZ[OL*[LYTPUHSKVTHPU"PPNLULYH[PVUVMHUL^5,:TV[PM
^OPJO YLPUMVYJLZ [OL U\JSLHY L_WVY[ VM 574 WYV[LPU )V[O HS[LYH[PVUZ HYL JYP[PJHS
MVYWLY[\YIPUN574T\[HU[[YHMÄJHUKMVY[OL\UKLYS`PUNHILYYHU[HJJ\T\SH[PVUVM
574PU[OLJ`[VWSHZTVMSL\RLTPJJLSSZ
5\JSLVSPHYLJVTWSL_Z[Y\J[\YLZHZZLTISLKHYV\UK[HUKLTHYYH`ZVMY+5(NLULZHUK
OLYL^LPU]LZ[PNH[L[OLZ[Y\J[\YHSKL[LYTPUHU[ZVM574U\JSLVSHYSVJHSPaH[PVU>L
ZOV^[OH[HKVTHPULUJVTWHZZPUN[OLSHZ[YLZPK\LZVM[OLWYV[LPU574*
IPUKZ^P[OOPNOHMÄUP[`VSPNVU\JSLV[PKLZLX\LUJLZ^P[O.X\HKY\WSL_Z[Y\J[\YLMV\UK
H[[OLUVUJVKPUNZ[YHUKVMYPIVZVTHS+5(0TWVY[HU[S`[OLTVZ[JVTTVUSL\RLTPJ
574]HYPHU[*[LYTPUHSKVTHPUPZ\UMVSKLKHUKJVTWSL[LS`SVZLZ[OPZHJ[P]P[`;OPZ
PZ[OLJVUZLX\LUJLVM.X\HKY\WSL_IPUKPUNKVTHPUKLZ[HIPSPaH[PVUZPUJLT\[H[PVUZ
HPTLK H[ YLMVSKPUN [OL SL\RLTPJ ]HYPHU[ HSZV YLZ\S[ PU YLZJ\PUN [OL .X\HKY\WSL_
IPUKPUN HJ[P]P[` HUK U\JSLVSHY SVJHSPaH[PVU 0U JVUJS\ZPVU [OPZ ^VYR LZ[HISPZOLZ [OL
Z[Y\J[\YHSIHZLZMVY\UKLYZ[HUKPUN574U\JSLVSHYSVJHSPaH[PVUHUKP[ZPTWHPYTLU[I`
(43HZZVJPH[LKT\[H[PVUZ
007VZ[LY:LZZPVU
PRO 19
Proteine
7YV[LVTPJWYVÄSLVMILUPNU[\TV\YZVMWHYV[PKNSHUKZ
+65(+06,3,5(.0<:;03(<9(=,5;9650;0A0(5(+(=(33,@3,50(
:,**0(=,9650*(:,33(90-9(5*,:*/050:;,-(56
3<*(**/050(5;6506
+LWHY[TLU[VM7Z`JOPH[Y`5L\YVIPVSVN`7OHYTHJVSVN`HUK)PV[LJOUVSVN`
<UP]LYZP[`VM7PZH7PZH0[HS`
+LWHY[TLU[VM6[VYOPUVSHY`UNVSVN`Z[,5;<UP[(aPLUKH6ZWLKHSPLYV
<UP]LYZP[HYPH7PZHUH7PZH0[HS`
0U[OLNYLH[]HYPL[`VMILUPNU[\TV\YZVM[OLWHYV[PKNSHUKZHWYLVWLYH[P]LKPHNUVZPZ
[OH[KPZ[PUN\PZO[OLKPMMLYLU[PZV[`WLZPZVM[LUKPMÄJ\S[0UWHY[PJ\SHY>HY[OPU»Z[\TV\Y
ZOV^ZJSPUPJHSMLH[\YLZ[OH[KVUV[WYV]PKLHZ\MÄJPLU[IHZPZMVYHWYLJPZLKPHNUVZPZ
0KLU[PÄJH[PVU VM WV[LU[PHS WYVNUVZ[PJ IPVTHYRLYZ ^V\SK Z\YLS` PTWYV]L [OL YLX\PYLK
HJJ\YHJ` PU KPHNUVZPZ 7YV[LVTL HUHS`ZPZ OHZ ILLU ZOV^U [V IL H \ZLM\S [VVS MVY
Z[\K`PUN O\THU KPZLHZLZ HUK MVY PKLU[PM`PUN UL^ WYVNUVZ[PJ KPHNUVZ[PJ HUK
[OLYHWL\[PJTHYRLYZ!^L[OLYLMVYLKLJPKLK[VL_WSVYLWYV[LVTPJWH[[LYUVM[OLÄUL
ULLKSLHZWPYH[PVUÅ\PKVMWSLVTVYWOPJHKLUVTH7(HUK^HY[OPU»Z[\TV\Y>;
;OPY[`[^VWH[PLU[Z^LYLLUYVSSLK ^LYLJSHZZPÄLKHZ7(HZ>;0TTLKPH[LS`
HM[LYZ\YNPJHSYLTV]HSVM[OLNSHUKHÄULULLKSLIPVWZ`^HZTHKLVU[OL[\TV\YHS
UVK\SL HUK VU [OL Z\YYV\UKPUN [PZZ\L ;OPZ Å\PK ^HZ JLU[YPM\NLK HUK [OL WYV[LPUZ
^LYLWYLJPWP[H[LK\ZPUN[OL;*((JL[VULTL[OVK;^VWVVSZILSVUNPUN[V[OL7(HUK
>;ZHTWSLZ^LYLTHKLHUKLHJOWVVS^HZSVHKLKPU[YPWSPJH[L(M[LY+ZLWHYH[PVU
[OLWYV[LPUWH[[LYUZVM[OL7(HUK>;ZHTWSLZ^LYLJVTWHYLKI`\ZPUN7YVNLULZPZ
:HTL:WV[ 5VU3PULHY +`UHTPJZ [V ZLHYJO KPMMLYLU[PHSS` L_WYLZZLK WYV[LPU ZWV[Z
7YV[LPUZ VM PU[LYLZ[ ^LYL Z\ITP[[LK [V UHUV3*,:04:4: HUHS`ZPZ MVY WYV[LPU
PKLU[PÄJH[PVU([V[HSVMZWV[ZKPMMLYLU[PHSS`L_WYLZZLK^LYLMV\UK^P[OMVSK]HYPH[PVU
%(TVUN[OLWYV[LPUZPKLU[PÄLKI`THZZZWLJ[YVTL[Y`THU`PTT\UVNSVI\SPUZHZ
0NHSWOHJOHPU*YLNPVU0NRHWWHJOHPU*YLNPVU0NSHTIKHJOHPU*YLNPVUZ0N
RHWWHJOHPU*YLNPVUYLZ\S[LK[VIL\WYLN\SH[LKPU>;WVVS^P[OYLZWLJ[[VHKLUVTH
WYVIHIS`K\L[V[OLS`TWOVTH[VZ\TVYPNPUVM[OL>;[\TV\Y
6\Y WYLSPTPUHY` YLZ\S[Z TH` Z\NNLZ[ [OL WV[LU[PHS HWWSPJHIPSP[` VM [OL WYV[LVTPJ
HWWYVHJO[VKLÄULHWV[LU[PHSWHULSVMWYV[LPUIPVTHYRLYZ\ZLM\SPU[OLKPHNUVZPZVM
ILUPNU[\TV\YVMWHYV[PKNSHUKZ
007VZ[LY:LZZPVU
PRO 20
Proteine
0U[LYHJ[PVUVMIHJ[LYPHSNS\[H[OPVUL[YHUZMLYHZL^P[OOLTPU
(336*(;05,90564(:<33040*/,3,-,+,90*03<*(
+00306*(9405,
+PWHY[PTLU[VKP:JPLUaL:WLYPTLU[HSPL*SPUPJOLHUK*L:0*LU[YV:JPLUaL
KLSS»0U]LJJOPHTLU[V<UP]LYZP[nº.K»(UU\UaPV»*OPL[P0[HS`
.S\[H[OPVUL [YHUZMLYHZLZ .:;Z ,* HYL WP]V[HS LUa`TLZ MVY KL[V_PÄJH[PVU
WYVJLZZLZPUL\RHY`V[LZHUKWYVRHY`V[LZ.:;ZJH[HS`ZLNS\[H[OPVULJVUQ\NH[PVU
[VHIYVHKZWLJ[Y\TVMJVTWV\UKZ0UHKKP[PVUM\UJ[PVUZUV[YLSH[LK[VKL[V_PÄJH[PVU
OH]L ILLU KPZJV]LYLK MVY .:;Z PUJS\KPUN Z[LYVPK IPVZ`U[OLZPZ ZPNUHS [YHUZK\J[PVU
HUKPU[YHJLSS\SHY[YHUZWVY[
/LTPU OHZ ILLU ZOV^U [V IL OHaHYKV\Z [V [OL JLSS TLTIYHUL ILJH\ZL P[ PUZLY[Z
PU[V[OLSPWPKIPSH`LYHUKIPUKZ[VTLTIYHULWYV[LPUZ;OLZLPU[LYHJ[PVUZTH`JH\ZL
TLTIYHULJOHUNLZ^OPJOTH`SLHK[VJLSS\SHYS`ZPZ
;OL HIPSP[` VM ZL]LYHS L\RHY`V[PJ .:;Z MYVT WS\YPJLSS\SHY HUK \UPJLSS\SHY VYNHUPZTZ
[VPU[LYHJ[^P[OOLTPUOHZILLU^PKLS`PU]LZ[PNH[LKHUK[OPZJVTWV\UK^HZ
ZOV^U[VIPUK[OLLUa`TL^P[OOPNOHMÄUP[`
;V ]LYPM` PM [OPZ PU[LYHJ[PVU PZ H WYLYVNH[P]L VM L\RHY`V[PJ .:;Z VY P[ OHZ H NLULYHS
ZPNUPÄJHUJL ^L Z[\KPLK [OL IPUKPUN VM OLTPU [V 7YV[L\Z TPYHIPSPZ 7T.:; H
YLWYLZLU[H[P]LIHJ[LYPHS.:;ILSVUNPUN[V[OL)L[HJSHZZ;OLPU[LYHJ[PVU^HZZ[\KPLK
I` TLHUZ VM PUOPIP[PVU HZZH`Z HUK Å\VYLZJLUJL ZWLJ[YVZJVW`9LZ\S[Z VI[HPULK
PUKPJH[L [OH[ 7T.:; IPUKZ OLTPU LMÄJPLU[S` ^P[O H JVUZPKLYHISL X\LUJOPUN VM [OL
PU[YPUZPJ Å\VYLZJLUJL VM 7T.:; -\Y[OLYTVYL OLTPU ^HZ MV\UK [V IL H Z[YVUN
PUOPIP[VYVM.:;JVUQ\NH[PVUHJ[P]P[`
/H`LZ1+-SHUHNHU1<1V^ZL`09(UU\9L]7OHYTHJVS;V_PJVS!
(SSVJH[P5-LKLYPJP34HZ\SSP4+P0SPV* -,):1!
