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Biochemistry, Basic Carbohydrates

prepared by Mr. Chroeng Sopheap, phone 012864353/011717361, e-mail: [email protected] of 20

CARBOHYDRATES

vtßúbMNgemeron 1-Bnül;BIPaBxusKñarvag carbohydrates cMruH (complex) nig carbohydrates samBaØ nig dwgBIcnYnrbs; BYkvanimYy²EdleKGnuBaØt[manenAkñúgrbbGaharRbcaMéf¶ .

2-GnuvtþRbB½n§énkareFIVcMNat;fñak; nig kar[eQμaHBYk monosaccharides GnuelameTAtambgÁúMnaTI nig cMnYn rbs;GatUmkae)anenAkñúgExS .

3-kMNt;faetIm:UelKulmYymancMNuckNþalEdlmanéd (chiral center) rebobdUcemþc .

4-Bnül;GMBI stereoisomerism .

5-kMNt;GtþsBaØaNrbs; monosaccharides faCaes‘rI D- b¤ L- .

6-KUs nig pþl;eQμaH monosaccharides EdlCYbjwkjab;edayeRbIrUbmnþeRKag (structueal formulas)

7-edaypþl;nUvrUbmnþCaExSlat (linear structure) rbs; monosaccharides KUsnUvrUbmnþ Haworth énTMrg; α- nig β- nig pÞúymkvij .

8-edayBinitüeTAelIrcnasm<½n§ TayfaetIsár (sugar) KWCasárer:dukr b¤ minEmnsárer:dukr (reducing or non-

reducing sugar) .

9-BiPakSaBIkareRbIRbtikr Benedict edIm,Ivas;kMriténKøúykUskñúgTwkenam .

10-KUs nig [eQ μaHBYk disaccharides nig BiPakSaBIsar³sMxan;rbs;BYkvaenAkñúgRbB½n§CIvsa®sþ .

11-BiB’NnaBIPaBxusKñarvag galactosemia nig lactose intolerance .

12-BiPakSaBIlkçN³énrcnasb<½n§ KImI nig CIvKImIrbs;GamIdug (startch) , glycogen nig cellulose .



KMeragemeron 1-RbePTrbs; CARBOHYDRATES

2-MONOSACHARIDES

namvlI Stereoisomers

Monosaccharides Edlmansar³sMxan;mYycMnYn

sárer:dukr (Reducing sugars)

3-DISACCHARIDES

Maltose Lactose Sucrose

4-POLYSACCHARIDES

GamIdug (startch)

Glycogen Cellulose

Bioch

prepar

esc carbo

RbPBsMrab;eday(kcal

cellu

CaBies(obes

)araMg eyIgnU carbo

mCÆwmGnuBaØénkaLeKGnu)ay TaMgKW

hemistry, Basic

red by Mr. Chro

ckþIepþIm Carbohy

ohydrates mBsMxan;énfambxYrk,al nig

yekasikasMrab;l) .

rbbGahaulose nig sáss sucrose

sity) nig karCarbohy

nMb½ug nig RKanUvkarpÁt;pÁg;sM

vamankarlohydrates .

mbUN’BaEdl)aaØatfa RbEhlLÚrIRbcaMéf¶RtUnBaØatenAkñúgrb pasta¦ nig

KCaRbPBsMxan

c

oeng Sopheap, p

ydrates RtUv)mandUcCa RKab;Blrbs;BYksRbB½n§RbsaT

bbegáItfamBl

arEdlpþl;suxrsamBaØ (sim

e RtUvEtmancMxUceF μj (toot

ydrates cMruHvab;FBaØCati KWC

sMxan;énséskñlM)akkñúgkarkM. PaKryFmμ)ayKWCaFatusMl 58° énkaLv)anmkBI su

bbGahar ¬rUbkMritbnÞab; Epø

n;én carbohy

phone 0128643

)anplitenAkñúgb;FBaØCati (gra

stV . Carbo

nig GacRtUvl 1 Rkam²

PaBl¥ (heal

mple sugars

cMnYnticGaRs½yth decay) .val¥cMeBaHeyIgCaRbPBfamBkñúgrbbGaharkMNt;[)anc,ata (actual p

sxan;rbs;rbbGLÚrIenAkñúgrbbcrose yl;RBbTI 17-2¦ pøeQI nig bEnøydrate Edlm

353/011717361,

grukçCatiedayains and cer

ohydrates Cv)aneRbIedayC

rbs; carbo

rUbTI 17

lthy diet) Rs) dUcCa fruc

yedaybrimaN

gCagsársamBld¾l¥ . bEnß

r .

as;las;nUvGIV percentage)

Gahar eTA 4bGaharRtUvEtRBm[eRbI {Ba. GaharenA)nø RtUvEtsMbUr

mankñúgrbbGah

e-mail: sophea

ysarrs μ IsMeyareals) nMb½ug GCaBiessKøúykCalikaepSgeTohydrate b

7-1

RtUvEtmanTaMg ctose nig su

NeRcInén suc

BaØ . GamIdugnmBIenH carbo

EdlPaK) ERbRbYlya:40°-50° e

t)anmkBI carb

r:amIdcMNIGah)atxageRkamCageKbMputenharrbs;eyIg

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aK ¬rUbTI 17GMeBA EpøeQI TwkUs (glucose

TotCaeRcIn .eBa©jfamBl

carbohydr

ucrose . b:ucrose enAkñúg

gRtUv)aneKeXIohydrates cM

ryénrbbGahagxøaMgenAkñúgBienAkñúgshrdæGrbohydrates

har} edIm,IbgðaménBara:mId gr

nAkñúgrbbGah.

Carboh

com Pag

7-1¦ . RbPTwkedaH nig Twe) KWCaRbPB

. enAeBlRtUvlRbEhl 4 k

ates cMruHdUcCbEnþ brimaNrbgrbbGaharbg

jmanenAkñúg cruH dUcCa cell

harRbcaMéf¶ BiPBelak BI Gaemrik . bnig minRtUveR

ajBIcMnYnGahrains ¬nMb½ug

harrbs;eyIg

hydrates

ge 3 of 20

BFmμCatirbs;TwkX μ úM EdlKWCafamBlTImYy)an {duteqH}kilocalories

CaGamIdug nigbs;sársamBaØgá[elIsTMgn;

)ay dMLÚg-lulose pþl;[

RtUvEtman80° enAkñúg

bc©úb,nñenH eKRcInCag 10°

harepSg²EdlRKab;FBaØCati

. RkumGahar

s a

y } s

g aØ n

-[

n g K ° l ti r

Bioch

prepar

1-Rb BIeRBalacto

Glyc

glyco

Edlvglyco

2-M

eday UkarsMmono

ekasicarbo

DNA

hemistry, Basic

red by Mr. Chro

bePTrbs; C

Monosa

aHvamanÉktasose manBIr m

cosidic bon

osidic bond

vamanExSEvg ogen nig cel

MONOSACHA

Monosac

yrUbmnþTUeTA (CsrYl[RsYl

osaccharide

ska)ak;etrIRtUvohydrate ma

A manGatUmGu

c

oeng Sopheap, p

CARBOHYD

accharides

sár sugar (s

monosaccha

nd . Olig

ds . Carboh

Cajwkjab;EluloseKWCa]TARIDES

ccharides ekICH2O)n EdlGaRs½yeday

es . edays)anCMnYseTAeanbgÁúMhVÚsVat (

GksIuEsnmYyti

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DRATES

dUcCa gluco

saccharide)

arides cgP¢agosaccharid

hydrates EdEbkecjCaExñgTahrN_énBYk

kIteLIgBI kael n KWCacMnYnKy monosa

sarehtuenH m

day amin

(phosphate

ticCagrUbmnþE

353/011717361,

rUbTI 17

ose nig fruc

EtmYyKt; (

ab;Kñatamry³des man m

lFMCageKbMpugCaeRcInénBYk

k polysaccha

)an GIuRdUEsnKt;BIelx 3 eTaccharides

monosacchar

no group

group) . DdleKTajTuk

e-mail: sophea

7-2

ctose KWCa c

(mono) . D

s<anGatUmGukmonosacchar

t nig s μ úRKs μk monosa

arides .

nig GuksIuEsTA elx 7 .

Edlmansarrides én blo

ps . FDeoxyribose

k .

[email protected]

carbohydrate

Disaccharid

ksIuEsn . srides BI 3

s μajCageKbMpuccharides.

sn nig PaKeRcIdUcEdleyIgnwr³sMxan;kñúgCIvood group a

FatukNþalCaee Ca monosa

Carboh

com Pag

es Edlsamdes rYmman sm<½n§enHRtUv)a3-10 cgP¢tKWCa Polys

GamIdug (

cInRtUv)anbBa¢nwgeXIj rUbmvsa®sþKWCaTMrgantigen CaeReRcInenAkñúgemaccharide Ed

hydrates

ge 4 of 20

BaØCageKbMputsucrose nig

)aneKehAfaCaP¢ab;Kñatamry³

accharides

(startch) ,

a¢ak;lkçN³eTAnþTUeTAenHKWCag;EksMrYlrbs;RcIn nig Pñas

mtabUlIsrbs;dlmanenAkñúg

t g a ³ s

,

A a

s s s g



1-namvlI Monosaccharides GacRtUv)aneK[eQμaHedaysMGageTAelImUldæanénbgÁúMnaTI EdlBYkvaman . monosaccharides bgÁúM ketone (carbonyl) KWCa Ketose . RbsinebImanbgÁúM aldehyde (carbonyl) vaRtUv )aneKehAfaCa Aldose . GaRs½yeday monosaccharides manEdrnUvbgÁúM Hydroxyl CaeRcIn BYkvaCYnkal RtUv)aneKehAfaCa Polyhydroxyaldehydes b¤ Polyhydroxyketones .

H CH2OH │ │ C═O C═O │ │ H─ C─OH H─C─OH │ │ H─C─OH HO─C─H │ │ CH2OH CH2OH

H CH2OH │ │ C═O C═O │ │ Aldehyde functional group Ketone functional group

RbB½n§mYyepSgeToténkarehAeQ μaHR)ab;eyIgBIcMnYnGatUmkae)anenAkñúgExSem (main skeleton) . monosaccharides EdlmanGatUmkae)anbIeKehAvafaCa Triose sárEdlmanbYnGatUmkae)anKWCa Tetrose

sárEdlmanR)aMGatUmkae)anKWCa Pentose sárEdlmanR)aMmYyGatUmkae)anKWCa Hexose nig bnþeTAeTot . kar dak;rYmKñanUvRbB½n§ehAeQμaHBIrpþl;B½t’manrwtEteRcInGMBIrcnasm<½n§ nig smasPaBrbs;sármYy . Ca]TahrN_ aldotetrose KWCasárEdlman 4 GatUmkae)anEdlCa aldehyde .

bEnßmBIelIeQμaHTUeTATaMgenH monosaccharide nimYy²maneQμaHEtmYyrbs;vapÞal; . eQμaHTaMgenH RtUv)anbgðajCamYynwg rcnasm<½n§xageRkam . GaRs½yeday monosaccharides GacmanTMrg;GIusUEmrxus²Kña CaeRcIn (isomeric forms) vamansar³sMxan;kñúgkarpþl;eQμaHeBjelj . ehtudUecñH eQ μaHeBjeljénrcna-sm<½n§enATMB½rbnÞab;KW D-glyceraldehyde, D-glucose, nig D-fructose . eQμaHTaMgenHR)ab;eyIgfarcna-sm<½n§bgðajnUvsárBiessmYy nig k¾kMNt;GtþsBaØaNsárfaCaTMrg;GIusUEmmYyNapgEdr (D- b¤ L-) .

Bioch

prepar

edIm,IksmaCiGIusUEmrbs;G

KUrGIuslMhrm:UelKRkumTsm<½n§ YsIuKñaCTMnak;17-3ehABY

hemistry, Basic

red by Mr. Chro

2-Stereo

rcnasm<½n§buBVbT D

kMNt;GtþsBaØCiknimYy²énmYmres‘rI D- xusGatUmenAkñúgm:U

Stereoch

sUEmr GatUmeT . ]TahKulTUeTA C-a

TaMg 4 xus²KñrYmKñatamry³

CamYybgÁúMnaTImkTMngrUbPaBqøú3 b¦ . Ste

BkvamYyKUenHf

c

oeng Sopheap, p

oisomers

n§ - nig L- RtUvaØaNTMrg;GIusUEmmYyKUr stereo

sBIGIusUEmres‘m:UelKul .

hemistry WmTaMgGs;cgP¢ahrN_TUeTAénKUabcd RtUv)an

Ka a, b, c nig³KMrUsm<½n§KImIdUcmYyepSgeTotHkBa©k;KñaEdl

ereoisomers

faCa enantio

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v)aneKBinitüemrmYykñúgcMeNoisomers RtU

s‘rI L- y:agdUc

KWCakarsikSaGab;KñaedayKMrUs

KUr stereoiso

nbegáIteLIgBIkg d . karenHcKña eTaHbICa

t . ehtudUecñHldUceTAnwgTMna

BIEdlminRtYomers . mYy

353/011717361,

eXIjenAkñúgeQNamGIusUEmrBItUvEtmanrUbm þcemþc ? GIusUEm

GMBIkarerobcMkñúsm<½n§KImIRtwmRomers RtUv)akarcgsm<½n§KIHbNþal[maBYkvaTaMgGs;miH BYkvaKWCa st

ak;TMngrUbPaBtsIuKñadUcrUbP

yCasmaCikénrUbTI 17

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Q μaHeBjeljrEdleKehAfnþm:UelKuldUcKñmr D- nig GIu

kñúglMhrepSg²RtUvdUcKña BYkanbgðajenAkñúmIrbs;kae)annBIrm:UelKulCmindUcKñak¾edatereoisomer

qøúHKñaénRbGb;PaBqøúHkBa©k;rbn D-family

-3

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jrbs; mono

faCa Stereoi

Kña nig manksUEmr L- xus

²rbs;GatUm kvaxusKñaEtkagrUbTI 17-3enAkNþal (

CaCagmYym:Uely . RbsinebIrs nwgKña . s

b;édeqVg nig bs; stereoiso

nig mYyeTotC

Carboh

com Pag

saccharide

isomers . karcgsm<½n§KImIsKñaEtkarerob

.enAkñúgsmaarerobcMGatUm3 . enAkñúgcentral carb

lKul . GIusUEbIBYkvadUcKña Btereoisomer

RbGb;édsþaMomer mYyepSCasmaCikén

hydrates

ge 6 of 20

RtUv)aneKeRbItamniymn½y

mIdUcKña . etIbcMenAkñúglMhr

aCiknimYy²énmrbs;vaenAkñúgg]TahrN_enHrbon) eTAnwgEmrnimYy²cgBYkvaKWCa RtYtrs TaMgBIrmanaM ¬emIlrUbTISgeTotRtUveKL-family .

bI y tI r

n g H g g t n TI K

Bioch

prepar

mole

(3-D

RtYtsI carb

cUrBiccUrkt;eyIgKU

TI 17dUecñHvx<s;bMxøaMgCasres

hemistry, Basic

red by Mr. Chro

m:UelKulEdecules . B) rbs;GatUm

sIuKña (enantio

GatUmkae)bone atom

carNa carb

tsMKal;faGatUKUs)an enant

3-Mono

a- Glyce

Carbohy

7-4¦ . D-gl

vaKWCa aldotr

put (the mo

ageKbMputCaTUsr . kae)ane

c

oeng Sopheap, p

dlGacmanrUbBaküenHmann½y

m:UelKulGacomers) CaTUeT)an Edlman

. m:UelKulohydrate

tUmkae)anRtUvctiomers BIrrb

osaccharide

eraldehyde

ydrate EdltUlyceraldehy

iose . CadMbUost oxidized

TUeTAsßitenAcugenHRtUv)aneKd

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bPaBqøúHkBa©k;yedaysamB ØcmanTMrgrUbPTAeKerobcMsMrnbgÁúMnaTIbYnxuslNamYyEdlEdltUcCageKcgP¢ab;eTAnwgbs; glyceral

es Edlmans

cCageKbMputKWyde KWCa trios

bUg cUrktsMKad carbon) bgExSxagelIbMpdak;elx[Ca

353/011717361,

kKñaminRtYtsIuKñBaØfaCalT§plPaBqøúHkBa©k;BIrrab; D- nig L-

s²KñaP¢ab;CamYman chiral

KbMput glyc

bgÁúMnaTI 4 xuldehyde Edl

rUbTI 17

sar³sMxan;mYy

KWCasárEdlmase nig Ca ald

al;farcnasm<½bgÁúMGal;edGIud putenAeBlrcC1. C2 rb

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Kña TMrg; enan

lénkarerobcMCr . cMeBaHKUrN- epSgeTot .

mYyvaRtUv)aneKcarbon mY

ceraldehyd

s²Kña . dUecñlrUbPaBqøúHkB7-4

ycMnYn

an 3 GatUmkdose ¬manbgÁúMm<½n§enHRtUv)anKUenAcugxagelInasm<½n§rbs;

bs; glycerald

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ntiomers RtUCarUbPaBkñúglMNamYykñúgcMeN.

KehAfaCa a

myGacmanmYye EdlbgðajcñHvaKWCa chir

Ba©k;minRtYtsIuKñ

kae)an ³ D-gl

gÁúMGal;edGIudenKUrCamYyBIkae

lI . GatUmkaecarbohydr

dehyde RtUv)

Carboh

com Pag

tUv)aneKehAflMhr three-d

NamTMrg;rUbPa

asymetric

yKUrén enan

jenAkñúgrUbTI ral carbon

Kña .

lyceraldehy

nAcugxagelIébe)anEdlTTYle)anEdlTTYlrates CaExSe)ancgtamsm<½

hydrates

ge 7 of 20

faCa Chiral

dimensional

aBqøúHkBa©k;min

(or chiral)

ntiomers .17-4 .

. CalT§pl

yde ¬emIlrUbbExS¦ . ehtu

lrgGuksIutkmμlrgGuksIutkmμebIkRtUv)aneKm<½n§kUva:Lg;eTA

l

l

n

)

.

. l

b t mμ mμ K A



nwgbgÁúMnaTI 4 xus²Kña dUecñH C2 vaKWCa chiral carbon . karqøúH D-glyceraldehyde eTAnwgkBa©k;pþl;nUvrUbPaB rbs; L-glyceraldehyde .

edaykarsn μti vaKWCaTItaMgénbgÁúM hydroxyl enAelI chiral carbon EdlenAq¶ayBIbgÁúM carbonyl Cag eKbMput ¬q¶ayBIcugEdlTTYlrgGuksIutkm μCageKbMput¦ EdlkMNt;faetI monosaccharide sßitenAkñúgTMrg; D-

b¤ L- . RbssinebI ─OH sßitenAxagsþaM m:UelKulenHsßitkñúgTMrg; D- . RbsinebI ─OH sßitenAxageqVg m:UelKul sßitkñúgTMrg; L- . esÞIrEtRKb; carbohydrates EdlmanenAkñúgRbB½n§manCIvitKWCasmaCikrbs;GMbUr es‘rI D- (D-family) .

b-Glucose

Glucose KWCasársMxan;CageKbMputenAkñúgsBa’gÁkaymnusS . eKeXIjvaenAkñúgGaharCaeRcIn nig man eQμaHEdleRbIjwkjab;CaeRcIn rYmman destrose, grap sugar, nig blood sugar . Glucose RtUv)anbMEbkenA kñúg glycolysis nig pøÚvepSgeTotedIm,IbeBa©jfamBlsMrab;karbMeBjmuxgarrbs;dgxøÜn .

kMhab;rbs; glucose enAkñúgQamvamansar³sMxan;xøaMgcMeBaHkarbMeBjmuxFmμtarbs;sBa’gÁkay . Ca lT§pl vaRtUv)aneKRtYtBinitüedayRbugRby½tñeday hormon insulin nig glucagon . kMritFmμtarbs; blood

glucose RbmaNBI 100-120mg/100mL (100-120mg/dL) rYmCamYynwgkMhab;x<s;CageKbMputekIteLIg bnÞab;karbriePaKGahar . Insulin eP¶ackareRbIR)as; (uptake) én glucose EdlelIsedayekasikaPaKeRcIn bMputrbs;sBa’gÁkay nig bnÞab;BIry³eBl 1-2 em:ag kMritenHk¾vilRtLb;eTAFm μtavij . RbsinebIkMhab; blood

glucose Føak;cuHTab buKÁlmñak;TTYlGarmμN_minl¥ (lightheaded) nig j½rsac; (shaky) . enAeBlkarFøak;enH ekIteLIg Glucagon eP¶aceføIm[beBa©j glucose eTAkñúgQamedaybegáItkMritFmμtasarCaf μ I . eyIgnwgBinitü emIlBItulükarenH[)ankan;Etl¥enAkñúgemeronTI 23-6 .

rUbmnþm:UelKulrbs; glucose, aldohexose KW C6H12O6 . rcnasm<½n§rbs; glucose RtUv)aneKbgðaj enAkñúgrUbTI 17-5 nig viFIEdleKeRbIedIm,IKUsrcnasm<½n§RtUv)anBiB’Nnakñúg]TahrN_ TI 17-1 .

]TahrN_ TI 17-1 karKUsrcnasm<½n§rbs; monosaccharide

enAkñúgPaBBit TMrg;ExSebIkrbs; glucose manvtþmanenAkñúgkMhab;ticbMputenAkñúgekasika . vamanTMrg; ExSbiT (cyclic form) sßitenAeRkamlkçxNÐsrIr³viTüa (physiological conditions) GaRs½yedaybgÁμM Carbonyl enA C-1rbs; glucose manRbtikmμCamYynwgbgÁúM hydroxyl enA C-5 edIm,Ipþl;[CaExSbiTEdlman GatUmkae)an 6 . kñúgkarBiPakSaBI aldehydes eyIgkt;sMKal;eXIjfaRbtikmμrvag aldehyde nig alcohol

[eTACa hemiacetal . enAeBl aldehyde portion rbs; glucose manRbtikmμCamYynwg bgÁúM hydroxyl enA C-5 plitplénRbtikmμenHKWCa Cyclic intramolecular hemiacetal . cMeBaH D-glucose GIusUEmrBIrGac RtUv)anbegáIteLIgenAkñúgRbtikmμenH ¬rUbTI 17-5¦ . GIusUEmrTaMgBIrenH RtUv)aneKehAfaCa α- nig β-D-

glucose . GIusUEmrTaMgBIrRtUv)anbegáIteLIgGaRs½yedaysarRbtikmμénkarbegáItCaExSbiTbegáItnUv chiral

Bioch

prepar

carbo

enAkñúg

RtUv)aNas;2.

eKeXIrbs; CHO

hemistry, Basic

red by Mr. Chro

on fμ ImYyeTotgGIusUEmr β b

enAkñúgrUbTIanbgðaj . e

skñúgkarTajy

]TahrN_ karKUr Ha

c-Fructo

Fructose

XIjmanvaCacMnYglucose .

OH enAelI C-

karbegáItC

c

oeng Sopheap, p

t kñúgkrNIenHbgÁúM hydroxyl

TI 17-5 ReTaHbICa enAkñ

yk Haworth

17-2 aworth proje

ose

e RtUv)aneKehnYneRcInenAkñúgenAeBlman -2 dUecñHsárenCaExSbiTén fr

phone 0128643

HKW C-1 . enrbs; C-1 sßi

RbePTfμ ImYyénkñúgkarBinitüCah projection

ection rbs; m

hApgEdrfaCaTwkX μ ú M corn

bgÁúM ─CH2O

HKWCa Ketose

ructose begáIt

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nAkñúg GIusUEmr sßitenAxagelIE

rUbTI 17

nrUbmnþrcnasabzm vahak

n BIrUbmnþrcn

monosaccha

a levulose nisyrup nig Epø

OH CMnYs[bge kñúgkrNIenHt)anCa α- nig

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α bgÁúM hydro

xS .

7-5

sm<½n§ RtUv)anek;dUcCas μ úRKsμaasm<½n§lat dU

arides BIrUbm

nig fruit sug

pøeQIEp¥m . rgÁúM ─CHO envaKWCa D-fruc

g β-D-fructo

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oxyl rbs; C

eKehAfaCa ajk¾eday b:u

dUcEdl)anbgð

mnþrcnasm<½n§l

ar manrsCatircnasm<½n§ frunAelI C-1 nictose .

ose :

Carboh

com Pag

-1sßitenAxag

Haworth

EnþvamanlkçNgðajenAkñúg[T

lat

tiEp¥mdUcsárdéuctose dUceTAg bgÁúM ─C═

hydrates

ge 9 of 20

geRkamExS nig

projection

N³samBaØxøaMgTahrN_ 17-

éTeTotEdr .Anwgrcnasm<½n§═O CMnYn[bgÁúM

g

n

g -

. n§ gÁM



α-D-fructose

β-D-fructose

α-D-G l t

β-D-Galactose

CH2OH │ C═O │ HO─C─H │ H─C─OH │ H─C─OH │ CH2OH

Fructose KWCa ketose b¤ Ca ketone sugar . cUrrMlwkfaRbtikm μrvag alcohol nig ketone pþl;[Ca Hemiacetal . ehtudUecñH Rbtikm μrvag C-2 keto group nig C-5 hydroxyl group enAkñúg fructose

molecule begáIt)anCa Intramolecular hemiacetal . Fructose begáIt)anCarcnasm<½n§ExSbiTEdlman R)aMRCug .

d-Galactose

hexose EdlsMxan;mYyeTotKW Galactose . rcnasm<½n§ExSlatrbs; D-galactose nig Haworth

projection rbs; α-D-galactose nig β-D-galactose RtUv)anbgðajdUcxageRkam ³

CHO │ H─C─OH │ HO─C─H │ HO─C─H │ H─C─OH │ CH2OH

O

H HO

OH H

OH

CH2OH

H

HOH2C HO

O

H HO

OH H

OH

CH2OH H

HOH2C HO

O

OH H

H OH

OH H

HOH2C HO

H

HO

H

O

OH H

H OH

OH

H

HOH2C HO

H

HO H



β-D-N-acetyl-galactosamine

α-D-Ribose

β-D-Ribose

Galactose RtUv)aneKeXIjmanenAkñúgRbB½n§CIvsa®sþsmaFaturbs; disaccharide KW Lactose b¤ milk

sugar . vaenH (lactose) KWCasársMxan;EdleKeXIjmanenAkñúgTwkedaHénRKb;BYkfnikstV . β-D-galactose

nig TMrg;bMErbMrYl β-D-N-acetyl-galactosamine k¾CasmasFaturbs; blood group antigen .

e-Ribose and Deoxyribose, Five Carbon Sugar

Ribose KWCasmasFatuémm:UelKulsMxan;CIvsa®sþCaeRcIn rYmman RNA nig Coenzymes epSg² ¬smasPaBmYyRkumEdlRtUvkarcaM)ac;edayGg;sIumCaeRcInEdlGnuvtþnUvRbtikm μCIvKImIenAkñúgsBa’gÁkay . rcna sm<½n§énsárman 5 GatUmkae)an D-ribose RtUv)anbgðajCaTMrg;ExSlat nig CaTMrg;ExSbiT α- nig β- .

O ║ C─H │ H─C─OH │ H─C─OH │ H─C─OH │ CH2OH

DNA m:UelKulEdlmanpÞúkB½t’manEhSnrbs;ekasika man 2-deoxyribose . enAkñúgm:UelKulenHbgÁúM ─OH enAelI C-2 RtUv)anCMnYseday Hydrogen ehtudUecñH namktþa (designation) “2-deoxy” bBa¢ak;BI Gvtþmanrbs;GuksIuEsn .

O

OH H

H

OH H

HOH2C HO

H

HO

H

NH │ C═O │ CH3

O

H H

OH OH

OH

H

H

HOH2C HO

O

H H

OH OH

OH

H H

HOH2C HO



β-D-2-Deoxyribose

4-sárer:dukr (Reducing sugars)

bgÁúMGal;edGIudrbs; Aldose TTYlrgGuksIutkmμedayRbtikebeNDik (Benedict’s reagent) . cUr rMlwkfa RbtikrebeNDik KWCasUluysüúgRTnab;)as (basic buffer solution) EdlmanGIuy:ug Cu2+ . GIuy:ug Cu2+ RtUv)anTTYlrger:dukkmμeTACa Cu+ EdlenAkñúgsUluysüúg)as [Cakkr Cu2O B’NRkhmdUcek,Óg . bgÁúM aldehyde rbs; aldose RtYv)anTTYlrgGuksIutkm μeTACa carboxylic acid EdlTTYlrgnUvRbtikmμ GasIut-)as (acid-base reaction) edIm,IbegáItCa carboxylate anion .

O O ║ ║ C─H C─O │ │ H─C─OH + 2Cu2+ (buffer) + 5OH → H─C─OH + Cu2O + 3H2O │ │ CH2OH CH2OH

eTaHbICaTUeTA ketones mingayRsYlkñúgkarTTYlGuksIutkmμ b:uEnþ Ketones KWCakelIkElgcMeBaH viFan enaH . GaRs½yedaybgÁúM ─OH EdlenAelIGatUmkae)anenACab;nwg bgÁúM Carbonyl , Ketones GacRtUv)an bMElgeTACabgÁúM Aldehyde enAeRkamlkçxNÐ)as tamry³Rbtikmμ Enediol nediol Reaction) :

H │ CH2OH HO─C─H C═O │ ║ │ C═O C─OH H─C─OH │ │ │ HO─C─H HO─C─H HO─C─H │ │ │ H─C─OH H─C─OH H─C─OH │ │ │ H─C─OH H─C─OH H─C─OH │ │ │ CH2OH CH2OH CH2OH D-fructose ` Enediol D-gucose

eQμaHRbtikmμ enediol manRsLayBIrcnasm<½n§énFatukNþalEdltamry³ ketose RtUv)anbMElgeTA Ca aldose : vamansm<½n§BIrCan; (ene) nig vamanbgÁúM hydroxyl BIr (diol) . GaRs½yedayRbtikm μ enediol enH ketoses k¾GacmanRbtikmμCamYynwgRbtikrebeNDik EdlmanlkçN³Ca)as . GaRs½yedayGIuy:ugelah³enA

O

H H

OH H

H

OH

H

HOH2C HO



kñúgsUluysüúgTTYlrger:dukkmμ sárRtUv)aneRbICaPñak;garer:dukm μ nig RtUv)aneKehAfaCa sárEdleFIVer:dukkm μ (Reducing sugars) . RKb; monosaccharides nig RKb; disaccharides EdlCYbjwkjab;elIkElgEt sucrose ¬sárs¦ KWCasárEdleFIVer:dukkmμ .

Gs;ry³eBlCaeRcInqμaM RbtikrebeNDikRtUv)aneKeRbIsMrab;eFIVkarviPaKrk Glucosuria ¬vtþmanén karelIs glucose enAkñúgTwkenam¦ . buKÁlEdlmanCMgWTwkenamEp¥mRbePTTI 1 (Type I insulin-dependent

diabetes mellitus) minbegáIt Hormone Insulin EdlCaGñkRtYtBinitükareRbIR)as; glucose BIkñúgQam . enAeBlkMrit glucose kñúgQamekIneLIgelIsBI 160 mg/dL tMrgenamminGacRsUbyk glucose EdlelIsGs; eT nig ehtudUecñH glucose RtUv)aneKrkeXIjenAkñúgTwkenam . eTaHCakMrit glucose enAkñúgQamGacRtUv)an RtYtBinitüedaykarcak; insulin k¾eday k¾b:uEnþkMrit glucose enAkñúgTwkenamRtUv)aneKtamdanedIm,IR)akdfacMnYn insulin cak;)anRtwmRtUv . RbtikrebeNDikKWCa]bkrN_mansar³sMxan;GaRs½yedaycMnYn Cu2O ekIteLIg nig ehtudUecñH kMritB’Npøas;bþÚrenAkñúgRbtikmμsmamaRtedaypÞal;eTAnwgcMnYnsárEdleFIVer:dukkmμenAkñúgTwkenam . B’NRkhmdUcek,ÓgbgðajfakMhab; glucose kñúgTwkenamx<s;xøaMgNas; . sUluysüúgB’NelOg ébtg exov-ébtg bgðajBIcMnYnfycuHtamlMdab;én glucose EdlmankñúgTwkenam nig sUluysüúgB’Nexov bgðajfaminman glucose enAkñúgTwkenameT . kareFIVkarviPaKrk glucose kñúgTwkenam)anRtUvCMnYsedaykarkMNt;brimaN glucose enAkñúgQamEdlpþl;nUvkarbgðajcMTisedACagfaetIGñkCMgWTwkenamEp¥mRtYtBinitürbbGaharrbs;Kat; y:agdUcemþc .

3-DISACCHARIDES

cUrrMlwkfa disaccharides man monosaccharides BIcgP¢ab;Kñatamry³{s<anGuksIuEsn} . enAkñúg RbB½n§CIvsa®sþ monosaccharides sßitenACaTMrg;ExSbiT nig dUcEdleyIg)aneXIj CaFmμtaBYkvaKKWCa hemiacetals b¤ hemiketals . cUrrMlwkfa enAeBl hemiacetal manRbtikmμCamYyGal;kul plitplKWCa acetal nig enAeBl hemiketal manRbtikmμCamYynwgGal;kul plitplKWCa Ketal . enAkñúgkrNI disaccharides Gal;kulEdl)anmkBI monosaccharide TI 2 . Acetal b¤ Ketal EdlekIteLIgRtUv)aneKpþl; eQμaHTUeTAfaCa Glyosides nig sm<½n§rvag C-O RtUv)aneKehAfaCa Glycosidic bonds .

karbegáItsm<½n§ Glycosidic minBiesseT dUecñH vaGacekIteLIgrvag hemiacetal b¤ hemiacetal nig bgÁúM hydroxyl NamYyenAelI monosaccharide TI 2 . b:uEnþ enAkñúgRbB½n§CIvsa®sþ eyIgeXIjedayjwkjab;Et disaccharides BiessdUcCa maltose ¬rUbTI 17-6¦ Lactose ¬rUbTI 17-8¦ b¤ Sucrose ¬rUbTI 17-9¦ . Disaccharides BiessTaMgenHRtUv)anbegáIteLIgenAkñúgekasika GaRs½yedayRbtikmμRtUv)ansMrYleday Gg;sIum. Gg;sIumnimYy²sMrYlkarsMeyaK disaccharide BiessmYy edayFanafa mYyKUrénbgÁúM hydroxyl enA elI monosaccharides EdlcUlrYmRbtikmμcUlrYmkñúgkarbegáItsm<½n§ glycosidic .

Maltose

RbsinebI α-D-glucose nig glucose TI 2 RtUv)ancgP¢ab;Kña dUcEdl)anbgðajenAkñúgrUbTI 17-6 disaccharides enHKWCa Maltose b¤ Malt sugar . vaenHKWCaplitplkNþalénkareFIV hydrolysis rbs;



α-D-Glucose β-D-Glucose

GamIdug . GaRs½yedaybgÁúM hydroxyl enAelI C-1 rbs; α-D-glucose RtUv)anP¢ab;eTAnwg bgÁúM hydroxyl enA elI C-4 rbs; glucose mYyepSgeTot disaccharide enHRtUv)anP¢ab;tamry³sm<½n§ glycosidic α (1→4) .

Maltose KWCasárEdleFIVer:dukkm μ (reducing sugar) . disaccharide mYyNaEdlmanbgÁúM hemiacetal hydroxyl ¬bgÁúM ─OH esrIenAelI C-1¦ KWCasárEdleFIVer:dukkmμ . karenHGaRs½yeday rcnasm<½n§ExSbiTGacebIkenAkñúgTItaMgenHedIm,IbegáItCa aldehyde esrI . Disaccharides EdlminmanbgÁúM hemiacetal enAelI C-1 minmanRbtikm μCamYynwgRbtikrebeNDik nig RtUv)aneKehAfaCa sárEdmineFIVer:dukkmμ (nonreducing sugar) .

Lactose

sárTwkedaH b¤ Lactose KWCa disaccharides EdlekIteLIgBImYym:UelKulrbs; β-D-galactose nig mYyeTotGacCa α- b¤ β-D-glucose . Galactose xusBI glucose edaysarEtTItaMgrbs;bgÁúM hydroxyl

enAelI C-4 ¬rUbTI 17-7¦ . enAkñúgTMrg;ExSbiTrbs; glucose bgÁúM hydroxyl enAelI C-4 enAxag {eRkam} nig enAkñúg galactose enAxag {elI} . enAkñúg lactose bgÁúM hydroxyl enAelI C-1 rbs; β-galactose cgeTAnwg bgÁúM hydroxyl enAelI C-4 TaMg α- b¤ β-D-glucose . sm<½n§rvag monosaccharides TaMgBIrdUecñHvaKWCa sm<½n§ glycosidic β-(1→4) ¬rUbTI 17-8¦ .

rUbTI 17-6

Maltose

O

OH H

H OH

OH H

HOH2C HO

HOOO

H

H

O

OH H

H OH

OH

H

HOH2C HO

HO

H H

O

OH H

H OH

OH H

HOH2C HO

H

H

O

OH H

H OH

H

HOH2C HO

HO

H H

O



rUbTI 17-7

rUbTI17-8

Lactose KWCasársMxan;énTwkedaHBYkfnikstV . edIm,IeRbIenAkñúgsBa’gÁkayCaRbPBfamBl Lactose

RtUvEtTTYlrgnUv hydrolysis edIm,IbegáItCa glucose nig galactose . cUrkt;sMKal;fa karenHvaRsYl Edl RbtikmμRtLb;RtUv)anbgðajenAkñúgrUbTI 17-8 . Glucose beBa©jeday hydrolysis én lactose RtUv)aneRbI edaypÞal;enAkñúgRbtikmμRbmUlfamBlrbs; glycolysis (energy-harvesting reaction of glycolysis) . b:uEnþ RbtikmμmYyes‘rIRtUvkarcaM)ac;edIm,IbMElg galactose eTACa glucose eRkamTMrgman Phosphate

(phosphorylated form of glucose) EdlGacRtUv)aneRbIenAkñúgRbtikmμemtabUlIsekasika . cMeBaHmnusS

α-D-Glucose β-D-Galactose

O

OH H

H OH

OH H

HOH2C HO

H

HO

H

O

OH H

H OH

H

H

HOH2C HO

HO

H OH

α-D-Glucose β-D-Galactose

O

OH H

H OH

OH H

HOH2C HO

H

HO

H

O

OH H

H OH

H

H

HOH2C HO

HO

H OH

β-Lactose

O

OH H

H OH

H

HOH2C HO

H

HO

H

O

OH H

H OH

H

H

HOH2C HO

H OH O



CMgWEhSn (genetic disease) Galactosemia bgáedayGvtþmanGg;sIummYy b¤ eRcInEdlRtUvkarsMrab;karbMElg enH . smasPaBBulEdlbegáIteLIgBI galactose RbmUlpþúMeRcIncMeBaHmnusSEdlTTYlrgCMgW galactosemia . RbsinebICMgWenHminRtUv)aneKBüa)al Galactosemia bNþal[mankaryWty:avxags μartIF¶n;F¶r (severe

mental retardation) eLIg)ayEPñk (cataract) nig søab;munGayu (early death) . b:uEnþ plb:HBal;énCMgWenH GacRtUv)aneCosvagTaMgRsugedaykarpþl;[kumarEdlmanCMgW galactosemia (galactosemic infants) nUvrbb GaharEdlminman galactose . dUecñH rbbGaharrebobhñwg minGacman lactose nig ehtudUecñHRtUvEtminman TwkedaH nig plitpleFIVBITwkedaH .

mnusSeBjv½yCaeRcIn nig kumarmYycMnYn minGaceFIV hydrolyse lactose eTGaRs½yedayBYkeKminGac plitGg;sIum Lactase . CMgWenH Edlb:HBal; 20° énRbCaCnenAkñúgshrdæGaemrik RtUv)aneKsÁal;faCa Lactose intolerance . Lactose EdlminRtUv)anbMEbksßitenAkñúgeBaHevon nig bgá[mankarQWeBaH nig raK EdlCacugbBa©b;bNþal[mankar)at;bg;CatiTwk (dehydration) . Lactose mYycMnYnRtUv)anTTYlrg emtabUlIseday)ak;etrIenAkñúgeBaHevonedaybeBa©jGasIutsrIragÁ nig CO2 enAkñúgeBaHevon edaybegáIt[man PaBminRsYleRcIneTot . Lactose intolerance KWCakarminRsYlxøÜn b:uEnþkarb:HBal;rbs;vaGacRtUveCosvag)an edayrbbGaharEdlminmanTwkedaH b¤ plitpleFIVBITwkedaH . m:üagvijeTot Gg;sIumEdleFIV hydrolyse

lactose maneRkamTMrg;Ca»sRKab; (tablet form) . enAeBlRtUv)anrMlayCamYynwgplitplTwkedaH vabMEbk lactose edaykarmin[maneraKsBaØaTaMgenH .

Sucrose

Sucrose k¾RtUv)aneKehApgEdrfaCasárs (table sugar) sárGMeBA (cane sugar) b¤ sárEbttar:av (beet sugar) . Sucrose KWCa carbohydrate Edlmansar³sMxan;kñúgrukçCati . varlaykñúgTwk nig Gacgay RsYldwknaMtamry³RbB½n§rukçrs (circulatory system of the plant) . vaminRtUv)ansMeyaKedaystVeT . kMhab;x<s;rbs; sucrose begáIt[)ansMBaFGUsμ Úx<s; EdlraraMgkarlUtlas;rbs;BYkemeraK (microorganism)

dUecñH vaRtUv)aneKeRbICasarFatuEfrkSa (preservative) . ehtudUecñH vak¾RtUv)aneKeRbIedayTUlMTUlayCa sarFatueFIV[Ep¥m . CakarBit eK)a:n;sμanfaCamFüm CnCatiGaemrikaMgRtUvkarBI 100-125 pounds én sucrose

kñúgral;qñaM . eKpþl;Caeyablfa sucrose enAkñúgrbbGaharminKab,IR)afñaeT BIeRBaHvatMNagRbPBmYyminman kaLÚrI (a source of empty calories) dUecñH vaminmanvItamIn b¤ BYkEr:eT . b:uEnþ karrYmKñaGviC¢manEtmYyKt; (the only negative association) EdlRtUv)anepÞógpÞat;tamlkçN³viTüasa®sþKWCaTMnak;TMng (the link) rvag sucrose enAkñúgrbbGahareTAnwgCMgWdgáÚvsIueF μj b¤ RbehageFμj .

Sucrose KWCa disaccharide Edl α-D-glucose RtUv)anP¢ab;eTAnwg β-D-fructose ¬rUbTI 17-9¦ . sm<½n§ glycosidic rvag α-D-glucose nig β-D-fructose vaxusKñay:agxøaMgBIsm<½n§EdleyIg)ankMNt;enAkñúg lactose nig maltose . kae)anTaMgBIrEdlBImunmkCaEpñkén hemiacetal b¤ hemiketal manRbtikmμCamYyKña edIm,IbegáItCasm<½n§ . sm<½n§rebobenHRtUv)aneKehAfaCa sm<½n§ glycosidic (α1→β2) kñúgkrNIenHvaCab;Tak;Tg eTAnwg C-1 rbs; glucose nig C-2 rbs; fructose ¬sMKal;CaB’NexovkñúgrUbTI 17-9¦ . CalT§plénsm<½n§



glycosidic (α1→β2) rcnasm<½n§ExSbiTminGacebIkCaExSlat)aneT nig minmanbgÁúM aldehyde b¤ Ketone

RtUv)anekIteLIgeT . ehtudUecñH sucrose minmanRbtikmμCamYynwgRbtikrebeNDikeT nig k¾minEmnCasárEdl eFIVer:dukmμEdr .

rUbTI 17-9

4-POLYSACCHARIDES

GamIdug (startch)

Carbohydrates PaKeRcInEdleXIjenAkñúgFmμCatiKWCa polymers FM²rbs; glucose . dUecñH plysaccharide mYyKWCam:UelKulFMmYyénEdlekIteLIgBI monosaccharodes CaeRcIn (monomers) tP¢ab;Kña CaExSEtmYy b¤ eRcInExS .

dUcEdl)aneXIjenAkñúgrUbTI 17-1 rukçCatimanlT§PaBkñúgkareRbIfamBlBn¶WRBHGaTitedIm,IbegáItCa monosaccharides CaBiess glucose BI CO2 nig H2O . eTaHbICa sucrose KWCaTMrg;énkardwknaMrbs;sár enAkñúgrukçCatik¾eday b:uEnþGamIdug (polysaccharide) KWCaTMrg;sþúkTuksMxan;rbs;rukçCatiPaKeRcIn . rukçCati TaMgenH sþúkTuk glucose enAkñúg glucose granules . esÞIrEtRKb;ekasikarukçCatiman glucose granules mYy cMnYn b:uEnþenAkñúgRKab; (seeds) dUcCaRKab;eBatman 80°énTMgn;s¶Ütrbs;ekasikaKWCaGamIdug .

GamIdugKWCa heterogenous material EdlpSMeLIgBI glucose plymers : Amylose nig Amylopectin . Amylose EdlmanRbEhl 80°énGamIdugrbs;ekasikarukçCati KWCa polymer ExSRtg;én m:UelKul α-D-glucose bnþCab;Kñaedaysm<½n§ glycosidic rvag C-1 rbs; glucose mYyeTAnwg C-4 rbs; glucose TI 2 . ehtudUecñH glucose enAkñúg Amylose cgP¢ab;Kñatamsm<½n§ glycosidic α(1→4) . ExSEtmYy

α-D-Glucose

O

OH H

H OH

H

H

HOH2C HO

HO

H OH

O

H HO

OH H

OH

CH2OH H

HOH2C HO

β-D-fructose

O

O

OH H

H OH

H

HOH2C HO

HO

H H

O

H HO

OH H

CH2OH H

HOH2C HO

Sucrose

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phone 0128643

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KlEdlsþúkTukectin . ExSEdlpþl;[cMN. ehtudUecñHm:U

kTukenAkñúgeføImcEdl)aneXIjkñúgQam[esß

353/011717361,

00 m:UelKul ose RtUv)anbgð

rUbTI 17-

sIumBIrRbePT mat;EdlBYkvakñúgExSEvg² edide maltose

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gðajenAkñúgrUbTI-10

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elKulrbs; cnpliteday a

Faturcnasm<½n§dic β (1→4

aCa Fibrils

TEdl cellulo

353/011717361,

KbMput edaymer én β-D

cellulose manalga Valonia

rUbTI 17-

n§rbs;Pñaseka4) GnuBaØati[. Fibrils rwose KWCaFatup

e-mail: sophea

yBitR)akdm:UeD-glucose EnRbEhl 30a man 2600-11

asikarukçCati [m:UelKul ce

rwgxøaMg nig RtUpSMénrcnasm<½

[email protected]

elKulsrIragÁEEdltP¢ab;Kñat000 m:UelKul 0 m:UelKul glu

. rcnasm<½n§miellulose begáItUvcgP¢ab;Kñay:n§Pñasekasika

Carboh

com Page

EdlsMbUrCagetamry³sm<½n§

glucoses b:uEucose .

minmanExñgrbgáItCaExSRtg;Ev

y:agCab;edaysarukçCati .

hydrates

e 19 of 20

eKbMputenAkñúgglycosidic

Enþ cellulose

s; cellulose

vgénm:UelKulsm<½n§GIuRdUEsn

g c

e

e

l n



pÞúyeTAnwg glycogen , amylose nig amylopectin , cellulose minGacRtUv)anbMEbkedayRbdab; rMlayGaharrbs;mnusSeT . ehtuplKWfa mnusSeyIgminGacsMeyaKGg;sIum Cellulase EdlCaGñkeFIV hydrolyze sm<½n§ glycosidic β (1→4) rbs; cellulose polymer . CakarBit manEtBYkstVmYycMnYn dUcCa keNÐór eKa nig BEBGacrMlay cellulose . stVTaMgenHmanenAkñúgRbdab;rMlayGaharnUvBYkmIRkUsrIragÁEdlGac begáItGg;sIum cellulase . sárEdlRtUv)anbeBa©jedaykarbMEbkedaymIRkUsrIragÁenAeBlenaHRtUv)anRsUb nig RtUv)aneRbIedayekasikarbs;stVTaMgenH . cMeBaHmnusS cellulose EdlmanenAkñúgEpøeQI nig bEnøRtUv)aneRbICa sés (fiber) enAkñúgrbbGahar .

Fin

Reference : General, Organic, and Biochemistry 3rd edition, 2001 by Denniston, Topping, Caret page 467-494

CIvKImIviTüa lIBId nig muxgarrbs;BYkvaenAkñúgRbB½n§CIvKImIsaRsþ

erobcMnigbkERbeday»sfkarIRcwgsuPaBTUrsB½Þ 012864353 b¤ 011717361 Page 1 of 42

lIBId nig muxgarrbs;vaenAkñúgRbB½n§CIvsaRsþ (LIPIDS AND THEIR FUNCTIONS IN BIOCHEMICAL SYSTEMS)

KMeragemeron

1- muxgarCIvsaRsþrbs;lIBId

2- GasIutRka (Fatty acids)

k- rcnasm<½n§ nig lkçN³ x- Rbtikm μKImIrbs;GasIutRka K- Eicosanoids : Prostaglandines, Leukotrienes and Thromboxanes

3- KøIesrId (Glycerides)

k- KøIesrIdNWt (Neutral glycerides)

x- hVÚsVÚKøIesrId (Phosphoglycerides)

4- lIBIdminEmnCaKøIesrId (Non-glyceride lipids)

k- sVaMgehÁalIBId (Sphingolipids)

x- esþr:UGIud (Steroids)

5- lIBIdkMupøic (Complex lipids)

6- rcnasm<½n§énPñasCIvsaRsþ k- Fluid Mosaic Structure of Biological Membranes

x- kardwkCBa¢Únqøgkat;Pñas (Membrane Transport)

K- tMrUvkarfamBlsMrab;kardwkCBa¢Ún



vtßúbMNgemeron 1- BiPakSaGMBIlkçN³rUb nig KImI nig muxgarCIvsaRsþénGMbUrnimYy²rbs;lIBId . 2- sresrrcnasm<½n§rbs;GasIutRkaEq¥t nig minEq¥t . 3- eRbobeFob nig EjkKña[c,as;énrcnasm<½n§rbs;GasIutRkaEq¥t nig GasIuRkaminEq¥t . 4- sresrsmIkarKImItMNag[Rbtikm μKImIEdlGasIutRkaTTYlrg . 5- BiB’NnaBImuxgarrbs; Prostagladines . 6- BiPakSaGMBIynþkarEdl Aspirin bnßyPaBQWcab; . 7- KUsrcnasm<½n§rbs; Prostaglandines nig BiPakSaGMBI Amplipathic nature rbs;va . 8- BiPakSaBIcMNat;fñak;TUeTArbs; Sphingolipids nig muxgarrbs;va . 9- KUsrcnasm<½n§énéNVy:Uesþr:UGIud nig BiPakSaGMBImuxgarrbs;G½rm:Unesþr:UGIud . 10- BiB’NnaGMBImuxgarrbs;lIb:URbUetGIunenAkñúgkardwknaM RTIKøIesrId nig kUelesþr:Ul kñúgsBa’gÁkay. 11- KUsrcnasm<½n§Pñasekasika nig BiPakSaGMBImuxgarrbs;va . 12-BiPakSaGMBIkarsayPayedayÉkÉg (passive and facilated diffusion) énsarFatuqøgkat; Pñas ekasika . 13- Bnül;GMBIdMeNIrkarénGUsμ Ú (osmosis) . 14- BiB’NnaGMBIynþkarénGMeBIrbs; Na+-K+ ATPase .



lIBId nig muxgarrbs;vaenAkñúgRbB½n§CIvsaRsþ (LIPIDS AND THEIR FUNCTIONS IN BIOCHEMICAL SYSTEMS)

1- muxgarCIvsaRsþrbs;lIBId (Biological Functionals of Lipids)

CaFm μtaBaküfa lIBId (Lipids) sMedAcMeBaHRbCMuénmU:elKulsrIragÁénsmasFatuKImIepSg² . BykvaRtUv)andak;CaRkumrYmKñaedaysMGageTAelIPaBrlayrbs;enAkñúgFaturMlayEdlK μan (Non-polar

solvents) . edayjwkjab; lIBIdRtUv)ancat;fñak;CabnþeTACabYnRkumsMxan; ³ 1- GasIutRka (fatty acids) ¬Eq¥t nig minEq¥t¦

2- KøIesrId (Glycerides)

3- lIBIdminEmnKøIesrId (Nonglyceride lipids)

4- lIBIdkMupøic (Complex lipids)

enAkñúgCMBUkenHeyIgBinitüGMBIrcnasm<½n§ lkçN³ RbtikmμKimI nig muxgarCIvsaRsþrbs;RkumnimYyénlIBId dUcEdl)anbgðajenAkñúgrUbTI 18-1 .

rUbTI 18-1 cMNat;fñak;lIBId

Lipids

Fatty acids Glycerides

Saturated Unsaturated Natural Glycerides

Phospho-Glycerides

Nonglycerides Lipids

Complex Lipids

Waxes Steroids Sphincgolipids Lipoproteins

Glycolipids Sphincgomyelins



CalT§plénPaBxusKñaenAkñúgrcnasm<½n§rbs;BYkva lIBIdmanmuxgarepSg²CaeRcIn²enAkñúg sBa’gÁkaymnusS . karerobrab;segçbxageRkamenH nwgpþl;[GñknUvKMnitGMBIsar³sMxan;rbs;lIBId enAkñúgdMeNIrkarCIvsaRsþ .

RbPBfamBl (Energy source) dUcCakabUGIuRdat b¤ KøúysIut lIBIdKWCaRbPBfamBld¾l¥ sMrab;sBa’gÁkay . enAeBlTTYlrgGuksIutkm μ 1 g rbs;xøaj;beBa©jfamBl 9 kcal b¤ eRcInCag BIrdgén famBlEdlbeBa©jedayGuksIutkm μén 1 g rbs;KøúysIut .

sþúkTukfamBl (Energy storage) PaKeRcInénfamBlEdlsþúkTukenAkñúgsBa’gÁkay KW sßitenAeRkamTMrg;CalIBId ¬RTIKøIesrId : [Triglycerides]¦ . edayRtUv)ansþúkTukenAkñúgekasikaxøaj; RtUv)aneKehAfaCa GaDIb:UsIut (Adipocytes) xøaj;TaMgenHKWCaRbPBfamBlEdlsMbUrCaBiesssMrab; sBa’gÁkay .

CaFatupSMénrcnasm<½n§Pñasekasika (Cell membrane structural components) Phospho-

glycerides, sphincgolipids nig steroids begáItnUvrcnasm<½n§RKwHénRKb;Pñasekasika . Pñas TaMgenHRtYtBinitükarhUrecjcUlénm:UelKuleTAkñúg nig ecjBIekasika nig GnuBaØati[mankarTak;Tg BiIekasikamYyeTAekasikamYyeTot .

G½rm:Un (Hormone) G½rm:Unesþr:UGIud (steroid hormones) CaGñknaMsarKImId¾sMxan; (Critical chemical messengers) EdleFIV[CalikaénsBa’gÁkayedIm,IeFIVTMnak;TmngCamYyCalikaepSgeTot . Prostaglandines man\T§iBlCIvsaRsþxøaMgeTAelITaMgekasikaEdlplitBYkva nig ekasikaepSgeTot rbs;sBa’gÁkay .

vItamIn (Vitamin) vItamInrlayenAkñúgxøaj; (lipi-soluble vitamins) A, D, E nig K man tYnaTIsMxan;enAkñúgkarEktMrUv (regulation) éndMeNIrkarCIvsaRsþsMxan;²CaeRcIn rYmmankarkkQam nig karemIl (Blood clotting and Vision) .

karRsUbcUlénvItamIn (Vitamin absorption) xøaj;éncMNIGaharRtUv)aneRbICaGñkdwknaM vItamIn EdlrlayenAkñúgxøaj; . vItamInTaMgGs;enHRtUv)andwknaMcUleTAkñúgekasikarbs;eBaHevontUc edayrYm CamYym:UelKulxøaj; . ehtucUecñHGaharEdlmanxøaj;ticeBkGacbNþal[mankarxVHxatén vItamInTaMg bYnRbePTenH .



karkarBar (Protection) xøaj;RtUv)aneRbICaRsTab;karBarsMrab;srIr³manCIvit (Vital organs) . RbEhl 4 % énxøaj;srubrbs;sBa’gÁkaybMrugTuksMrab;muxgarsMxan;enH .

eRbICaGIusULg; (Insulation) xøaj;EdlsþúkTukenAeRkamEs,k (subcutaneous fat) eRbIsMrab; karBarsBa’gÁkayBIsItuNðPaBRtCak;xøaMg .

2- GasIutRka (Fatty Acids)

k- rcnasm<½n§ nig lkçN³ (Structure and Properties)

GasIutRka (Fatty acids) KWCaGasIutm:UNUkabuksIulik (Monocarboxylic acids) manExSEvg . Caplvi)ak CivsMeyaKrbs;va CaTUeTAGasIutRkamancMnYnGatUmkae)anKt;KU . rUbmnþTUeTAén GasIutRka Eq¥t KW CH3(CH2)nCOOH Edl “n” enAkñúgRbB½n§CIvsaRsþKWCacMnYnKt;KU (even integer) . RbsinebI n=16 lT§plKWCaGasIutRkaEq¥tEdlman 18 GatUmkae)an maneQμaHfaCa GasIutesþGarik (Stearic

acid) EdlmanrUbmnþdUcxageRkam ³ CH3—CH2—CH2—CH2—CH2—CH2—CH2—CH2—CH2—CH2—CH2—CH2—CH2—CH2—CH2—CH2—CH2—COOH

cUrkt;sMKal;fa GatUmkae)annimYy²enAkñúgExSRtUv)ancgP¢ab;KñaeTA nwgcMnYnGtib,rmaénGatUm GIuRdUEsn. edIm,ICYy[caMnUvrcnasm<½n§rbs;GasIutRkaEq¥t GñkKYrEtKitGMBIGatUmkae)annimYy²enAkñúg ExSfava Eq¥t eTAedayGatUmGIuRdUEsn . ]TahrN_énGasIutRkaEq¥tRtUv)aneKpþl;[enAkñúgtaragTI 18-1 . ]TahrN_én GasIutRkaminEq¥t Edlman 18 GatUmkae)an KWCaGasIuGUelGiuk (Oleic acid) EdlmanrUbmnþdUcxageRkam ³

CH3—CH2—CH2—CH2—CH2—CH2—CH2—CH2—CH2═CH2—CH2—CH2—CH2—CH2—CH2—CH2—CH2—COOH

enAkñúgkrNIenH GasIutRkaminEq¥t manya:gehacNas;mansm<½n§BirCan;mYy EdltP¢ab;BI GatUm kae)anmYyeTAGatUmkae)anmYyepSgeTot . GaRs½yedaysm<½n§BIrCan; GatUmkae)anEdlCab;Tak;Tg enAkñúgsm<½n§enHKW minEq¥t eTAedayGatUmGIuRdUEsn . sm<½n§minEq¥tRtUv)aneKeXIjmanenAkñúg esÞIrEt RKb;GasIutRkaminEq¥tEdlekIteLIgenAkñúgFm μCatieRkamTMrg;Ca cis configuration . müa:gvijeTot sm<½n§BIrCan;minEmnsßitenAedayécdnüenAkñúgExSGIuRdUkabYreT . TaMgkartaMgenA nig TMrg;FrNImaRt saRsþ (the placement and the geometric configuration) rbs;sm<½n§BIrCan;RtUv)aneFIVtamGg;sIum EdlsMrYlCIvsMeyaKénGasIutRkaminEq¥tenaH . ]TahrN_énGasIutRkaminEq¥tEdleKCYbjwkjab; k¾RtUv)anbgðajenAkñúgtaragTI 18-1 Edr .



taragTI 18-1

GasIutRkaEq¥t nig minEq¥tEdleKCYbjwkjab; Common Saturated Fatty Acids

Common Name

I.U.P.A.C. Name

Melting Point (oC)

RCOOH Formula

Condensed Formula

Carpric Decanoic 32 C9H19COOH CH3(CH2)8COOH Lauric Dodecanoic 44 C11H23COOH CH3(CH2)10COOH Myristic Tetradecanoic 54 C13H27COOH CH3(CH2)12COOH Palmitic Hexadecanoic 63 C15H31COOH CH3(CH2)14COOH Stearic Octadecanoic 70 C17H35COOH CH3(CH2)16COOH Arachidic Eicosanoic 77 C19H39COOH CH3(CH2)18COOH

Common Unsaturated Fatty Acids

Common Name

I.U.P.A.C. Name

Melting Point (oC)

RCOOH Formula

Number of Double

bonds

Position of Double bonds

Palmitoleic Cis-9-Hexadecanoic 0 C15H29COOH 1 9 Oleic Cis-9-Octadecanoic 16 C17H33COOH 1 9 Linoleic Cis-9,12-Octadecadienoic 5 C17H31COOH 2 9,12 Linolenic All cis-9,12,15-Octadecatrienoic -11 C17H29COOH 3 9,12,15 Arachidonic All cis-5,8,11,14-Eicosatetranoic -50 C19H31COOH 4 5, 8, 11, 14

karBinitüemIltaragTI 18-1 nig rUbTI 18-2 bgðajBIcMNucsMxan;GMBIlkçN³rUbrbs; GasIutRka .

cMNucrlayrbs;GasIutRkaEq¥t ekIneLIgCamYynwgkarekIneLIg éncMnYnGatUmkae)andUc enAkrNI Gal;kan (Alkanes) . GasIutRkaEdlmanGatUmkae)an 10 b¤ elIsBIenHKWCaGgÁFaturwgenA sItuNðPaB Fmμta (Room temperature) .

cMNucrlayrbs;GasIutRkaEq¥t FMCagcMNucrlayrbs;GasIutRkaminEq¥t EdlmancMnYn GatUmkae)ankñúgExSes μ IKña . ehtuplEdlfa ExSGasIutRkaEq¥tman ninñakareTArkkar cgKñaya:gCab; enA kñúgrcnasm<½n§eTogTat; EdlCamUlehtunaM[mankarTajcUlrvagmU:elKulekIneLIg . karbgðajnUv sm<½n§BIrCan; cis enAkñúgExSGIuRdUkabYr begáItnUvmMu (bend) 30o . m:UelKulrmYlEbbenHminGactMerobKña [maM nig dUecñHmankarTajcUlénm:UelKulexSay nig cMNucrlayTab .

dUckñúgkrNIGasIutRkaEq¥tcMNucrlarbs;GasIutRkaminEq¥tekIneLIgCamYy nwgRbEvgExS GIuRdUkabYr.



rUbTI 18-2 cMNucrlayrbs;GasIutRka

karBinitüemIlPaBdUcKña nig xusKñarvagGasIutRkaEq¥t nig minEq¥t

lkçN³ GasIutRkaEq¥t GasIutRkaminEq¥tsmasPaBKImI kae)an GIuRdUEsn GuksIuEsn kae)an GIuRdUEsn GuksIuEsn Rcnasm<½n§KimI ExSGIuRdUkabYrEdlman COOH

enAcugExS ExSGIuRdUkabYrEdlman COOH

enAcugExS sm<½n§rvag C-C enAkñúgExS GIuRdUkabYr manEtsm<½n§mYyCan; ya:gehacmansm<½n§BirCan;mYyExSGIuRdUkabYrKWCalkçN³rbs;Rkum GiuRdUkabYr

Gal;kan Gal;Esn

TMrg;énExSGIuRdUkabYr ExSlatRtg; CaExSekagenAnwgkEnøgmansm<½n§BIr Can; .

sPaBrUbsaRsþenAkñúgsItuNðPaB Fmμta

GgÁFaturwg GgÁFaturav

cMNucrlaycMeBaHGasIutRka EdlmancMnYnGatUmkae)andUcKña

x<s; Tab

Tmnak;TmngrvagcMNucrlay nig RbEvgExS

ExSkan;EtEvgcMNucrlay kan;Etx<s;

ExSkan;EtEvgcMNucrlay kan;Etx<s;

oC 90

80

70

60

50

40

30

20

10

0

-10

-20

4 6 8 10 12 14 16 18 20 22 24 Carbon atom

Room temperature

Oleic acid, 1 double bons

Linoleic acid, 2 double bonds Linolenic acid, 3 double bonds

Saturated fatty acid



x- RbtikmμKimIénGasIutRka RbtikmμénGasIutRkadUceTAnwgGasIutkabuksIulikEdlmanExSxøI . RbtikmμsMxan;²EdlBYkva TTYlrgrYm maneGEsÞrkm μ (esterification) nigGIuRdUlIsGasIuténeGEsÞr saponification nig karbEnßmeTAelI sm<½n§BIrCan; .

RbtikmμeGEsÞrkm μ (Esterification)

enAkñúg eGEsÞrkmμ GasIutRkamanRbtikmμCamYyGal;kuledIm,IbegáItCa eGEsÞr nig Twk RsbeTA nwgRbtikmμTueTAdUcxageRkam ³

H+, Heat

R1—CO—OH + HO—R2 → R1—CO—O—R2 + H2O

RbtikmμGiuRdUlIsGasIut (Acid hydrolysis)

rMlwkfa GIuRdUlIsKWCaRbtikm μRtLb;énRbtikmμeGEsÞrkmμ edaybegáItCaGasIutRka BIeGEsÞr ³ H+, Heat

R1—CO—O—R2 + H2O → R1—CO—OH + R2—OH

Rbtikmμ Saponification

Saponification KWCaGIuRdUlIsEdlsMrYleday)aséneGEsÞr ³ R1—CO—O—R2 + NaOH → R1—CO—O−Na+ + R2—OH

plitplénRbtikmμenHKWCaGMbilEdlGacEbkCaGIuy:ug (Ionized salt) KWCasab‘U . GaRs½y eday sab‘UmanknÞúyEdlKμanbnÞúkGKÁisnI nig cugExSEdlmanbnÞúGKÁisnIGviC¢man ¬bgÁúMkabuksIulik¦ . vabegáItCabNþúMCab;Kña (Micelles) EdlGacrMlayeRbg nig PaKl¥itEdlRbLak; . ehtudUecñH FUlIRtUv )anlay (emulsify) EbkeTACaPaKl¥ittUc² nig GaclagsMGatecj . bBaðaGacekItmaneLIgenAeBlTwkrwg (hard water) RtUv)aneKeRbIsMrab;lagsMGat BIeRBaH kMhab;x<s;én Ca2+ nig Mg2+ enAkñúgTwkrwgenaH bgá[GMbilénGasIutRkaeLIgCakkr . minRtwmEtkar eRCotERCkCamYynwgGMeBIeFIV[rlayrbs;sab‘UeT b:uEnþ vak¾Tuk[BBuHFUlIrwgenAelIépÞFug b¤ PaCn_ Edldak;sMPar³e)akKk; .