*HJJ\YP(4(JL[V(7PLTVU[L-+P0SPV*9VZH[V5-LKLYPJP. ,\Y1)PVJOLT ! 3PLIH\,+H^VVK2--HIYPUP9-PZJOLY9PLWL37LYIHUK[4:[LSSH37LKLYZLU1A)VJLKP(7L[YHYJH
7-LKLYPJP.9PJJP. 1)PVS*OLT! 007VZ[LY:LZZPVU
PRO 21
Proteine
*OHYHJ[LYPaH[PVUVM:[HWO`SVJVJJ\ZS\NK\ULUZPZIPVÄST
MVYTH[PVUPUKPMMLYLU[NYV^[OJVUKP[PVUZ!YVSLVM0ZK
WYV[LPUZVUIPVÄSTMVYTH[PVU
40::05,6(5;65056+076;6(5;65,33(*07633(305(-<.(AA(
(5.,3(),9;(05(=(3,5;05(=,5<;6(55<5A0(;(70,;96*63(
.0(470,96905+0:0465((5+:7,A0(3,70,;96
4VSLJ\SHY4LKPJPUL+LWHY[TLU[<UP]LYZP[`VM7H]PH¸(*HZ[LSSHUP¹
)PVJOLTPZ[Y`:LJ[PVU
: S\NK\ULUZPZ PZ H ZRPU JVTTLUZHS [OH[ HJ[Z HZ HU H[`WPJHSS` ]PY\SLU[ WH[OVNLU
JH\ZPUN PUMLJ[PVUZ ^OPJO YLZLTISL [OVZL JH\ZLK I` : H\YL\Z L_WLJPHSS` HJ\[L
HUKOPNOS`KLZ[Y\J[P]LJHZLZVMUH[P]L]HS]LLUKVJHYKP[PZ[OH[VM[LUYLX\PYLZ\YNPJHS
[YLH[TLU[PUHKKP[PVU[VHU[PTPJYVIPHS[OLYHW`0[PZHSZVYLZWVUZPISLMVYZL]LYHS[`WLZ
VMIPVÄSTYLSH[LKPUMLJ[PVUZ+LZWP[L[OLNLUL[PJWYLZLUJLVMPJH(+)*OVTVSVN\LZ
WVS`5HJL[`SNS\JVZHTPUL75(.PZUV[HTHQVYJVTWVULU[VM:S\NK\ULUZPZIPVÄST
TH[YP_ Z\NNLZ[PUN [OH[ IPVÄST PZ THPUS` JVTWVZLK I` WYV[LPUHJLV\Z MHJ[VYZ ;OL
NLUVTLZLX\LUJLHUHS`ZPZVM:S\NK\ULUZPZZ[YHPU5 ZOV^Z[OH[P[PZ[OLVUS`
Z[HWO`SVJVJJHSZWLJPLZV[OLY[OHU:H\YL\Z[OH[WVZZLZZLZHSVJ\ZLUJVKPUNMV\Y
PYVUYLN\SH[LKZ\YMHJLKL[LYTPUHU[WYV[LPUZ0ZK20ZK10ZK*0ZK)PU]VS]LKPUPYVU
HJX\PZP[PVUMYVTOHLTVNSVIPU0U[OPZZ[\K`HJVSSLJ[PVUVM:S\NK\ULUZPZZ[YHPUZ^HZ
HUHS`ZLK\WVU[OLPYIPVÄSTMVYTH[PVUHIPSP[`PUHYPJOTLKP\T;:);Y`W[PJ:V`)YV[O
VYPUPYVUZ[HY]H[PVU9740(SS[OLZ[YHPUZL_HTPULKTHRLHWYV[LPUHJLV\ZIPVÄST
ZLUZP[P]L[V[Y`WZPUWYV[LPUHZL2HUK+5HZL0[YLH[TLU[^OPSLZVKP\TTL[HWLYPVKH[L
HUKKPZWLYZPU)OH]LUVLMMLJ[;OLHKKP[PVUVM›4-L*SKPTPUPZOLZ[OLHTV\U[
VMIPVÄSTMVYTH[PVUPU9740Z\NNLZ[PUNHWVZZPISLYVSLVM0ZKWYV[LPUZPU[OLWYVJLZZ
;OLL_WYLZZPVUVM0ZKWYV[LPUZVUIHJ[LYP\TZ\YMHJL^HZKLTVUZ[YH[LKI`>LZ[LYU
ISV["TVYLV]LY0N.YHPZLKHNHPUZ[[OLTZPNUPÄJHU[S`KLJYLHZLPUHKVZLKLWLUKLU[
THUULY IPVÄST MVYTH[PVU ;OL KLSL[PVU VM [OL LU[PYL 0ZK SVJ\Z VY 0ZK* NLUL HSVUL
JH\ZLHZ[H[PZ[PJHSZPNUPÄJHU[KLJYLHZL7#VMIPVÄSTMVYTH[PVU^OPSL0ZK20ZK1
VY0ZK)KLSL[PVUZOH]LUVLMMLJ[VUP[;VNL[OLY[OLZLÄUKPUNZVMMLYPUZPNO[ZPU[V[OL
YVSLVM0ZKWYV[LPUZPUWHY[PJ\SHY0ZK*VU:S\NK\ULUZPZIPVÄSTMVYTH[PVU^OLU[OPZ
WH[OVNLUPZNYV^PUNPUPYVUZ[HY]H[PVUJVUKP[PVUZ
007VZ[LY:LZZPVU
PRO 22
Proteine
5HZ\/HRVSHKPZLHZL!HWPSV[Z[\K`MVY[OLYLZLHYJOVM
WVZZPISLIPVTHYRLYZ
.0<30(56:,9,5(465;(3),;;0369,5A(-3690(50(55(4
)(9+650(55(:(3=05096),9;(
+LW[VM4VSLJ\SHY4LKPJPUL<UP]LYZP[`VM7H]PH
¹
-VUKHaPVUL3H\YH-VZZH[P¹653<:
+LW[VM7\ISPJ/LHS[O`5L\YVZJPLUJL,_WLYPTLU[HSHUK-VYLUZPJ4LKPJPUL
<UP]LYZP[`VM7H]PH
7VS`J`Z[PJSPWVTLTIYHUV\ZVZ[LVK`ZWSHZPH^P[OZJSLYVZPUNSL\RVLUJLWOHSVWH[O`VY
5HZ\/HRVSH KPZLHZL 5/+ PZ H NLUL[PJHSS` OL[LYVNLULV\Z YLJLZZP]LS` PUOLYP[LK
KPZVYKLY BD ;OL TVSLJ\SHY NLUL[PJ IHJRNYV\UK VM 5/+ ^HZ JOHYHJ[LYPaLK I`
PKLU[PM`PUN T\[H[PVUZ PU [^V NLULZ +(7 VY ;9,4 +(7 PZ H [YHUZTLTIYHUL
HKHW[VYWYV[LPU[OH[PZHZZVJPH[LK^P[O[OLJLSSZ\YMHJLYLJLW[VY;9,4BD
;OLOPZ[VSVNPJHSOHSSTHYRZVM5/+HYLJ`Z[PJIVULSLZPVUZVZ[LVWVYV[PJMLH[\YLZHUK
SVZZVM^OP[LTH[[LYPU[OLIYHPUBD;OPZJVTIPUH[PVUVMZ`TW[VTZPZ\UPX\L[V[OPZ
KPZLHZL^OPJOHZPUKPJH[LKI`4VU[HSIL[[PL[HSBDPZ\UKLYLZ[PTH[LKPU0[HS`7H[PLU[Z
HYLSPRLS`[VNV\UYLJVNUPaLKWHY[S`ILJH\ZL[OL`HYLJVUZPKLYLK[VILHMMLJ[LKI`
V[OLY RPUKZ VM KLTLU[PH VY I` ÄIYV\Z K`ZWSHZPH HUK WHY[PHSS` ILJH\ZL [OL OPNOLZ[
WYL]HSLUJLVM5/+PZMV\UKPU-PUSHUKHUK1HWHUHUKPZNLULYHSS`\URUV^UV\[ZPKL
[OLZLJV\U[YPLZ
;VMHJPSP[H[L[OLYLJVNUP[PVUVM5/+WH[PLU[ZPZULJLZZHY`VI]PV\ZS`[VPKLU[PM`WVZZPISL
IPVTHYRLYZVM[OLKPZLHZL;V[OPZHWYV[LVTPJHWWYVHJO+,HUK4:^HZ\ZLK
VUWSHZTHZHTWSLZVMZP_PUKP]PK\HSZMYVTHU0[HSPHUMHTPS`+,HUHS`ZPZ^HZ[O\Z
WLYMVYTLKVUWSHZTHMYVTHZ\IQLJ[^P[OOVTVa`NV\ZT\[H[PVUPU;9,4Z\IQLJ[Z
^P[OOL[LYVa`NV\ZT\[H[PVUPU[OLZHTLNLULHUKOLHS[O`Z\IQLJ[Z
;OLJVTWHYH[P]LHUHS`ZPZZOV^LK[OLWYLZLUJLVMZVTLWYV[LPUZKPMMLYLU[S`L_WYLZZLK
;OL` ^LYL PKLU[PÄLK I` NLS TH[JOPUN HUK THZZ ZWLJ[YVTL[Y` ( ML^ PUÅHTTH[VY`
WYV[LPUZ ^LYL JSHZZPÄLK /HW[VNSVIPU (SWOHHJPK NS`JVWYV[LPU HUK V[OLYZ
PU]VS]LK PU KPMMLYLU[ UL\YVKLNLULYH[P]L KPZLHZLZ ;YHUZY[`YL[PU *S\Z[LYPU JV\SK IL
KL[LJ[LK
;OPZ^VYR^HZZ\WWVY[LKI`¹-VUKHaPVUL3H\YH-VZZH[P¹653<:
BD=LYSVLZ(L[HS14LK.LUL[ "!
BD7HSVUL]H1L[HS5H[.LUL["!
BD7HSVUL]H1L[HS(T1/\T.LUL["!
BD4VU[HSIL[[P3L[HS-\UJ[5L\YVS" ! 007VZ[LY:LZZPVU
PRO 23
Proteine
(7YV[LVTPJ(WWYVHJO;V:[\K`4HSPNUHU[7SL\YHS
4LZV[OLSPVTH
+(=(33,@3,50(.0<:;03(<9(+65(+06,3,5(*09,.0(-,+,90*(
=,5;9650;0A0(5()656;;0(3,::(5+9(-6++0:9<+@
*90:;(<+6(3-65:63<*(**/050(5;6506
+LWHY[TLU[VM7Z`JOPH[Y`5L\YVIPVSVN`7OHYTHJVSVN`HUK)PV[LJOUVSVN`
<UP]LYZP[`VM7PZH7PZH0[HS`
6JJ\WH[PVUHS4LKPJPULVM<UP]LYZP[`/VZWP[HSVM7PZH0[HS`
+LWHY[TLU[VM,UKVJYPUVSVN`HUK4L[HIVSPZT6Y[OVWHLKPJZHUK;YH\TH[VSVN`
<UP]LYZP[`VM7PZH0[HS`
4HSPNUHU[TLZV[OLSPVTHPZHOPNOS`HNNYLZZP]LJHUJLY\UYLZWVUZP]L[VJOLTV[OLYHW`
YHKPV[OLYHW`VYZ\YNPJHSYLZLJ[PVUHUKP[JHUHMMLJ[HU`RPUKVMZLYVZHSTLTIYHUL
WSL\YH WLYP[VUL\T WLYPJHYKP\T HUK L]LU [OL [\UPJH ]HNPUHSPZ [LZ[PZ 4HSPNUHU[
WSL\YHS TLZV[OLSPVTH 474 PZ H YHYL [OVYHJPJ THSPNUHUJ` [OH[ PZ KPYLJ[S` SPURLK
[V HZILZ[VZ L_WVZ\YL ZV [OL THSL NLUKLY PZ HZZVJPH[LK ^P[O [OL THQVYP[` VM 474
KPHNUVZLZ 474 YLWYLZLU[Z SLZZ [OHU VM RUV^U JHUJLYZ I\[ P[Z PUJPKLUJL ^PSS
JVU[PU\L[VPUJYLHZLZPNUPÄJHU[S`^P[OZL]LYLW\ISPJOLHS[OJVUZLX\LUJLZ
>L HWWSPLK H WYV[LVTPJ HWWYVHJO [V KLÄUL WYV[LVTPJ WYVÄSLZ VM 474 [PZZ\LZ
)PVWZPLZ^LYLVI[HPULKMYVTWH[PLU[ZHUKJSHZZPÄLKPU474U$HUKO`WLYWSHZPH
U$7YV[LPUL_[YHJ[Z^LYLZLWHYH[LK\ZPUN[^VKPTLUZPVUHSLSLJ[YVWOVYLZPZ(M[LY
JVTWHYPZVUVM474HUKO`WLYWSHZPHWYV[LPUWYVÄSLZH[V[HSVMWYV[LPUZWV[Z^LYL
MV\UK[VILKPMMLYLU[PHSS`L_WYLZZLKLHJOL_OPIP[PUN™MVSKJOHUNLLP[OLYPUJYLHZL
VY KLJYLHZL VM TLHU ]HS\L ZWV[ PU[LUZP[` :WV[Z VM PU[LYLZ[ ^LYL J\[ HUK WYV[LPUZ
PKLU[PÄLK I` UHUV3*,:04:4: *HSYL[PUPU 7YLSHTPU (* +LZTPU -Y\J[VZL
IPZWOVZWOH[L HSKVSHZL ( T`VZPU SPNO[ JOHPU T`VZPU SPNO[ JOHPU ]LU[YPJ\SHY
JHYKPHJT\ZJSLPZVMVYTT`VZPUSPNO[JOHPU)HUK*YLH[PULRPUHZL4[`WLL_OPIP[LK
™ MVSKJOHUNL IL[^LLU 474 HUK O`WLYWSHZPH ZHTWSLZ ;OL KPMMLYLU[ L_WYLZZPVU
^HZ]HSPKH[LKI`^LZ[LYUISV[HUHS`ZPZHUK^HZJVTWHYLK^P[O*HYJPUVTHZHTWSLZ
U$HZHWVZP[P]LJVU[YVS;OPZPZ[OLÄYZ[Z[\K`WLYMVYTLKVU474IPVWZPLZHUK
ZOV^Z[OLWV[LU[PHSP[`VMWYV[LVTPJHWWYVHJO[VKLÄULHWV[LU[PHSWHULSVMWYV[LPU
IPVTHYRLYZ\ZLM\SPU[OLKPHNUVZPZVM474
;OPZ YLZLHYJO ^HZ Z\WWVY[LK PU WHY[ I` NYHU[ MYVT [OL ¸4PUPZ[LYV :HS\[L 9PJLYJH
-PUHSPaaH[H ¹
007VZ[LY:LZZPVU
PRO 24
Proteine
7V[LU[PHS[O`YVPKJHUJLYIPVTHYRLYZMYVTÄULULLKSL
HZWPYH[PVUPU]P]VHUKPUWVZ[Z\YNLY`SLZPVUZ
*09,.0(-,+,90*(.0<:;03(<9(+(=(33,@3,50(9(.6;0A0(5(
50**63(0-0307764630:(56(5.,36+0*6:*06.0(5*(9360(**650
70,;96;65(**/,9(4(::0463<*(**/050(5;6506
+LWHY[TLU[VM7Z`JOPH[Y`5L\YVIPVSVN`7OHYTHJVSVN`HUK)PV[LJOUVSVN`
<UP]LYZP[`VM7PZH"7PZH0[HS`+LWHY[TLU[VM,UKVJYPUVSVN`HUK4L[HIVSPZT
6Y[OVWLKPJZHUK;YH\TH[VSVN`6JJ\WH[PVUHS4LKPJPUL<UP]LYZP[`VM7PZH7PZH0[HS`
+LWHY[TLU[VM6UJVSVN`:LJ[PVUVM*`[VWH[OVSVN`<UP]LYZP[`VM7PZH7PZH0[HS`
+LWHY[TLU[VM:\YNLY`<UP]LYZP[`VM7PZH7PZH0[HS`
;OLHUU\HSPUJPKLUJLVM[O`YVPKJHUJLYOHZILLUYLWVY[LK[VYHUNLIL[^LLUHUK
JHZLZWLYPUTLUHUKIL[^LLUHUKJHZLZWLYPU^VTLU
-PULULLKSL HZWPYH[PVU -5( IPVWZ` PZ [OL [LJOUPX\L \Z\HSS` HWWSPLK [V KPZ[PUN\PZO
ILUPNU MYVT THSPNUHU[ [O`YVPK UVK\SLZ /V^L]LY J`[VSVNPJHS HUHS`ZPZ JHU»[ HS^H`Z
HSSV^ H WYVWLY KPHNUVZPZ VM [O`YVPK UVK\SLZ ;OLYLMVYL HU PTWYV]LTLU[ VU [OL
KPHNUVZ[PJJHWHIPSP[`VM-5(JV\SKH]VPK\UULJLZZHY`[O`YVPKLJ[VT`
0UHWYL]PV\ZZ[\K`^LL_HTPULK-5(JVSSLJ[LKPTTLKPH[LS`HM[LY[V[HS[O`YVPKLJ[VT`
>L PU]LZ[PNH[LK [OL NSVIHS JOHUNLZ VM -5( WYV[LPU WH[[LYUZ VM [^V ]HYPHU[Z VM
THSPNUHU[ WHWPSSHY` [O`YVPK JHUJLY 7;*! JSHZZPJHS ]HYPHU[ 7;* J7;* HUK [HSS JLSS
]HYPHU[ 7;* ;*= +PMMLYLUJLZ PU WYV[LPU L_WYLZZPVU IL[^LLU JHUJLY HUK JVU[YVSZ
^LYLPKLU[PÄLK\ZPUN[^VKPTLUZPVUHSLSLJ[YVWOVYLZPZHUKTHZZZWLJ[YVTL[Y`>P[O
[OLWYLZLU[^VYR^L[LZ[LK[OLWVZZPIPSP[`[V[YHUZMLY[OLZLÄUKPUNZVUHSHYNLYU\TILY
VMWYLZ\YNPJHS[O`YVPK-5(ZHTWSLZ>LJVSSLJ[LK-5(PU]P]V-VYLHJOZHTWSL
HJ`[VSVNPJHSHUHS`ZPZ^HZWLYMVYTLKHUKHOPZ[VSVNPJHSHUHS`ZPZ^HZHSZVJHYYPLKV\[
MVY[OLWH[PLU[Z[OH[\UKLY^LU[Z\YNPJHSPU[LY]LU[PVU-5()^LYLJSHZZPÄLKHZILUPNU
J7;*HUK;*=
>L L]HS\H[LK I` ,30:( HZZH` HUK ^LZ[LYU ISV[ HUHS`ZPZ SL]LSZ VM Ä]L WYV[LPUZ
WYL]PV\ZS` MV\UK \WYLN\SH[LK PU JHUJLY YLZWLJ[ [V JVU[YVSZ" [OLZL WYV[LPUZ ^LYL!