2 R—CO—O− + Ca2+ → (R—CO—O−)2Ca2+ (s)



RbtikmμenAnwgsm<½n§BICan; ¬GasIutRkaminEq¥t¦ RbtikmμbEnßmGIuRdUEsn (Hydrogenation) KWCa]TahrN_énRbtikmμbEnßm . xageRkamenHKWCa ]TahrN_KmrUénkarbEnßmGIuRdUEsnsm<½n§BirCan;énGasIutRka ³

2H2, Ni

CH3(CH2)4CH═CHCH2CH═CH(CH2)7COOH → CH3(CH2)16COOH Linoleic acid Searic acid

RbtikmμbEnßmGIuRdUEsnRtUv)aneKeRbIenAkñúg]sSahkmμcMNIGahar edIm,IbMElgeRbgrukçCatiEdl manGasIutRkaminEq¥t (Polyunsaturated vegetable oils) [eTACaxøaj;rwgEdlCaGasIutRkaEq¥t . kar bEnßmGIuRdUEsnedayEpñk RtUv)aneKGnuvtþedIm,IbEnßmeTAelIsm<½n§BIrCan;mYycMnYn minEmnTaMgGs;eT enAkñúgeRbgEdlmanGasIutRkaminEq¥t . enAkñúgviFIenH eRbgrukçCatiravRtUv)anbMElgeTACaTMrg;rwg . Crisco KWCa]TahrN_éneRbgrukçCatiEdleKbEnßmGIuRdUEsn .

Margarine k¾RtUv)anplitedaykarbEnßmGIuRdUnEsnedayEpñkéneRbgrukçCati dUcCaeRbgeBat b¤ eRbgsENþkesog . TMhMénkarbEnßmGIuRdUEsnRtUv)anRtYtBinitüedayRbugRby½tñ dUecñHxøaj;rwgnwgGac ray)ay nig mansPaB (consistency) dUc b½r enAeBleKbriePaK . RbsinebImansm<½n§BIrCan;eRcIn eBkRtUv)anbEnßmGIuRdUEsn plitplEdlekIteLIgnwgmansPaBdUcxøaj;stVEdleKmincg)an . B’Nsib,nimμ itRtUv)aneKbEnßmeTAelIplitpl nig GacRtUv)anlayCamYynwgTwkedaHedIm,IbegáItCa TidæPaBdUcb½r nig mankøin (butterlike and flavor) .

K- Eicosanoids : Prostaglandins, Leukotrienes nig Thromboxanes

GasIutRkaminEq¥tmYycMnYnEdlmansm<½n§BIrCan;eRcInCagmYy minGacRtUv)ansMeyaKeday sBa’gÁkay . Gs;ry³eBlCaeRcInqñaM eKdwgfa Linolenic acid nig Linoleic acid RtUv)aneKehAfaCa GasIuRkacaM)ac; (essential fatty acids) caM)ac;sMrab;muxgarKImICIvsaRsþ (Biochemical functions) nig RtUvEtpþl;enAkñúgGahar ¬emIltaragTI 18-1¦ . muxgarrbs; Linoleic acid eKsÁal;ya:gc,as;enAkñúg kMLúgTsvtSqñaM 1960 enAeBlvaRtUv)aneKrkeXIjfa Linileic acid RtUvkarcaM)ac;sMrab;CIvsMeyaKén Arachidonic acid EdlCa Precursor énsarFatumYyfñak;dUcG½rm:Un RtUv)aneKsÁal;faCa Eicosanoids. eQ μaHenHmanRsLayBIBaküRkicfa “eikos” mann½yfa “20” BIeRBaHBYkvaTaMgGs; manRsLayBIGasIutRkaEdlman 20 GatUmkae)an . Eicosanoids rYmmanbIRkuménsmasFatuEdl rcnasm<½n§Cab;Tak;Tg BYkvaKW ³ Prostaglandins, Leukotrienes nig Thromboxanes .



Prostaglandins KWCam:UelKulEdlmanGnuPaBCIvsaRsþxøaMgbMputedaymanskmμPaBdUcCa G½rm:Un . BYkvaTTYl)aneQ μaHfaCa Prostaglandins GaRs½yedaysar BYkvaRtUv)aneKyk ecjTwkkam (seminal fluid) EdlplitedayRkeBjrbs;RbUsþat (Prostate glande) b:uEnþfμ I²enH BYkva k¾RtUv)an eKykecjBICalikastVPaKeRcInbMput . Prostaglandins KWCaGasIutkabuksIulikminEq¥t

(Unsaturated carboxylic acids) EdleRKagrbs;vaman 20 GatUmkae)an ehIyenAkñúgeRKagman ExSbiTEdlman 5 GatUmkae)an .

mancMNat;fñak;TUeTArbs; prostaglandins CaeRcInRtUv)aneKdak;CaRkumenAeRkamkar[eQ μaH A,

B, E nig F kñúgcMeNamBYkvaepSgeTot . namvlIvén Prostaglandins sMGageTAelIkarerobcMéneRKag GatUmkae)an nig cMnYn nig karcg¥úlTis (orientation) énsm<½n§BIrCan; bgÁúMGIuRduksIul nig bgÁúMGaestUn (ketone group) . ]TahrN_ enAkñúgeQ μaH PGF2, PG eRbICMnYs Prostaglandin ehIy “F” bgðajBIRkum BiessmYyén Prostaglandins EdlmanbgÁúMGIuRduksIulenAelI C9 nig “2” bgðajfaman sm<½n§BIrCan;BIrrvagGatUmkae)anenAkñúgsmasFatu . ]TahrN_enAkñúgrUbTI 18-3 bgðajBIrcnasm<½n§ TUeTArbs; Prostaglandins nig RbB½n§namvlIbc©úb,nñ .

Prostaglandins RtUv)aneFIVenAkñúgCalikaPaKeRcIn nig man\T§iBlCIvsaRsþeTAelIekasikaEdl plitBYkva nig eTAelIekasikaepSgeTotenAkñúgtMbn;Citxag . cenøaHminFmμtaénmuxgarrbs; Prostaglandins rYmman ³

kareP¢acénsac;dMurelag (smooth muscle) karEktMrUvénCIvsMeyaKrbs;esþr:UGIud karraraMgénkarbeBa©jrsRkBH karraMgénGg;sIum Lipase EdlrYsCamYynwgG½rm:Un karraMgénkarCab;Kñarbs;)aøEkt kareP¢acénkarCab;Kñarbs;)aøEkt karEktMrUvénkarbBa¢ÚnB½t’mantamsésRbsaT karTTYlGarmμN_eTAnwgkarQWcab; nig karsMrYlénkareqøIytbeTAnwgkarrlak

GaRs½yeday Prostaglandins nig sarFatuCab;Tak;TgCitsñiT§ Leukotrienes nig Thromboxanes man\T§iBleTAelIdMeNIrkarénsBa’gÁkayya:geRcIn nig Cajwkjab; GaRs½yeday



BYkvabgá[man \T§iBlRbQmKñaenAkñúgCalikaepSg² vamankarlM)akkñúgkarrkSaKMrUénmuxgarEktMrUv CaeRcIn . xag eRkamenHKWCakarsegçbéndMeNIrkarCIvsaRsþmYycMnYnEdleKKitfaRtUv)an EktMrUveday

Prostaglandins, leukotrienes nig Thromboxanes .

1- karkkQam (Coagulation du sang) karkkQamekIteLIgenAeBlsésQamRtUv)anTTYlrgrbYs b:uEnþkarkkQamrebobenHenA tambeNþayénépÞxagkñúgsésQamGacbNþal[manCMgWKaMgebHdUg (Heart attack or stroke) . Thromboxane A2 ¬rUbTI 18-4¦ RtUv)anpliteday)aøEktenAkñúgQam nig CMruj[mankarrYménsés Qam nig karkþaMgCab;Kñaén)aøEkt .

rUbTI 18-3 rcnasm<½n§rbs; Prostaglandines TaMgbYn

COOH

OH OH

COOH

OH OH

OH

Prostaglandine E1

Prostaglandine F1

COOH

OH OH

COOH

OH OH

OH

Prostaglandine E2

Prostaglandine F2



rUbTI 18-4 rcnasm<½n§rbs; Thromboxane A2 nig Leucotriene B4

pÞúymkvij PGI2 (Prostacycline) RtUv)anplitedayekasikaEdlRkalépÞsésQam nig

man\T§iBl RbQmnwg Thromboxane A2 . Prostacycline raraMgkarkkkþaMg (aggregation)

rbs;)aøEkt nig bNþal[mankarrIknUvsésQam ehtudUecñHvakarBarmin[mankarbegáItnUvkMNkQam (Caillot) .

2- kareqøIytbeTAnwgPaBrlak (Réponse inflammatoire)

kareqøIytbeTAnwgPaBrlakKWCaynþkarkarBarrbs;sBa’gÁkaymYyepSgeTot . enAeBlCalika TTYlrgkarrbYsedaykarb:HTgÁic karrlak (inflammation) b¤ karQøanBanedayemeraK (Invasion

par microorganisms) RbePTepSg²énRKab;Qams (Globules blancs) cuHeTAkEnøgmankarxUcxat edIm,IBüayamedIm,IeFIV[fycuHnUvkarbMpøajCalika . lT§plénkareqøIytbenH KW karehIm eLIg Rkhm RKunekþA nig QW . eKKitfa Prostaglandines begáInnUvTidæPaBmYycMnYnénkareqøIytb eTAnwgPaBrlak CaBiess QW nig RKunekþA . »sfmYYycMnYndUcCa Aspirine raraMgkarsMeyaK Prostaglandines nig CYybnßynUveraKsBaØa (symptomes) . eyIgnwgGacBinitüemIl BIynþkarénGMeBI rbs;»sfTaMgenHenAeBleRkaykñúgEpñkenH .

3- RbB½n§bnþBUC (Système reproductive) PGE2 eP¶ackarkRnþak;énsac;dMurlIj (muscle lisse) CaBiesskarkRnþak;éns,Ún (Utérus) .

karekIneLIgénkMrit Prostaglandines RtUv)aneKkt;sMKal;eXIjmuneBlPøam²nUvkarcab;epþIménkar qøg Tenø . PGE2 k¾RtUv)aneKeRbIedIm,IeFIVkarrMlUtenARtImasTI 2 (second trimester abortion) . vaman

karbgðajya:gc,as;fa Dysmenorrhée ¬karmkrdUvedaymankarQWcab;¦ RtUv)anTTYlrgedayRsþI

COOH

OH

COOH

OHOH

Thromboxane A2

Leucotriene B4



CaeRcInGacCalT§plénkarekIneLIgelIsén Prostaglandines TaMgBIr . BitR)akdehIy »sf dUcCa Ibuprofene EdlraraMgkarsMeyaK Prostaglandine Edl)anÉkPaBeday FDA nig RtUv)aneKeXIjfa pþl;ya:gc,as;nUvkarbnßykarQWcab;BIeraKsBaØaTaMgenH .

4- pøÚvGahar (Voie digestive : Gastrointestinale)

Prostaglandines )anbgðajTaMgraraMgkarbeBa©jGasIut nig begáInkarbeBa©jRsTab; mucus

EdlkarBarenAkñúgRkBH . enAkñúgviFIenH Prostaglandines CYykarBarRkBH . cUrBicarNacMeBaH ry³eBlmYy plvi)akbnÞab;bnSMGacekItmaneLIgedaysarBnüarkareRbInUv»sfdUcCa Aspirine Gac man\T§iBleTAelIRkBH ³ dMe)ARkBH (ulcère gastrique). GasRs½yeday Aspirine vaeTAraraMgkar sMeyaK Prostaglandines CaFm μtavaGac[CMru[man dMe)ARkBH (ulcère gatrique) edaykarraraMgmin [mankarbegáItnUvRsTab; mucus karBarFm μta nig kñúgxN³CamYyKñaenHvanaM[mankarekIneLIgnUvkar beBa©jGasIutrbs;RkBH .

5- tMrgenam (Rein)

Prostaglandines RtUv)anbegáItenAkñúgtMrgenambgá[sésQaméntMrgenamrIk . karhUrcUl ya:geRcInénQamtamry³tMrgenambNþal[mankarbeBa©jTwk nig eGLikRtUlItekIneLIg .

6- pøÚvdegðIm (Voie respiratoire)

Eicosanoides RtUv)anplitedayekasikaRKab;QamsmYycMnYn Leucotrienes ¬emIlrUbTI 18-4¦ begáInnUvkarrYmbMBg;xül; (Bronches) EdlnaM[man hWt (Asthme) . Prostaglandines epSgeTotnaM[mankarrIkbMBg;xül; .

dUcEdlkarsikSaedaysegçbpþl;Caeyabl; Prostaglandines Cajwkjab;man\T§iBl anta-

goniste CaeRcIn . eTaHbICaBYkvaminsmRsbeTAnwg niymn½yrbs;G½rm:Un k¾eday ¬sarFatuEdl plitenAkñúgCalikaBiess nig RtUvdwknaMedaycrnþQameTAdl;CalikacMNuc (tissue cible) enAkEnøg NamYyenAkñúgdgxøÜn¦ k¾b:uEnþ Prostaglandines KWCaGñkEktMrUv (Régulateur) ya:gc,as;las;eday man\T§iBleTAq¶ay . dUcEdl)anKUsbBa¢ak; Prostaglandines eP¶ackareqøIytbPaBrlak nig CalT§pl TTYlxus RtUvCaBiesscMeBaHkarekIteLIgCabnþbnÞab;nUvvibtþiEdlbgá[manPaBQW . Aspirine RtUv)aneKsÁal; taMgyUrNas;mkehIykñúgkareFIVfycuHnUvPaBQWrebobenH nig \LÚveyIg)andwgfa vaeFIVdUecñHGaRs½y edaykarraraMgkarsMeyaK Prostaglandines .



dMNak;karBIrdMbUgénkarsMeyaK prostaglandines ¬rUbTI 18-5¦ karbeBa©jnUv acide

arachidonique BIPñasekasika nig karbMElgrbs;vaeTACa PGH2 edayGg;sIum cyclooxygenase ekIteLIgenARKb;CalikaEdlGacbegáIt Prostaglandines . karbMElg PGH2 eTACaTMrg;CIvsaRsþ skmμepSgeTotKWCalkçN³Biessrbs;Calika nig RtUvkarGg;sIumsmRsb EdleKeXIjmanenAkñúgEt CalikamYycMnYn . Aspirine RbRBwtþieTAedaykarraraMgnUvGg;sIum Cyclooxygenase EdlsMrYlRbtikmμ dMNak;kardMbUgénpøÚvEdlnaM[mankarbMElgBI Acide arachidonique eTACa PGH2 . bgÁúMGaesTIl (acetyl) rbs; Aspirine cgtamsm<½n§kUva:Lg;eTAnwgGg;sIum EdlenATIenaHGg;sIumkøayeTACaGkmμ (inactive) ¬rUbTI 18-6¦ . GaRs½yedayRbtikm μRtUv)ansMrYledayGg;sIum Cyclooxygenase

ekIteLIgenAkñúgRKb; ekasika Aspirine raraMgedaymanRbsiT§PaBnUvkarsMeyaK Prostaglandines RKb;RbePT .

rUbTI 18-5 segçbénkarsMeyaK Prostaglandines CaeRcInBI Acide arachidinique

COOH

Acide arachidonique

PGH2Leucotrienes

PGF2α

1. Relaxation de muscle lisse 2. Vasodilatation

TXA2 PGI2 1. Inflammation 2.Broncho-constriction; Vasoconstriction; perméabilité capillaire

1. Anti-plaquette aggregation 2. Vasodilatation

PGE2

1. Plaquette aggregation 2. Vasoconstriction

1. Contraction de muscle lisse 2. Vasconstriction



rUbTI 18-5 Aspirine raraMgkarsMeyaKrbs; Prostaglandines

O ║ O—C—CH3

COO−

Aspirine (Acide acétyl salicylique)

+Enzyme NH2

Enzyme

O ║ N—C—CH3 І H

+

OH

COO−

Enzyme inactive

Salicylate



3-KøIesrId (GLYCÉRIDES)

k- KøIesrIdNWt (Glycérides neutres)

Glycérides KWCaeGEsÞrénlIBIdEdlmanm:UelKulKøIesr:Ul (Glycérol) nig GasIutRka . BYkva GacRtUv)anEckBIrRkumCabnþeTot ³ KøIesrIdNWt nig hVÚsVÚKøIesrId (glycéride neuter et Phosphoglycérides) . Glycérides neuters minEbkCaGIuy:ug nig minmanb:Ul . Phosphoglycérides mantMbn;manb:Ul ¬bgÁúM Phosphoryle¦ edaybEnßmnUvknÞúyCaGasIutRkaK μanb:Ul. rcnasm<½n§rbs;va nimYy²mansar³sMxan;cMeBaHmuxgarrbs;va .

eGEsÞrkmμén glycérol CamYynwgGasIutRkabegáItnUv glycérides neutres . eGEsÞrkm μGaceFIV enAelITItaMg mYy BIr b¤ TaMgbIedaybegáIt)anCa Monoglycérides, diglycérides b¤ triglycérides. Gñkk¾GacehAvapgEdrfaCa mono, di b¤ triacylglycérol . eTaHbICa monoglycérides nig diglycérides manenAkñúgFm μCatik¾eday k¾b:uEnþ glycérides

EdlsMxan;CageKbMputKW triglycérides EdlCasmasFatusMxan;énekasikaxøaj; (adipocytes) . Triglycéride manExSsMxan;Ca glycérol nigcgP¢ab;eTAedayGasIutRkabIExStamry³sm<½n§eGEsÞr (ester). karbegáIt Triglycéride RtUv)anbgðajenAkñúgRbtikmμxageRkam ³

Triglycérides (Triacylglycérol) RtUv)aneK[eQμaHeTAtamExSsMxan;EdlmaneQμaHfaCa Glucérol Cabc©wmbT . eQ μaHénbgÁúMGasIutRka (Acyl) RtUv)aneKdak;enAxagmuxva . bgÁúMGasIutRka Acyl RtUv)aneK[eQ μaHedaykat; Bakü “ique” nig CMnYsvaeday oyl enAkñúg]TahrN_xageRkam eQ μaH Triglycéride enAxageqVg KWCa tri-stearoylglycérol . buBVbT tri R)ab;eyIgfamanbgÁúM acyl bIEdlCa stéaroyl P¢ab;eTAnwg glycérol . Triglycéride enAxagsþaMKWCa Triglycéride cMruH edayehtufavaman bgÁúM acyl bIxus²KñaEdlP¢ab;eTAnwg glycérol .

CH2—OH І CH—OH + 3 R—CO—OH І CH2—OH Glycérol Acide gras

CH2—O—CO—R І CH—O—CO—R + 3 H2O І CH2—O—CO—R Triglycéride eau

CH2—O—CO—(CH2)16CH3 І CH—O—CO—(CH2)16CH3 І CH2—O—CO—(CH2)16CH3

Tristearoylglycérol

CH2—O—CO—(CH2)14CH3 І CH—O—CO—(CH2)7CH═CH(CH2)7CH3 І CH2—O—CO—(CH2)16CH3 1-Palmitoyl-2-oleoyl-3-stearoylglycerol



GaRs½yBMumanbgÞúkGKÁisnI viC¢man b¤ GviC¢manenAelIm:UelKulenH BYkvaRtUv)aneKehAfaCa Glycérides neutres . m:UelKulEvg²TaMgenHdMerobKñaBImYyeTAmYyepSgeTotbgá[)anPaKeRcInén lIBIdEdlRtUv)ansþúkTukenAkñúgekasikaxøaj;énsBa’gÁkay .muxgarsMxan;rbs; Triglycérides enAkñúg RbB½n§CIvKImIKWCakarsþúkTukénfamBl . GaharsMbUr famBleRcInRtUv)aneKbriePaKelIsBItMrUvkar sMrab;dMeNIrkaremtabUlIs PaKeRcInénfamBlEdlelIs enHRtUv)anbMElgCa glycérides neutres nig RtUv)ansþúkTukCa Triglycérides enAkñúgCalikaxøaj; (Tissue adipeux) . enAeBlRtUvkarfamBl Triglycérides RtUv)anbMEbkedaysarsBa’gÁkay ehIy famBlRtUv)anbeBa©j . cMeBaHehtuplenH karhat;R)aN rYmCamYykarkat;bnßylμmnUvGaharsMbUr kaLÚrIRtUv)aneKpþl;[eRbIcMeBaHbuKÁlEdl elIsTMgn; . kahat;R)aN ¬dMeNIrkarEdlRtUvkarfamBl¦ begáInnUvGRtaemtabUlIsrbs;xøaj; nig bNþal[cuHTMgn; . x- Phosphoglycérides

Phospholipides KWCaRkummYyrbs;lIBIdEdlman ester phosphate . vtþmanrbs;bgÁúM Phosphoryle bNþal[m:UelKulmank,alman b:Ul ¬bgÁúM phosphoryle¦ nig knÞúy minmanb:Ul ¬ExS Alkyle rbs;GasIutRka¦ . GaRs½yedaybgÁúM phosphoryle EbkCaGIuy:ugenAkñúgsUluysüúg CalT§plKW lIBImanbnÞúkGKÁisnI . lIBIdénPñasekasikaEdlsMbUrCagbMputmanRsLayBI Glycérol-3-phosphate nig RtUv)aneK sÁal;faCa Phosphoglycérides . Phosphoglycérides manbgÁúM Acyle EdlmanRsLayBIGasIutRka EdlmanExSEvgenAnwg C1 nig C2 én Glycérol-3-Phosphate . enAnwg C3 bgÁúM Phosphoryle

RtUv)an P¢ab;eTAnwg Glycérol edaysarsm<½n§ Phosphoesters . Phosphoglycérides

samBaØCageKbMput man bgÁúM Phosphoryle esrI nig RtUv)aneKsÁal;faCa Phosphatidate ¬rUbTI 18-7¦ . enAeBlbgÁúM Phosphoryle RtUv)anP¢ab;eTAnwgm:UelKulrlaykúñgTwk (molecules hydrophylique)

Phosphoglycérides complexes RtUv)anbegáIteLIg . ]TahrN_ Phosphatidyl choline (lechithine) nig Phosphatidylethanolamine (Cephaline) RtUv)aneKeXIjmanenAkñúgPñasekasika PaKeRcInbMput ¬rUbTI 18-7¦. Lecithine man k,al manb:Ul nig knÞúyminmanb:Ul . ehtudUecñHvaKWCam:UelKul Amphotère

(Ampholyte) . rcnasm<½n§enHvadUceTAnwgrcnasm<½n§rbs;sab‘U nig m:UelKul detergent Edl)anBiPakSa BIxagelIrYcmkehIy . k,al EdlCaGIuy:ugrlayenAkñúgTwk nig manRbtikmμCamYymU:elKulTwk rIÉ knÞúy Edlminmanb:Ul minrlaykñúgTwk nigmanRbtikmμCamYym:UelKulminmanb:Ul . FmμCati



Amphotère sßitenAkNþalénrcnasm<½n§nigmuxgarrbs;Pñasekasika . eRkABICasmasPaBenAkñúgPñas ekasika lecithine KWCa Phospholipides enAkñúgsarFatuEdl skm μenAelIépÞénsYt (Surfactant pulmonaire) . vak¾RtUv)aneKeXIjmanenAkñúgEpñkRkhméns‘ut nig sENþkesog nig RtUv)aneKeRbICa Agent émulsionnant enAkñúgkaer:m . Agent émulsionnant CYyenAkñúgkareFIVCa Suspension én Triglycérides enAkñúgTwk . Lecithine EdlCa Amphotère eRbICas<ancgP¢ab;Kñarvag m:UelKulTwkEdlmanb:Ulx<s; nig Triglycérides Edlminmanb:Ul . kareFIV EmulsionekIteLIg BIeRBaH k,alEdlGacrlaykñúgTwkrbs; lecithine rlayenAkñúgTwk nig knÞúyEdlminrlay enAkñúgTwk rlayenAkñúg Triglycérides . Céphaline CaTUeTAmanrcnasm<½n§dUceTAnwg lécithine vaxusKñaEtbgÁúM Amine RtUv)ancgP¢ab; eTAnwg bgÁúM Phosphoryle .

rUbTI 18-7

rcnasm<½n§rbs; Phosphoglycérides

O ІІ −O—P—O І O−

CH2—O—CO І CH—O—CO І CH2

NH3+ O

І ІІ H—C—CH2—O—P—O І І COO− O−


Phosphatidate

O ІІ (CH3)3N

+—CH2—CH2—O—P—O І O−


Phosphatidylcholine (lecithine)

O ІІ H3N

+—CH2—CH2—O—P—O І O−


Phosphatidylethanolamine (Céphaline)

Phosphatidylserine



4- lIBIdminEmnKøIesrId (LIPIDES NON-GLYCÉRIDES) k- Sphingolipides

Sphingolipides KWCalIBIdminmanRsLayBI glycérol eT . dUcCa Phospholipides, Sphingolipides KWCasarFatu amphotère EdlmanbgÁúMk,almanb:Ul nig manknÞúyBIrCa GasIutRka nig CasmasFatuénrcnasm<½n§rbs;Pñasekasika . BYkvamanRsLayBI Sphingosines EdlCaGal;kul manExSEvgnigmanGasUtenAkñúgExS .

Sphingolipides rYmman Sphingomyéline nig Glycosphingolipide . Sphingomyéline KWCa cMNat;fñak;EtmYyén Sphingolipides EdlGacCa Phospholipides Edr .

Sphingimyéline sßitenABaseBjsBa’gÁkayb:uEnþvaCasmasFatusMxan;CaBiessénrcnasm<½n§ rbs;PñasekasikasésRbsaT . BYkvaRtUv)aneKeXIjsMbUrenAkñúg Myéline EdlB½T§CMuvij nig karBar ekasikaRbB½n§sésRbsaTkNþal (Central nervous system) . cMeBaHmnusS REhl 25 % énlIBId rbs; Myéline KWCa Sphingomyélines . tYnaTIrbs;BYkvasMxan;cMeBaH muxRtwmRtUvénxYrk,al (function

cerebral proper) nig Transmission nerveux . Glycosphingolipides b¤ Glycolipides rYmman Cérébrosides, Sulfatides nig Gangliosides nig RtUv)anbegáIteLIgedaysMGagelIrcnasm<½n§rbs; Céramide EdlKWCa GasIutRkaGamId (Acides gras amides) EdlmanRsLayBI Sphingosine .

OH І CH3(CH2)12CH═CH—C—H І H2N—C—H І CH2OH Sphingosine

OH І CH3(CH2)12CH═CH—C—H І R—CO—HN—CH O І ІІ CH2—O—P—O—CH2CH2N

+(CH3)3 І O−

Sphingomyéline

Acide gras

Sphingosine Groupement phosphoryle

Choline



Cérébroside RtUv)anbBa¢ak;edayvtþmanénbgÁúMk,alCam:UNUsak;karIdEtmYyKt; . Cérébro-

sides BIrEdleKCYbjwkjab; Glucocérébroside EdleKeXIjmanenAkñúgPñasrbs; Macrophages

¬ekasikakarBardgxøÜnedaykarrMlay nig bMpøajemeraKmkBIxageRkA¦ nig Galactocérébroside eK eXIjmanCaBiessesÞIrEtenAkñúgPñasekasikaxYrk,al . Glucocérébroside man céramide RtUv)ancg P¢ab;eTAnwg Glucose ehIy Galactocérébroside man céramide cgP¢ab;eTAnwg Galactose .

Sulfatides manRsLayBI Galactocérébroside EdlmanbgÁúM Sulfate . cUrbBa¢ak; BYkvaman bnÞúkGKÁisnIGviC¢manenAkñúg pH Fmμta (pH physiologique) .

OH І CH3(CH2)12CH═CH—C—H І H2N—C—H І І O═C CH2OH І (CH2)n І CH3 Céramide

CH3 І (CH2)n Glucocérébroside І CO І NH OH І І O—CH2—C—C—CH═CH—(CH2)12—CH3 І H HO

CH2OH

OH

OH

CH3 І (CH2)n Galactocérébroside І CO І NH OH І І O—CH2—C—C—CH═CH—(CH2)12—CH3 І H

HO

CH2OH

OH

OH



Gangliosides KWCa Glycolipides EdlmanbgÁúM oligosaccharides EdlrYmmanm:UelKulmYy b¤ eRcInén Acide N-acetlyneuraminque (Acide sialique) . CabzmRtUv)aneKykecjBIPñasénCalika sésRbsaT Gabgliosides RtUv)aneKeXIjmanenAkñúgCalikaPaKeRcInénsBa’gÁkay .

CH3 І Sulfatide de (CH2)n Galactocérébroside І CO І NH OH І І O—CH2—C—C—CH═CH—(CH2)12—CH3 І І H H

HO

CH2OH

OSO3

OH

HO

CH2OH

OH

OH

CH3 І (CH2)n Ganglioside І CO І NH OH І І O—CH2—C—C—CH═CH—(CH2)12—CH3 І І H H

CH2OH

OH

OH

O O

CH2OH

O

OH

COO− CH2CONH

HO

Glycerol

OH



x- Steroides

Stéroides ekIteLIgCaFm μtanUvGMbUrm:UelKulsrIragÁénkarcab;GarmμN_enAkñúg CIvKImI nig evC¢saRsþ. vaTb,divaT (controversy) CaeRcInmanenACMuvij BYk Stéroides epSg² . eyIgmankar)armÖ GMBIcMnYn Cholestérol enAkñúgrbbGahar nig \Ti§Blrbs;vaeTAelIsuxPaB . eyIgmankar)armÖGMBI kareRbI stéroides anaboliques edayBYkGtþBlkmμ (Athletes) edIm,IksagnUvma:s;sac;dMu nig EklMG nUvkarsMEdgrbs;BYkeK . b:uEnþsmaCikénGMbUrm:UelKUlenHmanRsLayBI Cholestérol manmuxgarsMxan; CaeRcInenAkñúgsBa’gÁkay . Sels biliares EdlCYykñúgkarlay[sBV (emulsion) nig rMlaylIBId (digestion) Cam:UelKul stéroides dUcKñaEdr Hormones sexuelles , Testosterone nig estrogene . Steroides KWCasmaCikénRbCMuénlIBIdd¾FMnigmaneRcInRbePT EdleKehAfaCa Isoprenoi-

des. RKb;smasFatuTaMgenHRtUv)anksageLIgBIÉktaEdlman 5 GatUmkae)an ¬EdleKehAvafa Isoprène ¦ mYy b¤eRcIn . Terpène KWCaBaküTUeTAcMeBaHlIBIdEdlRtUv)ansMeyaKBI Isoprène . ]TahrN_én Terpènes

rYmman Stéroide nig Sels biliaires vItamInrlayenAkñúgxøaj; (Vitamines liposolubles) Chlorophylle

nig G½rm:UnrukçCatimYycMnYn (Hormones végétaux) . RKb; Stéroides manéNVy:U stéroides dUcEdl)anbgðajxageRkam ³

eRKagkae)anénBYk stéroide manExSbiTbYnEdlrlaCab;Kña . ExSbiTmYyKU²manGatUmkae)anBIrrYmKña . ehtudUUecñHExSbiTBIrEdlrlayCab;Kñamansm<½n§mYy b¤ eRcInrYmKña EdlCaEpñkmYyéneRKagq¥wgrbs;va . ]TahrN_ExSbI A nig B , B nig C, nig C nig D EdlvaTaMgGs;Cab;KñaenAkñúgrcnasm<½n§bnþbnÞab; . Stéroides PaKeRcInmanbgÁúM Méthyle EdlP¢ab;eTAnwg C10 nig C13 k¾dUcCa bgÁúM Alkyle , Alcool b¤ k¾ Cétone enA C17 . Cholestérol Ca stéroide EdleKCYbjwkjab; eXIjmanenAkñúgPñasrbs;ekasikastV PaKeRcIn . vaKWCam:UelKul Amphotère ehIyvarlayenAkñúgtMbn;manTwkénPñasekasika . vaCab;Tak;

12 17 11 13 16 1 9 15 2 10 14 8 3 5 7 4 6

C

B A

D

Noyau stéroidique



TgenAkñúgkarEktMrUvénsnÞIy_PaB (fluidity) énPñasekasikaCalT§plénExSbiTEdlCab;Kñaminmanb:Ul . b:uEnþbgÁúM Hydroxyle manb:Ul nig manmuxgardUc k,alEdlmanb:Ulrbs; Sphingolipides nig Phospholipides . vamanTMnak;TMngya:gxøaMgrvagkMhab;rbs; Cholest;erol EdleXIjmanenAkñúgQam nig CMgWebHdUg CaBiess Arthérosclérose ¬karrwgén artère ¦ . Cholestérol enAeBlrYmCamYy sarFatuepSgeTot bNþal[rYmtUcnUvpøÚvxagkñúgsésQam . karrYmsésQamkan;EtekIneLIg sMBaF RtUvEtekIneLIgedIm,IeFIV[R)akdfakarhUrQam)anRKb;RKan; nig sMBaFQamx<s; (Hypertension) ekIt eLIg . Hypertesion k¾RtUv)anCab;Tak;TgeTAnwgCMgWebHdUgEdr . Cholestérol

EpñkRkhméns‘utmankMhab;x<s;én Cholestérol k¾dUcCaplitplEdleFIVBITwkedaH nig xøaj;stV. CalT§pl eKyl;RBmfacMnYnénplitplTaMgenHenAkñúgrbbGaharRtUv)aneKEktMrUv[ manl μm nUvGaharEdlman Cholestérol . Sels biliaires KWCaRsLay amphotère rbs; Cholestérol EdlRTUv)ansMeyaKenAkñúgeføIm nig sþúkTukenAkñúgRbma:t; . Sels biliaires sMxan;²enAkñúgmnusSKW Cholate nig Chenodeoxycholate. Cholate Chenodeoxycholate

Sels biliaires KWCa Agent émulsionnant EdlbgÁúM hydroxyle rbs;vamanRbtikmμCamYynwgTwk nig tMbn;minRCabTwkrbs;vacgP¢ab;eTAnwglIBId . bnÞab;BIkarbriePaKGahar TwkRbma:t;hUrBIfg;Rbma:t;

CH3 І CHCH2CH2COO−

CH3

CH3

HO

HO

OH

CH3 І CHCH2CH2COO−

CH3

CH3

HO

OH

CH3 І CHCH2CH2CH2CHCH3 І CH3

CH3

CH3

HO



cUleTAkñúg duodénum ¬tMbn;xagelIbMputéneBaHevontUc¦ . enATIenH Sels biliaires rMlayxøaj; Edl)anmkBIGahareTACadMNk;tUc²EdlGacRtUv)anbMEbkrYcCaeRsceday Lipases ¬Gg;sIumrMlay xøaj;¦ k¾GaceXIjmanenAkñúgeBaHevontUcEdr . Stéroides mantYnaTIenAkñúgvdþbnþBUC (cycle reproductif) . enAkñúges‘rIRbtikm μKImI Cholestérol RtUv)anbMElgeTACa Progestérone KWCaG½rm:UnsMxan;EdlrYmCamYynwgPaBmanK’P . edayplitenAnwgGUEvr (Ovaire) nig suk (Placenta) progestérone TTYlxusRtUvcMeBaHTaMgkarcab; edayeCaKC½y nig karbBa©b;edayeCaKC½yénPaBmanK’P . vaerobRsTab;éns,ÚnedIm,ITTYlyk oeuf

fertilisé . enAeBl oeuf RtUv)anP¢ab; Progestérone Cab;Tak;TgenAkñúgkarrIkcMerInrbs; foetus nig mantYnaTIenAkñúgkarkat;pþac;min[man Ovulation enAkñúgry³eBlénPaBmanK’P . Testotérone CaG½rm:Unburs (hormone male) eXIjmanenAkñúgBgsVas nig estrone KWCa G½rm:UnRsþI sarFatuTaMgBIrRtUv)anplitedaykarERbRbYlKImIén Progestérone . G½rm:UnTaMgenHCab; Tak;TgenAkñúgkarrIkcMerInénlkçN³ePTburs nig RsþI . Stéroides CaeRcIn rYmman Progestérone mantYnaTIsMxan;karrIkcMerInénPñak;garRtYtBinitü kMeNIt (Agents de control de la néssance) . 19-Norprogestérone KWCaPñak;garsMeyaKdMbUgEdl RtYtBintükMeNIt . vamanRbsiT§PaBRbEhl 10 dgxøaMgCag Progestérone kñúgkarpþl;karRtYtBinitü kMeNIt . b:uEnþkareRbIrbs;vaRtUv)aneKkMNt;ya:gxøaMg BIeRBaHsmasFatuenHminGaceRbItammat; nig GaceRbI)anEttamkarcak; . smasFatuEdlCaRsLayva Norlutin (17-α-ethyl-19-nortestostérone)

RtUv)aneXIjvapþl;nUvTaMg kMlaM nig RbsiT§PaBrbs; 19-norprogestérone nig GaceRbItammat; . bc©úb,nñenH karrYmbBa©ÚlKñaén»sfBnüarkMeNIteRbItammat; (Contraceptifs utilisés par voie

orale) RtUv)aneKeRbICajwkjab;bMput . BYkvaenHrYmman Progestérone nig estrogène . plitplfμI CageKTaMgenH pþl;[nUvkarkarBar)anl¥bMputénkareFIVkMu[mankUn (Contraceptive protection) Cag Pñak;garepSgeTotEdlRtUv)aneRbIEtÉg . BYkvak¾RtUv)aneKeRbIsMrab;EktMrUvnUvdMeNIrmanrdUv (Menstruation) cMeBaHGñkCMgWEdlmankarhUrQamrdUveRcIn . eday)aneFIVkarRsavRCavCaelIkdMbUg enAkñúgcugTsSvtqñaM 1950 (1950s) nig RtUvGnuBaØatieday FDA enAkñúgqñaM 1961 vamanya:gehac Nas; »sfrYmbBa©ÚlKñarebobenHEdlmanrhUtdl;bc©úb,nñ 30 mux . müa:gvijeTot TMrg;»sfEdlbiT elIEs,k (transdermal patch) sMrab;Büa)al Ostéoporoses post-ménopause RtUv)aneKkMBug RsavRCav . RKb;smasFatuTaMgenHmanGMeBIedaykarbgá[manK’Pedayxus (False pregnancy) EdlraraMg kM[man ovulation . enAeBlkareRbI»sfBnüarkMeNItminCab;lab; CaFmμta ovulation vaRtLb;mk



vijenAkñúgbIvdþrdUv (three menstrual cycle) . eTaHbICamanbBaðarYmCamYy »sfenHk¾eday vahak;dUc CaenAEtCaviFImanRbsiT§PaB nig suvtßiPaBénEpnkarRKYsarsMrab;RbCaCnCaeRcIn .

Progetérone Testostérone Estrogène 19-Norprogestérone Norlutin Aldostérone Cortisone

Cortisone k¾mansarsMxan;kñúgkarEktMrUvéndMeNIkarCIvKImICaeRcIn . ]TahrN_ vaCab;Tak;Tg enAkñúgemtabUlIsrbs;KøúsIut . Cortisone k¾RtUv)aneKeRbIsMrab;karBüa)alCMgWrlak;snøak; (Artrite

rhumatoides) hWt (asthme) , disorders gastrointestinals CMgWEs,kCaeRcIn (dermatoses) nig CMgWepSg² . b:uEnþkarBüa)aleday Cortisone minGt;maneRKaHfñak;eT . karBüa)almYycMnYnGacman \T§iBlbnÞab;bnSM (effets secondaires) rbs; Cortisone rYmman Rétention (rétention Na+) nig kar)at;bg;nUv K+ EdlGacbNþal[man Insuffisance cardiaque congestive . \T§iBlbnÞab;bnSM epSgeTot rYmPaBexSayénsac;dMu ostéoporose vibtþipøÚvGahar rYmman dMe)ARkBH (ulcère) nig eraKsBaØasésRbsaTrYmman vilmux (vertige) QWk,al (céphalée) nig Rbkac; (Convulsion) .

H3C CO—CH3

CH3

O

H3C OH

CH3

O

H3C O

HO

H3C CO—CH3

O

H3C

O

O ║

H—C CO—CH2OH

CH3

O

HO H3C

CO—CH2OH

CH3

O

OH O



Aldostérone Ca hormone stéroide pliteday Cortex surrénale nigbeBa©jeTAkñúg crnþQamenAeBlkMriténGIuy:ug Na+ TabeBk . GaRs½yedayeTAdl;CalikacMNucenAkñúgtMrgenam Aldostérone eFIV[RbtikmμmYycMnYnmanskmμPaBEdlbgá[ Na+ nig TwkRtLb;cUleTAkñúgQamvij . RbsinebIkMritén Na+ ekIneLIgx<s; Aldostérone minRtUv)anbeBa©jBI Cortex surrénale eT nig Na+

RtUv)aneRcaHecjeRkAQamedaytMrgenam nig beBa©jecaleTAxageRkA . K- RkmYn (Cires)

RkmYnmanRsLayBIRbPBepSg² nig mansmasPaBKImIepSg² GaRs½yeTAtamRbPBrbs;va. ]TahrN_ Paraffine ekIteLIgBIl,ayénGIuRdUkabYrrwg ¬CasmasFatuExSRtg;¦ . CaFm μtaRkmYnFmμCati pSMeLIgBIGasIutRkaEvg²P¢ab; (estérifier) eTAnwgGal;kulEdlmanExSEvg² . GaRs½yedayknÞúy GIuRdUkabYrEdlmanExSEvgmincUlcitþTwk (hydrophorbe) RkmYnminrlaykñúgTwkesaH . RkmYnk¾Ca GgÁFaturwgenAsItuNðPaBFmμta GaRs½yma:s;m:UlelKulrbs;vax<s; . ]TahrN_énRkmYnBIr KW Myricyl

palmitate KWCasmsPaBsMxan;rbs;RkmYnX μ úM nig xøaj;RtI)aELnEdlekItBI Cetyl palmitate . CaFm μta RkmYnEdlekIeLIgmankareRbIR)as;epSg² . Lanoline RtUv)aneKeRbICaeRKOgkarBar sk; nig Es,k RtUv)aneRbIeRkamTMrg;Ca ERkmlabEs,k nig Ointments . RkmYn Carnauba RtUv)aneK eRbIkúñgkarb:Ularfynþ . eRbgRtI)aELn enAeBlmYyenaHRtUv)aneKeRbICaeRbg\n§³ enAkñúg ointments

nig kñúgkareFIVeTon . b:uEnþRkmYnsMeyaKRtUv)aneKeRbICMnYsxøaj;RtI)aELn enAkñúgvis½yCaeRcIn GaRs½y edaykarhamXat;karensaTRtI)aELn .

5- lIBIdkMupøic (LIPIDES COMPLEXES)

O ║ CH3(CH2)14—C—O—(CH2)29CH3

Myristyl palmitate (Beeswax)

O ║ CH3(CH2)14—C—O—(CH2)15CH3

Cetyl palmitate (Whale oil)



rUbTI 18-8 KMrUénrcnasm<½n§rbs; Lipoprotéines enAkñúg)aøs μa

Lipides complexes KWCalIBIdEdlRtUv)ancgsm<½n§CamYym:UelKulRbePTepSgeTot . Lipides complexes EdlCYbjwkjab; nig sMxan;CageKbMputKW Lipoprotéines enAkñúg Plasma Edl TTYlxusRtUvkñúgkardwknaMlIBIdepSgeTotenAkñúgsBa’gÁkay . lIBIdrlayedayticbMputenAkñúgTwk nig clnarbs;lIBIdBIsrIr³mYy (organe) eTAsrIr³epSg eTottamry³crnþQamRtUvkarRbB½n§dwknaMEdleRbI Lipoprotéines enAkñúg plasma . PaKl¥itén Lipoprotéines mansñÚl (core) rbs;lIBIdEdlmincUlcitþTwkB½n§CMuvijeTAeday Protéines,

Phospholipides, nig Cholestérol EdlBYkvaTaMgenHmanlkçN³ Amphotère ¬rUbTI 18-8¦ . vamanbYnfñak;sMxan;én Lipoprotéines enAkñúg)aøsμarbs;mnusS . Chylomicrons Edlmandg;sIuetTabCag 0.95 g/mL dwknaM Triglycérides éncMNIGahar BIeBaHevontUceTACalikaepSgeTot . Lipoprotéines epSgeTotRtUv)aneKeFIVcMNat;fñak;eTAtamdg;sIuet rbs;BYkva . Very Low Density Lipoprotéins (VLDL) mandg;sIuetBIr (0.95-1.019) . BYkvaP¢ab; eTAnwg Triglycérides EdlsMeyaK)anenAkñúgeføIm nig dwknaMBYkvaeTA Adipocytes nig CalikasþúkTuk epSgeTot . Low Density Lipoproteins (LDL) RtUv)anbBa¢ak;lkçN³edaydg;sIuetrbs;enAcenøaH BI 1.019-1.036 g/mL . BYkvadwknaM Cholestérol eTACalikaCay² (Tissues périphériques) nig CYy



EktMrUvkMritrbs; Cholestérol enAkñúgCalikaTaMgenaH . BYkvaTaMgenHsMbUr Cholestérol Cajwkjab; Edlvaman 40 % én Cholestérol enAkñúg)aøsμa . High Density Lipoproteins (HDL) mandg;sIuetcenøaHBI 1.063-1.210 g/mL . BYkva cgP¢ab;eTAnwg Cholestérol enAkñúg)aøs μa b:uEnþBYkvadwknaM Cholestérol BICalikaenACay² mkeføImvij . rUbTI 18-9 bgðajedaysegçbBIsmasPaBrbs; Lipoprotéines enAkñúg)aøsμa . Chylomicron KWCakMNkcgP¢ab;Kña (aggrégats) én Triglycérides nig RbUetGIunEdldwknaM Triglycérides cMNIGahareTAekasikaBaseBjsBa’gÁ . minEmnlIBIdTaMgGs;enAkñúgQammanRsLay BIGahar . Triglycérides nig Cholestérol k¾RtUv)ansMeyaKenAkñúgeføImEdr nig k¾RtUv)andwknaMtam ry³QamenAkñúPaKl¥itrbs; Lipoprotéines Edr . Triglycérides RtUv)ancUleTAkñúgPaKl¥it VLDL

EdldwknaMm:UelKullIBIdsMbUrfamBleTACalikaEdlRtUvkarRbPBfamBl b¤ eTACalikaxøaj; (tissue

adipeux) sMrab;karsþúkTuk . dUcKñaenHEdr Cholestérol cUleTAkñúgPaKl¥it LDL sMrab;kardwknaMBIeføIm eTACalikaCay² .

rUbTI 18-9 segçbéncMnYnRbEhlén Cholestérol, Phospholipides,Protéines, Triglycérides nig

Cholestérol estérifié enAkñúgfñak;TaMgbYnrbs; Lipoprotéines



karcUlénPaKl¥it LDL eTAkñúgekasikaTak;TgeTAnwg karTTYlsÁal; nig karcgmanlkçN³ Biess (Spécifique) rvagPaKl¥it LDL nig GñkTTYlEdlCaRbUetGIun (Recepteur protéique) EdlCab; enAelIPñasekasika . GñkTTYl LDL (LDL récepteur) RtUv)aneKeXIjenAkñúgPñasekasikaEdlenAeRkA eføIm nig TTYlxusRtUvcMeBaHkareRbIR)as;én Cholestérol edayekasikaénCalikaepSg² . LDL

¬Lipoprotéines EdlcgP¢ab;eTAnwg Cholestérol¦ P¢ab;edaymanlkçN³BiesseTAnwg LDL récepteur nig bNúþM (complexe) enHRtUv)anykcUleTAkñúgekasikaedaydMeNIkarmYyEdleKehAfa Endocytose

EdlsMrYYledayGñkTTYl (Receptor-mediated endocysis) rUbTI 18-10 . Pñasekasikacab;epþIm ptcUl (invaginate) Gacniyaymüa:geTotfa PñasekasikaTajcUleTAkñúgekasikaenATIkEnøgrbs; bNþúM LDL récepteur . karenHTajPaKl¥it LDL TaMgmUlcUleTAkñúgekasika . cugbBa©b; karptcUlk¾kan;Et eRCA ehIyPñasekasikaEdlB½T§CMuvijPaKl¥it LDL cakecjBIPñasekasika nig begáItnUvPñasCMuvij PaKl¥it LDL . dUcEdleyIgnwg)aneXIjenAkñúgEpñkTI 6 énemeronenH PñasekasikaCasnÞIy_ nig GachUrecjcUl)an . ehtudUecñH vaGacbegáItCa vésicule b¤ endosome EdlmanPaKl¥it LDL . PaKl¥itekasikaRbdab;rMlayGahar (Organelles de cellules digestives) RtUv)aneKsÁal;fa Ca Lysosomes rlayCamYynwg Endosomes . karrlayenHsMerceTA)anenAeBlPñasrbs; Endosome nig Lysosome hUrcUlKñaedIm,IbegáItGgÁFatuEdlcgP¢ab;KñaFMMmYyb¤ Vésicule . Enzyme

hydrolytiques BI lysosome enAeBlenaHrMlaybNþMúTaMgmUledIm,IbeBa©j Cholestérol eTAkñúg Cytoplasme rbs;ekasika . enATIenaH Cholestérol raraMgCIvsMeyaKrbs;va nig eFIV[skm μGg;sIum EdlsMrYlkarsþúkbMrugCadMNk; Cholestérol ester . kMhab;x<s;én Cholestérol enAkñúgekasikak¾ raraMg karsMeyaKrbs; LDL récepteur EdredIm,IeFIV[R)akdfaekasikanwgmincab;yk Cholestérol eRcIneBk. RbCaCanEdlmanPaBminl¥énEhSn (gène) enAkñúgEhSn EdlftcMLg (coder) nUv LDL récepteur

minGaccab;yk Cholestérol )aneRcIneT . CalT§pl BYkvanaM[mankarekIneLIg LDL cholestérol

enAkñúg)aøsμa . karelIsén Cholestérol enAkñúg)aøs μa bnÞab;mkKrenAelIépÞénGakETr (artère) bgá[ man Arthérosclérose . CMgWenHRtUv)aneKehAfa Hypercholestérolémie . Lipoprotéines récepteurs rbs;eføImmanlT§PaBeFIV[man Cholestérol cMnYneRcInRtUv)andk ecjBIkñúg Qam edayFana[mankMhab;Tabén Cholestérol enAkñúgQam . ktþaepSgeTotEdlesμ IKña buKÁlEdlman Récepteurs lipoprotéines eRcInbMputnwgTTYlrgkarekIneLIgkMrit Cholestérol nig kar ekItCMgW Arthérosclérose ticbMput . vak¾mankarbgðajEdrfa kMritx<s;én HDL enAkñúgQamCYykat;bnßyeRKaHfñak;én Arthéro-

sclérose RbEhlGaRs½yeday HDL dwknaM Cholestérol BICalikaCay²eTAeføIm . enAkñúgeføIm



Cholestérol mYycMnYnRtUv)aneRbIsMrab;karsMeyaKTwkRbma:t; nig beBa©jcUleTAkñúgeBaHevontUc BIGIV EdlvaRtUv)anbeBa©j . TMnak;TMngcugbBa©b;RtUv)aneFIVeLIgrvagrbbGahar nig Arthérosclérose . RbCaCnEdlrbb Gaharrbs;BYkeKmanGasIuRkaEq¥teRcInmanninñakareTArkkMritx<s;én Cholestérol enAkñúgQam . eTaHbICaTMnak;TMngrvagGasIutRkaEq¥t nig emtabUlIs cholestérol minc,as;k¾eday b:uEnþeKdwgfa rbb GaharEdlsMbUrGasIutRkaminEq¥tbNþal[mankarfycuHénkMrit LDL nig ekIneLIgnUvkMrit HDL . CamYynwg TMnak;TMngviC¢manrvagCMgWebHdUg nig kMritx<s;én Cholestérol Gnusan_énrbbGaharbc©úb,nñ cat;bBa©ÚlrbbGaharEdlmanxøaj;tic nig karCMnYsénxøaj;Eq¥t ¬xøaj;stV¦ edayxøaj;minEq¥t ¬eRbgrukçCati . 6- rcnasm<½n§énPñasCIvsaRsþ (LA STRUCTURE DE MEMBRANES BILOGIQUES)

PñasCIvsaRsþKWCaRsTab;lIBIdBIrCan; EdlknÞúyminRCabTwkRbmUlpúþMenAEpñkxagkúñgénRsTab;BIr Can; (bicouches) nig k,alEdlCabgÁúMGIuy:UniksßitenAelIépÞedIm,ImanRbtikmμCamYynwgTwk ¬rUbTI 18-11¦ . knÞúyGIuRdUkabYén Phospholipides rbs;Pñaspþl;nUvsbkesþIg (thin shell) énsarFatuminman b:UlEdlkarBarmin[mankarlaycUlKñaénm:UelKulTaMgsgxag . ehtudUecñH knÞúyminmanb:Ulrbs; phospholipides énPñaspþl;Car)aMgrvagEpñkxagkúñgrbs;ekasika nig mCÄdæanCMuvijva . Epñkk,alman b:Ulrbs;lIBIdRtUv)anb:HeTAnwgTwk nig BYkvaRCabTwkxøaMg (Solvate) . karpøas;bþÚrtictYc RtUv)aneKsÁal; CaFm μtafaCa {RtLb;RtLin} ekIteLIgrvaglIBIdenAelIBak;kNþalxageRkA nig lIBIdenABak;kNþal xagkñúg énRsTab;eTVr (bichouche) rUbTI 18-12 . clnarbs;m:UelKullIBIdenAkñúgRsTab;mYy (seath) énRsTab;eTVr vaelOn . ekasika)ak;etrImanRbEhlbeNþayRbEhl 2 µm nig m:UelKullIBId sayPayBIcugmçagénekasikaeTAcugmçagepSgeTotkñúgmYyvinaTI . RsTab;TaMgBIrrbs; RsTab;eTVrénPñas Phospholipides mansmasPaBmindUcKñaeT . GaRs½y ehtuenH enAkñúgekasikaRKab;QamRkhm RbEhl 80 % én Phospholipides enAkñúgRsTab;xageRkA énPñasKWCa Phosphatidylcholine nig Sphingomyéines rIÉ Phosphatidylethanolamine nig Phosphatidylserine manRbEhl 80 % cMeBaHRsTab;Epñkxagkñúg . epSgBIenH BYkKøúysIut P¢ab;eTAnwg Et Phospholipides EdleXIjmanenAelIRsTab;xageRkArbs;Pñas . enATIenH BYkvacUlrYmenAkñúg muxgar Récepteur nig muxgarTTYlsÁal; (Reconnaissance) .



k- rUbKMnUrbMRBYjrcnasm<½n§rbs;PñasCIvsaRsþ (Fluid Mosaic Structure of Biological Membranes)

dUcEdleTIbEt)anbBa¢ak; PñasminEmnCafamvnþ (static) BYkvapSMeLIgGMBIm:UelKulEdlman clna . snÞnIy_PaB (fluidity) énPñasCIvsaRsþRtUv)ankMNt;edaysmamaRténbgÁúMGasIuRkaEq¥t nig GasIutRkaminEq¥tenAkñúg Phosphlipides énPñas . RbEhlBak;kNþalénGasIutRkaEdlcMraj;ecjBI lIBIdénPñasBIRKb;RbPBKWCaGasIutRkaminEq¥t . knÞúyGasIutRkaminEq¥tén Phospholipides bgá[mansnÞnIy_PaBénPñas GaRs½yedaymMu (bends) EdlmanenAkñúgExSGIuRdUkabYedaysarsm<½n§BIrCan; . edaysar {karrmYl} énknÞúyGasIutRka TaMgenHminrYmcUlKñaedaytwgxøaMgeT . eyIgk¾eXIjEdrfa PaKryénGasIutRkaminEq¥tenAkñúglIBIdrbssPñasvaRcassmamaRteTAnwg sItuNðPaBrbs;briyakas . )ak;etrI Ca]TahrN_ manpleFob (ratios) énGasIutRkaEq¥t nig minEq¥t enAkñúglIBIdénPñasrbs;BYkva edayCab;Tak;TgeTAnwgsItuNðPaBenACMuvijxøÜnBYkva . GaRs½yehtuenH Pñasrbs;)ak;etrIEdllUtlas;enAkñúgmhasmuRTGakTik (Artic Ocean) mankMritGasIutRkaminEq¥tx<s; dUecñHPñasvaenAEtERbRbYleTaHbICaenAkñúgsItuNðPaBRtCak;xøaMgk¾eday . pÞúykmvij srIragÁ (organism) Edlrs;enAkñúgTwkekþAén Yellowstone National Park EdlmansIutuNðPaBCitBuH manPñas EdlmankMritGasIutRkaEq¥tx<s; . PaBGacbt;Ebn (flexicibility) énGasIutRkaenHbNþal[)ak;etrIrkSa nUvsnÞnIy_PaBdUcKñaenAkñúgcenøaHsItuNðPaBesÞIrEt 100oC .

rUbTI 18-11 (a) rUbPaBrbs; Phospholipides (b) KMrUéncenøaHrbs; Phospholipides

(c) rUbPaBénPñasRsTab;eTVr Phospholipides (d) karbgðajnUvrUbmnþCaExSénPñas RsTab;eTVrEdlpSMeLIgBI Phospholipides, cholestérol nig Sphingolipides



rUbTI 18-12

CaTUeTAsItuNðPaBdgxøÜnénBYkfnikstVKWefrxøaMgNas; nig smasPaBGasIutRkaénlIBIdrbs; PñasCaFm μtavamanÉksNæanxøaMgNas; . karelIkElgmYyEdlKYr[cab;Garm μN_KWstVeRbIsTwkkk (Reindeer) . eRcInqñaM eRbIsTwkkkRtUvEteFIVdMeNIrqøgkat;Twkkk nig RBil . ehtudUecñHRkck nig Epñk xageRkaméneCIgbnþbMeBjmuxgarenAkñúgsItuNðPaBRtCak;CagEpñkepSg²eToténdgxøÜn . GaRs½yehtu enH PaKryénPaBminEq¥tenAkñúgPñasERbRbYleTAtambeNþayeCIgénstVeRbITwkkk . eyIgrkeXIjfa smamaRténGasIutRkaminEq¥tekIneLIgenAEpñkEdlenACitRkck . cMNucEdlenAkan;EtCitPaBRtCak; kan;EtxøaMg nig snÞnIy_PaBénlIBIdkan;EtxøaMgEdlmansmamaRtGasIuRkaminEq¥tkan;Etx<s; GnuBaØati[ PñasbMeBjmuxgarenAkñúgsItuNðPaBTabénTwkkk nig RBilEdlEpñkxageRkaméneCIgrbs;vaCan;elI . ehtudUecñH PñasvaERbRbYl edayminKitGMBIlkçx½NÐsItuNðPaBénbriyakas . CakarBit eK)an )a:n;s μanfaPñasmanPaBhab; (Consistency) dUceRbgGUlIv . eTaHbICarnaMgminRCabTwkRtUv)anbegáIteLIgedayRsTab;eTVrlIBIdEdlERbRbYl (fluid lipid

bilayer) KWCaTidæPaBsMxan;rbs;Pñas k¾b:uEnþ RbUetGIunEdlmanenAkñúgRsTab;eTVlIBIdk¾manlkçN³sMxan; Edr nig TTYlxusRtUvcMeBaHmuxgarsMxan;rbs;ekasika . vtþmanénRbUetGIunrbs;PñasTaMgenHRtUv)angðaj eday bec©keTsmIRkUTsSn_eGLikRtug (electron microscopic technique) EdleKehAfa "Free-

Fracture) . ekasikaRtUv)aneKykeTAbgákenAkñúgsItuNðPaBTabya:gxøaMg nig bnÞab;mkRtUv)anEjk edayeRbIkaMbiteFIVBIeBRCya:gtUcbMput . ekasikamYycMnYnRtUv)anEjkrvagRsTab;TaMgBIrénRsTab;eTVr lIBId . enAeBlRtUv)anbgðajedaymIRkUTsSn_eGLikRtug PñaselcecjCarUbPaB (mosaic) RtUv)an P¢ab;CamYynwgRbUetGIun . GaRs½yedaysnÞnIy_PaBrbs;Pñas nig karelcecjnvRbUetGIunRtUv)anemIl



eXIjedaymIRkUTsSn_eGLikRtugKMnitrbs;eyIgGMBIrcnasm<½n§RtUv)aneKehAfaCa KMrUrUbPaBEdl ERbRbYl (Fluid mosaic model) ¬rUbTI 18-13¦ . RbUetGIunmYycMnYEdleKBinieXIj RtUv)aneKehAfaCa RbUetGIunenACayénPñas (Peripheral

membrane proteins) RtUv)ancgP¢ab;eTAépÞmYyénPñasedaysarGMeBIeTAvijeTAmkénrvagbgÁúMk,al EbkCaGIuy:ugénlIBIdrbs;Pñas nig GasIutGamIneNEdlEbkCaGIuy:ugenAépÞrbs;RbUetGIunenACay (peripheral proteins) . RbUetGIunrbs;PñasepSgeTot RtUv)aneKehAfaCa RbUetGIunqøgkat; (trans-

membrane proteins) RtUv)ancgenAkñúgPñas niglatsn§wgeBjkMras;Pñas ehIyb:HTaMgEpñkxagkñúg nig EpñkxageRkArbs;ekasika . k¾dUceTAnwg smasPaBrbs; Phospholipides énPñasminsIuemRTI (asymmetric) dUecñHEdr KWCa karcgú¥lTisedAénRbUetGIunqøgkat; (transmembrane proteins) . RbUetGIunqøgkat;nimYy²mantMbn; minRCabTwkEdlrYmCamYyknÞúyGasIutRkaén Phospholipides rbs;Pñas . vanimYy²k¾manEpñkEdl RCabTwkEdlCaTUeTAeKeXIjrYmCamYynwgRsTab;xageRkAénPñas nig sßitenAelIEpñkxageRkAén ekasika. tMbn;enHénRbUetGIunCaBiessmannUv Oligosachharides EdlRtUv)anP¢ab;tamry³ sm<½n§kUva:Lg; . ehtudUecñHBYkvaenHKWCa Glycoproteins . dUcKñaenHEdr RbUetGIunqøgkat;nimYy²man EpñkRCabTwkTIBIr EdlCaTUeTAeKeXIjrYmCamYyRsTab;xagkñúgénPñas nig lycUleTAkñúgsIutU)aøsrbs; ekasika . edaylkçN³Biess tMbn;enHénRbUetGIunqøgkat;RtUv)anP¢ab;eTAnwg Filaments éneRKagrbs; sIutU)aøs . GaRs½yedayRsTab;eTVrlIBIdvaERbRbYl vaGacmankarsayPaytamExSepþkya:gelOn (latheral diffusion) énRbUetGIunrbs;Pñastamry³RsTab;eTVrlIBId b:uEnþRbUetGIunrbs;Pñas k¾dUcCalIBId rbs;Pñas minRtLb;cuHeLIg (flip-flop) enABaseBjPñas b¤ vilRtLb;cUleTAkñúgdUceTAnwgRtecokTVar énGaKar . PñasKWCarcnasm<½n§Edlpøas;bþÚrCaRbcaM (dynamic structure) dUcEdleyIgGacTajecjBI cMeNHdwgrbs;eyIgGMBIPaBmanclnarbs;RbIetGIun nig lIBIdénPñas . PaBmanclnaénRbUetGIunEdl sßitenAkñúgPñasCIvsaRsþRtUv)ansikSaedaykarbiTpøakRbUetGIunmYycMnYncMeBaHPñasekasikamnusS nig kNþúrCamYynwgB’N fluorescent Rkhm nig ébtg . ekasikamnusS nig kNþúrRtUv)anrlaycUlKña Gac famüa:geTot bec©keTsBiessRtUv)aneKeRbIedIm,IeFIV[Pñasrbs;ekasikakNþúr nig ekasikamnusShUr cUlKñaedIm,IbegáItCaekasikaEtmYy . ekasikaf μIRtUv)aneKBinitüeXIjtamry³mIRkUTsSn_BiessEdleRbI BnøW UV b¤ Fluorescence . r)aMg (patches) B’NRkhm nig ébtgRtUv)aneKdak;enAkñúgtMbn;énPñas ekasikaedImeBlkarBiesaFn_cab;epþIm . mYyema:geRkaymk r)aMgB’NRtUv)anEbgEckedayminman



ÉksNæanenAkñúgPñasekasikaEdlrlay ¬rUbTI 18-14¦ . karBiesaFn_enH)anpþl;Caeyabl;fa eyIgGacKitGMBI PñasEdlCarUbGacERbRbYl (fluid mosaic membrfane) faCamhasmuRTRtUv)an bMeBjedaypÞaMgTwkkkGENþteTAmk . x- kardwkCBa¢Únqøgkat;Pñas (Membrane transport)

PñasekasikasMrYlRbtikmμeTAvijeTAmkénekasikaCamYybriyakasCMuvijva nig TTYlxusRtUv cMeBaHkarRtYtBiniükarqøgkat;ecjBI nig cUleTAkñúgekasikaénsarFatu . PñasxageRkAénekasikaRtYt BinitükarcUlén\n§³ nig karbeBa©jnUvplitplkaksMNl; . PñasxagkñúgénekasikabMEbkemtabUlIt kñúgcMeNamPaKli¥trbs;ekasika (cell organelles) . PaKeRcInéndMeNIrkarénkardwkCBa¢ÚnTaMgenHRtUv )anRtYtBinitüRbUetGIundwknaMqøgkat;Pñas (transmembrane transport proteins) . RbUetGIundwknaM TaMgenHKWCaGñkcaMTVar (gatekeeper) Edlmuxgarrbs;vaenAkñúgkardwknaMénPñasdUceTAnwgmuxgarrbs; Gg;sIumkñúgkarGnuvtþn_RbtikmμKImI . b:uEnþ m:UelKulmYycMnYnqøgkat;tamPñasedayK μanCMnYy edaysar dMeNIrkardwknaMGkm μ (passive transport processes) énkarsayPay nig GUs μ Ú (Osmosis) . dMeNIr karTaMgenHsMedAcMeBaHdMeNIrkarGkmμ BIeRBaHBYkvaminRtUvkarcMNayfamBlNamYyedayekasikaeT.