3SHJ[H[L KLO`KYVNLUHZL ) JOHPU -LYYP[PU OLH]` JOHPU -LYYP[PU SPNO[ JOHPU (UUL_PU
(HUK4VLZPU
>LJVUÄYTLK[OLZPNUPÄJHU[KPMMLYLUJLZPUL_WYLZZPVUVM[OLZLWYV[LPUZZ\WWVY[PUN
[OLKPHNUVZ[PJYLSL]HUJLVM[OLZLW\[H[P]LIPVTHYRLYZ4VYLV]LY[HRLU[VNL[OLYV\Y
Z[\KPLZPUKPJH[L[OH[[O`YVPK-5(HUHS`ZPZPZH\ZLM\S[VVSPU[OLKPHNUVZ[PJJSHZZPÄJH[PVU
VM[OLKPZLHZL
;OPZ YLZLHYJO ^HZ Z\WWVY[LK I` NYHU[ MYVT [OL ¸7YVNL[[V :HS\[L 9LNPVUL
;VZJHUH¹
.P\Z[P3L[HS17YV[LVTL9LZ" ! 007VZ[LY:LZZPVU
PRO 25
Proteine
/VTVJ`Z[LPULPUK\JLZPUZ\SPUÄIYPSSH[PVUHUKSVZZVM
M\UJ[PVU!(WVZZPISLSPUR^P[OIV[OPUZ\SPUYLZPZ[HUJLHUK
J\[HULV\ZHT`SVPKVZPZ
7(6307(636 *(:,330(55(*09907(636:(5;0(30*,3<;0
:0465,.069.0*3(<+0(*06-0:0465,:)9(5(-9(5*,:*(
;090)0330)9<56*(40*0.<0+6
+LWHY[TLU[VM)PVJOLTPJHS:JPLUJLZ<UP]LYZP[`VM-SVYLUJL=PHSL4VYNHNUP-PYLUaL
+LWHY[TLU[VM*OLTPZ[Y`<UP]LYZP[`VM-SVYLUJL=PHKLSSHSHZ[Y\JJPH :LZ[V-UV
*590Z[P[\[VKP)PVÄZPJH]PHKL4HYPUP .LUV]H0Z[P[\[VKLP:PZ[LTP*VTWSLZZP·
0Z[P[\[V5HaPVUHSLKLSSL9PJLYJOL]PH4HKVUUHKLSWPHUV :LZ[V-UV
(T`SVPKVZLZ HYL WH[OVSVNPLZ JOHYHJ[LYPaLK I` L_[YH VY PU[YHJLSS\SHY ÄIYPSSHY WYV[LPU
HJJ\T\SH[PVU 7YV[LPU ÄIYPSSH[PVU PUP[PH[LZ ^OLU MVSKLK WYV[LPUZ HYL KLZ[HIPSPaLK
HKVW[PUN UVUUH[P]L JVUMVYTH[PVUZ TVYL WYVUL [V HNNYLNH[L ;OL HTVYWOV\Z
HNNYLNH[LZL]VS]L[VSLHKOPNOVYKLYLKÄIYPSZ
(WHY[PJ\SHYMVYTVMZ\IJ\[HULV\ZHT`SVPKVZPZ^HZVIZLY]LKPUKPHIL[PJZ^OVPUQLJ[
PUZ\SPU YLWLH[LKS` PU [OL ZHTL ZRPU ZP[L ;OL HUHS`ZLZ WLYMVYTLK VU [OL UVK\SLZ
MVYTLKH[[OLPUQLJ[PVUZP[LZOV^LK[OLWYLZLUJLVMHT`SVPKÄIYPSZJVU[HPUPUNPUZ\SPU
I\[Z\IQLJ[ZKPKUV[KL]LSVWZPNUZVMZ`Z[LTPJHT`SVPKVZPZ<W[VUV^[OLJH\ZLZ
[OH[PUK\JL[OL[YHUZMVYTH[PVUVM[OLUH[P]LZ[Y\J[\YLVMPUZ\SPUPU[OLÄIYPSSHYMVYTHYL
\URUV^U
:VTL H\[OVYZ KLTVUZ[YH[LK [OH[ PUZ\SPU KV^UYLN\SH[LZ [OL L_WYLZZPVU VM
J`Z[H[OPVUPUL ơZ`U[OHZL MH]V\YPUN [OL HJJ\T\SH[PVU VM OVTVJ`Z[LPUL /J` HU
HTPUVHJPK NLULYH[LK PU [OL WH[O^H` VM J`Z[LPUL Z`U[OLZPZ /LUJL [OL JOYVUPJ
L_JLZZ VM JPYJ\SH[PUN PUZ\SPU MYLX\LU[S` VIZLY]LK PU KPHIL[PJZ JV\SK JVU[YPI\[L [V
KL]LSVW O`WLYOVTVJ`Z[LPULTPL ^OPJO PZ HZZVJPH[LK ^P[O PUZ\SPU YLZPZ[HUJL ;OPZ
Z\NNLZ[Z[OH[OPNOWSHZTHSL]LSZVM/*`PU[LYMLYL^P[O[OLPUZ\SPUM\UJ[PVU
6\Y L_WLYPTLU[Z KLTVUZ[YH[LK [OH[ /*` IPUKZ [V PUZ\SPU HUK [OH[ [OPZ JVTWSL_
KPZWSH`ZHSV^LYHJ[P]P[`[OHU[OH[VMUH[P]LPUZ\SPUVU[OLPUZ\SPUYLJLW[VY0UHKKP[PVU
^L VIZLY]LK [OH[ /J` JH\ZLZ H WHY[PHS PUZ\SPU \UMVSKPUN MH]V\YPUN P[Z HNNYLNH[PVU
;OLVI[HPULKHNNYLNH[LZIPUK;OPVÅH]PUL;HUK*VUNV9LKHUKHYLJOHYHJ[LYPaLKI`
HUOPNOơZOLL[JVU[LU[-PUHSS`(-4HUHS`ZPZVM/*`[YLH[LKPUZ\SPUKLTVUZ[YH[LK
[OH[[OLZLHNNYLNH[LZJVU[HPU^VYTSPRLWYV[VÄIYPSZ;OLZLÄUKPUNZZ\NNLZ[[OH[/J`
JVU[YPI\[L[V[OLKL]LSVWTLU[VMPUZ\SPUYLZPZ[HUJLPU]P]V>L[OPUR[OH[OPNOPUZ\SPU
SL]LSZK\L[VYLWLH[LKPUZ\SPUPUQLJ[PVUPU[OLZHTLZP[LKL[LYTPULHSVJHSPUJYLHZLVM
/J`SL]LS[OH[PU[\YUSLHKZ[VPUZ\SPUHNNYLNH[PVU
>VYRZ\WWVY[LKI`[OL-VUKHaPVUL*HZZHKP7PZ[VPHHUK7LZJPHMVYM\UKPUN
007VZ[LY:LZZPVU
PRO 26
Proteine
*,9(;673(;(505(5+*,9(;6767<30505+<*,
4(72(*;0=(;0650573(5,(5+(9()0+67:0:3,(=,:
3<;0:)9<56.)(**,3300),95(9+094(9;,33050-
:*(3(((5+7(AA(.303
+PWHY[PTLU[VKP:JPLUaL)PVJOPTPJOL<UP]LYZP[nKLNSP:[\KP=PHSL4VYNHNUP
-PYLUaL0[HS`,U[VTVUZHZ]PHKP9PWVSP-PYLUaL
0[HS`+PWHY[PTLU[VKP)PVSVNPHKLSSL7PHU[L(NYHYPL<UP]LYZP[nKLNSP:[\KP=PHKLSSH
7PHNNL7PZH0[HS`+PWHY[PTLU[VKP)PV[LJUVSVNPL(NYHYPL<UP]LYZP[nKLNSP
:[\KP=PHKLSSH3HZ[Y\JJPH :LZ[V-PVYLU[PUV0[HS`
*LYH[VWSH[HUPU*7HUKJLYH[VWVW\SPU7VWHYLZTHSSWYV[LPUZWYVK\JLKI`[OL
WO`[VWH[OVNLUPJM\UNP*LYH[VJ`Z[PZWSH[HUPHUK*WVW\SPJVSHYLZWLJ[P]LS`*7HUK
7VW ILOH]L HZ 7(47Z 7H[OVNLU HZZVJPH[LK 4VSLJ\SHY WH[[LYU ZPUJL [OL` LSPJP[
[`WPJHSKLMLUZLYLZWVUZLZPU]HYPV\ZOVZ[HUKUVUOVZ[WSHU[ZZ\JOHZ[OLPUK\J[PVU
VM Z`U[OLZPZ VM WO`[VHSL_PUZ HUK [OL PUK\J[PVU VM [OL O`WLYZLUZP[P]L YLZWVUZL [OH[
J\STPUH[LZ^P[OHWYVNYHTTLKJLSSKLH[OVM[OLPUMLJ[LK[PZZ\LZ*7Z[Y\J[\YLOHZ
ILLUZVS]LKI`549HUKP[PZHKV\ISLƶơIHYYLSWYV[LPU7VWOHZHZPTPSHYMVSKI\[
JPYJ\SHYKPJOYVPZTZOV^ZKPMMLYLUJLZPU[OLZLJVUKHY`Z[Y\J[\YL9LJLU[S`P[OHZILLU
ZOV^U[OH[*7HUK7VWHYLHISL[VPUK\JL/6HUK56Z`U[OLZPZHUKLUOHUJL
[OL [YHUZJYPW[PVU VM ZL]LYHS WYV[LPUZ YLSH[LK [V KLMLUZL Z\JO HZ! 79 \URUV^U"
79 ơ NS\JHUHZL 79 [OH\TH[PU (7? (ZJVYIH[L WLYV_PKHZL ;OP- WYV[LPU
HKLU`S`S[YHUZMLYHZLHUK[OL>92@[YHUZJYPW[PVUMHJ[VY
;OLHPTVM[OPZ^VYRPZ[VPU]LZ[PNH[L[OLHJ[P]H[PVUVMHUV[OLY[`WPJHSKLMLUJLYLZWVUZL!
[OLHJ[P]H[PVUVM4(72JHZJHKLPUOVZ[WSHULHUKUVUOVZ[(YHIPKVWZPZSLH]LZ
;OL SLH]LZ ^LYL [YLH[LK ^P[O UTVS ›3 VM *7 HUK 7VW (M[LY TPU TPU
HUKHUKOV\YZVMPUJ\IH[PVUWSHULSLH]LZ^LYLZ\IQLJ[LK[VWYV[LPU
L_[YHJ[PVUI`H[OYLLZ[LWZTL[OVK\ZPUNHJL[VULHUKWOLUVS:HTWSL^LYLHUHS`aLK
I`:+:7(.,HUK^LZ[LYUISV[\ZPUNHU[P7OVZWOV,92O\THUHU[PIVKPLZ^OPJO
ZWLJPÄJHSS`YLJVNUPaLK[OL4(72HUK4(72VMWSHU[Z7YV[LPU^LYLL_[YHJ[LKMYVT
(YHIPKVWZPZ[YLH[LKSLH]LZI`;90:I\MMLYJVU[HPUPUN[^LLUHUKWYV[LHZLPUOPIP[VYZ
:HTWSLZ^LYLHZZH`LKHZHIV]LYLWVY[LK9LZ\S[ZZOV^[OLHJ[P]H[PVUVM4(72
HM[LYOV\YZVMPUJ\IH[PVUPU*7[YLH[LKSLH]LZHUKHM[LY OV\YZPU7VW[YLH[LK
SLH]LZ9LZ\S[ZJSLHYS`ZOV^[OH[*7HUK7VWKL[LYTPULHJ[P]H[PVUVM4(72Z^OPJO
WLJ\SPHY[VKLMLUZLYLZWVUZLZLP[OLYPUWSHU[HUKPUHUPTHS[PZZ\LZ
007VZ[LY:LZZPVU
PRO 27
Proteine
7YV[LPUTHJOPULY`PU]VS]LKPUJ`[JTH[\YH[PVU
PUNYHTULNH[P]LIHJ[LYPH
,=(+0:03=06(+,3,+04(;;,6*(936;9(=(.3050(336*(;,330
+PW:JPLUaL)PVJOPTPJOL¸(9VZZP-HULSSP¹:HWPLUaH<UP]LYZP[nKP9VTHHUK
0Z[P[\[VKP)PVSVNPHL7H[VSVNPH4VSLJVSHYPKLS*599VTH
;OLJ[`WLJ`[VJOYVTLZHYL^PKLZWYLHKLSLJ[YVUJHYYPLYWYV[LPUZMV\UKPUHSSKVTHPUZ
VM SPML HUK HYL JOHYHJ[LYPaLK I` H JV]HSLU[S` IV\UK OLTL -LWYV[VWVYWO`YPU 0?