rUbTI 18-13 KMrUrUbPaBEdl ERbRbYl (Fluid mosaic model)



1-karsayPayGkmμ ³ TMrgsamBaØCageKbMputénkardwknaMqøgkat;Pñas RbsinebIeKdak;Etkk (instant iced tea) mYYysøaRBakaehVenAelIépÞénTwkmYyEkv mU:UelKulEt vasayPayya:gelOnBaseBjsUluysüúg . m:UelKul Faturlay (solute) ¬Et¦ nig FaturMlay (solvent) ¬Twk¦ RtUv)anrulcUlKñaedaysarclnaedayécdnürbs;m:UelKul . EtEdlxab;BIdMbUgk¾cab; epþImraveTA² . dMeNIrkarénclnasrubrbs;FaturlayCamYynwgFatupSM ¬BIkEnøgEdlmankMhab;x<s; eTA kEnøgEdlmankMhab;Tab¦ RtUv)aneKehAfa karsayPay (diffusion) ¬rUbTI 18-15¦ karsayPay KWCameFüa)aymYyénkardwknaMGkmμqøgkat;Pñas . \LÚveyIgsnμtfaPñas CIvsaRsþman nig sarFatuRtUv)aneKeXIjmankMhab;mYyenAxageRkAPñas nig Bak;kNþalénkMhab;enA xagkñúgPñas . RbsinebIFaturlayGacqøgkat;Pñas karsayPaynwgekIteLIg rYmCamYynwgkardwknaM srubénsarFatuBItMbn;EdlmankMhab;BIdMbUgx<s;eTAtMbn;EdlmankMhab;BIdMbUgTab nig sarFatunwgman lMnwgkñúgkarqøgkat;Pñasekasika ¬rUbTI 18-16¦ . bnÞab;BIry³eBlmYy kMhab;rbs;sarFatunwgdUcKña TaMgsgxagPñas RbB½n§nwgmanlMnwg nig Elgmankarpøas;bþÚrsrubekIteLIgeTotehIy .

rUbTI 18-14 karbgðajfaPñasERbRbYl nig RbUetGIumanclnaesrIenAkñúgépÞrabénRsTab;eTVlIBId

rUbTI 18-15 karsarPaybNþal[manclnasrubénm:UelKulsárnigTwkBItMbn;EdlmankMhab;x<s;

eTAtMbn;EdlmankMhab;Tab . CacugbBa©b;kMhab;énsárnigTwkBaseBj Beaker es μ IKña



etItMélkñúgcMeNaménkarGnuvtþn_KWCadMeNIrkarsayPaycMeBaHekasikab¤ ? edayBitR)akd karsayPayGacEbgEckemtabUlItedayRbsiT§PaBBaseBEpñkxagkñúgénekasika . b:uEnþ etIGIVEdl niyayGMBIclnarbs;m:UelKulqøgkat;Pñas ? GaRs½yedayrcnasm<½n§énRsTab;eTVrlIBIdrbsPñas manEtm:UelKultictYcGacsayPayqøgkat;Pñas . m:UelKulTaMgenHrYmmandUcCa O2, CO2, nig H2O . m:UelKulEdlFM b¤ manbnÞúkGKÁsnIx<s; b¤ GIuy:ug minGacqøgkat;RsTab;eTVrlIBIdedaypÞal;eT . dUecñH mU:elKulRtUvkarCMnYyBIRbUetGIunénPñasekasika . PñasNamYyEdlGnuBaØati[mankarsayPaynUv m:UelKulmYycMnYn b:uEnþm:UelKulepSgeTotminGacsayPayeKniyayfavaCa kareRCabedayeRCIserIs

(slectively permeable) . rUbTI 18-16

karsarPayénFaturlaytamry³Pñas

2- karsayPayedaymankarsMrYl ³ lkçN³BiessénkardwknaMqøgkat;Pñas (Facilitated Diffusion : Specificity of Molecular Transport)

PaKeRcInénm:UelKulRtUv)andwknaMqøgkat;Pñasedaysar GñkdwknaMBiessEdlCaRbUetGIun (specific protein carriers) EdleKsÁal;faCa Permeases . enAeBlFatrlaysayPayqøgkat;



PñasBItMbn;mankMhab;x<s;eTAtMbn;mankMhab;Tabedaykarqøgkat;tampøÚvenAkñúg Permease dMeNIrkar enHRtUv)aneKsÁal;faCa karsarPayedaymankarsMrYl (Facilitated diffusion) . minmanfamBlRtUv )aneRbIR)as;edaykarsayPayedaymankarsMrYleT . ehtudUecñHvaKWCameFüa)ayeToténkardwknaM Gkmμ nig TisedAénkardwknaMGaRs½yeTAnwgkMhab;rbs;emtabUlItEdlmanenAkñúgEpñknimYy²énPñas. kardwknaMqøgkat;Pñasrbs;ekasikaedaysar karsayPayedaymankarsMrYlekIteLIgeTAtam ry³rn§enAkñúg Permease EdlmanTMrg; b¤ rUbrag EdlbMeBjKñaeTAnwgTMrg; b¤ rUbragm:UelKulEdlRtUv dwknaM . bnÞúkGKÁisnI nig TMrg;énrn§kMNt;nUvPaBBiess (specificity) rbs;GñkdwknaM ¬rUbTI 18-17¦. manEtm:UelKulEdlmanrUbragRtUvKñaEdlGaccUleTAkñúgrn§ . CalT§pl GRtaénkarsayPaycMeBaH m:UelKulNamYyRtUv)ankMNt;edaycMnYnm:UelKul permease enAkñúgPñasEdlTTYlkarxusRtUvcMeBaH karqøgkat;énm:UelKulenaH . karqøgkat;rbs; glucose KUsbBa¢ak;BIPaBBiessén permease EdlCaGñkdwknaM . D-glucose

RtUv)andwknaMedayGñkdwknaM glucose b:uEnþ enantiomer rbs;va L-glucose minGacRtUv)andwknaMeT . ehtudUecñH GñkdwknaM glucose bgðajBI stereospecificity . müa:gvijeTot FaturlayRtUv)annaMcUleTA kñúgekasikaRtUvEtRtUvKñaedayCak;lak; ¬dUcCaéd nigeRsamédGBa©wgEdr¦ eTAkñúgTIkEnøgén karTTYlsÁal;enAkñúgrcnasm<½n§rbs; permease . enAkñúgCMBUkTI 20 eyIgnwgeXIjfaGg;sIumEdlsMrYl RbtikmμKImIenAkñúgekasikabgðajBIRbePTénPaBBiessdUcKñaenH . GRtaénkardwknaMrbs;emtabUlItcUleTAkñúgekasikaman\T§iBlya:gxøaMgeTAelIGRta emtabUlIs srubén ekasikaCaeRcIn . Insulin KWCa Polypeptide hormone RtUv)ansMeyaKedayekasika β

énlMEBg (panceas) begáInGRtaGtib,rmaénkardwkCBa¢Ún Glucose edayplKuN (factor) BI 3 eTA 4. lT§plKWfaskmμPaBemtabUlIsrbs;ekasikaekIneLIgya:gxøaMg .

rUbTI 18-17 kardwknaMedaykarsayPayekIteLIgtamry³rn§enAkñúgRbUetGIundwknaMEdlTMhM nig

rUbragrbs;vaRtUvKñaeTAnwgTMhM nig rUbragrbs;m:UelKulEdlRtUvdwknaM



rUbTI 18-18 kardwknaMén Cl- nig HCO3

- enAkñúgTisedApÞúyKñaqøgkat;PñasekasikaRKab;QamRkhm

ekasikaRKab;QamRkhmeRbIpøÚv anion dUcKñakñúgkardwkCBa¢Ún Cl- cUleTAkñúgekasikakñúgkar bþÚrCamYynwgGIuy:ug HCO3

- ¬rUbTI 18-8¦ . kardwknaMeTAmkrebobenH (Two-way transport)

RtUv)aneKsÁal;faCa Antiport . kardwknaMrebobenHekIteLIgedaykarsayPayedaymankarsMrYl dUecñH Cl- hUrBIxageRkAEdlmankMhab;x<s;cUleTAxagkñúgEdlmankMhab;Tab nig HCO3

- hUrBIxagkñúg EdlmankMhab;x<s; eTAEpñkxageRkAEdlmankMhab;Tab . 3- GUs μ Ú ³ clnaGkmμénFaturMlayqøgkat;Pñas (Osmosis : Passive Movement of a Solvent Across a Membrane)

GaRs½yedayPñasekasikaGacRCabedayeRCIserIs CaTUeTAvaminGacRCabcMeBaHFaturlay kñúg karqøgkat;PñaskñúgkareqøIytbeTAnwgkarERbRbYlénkMhab;rbs;. kñúgkrNIrebobenHFaturMlay sayPay qøgkat;Pñas . PñasrebobenH GacRCabcMeBaHFaturMlay minRCabcMeBaHFaturlay eTRtUv)aneKehA edayBiessfaCa Semipermeable membrane . Osmosis KWCakarsayPay énFaturMlay ¬TwkenA kñúgRbB½n§CIvsaRsþ¦ qøgkat; semipermeable membrane kñúgkareqøIytbeTAnwg karERbRbYlénkMhab;rbs;Twk . cUrsnμtfa eyIgdak; 0.5 M énsUlusüúg glucose enAkñúgeTAkñúgfg; Dialysis EdlpSMeLIgBIPñas manrn§EdlGnuBaØati[mankarqøgkat;énm:UelKulTwkb:uEnþm:UelKul glucose minGacqøgkat;)aneT . cUr BicarNafaGIVnwgekItmaneLIg enAeBleYIgdak;fg;enaHcUleTAkñúg Beaker TwksuT§ (pure water) . eyIg )anbegáItnUvcMnYnsarFatu (gradient) EdlvamankMhab;x<s;én glucose enAEpñkxagkñúgCagEpñkxageRkA fg; b:uEnþ glucose minGacsayPayqøgkat;Pñasénfg;edIm,ITTYl)ankMhab;es μ IKñanUvEpñkTaMgsgxag énPñaseT .



\LÚveyIgKitBIsßanPaBenHenAkñúgrebobepSgmYyeTot . eyIgmankMhab;x<s;énm:UelKulTwkenA EpñkxageRkAfg; ¬EdlenATIenaHmanEtTwksuT§¦CagEpñkxagkñúgfg; ¬EdlenATIenaHmanmU:elKulTwkmYy cMnYnrYmCamYynwgPaKl¥iténFaturlayEdlminGaceFIVclnaedayesrIenAkñúgRbB½n§ . GaRs½yedayTwk GacsayPayqøgkat;Pñas karsayPaysrubénTwknwgekIteLIgtamPñascUleTAkñúgfg; . karenHKWCa dMeNIrkarén GUs μ Ú (Osmosis) ¬rUbTI 18-19¦ . dUcEdlGñkRbEhlCa)annrYcmkehIy RbB½n§enHminGacQandl;lMnwgeT ¬kMhab;esμ IénEpñk TaMgBIr¦ BIRBaHedayminKitBIfaetITwkcMnYnb:unμansayPaycUleTAkñúgfg; edayBRgavsUluysüúg glucose kMhab;rbs; glucose nwgCaTUeTAx<s;enAEpñkxagkñúgfg; ¬nigrYmedaykMhab;TwkesrITab¦. etIGIVekIteLIgenAeBlfg;cab;ykTwkeRcIneBk enAeBlenaHvaGacrIkFMeLIg ? \LÚvCBa¢aMgénfg; eRbIkMlaMgEdlnwgbBaÄb;karhUrsrubénTwkqøgkat;PñascUleTAkñúgfg; . sMBaFGUs μ Ú (Osmotic

pressure) KWCasMBaFEdlRtUvEt)aneFIVeLIgedIm,IbBaÄb;karhUrénTwkqøgcUledayeRCIserIsénPñas permeable edaysarGUsμÚ . edayEføgedayRtwmRtUv sMBaFGUs μ Ú énsUluysüúg KWCasMBaFsrubrYmCa mYyGIVEdlTwkcUleTAkñúgvaedayGUs μ ÚBIEpñkTwksuT§ enAeBlEpñkTaMgBIrRtUv)anx½NÐeTAedayPñas semipermeable .

rUbTI 18-19 GUs μ Úqøgkat;Pñas . FaturMlay ¬Twk¦ sayPayBItMbn;mankMhab;FaturlayTab

eTAtMbn;EdlmankMhab;Faturlayx<s;

More dilute More concentrated

Solute

Water



kMhab;GUs μ Ú (Osmotic concentration) b¤ Osmolarity KWCaBaküeKeRbIkñúgkarBiB’NnakMlaMgGUsμ Ú rbs;sUluysüúg . vaTak;TgEtelIpleFobéncMnYnPaKl¥iténFaturlayeTAnwgcMnYnPaKl¥iténFatu rMlay . ehtudUecñH Fm μCatiénCatiKImI nig TMhMrbs;FaturlayminsMxan; manEtkMhab;EdlRtUvsMEdgCa Molarity . GaRs½yehtuenHsUluysüúg 2 M rbs; glucose ¬m:as;m:Ulrbs; glucose : 180¦ maN Osmolarity dUcKñaeTAnwgsUluysüúg 2 M rbs; Albumin ¬ma:s;m:Ulrbs; Albumin : 60,000¦ . )aøs μaman Osmolarity smmUleTAnwgsUluysüúg 0.30 M rbs; glucose b¤ sUluysüúg 0.15

M rbs; NaCl . sUluysüúgcugeRkayenHKWvaBit BIeRBaHenAkñúgsUluysüúg NaCl bMEbkeTACa Na+ nig Cl- nig ehtudUecñHvabNþal[mancMnYnPaKl¥iténFaturlayBIrdgénm:UlelKulEdlminEbkCaGIuy:ug . RbsinebIRKab;QamRkhmEdlman osmolarity es μ IeTAnwg)aøsμa RtUv)andak;eTAkñúgsUluysüúg glucose

0.30 M minman)atuPUt osmosis ekIteLIg BIeRBaH osmolarity nig kMhab;TwkenAkñúgekasikaRKab;Qam Rkhmes μ IeTAnwg osmolarity rbs;sUluysüúg glucose 0.30 M . sUluysüúgenAxagkñúg nig xageRkA énekasikaRKab;QamRkhmRtUv)aneKehAfa Isotonic solution ¬iso mann½yfadUc nig tonic man n½yfakMlaMg¦ . GaRs½yeday Osmolarity dUcKñaTaMgxagkñúg nig xageRkA ekasikaQamRkhm nwgsßit enAkñúgTMhMdEdl ¬rUbTI 18-20b¦ . etIGIVEdlekIteLIg RbsinebI\LÚvenHeyIgdak;ekasikaQamRkhmeTAkñúg Hypotonic

solution Gacfamüa:geTotfaCasUluysüúgman Osmolarity TabCag osmolarity rbs;sIutU)aøs; rbs;ekasika ? enAkñúgsßanPaBenHclnaénTwkcUleTAkñúgekasikaenAeBlEdlTwksayPaybBa©úHkMhab; éncMnYnsarFatu . PñasekasikaRKab;QamRkhmminmankMlaMgkñúgkarTb;Tl;nwgsMBaFRKb;RKanedIm,I bBaÄb;karhUrrbs;TwkenH nig ekasikaeLIge)a:g nig Ebk ¬rUbTI 18-20c¦ . müa:gvijeTot RbsinebI eKdak;RKab;QamRkhmcUleTAkñúg Hypertonic solution ¬sUluysüúgEdlman osmolarity FMCag ekasika¦ TwknwgecjBIekasika nig BYkvarYmtUcya:gxøaMg ¬rUbTI 18-20a¦ . eKalkarN_TaMgenHmankarGnuvtþn_ya:gxøaMgkñúgkareRbIsUluysüúgtamsés (Intravenous

solution :IV) eTAelIbuKÁlNamñak; . CaFmμtaGgÁFaturavNamYy (fluids) RtUv)anbBa©ÚltamsésRtUvEt man osmolarity RtwmRtUv BYkvaRtUvEt Isotonic CamYynwgekasikaQam nig )aøsμa . karcak;bBa©Úl rebobenHCajwkjab;man 5.5 g én glucose enAkñúg 100 mL énsUluysüúg (0.30 M) nig 0.9 g én NaCl kñúg 100 mL énsUluysüúg (0.15 M) . kñúgkrNITaMgBI BYkvaTaMgBIrmansMBaFGUsμ ÚdUcKña nig Osmolarity dUceTAnwg )aøs μa nig ekasikaQam nig ehtudUecñHBYkvaGacRtUv)aneRbIedaysuvtßiPaB edaymineFIV[xUctulüPaBrvagQam nig ekasikaQam .



rUbTI 18-20 rUbfténRKab;QabRkhmdak;enAkñúg (a) Hypertonic, (b) Isotonic (c) hypotonic solution

rUbTI 18-21

KnUsbMRBYjénRbtibtþikarrbs; Na+-K+ ATPase

K- tMrUvkarfamBlsMrab;kardwknaM (Energy Requirements for Transport)

karsayPaysamBaØ karsayPayedaymankarsMrYl nig GUsμ Ú Cab;Tak;TgeTAnwgkarhUreday ÉkÉgnUvsarFatuBItMbn;EdlmankMhab;x<s;eTAtMbn;EdlmankMhab;TabCag ¬PaBeTrénkMhab;¦ . edIm,I rs; ekasikaCajwkjab;RtUvEtbBa¢ÚnsarFatu {eLIgPñM} RbQmnwgPaBeTrénkMhab; (concentration

gradient) . )atuPUtuenH RtUv)aneKehAfaCa kardwknaMskmμ (Active transport) RtUvkarfamBl . GIuy:ug nig m:UelKulcMNIGaharCaeRcInRtUv)annaMcUlqøgkat;Pñasekasikatamry³kardwknaMskmμ . famBlEdleRbIsMrab;dMeNIrkarenHGacRtUvkareRcInCagBak;kNþal énfamBlsrubEdlRbmUl)an edayemtabUlIsrbs;ekasika . ]TahrN_Edll¥énkardwknaMskm μKW Na+-K+ ATPase EdleFIV[GIuy:ug



TaMgenHmanclnacUleTAkñúg nig ecjeRkAekasikaRbQmeTAnwgPaBeTrrbs;BYkva ¬rUbTI 18-21¦. ekasikaRtUvEtrkSakMhab;x<s;én Na+ enAxageRkAekasika nig kMhab;x<s;én K+ enAxagkñúgekasika . karenHRtUvkarkarpþl;Canic©rnþnUvfamBlekasikaenAeRkamTMrg;Ca Adenosine Triphosphate (ATP) . eRcInCag 1/3 én ATP srubRtUv)anplitedayekasikaRtUv)aneRbIsMrab;rkSaPaBeTrénkMhab; Na+ nig K+ TaMgenH Edlqøgkat;PñasekasikaenH . ehtudUecñHeQ μaH Na+-K+ ATPase sMedAcMeBaHskm μPaB Gg;sIumEdlGIuRdUlIs ATP . GLuRdUlIs ATP beBa©jnUvfamBlEdlRtUvkaredIm,IeFIVclnaGIuy:ug Na+

nig K+ qøgkat;Pñasekasika . cMeBaHm:UelKul ATP nimYy²EdlRtUvGLuRdUlIs 3 GIuy:ug Na+ecjBI ekasika nig 2 GIuy:ug K+ cUleTAkñúgekasika .

Fin Reference : General, Organic, and Biochemistry 3rd edition, 2001 by Denniston, Topping, Caret page 495-552

Biochemistry, Basic Protein Structure and Function

prepared by Mr. Chroeng Sopheap, phone: 012864353 or 011717361, e-mail: [email protected] Page 1 of 30

rcnasm<½n§RbUetGIun nig muxgar (Protein Structure and Function)

vtßúbMNgemeron 1-rab;muxgarrbs;RbUetGIun 2-KUrrcnasm<½n§TUeTArbs;GasIutGamIneN nig eFIVcMNat;fñak;GasIutGamIneNedaysMGagelIbgÁúM R rbs;BYkva . 3-BiB’NnaBIrcnasm<½n§TI 1 rbs;RbUetGIun nig KUrrcnasm<½n§énsm<½n§ peptide . 4-KUrrcnasm<½n§ peptides tUc² nig [eQ μaHBYkva . 5-BiB’NnaBIRbePTénrcnasm<½n§TI 2 rbs;RbUetGIun . 6-BiPakSaBIkMlaMgEdlrkSarcnasm<½n§TI 2 7-BiB’NnaBIrcnasm<½n§ nig muxgarrbs;RbUetGIunsés (fibrous proteins)

8-BiB’NnaBIrcnasm<½n§TI 3 nig TI 4 rbs;RbUetGIun 9-rab;BIGnþrGMeBI (interactions) EdlrkSaTMrg;RtwmRtUvrbs;RbUetGIun (protein conformation)

10-rab;]TahrN_én conjugated proteins nig prosthetic groups nig Bnül;BIviFIEdlBYkvabMeBjmuxgar 11-BiPakSaBIsar³sMxan;énrcnasm<½n§kñúglMhr (three-dimensional structure) rbs;RbUetGIuncMeBaHmuxgar rbs;va . 12-BiB’NnaBImuxgarrbs; hemoglobin nig myoglobin

13-BiB’NnafaetI pH nig sItuNðPaBxøaMg b¤ exSayb:uNÑaEdlbgá[mankarxUcxatRbUetGIun 14-Bnül;BIPaBxusKñarvagGasIutGamIneNcaM)ac; nig GasIutGamIneNmincaM)ac; .




KMeragemeron 1-muxgarRbUetGIunenAkñúgekasika (Cellular Function of Proteins)

2-GasIutGamIneNGal;hVa (The α-Amino Acid)

3-sm<½n§ PEPTIDES (The Peptide Bond)

4-rcnasm<½n§TI 1 rbs;RbUetGIun (The Primary Structure of Proteins)

5-rcnasm<½n§TI 2 rbs;RbUetGIun (The Secondary Structure of Proteins)

k-α-Helix

x-β-Pleated Sheet

6-rcnasm<½n§TI 3 rbs;RbUetGIun (The Tertiary Structure of Proteins)

7-rcnasm<½n§TI 4 rbs;RbUetGIun (The Quaternary Structure of Proteins)

8-TidæPaBTUeTAénrcnasm<½n§ nig muxgarRbUetGIun (An Overview of Protein Structure and Function)

9-MYOGLOBIN nig HEMOGLOBIN

k-Myoglobin nig karsþúkTukGuksIuEsn x-Hemoglobin nig kardwknaMGuksIuEsn K-kardwknaMGuksIuEsnBImþayeTA[TarkkñúgépÞ X-Sickle Cell Anemia

10-kareFIV[xUcxaténRbUetGIun 11-RbUetGIunkñúgrbbGahar nig karrMlayRbUetGIun




enAkñúgTsSsvtSqñaM 1800s Mulder )annwkeXIjeQμaH RbUetGIun BaküenHmanRsLayBIBaküRkic Edlmann½yfa {sar³sMxan;TI 1} . karBit RbUetGIunKWCacMNat;fñak;énm:UelKulGaharsMxan;xøaMgNas;GaRs½y edayBYkvapþl;[srIragÁminRtwmEtkae)an nig GIunRdUEsn b:uEnþ GasUt nig s<an;F½rpgEdr . FatuTaMgBIxageRkay enHminGacmanenAkñúgxøaj; nig carbohydrates eT EdlCafñak;énm:UelKulGaharsMxan;epSgeTot . epSgBIsar³sMxan;énrbbGahar RbUetGIunKWCa macromolecules EdlsMbUrCageKbMputenAkñúgekasika nig BYkvaGnuvtþPaKeRcInénkargarenAkñúgekasika . karkarBarénsBa’gÁkayBICMgWqøg (infection) karKaMRT nig kMlaMgemkanik nig karsMrYlénRbtikmμemtabUlIs TaMgGs;enHKWCamuxgarrbs;RbUetGIunEdlsMxan;sMrab;CIvit . 1-muxgarRbUetGIunenAkñúgekasika (Cellular Function of Proteins)

RbUetGIunmanmuxgarCIvsa®sþCaeRcIn dUcEdl)anbgðajedaysegçbxageRkamenH ³ Gg;sIum (Enzymes) katalIkrCIvsa®sþ KWCaRbUetGIun . Rbtikm μEdlRtUvkarry³eBleRcInéf¶ b¤ eRcIn GaTitü b¤ RtUvkarsItuNðPaBx<s;edayminmanGg;sIumRtUv)anbBa©b;kñúgry³Pøam² (an instant) . edayminman Gg;sIumKYr[cab;Garm μN_TaMgenH CIvitnwgminGacmaneLIy . GgÁbdibkç (Antibodies) k¾GacehAEdrfa Immunoglobulins KWCam:UelKulRbUetGIunBiess (specific protein molecules) EdlRtUv)anplitedayekasikaÉkeTsénRbB½n§PaBsaMu (specialized cell of

immune system) kñúgkareqøIytbeTAnwgsarFatumkBIeRkA (foreign) KW Antigen . GñkQøanBanBIeRkA (foreign invaders) rYmman)ak;etrI nig virus EdlbgáeraKdl;sBa’gÁkay . GgÁbdibkçnimYy²mantMbn;EdlRtUv Kñay:agxøaMg nig cgP¢ab;eTAnwg Antigen EtmYyKt; . vaCYybBa©b;karbgáeraKedaykarcgP¢ab;eTAnwg antigen nig CYybMpøaj antigen b¤ dk antigen ecjBIsBa’gÁkay . RbUetGIundwknaM (Transport Proteins) dwknaMsarFatu (materials) BITIkEnøgmYyeTAkEnøgmYyeTot enAkñúgsBa’gÁkay . Transferrine dwknaMCatiEdl (Fe) BIeføImeTAxYrq¥wg EdlenATIenaHEdkRtUv)aneRbIedIm,I sMeyaKCa heme group sMrab; hemoglobin . Hemoglobin nig myoglobin TTYlxusRtUvsMrab;kardwknaM nig sþúkTuknUvGuksIuEsnenAkñúgsrIragÁEdlx<s;Cag . RbUetGIunEktMrUv (Regulatory Proteins) RtYtBinitüTidæPaBCaeRcInénmuxgarekasika rYmman emtabU-lIs nig karbnþBUC . eyIgGacbMeBjmuxgarEtenAkñúglkçxNÐmYyRkumEdlmanRBMEdnkMNt; (a limited set of

conditions) . cMeBaHCIvitEdlenArs; sItuNðPaBsBa’gÁkay pH rbs;Qam nig kMrit glucose enAkñúgQamRtUv Et)anEktMrUvedayRbugRby½tñ . G½rm:Un (hormones) CaeRcInEdlEktMrUvmuxgarsBa’gÁkay dUcCa Insulin nig Glucagon KWCaRbUetGIun . RbUetGIunrcnasm<½n§ (Structural proteins) pþl;nUvkarKaMRTemkanik (mechanical support) cMeBaH stVFM² nig pþl;[BYkvarYmCamYynwgsMbkxageRkA (outer coverings) . sk; nig RkckédekIteLIgBIRbUetGIun



eQμaH Keratin . RbUetGIunepSgeTotpþl;nUvkMlaMgemkanik (mechanical strength) cMeBaHq¥wg sés (tendon)

nig Es,krbs;eyIg . BMumankarKaMRTrebobhñwg srIragÁEdlmanmaDFMdUcCarUbeyIgminGacmaneT . RbUetGIunEdlmanclna (Movement Proteins) caM)ac;cMeBaHRKb;TMrg;énclna . sac;dMurbs;eyIg rYmTaMgsac;dMusMxan;CageKbMput sac;dMuebHdUg sac;dMuk®nþak; nig sac;dMuBnøatamry³Gnþr³GMeBIrbs;RbUetGIun Actin

nig Myosin . Smerm GacEhl)anGaRs½yedayman flagella EvgEdlekIteLIgBIRbUetGIun . RbUetGIunbMrugCaGahar (Nutrient Proteins) eRbICaRbPBrbs;GasIutGamIneNsMrab;Tarkkñúgs,Ún (embryos) b¤ kumar . Albumin énEpñksrbs;s‘ut nig Casein enAkñúgTwkedaHKWCa]TahrN_énRbUetGIunbMrug CaGaharEdlsþúkTuk . 2-GasIutGamIneNGal;hVa (The α-Amino Acid)

RbUetGIunénsBa’gÁkayRtUv)aneLIgBIkarrYmKñaén subunits xus²KñacMnYn 20 EdleKehAfaCa GasIut GamIneNGal;hVa (α-amino acid) rcnasm<½n§TUeTAénGasIutGamIneNGal;hVaRtUv)anbgðaenAkñúgrUbTI 19-1. eyIgeXIjfa 19 kñúgcMeNamGasIutGamIneNTaMg 20 EdlRtUv)andkecjedayjwkjab;BIRbUetGIunmanrcnasm<½n§ TUeTAdUcKña . vamanbgÁúMGamInTI 1 (primary amine) enAelIkae)anGal;hVa (α-C) nig GasIutGamIneNEdlenA sl; Proline KWCaGamInTI 2 (secondary amine) .

rUbTI 19-1 H │ (α-amino group) H3N

+─C─C─O- (α-Carboxylate group) │ ║ R O (Side-chain R group)

kae)anGal;hVa (α-C) enAkñúgrUbmnþTUeTARtUv)anP¢ab;eTAnwg Carboxylate group ¬Carboxylate

group Edl)at;RbUtugmYy ³ ─COO-¦nig bgÁúMGamInEdlelIsRbUtugmYy (─NH3+) . enAkñúgsUluysüúgTwk

(aqueous solution) Edlman pH ≈ 7 EdlCalkçxNÐtMrUvsMrab;CIvitbMeBjmuxgar GasIutGamIneNEdlbgÁúM carboxylate cab;ykRbUtugmYy (─COOH) nig bgÁúMGamIne)aHbg;ecalRbUtugmYy (─NH2) minGacekItman )aneT . enAkñúglkçxNÐenH bgÁúM Carboxyl EbkCaGIuy:ug nig bgÁúMGamInEdlCa)ascab;ykRbUtugEdlRtUv)an beBa©jenaH . CalT§pl GasIutGamIneNenAkñúgTwksßitenACaGIuy:ugmanb:UlBIr (dipolar ions) EdleKehAfaCa zwitterions . kae)anGal;hVaRtUv)anP¢ab;eTAnwgbgÁúMnaTIbYnxusKñaenAkñúgRKb;GasIutGamIneNelIkElgEt glycine . ehtudUecñH kae)anGal;hVarbs;GasIutGamIneNGal;hVaKWCa chiral carbon . dUecñHGasIutGamIneNGal;hVa Edl EjkecjBIRbUetGIunmYyminGacRtYtsIuKñaenAelIrUbPaBqøúHkBa©k;eT . Glycine manGatUmGIuRdUEsnBIrP¢ab;eTAnwg GatUmkae)anGal;hVa nig KWCaGasIutGamIneNEtmYyKt;EdlCYbjwkjab;enAkñúgRbUetGIunEdlminman chiral

carbin (asymetry carbon) .

α



TMrg;rbs;GasIutGamIneNEdlEjkecjBIRbUetGIunmYyKWCaTMrg; L- . karenHsMGageTAelIkareRbóbeFob GasIutGamIneNCamYynwg D-glyceraldehyde ¬rUbTI 19-2¦ . TMrgGasIutGamIneNEdlEjkecjBIRbUetGIun KWpÞúyeTAnwgTMrg;rbs; D-glyceraldehyde dUecñH karrkTisénbgÁúMnaTITaMgbYnenACMuvij L-alanine dUceTAnwg karrkTisrbs;bgÁúMCMnYsTaMgbYn (substituents) enACMujvi chiral carbon rbs; L-glyceraldehyde .

rUbTI 19-2 CHO CHO │ │ HO─ C─H H─C─OH │ │ CH2OH CH2OH L-glyceraldehyde D-glyceraldehyde Carbohydrate pair COO- COO- │ │ H3N

+─C─H H─C─+NH3 Amino acid pair │ │ CH3 CH3 L-alanine D-alanine

GaRs½yedayGasIutGamIneNmanbgÁúM Carboxyl nig bgÁúMGamIn RKb;PaBxusKñarvagGasIutGamIneN GaRs½yeTAnwgbgÁúM R rbs;va . GasIutmIneNRtUv)ancat;CaRkumenAkñúgrUbTI 19-3 GnuelameTAtamPaBmanb:Ul (polarity) rbs;bgÁúM R . ExSenAxagcug (R) rbs;GasIutGamIneNmYycMnYnminmanb:UleT . BYkvacUlcitþb:HCamYynwgm:UelKulmYy epSgeTotminRCabTwk nig RtUv)aneKehAfaCa hydrophobic amino acids . BYkvaCaFmμtaRtUv)aneKeXIjman kb;enAEpñkxagkñúgrbs;RbUetGIunEdlenATIenaHBYkvaGacrYmCamYyGasIutGamIneNmYyepSgeTot nig enAEtenAdac; BITwkdEdl . manGasIutGamIneN 9 EdlRtUvsßitenAkñúgcMNat;fñak;enH ³ alanine, valine, leucine,

isoleucine, proline, glycine, methionine, phenylalanine nig tryoptophane . bgÁúM R rbs; proline

manEtmYyKt; CaFmμtavaRtUvcgsm<½n§eTAnwg α-amino group edaybegáItCaGamInTI 2 . ExSenAxagcug (R) rbs;GasIutGamIneNEdlenAsl;man;b:Ul . GaRs½yedayBYkvaRtUv)ancgP¢ab;CamYy nwgm:UelKulTwkEdlmanb:Ul BYkvaRtUv)aneKehAfaCa hydrophil amino acids . ExSxagEdlcUlcitþTwk (hydrophil) Cajwkjab;eXIjenAelIépÞrbs;RbUetGIun . GasIutGamIneNEdlmanb:UGacRtUv)aneKEckecj CabIfñak; ³ GasIuGamIneNNWtmanb:Ul (Polar, neutrai amino acids) manbgÁúM R EdlmancMNUlKImIx<s; (affinity) cMeBaHTwk b:uEnþminbMEbkCaGIuy:ugenAkñúgTwkEdlman pH = 7 eT . Serine, threonine, tyrosine,

cysteine, asparagine nig glutamine sßitenAkñúgGasIutGamIneNRbePTenH . PaKeRcInénGasIutGamIneN TaMgenHrYmCamYynwgGasIutGamIneNmYyeTotedaysm<½n§GIuRdUEsn b:uEnþ m:UelKUl cysteine begáIt)anCasm<½n§ disulfide CamYynwg cysteine dUcEdl)anBiPakSaenAkñúgEpñkTI 6 .



GasIutGamIneNmanbnÞúkGviC¢man (Negatively charged amino acids) manbgÁúM carboxyl EbkCa GIuy:ugenAxagcugExS. enAkñúg pH =7 GasIutGamIneNTaMgenHmanbnÞúksrubesμI -1 . Aspartate nig glutamate

sßitenAkñúgenAkñúgGasIutGamIneNRbePTenH . BYkvaCaGasIutGamIneNGasIut BIeRBaHkarbMEbkCaGIuy:ugrbs; carboxylic acid beBa©jRbUtugmYy . GasIutGamIneNmanbnÞúkviC¢man (Positively charges amino acids) enAkñúg pH = 7 lysine,

arginine nig histidine manbnÞúksrubviC¢manGaRs½yedaycugExSmanbgÁúMviC¢man . bgÁúMGamInTaMgenHKWCa)as GaRs½yeday cugExSmanRbtikmμCamYynwgTwkedaycab;ykRbUtugmYy nig beBa©jnUv hydroxide anions

(OH-) . rUbTI 19-3

(a) (b) Hydrophorbic amino acids H H H H │ │ │ │ H3N

+─C─COO- H3N+─C─COO- H3N

+─C─COO- H3N+─C─COO-

│ │ │ │ R H CH3 HC─CH3 │ CH3 Glycine (Gly) Alanine (Ala) Valine (Val) H H H H │ │ │ │ H3N



│ │ │ │ │ CH2 H─C─CH3 CH2 H2C CH2 │ │ │ CH2 HC─CH3 CH2 │ │ CH3 CH3 Leucine (Leu) Isoleucine (Ile) Phenylalanine (Phe) Proline (Pro) H H │ │ H3N


│ │ CH2 CH2

│ │ N CH S │ │ H CH2 Tryptophane (Trp) Methionine (Met)



(c) Polar, neutral amino acids H H H H │ │ │ │ H3N



│ │ │ │ H─C─OH H─C─OH CH2 CH2 │ │ │ │ H CH3 SH │ OH Serine (Ser) Threonine (Thr) Tyrosine (Tyr) Cysteine (Cys) H H │ │ H3N


│ │ CH2 CH2 │ │ C─NH2 CH2 ║ │ O C─NH2 ║ O Asparagine (Asn) Glutamine (Gln) (d) Negatively charged amino acids (e) Postively charged amino acids H H H H │ │ │ │ H3N



│ │ │ │ CH2 CH2 CH2 CH2 │ │ │ │ C─O- CH2 CH2 CH2 ║ │ │ │ O C─O- CH2 CH2 ║ │ │ O CH2 N─H │ │ NH2 C═NH2

+ │ NH2 Aspartate (Asp) Glutamate (Glu) Lysine (Lys) Arginine (Arg)



H │ H3N

+─C─COO-

│ CH2──CH │ │ HN+ NH C │ H Histidine (His) H │ H3N

+─ C ─COO- │ │ H2C CH2 CH2 Proline (Pro)

eQμaHGasIutGamIneNRtUv)ansresredaysegçbedaykUdbIGkSr . GkSrkat;enHRtUv)anbgðajenAkñúg taragTI 19-1 .

taragTI 19-1 Amino acid Abbreviation Alanine ala Arginine arg Asparasine asn Aspartate asp Cysteine cys Glutamine gln Glutamic acid glu Glycine gly Histidine his Isoleucine ile Leucine leu Lysine lys Methionine met Phenylalanine phe Proline pro Serine ser Threonine thr Tryptophane trp Tyrosine tyr Valine val



3-sm<½n§ PEPTIDES (The Peptide Bond)

RbUetGIunKWCa linear polymers rbs; L-α-amino acids . bgÁúM carboxyl rbs;GasIutGamIneNmYy RtUv)anP¢ab;CamYynwgbgÁúMGamInrbs;GasIutGamIneNmYyepSgeTot . sm<½n§GamIdRtUv)anekIteLIgenAkñúgRbtikmμ RtUv)aneKehAfaCa sm<½n§ Peptide ¬rUbTI 19-4¦ H O H │ ║ │ H3N

+─C─C─N─C─C─O- │ │ │ ║ R H R1 O

m:UelKulEdlekIteLIgBIkarrYmKñaBIGasIutGamIneNBIrRtUv)aneKehAfa dipeptide . GasIutGamIneN EdlmanbgÁúM α-NH3

+ esrIRtUv)aneKsÁal;faCa amino terminal b¤edaysamBaØ N-terminal amino acid

nig GasIutGamIneNEdlmanbgÁúM ─COO- esrIRtUv)aneKsÁal;faCa Carboxyl b¤ C-terminal amino acid . rcnasm<½n§rbs;RbUetGIunRtUv)aneKsresredaysn μtiedaydak; N-terminal amino acid enAxageqVg .

rUbTI 19-4

H O H O │ ║ │ ║ H3N

+─C─C─O + H3N+─C─C─O-

│ │ (a) H CH3

H O H O │ ║ │ ║ H3N

+─C─C──N─C─C─O- + H2O │ │ │ H H CH3 Peptide bond H O H O H O H O H │ ║ │ ║ │ ║ │ ║ │ H3N

+─C─C─N─C─C─N─C─C ─N─C─C ─N─C─COO- │ │ │ │ │ │ │ │ │ H H Rx H Ry H Rz H CH3 Peptide bonds Amino-terminal Carboxyl-terminal amino acid or amino acid or N-terminal C-terminal amino acid amino acid (b)



cMnYnGasIutGamIneNenAkñúg peptides tUc²RtUv)anbBa¢ak;edaybuBVbT di ¬manBIrGasIuGamIneN¦ tri

¬manbIGasIutGamIneN¦ tetra ¬manbYnGasIutGamIneN¦ nig bnþeTAeTot . Peptides RtUv)aneK[eQμaH edayyktam C-terminal amino acid EdlTTYlykeQμaHvaTaMgmUlEtmþg . cMeBaHRKb;GasIutGamIneN epSgeTotcugbBa©b; “-ine” RtUv)anpøas;bþÚreTCa “-yl” . ehtudUecñH dipeptide alanyl-glycine man glycine

Ca C-terminal amino acid EdlRtUv)anbgðajnUveQμaHeBjrbs;vaKW glycine . H O H O │ ║ │ ║ H3N

+─C─C──N─C─C─O- Alanyl-glycine (ala-gly) │ │ │ CH3 H H3

Dipeptide ekIteLIgBI alanine nig glycine Edlman alanine Ca C-terminal amino acid vaKWCa glycyl-alanine : H O H O │ ║ │ ║ H3N

+─C─C──N─C─C─O- Glycyl-alanine (gly-ala) │ │ │ H H CH3

Dipeptides TaMgBIrmansmasPaBGasIutGamIneNBIrdUcKña b:uEnþmanQutGasIutGamIneNxusKña . rcnasm<½n§rbs; peptides tUc²GacRtUv)aneKKUsedayRsYlkñúgkarGnuvtþ RbsinebIviFanBitR)akdmYy RtUv)aneKeFIVtam . dMbUg cUrkt;sMKal;fa ExSem (backbone) rbs; peptide manQutekIteLIgdEdl² . N─C─C─N─C─C─N─C─C 1 2 1 2 1 2

Edl N KWCa α-amino group , C-1 KWCa α-carbon nig C-2 KWCa Carboxyl group . C-1 CaFmμtaRtUv)an cgsm<½n§CamYynwgGatUUmGIuRdUEsneTAnwgcugExS R group EdlvadUcKñaeTAtamGasIutGamIneNnimYy² . bnþkarKUr dUcEdl)anKUbBa¢ak;enAkñúg]TahrN_TI 19-1 .



]TahrN_TI 19-1 karsresrrcnasm<½n§rbs; Tripeptide ³ cUrKUrcnasm<½n§rbs; Tripeptide alanyl-glycyl-valine

H O H O H O │ ║ │ ║ │ ║ H3N

+─C─C──N─C─C──N─C─C─O- │ │ │ │ │ CH3 H H H HC─CH3 │ CH3

eTaHbICaGñkGacKitBIkarviledayesrIénsm<½n§ peptide k¾eday b:uEnþkarenHminCakrNIEdlRtUvKiteT . GaRs½yedayeTVtaeGLikRtugEtmYyrbs;GatUmGasUtmanGMeBICamYynwgkae)an nig GuksIuEsnrbs; carbonyl

group m:UelKulbgðajBI GnurvPaB (resonance) . karenHpþl;[sm<½n§ peptide nUvlkçN³sm<½n§BIrCan;eday Epñk (partially double bond character) ³ ─Cα H ─Cα H C─N C═N+ O Cα─ O Cα─

CalT§pl sm<½n§ peptide enAelIbøg;rabes μ I nig bgÁúM R RtUv)ancgP¢ab;eTAnwg α-C enACitKñasßitenA RbQmKñaénExSRbUetGIun ¬rUbTI 19-5¦ . GIuRdUEsnrbs;GasUténbgÁúMGamIdk¾sßitenAkñúgTItaMg trans CamYynwg O

rbs;bgÁúM Carbonyl . esÞIrEtRKb;sm<½n§ peptides enAkñúgRbUetGIunsßitenAelbøg;rabes μ I nig manTMrg;Ca trans

(trans configuration) . karenHmansar³sMxan;Nas;enAkñúgsrIr³viTüaxøaMgNas; GaRs½yedayvaeFIV[rcna sm<½n§RbUetGIunmanPaBrwgmaMedayminR)akd (relatively rigid) . RbsinebIminmanrUbragrbs;vamYyR)akdeT BYkvaminGacbMeBjmuxgarrbs;BYkva)aneT .

rUbTI 19-5



4-rcnasm<½n§TI 1 rbs;RbUetGIun (The Primary Structure of Proteins)

rcnasm<½n§TI 1 (the primary structure) rbs;RbUetGIunKWCaQutrbs;GasIutGamIneNénExSRbUetGIun. va)anmkBIsm<½n§kUva:Lg;rvagGasIutGamIneNenAkñúgExS ¬sm<½n§ peptides¦ . rcnasm<½n§TI 1 rbs;RbUetGIun KWCakarbkERbnUvB½t’manEdlmanenAkñúgEhSn (gene) . RbUetGIunnimYy²man rcnasm<½n§TI 1 edaymanGasIut GamIneNxus²KñakñúgTItaMgxus²KñaenAtambeNþayExSRbUetGIun . EhSnGacpøas;bþÚredaydMeNIrkarénkarERbRbYl (mutation) kñúgry³eBlénkarvivtþn_ . karERbRbYlenA kñúgEhSnGacbNþal[mankarpøas;bþÚrQutCabzménGasIutGamIneNrbs;RbUetGIunmYy . kñúgry³eBlyUr kar- pøas;bþÚrrwtEteRcInnwgekIteLIg . RbsinebI BUCBIrénsrIragÁbgáat;Kña ¬edIm,IbegáItCaBUCfμ I¦ kñúgeBlPøam² PaBxusKña enAkñúgQutGasIutGamIneNénRbUetGIunnwgmantictYcNas; . mü:agvijeTot RbsinebIBYkvaRtUv)anbgáat;rab;lan qñaMknøgeTA vamanPaBxusKñay:agxøaMgenAkñúgQutGasIutGamIneNrbs;RbUetGIunBYkeK . CalT§pl eyIgGac eRbóbeFobPaBTMnak;TMngénkarvivtþn_rvagBUCepSg²edayeFIVkareRbóbeFob rcnasm<½n§TI 1 rbs;RbUetGIun Edlman enAkñúgBUCTaMgBIenH . 5-rcnasm<½n§TI 2 rbs;RbUetGIun (The Secondary Structure of Proteins)

QutTImYyrbs;RbUetGIun ¬ExSénGasIutGamIneNEdlcgP¢ab;Kñaedaysm<½n§kUv:aLg;¦ bt;cUlKñaCarcna sm<½n§sarcuHsaeLIgedayeTogTat;Edlhak;dUcCarUb)a:kenAelIkMral . rcnasm<½n§EdlekIteLIgdEdl²enHkNt; )anCa rcnasm<½n§TI 2 rbs;RbUetGIun . rcnasm<½n§TI 2 rbs;RbUetGIunKWCalT§plénkarcgsm<½n§GIuRdUEsn rvag GIuRdUEsnénbgÁúMGamId CamYynwg GuksIuEsnénbgÁúM carbonyl énsm<½n§ peptide . mansm<½n§GIuRdUEsnCaeRcInRtUv karedIm,IFana rcnasm<½n§TI 2 rbs;RbUetGIun nig EdlehtuenHKWCarcnasm<½n§TaMmUlrbs;RbUetGIun . tMbn;xusKñaén ExSRbUetGIunGacmanRbePTxusKñaén rcnasm<½n§TI 2 . tMbn;mYycMnYnénExSRbUetGIunrcnasm<½n§edayécdnü b¤ edaymineTogTat; b:uEnþ RbePTCYbjwkjab;CageKbMputBIrén rcnasm<½n§TI 2 KW α-helix nig β-pleated sheet

GaRs½yedayBYkvaeFIV[manGtib,rmanUvsm<½n§GIunRdUEsnenAkñúgExSem (backbone) . k-α-Helix

RbePTCYbjwkjab;CageKbMputén rcnasm<½n§TI 2 rbs;RbUetGIunKWCaTMrgrmUdUcsøabc½Rk (coiled, helical

conformation) EdleKsÁal;faCa α-helix ¬rUbTI 19-6¦ . α-helix manTidæPaBsMxan;CaeRcIn . ral;GIuRdUEsnénbgÁúMGamId nig GuksIuEsnénbgÁúM carbonyl rYmCamYynwgExSem peptide Cab;Tak;TgeTA nwgsm<½n§GIuRdUEsnenAeBlExSenHrmUrCa α-helix . sm<½n§GIuRdUEsnTaMgenHeFIV[ α-helix enAnwgkñúgkEnøg . GuksIuEsnénbgÁúM carbonyl RtUv)ancgP¢ab;CamYynwgGIuRdUEsneTAnwgGIuRdUEsnénbgÁúMGamIdénGasIut GamIneNbYnEdlenAq¶ayBIExSenH . sm<½n§GIuRdUEsnrbs; α-helix RsbeTAnwgbeNþayGkS½én helix ¬emIlrUbTI 19-6¦ . ExS polypeptide enAkñúg α-helix KWrmYleTAxagsþaMéd (right-handed) . vamanTisedAdUceTAnwg ex©AFmμta . RbsinebIGñkmYlex©ARsbeTAnwgdMeNIrRTniknaLikavanwgcUleTAkñúgCBa¢aMg ehIyRbsinebImYlbRBa©as; TisRTniknaLikedIr ex©AnwgecjBICBa¢aMg .



RbEvgsareLIgvijén helix b¤ mYyrgVg;rbs;va (pitch) KW 5.4 Ǻ nigman 3.6 GasIutGamIneNkñúg mYyCMu helix .

rUbTI 19-6

RbUetGIunsés (Fibrous proteins) KWCaRbUetGIunEdlrcnasm<½n§RtUv)anerobcMCasés b¤ bnÞHEdlman RbePTEtmYyénrcnasm<½n§TI 2 . α-keratins KWCaRbUetGIunsésEdlbegáItnUveRKÓgsMrab;RKb (covering) ¬dUc Cask; eramecom (wool)Rkckéd (nails) RkckesH (hooves) nig eramstV (fur)¦ énstVEdlrs;enAelIdIeRcIn bMput . sk;mnusSpþl;Ca]TahrN_KMrUmYyénrcnasm<½n§rbs; α-keratins . RbUetGIunénsk;manesÞIrEtTaMgGs;Ca ExS polypeptide EdlrmUCa α-helices . ExS α-helix EtmYyrmUCadMuCamYynwg helices BIrepSgeTotedIm,Ipþl; Ca three-stranded supper-structure EdleKehAfa Protofibril EdlKWCaEpñkmYyénlMdab;CYrEdlRtUv)an eKsÁal;faCa microfibril emIlrUbTI 19-7 . rcnasm<½n§TaMgenH EdldUceTAnwg {knÞúyRCUk} (molecular

pigtails) mankMlaMgemkanikFM nig BYkvaminrlayenAkñúgTwkTal;EtesaH . RbUetGIunsésrbs;sac;dMuekItLIgBIRbUetGIunEdlmancMnYnd¾eRcInén α-helices . Ca]TahrN_ Myosin

RbUetGIunsMxan;mYykñúgcMeNamRbUetGIunrbs;sac;dMumanrcnasm<½n§dUcrMBat; (rodlike) Edl α-helices BIrbegáIt)an CaExSrmUrmYy ¬rUbTI 19-8¦ .



rUbTI 19-7

rUbTI 19-8



lkçN³sMxan;énrcnam<½n§rbs; coiled coil supperstructure rbs; α-helices KWCakMlaMgemkanikFM . lkçN³enHRtUv)anGnuvtþRbkbedayRbsiT§iPaBTaMgenAkñúgRbUetGIunsésrbs;Es,k nig RbUetGIunsésrbs;sac;dMu. dUcEdlGñkGacRsm½yeXIj RbUetGIunTaMgenHRtUvEtmankMlaMgxøaMgedImGnuvtþkargarkñúgkarKaMRTemkanikrbs; BYk va nig kark®nþak;sac;dMu . x-β-Pleated Sheet

rcnasm<½n§TI 2 rbs;RbUetGIunEdlCYbjwkjab;TI 2 manragdUceTAnwgsñampñt;énvaMgnn nig RtUv)aneKsÁal; faCa β-pleate sheet rUbTI 19-9 . RKb;GuksIuEsnénbgÁúM carbonyl nig GIuRdUEsnénbgÁúMGamId enAkñúg β-

pleated sheet Cab;Tak;TgeTAnwgsm<½n§ nig ExS polypeptides esÞIrEtrlaTaMgRsug . ExS polypeptide enAkñúg β-pleated sheet GacmankMNt;TisedA (orientations) BIr . RbsinebI N-termini KWCak,alExSeTAk,alExS (head to head) rcnasm<½n§enHRtUv)aneKsÁal;faCa Parallel β-pleated sheet . RbsinebI N-terminus

rbs;ExSmYyRtUv)anerobCaCYrCamYynwg C-terminus rbs;ExSTI 2 ¬k,alExSeTAknÞúyExS¦ rcnasm<½n§enHRtUv)an eKsÁal;faCa Antiparallel β-pleated sheet .

rUbTI 19-9



RbUetGIunsésmYycMnYnRtUv)anekIteLIgBI β-pleated sheet . ]TahrN_ dgáÚvnag (silk-worm)

begáItnUv fibroinénséssURt (silk fibroin) ³ CaRbUetGIunmYyEdlrcnasm<½n§rbs;vaKWCa antiparallel β-

pleated sheet ¬rUbTI 19-10¦ . ExS polypeptide én β-pleated sheet esÞIrEtrlaTaMgRsug nig sURtmin gaysn§wgedayRsYleT . Glycine cUlrYmesÞIrEtBak;kNþalénGasIuGamIneNrbs; fibroin rbs;séssURt . Alanine nig serine cUlrYmeRcInbMputcMeBaRbUetGIunsésepSgeTot . bgÁúMemTIlrbs; alanines nig bgÁúM hydroxymethyl rbs; serines sßitenARbQmKñaén sheet . ehtudUecñH stacked sheet Cab;KñaedayRsYl dUc sheets rbs;Rkdas;katugEdlbt;GaRs½yedaybgÁúM R tUcl μmGnuBaØati[man stacked sheet supper-

structure. rUbTI 19-10



6-rcnasm<½n§TI 3 rbs;RbUetGIun (The Tertiary Structure of Proteins)

PaKeRcInénRbUetGIunsés dUcCasURt collagen nig α-keratins esÞIrEtminrlaykñúgTwkTaMgRsug . ¬Es,krbs;eyIgeFIV[eyIgminRsYlxøaMg RbsinebIvarlayenAkñúgTwkePøóg¦. b:uEnþ PaKeRcInénRbUetGIunrbs; ekasika rlayenAkñúgsIutU)aøs;rbs;ekasika . RbUetGIunrlayCaFmμtaKWCa globular proteins . Globular

proteins manrcnasm<½n§ 3-D (three-dimensional structures) EdlRtUv)aneKehAfaCa rcnasm<½n§TI 3 rbs; RbUetGIun (tertiary structure of protein) EdlxusBIrcnasm<½n§TI 2 . ExS polypeptide EdlmaneTAeday tMbn;énrcnasm<½n§TI 2 rbs;va α-helix nig β-pleated bt;eRcInpñt;EfmeTotelIxøÜnÉgedIm,ITTYl)anCarcnasm<½n§ TI 3 . eyIg)aneXIjfakMlaMEdlrkSarcnasm<½n§TI 2 rbs;RbUetGIunKWCasm<½n§GIuRdUEsnrvagGIuRdUEsnénbgÁúMGamId nig GuksIuEsnénbgÁúM carbonyl énsm<½n§ peptide . etIkMlaMgGIVEdlFanarcnasm<½n§TI 3 rbs;RbUetGIun ? Globular tertiary structure ekIteLIgedayÉkÉg nig RtUv)anrkSa EdlCalT§plénGnþrGMeBIkñúgcMeNamEpñk xagénExS ¬bgÁúM R¦rbs;GasIutGamIneN . rcnasm<½n§RtUv)anFanaedayGnþrGMeBIénm:UelKuldUcxageRkam ³ Van der Waals forces rvagbgÁúM R rbs;GasIutGamIneNminmanb:UlEdlKWCa Hydrophorbe . sm<½n§GIuRdUEsnrvagbgÁúM R Edlmanb:Ulrbs;GasIutGamIneNEdlmanb:Ul . sm<½n§GIuy:UnikrvagbgÁúM R rbs;GasIutGamIneNEdlmanbnÞúkpÞúyKña . sm<½n§kUva:Lg;rvagGasIutGamIneNEdlbgÁúM Thiol . Cysteines BIrGacRtUv)anTTYlGuksIutkmμeTACa dimer amino acid RtUv)aneKehAfaCa Cystine ¬rUbTI 19-11¦ . sm<½n§ disulfide rbs; cystine GacRtUv )anExVgKñarvagRbUetGIunxus²Kña b¤ GaccgEpñkBIrcUleTAkñúgRbUetGIunmYyrYmKña . sm<½n§EdlFanarcnasm<½n§TI 3rbs;RbUetGIunRtUv)anbgðajenAkñúgrUbTI 19-12 . sar³sMxan;én sm<½n§TaMgenHkøayeTACac,as;enAeBleyIgdwgfavaKWCarcnasm<½n§TI 3rbs;RbUetGIunEdlkMNt;BImuxgarCIvsa®sþ rbs;va . PaKeRcInbMput GasIutGamIneNminmanb:UlRtUv)ankb;)at; RtUv)anx©b;edayCit enAkñúgEpñkxagkñúg globular protein minb:HCamYynwgTwk . GasIutGamIneNEdlmanb:Ul nig bnÞúksßitenAelIépÞén globular

proteins . Globular proteins ENnxøaMgNas; . rcnasm<½n§TI 3 GacmantMbn; α-helix nig β-pleated

sheet . tMbn; {Rtecok} (hinge region) énrmUrExSP¢ab;tMbn;én α-helix nig β-pleated sheet . GaRs½y edayrcnasm<½n§ExSbiTrbs;va Proline bg¥ak; α-helix . CalT§pl proline Cajwkjab;RtUv)aneKeXIjmanenA kñúgtMbn; {Rtecok} TaMgenH . cMnYnBitR)akdénRbePTnimYy²rbs;rcnasm<½n§TI 2 ERbRbYlBIRbUetGIunmYyeTA RbUetGIunbnÞab;eTot .



rUbTI 19-11

H H O H H O │ │ ║ │ │ ║ -----N─C─C----- ------N─C─C----- │ │ CH2 CH2 │ │ S─H S │ + 2H+ + 2e- S─H S │ │ CH2 CH2 │ │ ------N─C─C----- ------N─C─C----- │ │ ║ │ │ ║ H H O H H O

rUbTI 19-12



7-rcnasm<½n§TI 4 rbs;RbUetGIun (The Quaternary Structure of Proteins)

cMeBaHRbUetGIunCaeRcInEdlTMrg;bMeBjmuxgarminEmnekIteLIgBIExS peptide EtmYyKt;enaHeT b:uEnþvaekIt eLIgBIkarCab;Kñaén globular peptide tUc² . cMeBaHehtuplenH Hemoglobin ekIteLIgBI globular

peptide subunits bYn ³ identical α-subunits BIr nig identical β-subunits BIr . enAeBlEt subunits

TaMgbYnRtUv)ancgP¢ab;KñaBImYyeTAmYyepSgeTotKWCaRbUetGIunEdlGacbMeBjmuxgar . karrYmKñaén polypeptides

CaeRcInedIm,IbegáItCaRbUetGIunEdlmanmuxgarkMNt;)annUv rcnasm<½n§TI 4 rbs;RbUetGIun (quaternary structure

of a protein) . kMlaMgEdlcgrcnasm<½n§TI 4 rbs;.RbUetGIunKWdUceTAnwgkMlaMgEdlcgrcnasm<½n§TI 3 Edr . kMlaMgTaMg enHrYmmansm<½n§GIuRdUEsnrvagGasIutGamIneNmanb:Ul sm<½n§GIuy:UnikrvagGasIutGamIneNmanbnÞúkpÞúyKña kMlaMg Van der Waals rvagGasIutGamIneNminmanb:Ul nig s<an disulfides . enAkñúgrNImYycMnYnrcnasm<½n§TI 4 rbs;RbUetGIunbMeBjmuxgarCab;Tak;TgeTAnwgkarP¢ab;eTAnwgbgÁúMEdl minEmnCaRbUetGIun . bgÁúMEdlbEnßmenHRtUv)aneKehAfaCa bgÁúMsMrab;CMnYs (prosthestic group) nig RbUetGIun bMeBjmuxgarEdlmanbgÁúMsMrab;CMnYs (prosthestic group) RtUv)aneKehAfaCa Conjugated protein . Receptor proteins CaeRcInenAelIépÞekasikaKWCa Glycoproteins . Conjugated proteins TaMgenHrYmCa mYynwg sugar groups tamry³sm<½n§kUva:Lg; . Hemoglobin k¾Ca conjugated protein Edr . subunits

nimYy²RtUv)anP¢ab;eTAnwg Heme group EdlmanCatiEdk . Heme group KWCaGamInsrIragÁExSbiTminEq¥t nig FMrYmCamYynwgGIuy:ugEdkEdlsMrbsMrYlenAkñúgva ¬emIlrUbTI 19-15¦ . dUcenAkñúgkrNI hemoglobin bgÁúMsMrab; CMnYs (prosthetic group) Cajwkjab;kMNt;muxgarrbs;RbUetGIun . Ca]TahrN_ enAkñúg hemoglobin KWCa heme group EdlmanEdk EdlmanlT§PaBkñúgkarcgP¢ab;eTAnwgGuksIuEsn . 8-TidæPaBTUeTAénrcnasm<½n§ nig muxgarRbUetGIun (An Overview of Protein Structure and Function)

\LÚvcUrsegçbnUvRbePTepSg²rbs;rcnasm<½n§RbUetGIun nig TMnak;TMngrvagBYkvanimYy² ¬rUbTI 19-13¦ rUbTI 19-13



rcnasm<½n§TI 1 ³ rcnasm<½n§TImYyrbs;RbUetGIunKWCaQutGasIutGamIneNrbs;RbUetGIun . rcnasm<½n§TI 1 bNþalmkBIkarbegáItnUvsm<½n§ peptide kUva:Lg;rvagGasIutGamIneN . sm<½n§ peptides KWCasm<½n§GamIdEdl begáItrvag α-carboxylate group rbs;GasIutGamIneNmYy nig α-amino group rbs;GasIutGamIneNmYy epSgeTot . rcnasm<½n§TI 2 ³ enAeBlEdlExSRbUetGIunekIneLIg »kasCaeRcInénGnþrGMeBIEdlminEmnkUva:Lg; enA kñúgExSem (backbone) rbs;ExS polypeptide ekItmaneLIg . karenHbgá[ExSbt; nig kMNt;TiedAedayxøÜnÉg nUvkarerobcMCaTMrg;FM²CaeRcInRbePT . kMritTI 2 énrcnasm<½n§rYmman α-helix nig β-pleated sheet EdlKWCa lT§plénkarcgsm<½n§GIuRdUEsnrvagGIuRdUEsnénbgÁúMGamId nig GuksIuEsnénbgÁúM carbonyl rbs;sm<½n§ peptide . EpñkxusKñaénExSGacCab;Tak;TgenAkñúgRbePTxus²KñaénkarerobcMrcnasm<½n§TI 2 . tMbn;mYycMnYnGacCa α-helix

nig tMbn;epSgeTotGacCa β-pleated sheet . rcnasm<½n§TI 3 ³ enAeBleyIgBiPakSaBIrcnasm<½n§TI 3 eyIg)ancab;GarmμN_enAkñúgkarbt;TaMgGs;én ExSTaMgmUl . m:üagvijeTot eyIgmankgVl;CamYynwgkarbt;kan;EteRcIneToténrcnasm<½n§TI 2 . etIcugExSTaMgBIr eTACitKña b¤ enAdac;q¶ayBIKña ? etIrUbragTUeTAmYyNaEdlCab;Tak;Tg ? TaMgGnþrGMeBIminEmnkUva:Lg;énbgÁúM R

rbs;GasIutGamIneN nig GnþrGMeBIkUva:Lg;éns<an disulfide mantYnaTIkñúgkarkMNt;rcnasm<½n§TI 3 . GnþrGMeBImin EmnkUva:Lg;rYmman sm<½n§GIuRdUEsn sm<n§GIuy:Unik nig kMlaMg Van der Waals . rcnasm<½n§TI 4 ³ dUceTAnwgrcnasm<½n§TI 3 rcnasm<½n§TI 4 Cab;Tak;TgeTAnwgkarerobcMkñúglMhtam zan³ (topological, spatial arrangements) rbs;ExS peptide BIr b¤ eRcInedayeKarBKñaeTAvijeTAmk . etI ExS peptide mYykMNt;TisedAedayeKarBeTAtamExS peptide mYyepSgeTottamrebobdUcemþc ? etIGIVCa rUbragcugbBa©b;rbs;RbUetGIunbMeBjmuxgar ? rcnasm<½n§TI 4 RtUv)anrkSaedaysarkMlaMgdUcKñaEdlTTYlxusRtUvcMeBaHrcnasm<½n§TI 3 . vaKWCarcna sm<½n§TI 3 nig TI 4rbs;RbUetGIunEdlkMNt;edaycugeRkaybMputnUvmuxgarrbs;BYkva . mYycMnYnmanrcnasm<½n§Ca sésrYmCamYynwgkMlaMemkanikxøaMg (the great mechanical strength) . BYkvaTaMgenHbegáItsmasFaturcna-sm<½n§sMxan;rbs;ekasika nig srIragÁ . Cajwkjab;BYkvak¾TTYlxusRtUvEdrcMeBaHclnarbs;srIragÁ . mYycMnYneTot bt;pñt;eTATMrg;mUl (globular shapes) . PaKeRcInbMputénRbUetGIundwknaM RbUetGIunEktMrUv nig Gg;sIumKWCa globular proteins . rcnasm<½n§ 3-D EdleTogTat;xøaMgrbs;RbUetGIundwknaMGnuBaØati[BYkvaTTYlsÁal;nUv m:UelKulBiessNamYy nig sMrYlkarcUleTAkñúgekasika . dUcKñaenHEdr KWCarUbrag 3-D Biessrbs;RbUetGIun-EktMrUvEdlGnuBaØati[BYkvaP¢ab;eTAnwgGñkTTYl (receptor) BYkvaenAelIépÞekasikacMNuc . enAkñúgrebobenH BYk vaGacTMnak;TMngCamYynwgekasika edaykarbeRgón[ekasikaeFIVGMeBImYycMnYn . enAkñúgCMBUkTI 20 eyIgnwgBinitü emIlBIrcnasm<½n§ 3-D énTIkEnøskm μrbs;Gg;sIum (enzyme active sites) EdlGnuBaØati[BYkvaP¢ab;CamYynnwg RbtikrBiess (specific reactants) nig begáInel,ÓnRbtikmμCIvKImI . dUcEdleyIgnwg)aneXIjenAkñúg]TahrN_én sickle cell hemoglobin enAkñúgEpñkbnÞab; karxUcxatEt GasIutGamIneNEtmYyKt;enAkñúgrcnasm<½n§TI 1 rbs;RbUetGIunGacmankarCMBak;CMBiny:ageRcIn . enAeBlGasIut



GamIneNmYyCMnYsGasIutGamIneNmYyepSgeTotenAkñúgExS peptide mYy vamankarpøas;bþÚrnUvbgÁúM R rbs;RbUet-GIun . karenHbNþal[mankarbt[pñt;rcnasm<½n§TI 3 nig RbEhlrcnasm<½n§TI 4 GaRs½yedayRbePTén GnþrGMeBIminEmnkUva:Lg;RtUv)anxUcedaysarkarpøas;bþÚrbgÁúM R EdlmansMrab;karcgsm<½n§ . dUcKñaenHEdr karCMnYs GasIutGamIneNmYyepSgeTotCamYynwg Proline GacraraMgtMbn;sMxan;énrcnasm<½n§TI 2 . ehtudUecñH karpøas;bþÚr QutGasIutGamIneNenAkñúgrcnasm<½n§TI 1 Gacpøas;bþÚrnUvkarbt;pñt; 3-D rbs;RbUetGIunkñúgrebobEdlbgá[man[ vaminGacbMeBjmuxgar)an . enAkñúgrNI sickle cell hemoglobin karenHGacbNþal[søab; . 9-MYOGLOBIN nig HEMOGLOBIN

k-Myoglobin nig karsþúkTukGuksIuEsn PaKeRcInbMputénekasikarbs;sBa’gÁkayrbs;eyIgRtUv)ankb;enAkñúgsBa’gÁkay nig minGacTTYlGahar edaypÞal;m:UelKulGahar b¤ beBa©jkaksMNl; . RbB½n§Qamrt;edaHRsaynUvbBaðaenHedaykarbeBa©jnUvsarFatu bMrug nig GuksIuEsneTA[ekasikasBa’gÁkay nig dwkkaksMNl;beBa©jecal . ekasikaeyIgRtUvkarpþl;edayman esßrPaBnUvGuksIuEsn b:uEnþGuksIuEsnrlayticenAkñúgsUluysüúgTwk . edIm,ICMnHbBaðaénPaBrlayenH eyIgman RbUetGIunsMrab;dwknaMGuksIuEsnKW Hemoglobin . Hemoglobin RtUv)aneKeXIjmanenAkñúgekasikaRKab;Qam Rkhm nig KWCaRbUetGIundwknaMGuksIuEsnrbs;stVfñak;x<s; . Myoglobin KWCaRbUetGIunsþúkGuksIuEsnrbs;sac;dMu eRKagq¥wg (skeletal muscle) . rcnasm<½n§rbs; myoglobin (Mb) RtUv)anbgðajenAkñúgrUbTI 19-14. Heme group ¬rUbTI 19-15¦ k¾CasmasFatusMxan;rbs;RbUetGIunEdr . GIuy:ug Fe2+ enAkñúg heme group KWCakEnøgcgP¢ab; GuksIuEsnrbs; TaMg myoglobin nig hemoglobin . edaymanPBVvasna myoglobin mankark®nþak;ylGuksIuEsnxøaMgCag hemoglobin . ehtudUecñHvamankarbBa¢ÚnedayRbsiT§iPaBnUvGuksIuEsnBIcrnþQameTAekasikarbs;sBa’gÁkay . x-Hemoglobin nig kardwknaMGuksIuEsn Hemoglobin (Hb) KWCa Tetramer EdlekIteLIgBI polypeptide subunits bYn ³ BIr α-subunits

nig BIr β-subunits ¬rUbTI 19-16¦ . GaRs½yeday subunits nimYy²rbs; hemoglobin man heme group

m:UelKul hemoglobin Gaccg 4 m:UelKulGuksIuEsn ³ Hb + 4O2 → Hb(O2)4

karpþl;GuksIuEsn[ Hemoglobin enAkñúgsYt nig karbBa¢ÚnGuksIuEsnBI hemoglobin eTA[ myoglobin enAkñúgCalikaCadMeNIrkarsaMujauM . eyIgcab;epþImkarGegáténRBwtþkarN_TaMgenHrbs;eyIgCamYynwg kardkdegðImykxül; . karpþl;GuksIuEsn[ Hemoglobin CYysMrYly:ageRcInedaysarPaBxusKñarvag Oxygen partial

pressure (pO2) enAkñúgsYt nig enAkñúgQam . pO2 enAkñúgxül;kñúgsYtmansMBaFRbmaN 100 mg Hg ehIy pO2 enAkñúgQamEdl)at;bg;GuksIuEsn (oxygen-depleted blood) manRbEhl 40 mm Hg . GuksIuEsn sayPayBItMbn;Edlman pO2 x<s;enAkñúgsYteTAtMbn;Edlman pO2 TabenAkñúgQam . enATIenaH GuksIuEsncUl



eTAkñúgRKab;QamRkhm nig cgP¢ab;eTAnwgGIuy:g Fe2+ rbs; heme group én deoxyhemoglobin edaybegáIt )anCa Oxyhemoglobin . CaFmμtakarcgsm<½n§enHCYynaM[)an O2 cUleTAkñúgQam .

rUbTI 19-14

rUbTI 19-15

RBwtþkarN_énkarcgsm<½n§GuksIuEsnmankars μ úRKs μajxøH² . Deoxyhemoglobin manlMhrenAkñúgEpñk kNþalEdlenATIenaHGajú:gsrIragÁ 2,3-biphosphoglycerate (BPG) cgP¢ab; . enAeBl deoxyhemo-

globin subunit mYykñúgcMeNam subunits TaMgbYn)ancgP¢ab;CamYynwgGuksIuEsn karpøas;bþÚrrUbragRbUetGIun)an beBa©j BPG . karenHpþÜcepþImnUvRBwtþkarN_EdlenAsl; Hb subunits bIeTotTTYlrgCabnþbnÞab;nUvkarpøas;bþÚr rUbragRbUetGIunEdlbegáInlT§PaBkñúgkarTTYlykm:UelKulGuksIuEsn . ehtudUecñH enAeBl subunit TI 1 TTYl O2 , subunits bIEdlenAsl;bnþeFIVy:agelOn .



rUbTI 19-16

kMhab; H+ k¾man\T§iBleTAelIlT§PaBrbs; deoxyhemoglobin kñúgkarcgP¢ab;GuksIuEsnEdr . enA eBl deoxyhemoglobin cgP¢ab;CamYynwg O2 vabeBa©jRbUtug . RbUtugTaMgenHRtUv)andkecjy:agelOneday RbtikmμsMrYledayGg;sIumeQ μaH Carbonic anhydrase . Carbonic anhydrase H+ + HCO3

- → CO2 + H2O

Rbtikm μenHKWCaRbtikmμRtLb; . karBit HCO3- EdlCab;Tak;TgeTAkñúgRbtikm μenH RtUv)anbegáItecjBI

RbtikmμRtLb;enAeBl CO2 ¬kaksMNl;énemtabUlIs¦ cUleTAkñúgcrnþQam BICalikaEdleFIVemtabUlIs y:agskmμ. Carbonic anhydrase H+ + HCO3

- ← CO2 + H2O

enAkñúgsYt RbUtugEdlRtUv)anbeBa©jeday deoxyhemoglobin {ruj} RbtikmμKImIeTAxagsþaMeday beBa©jnUvkaksMNl; CO2 . CO2 enHPayecjBItMbn;Edlman pCO2 x<s;enAkñúgQamenAkñúgtMbn;Edlman pCO2 TabenAkñúgxül;kñúgsYt . bnÞab;mkvak¾nRtUv)andkecjBIsBa’gÁkayedaykardkdegðImecj . Rbtikm μeTAmux (forward reaction) k¾GacCYykarP¢ab;GuksIuEsnCamYynwg Hb GaRs½yedaykardk-ecjnUvGIuy:ug H+ BIkñúgQameP¶ackarcgGuksIuEsneday deoxyhemoglobin . ehtudUecñH karbeBa©j H+

enAkñúgry³eBlénkarP¢ab;GuksIuEsnrbs; deoxyhemoglobin eP¶acTaMgkardkecjnUvkaksMNl; CO2 BIsBa’gÁ kay nig karcgén O2 eday hemoglobin . enAeBlQamEdlsMbUredayGuksIuyEsnenAdl;CalikaEdleFIVemtabUlIsedayskm μ kaksMNl; CO2

PaycUleTAkñúgQam . karenHpþÜcepþImnUvkarRtLb;éndMeNIrkardUcEdleTIb)anBiB’Nna . CO2 cUleTAkñúg ekasikaRKab;QamRkhmEdlenATIenaH carbinic anhydrase sMrYlkarrYmKñarbs;vaCamYynwgTwkedIm,IbegáIt )anCa HCO3

- niig H+ . b:uEnþ dUcEdlkardkecjén H+ BIQamenAkñúgsYteP¶ackarP¢ab;GuksIuEsnrbs; Hb nig karekInnUv H+ enAkñúgQamenAeBlCYbCamYyCalikaEdleFIVemtabUlIsbegáInkarbeBa©jnUv O2 BI Hb . bnÞab;mk GuksIuEsnEdlbeBa©jsayPayBIQam ¬tMbn;Edlman pO2 x<s;¦ cUleTAkñúgCalikaEdlRtUvkarva . BPG begáIneLIyRcIneTotnUvkarbeBa©j O2 BI Hb . enAeBlEdl O2 RtUv)anbeBa©jBI subunit mYy kñúgcMeNam subunits TaMgbYnrbs; Hb , BPG cab;epþImeFIVkartampøÚvrbs;vaedayRtLb;cUleTAkñúglMhrrbs;



m:UelKul Hb . karenHbgár[mankarpøas;bþÚrrUbragrbs; subunits EdlGnuBaØati[BYkvabeBa©jGuksIuEsnEdl vaman . Myoglobin enAkñúgCalikamancMNUlKImIx<s;Cag Hb cMeBaHGuksIuEsn GaRs½yedayvamincgP¢ab;eTA nwg BPG . Myoglobin cgy:agelOnCamYy O2 EdlsayPaycUleTAkñúgCalika nig beBa©jva enAeBlvaRtUv karsMrab; aerobic respiration . K-kardwknaMGuksIuEsnBImþayeTA[TarkkñúgépÞ TarkenAkñúgépÞTTYl O2 BImþayrbs;vaedaykarsayPaysamBaØqøgkat;suk . RbsinebITaMgTark nig mþaymanRbePT Hb dUcKñadMeNIrkarbBa¢ÚnminmanRbsiT§PaBeT BIeRBaH Hb rbs;TarkkñúgépÞ nig Hb rbs;mþay mancMNUlKImIcMeBaHGuksIuEsndUcKña . b:uEnþTarkman Hb EtmYyRbePTKt;EdleKehAfaCa fetal Hemo-

globin. m:UelKul Hemoglobin EtmYyRbePTenHmancMNUlKImIcMeBaHGuksIuEsneRcInCag Hb rbs;mþay GaRs½yedayvamincgeTAnwg BPG dUc Hb rbs;mnusSeBjv½yeT . dUecñH O2 RtUv)andwknaMedayRbsiT§PaB tam ry³RbB½n§Qamrt; BIsYtrbs;mþayeTATarkkñúgépÞ . CIvsMeyaKén fetal Hb Qb;PøambnÞab;BIekItenAeBlEhSn EdlCaGñkkMNt;kUdsMrab; fetal Hb RtUv)an {biT} ehIyEhSnEdlCaGñkkMNt;kUdsMrab; Hb rbs;mnusSeBjv½y RtUv)an{ebIk} . X-Sickle Cell Anemia

Sickle cell anemia KWCaCMgWEhSnrbs;mnusSEdleLIgdMbUgeKenAtMbn;RtUBikxaglic nig GaRhVik kNþal . vab:HBal;dl;RbEhl 0.4% énCnCatiGaemrikEs,lexμA . buKÁlTaMgenHbegáItnUv mutant Hb EdleK sÁal;faCa sickle cell Hb (Hb S) . Sickle cell anemia TTYleQ μaHBIkarelcecjnUv sickled rbs;ekasika QamRkhmEdlekItenAkñúglkçx½NÐenH ¬rUbTI 19-17¦ . Sickled cells minGacqøgkat;sésQamtUc²én RbB½n§Qamrt; nig crnþQamRtUv)anraMg . karenHbNþal[mankarxUcxateTAnwgsrIr³CaeRcIn CaBiessq¥wg nig tMrgenam nig bNþal[søab;munGayu .

rUbTI 19-17

Sickled cell Hb xusBI Hb FmμtaedaysarEtGasIutGamIneNEtmYyKt; . enAkñúgExS β rbs; sickled

cell Hb , Valine (hydrophorbic amino acid) )anCMnYs glutamic acid ¬GasIutGamIneNmanbnÞúkGviC¢-



man¦ . karCMnYsenHpþl;CamUldæanRKwHénkarcgsm<½n§rbs;m:UelKul Hb S eTAnwgm:UelKulmYyepSgeTot . enA eBl Oxyhemoglobin S beBa©j O2 rbs;va m:UelKul deoxyhemoglobin enHcgP¢ab;CamYynwg deoxyhemoglobin S mYyepSgeTotekIt)anCasés polymers Evg . karenHekIteLIgGaRs½yeday valine

RtUvlμmnwgRbeLaH hydrophorbic EdlmanenAelIépÞrbs;m:UelKul deoxyhemoglobin S TI 2 . sés Hb

EdlRtUv)anbegáIteLIgtamrebobenHeFIV[xUcy:agxøaMgdl;rUbragrbs;ekasikaRKab;QamRkhm edaybNþal[ man\T§iBlragdUckeNÐóvenH . Sickle cell anemia ekIteLIgcMeBaHbuKÁlTaMgLayEdlTTYlmrtkénEhSncMeBaH sickle cell Hb BI «BukmþayEdlman sickle cell Hb . buKÁlEdlmanCMgWbegáItnUvExS βRbEhlBI 90-100° . buKÁlEdlTTYl )anEhSnFmμtamYy nig EhSnmanPaBex©aHmYybegáItTaMgExS β Fmμta nig minFmμta . RbEhl 10°énGaemrik Es,kexμAmanEtmYy copy rbs;EhSnmanPaBex©aH lkçxNÐenHRtUv)aneKsÁal;faCa Sickle cell trait . eTaHbI CavamineFIVb:HBal;F¶n;F¶k¾eday b:uEnþBYkeKman»kas 50° kñúgkarcMLgEhSneTA[kUnrbs;BYkeK . PaBTMnak;TMngEdlcab;GarmμN_ekItmanrvag sickle cell trait nig PaBFn;eTAnwgCMgWRKuncaj; . enAkñúg tMbn;mYycMnYnénTVIbGaRhVik manrhUtdl; 20° éncMnYnRbCaCnman sickle cell trait . kñúgcMeNamBYkTaMgenaH kñúgEpñkdUcKñaénTVIbGaRhVik mUlehtumYykñúgcMeNammUlehtu[søab;TaMgLayKWCaCMgWRKuncaj; . vtþmanén sickle

cell trait Cab;Tak;TgeTAnwgPaBFn;eTAnwgCMgWRKuncajekIneLIg GaRs½yeday)a:ra:sitCMgWRKuncaj;minGac ciBa©wmCIvitedaymanRbsiT§iPaBenAelI sickled red blood cells . RbCaCnEdlman sickle cell disease )an søab;kñúgv½yekμg GñkEdlminman sickle cell trait manlT§PaBx<s;kñúgkarsøab;edayCMgWRKuncaj; . edayKitCa mFüm RbCaCnEdlman sickle cell trait minTTYlrgeRcInBI sickle cell anemia nig Fn;edayRBmKñaeTAnwg CMgWRKuncaj;Edlbgá[søab; . GaRs½yedayGñkEdlman sickle cell trait man»kaseRcInkñúgkarrs; nig karbnþ BUC sickle cell Hb gen RtUvrkSaTukenAkñúgcMeNamRbCaCn . 10-kareFIV[xUcxaténRbUetGIun eyIg)anbgðajfarUbragrbs;RbUetGIunmansar³sMxanCadac;xatcMeBaHmuxgarrbs;va . eyIgk¾)anbBa¢ak; pgEdrfaCIvitGacmanEtenAkñúgcenøaHtUcceg¥óténsItuNðPaB nig pH . etIKMnitTaMgBIenHCab;Tak;TgKñay:agdUc emþc ? dUcEdleyIgnwgeXIj PaBelIsRCulén pH nig sItuNðPaBman\T§iBly:agxøaMgeTAelITMrg;rbs;RbUetGIun edaybgá[m:UelKul)at;bg;nUvrUbrag 3-D EdlmanlkçN³Biessrbs;BYkva . Denaturation ekIteLIgenA eBlrcnasm<½n§EdlerobcMRtwmRtUvrbs;RbUetGIunmYyCa α-helix , β-pleated sheet nig karbt;pñt;TI 3 RtUv)an xUcxatrcnasm<½n§TaMgRsug . b:uEnþvamineFIV[xUcdl;rcnasm<½n§TI 1 rbs;RbUetGIuneT . kareFIV[xUcén α-helix

protein RtUv)anbgðajenAkñúgrUbTI 19-18 .



rUbTI 19-18

k-sItuNðPaB cUrBicarNaBI\T§iBlénsItuNðPaBekIneLIgeTAelIsUluysüúgRbUetGIun ¬]TahrN_ sUluysüúgEpñks éns‘ut¦ . BIdMbUg karekIneLIgsItuNðPaBbegáInnUvGRtaénclnam:UelKul ¬kareFIVclnaénm:UelKulnimYy²enAkñúg sUluysüúg¦ . bnÞab;mk enAeBlsItuNðPaBbnþekIneLIg sm<½n§enAkñúgRbUetGIuncab;epþImj½ry:agxøaMg . cugBa©b; GnþrGMeBIexSay dUcCasm<½n§GIuRdUEsn nig GnþrGMeBI hydrophorbic EdlrkSarcnasm<½n§RbUetGIunRtUv)andac; . m:UelKulRbUetGIunRtUv)anxUcxatenAeBlBYkva)anbg;nUvTMrg; 3-D EdlmanlkçN³Biess nig køayeTACaxUcrcna sm<½n§TaMgRsug . Coagulation ekIteLIgenAeBlm:UelKulRbUetGIunElgbt;Capñt; nig RtUv)anCMBak;sμ úRKs μaj . enAkñúgcMNucenHBYkvaElgmanenAkñúgsUluysüúgeTotehIy BYkvakþaMgCab;KñaehIykøayeTACaGgÁFaturwg ¬emIlrUb TI 19-18¦ . Epñkséns‘utcab;epþImCasUluysüúgxab;énGal;b‘uymInrbs;s‘ut b:uEnþenAeBlRtUv)aneKcMGin RbUetGIunRbUv)anxUcxat nig kkkøayeTACaGgÁFaturwg . PaKeRcInénRbUetGIunrbs;ekasikaeyIg ]TahrN_ Gg;sIum CaRbePTsUluysüúgxab;dUcenHenAkñúgsIutU)aøs; rbs;ekasika . edIm,IbnþbMeBjmuxgaredayRtwmRtUv BYkvaRtUvEtsßitenACasUluysüúg nig rkSanUvTMrg; 3-D

RtwmRtUv . RbsinebIsItuNðPaBsBa’gÁkayekIneLIgx<s;eBk b¤ RbsinebItMbn;NamYyéndgxøÜnTTYlrgnUvsIutNð-PaBx<s;xøaMg dUcCaenAeBlb:HnwgKMrb;qñaMgekþA RbUetGIunekasikak¾xUcxat . BYkva)at;bg;nUvmuxgarrbs;BYkva nig ekasika b¤ srIragÁk¾søab; . x-pH GaRs½yedaybgÁúM R rbs;GasIutGamIneN RKb;RbUetGIunmanbnÞúkGKÁisnIBiess . GaRs½yedayRbUetGIun nimYy²mansmasFatuGasIutGamIneNepSg² RbUetGIunnimYy²nwgmanbnÞúkGKÁisnIBiesssrubenAelIépÞrbs;va . bgÁúM R EdlmanbnÞúkGKÁisnIviC¢man b¤ GviC¢manenAelIépÞrbs;m:UelKulmanGnþrGMeBICamYynwgGIuy:ug nig m:UelKulTwk nig GnþrGMeBITaMgenHrkSaRbUetGIun[enACasUluysüúgenAkñúgsIutU)aøs; . RbUetGIunEdlbgðajenAkñúgrUbTI 19-19ª manbnÞúksrub 2+ GaRs½yedayvamanbgÁúM ─NH3

+ BIr . RbsinebIeyIgEfmmYy dUcCa NaOH bgÁúMGamInEdlmanRbUtug)at;RbUtugrbs;va nig ehtudUecñHvamabnÞúkGKÁisnI NWt . \LÚvbnÞúksrubrbs;RbUetGIunKW 0 . enAeBl Polypeptide mancMnYnbnÞúkviC¢man nig GviC¢manes μ IKña eKehA



faCa Isoelectric . RbUetGIunEdlRtUv)anbgðajenAkñúgrUbTI 19-19b manbnÞúksrubes μ I 2– GaRs½yedayman bgÁúM carboxyl bEnßmBIreTot . enAeBlGasIutRtUv)aneKbEnßm bgÁúM carboxyl cab;ykRbUtug . \LÚv vak¾NWt nig bnÞúksrubenAelIRbUetGIunesI μ 0 . dUc CaenAkñúg]TahrN_BImunmk \LÚvenHRbUetGIunman Isoelectric .

rUbTI 19-19

enAeBlRbUetGIunmimanbnÞúksrubenAelIépÞrbs;va BYkvaElgRcanKñaecjeTotehIy . CalT§pl BYkva kþaMgCab;Kña nig køayeTACakkrecjBIsUluysüúg . enAkñúgrNIenH RbUetGIunGackk eTaHbICaminxUcxatk¾eday . karenHKWCaRbtikm μEdlGñkGacBinitüeXIjenAkñúgpÞH)ayrbs;Gñk . enAeBlTwkedaHRtUv)aneKdak;enAkñúg TUrTwkkkenAkñúgry³eBlyUr )ak;etrIenAkñúgTwkedaHcab;epþImduH (grow) . )ak;etrIeRbIsárTwkedaH (lactose)

CaRbPBfamBlenAkñúgdMeNIrkarénkareFIV[eLIgCaem (fermentation) nig begáIt)anCa lactic acid Ca plitplcugbBa©b; . enAeBl)ak;etrIbnþduH kMhab; lactic acid kan;EtekIneLIg . GasIutEdlbEnß (additional

acid) bNþal[mankarcab;RbUtugrbs;bgÁúM carboxyl enAelIépÞénRbUetGIunEdlrlayenAkñúgTwkedaH . BYkvakøay eTACa Isoelectric nig kkeTACaTwkedaHkk (solid curd) . cUrRsm½ymYyEPøtBIGIVEdlnwgekIteLIg RbsinebI pH rbs;QamGasIutxøaMgeBk b¤ )asxøaMgeBk . Qam KWCasnÞnIy_ (fluid) EdlmanTwk nig eGLikRtUlItEdlrlay RbePTepSg²énekasika rYmmanRKab;QamRkhm EdlTTYlxusRtUvkñúgkardwknaM O2 nig RbUetGIunepSg²CaeRcIneTot . RbUetGIunTaMgenHrYmman fibrinogen Edl Cab;Tak;TgeTAnwgRbtikm μénkarkkQam Immunoglobulins EdlkarBarxøÜneyIgBICMgW nig Albumins Edl dwknaM hydrophorbic molecules enAkñúgQam . enAeBl pH QamFøak;cuHTabxøaMgeBk RbUetGIunkñúgQamk¾køayeTACa polycations . dUcKñaenHEdr enA eBl pH QamekIneLIgx<s;xøaMgeBk RbUetGIunk¾køayeTACa polyanions . kñúgkrNITaMgBIr RbUetGIunnwgrlapñt; GaRs½yedaykarRcanecjénbnÞúk nig )at;bg;kareFIV[manesßrPaBnUvGnþrGMeBIrbs;GIuy:ug . eRkamlkçxNÐRCul



hYsehtuTaMgenH RbUetGIunkñúgQamEdlxUcxatnwgElgbMeBjmuxgarEdlvaRtUvkareTotehIy . ekasikaQam k¾RtUvsøab;enAeBlGg;sIumsMxan;RtUv)anxUcxat . Hb enAkñúgekasikaRKab;QamRkhmk¾xUcxat nig ElgGacdwknaM GuksIuEsneTotehIy . edaymanPBVvasna sBa’gÁkaymanynþkarmYycMnYnedIm,IeCosvagkarpøas;bþÚrsMxan;nUv pH rbs;QamEdlGacekIteLIgCalT§plénPaBlM)akénemtabUlIs b¤ kardkdegðIm . K-FaturMlaysrIragÁ (Organic solvents)

FaturMlaysrIragÁEdlmanb:Ul dUcCa rubbing alcohol (2-propanol) bMEbkRbUetGIunedaykarpþac;nUv sm<½n§FGIuRdUEsnenAkñúgRbUetGIun bEnßBIelIkarbegáItnUvsm<½n§GIuRdUEsnCamYynwgFaturMlay Twk . tMbn;minmanb:Ul rbs;FaturlayTaMgenHeRCótERCkedayGnþrGMeBI hydrophorbic enAEpñkxagkñúgénm:UelKulRbUetGIun EdlenATI enaHeFIV[xUcTMrg;rbs;RbUetGIun (conformation) . tamRbéBNI sUluysüúg rubbing alcohol 70% Cajwk jab;RtUv)aneKeRbICa disinfectant b¤ antiseptic . b:uEnþ P½sþútagfμ I²pþl;Caeyabl;fa vaminEmnCaPñak;garman RbsiT§PaBenAkñúgsmtßPaBenH . X-Detergents

Detergents manTaMgtMbn; hydrophorbic ¬knÞúyGasIutxøaj;¦ nig tMbn; hydrophilic ¬bgÁúM carbonyl ¦ . enAeBl detergents manGnþrGMeBICamYynwgRbUetGIun BYkvapþac;GnþrGMeBI hydrophorbic eday bgá[ExSRbUetGIunrla . g-elah³F¶n; (Heavy Metals)

elah³F¶n;dUcCa)art (Hg2+) b¤ sMN (Pb2+) GacbegáItCasm<½n§CamYynwgbgÁúMcugExSEdlmanbnÞúk GviC¢-man . karenHeRCotERCkCamYynwgsm<½n§GMbil (salt bridge) EdlRtUv)anbegáItrvagbgÁúM R rbs;GasIutGamIneN rbs;ExSRbUetGIun edaynaM[)at;bg;nUvTMrg; (conformation) . elah³F¶n;k¾Gaccgsm<½n§CamYynwgbgÁúM sulfhydryl rbs;RbUetGIun . karenHGacbgá[mankarpøas;bþÚry:agxøaMgenAkñúgrcnasm<½n§ 3-D rbs;RbUetGIun EdlGmeTAedaykar)at;nUvmuxgar . c-kMlaMgemkanik (Mechanical Stress)

karkUr karvay karRkLúk GaceFIV[dac;GnþrGMeBIexSayEdlrkSanUvTMrg;RtwmRtUvRbUetGIun . karenHCa ehtupl EdlvayEpñkséns‘utedIm,IbegáItCanMdut (stiff meringue) . 11-RbUetGIunkñúgrbbGahar nig karrMlayRbUetGIun RbUetGIunk¾dUcCa carbohydrates nig xøaj; KWCaRbePTsMxan;énRbPBfamBlenAkñúgrbbGahar . dUcCa Carbohydrate nig xøaj; RbUetGIuneRbIenAkñúgkareFIVmðÚbCaeRcIn . BYkvaTTYlrgGuksIutkmμedIm,IbegáItCafamBl. m:üagvijeTot GasIutGamIneNEdlRtUv)anbeBa©jedaysarkareFIV hydrolyze RbUetGIunRtUv)aneRbIedaypÞal;enA kñúgCIvsMeyaK (biosynthesis) . ynþkarsMeyaKRbUetGIunrbs;ekasikaGaceFIVsmahr½NkmμGasIutGamIneN ¬EdlbeBa©jedaykarbMEbkénRbUetGIunkñúgGahar¦edaypÞal;eTACaRbUetGIunfμ Irbs;ekasika . GasIutGamIneNk¾ RtUv)aneRbIEdrenAkñúgCIvsMeyaKénm:UelKulsMxan;FM²CaeRcIn EdleKehAvafaCa Nitrogenous compounds .



RkumenHrYmman hormones, heme group rbs; Hb nig Myoglobin nig BYk)asEdlmanGasUt EdleKeXIj manenAkñúg DNA nig RNA . karbMEbkRbUetGIunenAkñúgcMNIGaharcab;epþImenAkñúgRkBH . Gg;sIumrbs;RkBH Pepsin cab;epþImbMEbk edayGIuRdUlIssm<½n§ peptides mYycMnYnrbs;RbUetGIun . karenHbMEbkRbUetGIuneTACa peptides tUc² . karplitén Pepsin nig Gg;sIumbMEbkRbUetGIunrbs;bMBg;GaharepSgeTotRtUvEtRtUv)anRtYtBinitüeday RbugRby½tñedaysarGg;sIumskmμRtUvrMlay nig bMpøajekasikaEdlplitGg;sIumbMEbRbUetGIunTaMgenaH . dUecñH ekasikaRkBH (stomach lining cells) Edlplit pepsin RtUv)ansMeyaKCaFmμta nig beBa©jCaTMrg;minskmμ EdleKehAfaCa Pepsinogen . Pepsinogen manGasIutGamIneN 42 bEnßmeTotenAkñúgrcnasm<½n§TI 1. BYk vaenHRtUv)andkecjBIkñúgRkBHedImplitCa Pepsin skmμ (active pepsin) . karbMEbkRbUetGIunbnþenAkñúgeBaHevontUcEdlenATIenaHGg;sIum Trypsin, chymotrypsin, elastase nig Gg;sIumepSgeTotsMrYlkarGIuRdUlIsénsm<½n§ peptides enAkñúgTItaMgepSg²enAkñúgRbUetGIun . ]TahrN_ chymotrypsin kat;sm<½n§ peptide enAcug carbonyl rbs; aromatic amino acid nig trypsin kat;sm<½n§ peptide enAelIcug carbonyl rbs; basic amino acids . edayrYmKña Gg;sIumbMEbkRbUetGIunTaMgenHbMEbk RbUetGIunkñúgrbbGaharCaeRcIneTACaGasIutGamIneNEdlGacRsUbedayekasikarbs;eBaHevontUc . GasIutGamIneNGacEckCafñak;rbbGaharsMxan;BIr . GasIutGamIneNcaM)ac; (Essential amino

acids) KWCaGasIutGamIneNEdlminGacsMeyaKedaysBa’gÁkay nig RtUvkar[manenAkñúgrbbGahar . GasIut GamIneNmincaM)ac; (Nonessential amino acids) KWCaGasIutGamIneNEdlGacsMeyaKedaysBa’gÁkay nig minRtUvkardak;bBa©ÚleTAkñúgrbbGahar . taragTI 19-2 bgðajBIGasIutGamIneNcaM)ac; nig GasIutGamIneN mincaM)ac; .

taragTI 19-2 Essential Amino Acids Nonessential Amino Acids

Isoleucine Alanine Leucine Arginine Lysine Asparagine Methionine Aspartate Phenylalanine Cysteine Threonine Glutamate Tryptophan Glutamine Valine Glycine Histidine Proline Serine Tyrosine

RbUetGIunk¾RtUv)aneKeFIVcMNat;fñak;Ca Complete nig Incomplete . RbUetGIunEdlTTYl)anBIRbPBstV CaTUeTACa Complete proteins . dUecñH vapþl;nUvRKb;GasIutGamIneNcaM)ac; nig GasIutGamInmincaM)ac;enAkñúg cMnYnRtwmRtUvsMrab;CIvsMeyaK . pÞúymkvij RbUetGIunEdlmanRsLayBIRbPBbEnøCaTUeTAKWCa Incomplete



protein GaRs½yedayedayvaminmancMnYnRKb;RKan;nUvGasIutGamIneNcaM)ac;mYy b¤ eRcIn . RbCaCnEdlBYkeK cg;rkSanUvrbbGaharEdlmanEtbénøsuT§ (strictly vegetarian diet) b¤ cMeBaHGñkEdlminmansac;stVbriePaK manbBaðaEdlfa minmanbEnøEdlmanRbUetGIunx<s;manRKb;GasIutGamIneNcaM)ac;edIm,IFanakarbriePaKGahar RbcaMéf¶RKb;RKan; . ]TahrN_ RbUetGIunsMxan;rbs;sENþkesogmansMbUreTAeday lysine nig Tryptophane

b:uEnþman methionine ticxøaMgNas; . rIÉeBatman methionine y:ageRcIn b:uEnþman tryptophane b¤ lysine

ticxøaMgNas; . b:uEnþ l,ayéneBat nig sENþkesogRKb;RKan;sMrab;kartMrUvTaMgeTVrenH . karrYmKñaenH RtUv)aneK ehAfaCa “Succotash” EdlKWCasmasFatusMxan;énrbbGaharrbs; Native Americans sMrab;RbeTsCa eRcIn . karbriePaKrbbGaharbEnøsuT§ (vegetarian meals) BIr b¤ bI dg ral;GaTitüGacpþl;nUvRKb;GasIutGamIn eNEdlRtUvkar nig edayÉkÉgCYykat;bnßynUvcMnYnxøaj;Eq¥t (saturated fat) enAkñúgrbbGahar . Gaharrbs; CatiBn§ (ethnic foods) CaeRcInerobcMeLIgtameKalkarN_énkarlaycUlKñaénRbPBRbUetGIun . Gaharrbs;BYk miksiukdUcCa Tortillas nig Refried beans, Cajun dishes of spy beans and rice, Indian cuisin

of rice and lentils nig sUm,IEt Traditional American peanut butter sandwich KWCa[TahrN_ én viFIlayGaharcUlKñaedIm,Ipþl;)annUv complet protein .

Fin

CIvKImIviTüa Gg;sIum

erobcMnigbkERbeday»sfkarIRcwgsuPaBTUrs½BÞ012864353b¤011717361 e-mail: [email protected] Page 1 of 33

Gg;sIum (ENZYMES)

vtßúbMNgemeron 1- pþl;nUv]TahrN_énTMnak;TMngrYmKñarvageQ μaHEdlCYbjwkjab;rbs;Gg;sIumnigmuxgarrbs;va . 2- eFIVcMNat;fñak;Gg;sIumeyageTAtamRbePTRbtikm μEdlvasMrYlnigRbePTénPaBBiess . 3- BiB’NnaGMBI\T§BlEdlGg;sIummaneTAelIfamBlEdleFIV[skmμénRbtikmμ . 4- yl;dwgGMBI\T§iBlrbs;kMhab; substrate eTAelIRbtikmμEdlsMrYledayGg;sIum . 5- BiPakSaGMBItYnaTIénTIkEnøgskmμ nig sar³sMxan;énPaBBiessrbs;Gg;sIum . 6- BiB’NnaBIPaBxusKñarvagkarbegáItbNþúMr enzyme-substrate tamKMrUemesa-kUnesa nig tamKMrUEdl eFIV[RtUvKña (induced fit model) . 7- BiPakSaBImuxgarrbs; Cofactors nig Coenzymes enAkñúgskmμPaBrbs;Gg;sIum . 8- Bnül;faetI pH nig sItuNðPaBb:uNÑaEdlman\T§iBleTAelIRbtikmμEdlsMrYledayGg;sIum . 9- yl;GMBIynþkarEdleRbIedayekasikaedIm,IEktMrUvskmμPaBGg;sIum . 10- BiPakSaGMBIynþkarEdlCatiKImIEdlraraMgskm μPaBrbs;Gg;sIum . 11- BiPakSaGMBImuxgarrbs;Gg;sIum chymotrypsin nig Serine proteases epSgeTot . 12- pþl;nUv]TahrN_énbMerIbMras;kúñgevC¢saRsþrbs;Gg;sIum .



KMeragemeron 1- namvlI nig cMNat;fñak; (Nomenclature and Classification)

k- namvlIrbs;Gg;sIum x- cMNat;fñak;rbs;Gg;sIum 2- \T§iBlrbs;Gg;sIumeTAelIfamBleFIV[skmμrbs;Rbtikmμ 3- \T§iBlrbs;kMhab; Substrate eTAelIRbtikmμEdlsMrYledayGg;sIum 4- bNþúM Enzyme-Substrate

5- PaBBiessénbNþúM Enzyme-Substrate

6- sßanPaBbeNþaHGasnñ nig karbegáItnUvplitpl 7- Cofactors and Coenzymes 8- \Ti§BlénbriyakasCMuvij k- \T§iBlén pH

x- \T§iBlsItuNðPaB 9- karEktMrUvénskmμPaBGg;sIum k- Allostreic Enzyme

x- Feedback Inhibition

K- Zymogens

10- karraraMgénskm μPaBGg;sIum k- GñkraraMgminRtLb; x- GñkraraMgRbECgRtLb;mkvij K- GñkraraMgminRbECgRtLb;mkvij 11- Gg;sIumbMEbkRbUetGIun 12- kareRbIGg;sIumenAkñúgevC¢saRsþ



Gg;sIum (ENZYMES)

esckþIepþIm Gg;sIumKWCam:UelKulCIvsaRsþEdleRbICaGñksMrYl (catalyst) RbtikmμKImI . PaKeRcInénGg;sIum KWCaRbUetGIun . minmanGg;sIumedIm,IbegáInel,OnRbtikmμCIvKImI CIvitnwgminGacmaneLIy . CIvitekasika Cab;Tak;TgeTAnwgkarelcecjdMNalKñanUvrab;ryRbtikmμKImIEdlRtUvEtekIteLIgya:gelOn sßiteRkam lkçx½NÐl μm . vaGac ¬]TahrN_¦ kñúgkarbEnßmTwleTAelI Gal;Esn (Alkene) . b:uEnþ RbtikmμenHCa TUeTAGnuvtþenAkñúgsItuNðPaB 100oC enAkñúgsUluysüúgGasIuts‘ulhVÜrik (aqueous sulfuric acid) . lkçx½NÐrebobenHnwgsMlab;ekasika . ekasikaEdlgayxUcRtUvEtGnuvtþRbtikmμKImIrbs;vaenAkñúg sItuNðPaBdgxøÜn (37o) nig kñúgvtþmanénGasIutxøaMg b¤ )asxøaMg . etIkarenHvaGacsMerceTA)antam rebobdUcemþc ? enAkñúgEpñkTI 3 eyIgnwg)aneXIjfa GñksMrYl (catalyst) bnßynUvfamBleFIV[skm μ (energy of activation) rbs;Rbtikm μKImI nigenATIenaHvabegáInnUvGRtaénRbtikmμ . karenHGnuBaØati[ RbtikmμekIeLIgeRkamlkçx½NÐlμm (milder conditions) . ekasikaeRbIGg;sIumedIm,IedaHRsaybBaða énkareFIV[manRbtikmμKImIedIm,Ielcecjya:gelOneRkamlkçx½NÐl μmEdleKeXIjmanenAkñúgekasika. Gg;sIumsMrYl (facilitate) RbtikmμCIvKImI edaykareFIV[fycuHnUvfamBleFIV[manskmμPaB nig begáIn nUvGRtaRbtikmμ . karbMeBjmuxgarEdlmanRbsiT§PaBrbs;Gg;sIumKWvasMxan;cMeBaHCIvitekasika nig srIragÁ (organism) . )atuPUTeTVrénPaBmanlkçN³Biessx<s; (high specificity) nigGRtaRbtikmμelOn (rapid

reaction rate) CamUldæandMbUg (cornerstone) énskmμPaBrbs;Gg;sIum nig CaRbFanbTrbs;CMBUk enH. ekasikaFmμtamanrab;Ban;m:UlelKulepSg² vanimYy²Edlmansar³sMxan;enAkñúgKImIéndMeNIrkar CIvit . Gg;sIumnimYy²TTYlsÁal; EtmYy b¤ mþgmþalBIb¤bIénm:UelKulTaMgenH . TidæPaBEdlKYr[kt;Cag eKbMputrbs;Gg;sIumKW PaBBiess (specificity) enH . Gg;sIumnimYy²TTYlsÁal; nig cgP¢ab;eTAnwg RbePTEtmYyén Substrate b¤ Reactant . TMhMm:UelKul rUbrag nig karEbgEckbnÞúkGKÁisnI (charge

distribution) rbs;TaMgGg;sIum nig Substrate RtUvEtcuHsMrugKña (compatible) edIm,I[dMeNIrkar énkarcgEdleRCIserIsenHekIteLIg . bnÞab;mkGg;sIumbMElg Substrate eTACa plitpl (product)

Edlmanel,OnelOn . CakarBitRbtikmμEdlsMrYledayGg;sIumCajwkjab;ekIteLIgBImYYylaneTA 100 landgelOnCagRbtikmμEdlRtUvKñaehIyminsMrYledayGg;sIum . Gg;sIum Catalase pþl;nUv]TahrN_EdlKYr[cab;Garm μN_énkarekIneLIgnUvGRtaRbtikmμEdl naMmkedaysarGg;sIum . Gg;sIumenHRtUvkarsMrab;CIvitenAkñúgbrisßanEdlmanGuksIuEsn . enAkñúg



brisßanenH dMeNIrkarénkarbMEbkm:UelKulGaharEdlRtUvkarGuksIuEsnbegáItnUv H2O2 . BIeRBaH H2O2

BulcMeBaHekasika dUecñHvaRtUvEtbMpøajecal . mYym:UelKulrbs; catalase bMElg 40 lanm:UelKul én H2O2 [køayeTACaTwkminpþl;eRKaHfñak; nig GuksIuEsnkñúgral;vinaTI .

karenHKWCaRbtikmμdUcKñaEdlGñkeXIjenAeBlGñkcak; H2O2 eTAelIdMe)A . Catalase Pay ecjBIekasikaEdlmanrbYsya:gelOnedaybMEbk H2O2 . BBuHEdlGñkeXIjKWCa O2 EdlPayCa plitplRbtikmμ . 1- namvlIv nig cMNat;fñak;Gg;sIum (NOMENCLATURE AND CLASSIFICATION)

k- namvlIvGg;sIum (Nomenclature of enzymes)

eQ μaHjwkjab;rbs;Gg;sIummanRsLayBIeQμaHrbs; substrate ¬RbtikrEdlP¢ab;CamYynwg Gg;sIum nig RtUv)anbMElgeTACaplitpl¦ EdlGg;sIummanRbtikmμCamYynig¼b¤RbePTénRbtikmμEdl vasMrYl . GaRs½yehtuenH muxgarrbs;Gg;sIumCaTUeTACab;Tak;TgedaypÞal;eTAnwgeQμaHsÁal;Cajwk jab;rbs;va . cUrBinitüemIl]TahrN_mYycMnYnenAkñúgKMnitsamBaØ . Urea KWCa substrate RtUv)anTTYlrg RbtikmμedayGg;sIumeQμaH Urease . Urea - a + ase => Urease

cUrtt;sMKal;faeQ μaHrbs;Gg;sIumenHCaFmμtaCaeQ μaHrbs; substrate rYmedaybc©wmbT "ase". rYmCamYykarelIkElgéneQ μaHCYbjwkjab;CaRbvtþisaRsþ karbBa©b;CaTUeTAcMeBaHeQ μaHrbs;Gg;sIum KW "ase" . Ca]TahrN_ Lactose KWCa sunstrate rbs; lactase . Lactose - ose + ase => Lactase

Gg;sIumepSgeTotGac[eQ μaHeTAtamRbtikm μEdlvasMrYl ]TahrN_ ³ Dehydrogenases dkGatUmGIuRdUEsn edaybBa¢ÚnBYkvaeTA[ Coenzyme . Decarboxylases dkbgÁúMkabuksIul . buBVbT "de" bgðajfabgÁúMnaTIRtUv)andkecj . müa:gvijeTot Hydrogenases nig Carboxylases bEnßmGIuRdUEsn b¤ bgÁúMkabuksIul . eQμaHGg;sIummYycMnYnrYmTaMgeQ μaHrbs; substrate

Catalase 2 H2O2 → 2 H2O + O2

RbtikmμekIteLIg 40 landgral;vinaTI



nig eQ μaHRbePTRbtikm μ . ]TahrN_ Lactate dehydrogenase dkGatUmGIuRdUEsnBIGIuy:ug lactate nig Pyruvate decarboxylase dkbgÁúMkabuksIulBI Pyruvate . dUcCaenAkñúgvis½yepSgeToténKImI eQμaHRbvtþiEdlKñanTMnak;TMngTal;EtesaHeTAnwgeQ μaHrbs; substrate nig Rbtikm μenAEtRtUv)aneKbnþeRbIR)as; . enAkñúgkrNIenH substrate nig Rbtikmμ RtUvEtcaM CaFm μta . ]TahrN_mYycMnYnéneQ μaHRbvtþisaRsþEdlCYbCajwkjab;rYmman catalase, pepsin,

chymotrypsin nig Trypsin . x- cMNat;fñak;rbs;Gg;sIum (Classification of Enzymes)

Gg;sIumGacRtUvcat;fñak;GnuelameTAtamRbePTénRbtikmμEdlvaCab;Tak;Tg . manGg;sIum 6 fñak; dUcxageRkam ³ 1- Oxidoreductases Oxidoreductases KWCaGg;sIumEdlsMrYlRbtikmμ oxidation-reduction (redox) . Lactate

dehydrogenase KWCa oxidoreductase EdldkGatUmGIuRdUEsnBIm:UelKul Lactate . fñak;rgepSgeTot én oxidoreductases rYmman oxidases nig reductases . LDH

CH3—CHOH—COO− + NAD+ ⇄ CH3—CO—COO− + NADH + H+ ` 2- Transferases

Transferases KWCaGg;sIumEdlsMrYlkarbBa¢ÚnénbgÁúMnaTIBIm:UelKulmYyeTAm:UelKulmYYyepSg eTot . ]TahrN_ Transaminase sMrYlkarbBa¢ÚnénbgÁúMGamIn nig Kinase sMrYlkarbBa¢ÚnbgÁúM Phosphate . Kinases edIrtYnaTIsMxan;dMeNIrkarRbmUlfamBlEdlCab;Tak;TgeTAnwg ATP . enAkñúg RkeBj Adrenal (Adrenal glands) Norepinephrine RtUv)anbMElgeTACa epinephrine edayGg;sIum eQ μaH Phenylethanolamine-N-methyltransferase (PNMT) KWCa Transmethylase . 3- Hydrolases

—CHCH2NH2 І OH

HO—

HO

Methyl group donor + —CHCH2NH—CH3

І OH

HO—

HO

PNMT

Epinephrine Norepinephrine



Hydrolases sMrYlRbtikmμGIuRdUlIs EdlvaenHKWCakarbEnßmm:UelKulTwkeTAelIsm<½n§Edl bNþal[mankardac;sm<½n§ . RbtikmμTaMgenHvasMxan;enAkñúgdMeNIrkarrMlayGahar . ]TahrN_ Lipase sMrYlRbtikmμénsm<½n§eGEsÞr (ester bonds) enAkñúg Triglycerides) . 4- Lyases

Lyases sMrYlkarbEnßmbgÁúMnaTIeTAelIsm<½n§BIrCan; b¤ kardkykénbgÁúMnaTIedIm,IbegáItCa sm<½n§ BIrCan; . Citrate lyase sMrYlkardkénbgÁúM Acetyl BIm:UelKul Citrate . plitplrbs;RbtikmμenHrYm man Oxaloacetate, acetyl CoA, ADP nig bgÁúM Phosphate minEmnsrIragÁ (Pi) . 5- Isomerases

Isomerases erobcMeLIgvijnUvbgÁúMnaTIenAkñúgm:UelKul nig sMrYlkarbMElgBIGIusUEmrmYyeTACa GIusUEmrmYyepSgeTot . ]TahrN_ Phosphoglyceromutase bMElgGIusUEmréncnasm<½n§ 3-phospho-

glycerate eTACaGIusUEmrmYyepSgeTotKW 2-phosphoglycerate . 6- Ligases

Ligases CaGg;sIumsMrYlkarrYmKña b¤ karP¢ab;Kñarbs;m:UelKulBIr . ]TahrN_ DNA ligases

sMrYlkarP¢ab;KñaénbgÁúMGIuRduksIul (OH) rbs; nucleotide enAkñúgExS DNA CamYynwgbgÁúM Phosphoryl

kñúgkarP¢ab; Nucleotide edIm,IbegáIt)anCasm<½n§ Phosphoester ³

CH2—O—CO—(CH2)n—CH3 І Lipase CH—O—CO—(CH2)n—CH3 + 3 H2O → І CH2—O—CO—(CH2)n—CH3 Triglycerides

CH2—OH І CH—OH + 3 CH3—(CH2)n—CH3 І CH2—OH Glycerol Fatty acids

CH2—COO−

І Citrate lyase HO—CH—COO− + ATP + CoenzymeA + H2O → І CH2—COO−

Citrate

CH2—COO−

І CO + CH3—CO~S—CoA +ADP +Pi І COO−

Oxaloacetate Acetyl CoA

COO− O І ІІ CH—O—P—O−

І І CH2 O

− І OH 3-Phosphoglycerate

COO− І CH—OH O І ІІ CH2—O—P—O−

І O−

3-Phosphoglycerate

Phosphoglyceromutase



2- \T§iBlrbs;Gg;sIumeTAelIfamBleFIV[skmμrbs;Rbtikmμ (THE EFFECT OF ENZYMES ON THE ACTIVATION ENERGY OF REACTION)

etIGg;sIumbegáInel,OnRbtikmμKImItamrebobdUcemþc ? vamankarpøas;bþÚrpøÚvEdlRbtikm μKImIekIt eLIg edaypþl;nUvpøÚvEdlRtUvkarfamBlTabCagsMrab;karbMElgnUv substrate eTACaplitpl ¬RbePT énCatiKImI [chemical species] EdlTTYl)anBIRbtikmμKImIEdlsMrYledayGg;sIum¦ . ehtudUecñH Gg;sIumbegáInRbtikm μedayeFIV[TabnUvfamBlEdleFIV[skmμénRbtikm μ (activation energy of

reaction) . cUrrMlwkfa RbtikmμKImInimYYy² RtUv)anbBa¢ak;lkçN³eday efrlMnwg (equilibrium constant) . cUrBicarNa ]TahrN_ lMnwgsamBaØmYy aA ⇄ bB

efrlMnwgcMeBaHRbtikmμGacsresr ³

CaFmμtaefrlMnwgenHKWCakarqøúHbBa©aMgénPaBxusKñanUvfamBlrvagRbtikr (reactant) nig plitpl (product) . vaKWCakarvas;énPaBsßitesßrEdlCab;Tak;Tg (Relative stabilities) rbs;Rbtikr nig plitpl . eTaHbICaRbtikmμKImIminekItk¾eday ¬pøÚvEdlvaRtUveFIVtam¦ b:uEnþPaBxusKñaénfamBl rvagRbtikr nig plitplCaTUeTAdUcKña . Gg;sIummineFIV[xUcefrlMnwgRbtikm μEdlvasMrYleT . b:Enþ Gg;sIumR)akdCaeFIV[pøas;bþÚrpøÚvEdldMeNIrkarekIteLIg edaypþl;nUvpøÚvEdlRtUvkarfamBlTabsMrab; karbMElgén substrate [eTACaplitpl . Gg;sIumbegáInnUvGRtaRbtikmμKImIedayeFIV[TabnUvfamBl

O ІІ DNA strand—3'—OH + O−—P—O—5'—DNA strand І O−

O ІІ DNA strand—3'—OH—P—O—5'—DNA strand І O−

DNA ligase

[B]b [Product]b Keq = ------ = ------------ [A]a [Reactant]a



eFIV[skmμ cMeBaHRbtikmμ ¬rUbTI 21-1¦ . ehtuenHGg;sIumbegáInGRtaRbtikm μEdlvasMrY[dl;man lMnwgRbtikm μ .

rUbTI 21-1 rUbKMnUsbgðajBIPaBxusKñaénfamBlrvagRbtikr (A nig B) nig plitpl (C nig D)

cMeBaHRbtikmμmYy . Gg;sIumminGacpøas;bþÚrPaBxusKñaénfamBlenAeT b:uEnþ vaedIrtYCa GñkeFIV[TabnUvfamBleFIV[manskm μPaB (Ea) rbs;Rbtikmμ EdlenATIenHbegáInel,OnRbtikm μ.

3- \T§iBlrbs;kMhab; SUBSTRATE eTAelIRbtikmμEdlsMrYledayGg;sIum (THE EFFECT OF SUBSTRATE CONCENTRATION ON ENZYME-CATALYZED REACTIONS)

GRtaRbtikmμKImIEdlminsMrYledayGg;sIumCajwkjab;ekIneLIgeTVrdg enAeBlkMhab;rbs; substrate RtUv)anbegáIneLIgeTVr ¬rUbTI 20-2a¦. ehtudUecñH enAeBlkMhab; substrate ekIneLIg vamankarekIneLIg edaypÞal;nUvGRtaRbtikmμ . b:uEnþ cMeBaHRbtikm μEdlsMrYledayGg;sIummindUckrNI enHeT . eTaHbICa BIdMbUgGRtaRbtikm μeqøIytbeTAnwgkMhab;rbs; substrate k¾eday b:uEnþsßitenAkñúg kMhab;mYycMnYnNaenaH rbs; substrate GRtaénRbtikmμQandl;kMritGtib,rma . ExSekagén GRtaRbtikmμ (V) pÞúyeTAnwg kMhab;rbs; substrate [S] dUcEdl)anbgðajenAkñúgrUbTI 20-2 b . eyIg )aneXIjfaBIdMbUgGRtaRbtikmμekIneLIgya:gelOn enAeBlkMhab;rbs; substrate ekIneLIg b:uEnþ eyIg eXIjEdrfaGRtaRbtikmμRtUvQb;enAkñúgkMrittMélGtib,rma . enAkñúgGRtaGtib,rmaenH TIkEnøgskmμ

Reactant A+B

Product C+D

Progress of reaction

Ene

rgy

Catalyzed reaction

Reactant A+B

Product C+D

Progress of reaction

Ene

rgy

Uncatalyzed reaction



(the active sites) rbsRKb;m:UelKulGg;sIumenAkñúgsUluysüúgRtUv)anP¢ab;eTAedaym:UelKul substrate. TIkEnøgskm μ KWCatMbn;rbs;Gg;sIumEdlP¢ab;CamYy substrate nig sMrYlRbtikmμrYmeday lkçN³ Biess . m:UelKulf μ Irbs; substrate minGacP¢ab;CamYym:UelKulGg;sIumEdleKpþl;[ rhUtdl; m:UelKul substrate EdlRtUv)anP¢ab;rYcehIyeTAnwgGg;sIumRtUv)anbMElgCalT§pl . ehtudUecñH vahak;Cafa RbtikmμEdlsMrYledayGg;sIumekIteLIgkñúgBIrdMNak;kar . dMNak;karTI mYy ¬dMNak;kar elOn¦ KWCakarbegáItnUv Enzyme-substrate complex . dMNak;karTI 2 ¬CadMNak;karyWt¦ nig ehtudUecñHvaRtYtBinitüGRtaRbtikmμEdlekIeLIg . eKniyayfavaCa GñkkMNt;RBMEdnGRtaRbtikm μ (rate-limiting) nig Cab;Tak;TgeTAnwgkarbMElgBI substrate eTACaplitpl nig karbeBa©jnUvplitpl nig Gg;sIumBIbNþúM Gg;sIum-plitpl (Enzyme-product) . ehtuenH GRtaénRbtikm μRtUv)ankMNt; edayel,OnEdl substrate RtUv)anbMElgeTACaplitpl nig plitplRtUv)anbeBa©j . ehtudUecñH GRtaénRbtikm μCab;Tak;TgeTAnwgPaBman (availability) énGg;sIum .

rUbTI 20-2 GRta b¤ el,On (V) énRbtikmμpÞúyeTAnwgkMhab;rbs; Substrate [S]

4- bNþúM ENZYME-SUBSTRATE (ENZYME-SUBSTRATE COMPLEX)

RbtikmμRtLb;mkvijmYyes‘rIxageRkambgðajdMNak;karénRbtikm μEdlsMrYledayGg;sIum . dMNak;karTImYy ¬B’Nexov¦ Cab;Tak;TgeTAnwgkarRbQmKñaénGg;sIumCamYynwg substrate rbs;va nig karbegáItnUv Enzyme-substrate complex .

Substrate concentration

Rat

e of

rea

ctio

n (V

elo

city

)

Substrate concentration

Rat

e of

rea

ctio

n (V

elo

city

) Vmax

Reaction reaches a maximum rate

Step I Step II Step III Step IV

E + S ⇄ ES ⇄ ES* ⇄ EP ⇄ E + P



EpñkénGg;sIumEdlP¢ab;V\CamYynwg substrate RtUv)aneKehAfaCa TIkEnøgskmμ (Active site). eyIgeXIjfalkçN³énTIkEnøgskmμvasMxan;cMeBaHmuxgarrbs;Gg;sIum nig manlkçN³TUeTAdUcxag eRkam ³ TIkEnøgskmμrbs;KWCa ehae):A b¤ esa (Pockets or clefts) enAkñúgépÞrbs;Gg;sIum . bgÁúM naTI R enAkñúgTIkEnøgskmμEdlCab;Tak;TgeTAnwgkarsMrYlRtUv)aneKehAfa Catalytic group . rUbragénTIkEnøgskm μvaRtUvKñaeTAnwgrUbragrbs; substrate . KWfa Substrate RtUvKñaya:g xøaMgeTAnwgTIkEnøgskmμrbs;Gg;sIum . Gg;sIumTaj nig cab; substrates rbs;vaedayGMeBIeTAvijeTAmkexSay nig minEmnsm<½n§ kUva:Lg; . bgÁúMnaTI R Cab;Tak;TgeTAnwgkarcg substrate nig minmankarsMrYledaycaM)ac; begáItnUv TIkEnøgcgP¢ab; . TMrg;énTIkEnøgskmμ (Conformation) kMNt;nUvPaBBiess (Specificity) rbs;Gg;sIum BIeRBaHmanEt substrate EtmYyKt;EdlRtUveTAnwgTIkEnøgskm μEdlnwgRtUveRbIenAkñúgRbtikmμ . KMrUemesanigkUnesa (Lock-and-key model) énskm μPaBrbs;Gg;sIum RtUv)anbgðajenAkñúgrUb TI 200-3a RtUv)anerobcMeday Emil Fischer enAkñúgqñaM 1894 . enAkñúgeBlenaH eK)anKitfa Substrate cUledayRsYleTAkñúgkEnøgdUcCaEpñkmYyénEl,gpÁúMrUbPaB (jigsaw puzzle) b¤ kUnesa cUl eTAkñúgemesa .

rUbTI 20-3



sBVéf¶eyIgdwgfa RbUetGIunKWCam:UelKulEdlmanlkçN³bt;Ebn . kareFIV[ Daniel E.

Koshland, Jr. esñIeLIgnUvd¾sμ úRKsμajénviFIEdlGg;sIum nig substrate manGMeBIeTAvijeTAmk . KMrUenH )anesñIeLIgenAqñaM 1958 )anRtUveKehAfaCa Induced fit model ¬rUbTI 20-3b¦ . enAkñúgKMrUenH TIkEnøgskm μrbs;Gg;sIumminEmnCakEnøgmaM (rigid pocket) Edl substrate dak;cUledayRtwmRtUv CaCag vaKWCakEnøgEdlmanlkçN³bt;Ebn EdlRbhak;RbEhleTAnwgrUbragrbs; substrate . enAeBl substrate cUleTATIkEnøg (pocket) TIkEnøgskmμ { Tn;} edayxøÜnÉgenACMvij substrate . karenHbegáItPaBRtUvKñarvag Enzyme-substrate )anya:gl¥\ex©aH . 5- PaBBiessénbNþúM ENZYME-SUBSTRATE

cMeBaHGMeBIeTAvijeTAmk enzyme-substrate EdlekIteLIg épÞrbs;Gg;sIum nig substrate

RtUvEtbMeBjKña (complementary) . vaenHKWCatMrUvkarcMeBaHPaBRtUvKñaCaBiessEdlkMNt;faetI Gg;sIumnwgcgP¢ab;nwg substrate BiessNamYy nig GnuvtþRbtikm μKImI . PaBBiessrbs;Gg;sIum (Enzyme specificity) KWCalT§PaBrbs;Gg;sIumkñúgkarP¢ab;eTAnwg EtmYy b¤ ticbMput nig ehtudUecñH vasMrYlEtRbtikmμEtmYyKt; . edIm,IbgðajPaBBiessrbs;Gg;sIum cUrBicarNaBIRbtikmμxageRkam ³ pÞúymkvij Methylurea eTaHbICamanrcnasm<½n§dUceTAnwg urea k¾eday k¾buEnþvaminTTYlrgGMeBI rbs; Urease eT .

minEmnGg;sIumTaMgGs;mankMriténPaBBiessdUcKñaeT . manPaBBiessrbs;Gg;sIum bYnfñak; EdRtUv)aneKBinitüeXIj . PaBBiessdac;xat (Absolute Specificity) Gg;sIummYyEdlsMrYlRbtikm μén substrate EtmYy manPaBBiessdac;xat . Aminoacyl tRNA synthetases manPaBBiessdac;xat . vanimYy²RtUvEt dak;P¢ab;GasIutGamIneNRtwmRtUvcMeBaHm:UelKulRtwmRtUvén tRNA . RbsinebIGasIutGamIneNmin RtwmRtUv RtUv)anP¢ab;eTAnwg tRNA vanwgkMhuskñúgkarbEnßmeTAelIExS peptides edaybegáIt)an CaRbUetGIunGsmtßPaB (nonfunctional protein) .