SPURLK[V[OL*??*/TV[PMVM[OLHWVWYV[LPU0U[OLO\THUWH[OVNLU7ZL\KVTVUHZ
HLY\NPUVZH[OLIPVNLULZPZVM*`[JPZTLKPH[LKI`*JTWYV[LPUZ*JT()*+,-./0
JHSSLK:`Z[LT0>LOH]LHSYLHK`JOHYHJ[LYPaLK7H*JT.HUK7H*JT/IV[OPU]VS]LK
PU [OPVYLK\J[PVU WH[O^H` VM HWVJ`[J MYVT H M\UJ[PVUHS HUK Z[Y\J[\YHS WVPU[VM
]PL^>LHYLUV^Z[\K`PUN7H*JT0HUK7H*JT-[^VWYV[LPUZPU]VS]LKPU[OLÄUHS
Z[LWZVMOLTLHWVJ`[JSPNH[PVUWH[O^H`HUKWYVIHIS`MVYTPUNHT\S[PZ\I\UP[OLTL
SPNH[PVU JVTWSL_ [VNL[OLY ^P[O 7H*JT/ /V^ [OL *JT-/0 JVTWSL_ YLJVNUPaLZ P[Z
Z\IZ[YH[LZHWVJ`[HUKOLTLYLTHPUZ\URUV^U)PVPUMVYTH[PJHUHS`ZLZZ\NNLZ[[OH[
[OLWLYPWSHZTPJKVTHPUVM7H*JT0PZJVTWVZLKVM[OYLL;79TV[PMZ[L[YH[YPJVWLW[PKL
YLWLH[WYVIHIS`PU]VS]LKPU[OLYLJVNUP[PVUVMHWVJ`[JHUKHUƠơKVTHPUH[[OL
*[LYTPU\Z0[OHZILLUWYVWVZLK[OH[7H*JT0TH`PU[LYHJ[^P[OHWVJ`[[OYV\NO[OL
;79TV[PMZHUKWYLZLU[P[[V*JT-/JVTWSL_YLZWVUZPISLMVY[OLMVYTH[PVUVM[OL
JV]HSLU[IVUKZ^P[OOLTL7H*JT0OHZILLUJSVULKHUKW\YPÄLKPUHZVS\ISLZ[H[L!
M\UJ[PVUHSJOHYHJ[LYPaH[PVU\ZPUNLX\PSPIYP\THUKRPUL[PJÅ\VYLZJLUJLZWLJ[YVZJVW`PZ
PUWYVNYLZZ[VKLTVUZ[YH[L[OLIPUKPUN^P[OHWVJ`[JHUKZ`U[OL[PJWLW[PKLZTPTPJRPUN
[OL*[LYTPUHSWVY[PVUVMHWVJ`[J*Y`Z[HSZVM[OLW\YPÄLK7H*JT0WYV[LPUOH]LILLU
VI[HPULK" ^L HYL J\YYLU[S` PU]VS]LK PU WYVK\J[PVU VM ZLSLUVTL[OPVUPUL Z\IZ[P[\[LK
*JT0JY`Z[HSZPUVYKLY[VZVS]LP[Z+Z[Y\J[\YL
7H*JT- PZ H SHYNL PU[LNYHS TLTIYHUL WYV[LPU ^P[O ZTHSS WLYPWSHTH[PJ KVTHPUZ
)PVPUMVYTH[PJZHUHS`ZPZTL[OVKZ\NNLZ[Z[OH[P[JVU[HPUZ[YHUZTLTIYHULOLSPJLZ
PU7HLY\NPUVZH0[OHZILLUWYVWVZLK[OH[*JT-TH`HJ[HZOLTLYLK\J[HZLHUK
*`[JZ`U[OHZL>LHYLJ\YYLU[S`MVJ\ZPUNVU[OLM\UJ[PVUHSYVSLWSH`LKI`[OLSHYNLZ[
ZVS\ISLWLYPWSHZTPJKVTHPUVM[OPZTLTIYHULWYV[LPU7YLSPTPUHY`YLZ\S[ZZ\NNLZ[[OH[
[OPZYLZPK\LZKVTHPUYLJVNUPaLZHUKIPUKZ[V[OLOLTLNYV\W(S[VNL[OLY[OLZL
YLZ\S[ZTH`ZOLKSPNO[[VV\Y\UKLYZ[HUKPUN[OLJ`[VJOYVTLJIPVNLULZPZWYVJLZZPU
IHJ[LYPH
007VZ[LY:LZZPVU
PRO 28
Proteine
*OHYHJ[LYPaH[PVUVMUV]LSHU[PTPJYVIPHSWLW[PKLZPZVSH[LK
MYVT[OLZHSP]HVM7VSPZ[LZKVTPU\S\Z
4(9;,33050-)9<56.7(5;,9()3<;0:7(.50,
*(99,:03;<9033(AA0:*(77<.0.(5+7(AA(.303
+PWHY[PTLU[V:JPLUaL)PVJOPTPJOL<UP]LYZP[nKLNSP:[\KPKP-PYLUaL0[HS`
:J\VSHZ\WLYPVYL:(UUH7PZH0[HS`
+PWHY[PTLU[V)PVSVNPH,]VS\aPVUPZ[PJH<UP]LYZP[nKLNSP:[\KPKP-PYLUaL0[HS`
,U[VTVUZHZ]PHKP9PWVSP-PYLUaL0[HS`
;OLWYVK\J[PVUVMHU[PTPJYVIPHSWLW[PKLZ(47ZPZVULVM[OLTVZ[PTWVY[HU[PUUH[L
KLMLUZL TLJOHUPZTZ HKVW[LK I` WYVRHY`V[LZ HUK L\RHY`V[LZ HNHPUZ[ PUMLJ[PVUZ
JH\ZLK I` TPJYVYNHUPZTZ 6U [OL IHZPZ VM [OLPY Z[Y\J[\YHS MLH[\YLZ (47Z HYL
JSHZZPÄLK PU[V MV\Y JSHZZLZ! P SPULHY WLW[PKLZ KL]VPK VM J`Z[LPULZ HUK MVYTPUN
ƠOLSPJLZZ\JOHZJLJYVWPUZ"PPWLW[PKLZ^P[OHUV]LYYLWYLZLU[H[PVUPUWYVSPULHUK
VY NS`JPUL YLZPK\LZ" PPP WLW[PKLZ Z[HIPSPaLK I` PU[YHTVSLJ\SHY KPZ\SÄKL IVUKZ Z\JO
HZKLMLUZPUZ"P]WLW[PKLZJVTWVZLKI`YHYLHUKTVKPÄLKHTPUVHJPKZ4HU`(47Z
OH]LILLUMV\UKPUPUZLJ[Z"[OL`HYL\Z\HSS`ZTHSSJH[PVUPJTVSLJ\SLZJOHYHJ[LYPaLK
I`HSV^TVSLJ\SHY^LPNO[VM[LUMVYTPUNHTWOPWH[OPJZ[Y\J[\YLZWYLZLU[IV[OPU[OL
OLTVS`TWOHUKPU[OLZHSP]H
/LYL^LYLWVY[[OLPZVSH[PVUVM7VS(HUK7VS)[^V(47ZMYVT[OLSHY]HSZHSP]HVM[OL
ZVJPHS^HZW7VSPZ[LZKVTPU\S\ZH]LY`JVTTVUZWLJPLPU,\YVWL4PJYVIPVSVNPJHS
[LZ[ZWYV]LK[OH[SHY]HSZHSP]HVM7KVTPU\S\ZOHZHUHU[PIPV[PJHJ[P]P[`HNHPUZ[)HJPSS\Z
Z\I[PSPZ .YHT HUK ,ZJOLYPJOPH JVSP .YHT ;OLZL HU[PTPJYVIPHS WLW[PKLZ ^LYL
W\YPÄLKI`97/73*HUKJOHYHJ[LYPaLKI`WLW[PKLZLX\LUJPUNHUKTHZZZWLJ[YVTL[Y`
7VS(OHZHTVSLJ\SHY^LPNO[VM +HHUKPZJVTWVZLKVMHTPUVHJPKZ0[
PZ YPJO PU WYVSPUL HUK P[Z ZLX\LUJL OHZ ILLU JVTWSL[LS` KL[LYTPULK 7VS) ^P[O H
THZZVM+HPZMVYTLKMYVTHTPUVHJPKZHUKOHZILLUHSTVZ[JVTWSL[LS`
ZLX\LUJLK0[ILSVUNZ[V[OLNYV\WVMKLMLUZPUZHUKZOV^ZZVTLZPTPSHYP[PLZ^P[OH
W\[H[P]L KLMLUZPU VM [OL KPW[LY *\SPJVPKLZ ZVUVYLUZPZ 0U V\Y Z[\KPLZ ^L L]HS\H[LK
TPUPT\TPUOPIP[VY`JVUJLU[YH[PVU40* VM7VS(JVUJLYUPUN)Z\I[PSPZNYV^UVU
3)<UKLY[OLL_WLYPTLU[HSJVUKP[PVUZ\ZLK^LMV\UK[OH[40* PZ TNT3
[O\ZPUKPJH[LZ[OH[WVS(WYLZLU[ZHYLTHYRHISLHJ[P]P[`HNHPUZ[.YHT
([OVYV\NOJOHYHJ[LYPaH[PVUVM[OLWYVWLY[PLZVM[OLZLWLW[PKLZJV\SKIYPUNZPNUPÄJHU[
NVHSZPU[OLYHWL\[PJ
;OLJVTWSL[LJOHYHJ[LYPaH[PVUVM[OLWLW[PKLZ^PSSIYPUN[VHUPUJYLHZLVMHU[PTPJYVIPHS
TVSLJ\SLZ[VIL[LZ[LKHZUL^[OLYHWL\[PJHNLU[Z
007VZ[LY:LZZPVU
PRO 29
Proteine
4LTIYHULJOVSLZ[LYVSPZHRL`TVK\SH[VYVM[OL
LUKVJHUUHIPUVPKO`KYVSHZL-((/
+(05,:,,590*6+,-()90;00:.0(550:()(;<**0(55(3(<9(
6++0:,9.06(5.,3<**0*36;03+,)+07(5*9(A06*/0(9(
.069.056;650:;(53,@5(;/(50,3*9(=(;;),51(4054(<964(**(9965,
+LWHY[TLU[VM)PVTLKPJHS:JPLUJLZ<UP]LYZP[`VM;LYHTV;LYHTV0[HS`,\YVWLHU*LU[LY
MVY)YHPU9LZLHYJO*,9*:HU[H3\JPH-V\UKH[PVU9VTL0[HS`*VTW\[H[PVUHSIPVJOLTPZ[Y`
HUKIPVWO`ZPJZSHIVYH[VY`.90)0404<UP]LYZP[`VM7VTWL\-HIYH)HYJLSVUH)PVTLKPJHS
9LZLHYJO7HYR79)))HYJLSVUH:WHPU+LWHY[TLU[ZVM*LSS)PVSVN`HUK*OLTPZ[Y`;OL
:RHNNZ0UZ[P[\[LMVY*OLTPJHS)PVSVN`;OL:JYPWWZ9LZLHYJO0UZ[P[\[L3H1VSSH*HSPMVYUPH<:(
-H[[`HJPKHTPKLO`KYVSHZL-((/PZHYLWYLZLU[H[P]LTLTILYVMTLTIYHULLUa`TLZ
[OH[OH]PUNHZZ\IZ[YH[LZSPWPKTVSLJ\SLZULLK[VNL[HSPWVWOPSPJJVTWV\UKMYVT
^P[OPU[OLTLTIYHULHUK[VJH[HS`aLP[ZO`KYVS`ZPZI`^H[LY4VYLV]LY-((/PZ[OL
LUa`TLWSH`PUNHJYP[PJHSYVSLPU[OLTVK\SH[PVU[OLLUKVJHUUHIPUVPK[VULPUTHTTHSZ
KLNYHKPUNPTWVY[HU[UL\YV[YHUZTP[[LYZZ\JOHZHUHUKHTPKLHUKVSLHTPKL;OLSH[[LY
JVTWV\UKZ HYL PU]VS]LK PU H U\TILY VM O\THU KPZLHZLZ YLWYLZLU[PUN [OLYLMVYL H
]HS\HISL[HYNL[MVYIPVJOLTPJHSHUKWOHYTHJVSVNPJHSYLZLHYJO
([[LTW[Z [V PU]LZ[PNH[L [OL YVSL VM [OL Z\YYV\UKPUN SPWPKZ PU TLTIYHULLTILKKLK
LUa`TLZOHZUV[ILLUX\P[LZ\JJLZZM\SZVMHY`L[SPWVWOPSPJJVTWV\UKZHYLL_WLJ[LK
[VPU[LYHJ[^P[O[OLTLTIYHULPUZL]LYHS^H`Z;OLHPTVM[OPZZ[\K`^HZ[VPU]LZ[PNH[L
PUKL[HPSZ[OLLMMLJ[VMTLTIYHULZVMKPMMLYLU[SPWPKJVTWVZP[PVUZ[OH[VU-((/HUK[V
HZJLY[HPU[OLPYLMMLJ[VUP[ZZ[Y\J[\YLRPUL[PJHUKTLTIYHULIPUKPUNWYVWLY[PLZ