Urease H2N—CO—NH2 + H2O → CO2 + 2 NH3

Urease H2N—CO—NH—CH3 + H2O → No reaction



PaBBiessCaRkum (Group Specificity) Gg;sIummYyEdlsMrYldMeNIrkarEdlCab;Tak;TgeTAnwg m:UelKulEdldUcKñaEdlmanbgÁúMmuxgardUcKñaman PaBBiessCaRkum . Hexokinase KWCaGg;sIumEdlman lkçN³BiessCaRkum EdlsMrYlnUvkarbEnßmnUvbgÁúM Phosphoryl eTAelI glucose kñúgdMNak;karTImYy én

Glycolysis . Hexokinase k¾bEnßmbgÁúM Phosphoryl eTAelIBYksárEdlman 6GatUmkae)anepSgeTot. PaBBiessénkarcgP¢ab; (Linkage Specificity) Gg;sIumEdlsMrYlkarbegáIt b¤ karpþac;Etsm<½n§ mYycMnYnEdlmanenAkñúgm:UelKulEdlmanPaBBiessénkarcgP¢ab; . Proteases dUcCa trypsin,

chymotrypsin nig elastase KWCaGg;sIumEdlGIuRdUlIsedayeRCIserIsEtsm<½n§ peptide . dUecñH Gg;sIum TaMgenHman PaBBiessénkarcgP¢ab; . PaBBiessénrcnasm<½n§KImIkñuglMhr (Stereochemical Specificity) Gg;sIummYyEdlGacEjk GIusUEmrBIrEdlmanlkçN³qøúHkBa©k;Kña man PaBBiessénrcnasm<½n§KImIkñúglMhr . PaKeRcInénGg;sIum rbs;Ba’gÁkaymnusSbgðajBIPaBBiessénrcnasm<½n§KImIkúñglMhr . GaRs½yEteyIgeRbIEt D-sugars

nig L-amino acids Gg;sIumEdlCab;Tak;TgkñúgkareraKvinicä½y nig emtabUlIsTTYlsÁal;EtGIusUEmr BiessTaMgenaHEtb:ueNÑaH . 6- sPaBbeNþaHGasnñ nig karbegáItplitpl (THE TRANSITION STATE AND PRODUCT FORMATION)

etIkarcg Enzyme-Substrate bNþal[manRbtikmμKImIelOnya:gdUcemþc ? cMeLIyeday RtwmRtUveTAnwgsMNYrenHRbEhlCaxus²KñacMeBaHbNþúM Enzyme-Substrate nimYy² nig Cabnþ eyIgyl; GMBIynþkarBitR)akdénkarsMrYlcMeBaHGg;sIumtictYcNas; . eTaHbICadUecñH eyIgGacBinitüemIlTidæPaB TUeTAénGMeBIeTAvijeTAmkénbNþúM Enzyme-Substrate EdlbNþal[manGRtaRbtikmμekIneLIg nig karbegáItnUvplitpl . edIm,IeFIVkarenH eyIgRtUvEtBinitüemIldMNak;karRbtikm μ Enzyme-Substrate

edayepþatkarykcitþTukdak;eTACMhanEdlenAsl;BImanB’Nexov ³

enAkñúgEpñkTI 4 eyIg)anBinitüBIRBwtþikarN_éndMNak;TI I EdlGg;sIum nig substrate manGMeBIeTA vijeTAmkedIm,IbegáItCabNþúM Enzyme-sunstrate . enAkñúgEpñkenHeyIgnwgBinitüemIlBIRBwtþikarN_Edl naM[mankarbegáItnUvplitpl . eyIg)anBiB’NnarYcmkehIyfaGg;sIumCam:UelKulbt;Ebn (flexible

molecule) ehIy substrate k¾mannUvkMriténPaBbt;Ebnrbs;vaEdr . karmanGMeBIeTAvijeTAmkCabnþ bnÞab;rvagGg;sIum nig substrate pøas;bþÚrrUbrag b¤ TItaMgrbs; substrate enAkñúgrebobEdlTMrg;én

Step I Step II Step III Step IV

E + S ⇄ ES ⇄ ES* ⇄ EP ⇄ E + P



m:UelKul (Molecular configuration) ElgsßitesßreTotehIy ¬dMNak;TI II¦ . enAkñúgsPaBenH sPaBbeNaþaHGasnñ (Transition state) rUbragrbs; substrate RtUv)anxUcxat ¬BIeRBaHEtGMeBIeTAvij eTAmkrbs;vaCamYyGg;sIum¦eTACaTMrg;kNþal (intermediate) EdlmanTidæPaBrbs;TaMg substrate nig plitplbBa©b; . sPaBbeNþaHGasnñenH Cabnþ pþl;nUvkarbMElgrbs; substrate eTACaplitpl ¬dMNak;TI III¦ . plitplenAEtCab;CamYynwgGg;sIumkñúgmYyry³eBlxøI bnÞab;mksßitkñúgdMNak;karTI IV plitpl nig Gg;sIumEbkecjBIKña edayTukGg;sIum[enAdEdlminmankarpøas;bþÚr . etIkarpøas;bþÚrsPaBbeNþaHGasnñRbePTNaGacekItmaneLIgenAkñúg substrate EdlnwgeFIV[ Rbtikmμbnþya:gelOn ? 1- Gg;sIumGacdak;sMBaFeTAelIsm<½n§ nig enATIenaH sMrYlnUvkarpþac;sm<½n§ . cUrBicarNaGMBI kareFIVGIuRdUlIsén sucrose (saccharase) edayGg;sIum sucrase . Gg;sIumenHsMrYlkareFIVGIuRdUlIs én disaccharide sucrose [eTACa monosaccharides BIrKW glucose nig fructose . karbegáItnUvbNþúM enzyme-substrate ¬rUbTI 20-4a nig b¦ bNþal[mankarpøas;bþÚrrUbragrbs;Gg;sIum . Cabnþ karenH GacTajsn§wg b¤ dak;sMBaFeTAelIsm<½n§mYyrbs; substrate . sMBaFrebobenHeFIV[exSay sm<½n§ edaybNþal[vadac;RsYlCagkñúgeBlGvtþmanrbs;Gg;sIum . karenHRtUv)anbgðajenAkñúgrUbTI 20-4c CaExSekagénsm<½n§ O-glycosidic rvag Fructose nig Glucose . enAkñúgsPaBbeNþaHGasnñ substrate manTMrg;m:UelKuldUcnwgTaMg disaccharide ¬substrarte edIm¦ nig monosaccharides

TaMgBIr ¬plitplcugbBa©b;¦ . edayc,as;las; sMBaFEdldak;eTAelIsm<½n§eFIV[vaexSay nig RtUvkar famBltickñúgkarpþac;sm<½n§edIm,IbegáItplitpl ¬rUbTI 20-4d nig e¦ . karenHk¾man\T§iBleTAelI karbegáInel,OnénRbtikm μ . 2- Gg;sIumGacsMrYlRbtikmμedaykarnaMRbtikmμBIrEdlmanPaBRbhak;RbEhlKñaxøaMg nig kñúgTisedARtwmRtUvcMeBaHRbtikmμEdlekIteLIg . \LÚvcUrBicarNa BIRbtikm μ dehydration rvag glucose nig fructose edIm,IbegáItCa sucrose (saccharose) rUbTI 20-5a . sárnimYy²manbgÁúM hydroxyle 6 EdlGacTTYlrgnUvkarrYmKña (condensation) edIm,IbegáItCa disaccharide . b:uEnþeKal bMNgKWedIm,IbegáIt sucrose minEmn disaccharides epSgeToteT . eTAtamkarRbQmKñaedayécdnü énm:UelKul vaman»kasmYykñúgcMeNam»kas 25 Edlm:UelKulTaMgBIrnwgRbQmmuxKñakúñgTisedA RtwmRtUvedIm,IbegáItCa sucrose . PaBGacekIteLIg (Probability) Edlm:UelKulTaMgBIrnwgmanRbtikm μ CaFm μtaekIteLIgticCagenHq¶ayBIeRBaHlkçx½NÐepSgkñúgkarbEnßmeTAelITisedARtUvEtRKb;RKan;sMrab; RbtikmμekIteLIg . ]TahrN_ sItuNðPaBdgxøÜn karRbQménmU:elKuleRcInbMput nwg minmancMnYnRKb; RKan;nUvfamBledIm,ICMnHfamBlEdleFIV[skmμ (energy of activation) sßitenAkñúgTisedARtwmRtUv



k¾eday . Gg;sIumGacsMrYlRbtikm μedaykarnaMm:UelKulBIrcUlCitKñakñúgkarerobcMRtwmRtUv ¬rUbTI 20-5b¦ EdlenATIenaH edaykarCMrujnUvbgÁúMEdlRtUvmanRbtikmμrbs;m:UelKulTaMgBIrrYmcUlKñakñúgsPaB beNþaHGasnñ nig karCMrujel,OnRbtikm μya:gxøaMg . 3- TIkEnøgskm μrbs;Gg;sIumGaceFIV[ERbRbYl pH rbs;mIRkUbrisßan (Microenvironment) enA CMuvij substrate . edIm,IsMerckarenH Gg;sIumGac Ca]TahrN_ bMerICaGñkpþl; b¤ cab;yk H+ . CalT§pl vanwgmankarpøas;bþÚrnUv pH enAkñúgtMbn;CMuvij substrate edayBMumaneFIV[b:HBal;dl; pH

FmμtaenAkñúgTIepSgeTotrbs;ekasika . rUbTI 20-5

Gg;sIumGaceFIV[cuHfamBlEdleFIV[skmμEdlRtUvkarsMrab;[manRbtikm μedaykarcgP¢ab; [enACitKñabMput nig kñúgkartMrg;TiedARtwmRtUv .



7- COFACTORS AND COENZYMES

enAkñúgEpñkTI 19-7 eyIg)aneXIjfaRbUetGIunmYycMnYn (ex. Conjugated proteins) RtUvkar bNþúMminEmnRbUetGIun (prosthetic group) edIm,IbMeBjmuxgar . PaBdUcKñaKWvaBitrbs;Gg;sIummYycMnYn. EpñkEdlCaRbUetGIunrbs;Gg;sIumRtUv)aneKehAfaCa Apoenzyme nig bgÁúMminEmnRbUetGIun RtUv)aneK ehAfaCa Cofactor . edayrYmKña BYkvabegáIt)anCaGg;sIumskmμ (active enzyme) RtUv)aneKehAfaCa Holoenzyme . CaTUeTA Cofactor KWCaGIuy:ugelah³ EdlRtUvEtP¢ab;CamYyGg;sIumedIm,IrkSaTMrg; RtwmRtUvénTIkEnøgskmμrbs;Gg;sIum ¬rUbTI 20-6¦ . enAeBl Cofactor cgCamYy nig enAeBl TIkEnøgskm μsßitenAkñúgTMrg;RtwmRtUv Gg;sIumGaccgP¢ab;CamYynwg substrate nig sMrYlRbtikmμ .

rUbTI 20-6

Gg;sIummYycMnYnRtUvkarkarcgP¢ab;CabeNþaHGasnñnUv Coenzyme . karcgrebobenHCaTUeTA RtUv)ansMrYledayGMeBIeTAvijeTAmkexSaydUcCasm<½n§GIuRdUEsn . Coenzymes KWCam:UelKulsrIragÁ EdlCaTUeTAeRbICaGñkdwknaMeGLikRtug b¤ bgÁúMKImI . enAkñúgRbtikm μKImI BYkvaGacCaTaMgGñkpþl;bgÁúMnaTI eTA[ substrate b¤ eRbICaGñkTTYlbgÁúMnaTIEdlRtUv)andkecjBI substrate rUbTI 20-7 . Cajwkjab; Coenzymes manvItamInEdlRtUv)anEksMrYl (modified vitamins) CaEpñkén rcnasm<½n§rbs;BYkva . vItamInKWCasarFatusrIragÁEdlRtUvkarcaM)ac;enAkñúgGaharkñúgcMnYntictYcEt



b:ueNÑaH . kñúgcMeNamBYkvItamInrlaykñúgTwk manEtvItamIn C minrYmCamYy Coenzyme ¬taragTI 20-1¦ segçbnUv Coenzymes mYycMnYn nig vItamInEdlrlaykñúgTwkEdlplitBYkva . Nicotinamide adenine dinucleotide (NAD+) RtUv)anbgðajenAkñúgrUbTI 20-8¦ KWCa ]TahrN_én Coenzyme Edlmansar³sMxan;xøaMgbMputenAkñúgRbtikm μGuksIutkmμéndMeNIrkarRbmUl famBlrbs;ekasika (Cellular energy-harvasting processes) . m:UelKul NAD+ manlT§PaBkñúgkar TTYl Hydride ions ¬GatUmGIuRdUEsnEdlmaneGLikRtugBIr¦ BI substrate énRbtikm μRbmUlfamBl . Substrate TTYlrgGuksIutkmμ nig Epñkén NAD+ EdlmanRsLayBI vItamIn Niacin TTYlrger:dukmμ køayeTACa NADH . CabnþeTAeTot NADH pþl;nUv hydride ion eTA[GñkTTYlTImYyenAkñúgExSén kardwknaMeGLikRtug . karenHpþl;[vijnUv NAD+ nig pþl;eGLikRtugsMrab;karbegáIt ATP famBlKImI EdlRtUvkaredayekasika . k¾RtUv)anbgðajenAkñúgrUbTI 20-8 KWCaGñkdwknaM Hydride ion NAD+ nig GñkdwknaMGatUmGIuRdUEsn FAD . vaTaMgBIrRtUv)aneKeRbIenAkñúgRbtikmμGuksIutkmμ-er:dukkmμ EdlRbmUl famBlBIekasika . mindUceTAnwg NADH nig FADH2 , NADPH eRbICaGñkkat;bnßyfamBl (reducing power) cMeBaHekasikaedaykarpþl;nUv hydride ions enAkñúgRbtikmμCIvKImI . dUceTAnwg NAD+ , NADP+ manRsLayBI Niacin . FDA ekIteLIgBIvItamIn Ribflavin .

rUbTI 20-7



taragTI 20-1 vItamInrlaykñúgTwknig coenzymes EdlBYkvasmasPaBénrcnasm<½n§

Vitamins Coenzymes Function Thiamine (B1) Thiamine pyrophosphate Decarboxylation reactions Riboflavin (B2) Flavin mononucleotide (FMN) Carrier of Hydrogen atom Flavin adenine dinucleotide (FAD) Niacin (B3) Nicotinamide adenine dinucleotide

(NAD+) Carrier of hydride ions

Nicotinamide adenine dinucleotide phosphate (NADP+)

Pyrodxine (B6) Pyridoxal phosphate Carrier of amino and carboxyl Pyridoxamine phosphate groups Cyanocobalamine (B12) Deoxyadenosyl cobalamine Coenzyme in amino acid metabolism Folic acid (B9) Tetrahydrofolic acid Coenzyme for 1-C transfert Pantothenic acid (B5) Coenzyme A Acetyl group carrier Biotin Biocytin Coenzyme in CO2 fixation Ascorbic acid Unknown Hydroxylation of proline and lysine in collagen

rUbTI 20-8 rcnasm<½n§én Coenzymes bI

8- \T§iBlénbriyakasCMuvij (ENVIRONMENTAL EFFECTS)

k- \T§iBl pH PaKeRcInénGg;sIumskm μEtkñúgcenøaH pH tUcxøaMgNas; . sIutU)aøs;ekasikaman pH = 7 nig PaKeRcInénGg;sIumrbs;sIutU)aøs;bMeBjmuxgarenAkñúgRbsiT§PaBGtib,rmaenAkñúg pH = 7 enH . karEbg EckénGRtaRbEhl (relative rate) EdlGg;sIumsIutU)aøs;sMrYlRbtikm μBiessrbs;vamYyRbQmeTAnwg pH RtUv)anpþl;[enAkñúgrUbTI 20-9 .

rUbTI 20-9 \Ti§Blrbs; pH eTAelIGRtarbs;Gg;sIum

pH EdlGg;sIumbMeBjmuxgarCaFm μtaRtUv)aneKehAfaCa pH optimum . kareFIVsUluysüúg EdlmanlkçN³)asxøaMg b¤ GasIutxøaMgbnßynUvGRtaénRbtikmμ . dUcEdl)anBiPakSaenAkñúgEpñkTI 19-10 enAkñúg pH xøaMgRCul Gg;sIumCaFm μta)at;bg;nUvTMrg;CIvsaRsþskmμrbs;va (biologically active

conformation) nig RtUv)anxUcxat . karenHBIeRBaHkarpøas;bþÚr pH eFIV[xUcnUvkMritkarEbkCaGIuy:ug

5 7 9

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rbss;bgÁúM R rbs;GasIutGamIneNenAkñúgExSRbUetGIun ¬Gg;sIum¦ k¾dUcCakarXøatKñaEdlBYkvaGaccgKña tamsm<½n§GIuRdUEsn . k¾dUcKñaenHEdr GMeBIeTAvijeTAmkenHGaceFIV[xUcya:gxøaMgnUvTMrg;rbs;RbUetGIun TaMgRsug dUecñH Gacmankarpøas;bþÚrenAkñúgbgÁúM R énTIkEnøgskm μrbs;Gg;sIumEdlbMpøajnUvlT§PaBkñúg karbegáItbNþúM enzyme-substrate . eTaHbICasIutU)aøs;rbs;ekasika nig vtßúravEdleRsacRsBekasikaman pH EdlRtUv)anRtYtBinitü edayRbugRby½tñ dUecñHvasßitenAEtRbEhl 7 b:uEnþvak¾manbriyakasenAkñúgsBa’gÁkayEdlGg;sIumRtUv EtbMeBjmuxgarenAkñúg pH EdlXøatq¶ayBI 7 Edr . QutRbUetGIun (Protein sequence)

)anbgðajfaGacrkSanUvrcnasm<½n§RtwmRtUvkñúglMhr (the proper three-dimensional structure)

enAeRkamlkçx½NÐ pH Tab b¤ x<s;RCul (extreme) . ]TahrN_ pH rbs;RkBHRbEhl 2 EdlCalT§pl énkarbeBa©j HCl BIekasikaBiessrbs;RkBH . Gg;sIumrMlayRbUetGIun (Proteolytic digestive

enzyme) Pepsin RtUvEtbMEbkRbUetGIunedayRbsiT§PaBenAkñúg pH = 2 enH . kñúgkrNIrbs; Pepsin

Gg;sIumbgðajkúñgrebobenHfavaGacrkSanUvrcnasm<½n§TI 4enAkñúg pH = 2 nig skmμxøaMgkñúgkarsMrYlén kareFIVGIuRdUlIsBYk peptides EdlGacRtUv)anbMEbkenAkñúg pH TabxøaMg . dUecñH Pepsin man pH

optimum = 2 . enAkñúgrebobdUcKña Gg;sIumbMEbkRbUetGIunmYyepSgeTot Trypsin bMeBjmuxgareRkamlkçx½NÐ én pH x<s;EdleKCYbenAkñúgeBaHevontUc . TaMg Pepsin nig Trypsin kat;pþac;nUvsm<½nß peptides eday ynþkardUcKñaya:gxøaMg b:uEnþQutGasIutGamIneN)anbgðajenAeBlEdlBYkvasßitesßr nig skm μenAkñúg briyakasxus²KñaxøaMg . sBa’gÁkay)aneRbIkarsMrbsMrYl (adaptation) rbs;Gg;sIumcMeBaHbriyakasepSg²edIm,IkarBar xøÜnvaRbqaMgeTAnwgynþkarkarBarEdlbMpøajxøÜnva . enAkñúgsIutU)aøs;rbs;ekasikaKWPaKl¥it (organelles)

EdleKehAfa Lysosomes . Christian de Duve EdlCaGñkrkeXIj lysosomes enAkñúgqñaM 1956 edayehABYkvafaCa {fg;GtþXat} (suicide bags) BIeRBaHBYkvaCafg; (vesicules) RtUv)ancgP¢ab;nwg PñasekasikaEdlmanRbEhl 50 RbePTxus²nUv Hydrolases b¤ Gg;sIumGIuRdUlIs . eKalbMNgrbs; Gg;sIumTaMgenHKWedIm,IbMEbkm:UelKulFM²eTACam:UelKultUc²EdlmansarsMxan;sMrab;Rbtikm μRbmUl famBl . GaRs½yehtuenH mYycMnYnénGg;sIumenAkñúg lysosomes GacbMEbkRbUetGIun[eTACa GasIutGamIneN nig BYk Polysaccharides epSg²eTot[eTACa monosaccharides . ekasikamYy cMnYnénRbB½n§PaBsaMu (immune defense sytem) RKbeTAelIsarFatuQøanBanBIxageRkA (foreign

invaders) dUcCa )ak;etrI nig virus . bnÞab;mkBYkvaeRbIGg;sIumGIuRdUlIsenAkñúg lysosomes edIm,IbMEbk



nig bMpøajBYksarFatuQøanBanTaMgenaH nig eRbIBYk simple sugars GasIutGamIneN nig lIBIdEdl plit)anCaRbPBfamBl . GIVEdlnwgekIteLIg RbsinebIGg;sIumGIuRdUlIsrbs; lysosome RtUv)anbeBa©jedayécdnüeTA kñúgsIutU)aøsrbs;ekasika ? edayBitR)akd lT§plnwgKWCakarbMpøaj(destruction) én macromolecules rbs;ekasika nig karsøab;rbs;ekasika . GaRs½yedayeRKaHfñak;enH ekasikaRbmUl famBlCaeRcInkñúgkarrkSanUvbUrNPaBénPñasrbs; lysosomes . ynþkarkarBarCabEnßmTak;TgeTAnwg karBitEdlfaGg;sIumrbs; lysosome bMeBjmuxgarCaFm μtaenAkñúg pH GasIut (pH = 4,8). RbsinebI Gg;enHmYycMnYnelcecjBI lysosome b¤ RbsinebI lysosomes Ebkedayécdnü pH rbs;sIutU)aøs; (7,0-7,3) eFIV[BYkva)at;bg;skm μPaB . x- \T§iBlsItuNðPaB Gg;sIumxUcya:gelOn RbsinebIsItuNðPaBrbs;sUluysüúgekIneLIgx<s;Cag 37OC b:uEnþBYkva enAEtsßitesßr (stable) eTAkñúgsItuNðPaBTabxøaMg. cMeBaHehtuplenH KWfasUluysüúgrbs;Gg;sIum EdlRtUv)aneKeRbIsMrab;karviPaKKøInikRtUv)aneKsþúkTukenAkñúgTUrTwkkk (refrigerators or freezers) mun eBleRbI . rUbTI 20-10 bgðajBI\T§iBlrbs;sItuNðPaBeTAelIRbtikmμEdlsMrYl nig minsMrYleday Gg;sIum . GRtaénRbtikmμEdlminsMrYledayGg;sIumekIneLIgedaynwg (steadily) rYmCamYynwgkarekIn eLIgénsItuNðPaBBIeRBaHmankarRbQmKñaeRcInEdlekIteLIgCamYynwgfamBlRKb;RKan;kñúgkarCMnH famBlraraMgcMeBaHRbtikmμ (energy barrier). GRtaénRbtikmμEdlsMrYledayGg;sIumk¾ekIneLIgCa mYykarekIneLIglμmnUvsItuNðPaBBIeRBaHvamancMnYnénkarRbQmekIneLIgrvagG;sIum nig substrate. b:uEnþmuneBlyUr sItuNðPaBEdlekIneLIgcab;epþImbegáInnUvfamBlrMj½rénsm<½n§enAkñúgGg;sIum. cug bBa©b;sm<½n§CaeRcIn nig GMeBIeTAvijeTAmkexSayRtUv)andac;EdlGg;sIumRtUv)anxUcxat nig Rbtikm μRtUv Qb; .



rUbTI 20-10 \T§iBlsItuNðPaBeTAelIRbtikmμ

edaysarkarkMedA Gg;sIum nig RbUetGIunepSgeTotxUcnUvrcnasm<½n§kñúglMhrEdllkçN³Biess rbs;BYkva nig dUecñHxUcnUvskmμPaBrbs;BYkva ekasikaminGacenAkñúgsItuNðPaBx<s; . GaRs½yeday ekasikaminGacbMeBjmuxgaredayKμanskmμPaBRtwmRtUvrbs;Gg;sIum kMedAKWCameFüa)ayEdlman RbsiT§PaBénkarsMGatemeraK (sterilizing) cMeBaHsMPar³evC¢saRsþ nig sUluysüúgsMrab;cak;bBa©ÚleTA kñúgxøÜn b¤ etsþKøInik . eTaHbICa]bkrN_GacRtUv)anrMgab;emeraKedaysarkMedAs¶Üt (160OC) Edl Gnuvtþya:gehacry³eBlBIrema:genAkñúgLsMgYtk¾eday b:uEnþ karrMgab;emeraKeday Autoclave

elOnCag nig KYr[TucitCag . Autoclave sMGageTAelIeKalkarN_énsMBaFcMhayTwk . xül;RtUv)an )aj;[ecjeRkAbnÞb; nig cMhayTwkeRkamsMBaFRtUv)an)aj;cUleTAkñgbnÞb;rhUtdl;sMBaFekIneLIg dl; 2 atmospheres . sMBaFbgá[mansItuNðPaBcMhayTwk Edlman 100OC enAkñúgsMBaFbriyakas nig ekIneLIgdl; 120OC . kñúgry³eBl 20 naTI RKb;)ak;etrI nig virusRtUv)ansMlab; . karenHKWCa meFüa)ayEdlmanRbsiT§PaBkñúgkarbMpøajBYks<½rEdlFn;nwgkMedAxøaMg . EdlbegáIteLIgeday)ak;etrI CaeRcInénkarcab;GarmμN_enAkñúgKøInik . )ak;etrITaMgenHrYmman BYk Bacillus nig Clostridium EdlTTYl xusRtUvenAkñúgPaBminl¥enH nig CMgWbgá[søab;dUcCa anthrax, gas gangrene, tetanus, nig karBulGaharEdlbgáeday Botulism . b:uEnþminEmnRKb;Gg;sIumeTEdlRtUv)at;skm μPaBedaysarkMedA ehIysUm,IEtsMBaFx<s;k¾ eday. )ak;etrImYycMnYnrs;enAkñúgTIkEnøgGNþÚgFüÚgf μcas; EdlCaFmμtaRtUv)aneqH . mYycMnYneTot rs;enAkñúgCMerAmhasmuRTenAeBlsItuNðPaB nig sMBaFx<s;xøaMg . b:uEnþ )ak;etrIepSgeTot lUtlas; enAkñúgRbPBTwkekþA (hot spring) én]TüaCati Yellowstone EdlenATIenaH lUtlas;enAkñúgsItuNðPaB Ek,rcMNucrMBuHrbs;Twk . BYksrIrsgÁmanCIvitTaMgenHrYmCamYyGg;sIumrbs;BYkva rs;enAeRkamlkçx½NÐ

10 30 50 70 Temperature (OC)

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10 30 50 70 Temperature (OC)

Vel

ocity



EdlminKYreCO BIeRBaHQutGasIutGamIneNénRbUetGIuneFIVtamrcnasm<½n§EdlsßitesßrenAkñúgsItuNðPaB EdlminKYr[eCOenH . 9- karEktMrUvénskmμPaBGg;sIum (REGULATION OF ENZYME ACTIVITY)

enAkñúgcMeNamviFIsMxan;EdlGg;sIumxusBIkatalIkrminEmnCIvsaRsþd¾éT KWfaskmμPaBrbs; Gg;sIumCajwkjab;RtUv)anEktMrUvedayekasika . vamanehtuplCaeRcIn sMrab;karRtYtBinitümuxgarrbs; Gg;sIumenH . mYycMnYnCab;Tak;TgeTAnwgkarBicarNarénfamBl . RbsinebI ekasikaGs;famBlKImI vanwgsøab; . GaRs½yehtuenH ynþkarCaeRcInmansMrab;rkSaTukfamBlekasika . cMeBaHehtuenH vaman karx¢Hx¢ayfamBlkúñgkarplitGg;sIumRbsinebI substrate minman . dUcKñaenHEdr RbsinebIplitpl énRbtikmμEdlsMrYledayGg;sIummanvtþmanedayelIs vamankarx¢Hx¢ayfamBlcMeBaHGg;sIumkñúgkar bnþsMrYlRbtikm μ enATIenHmankarplitnUvplitplmincg)anCaeRcIn . dUcEdleTIb)anniyay vamanehtuplCaeRcInkñúgkarEktMrUvskmμPaBGg;sIum vamanynþkarCa eRcInsMrab;karEktMrUvrebobenH . ynþkarsamBaØCageKbMputKWedIm,IbegáItGg;sIumEtenAeBlmanvtþman rbs; substrate Etb:ueNÑaH . ynþkarenHRtUv)aneRbIeday)ak;etrIedIm,IEktMrUvGg;sIumEdlRtUvkarkñúgkar bMEbknUvsárepSg²edIm,ITTYl)annUv ATP sMrab;kargarekasika . )ak;etrIminmankarRtYtBinitüeTAelI briyakasCMuvij b¤ eTAelIGIVEdlCaRbPBGahar ¬RbsinebIman¦ Gacman . vanwgKWCakarx¢Hx¢ayya:g eRcInnUvfamBlkñúgkarplitRKb;Gg;sIumEdlRtUvkaredIm,IbMEbkRKb;sárEdlGacman . dUecñH)ak;etrIsnSM famBledaykarplitGg;sIumEtenAeBlman speicific sugar substrate Edlman . ynþkarepSgeTot sMrab;kar EktMrUvskmμPaBrbs;Gg;sIumrYmmankareRbI allosteric enzymes, feedback inhibition nig karplitén zymogens . \LÚveyIgBinitüemIlynþkarEktMrUvmYycMnYnedayBisþar . k- Allosteric Enzymes

RbePTmYyénkarEktMrUvGg;sIumCab;Tak;TgGg;sIumEdlmanTIkEnøgcgP¢ab;eRcInCagmYy . Gg;sIumTaMgenH ¬RtUv)aneKehAfa Allosteric enzymes¦ mann½yfa {TMrg;epSgeTot} KWCaGg;sIum EdlTIkEnøgskmμrbs;vaGacRtUv)anxUcedaysarkarcgénm:UelKultUc EdleKehAfa effector

molecule . dUcEdl)anbgðajenAkñúgrUbTI 20-11 karcgén Effector eTAnwgTIkEnøgcgrbs;vaeFIV[ eGayxUcrUbragénTIkEnøgskm μrbs;Gg;sIum . lT§plGacRtUv)anbMElgTIkEnøgskmμeTACa TMrg;minskmμ (inactive configuration) EdlKWCa Negative allosterism b¤ bMElgTIkEnøgskmμeTACa TMrg;skkmμ (active configuration) KWCa Positive allosterism . enAkñúgkrNITaMgBIr karcgP¢ab;nUv effector molecule EksMrYlskmμPaBGg;sIumedaykarkMNt;faetIvanwg skm μ b¤ Gkm μ .



enAkñúgCMBUkeRkayEdlnwgmkdl;xagmux eyIgnwgcab;epþImkarsikSarbs;eyIgnUvpøÚvemtabUlIs. pøÚvemtabUlIsKWCaes‘rIénRbtikmμCIvKImIEdleFIV[sMerc)an nUvkarbMEbk b¤ sMeyaKm:UelKulCIvsaRsþ mYy b¤ eRcIn . cUrBicarNaBIpøÚv Glycolysis EdldMNak;kardMbUgénkarbMEbk Carbohydrate kñúgkar beBa©jfamBlEdlGacRtUv)aneRbIedayekasika (Adenosine triphosphate : ATP) . pøÚvrebobenH RtUvEtEksMrYltMrUvkarrbs;sBa’gÁkay . enAeBlRtUvkarfamBleRcIn Rbtikm μénpøÚvnwgekIteLIgya:g elOn edaybeBa©jfamBlya:geRcIn . b:uEnþRbsinebItMrUvkarTab RbtikmμTaMgenHnwgfycuH edaybegáIt famBltic (ATP) . RbtikmμTI 3 én glycolysis KWCakarbBa¢ÚnbgÁúM Phosphoryl BI ATP eTA[ Fructose-6-

phosphate . edaybgðajTIenH RbtikmμRtUv)ansMrYledayGg;sIumeQμaH Phosphofructokinase . Phosphofructokinase Fructose-6-phosphate → Fructose-1,6-biphosphate

skmμPaB Phosphofructokinase rYseTAnwgTaMg Positive nig Negative allosterism . GaRs½yedayehtuenH enAeBl ATP maneRcIn sBaØaEdlsBa’gÁkaymanfamBlRKb;RKan; vaP¢ab;eTAnwg TIkEnøgcgP¢ab; effector enAelI Phosphofructokinase . karenHraraMgskm μPaBrbs;Gg;sIum nig dUecñH eFIV[yWtpøÚv glycolysis TaMgmUl . PaBsMbUrén AMP (Adenosine monophosphate) EdlCa Precursor rbs; ATP KWCaskçIPaBbgðajfasBa’gÁkayRtUvkareFIV ATP edIm,I[mankarpþl;famBl RKb;RKan; . enAeBl AMP cgP¢ab;eTAnwgTIkEnøgcg effector enAelI Phosphofructokinase skm μPaB rbs;Gg;sIumekIneLIgedaybegáInel,OnRbtikmμ nig pøÚv glycolysis TaMgmUl .

rUbTI 20-11 Allosteric Enzymes



x- Feedback Inhibition

Allosteric enzymes KWCamUldæanRKwHén Feedback inhibition énpøÚvCIvKImI . RbB½n§enHbMeBj muxgarsMGagelIeKalkarN_dUceTA thermostat énbnÞb;manbMBak;]bkrN_EksMrYlsItuNðPaB (furnace) . Gñkdak;kMNt; [ thermostat enAsItuNðPaB 70O F ]bkrN_enaHk¾dMeNIrkar nig beBa©j kMedArhUtdl; sensor EdlmanenAkñúg thermostat rab;eXIjsItuNðPaB 70OF . bnÞab;mkvapþl;sBaØa []bkrN_enaHQb;dMeNIrkar . CaFm μta Feedback inhibition EktMrUvpøÚvrbs;Gg;sIumEdlCab;Tak;TgeTAnwgkarsMeyaK mU:elKulCIvsaRsþ . pøÚvenHGacRtUv)anbgðajdUcKMnUsbMRBYjxageRkam ³ E1 E2 E3 E4 E5 A → B → C → D → E → F

enAkñúgpøÚvenH sarFatuEdlcab;epþIm A RtUv)anbMElgedayGg;sIum E1 [eTACa B . Gg;sIum E2

bMElgya:gelOnBI B eTA C nig CabnþbnÞab;rhUtdl;plitplcugbBa©b; F RtUv)ansMeyaK . RbsinebI F

ElgRtUvkar vaKWCakarx¢Hx¢ayfamBlekasikakúñgkarbnþplitva . edIm,IeCosvagkarx¢Hx¢ayfamBlenH ekasikaeRbI Feedback inhibition EdlplitplGac bBaÄb;pøÚvTaMgmUlsMrab;sMeyaKxøÜnva . karenHKWCalT§plénkarBitEdlfa plitpl F edIrtYCa Negative allosteric effector eTAelIGg;sIumxagedImmYyrbs;pøÚvemtabUlIs . ehtudUecñHGg;sIum E1

GacmanTIkEnøgcgP¢ab; effector sMrab; F bEnßmeTAelITIkEnøgskmμEdlP¢ab;eTAnwg A . enAeBlman vtþman F edayelIsvaP¢ab;eTAnwgTIkEnøgP¢ab; effector . karP¢ab;enHbgá[TIkEnøgskmμbiT dUecñHvamin GacP¢ab;eTAnwg sustrate A . dUecñH A minGacbMElgeTACa B . RbsinebIminman B RtUv)anplit vamin man substrate sMrab;Gg;sIum E2 nig pøÚvemtabUlIsTaMgmUlRtUv)anbBaÄb;Rbtibtþikar . plitpl F

)anbBaÄb;RKb;dMNak;karTaMgGs;EdlCab;Tak;TgeTAnwgkarsMeyaKxøÜnvaenH dUceTAnwgkMedAEdlbegáIt eday]bkrN_EktMrUvkMedA (furnace) EdlCacugbBa©b;TTYlxusRtUvcMeBaHkarbiT]bkrN_edayxøÜnÉg . K- Zymogens

meFüa)aycugeRkaybMputénkarEktMrUvskmμPaBGg;sIumCab;Tak;TgeTAnwgkarplitGg;sIum enA kñúgTMrg;Gkmμ (inactive form) EdleKehAfaCa Zymogens .bnÞab;mkCaTUeTAvak¾bMElgeday Proteolysis ¬GIuRdUlIsRbUetGIun¦ [eTACaTMrg;skmμ enAeBlvaeTAdl;TIkEnøgskmμrbs;va . etIGIVCa eKalbMNgénynþkarRbePTenH ? enAelIkarBinitüelIkTI 1 vaTMngCax¢Hx¢aybEnßmeTAelIdMNak;kar én kasMeyaKrbs;Gg;sIum . b:uEnþcUrBicarNacMeBaHclna EdlmanFmμCatiEdlGacxUcxatmYycMnYnrbs;



Gg;sIumEdlcaM)ac;sMrab;CIvitrs;enA . Gg;sIum Pepsin , Trypsin nig Chymotrypsin KWCaGg;sIum proteolytic énpøÚvGahar . vacaM)ac;sMrab;karrs;enA BIeRBaHvabMEbkRbUetGIunEdlmankñúgGaharCa GasIutGamIneNEdlRtUv)aneRbIedayekasika . b:uEnþetIGIVEdlnwgekIteLIgcMeBaHekasikaEdlbegáIt Gg;sIumTaMgenH RbsinebIBYkvaRtUv)ansMeyaKkñúgTMrg;Gkm μ ? ekasikaTaMgenaHnwgRtUv)anbMpøaj . ehtu dUecñH ekasikarbs;RbBHEdlCaFm μtabegáItnUv pepsin plitnUv zymogen GkmμEdleKehAfa Pepsinogen . Pepsinogen man 42 GasIutGamIneNbEnßmeTot . enAeBlmanvtþmanGasIutrbs; RkBH nig pepsin Edl)aneFIVskm μBImunmk GasIunGamIneN 42 Edl)anbEnßmRtUv)ankat;pþac;ecj nig zymogen RtUv)anbMElgeTACaGg;sIumskm μ . taragTI 20-2 erobrab;GMBI zymogens epSg²eTotCa eRcIn nig Gg;sIumEdlbMElgBYkvaeTATMrg;skm μ .

taragTI 20-2 Zymogens rbs;pøÚvGahar

Zymogens Actvators Enzymes Proelastase Tryppsin Elastase Trypsinogen Trypsin Trypsin Chymotrypsinogen A Trypsin + Chemotrypsin Chymotrypsin Pepsinogen Acid pH + Pepsin Pepsin Procarboxypeptidases Trypsin Carboxypeptidase A Carboxypeptidase B

10- karraraMgénkmμPaBGg;sIum (INHIBITION OF ENZYME ACTIVITY)

CatiKImICaeRcInGaccgP¢ab;eTAnwgGg;sIum nig TaMgGaclubbM)at; (eliminate) b¤ kat;bnßyya:g xøaMgnUvsmtßPaBsMrYlrbs;va (catalytic ability) . sarFatuKImITaMgenH RtUv)aneKehAfaCa GñkraraMgGg;sIum (Enzyme inhibitors) RtUv)aneKeRbIGs;ry³eBlCaeRcInryqñaMmkehIy . enAeBl nagbMBulCnrgeRKaHrbs;nageday Arsenic, Lucretia Borgia min)andwgfavaP¢ab;eTAnwgbgÁúM Thiol

rbs; Cysteine amino acids enAkñúgRbUetGIunrbs;CnrgeRKaHrbs;nageT nig ehtudUecñH edaykarman kareRCotERCkedaykarbegáItsm<½n§ disulfures (disulfide bonds) EdlRtUvkarsMrab;eFIV[manlMnwg rcnasm<½n§TI 3 rbs;Gg;sIum . b:uEnþnagdwgya:gc,as;BIPaBBuldl;søab;rbs;BYkGMbilelah³F¶n; dUcCa Arsenic nig )art (Hg) . enAeBlGñkpwkfñaM Penicillin sMrab;karBüa)alCMgWbgáeday)ak;etrI GñkkMBug eRbIGñkraraMgGg;sIumepSgeTot . Penicillin raraMgGg;sIumCaeRcInEdlCab;Tak;TgeTAnwgkarsMeyaKPñas ekasika)ak;etrI . GñkraraMgG;sIum RtUv)aneFIVcMNat;fñak;edaysMGageTAelI faetIkarraraMgenaHvaRtLb;mkvij b¤ min RtLb;mkvij RbECgKña b¤ minRbECgKña . PaBRtLb;mkvij (Reversibility) mann½yfaetI



GñkraraMgcugbBa©b;nwgEbkecjBIGg;sIum edaybeB©ajvaeRkamTMg;CaGg;sIumskm μ . karRbECg (Competition) mann½yfaetIGñkraraMg manPaBdUcKñaénrcnasm<½n§ b¤ dUceTAnwg substrate . RbsinebI dUecñH GñkraraMg nig substrate nwgRbECgKñacMeBaHTIkEnøgskmμrbs;Gg;sIum . k- GñkraraMgminRtLb;mkvij (Irreversible inhibitors)

GñkraraMgGg;sIumminRtLb;mkvij (Irreversible enzyme inhibitor) dUcCa As, CaFm μtaP¢ab; ya:gxøaMg CYnkaledasm<½n§kUva:Lg; eTAnwgGg;sIum . CaTUeTAkarenHCab;Tak;TgeTAnwgkarcgénGñkraraMg eTAnwgbgÁúMmYykñúgcMeNambgÁúM R rbs;GasIutGamIneNenAkñúgTIkEnøgskmμ . karcgGñkraraMgGaceFIV[ KaMgbgÁúMEdlcgTIkEnøgskmμ dUecñHbgÁúM enzyme-substrate minGacbegáIt)aneT . müa:gvijeTot GñkraraMgGaccUleRCotERCkCamYynwgbgÁúMsMrYl (Catalytic group) énTIkEnøgkmμ ehtudUecñHkarlubbM)at; nUvkarsMrYledaymanRbsiT§PaB . GñkraraMgminRtLb;mkvijCaTUeTAraraMgGg;sIumepSg²CaeRcIn . ]TahrN_rYmmanBisstVeBaH nig ]sμ ½nsésRbsaT . x- GñkraraMgRbECgRtLb;mkvij (Irreversible, Competitive Inhibitors)

GñkraraMgRbECgRtLb;mkvij Cajwkjab;sMedAcMeBaHsarFatuEdlmanPaBdUcKñaénrcnasm<½n§ eTAnwg substrate KWfa BYkvaKWCamU:elKulEdlmanrcnasm<½n§ nig karEbgEckénbnÞúkGKÁisnIdUceTAnwg substrate cMeBaHGg;sIummYycMnYn . GaRs½yedayPaBdUcKñaenH GñkraraMgGaccUleTAP¢ab;nwgTIkEnøg skmμrbs;Gg;sIum . ehtudUecñH K μanRbtikmμekIeLIg nig skmμPaBrbs;Gg;sIumRtUv)anraraMg . karraraMg enHRtUv)aneKniyayfavaRbECgKñaBIeRBaHGñkraraMg nig Substrate RbECgkñúgkarcgP¢ab;eTAnwg TIkEnøg skmμrbs;Gg;sIumEtmYy . ehtudUecñH kMriténkarraraMgGaRs½yeTAnwgkMhab;rbs;BYkva . RbsinebIGñk raraMgelIs b¤ cgya:gxøaMgeTAnwgTIkEnøgskm μ vanwgcUleTAP¢ab;eTAelITIkEnøgskmμeRcIn nig skmμPaB rbs;Gg;sIumnwgRtUvfycuHya:gxøaMg . müa:gvijeTot RbsinebI substrate manelIs vanwgcUleTAcgP¢ab; nwgTIkEnøgskm μeRcIn nig vamankarraraMgEtbnþibbnþÜc . »sf Sulfa (Sulfa drugs) Ca»sfpSHdMbUgbMputRtUv)aneKrkeXIj KWCa GñkraraMgRbECg én Gg;sIum)ak;etrIEdlRtUvkarsMrab;karsMeyaKénvItamIn Folic acid . Folic acid KWCavItamInEdlRtUvkar sMrab;karbBa¢ÚnnUvbgÁúM methyl enAkñúgCIvsMeyaKén methionine nig nitrogenous bases . mnusSmin GacsMeyaK Folic acid nig RtUv)anEtTTYlva)anBIGahar . müa:gvijTot )ak;etrI RtUvEteFIV Folic acid

BIeRBaH minGaccab;yk folic acid BIbriyakasCMuvij . Para-Aminobenzoic acid (PABA) KWCa substrate sMrab;dMNak;kardMbUgbMputkñúgkarsMeyaK Folic acid . Sulfa drugs TMrg;dMbUgbMput (Prototype) EdlRtUv)anrkeXIjeday Gerhard Damagk



enAkñúgTsSvtSqñaM 1930 Edlmanrcnasm<½n§dUceTAnwg PABA nig ehtudUecñH vaCaGñkraraMgRbECgrbs; Gg;sIumEdleRbI PABA Ca substrate Fmμtarbs;va .

RbsinebI substrate RtwmRtUv)anP¢ab;eTAnwgGg;sIum RbtikmμekIteLIg nig )ak;etrIenArs; .b:uEnþ ebI Sulfa drugs manenAkñúgkMritelIs PABA Cajwkjab;vaP¢ab;CaeRcIneTAelITIkEnøgskmμrbs; Gg;sIum. minman Folic acid RtUv)anplit ehIy)ak;etrIRtUvsøab; . GaRs½yedayeyIgTTYl folic acid BIrbbGaharrbs;eyIg Sulfa drugs mineFIV[b:HBal;dl; xøÜneyIgeT . b:uEnþ )k;etrIRtUv)ansMlab;edayeRCIserIs . CasMNag eyIgGacman»kaskñúgkarTaj ykplRbeyaCn_BIlkçN³enHsMrab;karBüa)alénCMgWbgáeLIgBI)ak;etrI nig CalT§pl Sulfa drugs

)anCYyCIvitrab;minGs; . eTaHbICaCMgWbgáedayemeraKCamUlehtucMbgénmrN³PaBmunkarrkeXIj sulfa drugs nig »sfpSH (Antibiotics) k¾eday mrN³PaBEdlbgáedayCMgWngáeday)ak;etrIenAeBl bc©úb,nñkMrman . K- GñkraraMgminRbECgRtLb;mkvij GñkraraMgminRbECgRtLb;mkvij P¢ab;eTAnwgbgÁúM R rbs;GasIutGamIneN b¤ eTAnwg Cofactor

EdlCaGIuy:ugelah³ . mindUceTAnwgGñkraraMgminRtLb;mkvij b:uEnþkarP¢ab;rbs;vaexSay nig skmμPaB rbs;Gg;sIumRtUveFIV[l¥dUcedImvij enAeBlGñkraraMgRtUv)anpþac;ecjBIbNþúMGg;sIum -GñkraraMg . eTaHbICaedayTUeTAGñkraraMgTaMgenHmincgeTAnwgTIkEnøgskmμk¾eday b:uEnþBYkR)akdeFIV[ERbRbYlrUb ragTIkEnøgskmμEdlcgP¢ab;eTAkEnøgNamYyenAkñúgrcnasm<½n§rbs;RbUetGIun . cUrcgcaMfa TMrg;kñúglMhr (three-dimensional structure) rbs;Gg;sIum RtUvkarrkSarUbragRtwmRtUvénTIkEnøgskmμ . karcgén GIuy:ug b¤ karcgcMNgexSayrbs;GñkraraMgeTAkEnøgNamYy b¤ eRcInenAelIépÞGg;sIumGaceFIV[xUcrUbrag TIkEnøgskm μkñúgrebobdUceTAnwg allosteric effector GñkraraMgenHk¾eFIV[ElgskmμEdrnUvGg;sIum CaeRcIn. 11- Gg;sIumbMEbkRbUetGIun (PROTEOLYTIC ENZYMES)

Gg;sIumbMEbkRbUetGIun KWCaGg;sIumEdlkat;pþac;RbUetGIun KWfaBYkvapþac;sm<½n§ peptide EdlrkSa nUvrcnasm<½n§TImYy . Chymotrypsin Ca]TahrN_ KWCaGg;sIumEdlGIuRdUlIsRbUetGIuncMNIGaharenA kñúgeBaHevontUc . vaedIrtYedaylkçN³BiesseTAelIsm<½n§ peptide xagbgÁúM Carbonyl énsm<½n§ peptide. GasIutGamIneN C-terminal rbs; peptides RtUv)anbeBa©jedaykarkat;sm<½n§KW

H2N

O ║ C—OH H2N

O ║ S—NH2 ║ O



methionine, tyrosine, tryptophan, nig Phenylalanine . PaBBiess¬1¦ rbs; Chymotrypsin vaGaRs½yeTAnwgvtþmanrbs; Hydrophobic pocket ¬bNþúMénGasIutGamIneNEdmincUlcitþTwkEdlsßit enACamYyKñaedaysarkarbt;CaragkñúglMhr (three-dimentional) énExSRbUetGIun . ExSbiTénBYk m:UelKulRkGUb (flat aromatic side chains) rbs;GasIutGamIneNmYycMnYn (tyrosine, tryptophan,

phenylalanine) rGilcUleTAkñúgfg;enH (pocket) edaypþl;[CaPaBBiessénkarcgEdlRtUvkarsMrab; karsMrYl ¬rUbTI 20-12¦ . etIeyIgGackMNt;rebobdUcemþcfasm<½n§mYyNaEdlRtUv)ankat;eday protease dUcCa Chymotrypsin ? edIm,Idwgsm<½n§mYyNaRtUv)ankat; eyIgRtUvEtsresrnUvQutrbs;GasIutGamIneN enAkñúgtMbn;én peptide EdlRtUv)ankat; . karenHGacRtUv)ankMNt;edaykarBiesaFn_edaybec©keTs kMNt;QutGasIutGamIneN (amino acid squencing techniques) . cUrcgcaMfa GasIutGamIneN N-

terminal RtUv)ansresrenAxageqVg nig C-terminal enAxagsþaM . cUrBicarNaBIRbUetGIunEdlmanQut -Ala-Phe-Gly- . RbtikmμRtUv)anekIeLIgEdlGg;sIum Chymotrypsin RtUv)anlayCamYynwgRbUetGIun substrate . bnÞab;BIRbtikmμekIeLIg plitplRtUv)anbnSúT§ nig QutGasIutGamIneNrbs;BYkvaRtUv )aneKkMNt;rk . karBiesaFn_RbePTenHbgðajfa Chymotrypsin kat;pþac;sm<½n§rvag Phenylalanine

nig Glycine EdlCasm<½n§ peptide enAxagExSEdlmanm:UelKulRkGUb (aromatic side chain) . Pancreatic serine protease (Trypsin, chymotrypsin, and elastase) BYkvaTaMgGs; GIuRdUlIssm<½n§ peptide . Gg;sIumTaMgenH KWCalT§plén divergent evolution EdlEhSndUntaEt mYydMbUgRtUvftcMlg (duplicated) nig bnÞab;mk EhSnEdlftcMlg)aneFIVkarvivtþn_edayELk²BIKña. BYkvamanrcnasm<½n§TI1 TI 3Rbhak;RbEhlKña nig manynþkarénGMeBIdUcKña . bu:EnþCalT§plén karvivtþn_ Gg;sIumTaMgenHmanPaBBiessxus²Kña . Chymotrypsin kat;sm<½n§ peptide enAelIEpñkxag Carbonyl rbs;GasIutGamIneN Edl manExSmU:elKulRkGUb nig GasIutGamIneNmincUlcitþTwkdUcCa Methionine . Trypsin kat;sm<½n§ peptide enAxagEpñk Carbonyl rbs;GasIutGamIneN)as . Elastase kat;sm<½n§ peptide enAxagEpñk Carbonyl rbs; Glycine nig Alanine . Gg;sIumTaMgenHmanfg;epSg² (pocket) sMrab;ExSén substrate ¬emesaepSg²RtUveTAnwg kUnesaepSg²¦ . PaBxusKñaenHbgðajxøÜnvaenAkñúgPaBBiessrbs; substrate Edl)anbgðajxagelI . ]TahrN_ fg;Edlcgrbs; tyrosin KWEvg ceg¥ot nig manbnÞÚkGviC¢manedIm,IeFIV[RtUveTAnwgbgÁúM R rbs;



lysine b¤ Arginine . b:uEnþeTaHbICa fg;Edlcg)anTTYlrgnUvkarvivtþn_BIedImdMbUg (divergent

evolution) TIkEnøgsMrYl)ansßitenAdEdlminmankarpøas;bþÚr nig ynþkarénGMeBIbMEbkRbUetGIundUceTA nwgRKb; serine proteases . enAkñúgkrNInimYy² ynþkarCab;Tak;TgeTAnwgbgÁúM R rbs; serine . 12- kareRbIGg;sIumenAkñúgevC¢saRsþ (THE USES OF ENZYMES IN MEDICINE)

karviPaKQamedIm,IkMNt;kMritrbs;Gg;sIummYycMnYnGacpþl;nUvB½t’manya:geRcIn GMBIlkçx½NÐ evC¢saRsþ . Cajwkjab; etsþrebobenHRtUv)aneRbIedIm,IbBa¢ak;eraKvinicä½yBIdMbUgEdlsMGagelIeraKsBaØa rbs;CMgW b¤ TidæPaBKøInik . ]TahrN_enAeBlman Heart attack ekIteLIg ¬karxVHxatnUvQamEdlRtUv)anpþl;eTA[ sac;dMuebHdUgbgá[ekasikasac;dMumYycMnYnsøab; . ekasikaTaMgenHbeBa©jsarFatuEdlenAxagkñúgrbs; BYkva rYmmanTaMgGg;sIumrbs;BYkva cUleTAkñúgcrnþQam . etsþ_samBaØGacRtUv)aneKeFIVedIm,Ivas;cMnYn Gg;sIummYycMnYnenAkñúgQam . etsþ_rebobenHRtUv)aneKehAfa Enzymatic assays EdlmanlkçN³ RtwmRtUv nig BiessxøaMgBIeRBaHBYkvasMGageTAelIPaBBiessrbs;bNþúM enzyme-substrate . RbsinebI GñkcgeFIVetsþ_cMeBaHGg;sIum Lactate dehydrogenase (LDH) GñkRtUvkarEtbEnßmEt substrate RKb; RKan; enAkñúgkrNIenH Pyruvate nig NADH . Rbtikm μEdlekIteLIgKWGuksIutkmμrbs; NADH eTACa NAD+ nig er:dukmμ Pyruvate eTACa Lactate . edIm,Ivas;GRtaRbtikm μKImI eKGacvas;kar)at;én substrate b¤ k¾karekIneLIgeRcInénplitplNamYy . kñúgkrNIén LDH viFIeRbI spectrophotomer

¬sMGageTAelIlkçN³RsUbBnøWrbs; substrate b¤ plitpl¦ GaceRbIedIm,Ivas;GRtaénkarplitrbs; NAD+ . tYnaTIrbs; LDH nig Gg;sIumepSgeTot nig Gg;sIumepSgeTotenAkñúgkareFIVeraKviniqä½y RtUv)an BiPakSaenAkñúg Clinical Perspective : Enzyme , Isoenzymes and Myocardial infarction. kareRCIserIsén substrate kMNt;nUvGIVEdlskmμPaBrbs;Gg;sIumRtUv)aneKvas; . kMhab;enAkñúg serum én Amylase nig Lipase ekIneLIg KWCakarbgðajnUv Pancratitis

¬karrlakénlMEBg¦ . CMgWeføIm dUcCaRkineføIm (cirrhose) nig rlakeføIm (hepatitis) bNþal[mankar ekIneLIgmYykñúgcMeNam isoenzymes rbs; Lactate dehydrogenase (LDH5) nig ekIneLIgnUvkMrit Alanine aminotransferase/Serum glutamate-pyruvate transaminase (ALT/SGPT) nig Aspartate aminotransferase/Serum glutamate-oxaloacetate transaminase (AST/SGOT) enA kñúg serum . CakarBit Gg;sIumBIrxageRkayk¾mankMhab;ekIneLIgbnÞab;BIman heart attack Edr b:uEnþ RKUeBTüGacEjklkçx½NÐTaMgBIrenHedaykarBicarNakareLIgtictYcénGg;sIumTaMgBIrenH . RbsinebI ALT/SGPT ekIneLIgFMCag AST/SGOT vaGacbgðaj[eXIjfabBaðaKWCaPaBRcbUkRcbl;éneføIm .



Gg;sIumk¾RtUv)aneRbICaRbtikrsMrab;viPaKEdrenAkñúgmnÞIrBiesaFn_KøInikGaRs½yedayPaBBiess rbs;va . Cajwkjab;vamanRbtikmμedayeRCIserIseTAelIsarFatumYy eday[CaplitpEdlgayRsYl kñúgkarvas; . ]TahrN_énkarenHkarviPaKén Urea enAkñúgQam . karvas;énkMrit urea enAkñúgQam valM)ak BIeRBaHPaBsμ úRKs μajénQam . b:uEnþ RbsinebI urea RtUv)anbMElgeTACa NH3 edayeRbI Gg;sIum urease , Ammonia (NH3) køayeTACaFatbnÞal; (Indicator) rbs; urea BIeRBaHvaRtUv)anplit BI Urea nig vagayRsYlnwgvas; . etsþ_enHRtUv)aneKehAfaCa Urea Blood NItrogen (BUN) RtUv)an eKeRbIkñúgkarrkeraKvinicä½yénPaBminFmμtarbs;tMrgenam nig eRbICa]TahrN_mYykñúcMeNaménkareRbI Gg;sIumenAkñúg Clinical Chemistry . karBüa)alCMnYsedayGg;sIumk¾GacRtUv)aneKeRbIenAkñúgkarBüa)alénCMgWmYycMnYn . CMgWmYy Gaucher's disease KWCaCMgWEdlmanPaBRcbUkRcbl;énEhSnbNþalBIPaBfycuHénGg;sIum glucocerebrosidase . enAkñúgsßanPaBFmμta Gg;sIumenHbMEbk glycolipids EdleKehAfa glucocerebroside EdlCaFatukNþalénkarsMeyaK nig karbMEbkbNþúM glucosphingolipids Edl eXIjmanenAkñúgPñasekasika . Glucocerebrosidase eKeXIjmanenAkñúg lysosomes EdlvaGIuRdUlIs glucocerebrosides eTACa Glucose nig Ceramide . enAkñúg Gaucher's disease minmanvtþmanGg;sIum nig glucocerebroside RtUv)anbegáItenAkñúg macrophage EdleXIjmanenAkñúg eføIm spleen nig xYrq¥wg . ekasikaTaMgenHnwgrIkFMrYmCamYy karelIslIBIdEdlminGacRtUv)aneFIVemtabUlIs nig bnÞab;mkCMnYsekasikaFmμtaEdlmansuxPaBl¥ enAkñúgxYrq¥wg . eraKsBaØaén Gaucher's disease rYmmanesøksøaMgRsYcRsav thrombocytopenia ¬kar fycuHéncMnYn)aøEkt¦ nig Hepatosphlenomegaly ¬karrIkFMéneføIm nig spleen¦ . vak¾GacmanEdr nUvbBaðaeRKagq¥wg (skeletal problems) rYmmanPaBTnexSayénq¥wg nig kar)ak; (bone deterioration

and secondary fractures) . Recombinant DNA Technology RtUv)aneRbIeday Genzyme Corporation edIm,IbegáIt Gg;sIumén lysosome rbs;mnusSKW β-glucocerebrosidase . eday[eQ μaHBaNiC¢km μ Cerezyme

Gg;sIumGIuRdUlIs glucocerebroside eTACa glucose nig ceramide dUecñHpliplGacRtUv)anTTYlrg emtabUlIsCaFmμta . GñkCMgWTTYl Cerezyme tamsésEv:nkñúgry³eBlBImYyeTABIrem:ag . dosage

nig taragénkarBüa)alGacRtUv)aneFIV[smRsbeTAnwgbuKÁlnimYy² . lT§plénkarsakl,genH KWCakarCYyxøaMgNs; . GñkCMgWTTYl)annUvkarEklMGrekasikaRKab;QamRkhm nig )aøEkt nig hepatosplenomegaly RtUv)ankat;bnßy .



Fin




sMNYr 1. GgsIumcUlrYmsMrYlRbtikm μeday ³ A. begIánnUvfamBleFIV[skmμ B. bnßynUvfamBleFIV[skmμ C.mineFIV[pøas;bþÚrfamBleFIV[skm μ 2. lkçN³Biessrbs;Gg;sIumGacRtUv)ankMNt;edaylkçN³xageRkam ³ ¬rkXøaRtUvCageKbMput¦ A. TMhMm:UelKul B. TMrg;m:UelKul C. RbePTbnÞúkGgÁisnIrbs;m:UelKul D. A + B + C RtwmRtUv 3. eKGacehAeQ μaHrbs;Gg;sIumeTAtamlkçN³dUcxageRkam ³ ¬rkXøaRtUvCageKbMput¦ A. tameQμaHrbs; substrate edaybEnßmbc©wmbT "ase"

B. tameQμaHrbs;Rbtikm μEdlGg;sIumsMrYl edaybEnßmbc©wmbT "ase"

C. tameQ μaH substrate nig eQ μaHRbtikmμEdlGg;sIumsMrYl . D. A b¤ B b¤ C suT§EtRtwmRtUv 4. kñúgcMeNamsarFatuxageRkammanmYyEdlminEmnCaGg;sIum vaenaHKW ³ A. Urease B. Hydrogenase C. pepsin D. Chymotrypsin E. Pyruvate

5. kñúgcMeNamGg;sIumxageRkammanCaGg;sIumsßitkñúgBYk Oxidoreductases vaenaHKW ³ A. LDH B. SGOT C. Lipase D. Phosphoglyceromutase E. DNA ligase

6. kñúgcMeNamGg;sIumxageRkammanCaGg;sIumsßitkñúgBYk Transferases vaenaHKW ³ A. LDH B. SGPT C. Lipase D. Phosphoglyceromutase E. DNA ligase

7. kñúgcMeNamGg;sIumxageRkammanCaGg;sIumsßitkñúgBYk Hydrolases vaenaHKW ³ A. LDH B. SGOT C. Lipase D. Phosphoglyceromutase E. DNA ligase

8. kñúgcMeNamGg;sIumxageRkammanCaGg;sIumsßitkñúgBYk Isomerases vaenaHKW ³ A. LDH B. SGOT C. Lipase D. Phosphoglyceromutase E. DNA ligase

9. GgsuImbegáInel,ÓnRbtikm μeday ³ A. eFIV[pøas;bþÚrRbePTplitpl B. bnßynUvfamBleFVI[skm μ C. eFIV[xUcefrlMnwgRbtikm μ 10. kñúgkrNIeKmanbrimaNGg;sIumkMNt;mYy kalNakMhab;rbs; substrate ekIneLIgGRtaRbtikmμ ³ A. ekIneLIgrhUttamkMhab; substrate B. ekIneLIgRtwmkMritGtib,rma C. minmankarekIneLIg 11. Substrate mYyGaccUlrYmeFIVRbtikmμCamYynwgGg;sIumenA ³ A. RKb;TIkEnøgTaMgGs;énGg;sIum B. TIkEnøgskm μrbs;Gg;sIum 12. lkçN³Biessrbs; enzyme-substrate mandUcCa ¬rkXøaRtwmRtUvCageKbMput¦ ³ A. PaBBiessdac;xat B. PaBBiessCaRkum C. PaBBiessénkarcgP¢ab;

D. PaBBiessénrcnasm<½n§KImIkñúglMhr E. A + B + C + D

13. Gg;sIumskmμRtUv)aneKehAvafaCa ³ A. Apoenzyme B. Coenzyme C. Holoenzyme

14. \T§iBlbriyakasCMuvijeTAelIGg;sIum³ ¬rkXøaxus¦



A. pH B. Temperature C. solubility

15. karEktMrUvénskmμPaBGg;sIumman ³ ¬rkXøaRtUvCageKbMput¦ A. Allosteric enzymes B. Feedback inhibition C. Zymogens D. A+B+C

16. karraraMgénskmμPaBrbs;Gg;sIumman ³ A. irreversible inhibition B. reversible competitive inhibition C. reversible non-competitive inhibition D. A + B + C

17. Gg;sIumKWCam:UelKulCIvsa®sþEdlCaGñksMrYlRbtikm μKImI ehIymansar³sMxan; ³ A. CaGayuCIvit ¬KμanGg;sIumK μanCIvit¦ B. l μm ¬mincaM)ac;manGg;sIumk¾)an¦ 18. GRtaRbtikmμEdlsMrYledayGg;sIumKW ³ rkXøamYyEdlxus A. BIdMbUgekIneLIgeqøIytbnwgkMhab;rbs; Substrate

B. ekIneLIgdl;kMritGtib,rmaenAkñúgkMritmYyénkMhab;rbs; substrate

C. ekIneLIgeTVreTAtamkarekIneLIgénkMhab;rbs; Substrate

19. cMNucxageRkamTaMgenH manmYyCamUlehtuEdlnaM[GRtaRbtikmμEdlsMrYledayGg;sIumekIneLIg dl;kMritGtib,rma enAkñúgkMritmYyénkMhab;rbs; substrate vaenaHKW ³ A. Gg;sIumRtUv)anbMgøajGs; B. TIkEnøgskmμCab;rvl;Gs; C. TIkEnøgskmμRtUv)anbMpøajGs; 20. xageRkamenHKWCadMNak;énkarsMrYlRbtikmμKImIedayGg;sIumelIkElgEtmYy vaenaHKW ³ A. dMNak;karbegáItbNþúM Enzyme-Substrate

B. dMNak;karbeNþaHGasnñ (ES*)

C. dMNak;karbMElgeTACaplitpl nig Gg;sIum (E +P)

D. dMNak;karbMpøajGg;sIum 21. xageRkamenHKWCalkçN³TUeTAénTIkEnøgskmμrbs;Gg;sIumelIkElgEtmYy vaenaHKW ³ A. TIkEnøgskmμKWCaRbeLaH ¬ehae):A b¤ esa¦ enAelIépÞrbs;Gg;sIum . B. rUbragrbs;TIkEnøgskm μRtUvKñanwgrUbragrbs; Substrate . C. Gg;sIumTaj nig cab;yk Substrate eTAtamsm<½n§exSay D. TMrg;énTIkEnøgskm μkMNt;nUvlkçN³Biess (Specificity) rbs;Gg;sIum . E. Gg;sIumTaj nig cab;yk Substrate eTAtamsm<½n§kUva:Lg; . 22. Coenzymes KWCam:UelKulsrIragÁEdlCaTUeTARtUv)aneRbICa ³ ¬rkXøYamYyEdlxus¦ A. GñkdwknaMeGLikRtug b¤ bNþúMKImI B. Gñkpþl;bgÁúMnaTIeTA[ Substrate

C. GñkTTYlykbgÁúMnaTIBI Substrate D. eFIV[xUcrcnasm<½n§ substrate

23. kñúgcMeNamsarFatuTaMgLayxageRkamenHsuT§EtCa Coenzymes elIkElgEtmYy vaenaHKW ³ A. NAD B. FAD C. NADP D. LDH E. FMN



24. xageRkamKWCalkçx½NÐTUeTA nig smRsmedIm,I[Gg;sIummanskmμPaBeBjeljelIkElgEtmYy vaenaHKW ³ A. pH optimun (≈ 7) B. pH Base (> 10) C. pH Acid (<6) D. sItuNðPaB 37oC E. B +C

25. Gg;sIum Proteolytic mYyEdlGacmanskmμPaBeBjeljenAkñúg pH optimum = 2 vaenaHKW ³ A. Pepsin B. Trypsin C. Chymotrypsin D. Elastase

26. cUrGanvaküx½NÐxageRkam nig dak;BaküTaMgenH[cMTIkEnøg ³ A. Negative allosterism,

B. Allosteric enzymes, C. Positive allosterism, D. effector molecule RbePTmYyénkarEktMrUvGg;sIumCab;Tak;TgGg;sIumEdlmanTIkEnøgcgP¢ab;eRcInCagmYy . Gg;sIumTaMgenHRtUv)aneKehAfa 1. _______B________ mann½yfa {TMrg;epSgeTot} KWCaGg;sIum EdlTIkEnøgskmμrbs;vaGacRtUv)anxUcedaysarkarcgénm:UelKultUcEdleKehAfa 2. ____D_____. lT§plGacRtUv)anbMElgTIkEnøgskm μeTACa TMrg;minskmμ (inactive configuration) EdlKWCa 3.___A____b¤ bMElgTIkEnøgskm μeTACa TMrg;skkm μ (active configuration) KWCa 4. ______C____

CIvKImIviTüa emtabUlIsKøúysIut

erobcMnigbkERbeday»sfkarIRcwgsuPaBTUrs½BÞ012864353b¤011717361e-mail: [email protected] Page 1 of 35

emtabUlIsrbs;kabUGIuRdat (CARBOHYDRATE METABOLISM)

vtßúbMNgemeron 1- BiPakSaGMBIsar³sMxan;rbs; ATP enAkñúgdMeNIrkarbBa¢ÚnfamBlkñúgekasika . 2- BiB’NnaGMBIdMNak;karTaMgbIénkatabUlIsén RbUetGIun kabUGIuRdat nig lIBIdenAkñúgrbbGahar. 3- BiPakSaGMBI Glycolysis enAkñúgEpñksMxan;TaMgBIr . 4- BinitüemIlsmIkaEdltMNag[Rbtikm μKImIEdlekIeLIgenAkñúg Glycolysis BiB’NnaGMBIRbePTén RbtikmμEdlekIteLIg nig sar³sMxan;énRbtikmμTaMgenaHenAkñúgpøÚvemtabUlIs . 5- BiB’NnaGMBIynþkarrbs;karEktMrUvénGRta glycolysis . BiPakSaGMBI[TahrN_BiessénkarEktMrUv. 6- BiPakSaGMBItYnaTIGnuvtþ nig tYnaTIemtabUlIsGMBIRbtikmμeLIgem (fermentation reactions) . 7- erobrab;GMBIplitplCaeRcInrbs;pøÚvén pentose phosphate EdlRtUvkarenAkñúgCIvsMeyaK . 8- eRbobeFob glycolysis nig glyconeogenesis . 9- segçbénkarEktMrUvénkMritKøúykUsenAkñúgQam eday glycogenesis nig glycogenolysis .

Fin



KMeragemeron

1.ATP : Cellular Energy Currency 2. Overview of Catabolic Processes 2.1. Stage 1 : Hydrolysis of Dietary Macromolecules into Small Subunits 2.2. Stage 2 : Conversion of Monomers into a Form That Can Be Completely Oxidized. 2.3. Stage 3 : The Complete Oxidation of Nutrients and the Production of ATP 3. Glycolysis 3.1. An Overview 3.2. Reactions of Glycolysis 3.3. Regulation of Glycolysis 4. Fermentations 4.1. Lactate Fermentation 4.2. Alcohol Fermentation 5. The Pentose Phosphate Pathway 6. Gluconeogenesis : The Synthesis of Glucose 7. Glycogen Synthesis and Degradation 7.1. The Structure of Glycogen 7.2. Glycogenolysis : Glycogen Degradation 7.3. Glycogenesis : Glycogen Synthesis



emtabUlIskabUGIuRdat (CARBOHYDRATE METABOLISM)

esckþIepþIm RKan;EteyIgRtUvkarfamBlkñúgkar rt; elat nig Kit ekasikaRtUvkar karpþl;rYcCaeRscnUv famBlrbs;ekasikasMrab;muxgarCaeRcInEdlKaMRTskmμPaBTaMgenH . ekasikaRtUvkarfamBlsMrab; kardwknaMskm μ (active transport) edIm,IeFIV[m:UelKulmanclnarvagbrisßanCMuvij nig ekasika. famBlk¾RtUvkarsMrab; CIvsMeyaK (Biosynthesis) énm:UlelKultUc²rbs;emtalIs nig karplitén m:UelKulFM²BIsarFatukNþalTaMgenH (intermediates) . cugbBa©b; famBlRtUvkarsMrab; kargaremkanik (mechanical work) rYmmankarkRnþak;sac;dMu nig clnarbs; Es<mma:tUsUGIut (motility

of sperm cell) . taragTI 21-1 erobrab;GMBI]TahrN_mYycMnYnéndMeNIrkarEdlRtUvkarfamBl TaMgenH . eyIgRtUvkar karpþl;énm:UelKulGaharEdlsMbUredayfamBl EdlGacRtUv)anbMEbk b¤ TTYlrgGuksIutkmμ edIm,Ipþl;famBlEdlekasikaRtUvkarTaMgenH . rbbGaharrbs;eyIgrYmmanRbPB sMxan;bIénfamBl ³ kabUGIuRdat xøaj; nig RbUetGIun . RbePTnimYy²kñúgcMeNamBYkvaenHénm:UelKul CIvsaRsþFM² RtUv)anbMEbkCaÉktabnÞab; ¬sársamBaØ fatty acids nig glycerol nig GasIutGamIneN¦ muneBlBYkvaGacRtUv)ancab;cUleTAkñúgekasika nig RtUv)aneRbIsMrab;plitfamBlkñúgekasika . kñúg cMeNamcMNat;fñak;TaMgenHénm:UelKulcMNIGaharkabUGIuRdatRtUv)anbMEbkeKehAfaCa Glycolysis . eyIgGaceXIjpøÚvdUcKñaenAkñúgsrIragÁ (organisms) xusBI )ak;etrI nig mnusS . enAkñúgCMBUkenH eyIgnwgBinitüCMhanénpøÚvEdlRbmUlfamBlBImunmkenH . eyIgnwgeXIjfava TTYlxusRtUvcMeBaHkarcab;yks<anfamBl (bond energy) mYycMnYnrbs;kabUGIuRdat nig karsþúkTuk nUvfamBlenaHeRkamTMrg;Ca ATP (adenine triphosphate) . CaFm μta Glycolysis beBa©j nig sþukTuk cMnYntictYc (2.2 %) énfamBlEdlmanGnuPaB (potential energy) rbs; glucose b:uEnþpøÚvenH k¾eRbICa RbPBénTItaMgksagCIvsMeyaK (biosynthetic building blocks) . vak¾)anbMElgkabUGIuRdatkñúgrebob enHEdlpøÚvepSgeTotGacbeBa©jfamBlEdlmanGnuPaBeRcIn (40 %) .



taragTI 21-1 RbePTénkargarekasikaEdlRtUvkarfamBl

CIvsMeyaK ³ sMeyaKénemtabUlItkNþal nig m:UelKulFM²karsMeyaKKøuykUsBI CO2 nig H2O enAkñúgdMeNIrkarrs μ IsMeyaKenAkñúgrukçCati karsMeyaKGasIutGamIneNkarsMeyaKnuyekøGUTId karsMeyaKlIBId karsMeyaKRbUetGIunBIGasIutGamIneNkarsMeyaKGasIutnuyekøGiuckarsMeyaKsrIr³ (organelles) nig Pñas (Membranes)

kardwkCBa¢Únskmμ ³ clnaGIuy:ug nig m:UelKulkardwkCBa¢Ún H+ kñúgkarrkSa pH [enAesßrkardwkCBa¢Únm:UelKulGaharcUleTAkñúgekasikakardwkCBa¢Ún K+ nig Na+ cUl nig ecjBIekasikasésRbsaTsMrab;bBa¢Únrlk sésRbsaT karbeBa©j HCl BIekasika parietal cUleTAkñúgRkBHkardwkCBa¢ÚnkaksMNl;BIQamcUleTAkñúgTwkenamtamry³tMrgenamkardwkCBa¢ÚnGasIutGamIneN nig sárEdlmanGatUmkae)an 6 cUleTAkñúgQam BIeBaHevon karRbmUlpþúMénGIuy:ug Ca enAkñúgmItUkugRDIclna karkRnþak; nig karrlaénekasikasac;dMukarbMEbkénRkUm:UsUmenAeBlbMEbkekasikalT§PaBrbs;Es<mkñúgkarEhledaysarknÞúyclnaén foreign substances epSgBIpøÚvdegðImedaysareram enAelIRsTab; epithelial rbs;bMBg;xül; karpøas;TItaMgrbs;Bg (eggs) cUleTAkñúgbMBg; fallopian edaysareram enAkñúgpøÚvbnþBUCRsþI



1- ATP : rUbiyvtßúfamBlénekasika (ATP : The Cellular Energy Currency)

karbMEbkénm:UlelKulfamBl (fuel molecules) EdleKehAfaCa katabUlIs pþl;nUvfamBl sMrab;muxgarEdlRtUvkarfamBlénekasika rYmman GaNabUlIs b¤ CIvsMeyaK . CaFm μta famBlén RbPBBIGaharGacRtUv)anbeBa©jeTAtampøÚvmYykñúgcMeNampøÚvBIr CakMedA b£ vasMxan;CagenHeTA[ ekasika CafamBléns<anKImI . eyIgGacemIleXIjrebobBIr (two alternative modes) én karbMEbk Edlmanxül; (aerobic degradation) rbs;KøúykUs . edaykareRbobeFobn½yya:gsamBaØ eyIgGac dak;KøúykUsCaeRKOgeqH . karenHbNþal[manGuksIutkmμeBjelj (complete oxidation) [eTACa CO2 nig H2O nig GacbeBa©j 686 kcal/mol rbs;KøúykUs . \LÚvenH kúñglkçx½NÐénekasika etIGIVEdl nwgRtUvsMerc)an ? minmanGIVeT RKb;famBlEdlmanGnuPaBéns<anrbs;KøúykUsKWRtUv)an)at;bg;Ca kMedA nig BnøW . karbeBa©jd¾xøaMgénfamBlnwgeFIV[xUcGg;sIum nig RbUetGIunepSgeTot nig bMpøaj ekasika . cUrBicarNapøÚvepSgeTot GuksIutkmμbnþicmþg²rbs;KøúykUsnaMmkCamYyedaykarbeBa©jnUvcMnYn tictYcénfamBlenAelIcMNucCaeRcInenAkñúgpøÚvemtabUlIs . RbsinebIvaenHCayuT§saRsþd¾eCaKC½y cMeBaHekasika vanwgRtUvkarGg;sIummYyes‘rIedIm,IsMrYl (catalyze) GuksIutkm μbnþicmþg² (stepwise

oxidation) nig meFüa)ayénkarrkSafamBlKImIsMrab;kareRbIedayRbtikmμEdlRtUvkarfamBl . CaRTwsþI karenHGacsMerc)anedaykarrYmKñaedaypÞal;énRbtikm μbMPayfamBl (exergonic

reactions) nig Rbtikm μRsUbfamBl (endergonic reactions) . enAeBlekasikamantMrUvkarfamBl vaRtUveqHedayRsYlnUvRbPBfamBl nig famBlEdlRbmUl)ansMrab;kargar . yuT§saRsþenHRtUvkarGIV EdlCaTUeTARbPBfamBlman nig EdlGIVenAq¶ayBIkarBit . vak¾RtUvkarEdrnUvGIVEdlkarbeBa©jfamBl nig RbtikmμKImIEdlRtUvkarfamBlRtUv)anrYmKñaedayeKarBeTAtamcMnYn nig TMrg;énfamBl . mYyQut²énRbtikmμRtUvCYbRbTHkarraMg (custom) EdlerobcMeLIg ehIyRbEhlEdlRtUvkarnUvcMnYn d¾eRcInénRbtikmμRtLb; . karsMeyaKGg;sIumRtUvkarrkSaRbB½n§rebobenHnwgcMNayfamBld¾eRcInsMrab; ekasika . vamanRbsiT§PaBxøaMgCagq¶aykñúgkarman dMNak;karkNþal (go-between) EtmYyKt;edIm,I dak;rYmKñanUvRbtikmμbMPayfamBl nig RsUbfamBlenAkñúgekasika . m:UelKulEtmYyKt;sMrab;karsþúk TukfamBlKImIEdlGaceRbICa rUbiyvtßúfamBlskl (Universal energy currency) . karenHKWCaGIV EdlekIteLIgedayBitR)akdenAkñúgekasika . m:UelKulEdleRbIsMrab;muxgarenHKW Adenosine

triphosphate (ATP) . RbsinebI ATP KWCaTMrg;sþúksMxan;énfamBlKImI vaRtUvEtGacTTYlykfamBl



EdlRtUv)anmkBIpøÚvemtabUlIsEdlbeBa©jfamBl nig pþl;nUvfamBlcMeBaHpøÚvemtabUlIsEdlRtUvkar famBl . karsMgat;cMeBaHmuxgarrbs; ATP CaFatukNþalEdlsMGagelIrcnasm<½n§rbs;vadUcEdl)an bgðajenAkñúgrUbTI 21-1 .

rUbTI 21-1 rcnasm<½n§énrUbiyvtßúfamBlskl ³ ATP

ATP KWCanuyekøGUTIdpSMeLIgGMBI Adenine P¢ab;CamYynwg Ribose . Cabnþ Ribose P¢ab;eTAnwg mYy (AMP) BIr (ADP) bI (ATP) bgÁúMhVÚssVÚrIl (Phosphoryl group) edaysm<½n§ hVÚssVÚeGEsÞr . m:UelKulenaHKWCasmasFatuEdlmanfamBlx<s; BIeRBaHsm<½n§ phosphoanhydride manbgÁúM phosphoryl enAxagcugbBa©b; . enAeBlsm<½n§enHdac; b¤GIuRdUlIs (hydrolyzed) BYkvabeBa©jnUv famBlCaeRcInEdlGacRtUv)aneRbIsMrab;kargarrbs;ekasika . cMeBaHehtuplenH eKGacehAvafaCa sm<½n§EdlmanfamBlx<s; (high-energy bonds) .sm<½n§EdlmanfamBlx<s;rbs; ATP GacbgðajCa nimμ isBaØa “~”dUcenAkñúgrUbTI 21-1 . rcnasm<½n§rbs; ATP KWRKan;EtCaEpñkmYyénehtuplEdlm:UelKulKWCaFatukNþald¾l¥enAkñúg karbMElgfamBlenAkñúgekasika . ATP RtUvEtmanfamBlx<s;CagsmasFatuEdlvapþl;famBl nig

OH OH

O O O

װ װ װ ¯O─P~O─P~O─P~O─CH2 | | | ¯O ¯O ¯O

Ribose

Adenosine

AMP

ADP

ATP

Phosphoanhydride bonds

Phosphoester bonds

Adenine



manfamBltiCagsmasFatuEdlCab;Tak;TgkñúgkarbegáItva . enAkñúgrebobenH enARKb;Rbtikm μman sar³sMxan;BIeRBaHTaMgRbtikmμbegáIt ATP nig GIuRdUlIs ATP edIm,Ipþl;nUvfamBlsMrab;kargarekasika KWCaRbtikmμPayfamBl . rUbTI 21-2 bgðajBIfamBlEdlTak;TgénsmasFatuEdlmanbgÁúM phosphoryl rYmman ATP EdlCab;Tak;TgenAkñúgemtabUlIsfamBl .

rUbTI 21-2 kareRbobeFobfamBlénsmasFatuEdlman bgÁúM Phosphoryl Edlman

sarsMxan;enAkñúgdMeNIrkaremtabUlIsekasika

GIuRdUlIs ATP naM[)anCa ADP nig )anCabgÁúM Phosphate minEmnsrIragÁ (Pi) nig famBl ¬rUbTI 21-3¦ . famBlEdlbeBa©jedaykarGIuRdUlIs ATP enH bnÞab;mkRtUv)aneRbIedIm,ICMrujdMeNIr karCIvsaRsþ GaRs½yedaykarP¢ab;bgÁúM phosphate eTAelI glucose b¤ fructose . ]TahrN_énviFIenHEdlfamBlrbs; ATP RtUv)aneRbIGaceXIjmanenAkñúgdMNak;kardMbUgén glycolysis karbMEbkedayminmanxül; (anaerobic degradation) rbs; glucose edIm,IbegáItnUvfamBl KImI . CMhandMbUgCab;Tak;TgeTAnwgkarbBa¢ÚnénbgÁúM Phosphate (—PO3

2−) BI ATP eTAbgÁúMGIuRduksIul enAelI C-6 rbs; glucose ¬rUbTI 21-4¦ . Rbtikm μenHRtUv)ansMrYledayGg;sIum Hexokinase .

Phosphoenolpyruvate 1,3-Bisphosphoglycerate Creatine phosphate ATP

Glucose-1-phosphate Fructose-6-phophate Glucose-6-phosphate

-14.8 -11.8 -10.3 -7.5

-5.0 -3.3

ATP

ADP

ADP

~P

~P

~P

~P

~P ~P

Go between



rUbTI 21-2 GIuRdUlIsén ATP eTACa ADP dac;nUvsm<½n§ Phosphoanhydride nig

beBa©jbgÁúM Phosphate minEmnsrIragÁ nig famBl

Adenosine triphosphate

H2O

Phosphate

rUbTI 21-4 karbBa¢ÚnbgÁúM Phosphoryl BI ATP eTAelIbgÁúMGIuRduksIulenAelI C-6 rbs; Glucose

OH OH

O O O

װ װ װ ¯O─P~O─P~O─P~O─CH2 | | | ¯O ¯O ¯O Ribose

Adenine

OH OH

O O

װ װ HO─P~O─P~O─CH2 | | ¯O ¯O Ribose

Adenine

O¯ | ¯O─P─O¯ || O

CH2OH

OH

OH

HO

O−

І CH2—O—P═O І O−

OH

OH

HO

Hexokinase + ATP + ADP



RbtikmμenHGacRtUv)ankat;edIm,IbgðajBItYnaTIrbs; ATP CaRbPBénfamBl . eTaHbICa karenHKWCaRbtikmμrYmKña eyIgGacKitGMBIvafaCadMeNIrkarEdlmanBICMhan . CMhanTImYyKWCaGIuRdUlIs ATP [eTACa ADP nig Pi (Phosphate) . karenHKWCaRbtikmμbMPayfamBlEdlbeBa©jRbEhl 7

kcal/mol . ATP + H2O → ADP + Pi + 7 kcal/mol

CMhanTIBIr karsMeyaKén Glucose-6-Phosphate BI Glucose nig Phosphate KWCaRbtikmμRsUb famBlEdlRtUvkar 3 kcal/mol .

3 kcal/mol + Glucose + Pi → Glucose-6-Phosphate + H2O

bnÞab;mkRbtikmμTaMgKImITaMgBIrGacRtUvbEnßmedIm,Ipþl;nUvsmIkarEdlbgðajviFIEdlkareFIVGIuRdUlIs ATP

rYmCamYynwgkarP¢ab;bgÁúM Phosphate eTAelI glucose ³ ATP + H2O → ADP + Pi + 7 kcal/mol 3 kcal/mol + Glucose + Pi → Glucose-6-Phosphate + H2O ----------------------------------------------------------------------------------------------

srub ³ ATP + Glucose → Glucose-6-Phosphate + ADP + 4 kcal/mol

GaRs½yedaykareFIVGIuRdUlIsén ATP beBa©jfamBleRcInCagRtUvkarsMrab;karsMeyaK glucose-6-

Phosphate BI glucose nig phosphate . vamankarsrubfamBlEdlbeBa©jenAkñúgdMeNIrkarenH nig RbtikmμbnþedayÉkÉgeq<aHeTAxagsþaM . plitpl “Glucose-6-Phosphate” manfamBleRcInCag Rbtikr “glucose” BIeRBaHva\LÚvenHvamanpÞúkfamBlBIsm<½n§ Phosphoanhydride EdlCasm<½n§edIm rbs; ATP . muxgarCabzmrbs;RKb;pøÚvbMEbk (catabolic pathway) KWedIm,IRbmUlfamBlKImIBIm:UelKul EdlGaceqH (fuel molecules) nig sþúkTukfamBledaykarplit ATP . karplitCanic©rnþén ATPenH KWCaGIVEdlpþl;nUvfamBlEdlmanGnuPaBRtUv)ansþúkTukEdlRtUv)aneRbIenAkñúgkareFIV[dMeNIrkar nUvmux garekasikaPaKeRcInbMput . 2- TidæPaBTUeTAéndMeNIkarkatabUlIs (Overview of Catabolic Processes)

eTaHbICakabUGIuRdat xøaj; nig RbUetGIu BYkvaTaMgGs;RtUv)anbMEbkedIm,IbeBa©jfamBl kabUGIuRdatKWCaRbPBfamBlEdleRbIeRcInCageKbMputrYcmkehIy . eyIgnwgcab;epþImedaykarBinitü GuksIutkmμrbs; glucose . enAkñúgCMBUkTI 22 nig 23 eyIgnwgeXIjpøÚvGuksIutkmμrbs; glucose k¾RtUv )aneRbIsMrab;karbMEbknUvxøaj; nig RbUetGIun .



dMeNIrkarénkarbMEbkRtUvEtcab;epþImedaykarpþl;nUvcMNIGahar . enAeBleyIgbriePaKGahar eyIgbriePaKbrimaNénkabUGIuRdat xøaj; nig RbUetGIun . BIcMNucenH dMeNIrkarbMEbk (catabolic

processes) RtUv)anbMEbkCadMNak;karmYyes‘rI . dMNak;karTaMgbIénkatabUlIsRtUv)ansegçbenAkñúg rUbTI 21-5 .

rUbTI 21-5 dMNak;karTaMgbIénkarbMElgcMNIGahareTACafamBlkñúgekasikaeRkamTMrg;Ca ATP

k- dMNak;karTI 1 ³ GIuRdUlIsm:UelKulcMNIGahareTAm:UelKultUc² eKalbMNgéndMNak;karTImYyrbs;katabUlIsKWedIm,IbMEbkm:UelKulFM²rbs;cMNIGahar eTACa smasFatubnÞab;tUc²rbs;BYkva . bnÞab;mksmasFatubnÞab;tUc²TaMgenH ¬sársamBaØ GasIutGamIneN GasIutRka (fatty acid) nig KøIesr:Ul (glycerol) ¦ RtUv)ancab;ykedayekasikadgxøÜnsMrab;kareRbIR)as; CaRbPBfamBldMeNIrkarénkarrMlayGaharRtUv)ansegçbenAkñúgrUbTI 21-6 .

Food

CarbohydrateProteins Fats

Amino Simple sugars Fatty acids & glycerol

Glycolysi ATP

Pyruvate

AcetylCoA

Citric Acid cycle

Oxidative phosphorylation

ATP

1. Hydrolysis of macromolecules to subunits

2. Conversion of subunits to a form that can be complete oxidize, usually Acetyl CoA

3. Complete oxidization of Acetyl CoA and the production of ATP



b:UlIsak;karId_RtUv)anGIuRdUlIseTACam:UNUsak;karId_ . dMeNIrkarenHcab;epþImenAkñúgmat; EdlenATIenaH Amylase cab;epþImkareFIVGIuRdUlIsGamIdug (starch) . karrMlayGaharbnþeTAkñúg eBaHevontUc EdlenATIenaH Amylase rbs;lMEBgbnþeFIVGIuRdUlIsGamIdugbEnßmeTot[eTACa ma:l;tUs_ ¬DIsak;karId_EdlmanBIrm:UlelKul glucose ¦ . ma:l;tas_ (Maltase) sMrYlkareFIVGIuRdUlIsma:l;tUs_ edaybegáIt)anBIrm:UelKul glucose . dUcKñaenHEdr sak;kar:Us_ (sucrose) RtUv)anGIuRdUlIs[eTACa glucose nig fructose edayGg;sIumeQμaH sak;karas_ (sucrase) nig Lak;tUs_ ¬sárTwkedaH¦ RtUv)an bMEbkCa glucose nig galactose edayGg;sIum Lak;tas_enAkñúgeBaHevontUc . m:UNUsak;karId_RtUv)an cab;edayekasikaeGBIetlIya:l; (epithelial cells) rbs;eBaHevonenAkñúgdMeNIrkarEdlRtUvkarfamBl EdleKehAfaCa kardwkCBa¢Únskmμ .

rUbTI 21-6 karBinitüTUeTAéndMeNIrkarEdleFIVGIuRdUlIskabUGIuRdat xøaj; nig RbUetGIun

karrMlayénRbUetGIuncab;epþImenAkñúgRkBH EdlenATIenaH pH TabeFIV[xUcrcnasm<½n§rbs; RbUetGIundUecñHBYkvaRtUv)anGIuRdUlIsedayRsYledayGg;sIumeQμaH Pepsin . BYkvak¾RtUv)anbMEbkbEnßm



eTotenAkñúgeBaHevontUceday Trypsin, chymotrypsin, elastase nig Proteases epSgeTot . plitplénkarrMlayRbUetGIun ¬GasIutGamIneN nig oligopeptides¦ RtUv)ancab;yedayekasikaEdl enAelIépÞeBaHevontUc . karcab;ykenHk¾Cab;Tak;TgeTAnwgynþkarénkardwknaMskmμ . karrMlayénxøaj;minTan;cab;epþImeT RbsinebIGaharminTan;eTAdl;eBaHevontUc eTaHbICaman lipases enATaMgkñúgTwkmat; nigrsRkBH . xøaj;eTAdl;kñúgeBaHevon (duodenum) EpñkdMbUgén eBaHevontUc enAeRkamTMrg;CaeKalikaFM² . GMbilTwkRbma:t; (Bile salts) EdlplitedayeføIm bMEbkBYk xøaj;[eTACa emulsion éndMNk;xøaj;tUc² . GaRs½yedaydMNk;xøaj;tUc²manépÞFM \LÚvenHlIBId gaynwgTTYlnUvGMeBIrbs; lipase lMEBg . Gg;sIumenHGIuRdUlIsxøaj;[eTACaGasIutRka nig KøIesr:Ul EdlRtUv)ancab;ykedayekasikaeBaHevonedaydMeNIrénkardwknaMEdlminRtUvkarfamBl . dMeNIrkar enHRtUv)aneKehAfaCa kardwkCBa¢ÚnGkmμ (passive transport) . RbtikmμGIuRdUlIsTaMgenHRtUv)an segçbenAkñúgrUbTI 21-7 .

rUbTI 21-7 Rbtikm μGIuRdUlIsénkabUGIuRdat xøaj; nig RbUetGIun

CH2OH

OH

OH

O

CH2OH

OH

OH

HO + H2O →

CH2OH

OH

OH HO

CH2OH

OH

OH HO

+

Glycosidase

Disaccharide Monosaccharides

H H O H O H R O H R O І І ІІ І ІІ Peptidase І І ІІ І І ІІ H —N—C—C—N—C—C—OH + H2O → H—N—C—C—OH + H—N—C—C—OH І І І І І R H R H H Peptide Amino acids

C17H35—CO—O—CH2 CH2OH І Lipase І C17H35—CO—O—CH2 + H2O → 3 (C17H35—COOH) + CH2OH І І C17H35—CO—O—CH2 CH2OH Triglyceride Fatty acid Glycerol



x- dMNak;karTI 2 ³ karbMElgm:UNUEmr[eTACaTMrg;EdlGacTTYrgGuksIutkmμeBjelj \LÚvenH m:UNUsak;karIt GasIutGamIneN GasIutRka nig KøIesr:UlRtUvEt)anRsUbnigbMEbkeTAkñúg pøÚvénemtabUlIsfamBl . pøÚvsMxan;BIrKW glycolysis nig Citric acid cycle ¬emIlrUbTI21-5¦ . CaFm μtaBYksár cUleTAkñúgpøÚv glycolysis eRkamTMrg;Ca glucose nig Fructose . CacugbBa©b;BYkvaRtUv )anRtUv)anbMElgeTACa AcetylCoA EdlCaTMrg;GacTTYlrgnUvGuksIutkm μeBjeljenAkñúg Citric acid

cycle . Amino groups RtUv)andkecjBIGasIutGamIneN nigenAEtKMeragExSkae)ancUleTAkñúgdMeNIkar katabUlIsenAelIdMNak;karCaeRcInrbs; Citric acid cycle . GasIutRkak¾RtUv)anbMElgeTACa AcetylCoA Edr nigcUleTAkñúg Citric acid cycle kñúgeRkamTMrg;enH . KøIesr:Ul RtUv)anbegáIteLIgeday kareFIVGIuRdUlIsénxøaj; RtUv)anbMElgeTACa glyceraldehydes-3-phosphate EdlKWCasmasFatu kNþalmYykñúgcMeNamsmasFatukNþalCaeRcInrbs; glycolysis nig cUleTAkñúgemtabUlIsfamBl enAnwgkMritenaH . K- dMNak;karTI 3 ³ GuksIutkmμeBjeljrbs;Gaharnigkarplit ATP

AcetylCoA manGatUmkae)anBIrEdlsl;BIGahar ¬bgÁúMGaesTIl¦cUleTAkñúg Citric acid

cycle. AcetylCoA cUleTAkñúgvdþ (the cycle) nig eGLikRtug nig GatUmGIuRdUEsnRtUv)anRbmUlykenA kñúgry³eBlénGuksIutkm μeBjeljrbs;bgÁúMGaesTIleTACa CO2 . Coenzyme A RtUv)anbeBa©j ¬RtLb;mYyCMueTot¦ edIm,IdwkbgÁúMGaesTIlbEnßmeTotcUleTAkñúgpøÚvemtabUlIs . eGLikRtug nig GatUm GIuRdUEsnEdlRtUv)anRbmUlykRtUv)aneRbIenAkñúgdMeNIrkar Oxidative phosphorylation edIm,IbegáIt ATP . 3- KøIkUlIs (GLYCOLYSIS)

k- karBinitüTUeTA Glycolysis k¾RtUv)aneKsÁal;pgEdrfaCa pøÚv Embden-Meyerhof KWCapøÚvsMrab;karbMEbkBYk kabUGIuRdat Edlcab;epþImCamYynwg Substrate Ca D-glucose . karBitrebobenHEdlfaRKb;srIragÁ (organisms) GaceRbI glucose CaRbPBfamBlcMeBaH glycolysis [Caeyabl;fa glycolysis KWCapøÚv EdlRbmUlykfamBledayeCaKC½ydMbUgeK Edlvivtþn_bnþicmþg²enAelIdI . pøÚvEdlvivtþn_enAkñúgeBl mYyenAeBlbriyakasrbs;EpndIminmanGuksIuEsn ¬minmanGuksIuEsnesrI¦ . CalT§pl Glycolysis

minRtUvkarGuksIuEsnKWCadMeNIrkarminmanxül; (anaerobic process) . bEnßmelIenHeTot vaRtUvEt)an vivtþn_enAkñúgrebobya:gsamBaØ ¬srIragÁÉkekasika¦ dUcCa)ak;etrI . srIragÁTaMgenHminmansrIr³ (organelles) s μ úRKsμajenAkñúgsIutU)aøs;edIm,IGnuvtþn_muxgarBiessrbs;ekasika . dUecñH Glycolysis



KWCadMeNIrRtUv)anGnuvtþedayGg;sIumEdlenAedayesrIenAkñúgsIutU)aøs;. rhUtmkdl;bc©úb,nñ glycolysis

enAEtCadMeNIrkarminmanxül; (anaerobic process) EdlRtUv)anGnuvtþedayGg;sIumrbs;sIutU)aøs; eTaHbICaenAkñúgekasikasμ úRKsμajrbs;mnusSeyIgk¾eday . db;dMNak;karén Glycolysis RtUv)anKUsbBa¢ak;enAkñúgrUbTI 21-8 . Substrate TImYyenAkñúg pøÚvenHKWCa glucose . Gg;sIumdb;RtUvkaredIm,IGnuvtþRbtikm μrbs;pøÚvenH . RbtikmμTImYyén glycolysis

Cab;Tak;TgeTAnwgkarvinieyaKfamBl . m:UelKul ATP RtUv)anGIuRdUlIs famBlRtUv)anbeBa©j nig bgÁúM phosphoryl RtUv)anbEnßmeTAelIsárEdlman6 GatUmkae)an . enAkñúgCMhanEdlminERbRbYlrbs; glycolysis RtUv)anRbmUlykedIm,IbegáItkarTTYlsrubrbs; ATP . plitplsMxan;bIrbs; glycolysis RtUv)aneKeXIjenAkñúgrUbTI 21-8 . BYkvaTaMgenHKWCa famBlKImIenAeRkamTMrg;Ca ATP famBlKImIenAeRkamTMrg;Ca NADH nig BIrmU:elKul Pyruvate Edl manbIGatUmkae)an . plitplnimYy² TaMgenHRtUv)aneKBicarNaxageRkam ³ famBlKImIeRkamTMrg;Ca ATP ³ bYnm:UelKul ATP RtUv)anbegáItedayedaydMeNIrkarén karP¢ab; Phosphate enAelI Substrate . karenHmann½yfa bgÁúM phosphoryl EdlmanfamBlx<s; BI substrate mYykñúgcMeNam substrates enAkñúg glycolysis RtUv)anbMElgeTACa ADP edIm,IbegáIt)an Ca ATP . Substrates BIrCab;Tak;TgenAkñúgRbtikm μbMElgTaMgenHKW 1,3-biphosphoglycerate nig phosphoenolpyruvate ¬cUremIlrUbTI 21-8 CMhanTI 7 nig TI 10¦ . eTaHbICa bYnm:UelKul ATP

RtUv)anbegáItenAeBlman glycolysis karTTYl)ansrubEtBIrm:UelKul ATP BIeRBaHBIrm:UelKul ATP

RtUv)aneRbIBIedImdMbUgén glycolysis ¬rUbTI 21-8 CMhanTI 1 nig 3¦ . BIrm:UelKul ATP RtUv)anbegáIt tMNagEt 2. 2 % énfamBlEdlmanGnuPaBrbs;m:UelKul glucose . dUecñH glycolysis minEmndMeNIr karRbmUlfamBlEdlmanRbsiT§PaBxøaMgeT . famBlKImIeRkamTMrg;Ca reduced NAD+, NADH ³ Nicotinamide adenine

dinucleotide (NAD+) KWCa Coenzyme manRsLayBI Vitamin niacin . TMrg; reduced NAD+,

NADH man hydride anions (H:−) ¬GIuRdUEsnEdlmaneGLikRtugBIr¦ RtUv)andkecjenAkñúgry³eBl GuksIutkmμrbs; substrate mYykñúgcMeNam substrates (glyceraldehydes-3-phosphate) ¬cUremIl rUbTI 21-8 CMhanTI 6¦ . eRkamlkçx½NÐmanxül; (aerobic conditions) eGLikRtug nig GatUmGIuRdUEsnRtUv)andwknaMBIsIutU)aøs;cUleTAkñúgmItUkugRDI (Mitochondria) . enATIenH BYkvacUleTAkñúg RbB½n§dwknaMeGLikRtugsMrab;karbegáIt ATP eday Oxidative phosphorylation . sßitenAeRkam lkçx½NÐK μanxül; NADH RtUv)aneRbICaRbPBrbs;eGLikRtugenAkñúgRbtikmμeLIgem (fermentation

reactions) .



rUbTI 21-8 Rbtikmμrbs; Glycolysis ³ RbtikmμTaMgenHsßitkñúgsIutU)aøs;

Dihydroxyacetone Phosphate

Glyceraldehyde 3-Phosphate

5

Glucose

Glucose-6-Phosphate

Fructose-6-Phosphate

Fructose-1,6-Phosphate

Hexokinase ATP

ADP

Phosphoglucoisomerase

Phosphofructokinase

Aldolase

1

2

3

4

ATP

ADP

1,3-BiPhosphoglycerate

3-Phosphoglycerate

2-Phosphoglycerate

Phosphoenolpyruvate

Pyruvate

Glyceraldehyde-3-phosphate dehydrogense

Phosphoglycerokinase

Phosphoglyceromutase

Enolase

2ADP

2ATP

6

7

8

9

10

2ADP

2ATP

2H2O

Pyruvate kinase

2NAD+ 2Pi

2NADH + H+



BIrm:UelKul Pyruvates ³ enAcugbBa©b;én glycolysis Edlm:UelKul glucose Edlman 6 GatUmkae)anRtUv)anbMElgeTACaBIrmU:elKul Pyruvates Edlman3GatUmkae)an . vasna (the fate)

rbs; Pyruvate k¾Cab;Tak;TgEdreTAnwgfaetIRbtikm μRtUv)anekIteLIgenAeBlmanvtþman b¤ Gvtþman rbs;GuksIuEsn . sßitenAeRkamlkçx½NÐmanxül; vaRtUv)aneRbIedIm,IbegáIt AcetylCoA EdleRbIsMrab; Citric acid cycle nig kareFIVGuksIutkmμeBjelj . eRkamlkçx½NÐminmanxül; vaRtUv)aneRbICaGñkcab; eGLikRtugenAkñúgRbtikmμeLIgem (fermentation reactions) . enAkñúgsßanPaBNamYy plitplcugeRkayBIrenH RtUvEt)aneRbIenAkñúgrebobmYycMnYn dUecñH glycolysis

GacbnþbMeBjmuxgar nig plit ATP . vamanehtuplBIr cMeBaHsßanPaBenH . ehtuplTI 1 RbsinebI Pyruvate RtUv)anekIteLIg vanwgbgá[ glycolsis Qb;dMeNIrkar dUecñHenAeBlenH bBaÄb;kar plit ATP . dUecñH Pyruvate RtUvEt)aneRbIenAkñúgRbePTénRbtikm μtamdanmYycMnYn ¬kñúglkçx½NÐman xül; b¤ K μanxül; . ehtuplTI 2 enAkñúgCMhanTI 6 glyceraldehyde-3-phosphate TTYlrgnUv GuksIutkmμ nig NAD+ TTYlrgnUver:dukkmμ ¬cab;yk hydride anion¦ . ekasikamankarpÁt;pÁg; NAD+

EtcMnYntictYc . Rbsin RKb; NAD+ TTYlrger:dukm μ vaminman NAD+ sMrab;RbtikmμenH nig glycolysis

nwgRtUvQb; . ehtudUecñH NADH RtUvEtTTYlrgGuksIutkmμeLIgvij dUecñH glycolysis Gacbnþkarplit ATP sMrab;ekasika . x- RbtikmμKøIkUlIs (Reactions of Glycolysis)

rcnasm<½n§énFatukNþalrbs; glycolysis RtUv)aneKeXIjenAkñúgrUbTI 21-9 RsbCamYynigkar BIB’NnaBisþarénRbtikmμEdlekIteLIgenAkñúgdMNak;nimYy² nigeQμaHrbs;Gg;sIumEdlsMrYlRbtikmμ nimYy² . Glycolysis RtUv)anEckCaBIEpñksMxan; . EpñkTImYyKWCakarvinieyaKénfamBl ATP . Rbsin ebIKμankarvinieyaKenH glucose nwgminmanfamBlRKb;RKan;sMrab;bnþdMeNIrkar glycolysis nig vaminman karplit ATP . EpñkenHrYmmanRbtikmμ 5 dMbUgén pøÚv glycolysis . EpñksMxan;TI 2 Cab;Tak;TgeTAnwg RbtikmμEdlsßitesßr (remaining reactions) rbs;pøÚv glycolysis ¬dMNak;TI 6-10¦ ehIyRbtikmμ TaMgenHEdlbNþal[manfamBlekIteLIg .



rUbTI 21-9 FatukNþal nig Gg;sIumrbs;KøIkUlIs

(The intermediates and enzymes of glycolysis)

CH2OH

OH

OH

HO

CH2—O—P

OH

OH

HO

ATP ADP

Hexokinase (1)

CH2—O—P

OH OH

CH2— HO

1. Glucose RtUv)anP¢ab;bgÁúM phosphoryl edaycMNay

ATP edIm,IbegáIt G-6-P

Glucose

Phosphogluco-isomerase (2)

2. G-6-P RtUv)anerobcMeLIgvijedIm,IbegáItCa F-6-P

CH2—O—P

OH OH

CH2— HO—P

ATP ADP

Glucose-6-Phosphate (G-6-P)

Fructose-6-Phosphate (F-6-P)

Phospho-fructokinase (3)

3. F-6-P RtUv)anP¢ab;bgÁúM phosphoryl edIm,IbegáItCa F-1,6-

biP edaycMNay ATP mYyepSgeTot . karcMNay 2

ATP tMNag[karvinieyaKfamBledIm,IeFIV[ glucose

skmμsMrab;kareFIVGuksIutkmμ .

Fructose-1,6-biPhosphate (F-1,6-biP)

CH2—O—P І C═O І CH2OH

CH2—O—P І CHOH І C═O І H

Aldolase 4 nig 5. Adolase kat; F-1,6-biP eTACaBIr m:UelKulEdlman3GatUmkae)an dihydroxyacetone

phosphate nig glyceraldehyde-3-phosphate . dihydroxyacetone phosphate RtUv)anbMElgCa glyceraldehydes-3-Phosphate edayGg;sIum triose

phosphate isomerase .Dihydroxyacetone phosphate Glyceraldehyde-3-

phosphate



CH2—O—P І CHOH І C═O І H

2 Pi

2NAD+

2 NADH + H+

Glyceraldehyde-3-phosphate dehydrogenase (6)

Glyceraldehydes-3-phosphate

6. Glyceraldehyde-3-phosphate TTYlrgGuksIutkmμ nig NADH RtUv )anekIteLIg . bgÁúM Phosphate minEmnsrIragÁRtUv)anbBa¢Ún eTAelIbgÁúM carboxyle edIm,IbegáItCa 1-3-

biphosphglycerate .

2ADP

2 ATP

CH2—O—P І CHOH І C═O І O—P

3

2

1

CH2—O—P І CHOH І C═O І O−

2

2

2

1,3-Biphosphoglycerate

Phosphoglyce- rokinase (7)

7. ATP RtUv)anbegáItenAkñúgkarP¢ab;bgÁúM phosphate elIkTI 1 énpøÚv. bgÁúM phosphate

RtUv)anbBa¢ÚnBI substrate eTA ADP edIm,IbegáIt Ca ATP

3-Phosphoglycerate

CH2OH І CH—O—P І C═O І O−

2-Phosphoglycerate

Phospho glyceromutase (8)

8. bgÁúM Phosphoryl EdlenAelI C-3 rbs; 3-

Phosphoglycerate RtUv)an bBa¢ÚneTAkae)anTI 2 (C-2)

Enolase (9) 9. Dehydration én 2-phosphoglycerate eTACa phospho-enopyruvate EdlsMbUrfamBl

H2O

CH2 ІІ C—O~P І C═O І O−

2 ADP

2 ATP

CH3—CO—CO—O− Pyruvate

Phosphoenolpyruvate

10. karP¢ab;bgÁúM phosphate eTAelI substrate

cugeRkaybegáIt)anCa ATP nig Pyruvate

Pyruvate Kinase (10)



RbtikmμTI 1 Substrate (Glucose) RtUvP¢ab;CamYYy bgÁúM Phosphate edaysarGg;sIumeQμaH Hexokinase

enAkñúgRbtikm μP¢ab;bgÁúM Phosphate (Phosphorylation reaction) . RbPBénbgÁúM Phosphate KW ATP . BIdMbUg RbtikmμenHhak;dUcCapÞúyeTAnwgeKalbMNgrbs; katabUlIs (catabolism) ¬karplitén ATP¦. karcMNay ATP enAkñúgRbtikmμdMbUgenHRtUvEteKKitfaCakarvinieyaK . CaFm μta ekasikaFøak; cUleTAkñúg bMNulfamBl (energy debt) enAkñúgRbtikmμdMbUgTaMgenH . b:uEnþkarvinieyaKenH mansar³sMxan;Cadac;xatedIm,IeFIV[pøÚvKøIkUlIscab;epþIm (glycolysis pathway) . Hexokinase Glucose + ATP → Glucose-6-Phosphate (G-6-P) + ADP + H+

RbtikmμTI 2 G-6-P EdlekIteLIgenAkñúgRbtikmμTI 1 RtUv)anerobcMsareLIgvijedIm,IbegáItCaGIusUEmrén rcnasm<½n§KW Fructose-6-Phosphate (F-6-P) . Gg;sIumeQ μaH Phosphoglucoisomerase sMrYl GIusUEmrIkmμ (Isomerization) . lT§plKWfa C-1 rbs;sárEdlman6GatUmkae)anEdlTTYlrgRbtikmμ vaElgsßitenACarcnasm<½n§ExSbiTeTotehIy . karBinitüénrcnasm<½n§ExSlatbgðajfaGIusUEmrIkmμenH bMElg Aldose eTACa Ketose . Phosphoglucoisomerase

Glucose-6-Phosphate ⇄ Fructose-6-Phosphate

RbtikmμTI 3 karvinieyaKfamBlelIkTI 2 KWRtUv)ansMrYledayGg;sIumeQ μaH Phosphofructoskinase . s<an PhosphoanhydrideenAkñúg ATP RtUv)anGIuRdUlIs nig sm<½n§ Phosphoester rvagbgÁúM Phosphate

nig bgÁúM Hydroxyl EdlenAelI C-1 rbs; F-6-P RtUv)anbegáIteLIg . plitplKW Fructose-1,6-

bisphosphate (F-1,6-bisP) . Phosphofructokinase Fructose-6-Phosphate → Fructose-1,6-bisPhosphate + ADP + H+

RbtikmμTI 4 F-1,6-bisP RtUv)anbMEbkCaFatukNþalBIrEdlmanbIGatUmkae)an enAkñúgRbtikmμEdlsMrYl edayGg;sIumeQμaH Aldolase . plitplRbtikm μKW Glyceraldehyde-3-phosphate (G3P) nig Dihydroxyacetone phosphate . Adolase Fructose-1,6-bisPhosphate → Glyceraldehyde-3-phosphate + Dihydroxyacetone phosphate



RbtikmμT 5 BIeRBaHEt G3P KWCa substrate EtmYyEdlGacRtUv)aneRbIedayGg;sIumbnÞab;enAkñúgpøÚv emtabUlIs dihydroxyacetone phosphate RtUv)anerobcMsareLIgvij[eTACam:UelKulTI 2 KW G3P . Gg;sIumEdlsMrYlRbtikmμGIusUEmrenHKW Triose phosphate isomerase . Triose phosphate isomerase Dihydroxyacetone phosphate → Glyceraldehyde-3-Phosphate

RbtikmμTI 6 enAkñúgRbtikm μenH Gal;edGIud (Glyceraldehyde-3-Phosphate) TTYlrgGuksIutkmμeTACa GasIutkabuksIulik (Carboxylic acid) enAkñúgRbtikmμEdlsMrYleday Glyceraldehyde-3-

Phosphate dehydrogenase . RbtikmμenHKWCadMNak;kardMbUgkñúgkarRbmUlfamBl nig vaCab; Tak;TgeTAnwger:dukmμrbs; Coenzyme Nicotinamide adenine dinucleotide (NAD+) RbtikmμenH ekIteLIgBICMhan . CMhanTI 1 NAD+ TTYlrger:dukmμeTACa NADH enAeBlEdlbgÁúMGal;edGIud rbs; G3P TTYlrgGuksIutkm μeTACabgÁúMkabuksIul (Carboxyl group) . CMhanTI 2 bgÁúM Phosphate minEmn srIragÁRtUv)anP¢ab;eTAelIbgÁúMkabuksIuledIm,Ipþl;Ca 1,3-bisPhosphoglycerate . cUrkt;sMKal;fa sm<½n§fIμRtUv)anbgðajedaysBaØa “~” Edlbgðajfasm<½n§enHmanfaBlx<s; . RbtikmμenH nig RKb; RbtikmμEdlesßr (remaining reactions) rbs; glycolysis ekIteLIgBIrdgcMeBaH glucose mYym:UelKul

BIeRBaH glucose nimYy²)anRtUvbMElgeTACa BIrm:UelKul Glyceraldehyde-3-Phosphate . Glyceraldehyde- 3-Phosphate dehydrogenase Glyceraldehyde-3-Phosphate + NAD+ + Pi → 1,3-Bisphosphoglycerate + NADH + H+

RbtikmμTI 7 kñúgRbtikmμenH famBlRtUv)anRbmUlenAeRkamTMrg; ATP . Gg;sIum Phosphoglycero-

kinase sMrYlkarbBa¢ÚnénbgÁúM Phosphoryl BI 1,3-Bisphosphoglycerate eTA[ ADP . RbtikmμenHKW CakarP¢ab;bgÁúM Phosphate enAelI substrate TI 1 én glycolysis nig vabegáItCa ATP nig 3-

phosphoglycerate . vaKWCaRbtikm μrYm (a coupled reaction) EdlkñúgenaH sm<½n§ anhydride cMruHEdl manfamBlx<s;RtUv)anGIuRdUlIs nig famBlEdlbeBa©jRtUv)aneRbIedIm,ICMrujkarsMeyaK ATP ‘ Phosphoglycerokinase 1,3-Bisphophoglycerate + ADP + H+ → 3-Phosphoglycerate + ATP

RbtikmμTI 8 3-phosphoglycerate TTYlrgnUvRbtikmμGIusUEmrkmμ edIm,IbegáItCa 2-phosphoglycerate enA kñúgRbtikmμsMrYledayGg;sIum Phosphoglyceromutase . bgÁúM Phosphate EdlCab;nwg C-3 rbs;



3-phosphoglycerate RtUv)anbBa¢ÚneTA C-2 . Phosphoglyceromutase 3-Phosphoglycerate → 2-Phosphoglycerate

RbtikmμTI 9 enAkñúgdMNa;karenH Gg;sIum Enolase sMrYlRbtikmμdkTwk (dehydration) ecjBI 2-

phosphoglycerate . plitplEdlsMbUrfamBlKW Phosphoenolpyruvate CasmasFatuEdlman bgÁúM phosphate manfaBlx<s;CageKbMputenAkñúgemtabUlIs . Enolase 2-Phosphoglycerate → Phosphoenolpyruvate

RbtikmμTI 10 enATIenH eyIgeXIjkarP¢ab;bgÁúM Phosphate enAelI substrate cugeRkayeKbMputenAkñúgpøÚv emtabUlIsEdlRtUv)ansMrYleday Pyruvate kinase . Phosphoenolpyruvate eRbICaGñkpþl;nUv bgÁúM Phosphate EdlRtUv)anbBa¢ÚneTA[ ADP edIm,IbegáItCa ATP . Rbtikm μenHKWCaRbtikmμrYmmYy epSgeTot (another coupled reaction) EdlkareFIVGIuRdUlIsénsm<½n§ phosphoester enAkñúg Phosphoenolpyruvate pþl;nUvfamBlsMrab;karbegáItnUvsm<½n§ Phosphoanhydride rbs; ATP . plitplcugbBa©b;rbs; glycolysis KW Pyruvate . Pyruvate kinase Phosphoenolpyruvate → Pyruvate

eKKYrEtkt;sMKal;faRbtikmμBITI 6 dl;TI 10 ekIteLIgBIrdgcMeBaH glucose mYym:UelKul BIeRBaHeBlcab;epþImBIsárEdlman 6 GatUmkae)an RtUv)anbMEbkCaBIrm:UelKulEdlman 3 GatUm kae)an . dUecñHenAkñúgRbtikm μTI 6 BIrm:UelKul NADH RtUv)anbegáIteLIg nig mancMnYnsrub 4 ATP

RtUv)anpliteLIg ¬CMhanTI 7 nig CMhanTI 10¦ . karTTYl ATP srub BIpøÚvemtabUlIsenHKW manEt 2 m:UelKuleT BIeRBaHvamankarvinieyaK 2 m:UelKul ATP kñúgdMNak;dMbUgénpøÚvemtabUlIsenH . karvinieyaKenHRtUv)ansgmkvijeday 2 m:UelKul ATP EdlRtUv)anplitedaykarP¢ab;bgÁúM Phosphate enAelI substrate enAkñúgdMNak;karTI 7 . lT§plfamBlCaFm μtaRtUv)anpliteday karP¢ab;bgÁúM Phosphate enAelI substrate kúñgRbtikm μTI 10 . K- karEktMrUvénKøIkUlIs (Regulation of Glycolysis)

pøÚvEdlRbmUlfamBl dUcCa glycolysis eqøIytbeTAnwgesckþIRtUvkarfamBlrbs;ekasika . Rbtikmμrbs;pøÚvemtabUlIsekIneLIgenAeBlvamankarTamTar (demand) ATP) . Rbtikm μrbs;pøÚv emtabUlIsfycuHenAeBlvasMbUr ATP edIm,IeqøIytbeTAnwgtMrUvkarfamBlrbs;ekasika .



ynþkarmYykñúgcMeNamynþkarCaeRcInsMrab;karRtYtBinitüpøÚv glycolysis KWkarRbI Allosteric

enzymes . epSgBITIkEnøgskmμ (active site) EdlcgP¢ab;eTAnwg substrate Gg;sIum Allosteric man TIkEnøgcgP¢ab; effector (effector) EdlcgsBaØaCatiKImI (chemical signal) EdlmanlT§PaB[sBaØa dl;GRtaEdlGg;sIumsMrYlRbtikmμ . karcg effector GacbegáIn (positive allosteric) b¤ bnßyGRta rbs;Rbtikmμ (negative allosteric) . sBaØaCatiKImI (chemical signals) b¤ effector EdlbgðajBIesckþIRtUvkarrbs;ekasikarYmman m:UelKuldUcCa ATP . RbsinebIkMhab; ATP x<s; ekasikaRtUvEtmansþúkbMrugfamBlRKb;RKan; . dUcKñaenHEdr ADP nig AMP EdlCa precursors rbs; ATP KWCaFatubnÞal; (indicators) Edlfa ekasikakMBugRtUvkar ATP . CakarBit RKb;m:UelKulTaMgenHKWCa Allosteric effectors Edl[sBaØaBI GRtaRbtikmμRLb;mkvijEdlRtUvsMrYledayGg;sIumenAkñúgpøÚv glycolysis . Gg;sIum Hexokinase EdlsMrYlRbtikm μénkarP¢ab;bgÁúM Phosphate eTAelI glucose RtUv)an raraMgedaysar\T§iBl allosteric edaysarplitplrbs;Rbtikm μEdlvasMrYl (Glucose-6-

Phosphate) . karekIneLIg (buildup) énplitplbgðajfaRbtikmμ glycolysis minbnþenAkñúgGRta elOn edaysn μtBIeRBaHkarTamTarfamBlrbs;ekasikaRtUv)aneqøIytb . Phosphofructokinase Gg;sIumEdlsMrYlRbtikm μTI 3 enAkñúg glycolysis KWCaGg;sIumEktMrUvCa sMxan; (a key regulatory enzyme) enAkñúg glycolysis . ATP KWCa allosteric inhibitor rbs; Phosphofructokinase rIÉ ADP nig AMP KWCa Allosteric activators . Allosteric inhibitor rbs; Phsophofructokinase epSgeTot KW sIuRtat (citrate) . dUcEdleyIgnwg)aneXIjenAkñúgCMBUkbnÞab;enH sIuRtatKWCaFatukNþalTI 1 enAkñúg Citric acid cycle . Citric acid cycle KWCapøÚvemtabUlIs EdlnaM[ manGuksIutkm μeBjeljrbs; Pyruvate EdlRtUv)anpliteday glycolysis . kMhab;rbs;sIuRtatx<s; [sBaØafa substrate RKb;RKan;kñúgkarcUleTAkñúg Citric acid cycle . karraraMgén Phosphofructokinase edaysIuRtat KWCa]TahrN_mYyén Feedback inhibition ³ plitpl ¬sIuRtat¦ raraMgskm μPaBrbs;Gg;sIumeday\T§iBl allosteric BIdMbUgenAkñúgpøÚvemtabUlIs . Gg;sIumcugeRkayén glycolysis KW Pyruvate kinase RtUv)anTTYlrgnUvkarEktMrUv allosteric

Edr . enAkñúgkrNIenH Fructose-1,6-Bisphosphate KWCa allosteric activator . eKmankar cab;GarmμN_fa Fructose-1,6-Bisphosphate KWCaplitplrbs;Rbtikm μEdlsMrYleday Phosphofructokinase . dUecñH karCMruj[manskmμPaB (activation) én Phosphofructokinase

bNþal[mankarCMruj[manskmμPaBrbs; Pyruvate kinase Edr . karenHKWCa]TahrN_mYyén



Feedforward activation BIeRBaH plitplénRbtikmμBImun bgá[mankarCMrujskm μPaBrbs;Gg;sIum énRbtikmμEdlekIteLIgenAeBleRkayenAkñúgpøÚvemtabUlIs . 4- karkeLIgCaem (FERMENTATIONS)

enAkñúgkarBinitüCaTUeTAén glycolysis eyIg)anbBa¢ak;fa Pyruvate EdlekIteLIg RtUvEt)an eRbIGs;eTAtamviFImYycMnYn dUecñHpøÚvEdlnwgRtUvbnþedIm,IbegáIt ATP . dUcKñaenHEdr NADH EdlekIt eLIgedaysar glycolysis enAkñúgdMNak;karTI 6 ¬emIlrUbTI 21-9¦ RtUvEt)anTTYlrgGuksIutkmμ sarCafμ IenAkñúgeBleRkay b¤k¾ glycolysis nwgkat;bnßyrhUtdl;Qb;enAeBlEdl NAD+ RtUv)an eRbIGs; . RbsinebIekasikabMeBjmuxgarsßitenAeRkamlkçx½NÐEdlmanxül; (aerobic) NADH nwgRtUv TTYlrgGutsIutkmμsarCaf μI nig Pyruvate nwgRtUvTTYlrgnUvGuksIutkmμeBjeljedaykardkdegðImEdl manxül; (aerobic respiration) . b:uEnþ enAeRkamlkçx½NÐK μanxül; (anaerobic) RbePTepSg²én RbtikmμkeLIgCaem (fermentation) bMeBjnUveKabMNgTaMgenH . karkeLIgCaem (fermentation) KWCa RbtikmμkatabUlIs (catabolic reactions) EdlekIteLIgedayBMumanGuksIutkmμsrub (no net

oxidation) . Pyruvate b¤ smasFatusrIragÁEdlplitBI Pyruvate TTYlrger:dukmμ enAeBl NADH

TTYlrgGuksIutkm μ . eyIgnwgBinitüpøÚvénkarkeLIgCaemBIrRbePTedayBisþar KW lactate fermentation

nig alcohol fermentation . k- karkeLIgCaemCaLak;tat (Lactate fermentation)

Lactate fermentation Cab;Tak;Tgya:gCitsñiT§eTAGñkNamñak;Edl)aneFIVkarhat;R)aNya:g xøaMg . RbsinebIGñkeFIVlMhat;R)aNxøaMgrhUtdl;sYt nig RbB½n§Qamrt;rbs;minGacbeBa©jGuksIuEsn RKb;RKan;eTA[sac;dMuEdlkMBugeFIVkar pøÚvRbmUlfamBlEdlmanxül; (aerobic energy-harvesting

pathway) minGacpþl; ATP [)anRKb;RKan;cMeBaHsac;dMurbs;GñkeT . b:uEnþ sac;dMuenAEtTamTar famBl. sßitenAeRkamlkçx½NÐKñanxül;enH lactate fermentation )ancab;epþImeLIg . enAkñúgRbtikmμ enH Gg;sIum Lactate dehydrogenase eFIVer:dukkmμeTAelI Pyruvate [eTACa Lactate . NADH

KWCaPñak;garer :dukkm μcMeBaHdMeNIrkarenH ¬rUbTI 21-10¦ . enAeBl Pyruvate TTYlrger:dukkmμ NADH TTYlrgGuksIutkm μnig NAD+ RtUvTTYl)aneLIgvijedayGnuBaØati[ glycolysis bnþdMeNIr eTAeTot . Lactate EdlekIteLIgcMeBaHsac;dMuEdlkMBugeFIVkargarcUleTAkñúgQam . CacugbBa©b;RbsinebI lMhat;R)aNxøaMgenAEtbnþ kMhab ;rbs; Lactate ekIneLIgx<s;rhUtdl; fermentation enHGacElgbnþ eTAmuxeTotehIy . Glycolysis nig ehtudUecñH karplit ATP RtUv)anbBaÄb; . sac;dMuxVHfamBl



GacElgbMeBjmuxgareTotehIy . cMNucénkarGs;kMlaMgenH (exhaustion) RtUv)aneKehAfa kMritRTaM)anénPaBKμanxül; (anaerobic threshold) . vaBitR)akdehIy PaKeRcInénBYkeyIgmineFIVlMhat;R)aN[dl;cMNucenHeT . enAeBllMhat; RtUv)anbBa©b; sBa’gÁkaycab;dMeNIrkarénkareFIV[dUcedImvijnUvRKb;famBlEdlmanGnuPaBEdlRtUv )an)at;eRkamTMrg;Ca lactate . eføIm)ancab;yk lactate BIQam nig bMElgvaeTACa Pyruvate vij . \LÚv karpþl;GuksIuEsnRKb;RKan;man pyruvate GacTTYlrgGusIutkmμedayeBjeljenAkñúgRbtikmμ RbmUlfamBlEdlmanxül; EdlmanRbsiT§PaBeRcInCagq¶ay edIm,IbMeBjeLIgvijnUvsþúkbMrugrbs; ATP . edaytampøÚvNak¾eday Pyruvate GacRtUv)anbMElgeTACa glucose nig RtUv)aneRbIedIm,IeFIV[ l¥eLIgvijnUvkarpþl;rbs; glycogen éneføIm nig sac;dMu . karbþÚr (exchange) énemtabUlItenH rvagsac;dMunigeføImRtUv)aneKehAfa Cori cycle . manRbePT)ak ;etrIepSg²EdlGaceFIV lactate fementation enAkñúglkçx½NÐK μanxül; . karenH mansar³sMxan;xøaMNas;enAkñúgeragc½RkmðÚbGaharBIeRBaHsrIragÁ (organisms) TaMgenHRtUv)aneKeRbIedIm,I ya:GY (Yogurt) nig cheeses . køinmut (tangy flavor) rbs;ya:GYRtUv)anpþl;eday lactate Edlplit eday)ak;etrITaMgenH . b:uEnþKYr[GP½BV srIragÁ (organisms) dUcKñaenHk¾bgá[TwkedaHp¥Úm . x- karkeLIgCaemeTACaGal;kul (Alcohol fermentation)

karkeLIgCaemeTACaGal;kul RtUv)aneKvaytMélya:gx<s; ¬RbsinebIeKminyl;¦ cab;taMgBI edImdMbUgénGriyF’m (the dawn of civilization) . dMeNIrkarkeLIgCaemxøÜnÉgRtUv)anrkeXIjeday Louis Pasteur enAkñúgry³eBlénkarsikSaKImIviTüaénkareFIVRsa nig {CMgWedaysarRsa} . sßitenA eRkamlkçx½NÐKμanxül; pSit (yeast) GaceFIV[eLIgCaemnUvsárEdl)anmkBIEpøeQInigRKab;FBaØCati . sárRtUv)anbMEbkeTACa Pyruvate edaysar glycolysis . karenHRtUv)anbnþedayRbtikmμBIr énkar keLIgCaemeTACaGal;kul . RbtikmμTI 1 Pyruvate decarboxylase dkyk CO2 ecjBI Pyruvate

edaybegáItCa acetaldehyde ¬rUbTI 21-11¦ . RbtikmμTI 2 Alcohol dehydrogenase sMrYl Rbtikmμer:dukkmμrbs; Acetaldehyde [eTACa ethanol b:uEnþ ritEtsMxan;eTAeTot vaeFIV[manGuksIutkm μ NADH eLIgvijenAkñúgdMeNIrkarenH . karekIeLIgnUv NAD+ GnuBaØati[ glycolysis bnþeTAeTot dUcenA kñúgkrNIkarkeLIgCaemeTACaLak;tatEdr . plitplBIrénkarkeLIgCaemeTACaGal;kul KW eGtaNul nig CO2 . eyIgTajykpl RbeyaCn_BIkareLIgCaemenHenAkñúgkarplitRsa nig Rsaebor_epSgeTotenAkñúgdMeNIrkarénkareFIVnMbu½g.



rUbTI 21-10 RbtikmμcugbBa©b;énkareLIgCaemeTACaLak;tat

rUbTI 21-11 RbtikmμcugeRkayBIrénkarkeLIgCaemeTACaGal;kul

5- pøÚvb:g;tUhVÚsVat (THE PENTOSE PHOSPHATE PATHWAY)

pøÚvb:g;tUhVÚsVat KWCapøÚvepSgeTotsMrab;GuksIutkmμ glucose . vapþl;[ekasikarYmCamYy famBleRkamTMrg;énkarkat;bnßyfamBlsMrab;CIvsMeyaK . CaBiess NADPH RtUv)anbegáItenAkñúg dMNak;karGuksIutkmμénpøÚvenH . NADPH KWCaPñak;garer:dukmμEdlRtUvkarsMrab;pøÚvCIvsMeyaKCaeRcIn . karBiB’NnaBisþarénpøÚvb:g;tUhVÚsVat minRtUv)anerobrab;enAkñúgGtßbTenHeT . b:uEnþkarBinitüCa TUeTAénRbtikm μsMxan;²nwgGnuBaØati[eyIgyl;GMBIsarsMxan;rbs;pøÚvenH ¬rUbTI 21-12¦ . eyIgGac BicarNapøÚvenHkñúgbIdMNak; . dMNak;TI 1 KWCadMNak;GuksIutkmμ EdlGacRtUv)ansegçbdUcxageRkam ³ Glucose-6-Phosphate + 2 NADP+ + H2O → Ribulose-5-Phosphate + 2 NADPH + 2 CO2

RbtikmμTaMgenHpþl;nUv NADPH EdlRtUvkarsMrab;CIvsMeyaK .

CH3—CO—COO−

Pyruvate

CH2—CHO + CO2 Acetaldehyde

CH3—CH2OH Ethanol

Alcohol dehydrogenase

NADH

NAD+

CH2—CO—CO—O− Pyruvate

NADH

NAD+

CH2—CHOH—CO—O−

Lactate

Lactate dehydrogenase



dMNak;karTI 2 Cab;Tak;TgeTAnwgRbtikm μGIusUEmrkm μEdlbMElg Ribulose-5-Phosphate [ eTACa Ribose-5-Phosphate b¤ Xylose-5-Phosphate . eQμaHrbs;pøÚvenHqøúHbBa©aMgBIplitplénBYk sárEdlman5GatUmkae)anEdlmanP¢ab;eTAedaybgÁúM Phosphate (Pentose phosphate) . dMNak;karTI 3 KWCaes‘rIRbtikmμsμ úRKs μajEdlCab;Tak;Tgnwgkardac;nigkarkekItsm<½n§ C—C . lT§plénRbtikmμKWCakarkekItnUvBIrm:UelKul Fructose-6-Phosphate nig mYym:UelKul

Glyceraldehyde-3-Phosphate BIbIm:UelKul Pentose phosphate . rUbTI 21-12

segçbdMNak;karsMxan;²énpøÚvb:g;tUhVÚsVat

bEnßmBIelIkarpþl;karkat;bnßyfamBl pøÚvb:g;tUhVÚsVatk¾pþl;nUvsárEdlmanbgÁúMhVÚsVatEdlRtUv karsMrab;CIvsMeyaK . edaysarehtuplenH Ribose-5-Phosphate RtUv)aneRbIsMrab;karsMeyaK nuyekøGUTId dUcCa ATP . sárman 4GatUmkae)anEdlmanbgÁúMhVÚsVat (Erythrose-4-Phosphate) Edl ekIteLIgenAkñúgdMNak;karTI 3 énpøÚvb:g;tUhVÚsVatKWCa precursor rbs;GasIutGamIneN Phenylalanine

tyrosine nig tryptophan . pøÚvb:g;tUhVÚsVatvaskmμxøaMgbMput enAkñúgCalikaEdlCab;Tak;TgeTA nwgCIvsMeyaKrbs; Cholesterol nig GasIutRka . dMeNIrkarTaMgBIrenHRtUvkarPaBsMbUrén NADPH . dUecñH eføIm ¬EdlCakEnøgsMeyaK cholesterol nig CakEnøgsMxan;bMputénCIvsMeyaKrbs;GasIutRka nig Calikaxøaj; (adipose) EdlenATIenaHkarsMeyaKskm μénGasIutRkak¾ekIteLIgEdr ¦ mankMritx<s;xøaMgénGg;sIumrbs; pøÚvb:g;tUhVÚsVat .