/LYL^LKLZJYPIL[OLÄYZ[Z[Y\J[\YLPUZVS\[PVUVM-((/I`ZTHSSHUNSL?YH`ZJH[[LYPUN
:(?: HUHS`ZPZ HUK KPZJSVZL UL^ Z[Y\J[\YHS HUK M\UJ[PVUHS LMMLJ[Z VM TLTIYHUL
SPWPKZVU[OPZLUa`TL-\Y[OLYTVYL[OPZZ[\K`YLWYLZLU[ZHJVUJLW[\HSZ[LWMVY^HYK
[V \UKLYZ[HUK TLJOHUPZ[PJ KL[HPSZ VM [OL M\UJ[PVUHS YLN\SH[PVU VM H WHYHKPNTH[PJ
TLTIYHULLUa`TLSPRL-((/I`[OLZ\YYV\UKPUNSPWPKZ;V[OPZHPTHJVTIPUH[PVU
VM L_WLYPTLU[HS HUK JVTW\[H[PVUHS TL[OVKZ ^HZ PU[LNYH[LK PU H T\S[PKPZJPWSPUHY`
HWWYVHJO
6]LYHSSV\YÄUKPUNZZ[YLZZ[OLPTWVY[HUJLVMTLTIYHULSPWPKZHUKPUWHY[PJ\SHYVM
JOVSLZ[LYVS MVY [OL TVK\SH[PVU VM [OL LUKVJHUUHIPUVPK [VUL ^P[OPU H JLSS I\[ HSZV
ZOV^ HU PU[LYLZ[PUN L_HTWSL VM [OL L_[LUZP]L TVK\SH[PVU [OH[ [OL TLTIYHUL SPWPK
JVTWVZP[PVUL_LY[ZVUHJOHYHJ[LYPZ[PJHUKPTWVY[HU[TLTIYHULLUa`TLH[[OLSL]LS
VMZ[Y\J[\YLSVJHSPaH[PVUHUKHJ[P]P[`
;OPZ^VYR^HZZ\WWVY[LKI`4PUPZ[LYVKLSS»0Z[Y\aPVULKLSS»<UP]LYZP[nLKLSSH9PJLYJH.YHU[7905
007VZ[LY:LZZPVU
PRO 30
Proteine
+5(IPUKPUNWYVWLY[PLZVM[OLO\THUA-7WYV[LPU
+,*,:(9,3<*0()(.30=603(90(,:76:0;6:()905(
90=,33056(3,::0(.904(3+0.06=(55(90**06(5+9,((5+
7,+65,7(636=
+LWHY[TLU[VM,U]PYVTLU[HS:JPLUJL<UP]LYZP[`VM5HWSLZ*HZLY[H0[HS`
0UZ[P[\[LVM.LUL[PJZHUK)PVWO`ZPJZ()\aaH[P;YH]LYZV*595HWSLZ0[HS`
+5( TL[O`SH[PVU PZ VUL VM [OL LWPNLUL[PJ TVKPÄJH[PVUZ PU]VS]LK PU NLUL ZPSLUJPUN
PUJS\KPUN [OH[ VM [OL PTWYPU[LK NLULZ 0TWYPU[LK NLULZ HYL L_WYLZZLK MYVT VUS`
VUL WHYLU[HS HSSLSL HSVUN TL[O`SH[PVU WH[[LYUZ [OH[ HYL KL[LYTPULK K\YPUN NHTL[L
MVYTH[PVU0TWYPU[LKNLULZWSH`HRL`KL]LSVWTLU[HSYVSLHUKHYLNLULYHSS`VYNHUPaLK
PUJS\Z[LYZJVU[YVSSLKI`*W.YPJOZLX\LUJLZRUV^UHZ0TWYPU[PUN*VU[YVS9LNPVUZ
0*9Z0*9ZHYL[HYNL[LKI`WHYLU[HSHSSLSLZWLJPÄJ+5(4L[O`SH[PVUPTWYPU[Z[OH[HYL
LZ[HISPZOLKK\YPUNSH[LNHTL[VNLULZPZTHU[LPULKPUa`NV[LHUKZVTH[PJJLSSZHUK
LYHZLKPUWYPTVYKPHSNLYTJLSSZ
4L[O`S+5()PUKPUN7YV[LPUZ4)7ZPU[LYHJ[Z^P[OTL[O`SH[LK*W.HUKYLJY\P[VU
+5( KPMMLYLU[ JVYLWYLZZVY JVTWSL_LZ MVYTPUN [YHUZJYPW[PVUHSS` ZPSLUJLK JOYVTH[PU
Z[Y\J[\YLZ:LX\LUJLZWLJPÄJYLJVNUP[PVUVMTL[O`SH[LK+5(OHZILLUKLTVUZ[YH[LK
MVY O\THU aPUJ ÄUNLY WYV[LPUZ 2HPZV A);) A);) HUK A-7 ;OLZL WYV[LPUZ
PU[LYHJ[PUHZLX\LUJLZWLJPÄJTHUULY^P[O+5(JVU[HPUPUNTL[O`SH[LK*W.
>L OH]L KLTVUZ[YH[LK [OH[ [OL TV\ZL A-7 WYV[LPU TA-7 JVU[HPUPUN aPUJ
ÄUNLY KVTHPUZ A-Z YLJVNUPaLZ TL[O`SH[LK +5( [HYNL[ ZLX\LUJLZ \ZPUN [OL
ZLJVUKHUK[OL[OPYKA-Z0U[OPZZ[\K`^LWYLZLU[V\YYLZ\S[ZVU[OLO\THUA-7
OA-7WYV[LPUOA-7JVU[HPUZA-Z[OL[OPYKHUK[OLMVY[OILPUNOVTVSVN\L
[V [OL ZLJVUK HUK [OPYK VM [OL TA-7 >L KLTVUZ[YH[LK [OH[ [OL O\THU WYV[LPU
ZLX\LUJLL_[LUKPUNMYVTNS`JPUL[V]HSPULPUJS\KPUN[OL[OPYKHUK[OLMV\Y[O
A-ZPZZ\MÄJPLU[MVYOPNOHMÄUP[`IPUKPUN[V[OLTL[O`SH[LK+5(ZP[LZHUK^LTHWWLK
[OLJYP[PJHSA-HTPUVHJPKYLZPK\LZYLX\PYLKMVY[OLPU[LYHJ[PVU^P[O+5(
9LJLU[S`T\[H[PVUZPUOA-7THWWPUNPU[OLaPUJÄUNLYYLNPVUOH]LILLUHZZVJPH[LK
^P[O [YHUZPLU[ ULVUH[HS KPHIL[LZ ;5+" ^L KLTVUZ[YH[LK [OH[ ;5+ T\[H[PVUZ PU
OA-7A-HUKHIVSPZO+5(IPUKPUN
9LMLYLUJLZ!
8\LUUL]PSSL:L[HS4VSLJ\SHY*LSS5V]
4HJRH`+1.L[HS5H[\YL.LUL[PJZ (\N
007VZ[LY:LZZPVU
PRO 31
Proteine
:PTPSHYP[PLZHUKKPMMLYLUJLZVMYH[HUKO\THUMH[[`HJPK
HTPKLO`KYVSHZLZ
+0=,5,9,(34,905+(+(05,:,,590*6-,AA(-0364,5(
(5.,3<**0),(;90*,*36;03+,96:(;650*63(
*9(=(;;),51(405--05(AA0(.9×(3,::(5+964,0.0(470,96
4(**(9965,4(<96
-YVT[OL5L\YVTLK¶7VaaPSSP0[HS`HUK+LWHY[TLU[VM,_WLYPTLU[HS4LKPJPULHUK
)PVJOLTPJHS:JPLUJLZ<UP]LYZP[`VM9VTL¸;VY=LYNH[H¹9VTL0[HS`[OL
+LWHY[TLU[VM)PVTLKPJHS:JPLUJLZ<UP]LYZP[`VM;LYHTV;LYHTV0[HS`
[OL,\YVWLHU*LU[LYMVY)YHPU9LZLHYJO0Z[P[\[VKP9PJV]LYVL*\YHH*HYH[[LYL
:JPLU[PÄJV:HU[H3\JPH-V\UKH[PVU9VTL0[HS`[OL+LWHY[TLU[ZVM*LSS
)PVSVN`HUK*OLTPZ[Y`;OL:RHNNZ0UZ[P[\[LMVY*OLTPJHS)PVSVN`;OL:JYPWWZ
9LZLHYJO0UZ[P[\[L3H1VSSH*HSPMVYUPH HUK[OL*LU[LYMVY0U[LNYH[LK9LZLHYJO
*09<UP]LYZP[`*HTW\Z)PVTLKPJVKP9VTH9VTL0[HS`
-H[[`HJPKHTPKLO`KYVSHZL-((/PZHTLTIYHULIV\UKLUa`TL[OH[PU[LYHJ[Z^P[O
UL\YVTVK\SH[VY`MH[[`HJPKHTPKLZHUKLZ[LYZHUKPZL_WYLZZLKPUTHTTHSPHUIYHPU
HUKWLYPWOLYHS[PZZ\LZ0[KLNYHKLZPTWVY[HU[UL\YV[YHUZTP[[LYZZ\JOHZVSLHTPKLHUK
HUHUKHTPKLHUKP[OHZILLUPU]VS]LKPUHU\TILYVMO\THUWH[OVSVNPJHSJVUKP[PVUZ
YLWYLZLU[PUN [OLYLMVYL H ]HS\HISL [HYNL[ MVY IPVJOLTPJHS HUK WOHYTHJVSVNPJHS
YLZLHYJO0U[OPZZ[\K`^LOH]LPU]LZ[PNH[LKPU]P[YV[OLZ[Y\J[\YLM\UJ[PVUYLSH[PVUZOPW
VMYH[HUKO\THU-((/Z[OYV\NOZL]LYHSZWLJ[YVZJVWPJ[LJOUPX\LZ0UWHY[PJ\SHY^L
PU]LZ[PNH[LK [OL Z[Y\J[\YHS MLH[\YLZ VM [OL [^V WYV[LPUZ IV[O PU [OL WYLZLUJL HUK
HIZLUJL VM [OL PYYL]LYZPISL PUOPIP[VY TL[OV_`HYHJOPKVU`SÅ\VYVWOVZWOVUH[L ;OL
YLZ\S[Z KLTVUZ[YH[L [OH[ [OL Z[Y\J[\YHS WYVWLY[PLZ VM [OL [^V -((/Z HYL KPMMLYLU[
PU]VS]PUNIV[OZLJVUKHY`HUK[LY[PHY`Z[Y\J[\YLLSLTLU[ZVM[OLWYV[LPUZKLZWP[L[OLPY
OPNOZLX\LUJLOVTVSVN`HUKV]LYHSSZPTPSHYP[`PU[LTWLYH[\YLKLWLUKLUJL
(KKP[PVUHSS`TLTIYHULIPUKPUNHUKRPUL[PJHZZH`ZVMIV[O-((/ZPUKPJH[L[OH[HSZV
[OLM\UJ[PVUHSWYVWLY[PLZVM[OL[^VLUa`TLZHYLKPMMLYLU[PU[OLPYPU[LYHJ[PVU^P[OSPWPK
IPSH`LYZHUK^P[OL_VNLUV\ZPUOPIP[VYZ
007VZ[LY:LZZPVU
PRO 32
Proteine
9VSLVM(YNMVY[OLYTVZ[HIPSP[`HUKJH[HS`[PJHJ[P]P[`VM
SPWV_`NLUHZLZ
+(33(),95(9+05(:0465(+0=,5,9,(34,905+(4,0.0(470,96
2l/5/(9;4<;0=(56=0.69(5+96:(;650*63(
+LWHY[TLU[VM,_WLYPTLU[HS4LKPJPULHUK)PVJOLTPJHS:JPLUJLZ<UP]LYZP[`VM;VY
=LYNH[H=PH4VU[WLSSPLY9VTL0[HS`"0UZ[P[\[LVM)PVJOLTPZ[Y`*OHYP[t
<UP]LYZP[p[ZTLKPaPU)LYSPU6\KLUHYKLY:[Y+)LYSPU.LYTHU`"0UZ[P[\[L
VM;V_`JVSVN`HUK7OHYTHJVSVN`<UP]LYZP[p[ZTLKPaPU9VZ[VJR:JOPSSPUN(SSLL
+ 9VZ[VJR.LYTHU`
3PWV_`NLUHZL HYL TL[HSJVU[HPUPUN SPWPK WLYV_PKPaPUN LUa`TLZ [OH[ JH[HS`aL
KPV_`NLUH[PVU VM WVS`\UZH[\YH[LK MH[[` HJPKZ [V [OLPY JVYYLZWVUKPUN O`KYVWLYV_`
KLYP]H[P]LZ
4HTTHSPHU SPWV_`NLUHZLZ 36?Z OH]L ILLU PTWSPJH[LK PU [OL WH[OVNLULZPZ VM