3 Glucose-6-Phosphate

3 Ribulose-5-Phosphate

Ribose-5-Phosphate + 2 Xylulose-5-Phosphate

2 Fructose-6-Phosphate + Glyceraldehyde-3-Phosphate

6 NADPH + CO2

6 NADP+ + H2O dMNak;karTI 1

dMNak;karTI 2

dMNak;karTI 3



pøÚvb:g;tUhVÚsVatk¾RtUv)anEktMrUvpgEdredIm,IR)akdfaekasika)anpþl;RKb;RKan;nUv Ribose-5-

phosphate nig NADPH . Glucose-6-Phosphate dehydrogenase KWCaGg;sIumEdlsMrYl GuksIutkmμrbs; Glucose-6-Phosphate enAkñúgRbtikmμTI 1 énpøÚv . Gg;sIumenHRtUv)aneP¶aceday eday Glucose-6-Phosphate nig RtUv)anraraMgeday NADPH . dUecñH pøÚvenHvaskm μxøaMgbMput enAeBl substrate maneRcIn nig vamanesckþIRtUvkarcMeBaH NADPH . 6- karsMeyaKKøúykUs (GLUCONEOGENESIS)

sßitenAeRkamlkçx½NÐFm μta eyIgman glucose edIm,IbMeBjesckþIRtUvkarrbs;eyIg . b:uEnþ sßitenAeRkamlkçx½NÐmYycMnYn sBa’gÁkayRtUvEteFIV glucose . karenHvacaM)ac; bnÞab;BIkarhat;R)aN xøaMgkøa edIm,IbMeBjeLIgvijnUvkarpþl;én glycogen rbs;eføIm nig sac;dMu . vak¾ekIteLIgEdrkñgry³eBl énPaBxVHxatcMNIGahar dUecñHsBa’gÁkayGacrkSakMrit glucose kñúgQam)anRKb;RKan;edIm,Ipþl;[ ekasikaxYrk,al nig ekasikaQamRkhm . sßitenAeRkamlkçx½NÐFmμtaCalikaTaMgBIrenHeRbIEt glucose sMrab;famBl . Glucose RtUv)an)anpliteTAtamdMeNIrkarén Gluconeogenesis EdlCakarplit glucose

ecjBIsarFatuEdlminCaBYkkabUGIuRdat ¬rUbTI 21-13¦ . Gluconeogensis pøÚvsMeyaK (anabolic

pathway) ekIteLIgCabzmenAkñúgeføIm . Lactate RKb;GasIutGamIneN ¬elIkElgEt leucine nig

lysine ¦ nig Glycerol Edl)anmkBIxøaj; GacRtUv)aneRbITaMgGs;edIm,Iplit glucose . b:uEnþ GasIut GamIneN nig glycerol CaTUeTARtUv)aneRbIEtkñúgkrNIxVHxatcMNIGahar (starvation) . emIlBIdMbUgPøam Gluconeogenesis hak;dUcCakarRtLb;ya:gsamBaØrbs; glycolysis

¬cUreRbobeFobrUbTI 21-13 nig 21-8¦ BIeRBaHFatukNþalénpøÚvTaMgBIrenHdUcKña . b:uEnþkarenH mindUcKñaeT BIeTeRBaHdMNak;karTI 1/ 3 nig 10 rbs; glycolysis minGacRtLb;eTAvij)aneT . ehtudUecñHRbtikmμRtLb;eTAvijRtUvEt)anGnuvtþedayGg;sIumepSgeTot . enAkñúgdMNak;karTI 1 én glycolysis , Hexokinase sMrYlénkarP¢ab;bgÁúM Phosphate eTAelI glucose . enAkñúg Gluconeogenesis pþac;bgÁúMhVÚsVatBI Glucose-6-Phosphate EdlRtUv)ansMrYledayGg;sIumeQμaH Glucose-6-Phosphatase EdlRtUv)aneKeXIjmanenAkñúgeføIm b:uEnþminmanenAkñúgsac;dMueT . dUcKña enHEdr RbtikmμTI 3 karP¢ab;énbgÁúMhVÚsVatén Fructose-6-Phosphate sMrYleday Phosphafructo-

kinase EdlminRtLb;eTAvij . dMNak;karenHqøgcUleTAkñúg (bypass) gluconeogenesis edaykareRbI Gg;sIumeQμaH Fructose-1,6-bisphosphatase . CacugbBa©b; karP¢ab;bgÁúMhVÚsVatén ADPRtUv)an sMrYleday Pyruvate kinase dMNak;karTI 10 én glycolysis minGacRtLb;vijeT . karbMElgén Pyruvate eTACa Phosphoenolpyruvate CaFmμCab;Tak;TgeTAnwgGg;sIumBIrnig RbtikmμminFm μtamYy



cMnYn . Gg;sIumTI 1 KW Pyruvate carboxylase bEnßm CO2 eTAelI Pyruvate . plitplKWCa smasFatuEdlman4GatUmkae)anKW Oxaloacetate . bnÞab;mk Phosphoenolpyruvate

carboxykinase dk CO2 nig bEnßmbgÁúMhVÚsVat . Gñkpþl;bgÁúMhVÚsVatenAkñúgRbtikmμminFmμtaenHKW Guanosine triphosphate (GTP) . vaenHCanuyekøGUTIddUc ATP Edr elIkElgEt)asrbs;vaKWCa Guanine .

rUbTI 21-13 kareRbobeFobRbtikmμén glycolysis nig Gluconeogenesis

Glucose

Glucose-6-Phosphate

Fructose-6-Phosphate

Fructose-1,3-Bisphosphate

Glyceraldehyde- 3-Phosphate

Dihydroxyacetone phosphate

Glycerol

1,3-Bisphosphoglycerate

3-Phosphoglycerate

2-Phosphoglycerate

Phosphoenolpyruvate

Pyruvate

Pyruvate carboxylase

ADP

ATP

Oxaloacetate

CO2

Amino acids +

Fatty acids

CO2

GDP

GTP

ADP

ATP

Glucose-6-Phosphatase 1

2

3

4

5

6

7

8

9

10

Phosphoenol Pyruvate

carboxylase

dMNak;karminRtLb; EdlRtUvvag



RbtikmμmYyKUrcueRkayenHvasμ úRKs μajedaysarkarBitEdlfa Pyruvate carboxylase eKeXIj manenAkñúgmItUkugRDI rIÉ Phosphoenolpyruvate carboxykinase eKeXIjmanenAkñúgsIutU)aøs; . dUcEdl)aneXIjenAkñúgCMBUkTI 22 nig 23 mItUkugRDI KWCa seIr³ (organelles) EdlGuksIutkm μcugbBa©b; rbs;m:UelKulcMNIGaharekIteLIgenATIenaH nig cMnYnd¾eRcInén ATP RtUv)anplit . RbB½n§dwknaMEdl s μ úRKsμajdwknaM Oxaloacetate Edlplit)ancUleTAkñúgmItUkugRDI tamry³PñasBIrrbs;mItUkugRDI nig cUleTAkñúgsIutU)aøs; . enATIenaH Phosphoenolpyruvate carboxykinase sMrYlkarbMElg oxaloacetate [eTACa Phosphoenolpyruvate . RbsinebI glycolysis nig gluconeogenesis minRtUv)anEktMrUvenAkñúgrebobdUcKña pøÚvTaMgBIrnwg ekItdMNalKña rYmeTAeday\T§iBlEdlb:HBal;ya:gxøaMgEdlminmanGIVnwgRtUveFVI . TIkEnøgbIsMrab;karEk tMrUvenHKWCakarEktMrUvénpÚøvvagbI . dMNak;karTI 3 én glycolysis RtUv)ansMrYledayGg;sIum Phosphofructokinase . Gg;sIumenHRtUv)aneP¶acedaykMhab;x<s;én; AMP ADP nig bgÁúMhVÚsVat Edlpþl;CasBaØafaekasikaRtUvkarfamBl . enAeBlGg;sIummanskmμPaB (active) glycolysis k¾bnþ dMeNIrkar . müa:gvijeTot enAeBl ATP sMbUr Phosphofructokinase RtUv)anraraMg nig fructose-

1,6-bisphosphatase RtUv)aneP¶ac[manskmμPaB . lT§plsrub KWfa enAkñúgeBlEdlfamBlelIs ¬kMhab; ATP x<s;¦ gluconeogenesis nwgekIteLIg . dUcEdleyIg)aneXIjrYcmkehIy karbMElgrbs; lactate eTACa glucose mansar³sMxan;cMeBaH stV . enAeBlsac;dMueFVIkargar BYkvaplitnUv lactate EdlRtUv)anbMElgRtLb;eTACa glucose vijenA kñúgeføIm . Glucose Edlplit)anRtUv)andwknaMcUleTAkñúgQam nigecjBIkñúgQamRtLb;eTAkñúgsac;dMu vij . enAkñúgsac;dMu glucose enHRtUv)anbMEbkedIm,IbegáItCa ATP b¤ vaRtUv)aneRbIedIm,IbMeBjeLIgvij sþúkbMrug glycogen rbs;sac;dMu . dMeNIrkarvilCMurvageføIm nig sac;dMuenH (skeletal muscle) RtUv)aneK ehAfa vdþkUrI (Cori Cycle) nig RtUv)anbgðajenAkñúgrUbTI 21-14 . tamry³vdþenH gluconeogenesis

plit glucose RKb;RKan;edIm,IeFIV[)anl¥eLIgvijnUv glycogen rbs;sac;dMuEdl)an)at;bg; . 7- karsMeyaK nig karbMEbkénKøIkUEsn (GLYCOGEN SYNTHESIS AND DEGRADATION)

Glucose KWCaRbPTEtmYyénfamBl én ekasikaQamRkhm nig ekasikaxYrk,alrbs;BYk fnikstV . eTaHbICa ekasikaQamRkhm k¾dUcCaekasikaxYrk,alminGacsþúk glucose )an b:uEnþkarpþl; CaRbcaMCa glucose enAkñúgQamRtUvEtman . karenHRtUv)anpþl;eday glucose éncMNIGahar nig edaykarplitnUv glucose TaMg edaysar gluconeogenesis b¤ Glycogenolysis ¬karbMEbknUv glycogen¦ . Glycogen KWCab:UlIEmrén glucose EdlmanExSEvgnigmanExñg (a long-branched



chain polymer of glucose) . edaysþúkTukenAkñúgeføIm nig sac;dMuéneRKagq¥wg vaKWCaTMrg;sþúkmUldæan rbs; glucose .

rUbTI 21-14 vdþkUrI (Cori cycle)

cMnYnsrubén Glucose enAkñúgQamrbs;mnusSeBjv½y 0 KILÚRkam ¬RbEhl 150 lb¦ manRbEhl 20 g b:uEnþxYrk,alEtÉgRtUvkarBI 5-6 g én glucose kñúgmYyema:g . glycogen EdlbMEbk enAkñúgeføImeFIV[ glucose manclnaenAeBlsBaØarbs; hormone rkeXIjBItMrUvkarén glucose

ekIneLIg . sac;dMuéneRKagq¥wgk¾mansþúkTukya:gsMxan;én glycogen Edr Edlpþl;nUvkarkRnþak;sac;dMu ya:gelOn . b:uEnþ glycogen enHminGacbNþal[man glucose enAkñúgQameT BIeRBaHekasikasac;dMu minmanGg;sIum glucose-6-phosphate . BIeRBaH glucose minGacRtUv)anbegáItBI Glucose-6-

Phosphate dUecñHvaminGacRtUv)anbeBa©jcUleTAkñúgcrnþQam . k- rcnasm<½n§rbs;KøIkUEsn (The structure of glycogen)

Glycogen KWCab:UlIEmén glucose EdlmanExñg EdlExSsMxan;RtUv)ancgP¢ab;edaysm<½n§ glycoside α (1-4) . b:UlIEmrk¾mansm<½n§ glycoside (1-6) CaeRcInEdr Edlpþl;cMNucExñgCaeRcInenA tambeNþayExS . rcnasm<½n§enHRtUv)anbgðajenAkñúgrUbTI 21-15 . RKab;KøIkUEsn (Glycogen

granules) EdlmanGgát;p©itBI 10-40 nm RtUv)aneKeXIjmanenAkñúgsIutU)øas;rbs;ekasikaeføIm nig sac;dMu . RKab;KøIkUEsnTaMgenHsßitenArYmCabNúþMCamYynwgGg;sIumEdlTTYlxusRtUvsMrab;karsMeyaK nig karbMEbkKøIkUEsn . rcnasm<½n§énRKab;enHk¾RtUv)anbgðajEdrenAkñúgrUbTI 21-15 .

Glycogen

Exercise Rest 1

Glucose-6-Phosphate

2

Pyruvate

3

Lactate

Glucose

Glycogen

7

G-6-P

6

Pyruvate

5

Lactate

Blood

8

Blood

4



rUbTI 21-15 rcnasm<½n§rbs;KøIkUEsn nig RKab;KøIkUEsn

-

O

CH2OH

OH

OH

CH2OH

OH

OH

O

CH2OH

OH

OH

CH2

OH

OH

O O

CH2OH

OH

OH CH2OH

OH

OH

O

CH2OH

OH

OH O

O

α(1-4) glycosidic bond

α(1-6) glycosidic bond



x- karbMEbk Glycogen (Glycogenolysis)

manG½rm:UnBIrRtYtBinitü glycogenolysois ¬karbMEbk glycogen¦ . BYkvaKWCa Glucagon Edl CaG½rm:Un peptide RtUv)ansMeyaKenAkñúglMEBg nig Epinephrine EdlRtUv)anplitenAkñúgRkeBj Adrenal . Glucagon RtUv)anbeBa©jBIlMEBgedIm,IeqøIytbeTAnwgkMrit glucose kñúgQamTab nig Epinephrine beBa©jBIRkeBj Adrenal kñúgkareqøIytbeTAnwgkarKMramkMEhg nig PaBtantwg . sßanPaBTaMgBIr tMrUv[mankarekIneLIgnUvkMrit glucose kñúgQam nig G½rm:UnTaMgBIrbMeBjmuxgareday kareFIV[b:HBal;dl;skmμPaBrbs;Gg;sIumBIr KW Glycogen phosphorylase nig Glycogen synthase . Glycogen phisphorylase Cab;Tak;TgeTAnwgkarbMEbk glycogen nig RtUv)aneFIV[skm μ Glycogen synthase Cab;Tak;TgeTAnwgkarsMeyaK glycogen nig RtUveFIV[)at;skmμPaB . dMNak;karénkarbMEbk glycogen RtUv)ansegçbdUcxageRkam ³ dMNak;karTI 1 Gg;sIum glycogen phosphorylase sMrYl phosphorolysis én glucose

enAcugmYyén glycogen polymer ¬rUbTI 21-16¦ . Rbtikm μCab;Tak;TgeTAnwgCab;Tak;TgeTAnwgkar cakecjrbs; glucose BI glycogen edaymanbgÁúM Phosphate . CalT§plén Phosphorolysis Glucose-1-Phosphate RtUv)anplitedayminmaneRbI ATP CaGñkpþl;bgÁúM Phosphate . dMNak;karTI 2 Glycogen manExñgCaeRcInEdlcgP¢ab;eTAnwgExSRtg α(1-4) tamry³sm<½n§ α(1-6) . ExSExñgTaMgGs;enHRtUvEt)andkecjedIm,IGnuBaØati[mankarbMEbk glycogen TaMgRsug . GMeBITUlMTUlayrbs; glycogen phosphorylase begáIt)anCa Polysaccharides EdltUcCag nig Glucose mYyEdlcgtamry³sm<½n§ α(1-6) eTAnwgExSem . Gg;sIum α(1-6) glycosidase k¾GacRtUv )aneKehApgEdrfaCa debranching enzyme GIuRdUlIssm<½n§ α(1-6) glycosidic enAnwgcMNucExñg nig eFIV[mankarbeBa©jnUvmYym:UelKul glucose ¬rUbTI 21-17¦ . m:UelKulrbs; glucose enHGacRtUv )anP¢ab;edaybgÁúM P{hosphate nig RtUveRbIR)as;enAkñúg glycolysis b¤ k¾GacbeBa©jeTAkñúgcrnþQam sMrab;eRbIR)as;enATIkEnøgepSg²eTot . kareFIVGIuRdUlIsénsm<½n§ExñgbeBa©jnUvkarlatsn§wgnUvsm<½n§ α-

glucose epSgeTotsMrab;GMeBIrbs; glycogen phosphorylase . dMNak;karTI 3 Glucose-1-Phosphate RtUv)anbMElgeTACa Glucose-6-Phosphate eday sar Phosphoglucomutase ¬rUbTI 21-18¦ . Glucose edayedImdMbUgRtUv)ansþúkTukenAkñúg glycogen cUleTAkñúg Glycolysis tabry³GMeBIrbs; Phosphoglucomutase . müa:gvijeTot enAkñúg eføIm nig tMrgenamvaGacRtUv)andkbgÁúM Phosphate ecjsMrab;dwknaMcUleTAkñúgcrnþQam . K- karsMeyaK Glycogen (Glycogenesis)



G½rm:Un Insulin EdlplitedaylMEBgkñúgkareqøIytbeTAnwgkMrit glucose x<s;enAkñúgQam eP¢ackarsMeyaKén glycogen (glycogenesis) . Insulin RbEhlCaG½rm:UnmYykñúgcMeNamG½rm:UnEdl man\T§iBlCageKbMputkñúgsBa’gÁkay BIeRBaHvab:HBal;pÞal;eTAnwgemtabUlIs nig kareRbIR)as;TaMgGs; én glucose elIkElgEtekasikamYycMnYnénsBa’gÁkay . enAeBlkMritén glucose ekIneLIg bnÞab;BIbriePaKGahar ekasika β énlMEBgbeBa©jnUv Insulin . CabnÞan;vaCMrujkareRbIR)as;én glucose edayRKb;ekasikaénsBa’gÁkayelIkElgEtekasika xYrk,al nig ekasikaQam . enAkñúgekasikaTaMgenH kareRbIR)as; glucose KWminCab;Tak;TgeTAnwg Insulin . kareRbIR)as;én glucose ekIneLIgRtUv)ankt;sMKal;CaBiessenAkñúgeføIm ebHdUg sac;dMuén eRKagq¥wg nig Calikaxøaj; (adipose tissue) . enAkñúgeføIm Insulin CMruj[mankarsMeyaK nig sþúkTukén glycogen . vaeFIVdUecñH edaykarman \T§iBleTAelIGg;sIumEdlCab;Tak;TgenAkñúgemtabUlIsrbs; glycogen kúñgrebobEdlRbQmeTAnwg GMeBIrbs; glucagon . Insulin raraMg glycogen phosphorylase ehtuenHvararaMg glycogenolysis . vak¾CMrujpgEdrnUv glycogen synthase nig glucokinase EdlGg;sIumTaMgBIrenHCab;Tak;TgeTAnwg karsMeyaK glycogen . eTaHbICa Glycogenesis nig Glycogenolysis manRbtikmμmYycMnYnrYmKñak¾eday pøÚvTaMgBIrenH minRtLb;eTAvijeTAmk)aneLIy . Glycogenesis Tak;TgeTAnwg RbtikmμminTUeTAmYycMnYn EdleyIg nigBinitüedayBisþarkñúgeBl\LÚvenH . RbtikmμTImYyénkarsMeyaK glycogen enAkñúgeføImcab;yk glucose enAkñúgekasikaedaykar P¢ab;bgÁúM Phosphoryle eTAelI glucose . kñúgRbtikmμenHRtUv)ansMrYledayGg;sIumeQ μaH Glucokinase ehIy ATP RtUv)aneRbICaGñkpþl;bgÁúM Phosphoryle nig Glucose-6-Phosphate

RtUv)anbegáIteLIg . Glucokinase Glucose + ATP → Glucose-6-Phosphate + ADP + H+

RbtikmμTIBIrénkarsMeyaK glycogen KWCakarRtLb;mkvijénRbtikmμ glycogenolysis . Glucose-6-Phosphate EdlRtUv)anbegáItenAkñúgdMNak;TImYyRtUv)anTTYlrgGIusUEmrkmμeTACa Glucose-1-Phosphate . Gg;sIumEdlsMrYlRbtikmμKW Phosphoglucomutase . Phosphoglucomutase

Glucose-6-Phosphate ⇄ Glucose-1-Phosphate

\LÚv Glucose-1-Phosphate RtUvEtkøayeTACaskmμmuneBlvaRtUv)anbEnßmeTAelIExS glycogen . smasPaBEdlmanfamBlx<s;EdleFIV[sMerckic©karenHKW Uridine Triphosphate



(UTP). kñúgRbtikmμenHRtUv)ansMrYledayGg;sIum Pyrophosphorylase ehIybgÁúM Phosphoryle

EdlenAelI C-1 én glucose RtUv)anP¢ab;eTAnwg bgÁúM α phosphoryle rbs; UTP edIm,IbegáItCa UDP-

Glucose ³ Pyorophosphorylase

Glucose-1-Phosphate + UTP ⇄ UDP-Glucose + Pyrophosphate

karenaHnaMmkCamYynUvkarbeBa©jbgÁúM Pyrophosphate (Pi) . rcnasm<½n§én UDP-Glucose

RtUv)anbgðajenAkñúgrUbTI 21-19 . \LÚv UDP-Glucose GacRtUv)aneRbIedIm,IBnøÚtExS glycogen . Gg;sIum Glycogen sythase

pþac;sm<½n§ Phosphoester rbs; UDP-Glucose nig begáItsm<½n§ glucosidic α(1-4) rvag glucose nig ExS glycogen EdlRtUvlUt . UDP RtUv)anbeBa©jenAkñúgdMeNIrkarenH . Glycogen synthase UDP-glucose + Glycogen primer (n residues) → Glycogen (n+1 residues) + UDP

CacugbBa©b; eyIgRtUvEtbgðajsm<½n§ glycosidic α(1-6) edIm,IbegáItCaExñg . ExñgTaMgGs;man sar³sMxan;xøaMgkñúgkareRbIR)as; glycogen [)anRtwmRtUv . dUcEdlrUbTI 21-20 bgðaj Branching

enzyme dkEpñkEdlCa Glycogen ExSRtg; α(1-4) nig P¢ab;vaeLIgvijenAkñúgsm<½n§ glycosidic α(1-6)

enAkñúgTINamYyénExS glycogen . X- PaBcuHsMrugKñaénkarsMeyaK nig karbMEbk glycogen

(Compatibility of Glycogenesis and Glycogenolysis)

GaRs½edaykrNICamYy glycolysis nig glycogenesis vanwgminbegáItnUvlT§plEdlmansar³ sMxan;sMekasikakñúgkarGnuvtþn_karsMeyaK nig karbMEbk glycogen dMNalKñaeT . lT§plEdlsMerc )anedaysarGMeBIrbs;pøÚvmYy (glycogenesis) nwgminGaceFIV)anedaysarpøÚvepSgeToteT (glycogenolysis) . bBaðaenHRtUv)aneCosvag)anedaysarkarRtYtBinitürbs;G½rm:UnmYyes‘rI EdleFIV[ skmμGg;sIumrbs;pøÚvNamYykúñgxN³enHEdrmankareFIV[)at;skm μPaBrbslGg;sIuménpøÚvepSgeTot. enAeBlkMrit glucose kñúgQamx<s;xøaMgeBk lkçx½NÐEdleKsÁal;faCa Hyperglycemia Insulin eP¢ackareRbIR)as; glucose tamry³kardwknaMqøgkat;Pñas . vaeP¢acCabnþeTotnUvkarcab; glucose edayskm μPaBekIneLIgén Glucokinase . CacugbBa©b;vaeFIV glycogen synthase

manskmμPaB EdlCaGg;sIumcugeRkaybMputkñúgkarsMeyaKénExS glycogen .edIm,ICMrujkarsþúkTuk kan;EteRcInEfmeTot Insulin raraMgGg;sIumTI 1 enAkñúgkarbMEbk glycogen KW glycogen

phosphorylase . \T§iBlsrub RtUv)aneKeXIjmanenAkñúgrUbTI 21-21 KWfa glucose RtUv)andkecj



BIcrnþQam nig bMElgeTACa Glycogen enAkñúgeføIm .enAeBlEdlXøaMgsþúkTuk glycogen eBj glucose EdlelIsRtUv)anbMElgeTACaxøaj;EdlsúþkTukeRkamTMrg;CaCalikaxøaj; . Glucagon RtUv)anplitkñúgkareqøIytbeTAnwgkMrit glucose kñúgQamTab lkçx½NÐEdleKsÁal; faCa Hypoglycemia nig man\T§iBlpÞúyBI\T§iBlrbs; Insulin . vaeP¢ab glycogen phosphorylase

EdlsMrYldMNak;kardMbUgénkarbMEbk glycogen . karenHvabegáInnUv Glycogenolysis nig beBa©j glucose eTAkñúgcrnþQam . \T§iBlRtUv)anbegáInEfmeTot BIeRBaH Glucagon raraMg Glycogen

synthase . \T§iBlRbQmKñarbs; Insulin nig Glucagon RtUv)ansegçbenAkñúgrUbTI 21-21 . RbB½n§RtYtBinitürbs;G½rm:Und¾l¥enHFanafaRbtikmμEdlCab;Tak;TgenAkñúgkarbMEbk nig kar sMeyaK glycogen minbMeBjKñaeTAvijeTAmkeT . enAkñúgrebobenH BYkvapþl; glucose enAeBlkMrit glucose enAkñúgQamTabeBk nig BYkvabgá[mankarsþúkTuknUv glucose enAkñúgeBlEdlelIs .

Fin Reference : General, Organic, and Biochemistry 3rd edition, 2001 by Denniston, Topping, Caret page 599-633

CIvKImIvITüa kardkdegðImEdlmanxül; nig karplitfamBl

erobcMnigbkERbeday»sfkarIRcwgsuPaBTUrs½BÞ012864353 b¤ 011717361 e-mai: [email protected] Page 1 of 28

kardkdegðImEdlmanxül; nig karplitfamBl (AEROBIC RESPIRATION AND ENERGY PRODUCTION)

vtßúbMNgemeron 1- [eQ μaHtMbn;TaMgLayrbs;mItUkugRDI nig muxgarrbs;tMbn;nimYy² . 2- BiB’NnaGMBIRbtikmμEdlbNþal[mankarbMElgBI Pyruvate eTACa Acetyl CoA edayBiB’NnaBI TIkEnøgénRbtikm μ nig smasPaBrbs;bNþúM Pyruvate dehydrogenase . 3- segçbRbtikm μénkardkdegðImEdlmanxül; . 4- edayBinitüemIsmIkarEdltMNag[RbtikmμmYy kñúgcMeNamRbtikm μKImIEdlekIteLIgenAkñúg Citric

acid cycle BiB’NnaRbePTénRbtikmμEdlekIteLIg nig sar³sMxan;énRbtikmμTaMgenaHenAkñúgpøÚv . 5- Bnül;GMBIynþkarsMrab;karRtYtBinitüén Citric acid cycle . 6- BiB’NnaGMBIdMeNIrkarén Oxidative Phosphorylation . 7- BiB’NnaGMBIkarbMElgGasIutGamIneNeTACam:UelKulEdlGaccUleTAkñúg Citric acid cycle . 8- Bnül;GMBIsarsMxan;rbs; Urea cycle nig BiB’NnaBIdMNak;karsMxan;² . 9- BiPakSaGMBImUlehtu nig \T§iBlrbs; Hyperammonia . 10- segçbBItYnaTIrbs; Citric acid cycle enAkñúg Catabolism nig Anabolism .



KMeragemeron 1- mItUkugRDI (Mitochondria)

k- rcnasm<½n§ nig muxgar x- RbPBrbs;mItUkugRDI 2- karbMElgBI Pyruvate eTACa Acetyl CoA

3- karBinitüTUeTAénkardkdegðImEdlmanxül; 4- Citric Acid Cycle

Rbtikmμ Citric Acid Cycle

5- karRtYtBinitü Citric Acid Cycle

6- Oxidative Phosphorylation

k- RbB½n§dwknaMeGLikRtug nig karRCalénGIuy:ugGIuRdUEsn x- ATP Synthase nig karplit ATP

K- segçbnUvlT§plTTYl)anénfamBl 7- karbMEbknUvGasIutGamIneN k- kardkecjnUvbgÁúM α-amino : Transamination

x- kardkecjnUvbgÁúM α-amino : Oxidative Deamination

K- vasnarbs;eRKagExSkae)anrbs;GasIutGamIneN 8- Urea Cycle

Rbtikmμén Urea Cycle

9- karBinitüTUeTAén Anabolism : The Citric Acid Cycle CaRbPBsarFatukNaþal énCIvsMeyaK



kardkdegðImEdlmanxül; nig karplitfamBl (AEROBIC RESPIRATION AND ENERGY PRODUCTION)

esckþIepþIm dUcEdleyIg)aneXIjpøÚv Anaerobic glycolysis cab;epþImkarbMEbk glucose nig begáItnUvcMnYn tictYcén ATP nig NADH . b:uEnþ vaKWCapøÚv Aerobic catabolism EdlbEnßm[cb;sBVRKb;nUvGuksIutkmμ rbs; glucose [eTACa CO2 nig H2O nig pþl;nUv ATP eRcInbMputEdlRtUvkaredaysBa’gÁkay . karBit enAkñúgdMeNIrkarenH eKehAfaCa kardkdegðImEdlmanxül; (Aerobic respiration) begáItnUv 36

ATP edayeRbIfamBlEdlRbmUl)anBIm:UelKulnimYy²EdlcUleTAkñúg Glycolysis . RbtikmμTaMgenH ekIteLIgenAkñúgpøÚvemtabUlIsEdlsßitenAkñúg mItUkugRDI EdlCaeragc½RkplitfamBlrbs;ekasika . mItUkugRDIKWCaRbePTénPaKl¥itrbs;ekasikaEdlmanPñasCMuvij . enATIenH enAkñúgmItUkugRDI GuksIutkmμcugbBa©b;rbs; carbohydrate lIBId nig RbtUetGIunekIt eLIg . enATIenH eGLikRtugEdlRbmUl)anBIRbtikmμGuksIutkmμk¾RtUv)aneRbIedIm,IeFIV ATP Edr . enAkñúgRbtikm μEdlmanRbsiT§PaBya:gxøaMgTaMgenH Cit 40 % énfamBlEdlmanskþanuBlrbs; GlucoseRtUv)ansþúkTukeRkamTMrg;Ca ATP . 1- mItUkugRDI (MITOCHONDRIA)

mItUkugRDICaPaKl¥itmanragmUldUc)al; EdlmanTMhMRbEhleTAnwgekasika)ak;etrI . BYkvaRtUv )ancgeTAeday PñasxageRkA (outer mitichondrial membrane) nig Pñasxagkñúg (inner

mitochondrial membrane) rUbTI 22-1 . tMbn;enAcenøaHrvagPñasTaMgBIrRtUv)aneKehAfaCa cenøaHGnþrPñas (intermembrane space) nig tMbn;EdlrMuB½T§eTAedayPñasxagkñúgRtUv)aneKsÁal;faCa Matrix space . Gg;sIumrbs; Citric acid cycle , β-oxidation sMrab;karbMEbkGasIutRka nig sMrab;kar bMEbkGasIutGamIneNRtUv)aneKeXIjTaMgGs;enAkñúg Matrix space rbs;mItUkugRDI .

rUbTI 22-1



k- rcnasm<½n§ nig muxgar PñasxageRkArbs;mItUkugRDI RCabedayesrIeTAnwgsarFatuEdlmanma:s;m:UltUcCag 10,000

g/mol . m:UleKultUcRtUv)anTTYlrgGuksIutkmμsMrab;karplit ATP dUcCa Pyruvate EdlRtUv)aneday glycolysis GaccUledayesrIeTAkñúgcenøaHGnþrPñastamry³rn§enAkñúgPñasxageRkA . Pñasxagkñúgmanpñt;CaeRcInEdlbegáInépÞya:gFM . PñasEdlbt;enHRtUv)aneKsÁal;faCa Cristae. Pñasxagkñúgrbs;mItUkugRDIesÞIrEtminRCabeTAnwgsarFatuPaKeRcIn . GaRs½yehtuenH vamanRbUetGIundwknaMCaeRcInedIm,InaMmU:elKul\n§³BiesscUleTAkñúg Matrix space . edaymanenAkñúg Pñasxagkñúgrbs;mItUkugRDIKWCaRbUetGIundwknaMeGLikRtugén RbB½n§dwknaMeGLikRtug nig ATP synthase

. ATP synthase KWCabNþúMFMmYymanRbUetGIunCaeRcInEdlsMrYlkarsMeyaK ATP . x- RbPBrbs;mItUkugRDI (Origin of Mitochondria)

minRtwmEtmanTMhMRbEhlnwg)ak;etrIeT mItUkugRDImanTidæPaBCaeRcInepSgeTotEdl)aneFIV[Gñk RsavRCavsgS½yfaCa)ak;etrIEdlrs;enAedayesrIenAeBlNamYy EdlRtUv)ancab;ykedayekasika BYk Eucaryote . BYkvamanB½t’manEhSn (DNA) pÞal;rbs;va . BYkvak¾eFIV ribosomes rbs;vaEdldUc Kñaya:gxøaMgeTAnwg)ak;etrI . Ribosomes TaMgenHGnuBaØati[mItUkugRDIsMeyaKRbUetGIunrbs;vamYycMnYn. CacugbBa©b; mItUkugRDI CaTUeTAvaGaceFIVkarbMEbkedayxøÜnÉg . BYkvaGaclUtlas;rIkFM nig bMEbkxøÜnedIm,IbegáItCamItUkugRDIfμ I . RKb;lkçN³TaMgenHpþl;Caeyablfa mItUkugRDIEdlplitPaKeRcIn én ATP cMeBaHekasikarbs;eyIgmanRbPBBI)ak;etrIRtUv)anekasikacab;ykRbEhl 1.5 x 109 qñaMknøg eTA . 2- karbMElgBI PYRUVATE eTACa ACETYL CoA

dUcEdl)aneXIjenAkñúgCMBUk 21 eRkamlkçx½NÐminmanxül; glucose RtUv)anTTYlrge mtabUlIseTACaBIrmU:elKul Pyruvate EdlbnÞab;mkRtUvbMElgeTACaplitplénkareLIgCaemesßr (stable fermentation product) . karbMEbk glucose EdlmanRBMEdnkMNt;enHbeBa©jnUvfamBlEdl manskþanuBlrbs; glucose ticxøaMgNas; . enAeRkamlkçx½NÐEdlmanxül; ekasikaGaceRbI oxygen

nig eFIVGuksIutkm μedayeBjeljrbs; glucose [eTACa CO2 enAkñúgpøÚvemtabUlIs . EdleKehAfaCa Citric Acid Cycle . Cajwkjab;pøÚvenHeKsMedAcMeBaH Krebs Cycle kúñgkarpþl;kitþiysdl;elak Hans Krebs

Edl)anbegáItnUvdMNak;karTaMgLayén Citric acid cycle enHBITinñn½yBiesaFn_rbs;Kat; nig Tinñn½y rbs;GñkRsavRCavepSgeTot . vak¾RtUv)aneKehApgEdrfaCa Tricarboxylic acide(TCA) cycle BIeRBaH emtabUlItkNþaledImdMbUgCaeRcInénpøÚvmanbgÁúM Carboxyle bI .



\LÚveyIgbMEbrkarykcitþTukdak;rbs;eyIgeTAelIkarpliténmU:elKulEdlmanEpñkénBIrGatUm kae)an (two carbon fragments) KW bgÁúM Acetyl BI Pyruvate cUleTAkñúgpøÚvRbmUlfamBlEdlman xül;. m:UelKulenHKW Acetyl CoA . rcnasm<½n§rbs; Acetyl CoA RtUv)aneKeXIjenAkñúgrUbTI 22-2 . Coenzyme A Epñkénm:UelKul manRsLayBI ATP nig Vitamin pantothenic acid (B5) . vabMerICaGñkcab;ykbgÁúM Acetyl bgðajCa B’NRkhmenAkñúgrUbTI 22-2 EdlP¢ab;eTAnwgbgÁúM Thiol énm:UelKuledaysarsm<½n§ Thioester . eyIg Gaccat;Tuk Acetyl CoA faCaTMrg;skmμénbgÁúM Acetyl . RbtikmμEdlbMElg Pyruvate eTACa AcetylCoA RtUv)anbgðajenAkñúgrUbTI 22-3 . Pyruvate

RtUv decarboxylated EdlbeBa©jnUv CO2 . vak¾RtUvTTYlrgGuksIutkm μnig Hydride anion (H-)

EdlRtUv)andkecjRtUvTTYlykeday NAD+ EdlehtuenHvaTTYlrger:dukm μ . CacugbBa©benAsl;Et bgÁúM Acetyl (CH3CO—) RtUv)anP¢ab;eTAnwg Coenzyme A edaysm<½n§ Thioester . RbtikmμEdl s μ úRKya:gxøaMgenHRtUv)anGnuvtþedayGg;sIumbI nig Coenzyme R)aM . edIm,IsnSMsMécdMeNIrkarenH Gg;sIum nig Coenzyme TaMgGs;sßitenAkñúgkBa©b;mYyEdleKehAfaCa Pyruvate dehydrogenase

complex ¬emIlrUbTI 22-3¦ . enAkñúgpøÚvenH Substrate Gacqøgkat;BIGg;sIummYyeTAGg;sIumbnÞab; edaymankarERbRbYlekIteLIg . KMnUsbMRBYjbgðajenAkñúgrUbTI 22-3b .

rUbTI 22-2 rcnasm<½n§rbs; AcetylCoA

O O O H CH3 O O ║ ║ ║ │ │ ║ ║

CH3—C~S—CH2—NH—C—CH2—CH2—NH—C—C—C—CH2—O—P—O—P—O—CH2 │ │ │ │ OH CH3 O

- O-

P

Ribose

Adenine

Acetyl group

Beta-mercapto

ethylamine group

Pantothenate unit

Phosphorylated ADP



RbtikmμEtmYyenHRtUvkar Coenzymes bYnEdlplitecjBIvItamInepSg² epSgBI Coenzyme

lipoamide . BYkvaTaMgenHrYmman Thiamine pyrophosphate EdlmanRsLayBI thiamine (Vitamin

B1) . FAD manRsLayBI Riboflavin (Vitamin B2) . NAD+ manRsLayBI Naicin nig Coenzyme

A EdlmanRsLayBI Pantothenic acid (Vitamin B5) . edayCak;Esþg karfycuHénvItamInTaMgenH nwgeFIV[fycuHya:geRcInnUv Acetyl CoA Edlekasikarbs;eyIgGacplit . Cabnþ karenHnwgkMNt;nUv cMnYn ATP EdlxøÜneyIgplit nig bNþal[manCMgWkgVHvitamIn . edayP½BVsMNag rbbGaharEdlman tulüPaBl¥pþl;nUvkarpÁt;pÁg;RKb;RKan;nUvvItamInTaMgenH nig vItamIneepSg²eTot . rUbTI 22-4 eyIgeXIjfa Acetyl CoA KWCaFatukNþal (central character) enAkñúg emtabUlIsekasika . vaRtUv)anplitedaykarbMEbk glucose , Fatty acids, nig GasIutGamIneNmYy cMnYn . muxgarsMxan;rbs; Acetyl CoA enAkñúgpøÚvRbmUlfamBl KWedIm,IdwknaMbgÁúM Acetyl cUleTAkñúg Citric acid cycle EdlvanwgRtUv)aneRbIedIm,IplitnUvcMnYnd¾eRcInén ATP . bEnßBIPar³kic©katabUlIsenH bgÁúM Acetyl rbs; Acetyl CoA k¾GacRtUveRbIsMrab; GaNabUlIs (anabolism) b¤ Rbtikm μCIvsMeyaKedIm,I plit cholesterol nig Fatty acids . eKKitfa Actyl CoA CaFatukNþalEdlRKb;RbPBfamBl ¬xøaj; RbUetGIun nig carbohydrate¦ GacbMElgeTAvijeTAmk)an (inter-convertible) .

rUbTI 22-4 muxgarkNþalén Acetyl CoA enAkñúgemtabUlIsekasika

Acetyl CoA

Proteins

Amino acids

Lipids

Fatty acids

Carbohydrates

Monosaccharides

Cholesterol Ketone bodies Fatty acids Citric Acid Cycle

Phosphplipids Triglycerides Steroids Bile salts

CO2 + H2O +ATP



3- karBinitüTUeTAénkardkdegðImEdlmanxül; (An Overview of Aerobic Respiration)

rcnasm<½n§rbs; mItUkugRDIsμ úRKsμajxøaMgNs; b:uEnþt<Mbn;nimYy²énmItUkugRDI mantYnaTIsMxan;enA kñúgdMeNIrkarén kardkdegðImEdlmanxül; (aerobic respiration) . kardkdegðImEdlmanxül;KWCa karbMEbknUvm:UelKulcMNIGaharEdlRtUvkarGuksIuEsn nig begáIt ATP . Gg;sIumsMrab; Citric acid cycle RtUv)aneKeXIjenAkñúglMhr matrix rbs;mItUkugRDI . Gg;sIumTI mYykñúgcMeNamGg;sIumTaMgenHsMrYlRbtikmμEdlP¢ab;bgÁúM Acetyl rbs; Acetyl CoA ¬manBIrGatUm kae)an¦ eTAelIm:UelKulEdlmanbYnGatUmkae)an (Oxaloactate) edIm,IbegáItCa Citrate ¬6 GatUm kae)an¦ . Gg;sIumEdlenAsl;sMrYlnUvmYyes‘rIénRbtikm μerobcMeLIgvij RbtikmμdkbgÁúM CO2 nig RbtikmμGuksIudU-er:dukkmμ . plitplcugbBa©b;én Citric acid cycle KW CO2 mU:elKul nig Oxaloactetate ¬m:UelKulEdlcab;epþImdMbUg¦ . enAkñúgdMNak;karCaeRcInén Citric acid cycle , subsrtrate TTYlrgnUvGuksIutkm μ . kñúgdMNak; karbIkñúgcMeNamdMNak;karTaMgenH mYyKUéneGLikRtug (a pair of electron) RtUv)anbBa¢ÚnBI substrate

eTA NAD+ edaybegáIteTACa NADH ¬NADH 3 m:UelKulkñúgmYyCMuvdþ¦ . enAkñúgdMNak;karepSgeTot mYyKUeGLikRtug RtUv)anbBa¢ÚnBI substrate eTA[ FAD edaybegáIteTACa FADH2 ¬FADH2 1 m:UelKulkñúgmYyCMuvdþ¦ . eGLikRtugRtUv)anqøgBI NADH b¤ FADH2 tamry³RbB½n§dwknaMeGLikRtugEdlsßitenAkñúgPñas xagkñúgénmItUkugRDI nig cugbBa©b; eTAGñkcab;ykeGLikRtugcugeRkaybMput (terminal electron

acceptor) KWGuksIuEsnCam:UelKul (molecular oxigen [O2]) . karbBa¢ÚneGLikRtugtamry³RbB½n§dwk naMeGLikRtugbgá[ Protons (H+) bUmBI matrix rbs;mItUkugRDIcUleTAkñúgEpñkEdlenAcenøaHPñas (intermembrane compartment) . lT§plenHKWCaepIgpÞúk H+ EdlmanfamBlx<s; (high-energy H+

reservoir) . kñúgdMNak;karcugeRkay famBlénepIgpÞúkRbUtugRtUv)anaeRbIedIm,IbegáIt ATP . dMNak;karcug eRkaybMputenH RtUv)anGnuvtþedaybNþúMGg;sIum ATP synthase . enAeBlRbUtughUrRtLb;cUleTAkñúg matrix rbs;mItUkugRDIvijtamry³rn§enAkñúgbNþúM ATP synthase Gg;sIumenHsMrYlkarsMeyaK ATP . dMeNIrEvgenH ¬Cab;Tak;TgeTAnwgdMeNIrkarEdleKehAfaCa Oxidative phosphorylation)

BIeRBaHfamBléneGLikRtugBI GuksIutkmμ rbs; substrates enAkñúg Citric acid cycle RtUv)aneRbIedIm,I eFIVkarP¢ab;bgÁúM Phosphate eTAelI ADP edIm,IbegáItCa ATP . karBiB’NnaBisþaréndMNak;karnimYy² rbs;dMNak;TaMgGs;enHnwgRtUvBinitükñúgEpñkxageRkamenH .



4- THE CITRIC ACID CYCLE (THE KREBS CYCLE)

1- Rbtikmμén Citric acid cycle Citric acid cycle CYnkalRtUv)aneKehAfa Krebs cycle

Cakitþiysdl;GñkRsavRCaveXIj KWelak Hans Krebs . vaKWCadMNak;karcugeRkayénkarbMEbkén Carbohydrate, xøaj; nig GasIutGamIneNEdlbeBa©jBIRbUetGIunéncMNIGahar ¬rUbTI 22-5¦ . edIm,Iyl;GMBIvdþEdlsMxan; \LÚveyIgtamemIlBIvasnarbs;bgÁúM Acetyl rbs; Acetyl CoA enA eBlvaqøgkat; Citric acid cycle . dMNak;karCaeRcInRtUv)anerobrab;xageRkamRtUvKñaeTAnwglMdab;enA kñúg Citric acid cycle EdlRtUv)ansegçbenAkñúgrUbTI 22-5 . RbtikmμTI 1 - bgÁúM Acetyl rbs; Acetyl CoA RtUv)anbBa¢ÚneTAelI Oxaloacetate enAkñúg RbtikmμsMrYledayGg;sIumeQ μaH Citrate synthase . plitplEdlekIteLIgKW Citrate .

RbtikmμTI 2- Gg;sIum Aconitase sMrYlkardkm:UelKulTwkBI Citrate [køayeTACa cis-

aconitate . Gg;sIumdEdlenH Aconitase bnÞab;mkk¾sMrYlkarbEnßmm:UelKulTwkeTAelI cis-aconitate

edIm,IbMElgvaeTACa Isocitrate . lT§plsrubéndMNak;karTaMgBIrenHKWkaeFIVGIusUEmrkmμrbs; Citrate

[eTACa Isocitrate . RbtikmμTI 3- dMNak;karGuksIutkm μTImYyén citric acid cycle RtUv)ansMrYleday Isocitrate

dehydrogenase . vaKWCaRbtikmμsμ úRKs μajEdlmanplitplbIekIteLIg ³ k- bgÁúM OH rbs; isocitrate TTYlrgGuksIutkmμeTACa Ketone

x- ]s μ ½nkabUnik (CO2) RtUv)anbeBa©j nig K- NAD+ RtUv)anTTYlrger:dukm μeTACa NADH . plitplénRbtikmμdkbgÁúMkabuksIul (decarboxylation reaction) KWCa α-ketoglutarate .

COO- │ CH2 │ HO—C—COO- │ CH2 │ COO-

Citrate

COO- │ C=O │ CH2 │ COO-

Fumarate

O ║ CH3—C~S—CoA H2O + +

Citrate synthase

COO- │ CH2 │ H—C—COO- │ HO—C—H │ COO-

Isocitrate

Aconitase

COO- │ CH2 │ C—COO- ║ C—H │ COO-

cis-Aconitate

COO- │ CH2 │ HO—C—COO- │ H—C—H │ COO-

Citrate

Aconitase



RbtikmμTI 4- Coenzyme A cUlsarCafμ IeTAkñúgdMNak;karenHén citric acid cycle . bNþúMrbs; α-ketoglutarate dehydrogenase Gnuvtþes‘rIénRbtikm μTaMgenH . bNþúMenHvadUya:gxøaMgeTAnwg Pyruvate dehydrogenase nig RtUvkar Coenzyme A dUcKña . Caf μImþgeTotplitplTaMgbIekIteLIg ³ k- α-ketoglutarate )at;nUvbgÁúMkabuksIuleRkamTMrg;Ca CO2 x- NAD+ TTYlrger:dukm μeTACa NADH nig K- Coenzyme A rYmCamYyplitpl (succinate) edIm,IbegáItCa Succinyl CoA . ehtudUecñH sm<½n§EdlekIteLIgrvag succinate nig Coenzyme A KWCasm<½n§ thioester EdlmanfamBlx<s; .

RbtikmμTI 5- Succinyl CoA RtUv)anbMElgeTACa Succinate enAkñúgdMNak;karenHEdlCab;Tak; Tgya:gxøaMgedaylkçN³KImI . Gg;sIum Succinyl CoA synthase sMrYlRbtikmμmYyKUr (a coupled

reaction) Edlsm<½n§ thioester EdlsMbUrfamBlrbs; Succinyl CoA RtUv)anGIuRdUlIs nig bgÁúM Phosphate minEmnsrIragÁ (Pi) RtUv)anbEnßmeTAelI GDP edIm,IbegáItCa GTP .


Isocitrate

COO- │ CH2 │ CH2 │ C=O │ COO- α-ketoglutarate

+ NAD+

Isocitrate dehydrogenase

+ CO2 + NADH

α-ketoglutarate dehydrogenase complex


COO- │ CH2 │ CH2 │ C~S—CoA ║ O Succinyl CoA

+ CO2 + NADH + NAD+ + CoenzymeA

COO- │ CH2 │ CH2 │ C~S—CoA ║ O

Succinyl CoA

COO- │ CH2 │ CH2 │ COO-

Succinate

+ GDP + Pi + GTP + CoenzymeA Succinyl CoA synthase



Gg;sIumepSgeTot dinucleotide diphosphokinase sMrYlkarbBa¢ÚnbgÁúM Phosphate BI GTP eTA ADP edIm,IbegáItCa ATP .

RbtikmμTI 6- bnÞab;mk Succinate dehydrogenase sMrYRbtikmμGuksIutkmμrbs; succinate

[køayeTACa furmarate enAkñúgdMNak;karbnÞab; . Pñak;garGuksIutkmμ Flavin adenine dinucleotide

(FAD) TTYlrger:dukkm μenAkñúgCMhanenH .

RbtikmμTI 7 - karbEnßm H2O eTAelIsm<½n§BIrCan;rbs; furmarate pþl;[Ca Malate . Gg;sIum Furmarase CaGñksMrYRbtikmμenH .

RbtikmμTI 8 - enAkñúgdMNak;karcugeRkayrbs; Citric acid cycle , Malade dehydrogenase

sMrYler:dukm μrbs; NAD+ [eTACa NADH nig GuksIutkmμén Malate [eTACa Oxaloacetate . BIeRBaH Citric acid cycle cab;edaykarbEnßbgÁúM Acetyl eTAelI Oxaloacetate dUecñHeyIgmanrgVg;mYyCMu .

GTP +ADP GDP +ATP

Dinuclotide diphosphokinase

COO- │ CH2 │ CH2 + FAD │ COO-

Succinate

COO- │ C—H ║ H —C + FADH2 │ COO-

Fumarate

Succinate dehydrogenase

COO- │ C—H ║ H —C + H2O │ COO-

Succinate

COO- │ HO—C—H ║ H —C—H │ COO-

Malate

Furmarase

COO- │ HO—C—H │ CH2 + NAD │ COO-

Malate

COO- │ C=O │ H —C + NADH │ COO-

Oxalaacetate

Malate dehydrogenase



rUbTI 22-5 Rbtikmμén Citric Acid Cycle

rUbTI 22-5 Rbtikmμén Citric acid cycle

COO- │ CH2 │ HO—C—COO- │ CH2 │ COO-

Citrate


Isocitrate


O ║ C~S—CoA │ CH2 │ CH2 │ COO-

Succinyl CoA

COO- │ CH2 │ CH2 │ COO-

Succinate

COO- │ CH ║ HC │ COO-

Fumarate

COO- │ HO—C—H │ CH2 │ COO-

Malate

COO- │ C=O │ CH2 │ COO-

Fumarate

O ║ CH3—C~S—CoA

1. Citrate synthase

1. Aconitase

3. Isocitrate dehydrogenase

4. α-ketoglutarate dehydrogenase complex

5. Succinyl CoA synthase

6. Succinate dehydrogenase

8. Malate dehydrogenase

7. Fumarase

NAD+

CO2+NADH

FADH2

FAD

GTP + Coenzyme A GDP + Pi

NAD+

NADH

Coenzyme A + NAD+

CO2+NADH

H2O



5- karRtÜtBinitüén CITRIC ACID CYCLE

esÞIrEtdUc glycolysis, Citric acid cycle eqøIytbeTAnwgesckþIRtUvkarfamBlrbs;ekasika. pøÚvenHekIneLIgenAeBlmankarTamTar ATP ya:gxøaMg nig vafycuHenAeBl ATP elIs . enAkñúgCMBUk eRkaybMputeyIgnwgeXIjGg;sIumCaeRcInEdlsMrYlRbtikmμrbs; glycolysis KWCa Allosteric enzyme. dUcKaenHEdr Gg;sIumbYn b¤ bNþúMGg;sIumCab;Tak;TgeTAnwgkareFIVGuksIutkmμeBjeljrbs; Pyruvate

tamry³ Citric acid cycle k¾Ca Allosteric enzymes Edr . BYkvaGaccgP¢ab;eTAnwg effectors dUcCa ATP b¤ ADP EdleFIV[xUcTMrg;énTIkEnøgskmμrbs;Gg;sIum (Enzyme active site) rYmTaMgkareP¶ac GRtarbs;Rbtikmμ (Positive allosterism) b¤ k¾raraMgRbtikm μ (Negative allosterism) . GaRs½yedaykarRtYtBinitüpøÚvenHRtUvEteTogTat; (Precise) vamandMNak;karGg;sIumCaeRcIn EdlRtUv)anEktMrUv . dMNak;karTaMgenHRtUv)ansegçbenAkñúgrUbTI 22-6 xageRkamenH ³ 1- karbMElgBI Pyruvate eTACa Acetyl CoA - bNþúM Pyruvate dehydrogenase RtUv)an raraMgedaykMhab;x<s;rbs; ATP, acetyl CoA nig NADH . ehtuenH vtþmanrbs;smasFatuTaMgenH kñúgkarpþl;sBaØaPaBsMbUrEdlekasikamankarpÁt;pÁg;RKb;RKannUvfamBl nig dUecñH emtabUlIsfamBl (energy metabolism) RtUv)anfycuH . 2- karsMeyaK Citrate BI Oxaloacetate nig Acetyl CoA - Gg;sIum Citrate synthase

KWCa Allosteric enzyme . kñúgkrNIenH Negative effector KW ATP . Caf μ ImþgeTot karenHKWCakarBit BIeRBaHkarelIs ATP enHbgðajfaekasikamanPaBsMbUrnUvfamBl . 3- GuksIutkmμ nig kardkbgÁúMkabuksIulén Isocitrate eTACa α-ketoglutarate - Isocitrate

dehydrogenase k¾Ca Allosteric enzyme b:uEnþGg;sIumenHRtUv)anRtYtBinitüeday Positive allosteric

effector (ADP) . ADP KWCasBaØa EdlkMritén ATP RtUvEtTab nig ehtudUecñHGRtaén Citric acid

cycle nwgRtUvekIneLIg . KYr[cab;GarmμN_ Isocitrate dehydrogenase k¾RtUv)anraraMgpgEdreday kMritx<s;én NADH nig ATP . 4- karbMElgBI α-ketoglutarate eTACa Succinyl CoA - bNþúMGg;sIum α-ketoglutarate

dehydrogenase RtUv)anraraMgedaykMritx<s;énplitplrbs;RbtikmμEdlbNþúMGg;sIumenH)ansMrYl EdlmaneQ μaHfa NADH nig Succinyl CoA . vaRtUv)anraraMgCabnþeTotedaykMhab;x<s;rbs; ATP .



rUbTI 22-6 karEktMrUvénbNþúMGg;sIum Pyruvate Dehydrogenase nig Citric Acid Cycle

6- OXIDATIVE PHOSPHORYLATION

enAkñúgEpñkTI 22-3 eyIg)anbBa¢ak;faeGLikRtugEdldwknaMeday NADH GacRtUv)aneRbIedIm,I begáItbIm:UelKul ATP nig eGLikRtugEdldwknaMeday FADH2 GacRtUv)aneRbIedIm,IbegáItBIrm:UelKul ATP . etIdMeNIrkarGIVEdlfamBlrbs;eGLikRtugdwknaMeday Coenzymes TaMgenHRtUv)anbMElgeTA CafamBl ATP ? vaKWCaes‘rIénRbtikm μEdleKehAfa Oxidative Phosphorylation Edldak;rYmKña nUvGuksIutkm μrbs; NADH nig FADH2 edIm,I[køayCa Phosphorylation én ADP [)anCa ATP . 1- RbB½n§dwknaMeGLikRtug nig cMeNatGIuy:ugGIuRdUEsn (Electron transport system and the hydrogen ion gradient)

muneBleyIgBüayamyl;BIynþkarrbs; Oxidative Phosphorylation cUrBinitüemIlCadMbUgeTA elIm:UelKulEdlGnuvtþnUvdMeNIrkarsμúRKs μajenH . edayP¢ab;enAkñúgPñasxagkñúgénmItUkugRDI KW RbB½n§

Pyruvate

Acetyl CoA

Pyruvate

Oxaloacetate Citrate

Malate Isocitrate

α-ketoglutarate

Succinyl CoA

Succinate

Furmarate

Inhibited by NADH, ATP and Acetyl CoA

Inhibited by ATP

Inhibited by ATP NADH, Stimulated by ADP

Inhibited by Succinyl CoA, NADH, & ATP

4

3

2

1



dwknaMeGLikRtug (Electron Transport System) . BYkvaTaMgenHekIteLIgBIGñkdwknaMeGLikRtugmYy es‘rI rYmman Coenzymes nig Cytochromes . RKb;m:UelKulTaMgenHsßitenAkñúgPñaskñúgreboberobcM EdlGnuBaØati[BYkvaeFIV[eGLikRtugqøgkat;BImYyeTAmYybnÞab;eTot . CYrCalMdab;rbs;GñkdwknaM eGLikRtugRtUv)aneKehAfa RbB½n§dwknaMeGLikRtugénkardkdegðIm (Respiratory electron transport

system) rUbTI 22-7 . enAeBlGñknwgrMBwgkñúgrebobenHnUvRbtikmμGuksIutkmμ-er:dukm μekIteLIg eGLikRtug)at;bg;famBlmYycMnYnrYmCamYykarbBa¢ÚnnimYy² . mYycMnYnénfamBlTaMgenHnwgRtUv)an eRbIedIm,IbegáIt ATP . enAnwgTIkEnøgbI enAkñúgRbB½n§dwknaMeGLikRt RbUtug (proton) GacRtUv)anbUmBI matrix rbs;mItU-kugRDIeTAlMhcenøaHPñas (intermembrane space) . RbUtugTaMgenHbNþal[epIgpÞúkGIuRdUEsnEdlman famBlx<s; (high-enrgy H+ reservoir) . enAnwgTIkEnønimYy²kúñgcMeNamTIkEnøgTaMgbI GIuy:ug H+

RKb;RKan;RtUv)anbUmcUleTAkñúgepIgGIuRdUEsnedIm,IbegáIt ATP mYym:UelKul . TIkEnøgTImYyKW NADH

dehydrogenase . GaRs½yeGLikRtugBI NADH cUleTAkñúgRbB½n§dwknaMeGLikRtugedayRtUv)anbBa¢Ún eTA[ NADH dehydrogenase RKb;TIkEnøgTaMgbIbUmykRbUtugya:gskmμ nig bIm:UelKul ATP RtUv)an ekIteLIg ¬emIlrUbTI 22-7¦ . FADH2 KWCaGñkpþl;eGLikRtugEdlmanfamBltic . vabBa¢ÚneGLik-RtugeTA[GñkdwknaMEdlbnþBI NADH2 dehydrogenase . CalT§pl enAeBl FADH2 TTYlrgGuksIut-kmμ manEtkEnøgTIBIr nig TIbIeTEdlbUmRbUtug nig manEtBIrm:UelKul ATP RtUv)anekIteLIg . smasFatucugeRkaybMputEdlRtUvkarsMrab; oxidative phosphorylation KWCabNþúM multi-

protein EdleKehAfa ATP synthase eKk¾ehApgEdrfa F0F1 complex ¬emIlrUbTI 22-7¦ . eday latsn§wgenAkñúgPñasxagkñúgénmItUkugRDIKWbNþúM Protein (F0) Edlpþl;CapøÚvsMrab;RbUtugGacqøgkat; . edaylatsn§wgcUleTAkñúg matrix rbs;mItUkugRDIKWCabNþúMRbUetGIunragEsV‘ (F1) EdlmanlT§PaBCa Gg;sIumkñúgkarP¢ab;bgÁúM Phosphate eTAelI ADP edIm,IbegáIt ATP . 2- ATP synthase nig karplit ATP

etIynþkarEdls μ úRKs μajTaMgGs;enHbMeBjmuxgarrebobdUemþc ? NADH dwknaMeGLikRtugeday manRbPBBI glucose eTA[GñkdwknaMTImYyrbs;RbB½n§dwknaMeGLikRtug NADH dehydrogenase ¬emIl rUbTI 22-7¦ . enATIenaH NADH RtUv)anTTYlrgGuksIutkm μeTACa NAD+ EdlRtLb;cUleTAkñúgTIkEnøg rbs; Citric Acid Cycle EdlRtUvTTYlrger:dukmμsarCafμ I . dUcenAkñúgrUbTI 22-7 bgðaj ¬ExSdac;²B’N Rkhm¦ mYyKUrbs;eGLikRtug (a pair of electron) qøgkat;eTAGñkdwknaMeGLikRtugbnÞab; nig RbUtug RtUv)anbUmcUleTAkñúgEpñkenAcenøaHPñas (intermembrane compartment) . eGLikRtugqøgkat;Ca bnþbnÞab; tamry³RbB½n§dwknaMeGLikRtug nig enATIkEnøgBIrepSgeTot RbUtugmkBI matrix RtUv)anbUmcUl



eTAkñúgEpñkcenøaHPñas . rYmCamYynwgkarbBa¢ÚnnimYy² eGLikRtug)at;nUvfamBlmYycMnYn . famBl rbs;eGLikRtugmYycMnYnRtUv)aneRbIedIm,IdwknaM H+ Pñasxagkñúgrbs;mItUkugRDI nig cUleTAkñúgepIg H+ . cugbBa©b; eGLikRtug)aneTAdl;GñkdwknaMcueRkaybMput . \LÚvenHBYkvamanfamBltictYckñúgkareFIV[ sMerc)annUvkargarNamYyEfmeTot b:uEnþBYkvaRtUvEtTTYl)anBIGñkcab;ykeGLikRtugcugeRkaymYy cMnYn ehtudUecñH RbB½n§dwknaMeGLikRtugGacbnþbMeBjmuxgar)an . enAkñúgsrIragÁElmanxül; Gñkcab; eGLikRtugcugbBa©b; KWCam:UelKulGuksIuEsn (O2) nig plitplKWCa H2O .

rUbTI 22-7

ehtudUecñH RbB½n§dwknaMeGLikRtug GnuvtþnUvGuksIutkm μrbs; NADH nig FADH2 CamYynwgm:Uel-KulGuksIuEsnEdlCaGñkcab;ykeGLikRtugcugbBa©b; . dUcEdl)anbBa¢ak;BIxagmunmk FADH2 pþl; eGLikRtugrbs;vaeTA[GñkdwknaMEdlmanfamBlTab . lT§plénkarenHKWfaRbUtugtictYcRtUv)anbUm cUleTAkñúgepIg H+ nigman ATP tictYcRtUv)anbegáIteLIg . enAeBlRbB½n§dwknaMeGLikRtugenAbnþbMeBjmuxgar kMhab;x<s;énRbUtugekIteLIgenAkñúglMhr GnþrPñas (intermembrane space) . karenHbegáItnUvcMeNat (gradient) H+ BaseBjPñasxagkñúg



rbs;mItUkugRDI . cMeNatrebobenHKWCaRbPBfamBlya:gxøaMg dUcCaTwkRtUv)aneKsþúkenAxagkñúgTMnb;. mItUkugRDIeFIVkareRbIénfamBlEdlmansukþanuBlrbs;cMeNatedIm,IsMeyaKfamBl ATP . vaKWCa ATP synthase EdlRbmUlnUvfamBlrbs;cMeNatenHedIm,IbegáIt ATP . eTaHbICaPñas xagkñúgrbs;mItUkugRDIvaminRCabxøaMgk¾eday ATP synthase pþl;nUvpøÚv (F0) EdlRbUtugGacRtLb;eTAkñúg matrix vijtampøÚvenH . enAeBlRbUtugqøgkat;tamry³ F0 EpñkGg;sIumrbs; ATP synthase (F1) TTYl rgnUvkarpøas;bþÚrrUmrag nig sMrYlkarP¢ab;bgÁúM Phosphate eTAelI ADP edIm,IbegáIt ATP . enAkñúgrebob enHfamBlrbs;epIg H+ RtUv)anRbmUledIm,IeFIV ATP . 3- segçbénlT§plbegáItfamBl (Summary of the Energy Yield)

mYyCMuén Citric Acid Cycle bNþal[mankarplitnUv CO2 BIrmU:elKul NADH bIm:UelKul FADH2 mYym:UelKul nig ATP mYym:UelKul . Oxidative Phosphorylation bNþal[man ATP

bIm:UelKulkñúg NADH mYym:UelKul nig ATP BIrm:UelKulkñúg FADH2 mYym:UelKul . karelIkElgEt mYyKt;cMeBaHkarpþl;famBlTaMgenHKW NADH EdlRtUv)anplitenAkñúgsIutU)aøs;kñúgry³eBl glycolysis. Oxidative Phosphorylation pþl; ATP EtBIrm:UelKulkñúg cytoplasmic NADH mYym:Uel-Kul . ehtuplcMeBaHkarenHKWfafamBlRtUvEt)ancMeNayedIm,IeFIV[eGLikRtugeFIVdMeNIrBI NADH

enAkñúgsIutU)aøs;eTAelI FADH2 enAkñúgmItUkuRDI . edaydwgGMBIB’t½manenH nig caMenAkñgstiGarmμN_EdlfaeKRtUvkarBIrCMuén Citric acid cycle

eyIgGacbUksrubnUvfamBlsrubEdl)anmkBIkareFIVGuksIutkmμeBjeljén glucose mYym:UelKul. lT§plsrubrbs;GuksIutkmμeBjeljrbs; glucose KWCakarplitnUv ATP 36 m:UelKul ¬rUbTI 22-8¦ . karenHtMNag[karRbmUlfamBlRbEhl 40 % énfamBlEdlmansukþanuBlrbs; glucose . kaekMNt;lT§plrbs; ATP BIkardkdegðImEdlmanxül; cUrKNnacMnYnrbs; ATP Edlplit)anedaykareFIVGuksIutkm μeBjeljrbs; glucose mYy m:UelKul . dMeNaHRsay Glycolysis Substrate-level phosphorylation 2 ATP 2 NADH x 2 ATP/cytoplasmic NADH 4 ATP Conversion of 2 pyruvate molecules to 2 Acetyl CoA molecules 2 NADH x 3 ATP/NADH 6 ATP Citric Acid Cycle 2 GTP x 1 ATP/GTP 2 ATP 6 NADH x 3 ATP/NADH 18 ATP 2 FADH2 x 2 ATP/FADH2 4 ATP ------------------------ Total 36 ATP



rUbTI 22-8

Aerobic metabolism vamanRbsiT§Cag anaerobic metabolism eRcInCagxøaMgNas; . famBl d¾eRcInRtUv)anRbmUleday aerobic metabolism manplvi)akya:gFM (enormous consequences)

cMeBaHBiPBCIvsaRsþ . PaKeRcInénfamBlbeBa©jedayGuksIutkmμrbs;\n§³ (fuels) minRtUv)an)at;bg; enAeBlekþA b:uEnþRtUv)anrkSaTukeRkamTMrg;Ca ATP . srIragÁEdlmanfamBlsMbUrvivtþn_bnþicmþg²eTACa srIragÁBhuekasika (multicellular organism) nig begáItnUvmuxgarBiess . Caplvi)akéntMrUvkar famBlrbs;BYkvaRKb;srIragÁBhuekasikaKWCasrIragÁEdlRtUvkarGuksIuEsn (aerobic) . 7- karbMEbkGasIutGamIneN (THE DEGRADATION OF AMINO ACIDS)

Glucose

PGAL

Pyruvate

ATP

ATP

2 NADH 2 NADH

Citric Acid Cycle and Phosphorylation reaction

Glycolysis

Acetyl CoA

2 NADH

6 NADH

2 FADH2

2 GTP

2 ATP

2 ATP

32 ATP

Electron

Oxidative phosphorylation

36 ATP



Carbohydrate minEmnCaRbPBfamBlEtmYyKt;rbs;eyIgeT . dUcEdleyIg)aneXIjenAkñúg CMBUkTI 21 RbUetGIunenAkñúgrbbGahar (dietary protein) RtUv)anbMEbkeTACa GasIutGamIneN (amino

acids) Edlk¾GacRtUv)aneRbICaRbPBfamBlEdr eTaHbICakarenHminEmnCamuxgaremtabUlIssMxan; rbs;BYkvak¾eday . PaKeRcInénGasIutGamIneNRtUv)aneRbIsMrab;famBlEdl)anmkBIGahar . CakarBit vaCab;Tak;TgEtkñúglkçx½N§EdlmanTurÖPikS (starvation conditions) enAeBl sþúkbMrug glycogen nig lIBIdRtUv)anGs; EdlsBa’gÁkaycab;epþImdut (burn) RbUetGIunrbs;va ¬Edl)anBIsac;dMu¦ Ca\n§³ . vasnaénl,ayrbs;GasIutGamIneN EdlRtUvpþl;edaykarbMEbkRbUetGIunGaRs½yeTAnwgtulü-PaBrvagesckþIRtUvkarcMeBaHGasIuGamIneNsMrab;CIvsMeyaK nig esckþIRtUvkarsMrab;famBlekasika . manEtBYkGasIutGamInTaMgenaH EdlminRtUvkarsMrab;karsMeyaKRbUetGIunCacugbBa©b;RtUv)anbMElgeTACa FatukNþalén Citric acid cycle (Citric acid cycle intermediates) nig RtUv)aneRbICa\n§³ . karbMEbkGasIutGamIneNRtUv)anekIteLIgCabzmenAkñúgeføIm nig ekIteLIgCaBIdMNak;kar. dMNak;karTI 1 KWCakardkecjnUvbgÁúMGamInenATItaMg alpha (α-amino group) nigdMNak;karTI 2 KWkar bMEbkeRKagGatUmkae)an . cMeBaHfnikstVenAnwgdI (land mammals) bgÁúMGamInCaTUeTAbBa©b;kñúgTMrg; Ca Urea EdlRtUv)anbeBa©jmkeRkAtamTwkenam . eRKagGatUmkae)anGacRtUv)anbMElgeTACa smasFatuepSg² rYmmanFatukNþalén Citric Acid Cycle, pyruvate, acetyl CoA b¤ Acetoacetyl

CoA. karbMEbknUveRKagGatUmkae)anRtUv)ansegçbenAkñúgrUbTI 22-9 . RbtikmμdkbgÁúMGamIn nig vasnarbs;eRKaGatUmkae)anrbs;GasIutGamIneNKWCakarepþatkaryk;citþTukdak;enAkñúgEpñkenH . 1- kardknUvbgÁúMGamInenATItaMgGal;hVa :Transamination

(Removal of α-amino groups)

dMNak;kardMbUgénkarbMEbkGasIutGamIneN ¬kardknUvbgÁúMGamInenAelIkae)an α¦ RtUv)anGnuvtþ edayRbtikmμ Transamination . Aminotransferases k¾GacRtUv)aneKehAEdrfaCa Transaminases sMrYlkarbBa¢ÚnbgÁúMGamInenAelIkae)an α BI α-amino acid eTA[ α-keto acid ³

bgÁúMGamInEdlenAelIkae)an α énGasIutGamIneNCaeRcInRtUv)anbBa¢ÚneTA[ α-ketoglutarate

edIm,IbegáIt)anCa Glutamate nig keto acid fμ I . GMbUr glutamate rbs; Aminotransferases man

NH3 O │ ║ H—C—COO- + C—COO- І І R1 R2 Donor Acceptor amino acid keto acid

O NH3 ║ │ C—COO- + H—C—COO- І І R1 R2 Keto acid of New amino acid amino acid

Transaminase



sarsMxan;CaBiess BIeRBaH α-Keto acid EdlRtUvKñaeTAnwg glutamate KWCa α-ketoglutarate ¬EdlCa FatukNþalrbs; Citric Acid Cycle¦ . ehtudUcenH Glutamate aminotransferases pþl;nUvTMnak;TMng edaypÞal;rvagkarbMEbkGasIutGamIneN nig Citric Acid Cycle . Aminotransferase sMxan;epSgeTotKW Aspartic aminotransferase EdlsMrYlkarbBa¢ÚnbgÁúM GamInenAelIkae)an α BI Aspartate eTA[ α-ketoglutarate edaypþl;Ca Oxaloacetate nig glutamate³

Aspartate + α-Ketoglutarate ⇄ Oxalaacetate + Glutamate

Pyruvate

Acetyl CoA

Citrate Oxalacetate

α-ketoglutarate

Succinate

Fumarate

Citric Acid Cycle

Alanine Glycine Cysteine Serine

Isoleucine Leucine

Tryptophane

Fatty Acids

Acetoacetyl CoA

Keto bodies

Leucine Lysine

Phenylalanine Tyrosine

Aspartic acid

Glutamic acid Histidine Proline

Arginine

Isoleucine Methionine Threonine

Valine

Tyrosine Phenylalanine



Transaminase EdlsMxan;enAkñúgCalikafnikstVKW Alanine aminotransferase EdlsMrYlkarbBa¢Ún bgÁúMGamInenAelIkae)an α BI Alanine eTAelI α-ketoglutarate nig begáIt)anCa Pyruvate nig Glutamate .