PUÅHTTH[VY`HUKO`WLYWYVSPMLYH[P]LKPZLHZLZI\[[OLKL[HPSLKTLJOHUPZTVMHJ[PVU
PZZ[PSSHTH[[LYVMKPZJ\ZZPVU4\[HNLULZPZHUKZ[Y\J[\YHSTVKLSPUNZ\NNLZ[LK[OH[
MVY 36? MH[[` HJPK Z\IZ[YH[L WLUL[YH[LZ PU[V [OL HJ[P]L ZP[L ^P[O P[Z TL[O`S
[LYTPU\ZHOLHK^OLYLI`[OLMH[[`HJPKJHYIV_`SPJNYV\WTPNO[PU[LYHJ[^P[O(YN
[OH[ PZ SVJHSPaLK H[ [OL LU[YHUJL PU[V [OL W\[H[P]L Z\IZ[YH[L IPUKPUN WVJRL[ ?YH`
JY`Z[HSSVNYHWO`HUKTVSLJ\SHYK`UHTPJZZPT\SH[PVUZVMSPNHUKIPUKPUNH[[OLHJ[P]L
ZP[LVMYHIIP[36?Z\NNLZ[LK[OH[HJ[P]LZP[LPUOPIP[VYZHUKVYZ\IZ[YH[LMH[[`
HJPKZPUK\JLJVUMVYTH[PVUHSJOHUNLZPU[OLPTTLKPH[LZ\YYV\UKPUNVM[OLJH[HS`[PJ
JLU[LY 0U WHY[PJ\SHY L_[LYUHS ƠOLSP_ [YHUZSVJH[LZ H^H` MYVT P[Z VYPNPUHS WVZP[PVU
[VVWLU[OLLU[YHUJL[V[OLIPUKPUNJH]P[`JSVZL[V(YNWHY[S`^OLYLI`P[ZIHZL
YLZPK\LZ ILJVTL KPZVYKLYLK 0U SPNHUKMYLL 36? (YN MVYTZ HU
PVUPJUL[^VYR^P[O[OLZPKLJOHPUVMLP[OLY(ZWHZMV\UKPUJY`Z[HSHUKVY.S\
HZVIZLY]LKK\YPUN4+ZPT\SH[PVUZ(ZZ\NNLZ[LKI`4+ZPT\SH[PVUZ[OPZPU[LYHJ[PVU
TPNO[ILKPZY\W[LK\WVUSPNHUKIPUKPUN
;VVI[HPUTVYLKPYLJ[L]PKLUJLMVY[OLWVZZPISLYVSLVM(YNMVY[OLJH[HS`[PJHUK
Z[Y\J[\YHSWYVWLY[PLZVMTHTTHSPHU36?^LJYLH[LK[OL(YN3L\T\[HU[VM
[OL YHIIP[ 36? ;OL KH[H WYLZLU[LK OLYL Z\NNLZ[ [OH[ (YN3L\ L_JOHUNL
HMMLJ[ZJH[HS`[PJHJ[P]P[`HUKPUJYLHZLK[OLZ\ZJLW[PIPSP[`VM[OLLUa`TLMVYZ\IZ[YH[L
PUOPIP[PVU/V^L]LYP[OHYKS`PTWHPYLK[OLHMÄUP[`VM[OLT\[HU[[V^HYKZIV[OSPUVSLPJ
HJPK HUK HYHJOPKVUPJ HJPK 4VYLV]LY ^L MV\UK [OH[ (YN3L\ L_JOHUNL KVLZ
UV[ PUK\JL THQVY HS[LYH[PVUZ PU ZLUZP[P]P[` VM [OL LUa`TL MVY [LTWLYH[\YLPUK\JLK
\UMVSKPUN0UJVU[YHZ[[OL[LY[PHY`Z[Y\J[\YLVM[OLLUa`TLWYV[LPU^HZJVUZPKLYHIS`
KLZ[HIPSPaLKHZPUKPJH[LKI`Å\VYLZJLUJLZ[\KPLZ
007VZ[LY:LZZPVU
PRO 33
Proteine
;OLWVPU[T\[H[PVUPU[OLHWVW[VZPZPUK\JPUNMHJ[VY(0-
NLULJH\ZPUNLHYS`KLYHUNLTLU[PUO\THUIYHPU
KL]LSVWTLU[ZWLJPÄJHSS`HS[LYZ[OL5(+/YLHJ[P]P[`VM[OL
ÅH]VWYV[LPU
(30=,9;0(3,::(5+96)(9650:(9(7(5+050=0;;690670**636,3,5(
:699,5;0563<*(=(56504(90((5;650,;;(
+PWHY[PTLU[VKP)PVZJPLUaL<UP]LYZP[nKLNSP:[\KPKP4PSHUV
]PH*LSVYPH4PSHUV0[HS`
;OL HWVW[VZPZ PUK\JPUN MHJ[VY (0- PZ H TP[VJOVUKYPHS 5(+/IPUKPUN ÅH]VWYV[LPU
[OH[ ^OLU YLSLHZLK MYVT [OL VYNHULSSL PU YLZWVUZL [V ]HYPV\Z Z[PT\SP JH\ZLZ JLSS
KLH[O 0U HKKP[PVU [V P[Z YVSL PU JHZWHZLPUKLWLUKLU[ HWVW[VZPZ (0- HSZV WSH`Z
HU LZZLU[PHS HS[OV\NO WVVYS` \UKLYZ[VVK M\UJ[PVU PU THPU[HPUPUN [OL M\UJ[PVUHSP[`
VM V_PKH[P]L WOVZWOVY`SH[PVU >OPSL (0- NLUL PUHJ[P]H[PVU PZ LTIY`VUPJ SL[OHS PU
THTTHSZ KV^UYLN\SH[PVU VM P[Z L_WYLZZPVU JH\ZLZ T`VWH[OPLZ HUK UL\YVSVNPJHS
HIUVYTHSP[PLZ=LY`YLJLU[S`[^VKPZ[PUJ[WVPU[T\[H[PVUZPUO\THU(0-NLUL^LYL
PKLU[PÄLK HZ YLZWVUZPISL VM O\THU TP[VJOVUKYPVWH[OPLZ YLZ\S[PUN PU ZL]LYL LHYS`
LUJLWOHSVT`VWH[O` HUK WYLUH[HS ]LU[YPJ\SVTLNHS` YLZWLJ[P]LS` >L OH]L
YLWYVK\JLK [OL SH[[LY T\[H[PVU PU TV\ZL (0- ;OL WH[OVNLUPJ (0-., ]HYPHU[
^HZPZVSH[LKPUHYLJVTIPUHU[MVYTHUK^HZZOV^U[VILHZZ[HISLHZ[OL^PSK[`WL
WYV[LPU;OPZHTPUVHJ`SYLWSHJLTLU[^HZMV\UK[VZPNUPÄJHU[S`ZSV^KV^U[OLYLHJ[PVU
IL[^LLU[OLWYV[LPUHUK5(+/[OH[SLHKZ[V[OLMVYTH[PVUVMH5(+-(+/JOHYNL
[YHUZMLYJVTWSL_^OPJOOHZILLUWYL]PV\ZS`ZOV^U[VILSPURLK[VHTVUVTLYKPTLY
[YHUZP[PVUVM[OLWYV[LPUX\H[LYUHY`Z[Y\J[\YL;OLZLYLZ\S[ZZ\NNLZ[[OH[[OLZSV^LY
YLHJ[PVU^P[O5(+/VM(0-.,PUJVTWHYPZVU[V[OL^PSK[`WLWYV[LPUTPNO[ILH[
[OLIHZPZVM[OLWH[OVNLUPJTLJOHUPZTVM[OLT\[H[PVU-\Y[OLYTVYLPU[OLSPNO[VM
[OLWYVWVZLKYVSLVM(0-HZHZ^P[JOPUZLUZPUN[OLTP[VJOVUKYPHSYLKV_Z[H[LHUKVY
5(+/JVUJLU[YH[PVUV\YKH[H\UKLYZJVYL[OLPTWVY[HUJLVMHYLHK`YLHJ[P]P[`VM
(0-[V^HYK5(+/MVY[OLM\SSM\UJ[PVUHSPU[LNYP[`VM[OLTP[VJOVUKYPVU
;OPZ^VYROHZILLUZ\WWVY[LKI`H7905NYHU[H^HYKLKI`[OL0[HSPHU4PUPZ[Y`VM
,K\JH[PVUHUK<UP]LYZP[`40<9[V4(=HUVUP
:L]YPV\RV]H0-(U[PV_PK9LKV_:PNUHS"! .OLaaP+L[HS(T1/\T.LUL["! )LYNLY0L[HS4VS.LUL[4L[HI"!
007VZ[LY:LZZPVU
PRO 34
Proteine
;PZZ\LHUKZ\IJLSS\SHYSVJHSPaH[PVUVMTHTTHSPHUYLUHSHZL
H-(+JVU[HPUPUNWYV[LPUPU]VS]LKPU[OLWH[OVNLULZPZVM
JHYKPV]HZJ\SHYKPZLHZLZ
)(9650:(9( :/<.9<,*/90:;05,=,3ÍA8<,A/,056
(30=,9;0(3,::(5+96.69,30*2-9,+:+,:09.(9@=
+PWHY[PTLU[VKP)PV:JPLUaL<UP]LYZP[nKLNSP:[\KPKP4PSHUV
]PH*LSVYPH4PSHUV0[HS`
+LWHY[TLU[VM0U[LYUHS4LKPJPUL@HSL<UP]LYZP[`:JOVVSVM4LKPJPUL
=(*/:*LKHY:[YLL[5L^/H]LU*; <:(
9LUHSHZL PZ H ZLJYL[VY` WYV[LPU HUK ÅH]VLUa`TL [OH[ PZ \IPX\P[V\Z PU ]LY[LIYH[LZ
HUK JVUZLY]LK PU ZVTL V[OLY WO`SH 0U THTTHSZ P[ OHZ ILLU ZOV^U [V TVK\SH[L
JHYKPV]HZJ\SHYYLZWVUZLZILPUNWHY[PJ\SHYS`HJ[P]LPUKLJYLHZPUNJH[LJOVSHTPULYNPJ
[VULSV^LYPUNISVVKWYLZZ\YLHUKPUWYV[LJ[PUN[OLOLHY[HNHPUZ[PZJOLTPJKHTHNL
3V^LYLKYLUHSHZLSL]LSZPU[PZZ\LHUKWSHZTHTPNO[IL[OLIHZPZVM[OLJHYKPV]HZJ\SHY
JVTWSPJH[PVUZ VIZLY]LK PU JOYVUPJ RPKUL` KPZLHZL WH[PLU[Z 9LUHSHZL ZLJYL[PVU
PU[V[OLJPYJ\SH[PVUPZLUOHUJLKPUYLZWVUZL[VZ[YLZZVYZZ\JOHZO`WLY[LUZPVUI\[[OL
TVSLJ\SHYTLJOHUPZTYLN\SH[PUNP[ZIHZHSVYZ[PT\SH[LKZLJYL[PVUHYL\URUV^U>L
ÄUK[OH[YLUHSHZLOHZHZPNUHSZLX\LUJLI\[[OPZZLX\LUJLPZUV[JSLH]LKWYPVY[VP[Z
ZLJYL[PVUZ\NNLZ[PUN[OH[P[TH`[YHMÄJPUHUH[`WPJHSZLJYL[VY`WH[O^H`0UWPNRPKUL`
V\Y PTT\UVÅ\VYLZJLUJL Z[\KPLZ ZOV^LK [OH[ YLUHSHZL PZ L_JS\ZP]LS` L_WYLZZLK PU
[OLWYV_PTHS[\I\SL:PTPSHYZ[\KPLZPUO\THUPTTVY[HSPaLK/2JLSSZHZ^LSSHZVU
WPNHUKTV\ZLWYPTHY`JLSSSPULZPUKPJH[LK[OH[YLUHSHZLPZWYLMLYLU[PHSS`SVJHSPaLK
PU[OLJ`[VWSHZT^OLYLP[ZOV^ZHW\UJ[H[LKPZ[YPI\[PVUZ\NNLZ[P]LVMHUVYNHULSSL
HZZVJPH[PVU ;OL PKLU[PÄJH[PVU VM [OLZL Z\IJLSS\SHY JVTWHY[TLU[Z TLJOHUPZT VM
HZZVJPH[PVU HUK YLUHSHZL»Z TLJOHUPZT VM ZLJYL[PVU HYL \UKLY^H` ;OPZ RUV^SLKNL
JV\SKSLHK[VUV]LS[OLYHWPLZMVYJHYKPV]HZJ\SHYHUKRPKUL`KPZLHZLZ
;OPZ^VYROHZILLUZ\WWVY[LKI`[YH]LSMLSSV^ZOPWZNYHU[LKI`[OL0[HSPHU:VJPL[`VM
)PVJOLTPZ[Y`HUK4VSLJ\SHY)PVSVN`:0)HUKI`[OL*VUZVYaPV0U[LY\UP]LYZP[HYPVKP
)PV[LJUVSVNPL[V:)HYVUP
+LZPY.=*\YY6WPU5LWOYVS/`WLY[LUZ"!
4PSHUP4L[HS14VS)PVS"!
<UNLY;L[HS,\Y/LHY[1"! 007VZ[LY:LZZPVU
PRO 35
Proteine
:[Y\J[\YHSHUHS`ZPZVM5\JSLVWOVZTPU*[LYTPUHSKVTHPU
PUJVTWSL_^P[OH.X\HKY\WSL_ZLX\LUJLMYVT[OLJ4@*
WYVTV[LY
36:;,9A6*(936 .(336(5.,36469040926+04(;;,6(+,3,
),9;0500=(56)(5*0)9<56904(<90A06(5+-,+,90*03<*(
0UZ[P[\[LVM)PVSVN`HUK4VSLJ\SHY7H[OVSVN`VM[OL*59+LWHY[TLU[VM
)PVJOLTPJHS:JPLUJLZ¸(9VZZP-HULSSP¹¸:HWPLUaH¹<UP]LYZP[`VM9VTL
9VTL0[HS`4HNUL[PJ9LZVUHUJL*LU[LY+LWHY[TLU[VM*OLTPZ[Y`
<UP]LYZP[`VM-SVYLUJL :LZ[V-PVYLU[PUV0[HS`
*LU[LYVM,_JLSSLUJLVU(NPUNHUK+LWHY[TLU[VM)PVTLKPJHS:JPLUJLZ
<UP]LYZP[`VM*OPL[P¸.+»(UU\UaPV¹*OPL[P0[HS`
5\JSLVWOVZTPU574PZHU\JSLVJ`[VWSHZTPJZO\[[SPUNWYV[LPUTHPUS`SVJHSPaLKH[
U\JSLVSP^OPJOWHY[PJPWH[LZ[VZL]LYHSJLSS\SHYM\UJ[PVUZPUJS\KPUNYPIVZVTLTH[\YH[PVU
HUKL_WVY[JLU[YVZVTLK\WSPJH[PVUHUKYLZWVUZL[VZ[YLZZZ[PT\SP;OL574NLULPZ
[OLTVZ[MYLX\LU[S`T\[H[LKPU(J\[L4`LSVPK3L\RLTPH(43WH[PLU[Z(SST\[H[PVUZ
THW[V[OLSHZ[L_VUVM[OLNLULHUK[OLLUJVKLKT\[H[LKWYV[LPUZHYLHILYYHU[S`HUK
Z[HIS`SVJHSPaLKPU[OLJ`[VWSHZTK\L[VOPNOKLZ[HIPSPaH[PVUVM[OL574*[LYTPUHS
KVTHPU HUK [OL HWWLHYHUJL VM H UL^ U\JSLHY L_WVY[ ZPNUHS 7YL]PV\ZS` ^L OH]L
ZOV^U [OH[ [OL YLZPK\L 574 *[LYTPUHS KVTHPU 574* IPUKZ H ZWLJPÄJ
YLNPVUH[[OLJ4@*NLULWYVTV[LYJOHYHJ[LYPaLKI`WHYHSSLS.X\HKY\WSL_Z[Y\J[\YL
/LYL ^L WYLZLU[ [OL ZVS\[PVU Z[Y\J[\YL VM [OL 574* KVTHPU HUK HUHS`ZL P[Z
PU[LYHJ[PVU^P[OHJ4@*KLYP]LK.X\HKY\WSL_I`549;OLZLKH[H^LYL\ZLK[V
JHSJ\SH[LHUL_WLYPTLU[HSS`YLZ[YHPULKTVSLJ\SHYKVJRPUNTVKLSMVY[OLJVTWSL_;OL
574*[LYTPUHS[OYLLOLSP_I\UKSLIPUKZ[OL.X\HKY\WSL_+5(H[[OLPU[LYMHJL
IL[^LLUOLSPJLZ/HUK/[OYV\NOLSLJ[YVZ[H[PJPU[LYHJ[PVUZ^P[O[OL.X\HKY\WSL_
WOVZWOH[L IHJRIVUL :L]LYHS YLZPK\LZ JVU[HJ[PUN [OL .X\HKY\WSL_ +5( HYL
RUV^U[VILZ\IQLJ[LK[VWVZ[[YHUZSH[PVUHSTVKPÄJH[PVUZPUJS\KPUNHJL[`SH[PVUHUK
WOVZWOVYPSH[PVUZ\NNLZ[PUN[OLPYWVZZPISLYLN\SH[VY`YVSLPUTVK\SH[PUN574+5(
PU[LYHJ[PVUZ-PUHSS`HUKPU[LYLZ[PUNS`^LYLWVY[[OH[HYLZPK\LS`ZPULYPJOZLX\LUJL
H[[OL5[LYTPU\ZVM[OL[OYLLOLSP_I\UKSL[OH[WYV]LKULJLZZHY`[VPUJYLZLHMÄUP[`
MVY[OL.X\HKY\WSL_WHY[ULYPZKPZVYKLYLKHUKKVLZUV[WHY[PJPWH[LKPYLJ[S`[V[OL
JVU[HJ[Z\YMHJLPU[OLJVTWSL_
007VZ[LY:LZZPVU
PRO 36
Proteine
;/,94(3(5+*/,40*(3<5-63+05.6-
4,;/@3;/06(+,56:05,7/6:7/69@3(:,00-964;/,
(9*/(,65:<3-636)<::63-(;(90*<:
*(**0(7<6;0.06=(55(-<**06-9(5*,:*(+,3,6,:;,9
)(.(96364(90(30),9((5+769*,3304(905(
+PWHY[PTLU[VKP)PVJOPTPJHL)PVÄZPJH:LJVUKH<UP]LYZP[nKP5HWVSP=PH
*VZ[HU[PUVWVSP5HWVSP
»+LV_`»TL[O`S[OPVHKLUVZPUL WOVZWOVY`SHZL 00 MYVT :\SMVSVI\Z ZVSMH[HYPJ\Z
:Z4;(700PZHO`WLY[OLYTVWOPSPJHUKO`WLY[OLYTVZ[HISLLUa`TLILSVUNPUN[VW\YPUL
U\JSLVZPKL WOVZWVY`SHZLZ \IPX\P[V\Z LUa`TLZ [OH[ M\UJ[PVU PU [OL W\YPUL ZHS]HNL
WH[O^H`:Z4;(700PZH[PNO[S`WHJRLKOL_HTLYPJWYV[LPUVYNHUPaLKHZHKPTLYVM
[YPTLYZ^P[OVULHJ[P]LZP[LWLYTVUVTLY,HJOZ\I\UP[PZZ[HIPSPaLKI`[^VWHPYZVM
PU[YHZ\I\UP[KPZ\SÄKLIYPKNLZHUKJVU[HPUZPUP[Z*[LYTPUHSYLNPVUH*?*TV[PMHZH
[`WPJHSMLH[\YL;VNL[PUMVYTH[PVUVU[OLYVSLWSH`LKI`KPZ\SÄKLIVUKZPUZ[HIPSP[`
HUKMVSKPUN[OLJVUMVYTH[PVUHSZ[HIPSP[`VM:Z4;(700HUK*:HUK* :*:
T\[HU[Z ^HZ Z[\KPLK I` [OLYTHS HUK N\HUPKPUP\T JOSVYPKLPUK\JLK \UMVSKPUN HUK
HUHS`aLKI`Å\VYLZJLUJLZWLJ[YVZJVW`JPYJ\SHYKPJOYVPZTHUK:+:7(.,5V[OLYTHS
\UMVSKPUN[YHUZP[PVUVM:Z4;(700JHUILVI[HPULK\UKLYUVUYLK\JPUNJVUKP[PVUZ^OPSL
PU[OLWYLZLUJLVMYLK\JPUNHNLU[ZH;TVM‡*JHUILTLHZ\YLKKLTVUZ[YH[PUN
[OLPU]VS]LTLU[VMKPZ\SÄKLIYPKNLZPULUa`TL[OLYTVZ[HIPSP[`+PMMLYLU[S`MYVT[OL
^PSK[`WL *: HUK * :*: ZOV^ JVTWSL[L [OLYTHS KLUH[\YH[PVU J\Y]LZ
^P[OZPNTVPKHS[YHUZP[PVUZJLU[LYLKH[‡*HUK ‡*YLZWLJ[P]LS`<UKLYYLK\JPUN
JVUKP[PVUZ [OLZL ]HS\LZ KLJYLHZL I` ‡* HUK ‡* YLZWLJ[P]LS` OPNOSPNO[PUN [OL
PTWVY[HU[ YVSL L_LY[LK I` *?* KPZ\SÄKL VU LUa`TL [OLYTVZ[HIPSP[` *OLTPJHS HUK
[OLYTHS \UMVSKPUN VM :Z4;(700 HYL PYYL]LYZPISL WYVJLZZLZ 5L]LY[OLSLZZ \UKLY
YLK\JPUN JVUKP[PVUZ ^L ^LYL HISL [V OPNOSPNO[ [OL YL]LYZPISL KPZZVJPH[PVU VM [OL
OL_HTLY[VTVUVTLYZHUK[VKL[LJ[[OLYL]LYZPIS`\UMVSKLKZWLJPLZI`:+:7(.,
;OLZL YLZ\S[Z WYV]PKL JVU]PUJPUN L]PKLUJL [OH[ KPZ\SÄKL IVUKZ WSH` H RL` YVSL PU
[OLJVUMVYTH[PVUHSZ[HIPSP[`VM:Z4;(7000UMHJ[[OLYLTV]HSVM[OLZLJV]HSLU[SPURZ
^LHRLUPUN[OLOPNOS`JVTWHJ[Z[Y\J[\YLVM[OLLUa`TLHSSV^Z[OLYL]LYZPISL[YHUZP[PVU
MYVT [OL UH[P]L [V [OL KLUH[\YLK Z[H[L ILMVYL JV]HSLU[ TVKPÄJH[PVUZ PUK\JLK I`
OPNO [LTWLYH[\YLZ HUK JOLTPJHS KLUH[\YHU[Z TH` JH\ZL [OL PYYL]LYZPISL WYV[LPU
KLNYHKH[PVU;OPZPZ[OLÄYZ[YLWVY[VU[OLYL]LYZPISL\UMVSKPUNVMHO`WLY[OLYTVWOPSPJ
VSPNVTLYPJWYV[LPU^P[OKPZ\SÄKLIVUKZ
007VZ[LY:LZZPVU
PRO 37
Proteine
)PVJOLTPJHSJOHYHJ[LYPaH[PVUVM7ZL\KVTVUHZHLY\NPUVZH
/+.@7WOVZWOVKPLZ[LYHZLZPU]VS]LK
PUJKP.47KLWLUKLU[IPVÄSTMVYTH[PVU
:;,30;(56=(3,5;05(905(3+6:,9,5(.0(9+05(.069.06
*(:;0.3065,50*63,;;(*(9<:64(5<,3(*<;9<AA63±-9(5*,:*(
0Z[P[\[V7HZ[L\Y-VUKHaPVUL*LUJP)VSVNUL[[P+LWHY[TLU[VM)PVJOLTPJHS:JPLUJLZ
:HWPLUaH<UP]LYZP[`VM9VTL0[HS`
)HJ[LYPHL_PZ[PUUH[\YLHZWSHUR[VUPJZPUNSLJLSSZVYPUHZLZZPSLT\S[PJLSS\SHYZ[H[L
UHTLKIPVÄST0U[OLSH[[LY[OLIHJ[LYPHSJVTT\UP[`VW[PTPaLZ[OLJLSSLU]PYVUTLU[
HUKJLSS[VJLSSJVTT\UPJH[PVUZLUZPUNZ[YH[LNPLZ)PVÄSTZHYL^PKLS`KPMM\ZLPUTHU`
PUK\Z[YPHS LU]PYVUTLU[HS HUK JSPUPJHS ZL[[PUNZ HUK HYL SLZZ ZLUZP[P]L [V [YLH[TLU[Z
^P[O HU[PTPJYVIPHS HNLU[Z JVTWHYLK [V WSHUR[VUPJ JLSSZ )PVÄSTMVYTPUN WH[OVNLUZ
HYLYLZWVUZPISLVM%VMO\THUPUMLJ[PVUZ![OLYLK\JLKZLUZP[P]P[`[VJVU]LU[PVUHS
HU[PIPV[PJZJHSSZMVYUL^HU[PIPVÄST[OLYHWPLZ(RL`WSH`LYPUIPVÄSTKL]LSVWTLU[PZ
[OLZLJVUKTLZZLUNLYJ`JSPJKP.47"[OLLUa`TLZJVU[YVSSPUNP[ZPU[YHJLSS\SHYSL]LSZ
PL KPN\HU`SH[L J`JSHZLZ +.* HUK WOVZWOVKPLZ[LYHZLZ 7+, HYL MV\UK VUS` PU
IHJ[LYPH[O\Z[OL`YLWYLZLU[PKLHS[HYNL[Z[VKL]LSVWLMMLJ[P]LHU[PIPVÄSTZ[YH[LNPLZ
;OL HPT VM [OPZ WYVQLJ[ PZ [V Z[\K` [OL IPVJOLTPJHSZ[Y\J[\YHS IHZLZ VM JKP.47
TL[HIVSPZTPU[OLO\THUWH[OVNLU7ZL\KVTVUHZHLY\NPUVZHTHPUJH\ZLVMKLH[O
PUJ`Z[PJÄIYVZPZWH[PLU[Z+LZWP[L[OLYLSL]HUJLVM[OLZLLUa`TLZUVTVSLJ\SHYKH[H
VU [OL YLHJ[PVU TLJOHUPZT HYL H]HPSHISL >L OH]L JOVZLU [V Z[\K` YLWYLZLU[H[P]L
TLTILYZVMLHJOZ\IJSHZZVMW\[H[P]L+.*ZHUK7+,Z/LYL^LYLWVY[[OLYLZ\S[Z
VI[HPULKVU[^VW\[H[P]L7+,ZVM[OL/+.@7Z\I[`WL7(HUK7(^OVZL
HIPSP[`[VSV^LYJKP.47SL]LSZPU7HLY\NPUVZH^HZWYL]PV\ZS`Z\NNLZ[LK
6\YYLJLU[IPVJOLTPJHSZ[\KPLZVUW\YPÄLK7(KLTVUZ[YH[LK[OH[[OPZ/+.@7
WYV[LPUOHZJKP.47HUKW.W.JKP.47KLNYHKH[PVUWYVK\J[O`KYVS`[PJHJ[P]P[`
6U[OLV[OLYOHUKW\YPÄLK7(HUKP[ZPZVSH[LKJH[HS`[PJKVTHPUHYLPUHJ[P]LHZ
JKP.47 O`KYVSHZL ^OLYLHZ [OL SH[[LY WYV[LPU JHU JSLH]L [OL PU[LYTLKPH[L W.W.
;OLZL YLZ\S[Z Z\NNLZ[ [OH[ 7( TH` SV^LY [OL PU[YHJLSS\SHY SL]LSZ JKP.47
PUKPYLJ[S`^OLYLHZ7(HJ[ZHZHIVUHÄKL7+,:PUJL7(JH[HS`[PJKVTHPU
PZHISL[VYLK\JLJKP.47PU[YHJLSS\SHYSL]LSZHSZVHZYLJVTIPUHU[WYV[LPUPU,JVSP
[OLULLKVMHWO`ZPVSVNPJHSWHY[ULY[V[YPNNLY7(LUa`TH[PJHJ[P]P[`JHUUV[IL
Y\SLKV\[
007VZ[LY:LZZPVU
PRO 38
Proteine
(T`SVPKVZLZHZZVJPH[LK[V(WVSPWVWYV[LPU(0!
HUHS`ZPZVM[OLTVSLJ\SHYIHZLZVM[OLKPZLHZL
90;(+,3.0<+0*,(5.,3((9*0,336+(90(4(90(465;0
(5.,3((469,:(56(5+9,5(;(70**630
+LWHY[TLU[VM:[Y\J[\YHSHUK-\UJ[PVUHS)PVSVN`HUK+LWHY[TLU[VM*OLTPJHS
:JPLUJLZ<UP]LYZP[`VM5HWSLZ-LKLYPJV005HWSLZ0[HS`
=HYPHU[Z VM (WVSPWVWYV[LPU (0 (WV(0 HYL HZZVJPH[LK [V OLYLKP[HY` Z`Z[LTPJ
HT`SVPKVZLZJOHYHJ[LYPZLKI`HT`SVPKKLWVZP[PVUVML_[YHJLSS\SHYÄIYPSZPUWLYPWOLYHS
VYNHUZ VM WH[PLU[Z THPUS` JVUZ[P[\[LK I` 5[LYTPUHS MYHNTLU[Z VM [OL WYV[LPU ;OL
WVS`WLW[PKL JVYYLZWVUKPUN [V (WV(0 YLNPVU PZ [OL THPU JVUZ[P[\LU[ VM OLHY[
HT`SVPKKLWVZP[Z
>L WYVK\JLK H YLJVTIPUHU[ MVYT VM [OL ÄIYPSSVNLUPJ WVS`WLW[PKL UHTLK B D
(WV(0 [OH[ ZOHYLZ ÄIYPSSVNLUPJ WH[O^H` ^P[O P[Z UH[\YHS JV\U[LYWHY[ ( SPWPK
LU]PYVUTLU[ HMMLJ[Z [OL JVUMVYTH[PVUHS Z[H[L HUK HNNYLNH[PVU WYVWLUZP[` VM B D
(WV(0*OVSLZ[LYVSHUH[\YHS(WV(0SPNHUKHUKSPWVZVTLZVMKPMMLYLU[JVTWVZP[PVUZ
PUK\JLHUKZ[HIPSPaL[OLWVS`WLW[PKLƠOLSPJHSZ[H[LPUOPIP[PUNHNNYLNH[PVUB D
(WV(0YLJVNUPaLZZWLJPÄJIPUKPUNZP[LZVUYH[JHYKPVT`VISHZ[ZJLSSTLTIYHULHUK
PZ PU[LYUHSPaLK TVZ[S` [OYV\NO JOSH[YPUJVH[LK ]LZPJSLZ HUK SPWPK YHM[Z ^OPSL (WV(0
PZPU[LYUHSPaLKTVZ[S`[OYV\NOJOSH[YPUJVH[LK]LZPJSLZHUKTHJYVWPUVJ`[VZPZ<USPRL
(WV(0[OLWVS`WLW[PKLPZKLNYHKLKI`S`ZVZVTLZHUKWYV[LHZVTL-PIYPSZVMB D
(WV(0HYLPUZ[LHK\UHISL[VLU[LY[OLJLSSZ;OLPU[LYHJ[PVUVM[OLWVS`WLW[PKL^P[O
[HYNL[JLSSZHUKP[ZPU[YHJLSS\SHYYV\[LPZJLY[HPUS`YLSL]HU[[V[OLJVTWYLOLUZPVUVM[OL
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46067(:8<(3,
465;,*<**6*,:(9,
465;03<*(
469(**04(9*6
4690=(3,906
THYPSPZHZ\SWPaPV'`HOVVP[
46;;(*(93(
JHYSHTIH'SPILYVP[
4<9(<4),9;6
5(5,;;03(<9(
\T\YH'IPVSVNPH\UPWPP[
5(7630;(564(9*6
5(:;(:0.0(5*(936
THYJVUHWVSP[HUV'SPILYVP[
5,.90(94(5+6
630(96)6::6:0465,;;(
693(**/06(3+6V
HYTHUKVULNYP'\UPTPP[
6:;<50(5.,3(
7(*,33(:;,7/(50,
7(+<3((55(
7(340,90-,9+05(5+6
7(340,903<0.0
7(5,)0(5*6*65*,;;(
7(6307(636
HUNLSHVZ[\UP'\UPIHZP[
7(7(:,9.06
7(9(+0,:.0<:,77,
7(9,5;,(<.<:;6
7(996;;(96::,33(
7(:*(3,4(90(
ZLYNPVWHWH'\UPIHP[
7(::(96-()0(5(
7(::0(3),9;6
7,+65,7(636=05*,5A6
7,55,330(3-65:6
MHIPHUHWHZZHYV'\UPUHP[
7,9;05/,A;/,34(
[OLSTH'\UPWYP[
7,;9(.5(5050*63(
UWL[YHNUHUP'\UPJOP[
JLZHYLTVU[LJ\JJV'\UPWKP[
S\JHTVU[P'H[LULVW]P[
TTVYHJJP'PIWJUYP[
]TVYP'\UP]WTP[
SUHUL[[P'\UP]WTP[
NUHZ[HZP'O[VTHPSP[
ZPTVUHVSPHYV'\UP[VP[
YS`'\UPWNP[
WHJZ[LWOHUPL'`HOVVP[
HUUHWHK\SH'\UPIVP[
MWHST'MHYTIPVS\UPIHP[
S\PNPWHSTPLYP'\UPIHP[
JVUJL[[HWHULIPHUJV'\UPIHP[
WHVSVWHVSP'\UPÄP[
NWHYHKPLZ'IPVSVNPH\UPIHP[
H\N\Z[VWHYLU[L'\UPUHP[
YVZZLSSHWHYYV[[H'\UP[VP[
WHZJHSL'\UPZHP[
HSILY[VWHZZP'\UPUZ\IYPHP[
WHVSV]WLKVUL'\UPUHP[
HWLUULSSP'KZI\UPJOP[
7,;9<AA,3309(--(,3,
70**6309,5(;(
YWL[Y\aaLSSP'KZI\UPJOP[
70**6304(9*6
7050(3,::(5+96
WPJJVSPTHYJV'NTHPSJVT
7056;;040926
707056*(;,905(
TPYRVWPUV[[P'\UPMLP[
763*/0(30*,
7647<**0.0<:,77,
769;(9(--(,3,
HSPJLWVSJOP']PYNPSPVP[
769;(56=(7(;90A0(
76AA0=(3,5;05(
79060,;;0+»,47(09,3<*0(
9(--(,3305(+0(
9(--(,330-9(5*,:*(
9(=,9(:03=0(
9,(:03=0(5(
9,0*//(9;1,(54(9*1
90)6503(<9(
90**0;,330,3,5(
90.(569(*/,3,
90=,33056(3,::0(
90=0,336(5;65,33(
WH[YPaPHWVY['SPILYVP[
90AA04,50*6
90AA6),5,+,;;(
90AA6(5.,3(4(90(
96:(;0(3,::(5+9(
TLUPJVYPaaP'WOHYT\UPWTUP[
96:(;650*63(
96::0*3(<+0(
96::04(98<,A.06=(55(
96::050.0(57(636
9<..0,90:03=,906
UPJVSHYVZH[V'\UPYVTHP[
:()(;050:;,-(50(
Z[LMHUPHZHIH[PUP'\UPYVTHP[
WPJJVSP'\UPUHP[
WPUPH'\UPZPP[
JWPWPUV'\UPJOP[
WVTW\JJPF 'SPILYVP[
WVY[HYHM'\UPUHP[
]HSLU[PUHWVaaP'NTHPSJVT
OUZ'\UPMLP[
UYHMMHLSSP'\UP]WTP[
MYHUJLZJHYHMMHLSSP'HSPJLP[
ZPS]PHYH]LYH'NTHPSJVT
ZPS]PHUHYLH'\UPUHP[
49LPJOOHY['\UPZ[YHMY
SH\YHYPIVUP'\UPTPP[
LSLUHYPJJP[LSSP'\UPTPP[
YHJOLSLYPNHUV'PZZP[
HSLZZPHYP]LSSPUV'\UPUHP[
YP]PLSSV'OV[THPSP[
ILULKL[[HYPaaV'\UPIVP[
HUNLSHTHYPHYPaaV'\UPTPP[
HYVZH[P'\UPZHP[
JSH\KPHYVZZP'\UPJOP[
NPV]HUUHYVZZPTHYX\La'\UPUHP[
NPHUWHVSVYVZZPUP'\UPTVYLP[
Y\NNPLYP'\UP]WTP[
:(**/,;;(7(636
WZ'\UPJOP[
:(3(.0(53<*(
:(3646507(636
:(55(-()06
:(5;0(30*,
NPHUS\JHZHSH'\UPTVYLP[
:(5;<**0(55(30:(
:(9+(5,330(55(4(90(
:(9+(9650*63(
:(9;050+(=0+,
:(9;69.069.06
:*(**6:(3=(;69,
:*(9-0:650(
:*(905.09(--(,33(
:*/<9<*/040*/,3,
:*6;;03<*(
HUUHSPZHZHU[\JJP'\UPZPP[
:*904(96:,33(
YZJYPTH'\UPMNP[
:,770*3(<+06
JZLWWP'\UPW]P[
:,9050.<0+6
N\PKVZLYPUP'PYJJP[
:0.569,3364(90(.9(A0(
:0.56903,(55(
:03=,:;9003(90(4(90(;,9,:(
:050.(.30(-()063(
:63(050.0(5*(936
:70:50(3),9;6
:;,-(5,;;0*/0(9(
:;,30;(56=(3,5;05(
:;9(AA<330(5+9,(
:<**6064(90(5.,3(
:<370A064(9030:(
;(.30(A<**/0+(=0+,
;,:690,9,3<0:(
;0)650.0(54(906
THYPHNYHaPHZPNUVYLSSV'\UPNLP[
WZHSVTVUP'\JSHJ\R
ZHUUHMHIPV'`HOVVP[
HSPJLZHU[P'\UPÄP[
ZHYKHULSSPH'IPVJOLT\UPIHP[
ZHYKHYVUPJVSH'SPILYVP[
KH]PKLFZHY[PUP'SPILYVP[
NPVYNPVZHY[VY'\UPIVP[
ZHS]H[VYLZJHJJV'\UPIHP[
ZVUPHZJHYÄ'\UPNLP[
YHMMHLSSHZJHYPUNP'\UPTPP[
TZJ\Y\JOP'`HOVVP[
SZJV[[P'\UPJOP[
HUUHZPNUVYPSL'IPVJOLT\UPIHP[
PSHYPHZPS]LZ[YP'\UPTPP[
MHIPVSHZPUPNHNSPH'TLK\UPWTUP[
NPHUJHYSVZVSHPUP'\UPIVP[
HSILY[VZWPZUP'\UPWYP[
JOPHYHZ[LMHUL[[P'JLU[YV]VS[HP[
]HSLU[PUHZ[LSP[HUV'\UPYVTHP[
HZ[YHaa\SSP'PIWJUYP[
THYPHUNLSHZ\JJVPV'NTHPSJVT
THYPSPZHZ\SWPaPV'PZ[Y\aPVULP[
KH]PKL[HNSPHa\JJOP'\UPTVYLP[
S\PZH[LZVYPLYL'\UPWHP[
[PIVUP'\UPJOP[
;090)<A096),9;6
;69;04(<96
YVILY[V[PYPI\aP'`HOVVP[
;69;69(7(636
;6:(;;6:03=06
;6::0(3,::(5+96
;9(050:(9(
WHVSV[VY[VYH'\UPTPIP[
;9,5;050(3,::(5+96
;905*(=,3304(90(3,;0A0(
;905.(30*90:;05(
;96(+,*1,(5+,50:
;<9*64(90(*(;,905(
=(56504(9*6
=,5,9(5+6)9<56
=,5;69,5A(
HSLZZHUKYV[YLU[PUP'\UPMLP[
=0(507(63(
=0,*,30+(33(:,.(-9(5*,:*6
A(77(=0.5(:03=0(
A6**/0369,+(5(
A<**/,330409*6
A<336(3),9;6
WHVSH]PHUP'\UPTPP[
T[VY[P'\UPW]P[
ZPS]PV[VZH[[V'\UPWKP[
H[VZZP'\UP[ZP[
[YHPUPZ'\UPJOP[
S[YPUJH]LSSP'MHYT\UPWPP[
JYPZ[PUH[YPUNHSP'\UPTPP[
QK[YVHKLJ'\UP]JLaHUULMY
TJ[\YJV'\UPZHP[
THYJV]HUVUP'\UPTPIP[
IY\UV]LULYHUKV'\UPTPP[
Y]LU[V'\UPWHP[
MYHUJLZJV]PLJLSP'\UPIVP[
ZPS]PHaHWWH'SPILYVP[
SVYLKHUHFaVJJOP'`HOVVP[
Ta\JJOLSSP'\UPJOP[
HSIa\SSV'\UPZHUUPVP[
index
7YVNYHTTL7HN
(U[VUPUP3LJ[\YL7HN
:LZZPVU 7HN
:LZZPVU 7HN
:LZZPVU 7HN
:LZZPVU 7HN
:LZZPVU 7HN
:LZZPVU 7HN
:LZZPVU 7HN
:LZZPVU 7HN
(TTPUL)PVNLUL 7HN
)PVJOPTPJH4HYPUHLKLSS»(TIPLU[L 7HN
)PVJOPTPJH*LSS\SHYL 7HN
)PVJOPTPJHKLSSH5\[YPaPVUL7HN
index
0TT\UVSVNPH)PVJOPTPJH7HN
:]PS\WWV+PMMLYLUaPHTLU[VL(WVW[VZP7HN
)PV[LJUVSVNPLL)PVJOPTPJH0UK\Z[YPHSL7HN )PVPUMVYTH[PJH7HN
,UaPTVSVNPHL9LNVSHaPVUL4L[HIVSPJH7HN
.SPJVIPVSVNPH7HN
4LTIYHULL)PVLULYNL[PJH7HN
5\JSLV[PKP(JPKP5\JSLPJP.LUVTP7HN
5L\YVJOPTPJH7HN
7YV[LPUL7HN 7HY[PJPWHU[SPZ[7HN