RKb;én Aminotransferases eRcInCag 50 EdlRtUveKRsavRCaveXIjvaRtUvkar Coenzyme

Pyridoxal Phosphate . Coenzyme enHmanRsLayBI Vitamin B6 . Rbtikmμ Transamination )anbgðajBIxagelIhak;dUcCakarbBa¢ÚnFm μta b:uEnþ CakarBit Rbtikmμ enHvasμ úRKs μajCagenHq¶ayxøaMgNas; . Aminotransferase cgP¢ab;eTAnwgGasIutGamIneN ¬Aspartate enAkñúgrUbTI 22-11a¦ enAkñúgTIkEnøgskmμ . bnÞab;mk bgÁúMGamInenAelIkae)an α rbs; Aspartate RtUv)anbBa¢ÚneTA[ Pyridoxal phosphate edaybegáItCa Pyridoxamine phosphate nig Oxaloacetate ¬rUbTI 22-11b¦ . bnÞab;mkbgÁúMGamInRtUv)anbBa¢ÚneTA[ α-keto acid kñúgkrNIenH α-

ketoglutarate ¬rUbTI 22-11 c¦ edIm,IbegáItCaGasIutGamIneN Glutamate ¬rUbTI 22-11 d¦. bnÞab;mkeyIgnwgBinitüBIvasnarbs;bgÁúMGamInEdlRtUv)anbBa¢ÚneTA[ α-ketoglutarate edIm,IbegáItCa glutamate . 2- kardknUvbgÁúMGamInenATItaMgGal;hVa :Oxidative Deamination

enAkñúgdMNak;karbnÞab;énkarbMEbkGasIutGamIneN Ammonium ion RtUv)anbeBa©jBI glutamate EdlRtUv)anbegáIteday Aminotransferase . karEbkén glutamate enHRtUv)ansMrYl edayGg;sIum glutamate dehydrogenase ekIteLIgdUcxageRkam ³

Glutamate + NAD+ + H2O ⇄ NH4+ + α-ketoglutarate + NADH + H+

karenHKWCa]TahrN_mYyén Oxidative deamination dMeNIrkarénGuksIutkmμ-er:dukmμEdl NAD+ TTYlrger:dukmμeTACa NADH nig GasIutGamIneNRtUv)andkbgÁúMGamInecj . karsegçbén RbtikmμkardkbgÁúMGamIn (deamination reaction) RtUv)anBiB’NnaenAkñúgrUbTI 22-12 3- vasnaéneRKagGatUmkae)anGasIutGamIneN eRKagGatUmkae)anEdlRtUv)anplitedayRbtikm μxagelITaMgenH nig Rbtikm μ deamination

epSgeTotcUleTAkñúg glycolysis b¤ Citric acid cycle enAkñúgdMNak;karCaeRcIn . edaysarehtuenH eyIg)aneXIjfa Transamination bMElg Aspartate eTACa Oxaloacetate nig Alanine eTACa

Alanine + α-Ketoglutarate ⇄ Pyruvate + Glutamate



Pyruvate . TItaMgEdleRKagGatUmkae)anrbs;GasIutGamIneNepSg²cUleTAkñúgpøÚvRbmUlfamBlRtUv )ansegçbenAkñúgrUbTI 22-9 .

rUbTI 22-11 ynþkarénkarbBa¢ÚnbgÁúMGamIn

rUbTI 22-12 segçbénkardkbgÁúMGamInrbs;GasIuGamIneN nig vasnarbs; NH4

+



8- THE UREA CYCLE

Oxidative deamination begáItnUvcMnYnya:geRcInén NH4+ . RbsinebIkarenHminRtUv)aneFIV

smahrnkm μeTAkñúgmU:elKulCIvsaRsþ nig RtUv)andkecjBIsBa’gÁkay vanwgQandl;nUvkMritBul ya:gelOn . kMritén NH4

+ ekIneLIgvanaM[BulBIeRBaH NH4+ KWCa)asexSayEdlGacbNþal[man

karekIneLIgnUv pH EdlGacbNþal[søab; . ehtudUecñH vamansarsMxan;xøaMgNas;cMeBaHkarrs;enA rbs;srIragÁkñúgkarmanpøÚvmYysMrab;beBa©jecalnUv NH4

+ edayminKitGMBIkarRtUvkarfamBl . cMeBaH mnusS nig BYkstVEdlmanq¥wgxñgrs;enAelIdI meFüa)ayénkardk NH4

+ ecjKWCa Urea Cycle . dUcEdl)anbgðajBIxagelI NH4

+ CaCatiBulya:gxøaMg . BYkvaRtUv)aneFIV[ElgBulenAkñúgeføIm edaykarbMElg NH4

+ eTACa Urea . pøÚvenH RtUv)aneKehAfaCa Urea Cycle CaviFIEdl NH4+ BulRtUv

)andkecjBIkñúgQam . karelIsén NH4+ RtUv)aneFIVsmahrnkmμeTACa urea RtUv)anbeBa©jtam

Twkenam ¬emIlrUbTI 22-13¦ . Rbtikm μén Urea Cycle

Rbtikmμén Urea Cycle 5 RtUv)anbgðajenAkñúgrUbTI 22-13 nig karBiB’NnaBisþarrbs; RbtikmμRtUv)ansegçbdUcxageRkam . dMNak;karTI 1- dMNak;karTImYyénvdþKWCaRbtikmμEdl CO2 nig NH4

+ begáIt)anCa Carbamoyl

phosphate . Rbtikmμ ¬EdlRtUvkar ATP nig H2O Edr¦ ekIteLIgenAkñúgmItUkugRDI nig RtUv)ansMrYl edayGg;sIum Carbamoyl phosphate synthetase .

dMNak;karTI 2- enAeBlenaH Carbamoyl phosphate rYmCamYyGasIuGamIneN Ornithine

edIm,IbegáItCaGasIutGamIneN Citrulline . Rbtikm μenHk¾ekIteLIgenAkñúgmItUkugRDIEdr nig RtUv)ansMrYl edayGg;sIum Ornithine transcarbamoylase .

O O ║ ║ CO2 + NH4

+ + 2 ATP + H2O → H2N—C—O—P—O- + 2 ADP + Pi + 3 H+ │ O- Carbamoyl phosphate



rUbTI 22-13



dMNak;karTI 3- Citrulline RtUv)anbBa¢ÚneTAkñúgsIutU)aøs; nig \LÚvrYmCamYynwg Aspartate edIm,I begáItCa Argininosuccinate . Rbtikm μenH EdlRtUvkarfamBlbeBa©jedaykarGIuRdUlIs ATP sMrYl edayGg;sIum Argininosuccinate synthase .

dMNak;karTI 4 - \LÚv Argininosuccinate RtUv)ankat;pþac;edIm,IbegáItCaGasIutGamIneN Arginine nig FatukNþalrbs; Citric acid cycle KW Fumarate . RbtikmμTaMgenHRtUv)ansMrYleday Gg;sIum Argininosuccinate lyase .

NH3 │ H—C—H O O │ ║ ║ H—C—H + H2N—C—O—P—O- │ │ H—C—H O- │ H—C—H

│ COO- Ornithine Carbamoyl Phosphate

H—N—CO—NH3 │ H—C—H │ H—C—H + Pi │ H—C—H │ H—C—H

│ COO- Citrulline Phosphate

H—N—CO—NH3 │ H—C—H COO- │ │ H—C—H + H3

+ N—C—H

│ │ H—C—H CH2 │ │ H—C—H COO-

│ COO- Citrulline Aspartate

NH2+ COO-

║ │ H—N—C—NH——C—H │ │ H—C—H CH2 │ │ H—C—H COO- │ H—C—H │ H—C—H

│ COO- Argininosuccinate

NH2+

║ H—N—C—NH2 │ H—C—H COO- │ │ H—C—H + H3

+ + C—H

│ ║ H—C—H C—H │ │ H—C—NH3

+ COO-

│ COO- Arginine Fumarate

NH2+ COO-

║ │ H—N—C—NH——C—H │ │ H—C—H CH2 │ │ H—C—H COO- │ H—C—H │ H—C—H

│ COO- Argininosuccinate



dMNak;karTI 5 - CacugbBa©b; Arginine RtUv)an Hydrolyse edIm,IbegáItCa Urea EdlCa plitplRtUv)anbeBa©jecaltamTwkenam nig Ornithine CaRbtikredImdMbUgbMputenAkñúgvdþenH . Arginase KWCaGg;sIumEdlsMrYRbtikmμenH .

cUrkt;sMKal;fa bgÁúMGamInmYykñúgcMeNambgÁúMGamInTaMgBIrrbs; urea TTYl)anBI NH4 nig TIBIr )anmkBI Aspartate . CaTUeTAvamanvamankarcMLgCMgWEdlbNþalmkBIPaBfycuHénGg;sIummYykñúgcMeNamGg;sIum rbs Urea Cycle . sar³sMxan;én Urea Cycle KWvac,as;enAeBleyIgBicarNakenSameraKsBaØaEdl TTYlrgedaybuKÁlEd;lQW (afficted individuals) . PaBfycuHénGg;sIum Urea cycle bgá[mankar ekIneLIgnUvkMhab;rbs; NH4

+ Calkçx½NÐEdleKsÁal;faCa Hyperammonenia . RbsinvamanPaBcuH TaMgRsug (complete deficiency) énGg;sIummYykñúgcMeNamGg;sIumTaMgenaH lT§plKWeskþIsøab;mun Gayu (early infancy) . RbsinebIvamanPaBfycuHedayEpñkénGg;sIummYykñúgcMeNamGg;sIumén Urea

cycle lT§plGacCakaryWtya:v (retardation) Rbkac; (convulsions) nig kark¥Üt . enAkñúgTMrg;Rsal én hyperammonenia rbbGaharEdlmanRbUetGIunticbNþal[mankMhab;Tabén NH4

+ enAkñúgQam nig mankenSameraKsBaØamanPaBRsYcRsavtiNas; . 9- TidæPaBTUeTAénGaNabUlIs ³

THE CITRIC ACID CYCLE CaRbPBénFatukNþalCIvsMeyaK mkdl;eBlenH eyIg)anniyaGMBI Citric acid cycle RKan;EtCaynþkarRbmUlfamBl . eyIg )aneXIjehIyfa carbohydrates nig amino acids cUleTAkñúgpøÚvedaypÞal;enAkñúgdMNak;karepSg² nig TTYlrgGuksIutkm μedIm,IbegáItCa NADH nig FADH2 Edl ¬tammeFüa)ayén oxidative phospho-

rylation ¦ RtUv)aneRbIedIm,IbegáIt ATP .

NH2+

║ H—N—C—NH2 │ H—C—H │ H—C—H + H2O │ H—C—H │ H—C—NH3

+

│ COO- Arginine Water

NH3

+ │ H—C—H O │ ║ H—C—H + H3

+ + H2N—C—NH2

│ H—C—H │ H—C—NH3

+

│ COO- Ornithine Urea



b:uEnþtYnaTIrbs; Citric Acid Cycle enAkñúgemtabUlIsekasikaCab;Tak;TgeRcInCagEtkñúg Catabolism . vaedIrtYnaTIsMxan;enAkñúg Anabolism b¤ CIvsMeyaK pgEdr . rUbTI 22-14 bgðajBI tYnaTIkNþalrbs; gltcolysis nig Citric Acid Cycle CaRbtikmμEdlRbmUlfamBl k¾dUcCatYnaTIrbs; BYkvaCaRbPBén precursor CIvsMeyaK . dUcEdlGñkGacsgS½yrYcmkehIyBIkarBitEdlfaGasIutGamIneNGacRtUv)anbMElgeTACaFatu kNþalrbs; Citric Acid Cycle ehIyFatukNþalén Citric Acid Cycle dUcKñaTaMgenHk¾GacRtUv)aneRbI CasarFatucab;epþImsMrab;karsMeyaKénGasIutGamIneNEdr . Oxaloacetate pþl;nUveRKagGatUmkae)an sMrab;karsMeyaKmYydMNak; (one-step synthese) rbs;GasIutGamIneN Aspartate edayRbtikmμ Transamination reaction . Oxaloacetate + glutamate ⇄ Aspartate + α-ketaglutarate

epSgBIkarpþl; Aspartate sMrab;karsMeyaKRbUetGIun RbtikmμenHpþl;nUv Asartate sMrab; Urea

cycle . Asparasine RtUv)anplitBI Aspartate edayRbtikmμbEnßmbgÁúMGamIn (amination reaction). Aspartate + NH4 + ATP → Asparasine + AMP + PP1 + H+

α-ketoglutarate eRbICaExSkae)anedImdMbUg (the starting carbon chain) sMrab;GMbUrrbs;GasIut GamIneNrYmman glutamate, glutamine, proline, nig arginine . Glutamate mansar³sMxan;Ca Biess BIeRBaHvaeRbICaGñkpþl;nUvbgÁúMGamInenAelIkae)an α esÞIrEtRKb;GasIutGamIneNepSgeTot . va (glutamate) RtUv)ansMeyaKBI NH4

+ nig α-ketoglutarate enAkñúgRbtikmμEdlsMrYleday glutamate

dehydrogenase . karenHKWCakarRtLb;eTAvijénRbtikmμEdl)anbgðajenAkñúgrUbTI 22-12 nig kar BiB’NnaBImunmk . enAkñúgkrNIenH Coenzyme EdleRbICaPñak;garer:dukm μKW NADPH enAkñúgRbtikmμ .

NH4+ + α-ketoglutarate + NADPH ⇄ L-glutamate + NADP+ + H2O

Glutamine, proline nig Arginine RtUv)ansMeyaKBI glutamate . karBinitürUbTI 22-14 bgðajfa Serine, glycine nig cysteine RtUv)ansMeyaKBI 3-phospho-

glycerate . Alanine RtUv)ansMeyaKBI Pyruvate nig Tyrosine RtUv)anplitBI Phosphoenol-

pyruvate nig sárEdlmanbYnGatUmkae)an erythrose-4-phosphate EdlCabnþRtUv)ansMeyaKBI glucose-6-phosphate enAkñúgpøÚv Pentose phosphate . epSgBI Precursor CaGasIutGamIneN glycolysis nig Citric acid cycle k¾pþl;pgEdrnUv Precursors sMrab;lIBId nig Nitrogenous bases

EdlRtUvkarcaM)ac;edIm,IeFIV DNA EdlCam:UelKulmanpÞúkB½t’manénEhSn . BYkvak¾begáItpgEdrnUv



Precursors sMrab; Heme EdlCa prosthetic group EdlRtUvkarsMrab; hemoglobin, myoglobin nig cytochromes . edayc,as;las; Rbtikmμ glycolysis nig Citric acid cycle sßitenAkNþalrvag skmμPaB anabolism nig Catabolism rbs;ekasika . pøÚvemtabUlIsEdlbMeBjmuxgarTaMg anabolism nig Catabolism RtUv)aneKehAfa Amphibolic pathway . cUrBicarNarcMeBaHxN³mYy PaBxusKñaEdl FmμCatiénpøÚvBIr (dual nature of pathways) GacmancMeBaHekasikamYy . enAeBlekasikamYykMBug lUtlas;edayskmμ vamankarTamTarya:gxøaMgnUv precursors CIvsMeyaKedIm,Isagsg;rcnasm<½n§ ekasikafμ I . karTamTarGasIutGamIneNGacdkya:geRcInnUv oxaloacetate nig α-ketoglutarate ecjBI Citric acid cycle . bBaðaEdllM)akEdlekIteLIgKWfary³eBlénkarlUtlas;skm μrbs;ekasika k¾TamTarnUvcMnYnd¾eRcInénfamBlEdr b:uEnþkar)at;bg;nUv oxaloacetate rbs;mItUkugRDInwgkat;bnßy nUv smtßPaBkñúgkarplit ATP . karedaHRsaycMeBaHbBaðaenHKWedIm,ImanpøÚvepSgeTotsMrab; karsMeyaK Oxaloacetate EdlGac plit oxaloacetate )anRKb;RKankñúgkarpÁt;pÁg;esckþIRtUvkarén Anabolism nig Catabolism rbs; ekasika . eTaHbICa)ak;etrI nig rukçCatimanynþkarCaeRcIn b:uEnþpøÚvEtmYyEdlekasikafnikstVGacplit oxaloacetate )aneRcInKWedaysar carboxylation én pyruvate EdlCaRbtikmμsMxan;EdrenAkñúg gluconeogenesis . Rbtikm μenHKW ³ Pyruvate + CO2 + ATP → Oxaloacetate + ADP + P1

Gg;sIumEdlsMrYlRbtikmμenHKW Pyruvate carboxylase . vaKWCa Conjugated protein

EdlmanCabgÁúM prosthetic cgP¢ab;edaysm<½n§kUva:Lg;KWCa vitamin Biotin . Gg;sIumenHvadMeNIrkarenA eBlmankMrit Acetyl CoA x<s; . kMhab;x<s;én acetyl CoA KWCasBaØaEdlekasikaRtUvkarkMritx<s;én FatukNþalén Citric acid cycle CaBiess oxaloacetate EdlCa substrate edImdMbUgeK . RbtikmμEdlsMrYleday Pyruvate carboxylase RtUv)aneKehAfaCa anaplerotic reaction . Baküfa "Anaplerotic" mann½yfabMeBj[RKb; . dUecñH Gg;sIumsMxan;enHRtUvEtbMeBjbEnßm oxaloacetate Canic© nig edaypÞal;nUvRKb;FatukNþalrbs; citric acid cycle EdlRtUv)andkecjCa precursors CIvsMeyaKsMrab;RbtikmμEdlsegçbenAkñúgrUbTI 22-14 .

Fin



rUbTI 22-14

Glycolysis nig Citric acid cycle k¾pþl;nUv Precursors epSg²sMrab;CIvsMeyaK én GasIutGamIneN Nitrogenous bases nig Porphyrins

Glucose

G-6-P

Pyruvate

Triose-P

Phosphoenolpyruvate

3-Phosphoglycerate

Acetyl CoA

Citrate Oxaloacetate

Isocitrate

α-ketoglutarate

Succinyl CoA

Succinate

Glutamate

Glutamine Proline

Arginine

Nucleoside

Ribose-P Erythroe-4-P

Chrorismate

Tyrosine Serine

Cysteine Glycine

Purines

CO2 Lipids

Glycerol

Lipids

Aspartate

Pyrimidines

Asparagine

Porphyrines

Heme

CO2 CO2

emtabUlIsGasIuRka (FATTY ACID METABOLISM)

vtßúbMNgemeron 1- segçbBIkarrMlay nig karsþúkTuklIBId . 2- BiB’NnaBIkarbMEbkénGasIuRkaeday β-Oxidation . 3- Bnül;BItYnaTIrbs; Acetyl CoA enAkñúgemtabUlIsrbs;GasIutRka . 4- yl;GMBItYnaTIénkarplit keto body enAkñúg β-oxidation . 5- eRbobeFob β-oxidation rbs;GasIutRka nig CIvsMeyaKrbs;GasIutRka . 6- BiB’NnaBIkarEktMrUvénemtabUlIsrbs;lIBId nig Carbohydrate enAkñúgkarTak;TgeTAnwgeføIm Calikaxøaj; Calikasac;dMu nig xYrk,al . 7- segçbBI\T§iBlRbqaMgKñarbs; glucagon nig Insulin . KMeragemeron 1- emtabUlIslIBIdcMeBaHstV - karrMlay nig karRsUbén Triglyceride enAkñúgcMNIGahar - karsþúkTukénlIBId 2- karbMEbkGasIutRka - TidæPaBTUeTAénkarbMEbkGasIutRka - Rbtikm μén β-oxidation 3- Ketone bodies

- Ketosis

- Ketogenesis

4- karsMeyaKGasIutRka - kareRbobeFobkarsMeyaKGasIutRka nig karbMEbk 5- karEktMrUvénemtabUlIslIBId nig Carbohydrate

- eføIm Calikaxøaj; Calikasac;dMu xYrk,al 6- \T§iBlén Insulin nig Glucagon nig emtabUlIsekasika

emtabUlIsGasIuRka (FATTY ACID METABOLISM)

eskþIepþIm emtabUlIsrbs;GasIutRka nig lIBIdvivtþn_CMuvijvasnarbs; Acetyl CoA . eyIg)aneXIjenAkñúg CMBUkTI 22 fa enAeRkamlkçx½NÐmanxül; Pyruvate RtUv)anbMElgeTACa Acetyl CoA EdlRtUv)anpþl; [eTAkñúg Citric Acid Cycle . GasIutRkak¾RtUv)anbMEbkeTACa Acetyl CoA Edr nig TTYlrgGuksIutkmμ eday Citric Acid Cylcle dUcCaGasIutGamIneNmYycMnYn . müa:gvijeTot Acetyl CoA xøÜnÉgenHKWCa sarFatucab;epþImsMrab;CIvsMeyaKénGasIutRka Bak;kNþalGasIutGamIneN cholesterol nig steroid

hormones . ehtudUecñH Acetyl CoA CaemtabUlItsMxan;mYykñúgcMeNamemtabUlItsMxan;énemtabUlIs kNþal . 1- emtabUlIslIBIdcMeBaHstV (LIPID METABOLISM IN ANIMALS)

1- karrMlay nig karRsUbén Triglyceride EdlmanenAkñúgcMNIGahar Triglycerides KWCasarFatuEdlmincUlcitþTwkxøaMg . GaRs½yedaykarminrlaykñúgTwkenH BYk vaRtUvEt)anEkécñmuneBlBYkvaGacRtUv)anrMlay RsUb nig TTYlrgemtabUlIs . GaRs½yedaykarEk écñénlIBIdrbs;GaharekIteLIgenAkñúgeBaHevontUc Lipases EdlrlaykñúgTwk Gg;sIumGIuRdUlIs

Triglycerides EdleKeXIjmanvaenAkñúgRkBH nig kñúgTwkmat;minmanRbsiT§PaBxøaMg . karBit xøaj;enA kñúgcMNIGaharPaKeRcIn)anmkdl; duodenum EdlCaEpñkTImYyéneBaHevontUc eRkamTMrg;Ca eKalika xøaj; (fat globule) . eKalikaxøaj;TaMgenHCMrujkarbeBa©jkarbeBa©jTwkRbma:t;BIfg;TwkRbma:t; . Twk-Rbma:t; (Bile) pSMeLIgBIbMNþúM (micelles) én lecithines, cholesterol, protein, bile salt, inorganic

ions nig bile pigments . Micelles ¬rUbTI 23-1¦ KWCabNþúMkþaMgCab;Kñaénm:UelKulEdlmantMbn;man b:Ul nig tMbnlminmanb:Ul . cugminmanb:Ulrbs; bile salt manninñakareTArkkarcgP¢ab;KñaenAeBldak; vaenAkñúgTwk . tMbn;EdlcUlcitþTwkénm:UelKulTaMgenHmanRbtikm μCamYynwgTwk . Bile salt RtUv)aneFIV enAkñúgeføIm nig RtUv)ansþúkTukenAkñúgfg;TwkRbma:t; EdlrgcaMkareP¶acedIm,IbeBa©jTwkRbma:t;eTAkñúg duodenum . Bile salts EdlsMxan;cMeBaHmnusSKW cholate nig chenodeoxycholate ¬rUbTI 23-2¦. Cholesterol esÞIrEtminrlayTaMgRsugenAkñúgTwk b:uEnþkarbMElgén cholesterol eTACa bile

salts begáItnUv detergents Edlk,almanb:Ulrbs;vaeFIV[BYkvarlayenAkñúgEpñkTwkrbs;sIutU)aøs; nig knÞúyminrlaykñúgTwkP¢ab;eTAnwg Triglycerides . bnÞab;karbriePaKGahar TwkRbma:t;hUtambMBg;Twk-Rbma:t;rYmcUleTAkñúg duodenum EdlenATIenaH bile salts RtUvlayCamYynwgeKalikaxøaj;[eTACa

dMNk;tUc² . karenHbegáInnUvépÞrbs;m:UelKullIBId EdlGnuBaØatiBYkvaTTYlnUvkarGIuRdUlIsedayRsYl edaysar lipases ¬rUbTI 23-3¦ .

rUbTI 22-1 rcnasm<½n§rbs; Micelles

rUbTI 22-2 rcnasm<½n§rbs; Bile acids

rUbTI 23-3 dMNak;karénkarbMEbklIBIdenAkñúgeBaHevon

PaKeRcInénlIBIdenAkñúgdMNk;xøaj;enHsßitenAeRkamTMrg;Ca Triglycerides b¤ Triacyl-

lycerides EdlKWCa fatty acid esters of glycerol . RbUetGIunEdleKehAfa Colipase cgP¢ab;eTAnwg épÞrbs;dMNk;lIBId nig CYy pancreatic lipases kñúgkarP¢ab;eTAnwgépÞ nig GIuRdUlIssm<½n§ ester rvag glyceol nig GasIutRkarbs; triglyceride rUbTI 23-4 . enAkñúgdMeNIrkarenH GasIutRkaBIrkñúgcMeNam GasIutRkaTaMgbIRtUv)anbeBa©j nig monoglycerides nig GasIutRkaEdlplit)anlayKñaedayesrI CamYynwg micelles rbs;TwkRbma:t; . Micelles TaMgenHRsUbcUltamekasika epithelial Pñasrbs; eBaHevontUc ¬rUbTI 23-5¦ .

rUbTI 22-4 GMeBIrbs; Pancreatic lipase kñúgkarGIuRdUlIslIBIdenAkñúgcMNIGahar

rUbTI 22-5 karqøgkat;én Triglycerides enAkñúg Micelles cUleTAkñúgekasika epithelium eBaHevontUc

edayKYr[P¶ak;ep¥Il Monoglycerides nig GasIutRkabnÞab;mkk¾rYmKñabegáItCa Triglycerides

vij EdlRtUv)anrYmCamYyRbUetGIunedIm,IbegáItCafñak;én lipoproteinenAkñúg)aøs μaEdleKehAfa Chylomicron ¬rUbTI 23-5¦ . karRbmUlénlIBId nigRbUetGIunTaMgenHRtUv)anbeBa©jeTAkñúgsés lymph tUc² nig cugbBa©b;BYkvaeTAdl;crnþQam . enAkñúgcrnþQam triglycerides RtUv)anGIuRdUlIssar Cafμ IedIm,IbegáItCa glycerol nig GasIutRkaEdlbnÞab;mkRtUv)anRsUbedayekasika . RbsinebIdgxøÜn RtUvkarfamBl m:UelKulTaMgenHRtUv)anbMEbkedIm,IbegáIt ATP . RbsinebIsBa’gÁkay minRtUvkar famBl m:UelKulEdlsMbUrfamBlenHRtUv)ansþúkTuk . 2- karsþúkTuklIBId (Lipid storage)

GasIutRkaRtUv)ansþúkTukeRkamTMrg;Ca Triglycerides . PaKeRcInénm:UelKul Triglycerides

rbs;sBa’gÁkayRtUv)ansþúkTukeRkamTMrg;CadMNk;xøaj;enAkñúgsIutU)aøs;rbs; Adipocytes (fat cell) Edl begáIt)anCa Adipose tissue . Adipocyte nimYy²mandMNk;xøaj;FMEdlbegáIt)anesIÞrEtmaDTaMgmUl

rbs;ekasika . ekasikaepSgeTot dUcCaekasikasac;dMuebHdUgmandMNk;xøaj;tUc²tictYc . enAkñúg ekasikaTaMgenH dMNk;xøaj;RtUv)anrMuB½T§eTAedaymItUkugRDI . enAeBlekasikaRtUvkarfamBl Triglycerides RtUv)anGIuRdUlIsedIm,IbeBa©jCaGasIutRkaEdlRtUvdwknaMcUleTAkñúglMh matrix rbs;mItU-kugRDI . enATIenaHGasIutRkaRtUv)anTTYlrgGuksIutkm μTaMgRsug nig ATP RtUv)anpliteLIg . GasIutRkaEdlpþl;edaykarGIuRdUlIsén triglycerides KWCaRbPBsMbUrfamBlrbs;sBa’gÁkay . GuksIutkmμTaMgRsugrbs;GasIutRkabeBa©jfamBleRcInCagGuksIutkmμcMnYndUcKñaén glycogens . 2- karbMEbkGasIutRka (FATTY ACID DEGRADATION)

1- TidæPaBTUeTAénkarbMEbkGasIutRka enAedImstvtSTI 20 karBiesaFn_ya:gl¥RtUv)aneKeFIVedIm,IkMNt;faetIGasIuRkaRtUv)anbMEbk tamrebobdUcemþc . rMlwkBICMBUkTI 10 EdlFatuviTüúskm μ (radioactive elements) GacRtUv)anP¢ab;eTA nwgm:UelKulCIvsaRsþ nig eFIV[manBaseBjsBa’gÁkay . KImIviTUCnCatiGalWøm:g; Franz Knoop begáIt nUvRbePTdUcKñaénkarBiesaFn_EdlkarbMBak;pøak;ya:gyUrmuneBlsarFatuviTüúskm μ (racdioactive

tracers) man . Knoop )anpþl;nUvGasIutRka[EqáEdlCaTUeTAbgÁúM methyl enAcugExSmanbgÁúM phenyl

P¢ab;eTAnwgva . m:UelKulrebobenHRtUv)aneKehAfaCa ω-labeled (omega) fatty acid ¬rUbTI 23-6¦. enAeBlEdlKat;dkyk (isolate) GasIutRkaEdlTTYlrgemtabUlIsrYcBITwkenamrbs;Eqá Kat;)aneXIj fa phenyl acetate RtUv)anbegáItenAeBlGasIutRkamancMnYnGatUmkae)anKt;KUr . b:uEnþ Benzoate RtUv)anbegáIteLIgenAeBlGasIutRkamancMnYnGatUmkae)aness . Knoop bkRsayTinñn½y cMeBaHmeFüa)ayEdlfakarbMEbkGasIutRkaekIteLIgedaykardknUvbgÁúM Acetate EdlmanBIGatUm-kae)anBIxagcug carboxyl rbs;GasIutRka . \LÚvenHeyIgdwgfa EpñkEdlmanBIGatUmkae)anEdlRtUv )anplitedaykarbMEbkénGasIutRkaminEmnCa acetate b:uEnþCa Acetyl CoA . pøÚvsMrab;bMEbkGasIut-Rka[eTACa Acetyl CoA RtUv)aneKehAfaCa β-Oxidation .

rUbTI 23-6

2 CH3—COO-

CH2—CH2—CH2—CH2—CH2—COO-

CH2—COO-

+

CH2—CH2—CH2—CH2—CH2—CH2—COO-

3 CH3—COO-

COO-

+

ω-Phenyl-labeled fatty acid with an even number of carbon atoms

ω-Phenyl-labeled fatty acid with an odd number of carbon atoms

β-oxidation cylcle ¬dMNak;karTI 2-5 rUbTI 23-7¦ manmYysMrab;Edlman 4 Rbtikm μEdl TMrg;TaMgRsugrbs;vadUceTAnwgRbtikm μ 4 cugeRkayrbs; Citric Acid Cycle . dMeNIrnimYy²kñúgmYyQut énRbtikmμbeBa©j Acetyl CoA nig RtLb;m:UelKul fatty acyl CoA EdlmanGatUmkae)anBIrticCagmun cUleTAkñúg cycle vij . mYym:UelKulrbs; FADH2 smmUleTAnwgBIrm:UelKul ATP nig mYym:UelKul rbs; NADH smmUleTAnwgbIm:UelKul ATP RtUv)anbegáItkñúgvdþnimYy²én β-oxidation .

rUbTI 23-7 vdþénRbtikmμ β-oxidation rbs;GasIutRka

3- Rbtikmμ β-oxidations

Gg;sIumEdlsMrYl β-oxidation rbs;GasIutRkasßitenAkñúglMh matrix rbs;mItUkugRDI . ynþkar dwknaMBiessRtUvkaredIm,InaMm:UelKulGasIutRkacUleTAkñúg matrix rbs;mItUkugRDI . enAEpñkxagkñúg

Fatty acid—CH2—CH2—COO-

Fatty acid—CH2—CH2—CO~S—CoA

Fatty acid—CH═CH—CO~S—CoA

Fatty acid—CH(OH)—CH2—CO~S—CoA

Fatty acid—CO—CH2—CO~S—CoA

Fatty acid—CO~S—CoA +

Acetyl CoA

CH3—CO~S—CoA

Citric Acid Cycle →→ 12 ATP

1

ATP

AMP + PPi

CoA

2 FAD

FADH2 →→→ 2 ATP

3 H2O

4 NAD+

NADH →→→ 3 ATP

5 CoA

GasIutRkaRtUv)anbMEbkedayRbtikm μ β-oxidation . dUcEdleyIg)aneXIj Rbtikm μTaMgenHmanGMeBI eTAvijeTAmkCamYy Oxidative phosphorylation nig Citric Acid Cycle edIm,IbegáIt ATP . RbtikmμTI 1- dMNak;karTI 1 KWCa activation reaction EdlbNþal[mankarpliténm:Uel-Kul Fatty acyl CoA . sm<½n§ Thioester RtUv)anbegáItrvag Coenzyme A nig Fatty acid ³

RbtikmμenHRtUvkarfamBleRkamTMrg;Ca ATP EdlRtUv)ankat;pþac;Ca AMP nig Pyropho-

sphate . karenHCab;Tak;TgeTAnwgkareFIVGIuRdUlIsénsm<½n§ Phosphoanhydride BIr . enATIenHsarCafμI mþgeTot eyIgeXIjesckþIRtUvkarkñúgkarvinieyaKnUvcMnYntictYcénfamBl dUecñHcMnYnfamBld¾eRcInGac RtUv)anRbmUlenAeBleRkayenAkñúgpøÚvenH . Coenzyme A k¾RtUvkarsMrab;dMNak;karenHEdr . plit-pl (Fatty acyl CoA) mansm<½n§ Thioester EdlmanfamBlx<s;rvag Fatty acid nig Coenzyme A . Acyl-CoA ligase EdlsMrYlRbtikm μenH sßitenAPñasxageRkArbs;mItUkugRDI . ynþkarEdlnaM Fatty

acyl CoA cUleTAkñúg matrix énmItUkugRDICab;Tak;TgeTAnwgm:UelKuldwknaMmYyEdleKehAfa Carnitine. dMNak;kardMbUg EdlsMrYledayGg;sIum Carnitine acyltransferase I KWCakarbBa¢Únén bgÁúM fatty acyl eTA[ carnitine edaybegáIt)anCa acylcarnitine nig Coenzyme A . RbUetGIunCaGñk-dwknaMbnÞab;sßitenAPñasxagkñúgrbs;mItUkugRDIbBa¢Ún acylcarnitine cUleTAkñúg matrix rbs;mItUkugRDI EdlenATIenaH Carnitine acyltransferase II sMrYlkarbegáIteLIgvij (regeneration) én Fatty acyl

CoA . RbtikmμTI 2- RbtikmμTI 2KWCaRbtikmμGusIutkmμEdldkecjnUvGatUmGIuRdUEsnmYyKUrBI Fatty

acid . vaTaMgBIenHRtUv)aneRbIedIm,IeFIVer:dukmμeTAelI FAD edIm,IbegáIt)anCa FADH2 . Rbtikmμ Dehydrogenation enHRtUv)ansMrYleday Gg;sIum Acyl-CoA dehydrogenase nig bNþal[man karbegáItsm<½n§BIrCan;rvagGatUmkae)anBIr ³

Oxidative phosphorylation naM[)anBIrm:UelKul ATP sMrab;m:UelKul FADH2 Edlpliteday RbtikmμGuksIutkmμ-er:dukmμ .

CH3—(CH2)n—CH2—CH2—COOH Fatty acid

CH3—(CH2)n—CH2—CH2—CO~S—CoA Fatty acyl CoA

ATP AMP + PPi

Coenzyme A

CH3—(CH2)n—CH═CH—CO~S—CoA

CH3—(CH2)n—CH2—CH2—CO~S—CoA Fatty acyl CoA

FAD FADH2

RbtikmμTI 3- RbtikmμTI 3 Cab;Tak;Tg karbEnßmTwk (hydration) eTAelIsm<½n§BIrCan;EdlekIt eLIgenAkñúgRbtikm μTI 2 . CalT§pl β-carbon RtUv)anP¢ab;eTAedaybgÁúM OH . Rbtikm μenHRtUv)an sMrYledayGg;sIum enol-CoA hydrase .

RbtikmμTI 4 - enAkñúgRbtikm μ GuksIutkmμ enHbgÁúM OH rbs; β-carbon RtUv)anTTYlrgnUvkardk GIuRdUEsnecj (dehydrogenated) ehIy NAD+ RtUv)anTTYlrger:dukmμeTACa NADH EdlCacugbBa©b; RtUv)aneRbIsMrab;begáItm:UelKul ATP eday Oxidative phosphorylation ehIy L-β-Hydroxyacyl-

CoA dehydrogenase sMrYlRbtikmμenH .

RbtikmμTI 5 - RbtikmμcugbBa©b; RtUv)ansMrYleday Thiolase KWCakarkat; EdlbeBa©jnUv Acetyl CoA . karenHrYbeTAeday Thiolysis karb:HTgÁicénm:UelKul Coenzyme A eTAelI β-

carbon. lT§plKWCakarecjén Acetyl CoA nig acyl CoA EdlmanGatUmkae)anBIrticCag GasIut-RkaBIedImdMbUg ³

Fatty acyl CoA EdlRtUv)ankat;[xøIRtUv)anTTYlrgGuksIutkmμCabnþeTotedaykarvilCMutam ry³RbtikmμTI 2 dl;RbtikmμTI 5 rhUtdl;ExSkae)anrbs;GasIutRkaRtUv)anbMEbkTaMgRsugeTACa Acetyl

CoA . Acetyl CoA Edlpliteday β-oxidation énGasIutRka bnÞab;mkcUleTAkñúgRbtikm μén Citric

Acid Cycle . dUecñH karenHCacugbBa©b;bNþal[mankarplit 12 ATP kñúgmYym:UelKul Acetyl CoA EdlbeBa©jenAeBlman β-oxidation . dUcCa]TahrN_énfamBlekIteLIgBI β-oxidation karbBa¢ak;BItulüPaB (balance sheet)

cMeBaH Palmitic acid Edlman 16 GatUmkae)anenAeBlvaRtUv)anbMEbk eday β-oxidation RtUv)an segçbenAkñúgrUbTI 23-8 . kareFIVGuksIutkm μTaMgRsugrbs; Palmitic acid bNþal[mankarplit ATP

cMnYn 129 m:UelKul 3 dgknøHeRcInCaglT§pl)anBIGuksIutkmμTaMgRsugrbs; Glucose EdlmancMnYn smmUl .

CH3—(CH2)n—CH═CH—CO~S—CoA

CH3—(CH2)n—CH(OH)—CH2—CO~S—CoA

H2O

CH3—(CH2)n—CH(OH)—CH—CO~S—CoA CH3—(CH2)n—CO—CH2—CO~S—CoA

NAD+ NADH

CH3—(CH2)n-2—CH2—CH2—CO~S—CoA + H3—CO~S—CoA

CH3—(CH2)n—CO—CH2—CO~S—CoA

CoA

rUbTI 23-8 GuksIutkm μTaMgRsugén Palmitic Acid

Ci

3- KETONE BODIES

cMeBaH Acetyl CoA Edlpliteday β-oxidation rbs;GasIutRkaedIm,IcUleTAkñúg Citric acid

cycle edayRbsiT§PaB vaRtUvEtmankarpþl;edayRKb;RKan;én Oxaloacetate . RbsinebI glycolysis nig β-oxidation ekIteLIgkñúgGRtadUcKña vanwgmankarpþl;edaysßitesßrnUv Pyruvate ¬mkBI glycolysis¦ EdlGacRtUv)anbMElgCa Oxaloacetate . b:uEnþGIVEdlekIteLIg RbsinebIkarpþl;én Oxaloacetate

TabxøaMgeBkkñúgkarGnuBaØati[RKb; Acetyl CoA cUleTAkñúg Citric Acid Cycle ? sßienAeRkamlkçx½NÐ enH Acetyl CoA RtUv)anbMElgeTACasarFatuEdleKehAedaRcLMfaCa Ketone bodies : β-

hydroxybutyrate, acetone nig Acetoacetate emIlrUbTI 23-9 .

CH3—(CH2)14—COOH

Palmitic acid

CH3—(CH2)14—CO~S—CoA

Palmityl CoA

ATP

AMP + PPi

Coenzyme A

8 CH3—CO~S—CoA + 7 FADH2 + 7 NADH Acetyl CoA

14 ATP 21 ATP

Citric Acid

Cycle

72 ATP

16 ATP

8 x 3 NADH

8 x 1 FADH2 8 x 1 GPT

8 x 1 ATP

Two high-energy phosphate bonds input 2-ATP

Net ATP Production 129 ATP

1- Ketosis

Ketosis ¬kMritx<s;minFmμtaenAkñúgQamrbs; ketone bodies¦ KWCasßanPaBEdlekIeLIgeRkam lkçx½NÐCMgWmYycMnYn dUcCaPaBTurÖikS (starvation) GaharEdlxVHBYk carbohydrate ¬kñúgxN³EdleRbI rbbGaharravEdlsMbUrRbUetGIun¦ b¤ CMgWTwkenamEp¥mEdlminGacRtYtBinitü . kareRbIR)as;én carbohydrate rbs;GñkCMgWTwkenamEp¥mFm μta b:uEnþ carbohydrate minGaccUleTAkñúgekasikaedIm,IeRbI Ca\n§³ . ehtudUecñH GñkCMgWTwkenamEp¥mCYbnwgPaBTurÖikSkñúgkNþalénPaBsMbUr (the starvation in the

midst of plenty) . cMeBaHGñkCMgWTwkenamEp¥m kMhab;x<s;xøaMgén ketone acids enAkñúgQambNþal[ man Ketoacidosis . Ketone acids KWCaGasIutxøaMgKYrsm nig ehtudUecñHvaRtUv)anbMEbkedaybeBa©j nUv H+ . sßitenAeRkamlkçx½NÐTaMgenH pH rbs;QamkøayeTACaGasIut EdlGacbNþal[søab; .

rUbTI 23-10 RbtikmμEdlCab;Tak;TgenAkñúg Ketogenesis

2 CH3—CO~S—CoA 2 Aacetyl CoA

CH3—CO—CH2—CO~S—CoA Aacetoacteyl CoA

OH

│ -OOC—CH2—C—CO~S—CoA β-Hydroxy-β-methylglutaryl CoA │ CH3

-OOC—CH2—CO—CH3 Acetoacetate

CH3—CO—CH3 Acetone

-OOC—CH2—CH(OH)—CH3 β-hydroxybutyrate

CoA

Acetyl CoA + H2O

CoA

AcetylCoA H+

CO2

NAD+

NADH + H

2- Ketogenesis

pøÚvsMrab;karplitén Ketone bodies ¬rUbTI 23-10¦ cabepþImedaykarRtLb;eTAvijén dMNak;karcugeRkayrbs; β-oxidation . enAeBlkMritrbs; Oxaloacetate Tab Gg;sIumEdlCaFm μta GnuvtþeTAelIRbtikmμcugeRkaybMputén β-oxidation \LÚvsMrYlkarrlaycUlKñaén Acetyl CoA

BIrm:UelKul edIm,IbegáItCa acetoacetyl CoA .

Acetyl CoA ⇄ Acetoacetyl CoA

Acetoacetyl CoA GacmanRbtikmμCamYym:UelKul Acetyl CoA TIbIedIm,IbegáIt)anCa β-

hydroxy-β-methylglutaryl CoA (HMG-CoA) .

Acetoacetyl CoA + Acetyl CoA ⇄ β-Hydroxy-β-methylglutaryl CoA

RbsinebI HMG-CoA RtUv)anbegáItenAkñúgsIutU)aøs; vanwgeRbICa precursor sMrab;CIvsMeyaK Cholesterol . b:uEnþ Ketogenesis ¬dUc β-oxidation¦ ekIteLIgenAkñúg matrix rbs;mItUkugRDI nig enA TIenH HMG-CoA RtUv)ankat;edIm,IbegáItCa Acetoacetate nig Acetyl CoA ³

β-Hydroxy-β-methylglutaryl CoA ⇄ Acetoacetate + Acetyl CoA

cMeBaHcMnYnticxøaMg Acetoacetate edayÉkÉg)at; CO2 edayÉkÉgedIm,Ipþl;Ca Acetone . karenHKWCaRbtikmμEdlbgá[man kardkdegðImmanGaestUn EdlCajwkjab;ekItmanCamYynwgCMgWTwk enamEp¥mEdlminGacRtYtBinitü)an . edayjwkjab;CagenHeTAeTot vaTTYlrgnUvRbtikm μer:dukmμEdl Cab;Tak;TgeTAnwg NADH edIm,IbegáItCa β-hydroxbutyrate Acetoacetate + H+ → Acetone + CO2 Acetoacetate + NADH → β-hdroxybutyrate + NAD+

Acetoacetate nig β-hydroxybutyrate RtUv)anplitCabzmenAkñúgeføIm . emtabUlItTaMgenH sayPayeTAkñúgQam nig RtUv)aneFIVcracreTACalikaepSgeTotEdenATIenaHBYkvaGacRtUv)anbMElg[ eTACa Acetyl CoA sarCaf μ I nig RtUveRbIedIm,IbegáItCa ATP . CakarBit sac;dMuebHdUgTTYlnUvfamBl emtabUlIsPaKeRcInBIGuksIutkm μrbs; Ketone bodies minEmnmkBIGuksIutkmμrbs; glucose eT . CalikaepSgeTotEdlsMrbsMrYl)anl¥bMputeTAnwgkareRbIén glucose nwgCab;Tak;TgedayekIneLIgeTA nwg ketone bodies sMrab;begáItfamBlenAeBl glucose minGacrk)an b¤ mancMnYnkMNt; . karenH KWCaBitCaBiessmYyrbs;xYrk,al .

4- karsMeyaKGasIutRka (FATTY ACID SYNTHESIS)

RKb;dMeNIrkarénsrIragÁ lT§PaBkñúgkarsMeyaKGasIutRka . cMeBaHmnusS karelIsén Acetyl

CoA RtUv)anplitedaykarbMEbk Carbohydrate RtUv)aneRbIedIm,IeFIVGasIutRkaEdlbnÞab;mkRtUv)an sþúkTukeRkamTMrg;Ca Triglycerides . 1- kareRbobeFobénkarsMeyaK nig karbMEbkGasIutRka enAelIkarBinitüelIkTImYy karsMeyaKGasIutRkahak;dUcCakarRtLb;edaysamBaØén β-

oxidation . edaymanlkçN³Biess ExSGasIutRkaRtUv)anksageLIgedaykarbEnßmQuténbgÁúM Acetyl

EdmanBIrGatUmkae)an ¬rUbTI 23-11¦ . eTaHbICaKImIviTüaénkarsMeyaK nig karbMEbkGasIutRkadUc Kñak¾eday b:uEnþvamanPaBxusKñasMxan;²CaeRcInrvag β-oxidation nig CIvsMeyaKénGasIutRka . karenH RtUv)ansegçbdUcxageRkam ³ TItaMgenAkñúgekasika (Intracellular location) Gg;sIumTTYlxusRtUvcMeBaHCIvsMeyaKGasIutRka sßitenAkñúgsIutU)aøs;rbs;ekasika rIÉGg;sIumTTYlxusRtUvcMeBaHkarbMEbkGasIutRkasßitenAkñúgmItUkugRDI. GñkdwknaMbgÁúM Acyl (Acyl group carriers) FatukNþalénEdlRtUv)aneFIV[skmμénCIvsMeyaK RtUv)ancgP¢ab;eTAnwgm:UelKuldwknaM (a carrier molecule) EdleKehAfa Acyl carrier protein

(ACP) rUbTI 23-12 . . enAkñúg β-oxidation GñkdwknaMbgÁúM acyl KWCa Coenzyme A . b:uEnþvamanPaB sMxan;dUcKñarvagGñkdwknaMTaMgBIrenH . vaTaMgBIrmanbgÁúM Phosphopantetheine EdlekIteLIgBI vItamIn pantothenic acid nig bgÁúM β-mercaptoethylamine . enAkñúgkrNITaMgBIr bgÁúM Acyl

énGasIutRkaRtUv)ancgP¢ab;edaysarsm<½n§ Thioester eTAnwgbgÁúM phosphopantetheine . Gg;sIumEdlCab;Tak;Tg (Enzyme involved) karsMeyaKGasIutRkaRtUv)anGnuvtþedaybNþúM Gg;sIumcMruHEdlRtUv)aneKsÁal;faCa Fatty acid synthase . Gg;sIumEdlTTYlxusRtUvcMeBaHkarbMEbk GasIutRkaminrYmCamYyKñatamrebobenHeT . GñkdwknaMeGLikRtug ( Electron carriers) NADH nig FADH2 RtUv)anplitedayGuksIutkmμ GasIutRka rIÉ NADPH KWCaPñak;garer:dukmμcMeBaHCIvsMeyaKGasIutRka . Cac,ab;TUeTA NADH RtUv)an plitedayRbtikm μkatabUlik (catabolic reaction) nig NADPH KWCaPñak;garer:dukm μénRbtikmμCIv sMeyaK . Coenzymes TaMgBIrenHxusKñaedaysarEtbgÁúM Phosphate EdlcgP¢ab;eTAnwg Ribose én NADPH rUbTI 23-13 . b:uEnþ Gg;sIumEdleRbI Coenzyme TaMgenHGacgayRsYlEbkecjBIQelI eKalkarN_enH .

rUbTI 23-11 segçbkarsMeyaKGasIutRka

rUbTI 23-12 rcnasm<½n§én Phosphopantetheine

rUbTI 23-13 rcnasm<½n§rbs; NADPH

H3C—CO~S—ACP + -OOC—CH2—CO~S—ACP

H3C—CO—CH2—CO~S—ACP

H3C—CHOH—CH2—CO~S—ACP

H3C—CH═CH—CO~S—ACP

H3C—CH2—CH2—CO~S—ACP

ACP +CO2 Condensation

NADPH

NADP+

H2O

Reduction

Dehydration

ReductionNADPH

NADP+

Malonyl ACPAcetyl ACP

Acetoacetyl ACP

β-hydroxybutyryl ACP

Crotonyl ACP

Butyryl ACP

5- karEktMrUvénemtabUlIsrbs;lIBId nig KøúysIut (The Regulation of Lipid and Carbohydrate Metabolism)

emtabUlIsrbs;GasIutRka nig KøúysIutekIeLIgkñúgTMhMxusKñaenAkñúgsrIr³(organs) xusKña . dUc EdleyIgnwg)aneXIjenAkñúgEpñkenH karEktMrUvénénTidæPaBrbs;emtabUlIsEdlCab;Tak;TgTaMgBIrenH mansarsMxan;xagsrIrviTüaya:gxøaMg . 1- eføIm eføImpþl;nUvkarpþl;edaysßitesßrnUv glucose sMrab;sac;dMu nig xYrk,al nig edIrtYnaTIsMxan;enAkñúg karEktMrUvnUvkMhab; glucose enAkñúgQam . karEktMrUvenHsßitenAeRkamkarRtYtBinitürbs;G½rm:Un . rMlwkeLIgvijfaG½rm:Un Insulin bgá[ glucose enAkñúgQamRtUv)ancab;ykedaysareføIm nig sþúk TukCa glycogen (glycogenesis) . enAkñúgpøÚvenH eføImkat;bnßynUvkMrit glucose kñúgQam enAeBlEdl vaekIneLIgx<s;eBk . müa:gvijeTot G½rm:Un glucagon eP¶ackarbMEbk glycogen nig beBa©j glucose cUleTAkñúg crnþQam . Lactate Edlplitedaysac;dMueRkamlkçx½NÐK μanxül;k¾RtUv)ancab;ykedaysarekasika eføIm nig RtUv)anbMElgeTACa glucose eday gluconeogenesis . TaMgkarbMEbk glycogen

(glycogenolysis) nig Gluconeogenesis KWCapøÚvEdlplit glucose sMrab;karnaMecjeTAsrIr³epSg eTotenAeBlRtUvkarfamBl ¬rUbTI 23-14¦ . eføImk¾edIrtYnaTIkNþalénemtabUlIslIBId . enAeBlmankarelIsnUv\n§³ eføImsMeyaK GasIuRka. karenHRtUv)aneRbIedIm,Iplit triglycerides EdlRtUv)andwknaMBIeføImeTACalikaxøaj; (adipocyte tissues) edaybNþúM Very Low Density Liporotein (VLDL) . CakarBit bNþúM VLDL pþl; [ Adipose tissue rYmCamYyRbPBsMxan;énGasIutRka . kardwkCBa¢ÚnenHskmμCaBiessenAeBleK briePaKkaLÚrIeRcInCagkareRbIR)as; . b:uEnþ enAeRkamlkçx½NÐminTan;briePaKGahar b¤ TurÖikS eføImbMElg GasIutRkaeTACa Acetoacetate nig Ketone bodies epSgeTot . eføImminGaceRbI Ketone bodies enH eT GaRs½yedayvaxVHnUvGg;sIumsMrab;karbMElg Acetoacetate [eTACa Acetyl CoA . ehtudUecñH Ketone bodies Edlplit)anedayeføImRtUv)andwknaMeTAsrIr³epSgeTotEdlenATIenaHBYkvaTTYlrg GuksIutkmμedIm,IeFIV ATP . 2- Adipose Tissue

Adipose Tissue KWCakMNsþúkTukCasMxan;énGasIutRka . Triglycerides EdlRtUv)anplit edayeføImRtUv)andwknaMtamry³crnþQameRkamTMrg;CasmasPaBénbNþMú VLDL . Triglycerides RtUv )anGIuRdUlIseday Lipases dUcKña EdlmanGMeBIeTAelI Chylomicron ehIy GasIutRkaRtUv)anRsUb

edayCalikaxøaj; . karsMeyaK Triglycerides enAkñúgCalikaxøaj;RtUvkar glycerol-3-phosphate . b:uEnþ Calikaxøaj;minGaceFIV glycerol-3-phosphate nig Cab;Tak;TgeTAnwg glycolysis sMrab;karpþl;én m:UelKulenH . ehtudUecñHekasikaxøaj;RtUvEtmanrYcCaeRscnUvRbPBrbs; glucose kñúgkarsMeyaK nig sþúkTuk Triglycerides . Triglycerides RtUv)anGIuRdUlIs nig sMeyaKeLIgvijedaysßitesßrenAkñúgekasikaénCalika- xøaj;. Lipases EdlsßitenAeRkamkarRtYtBinitürbs;G½rm:UnkMNt;nUvGRtaGIuRdUlIsrbs; Triglycerides

eTACaGasIutRka nig Glycerol . RbsinebI glucose sßitenAkñúgkarpþl;EdlmanEdnkMNt; vanwgminman glycerol-3-phosphate RKb;RKan;sMrab;karsMeyaKeLIgvijén Triglycerides ehIyGasIutRka nig glycerol RtUv)annaMecjeTAeføImsMrab;karEkécñxøaMgCagenHeTot ¬rUbTI 23-15¦ .

rUbTI 23-14 eføImRtYtBinitükMhab; glucose kñúgQam

3- Calikasac;dMu karTamTarfamBlénsac;dMuEdlsMrak;enAes¶omCaTUeTARtUv)anpþl;eday β-oxidation én GasIutRka . CaFm μtasac;dMuebHdUgcUlcitþ ketone bodies Cag glucose . b:uEnþ cMeBaHsac;dMukMBugeFIVkargarTTYlfamBledaykarbMEbkénkarpþl; glycogen rbs;xøÜnva pÞal; . karbMEbk glycogen begáIt)anCa glucose-6-phosphate EdlhUrcUledaypÞal;eTAkñúg glycolysis . RbsinebIsac;dMukMBugeFIVkarxøaMgeBkrhUtdl;vaminGacmanGuksIuEsnRKb;RKan; vabegáItnUv lactate ya:geRcIn . plitplcugbBa©b;énkareLIgem k¾dUcCa alanine ¬EdlTTYl)anBIkarbMEbkén

Stimulated by insulin

Glycogen ←G-6-P ← Glucose ↓ ↓ Fatty acid synthesis Used as fuel

Glucose from the blood

Stimulated by insulin

Glycogen → G-6-P → Glucose Fatty acids used by the liver as fuel

Glucose released into the blood

Fatty acids from adipose

VLDL (contains fatty acids)adipose tissue

(a) After a meal (b) After an overnight

RbUetGIun nig transamination rbs; pyruvate¦ RtUv)annaMecjeTAeføIm . enATIenHBYkvaRtUv)anbMElg eTACa glucose eday gluconeogenesis ¬rUbTI 23-16¦ .

rUbTI 23-15 karsMeyaK nig karbMEbkén Triglyceride enAkñúgCalikaxøaj;

rUbTI 23-16 TMnak;TMngemtabUlIsrvageføIm nig sac;dMu

4- xYrk,al sßitenAeRkamlkçx½NÐFm μta xYrk,aleRbIR)as; glucose CaRbPBEtmYyKt;énfamBlemta-bUlIs . enAeBlsBa’gÁkaysßitenAkñúgsPaBsMrak; RbEhl 60 % én glucose esrIrbs;sBa’gÁkay RtUv)aneRbIR)as;edayxYrk,al . PaBTurÖikSeFIV[)at;bg;nUvsþúk glycogen nig cMnYn glucose Edlman sMrab;xYrk,alFøak;cuHya:gxøaMg . bnÞab;mk Ketone bodies (acetoacetate nig β-hydroxybutyrate)

RtUv)aneRbIedayxYrk,alCaRbPBfamBlepSgeTot . GasIutRkaRtUv)andwknaMenAkñúgQameRkamTMrg;

Glucose (from the liver

Fatty acids Glucose

VLDL (from the liver)

Glycerol-3-Phosphate

Fatty acyl CoA

Triglycerides

Glycerol Fatty acids

Glycerol (to the liver)

Fatty acids (to the liver)

Lipase

Glycogen → Glucose-6-Phosphate → Glucose ↑ Alanine → Pyruvate ↑ Lactate

Liver

Glucose ↓ Glycolysis Pyruvate ↓ ↓ Lactate Alanine ↑ Protein degradation

Muscle

bNþúMCamYyRbUetGIun nig minGacqøgr)aMgQam-xYrk,al (blood-brain barrier) EdlRtUv)aneRbIeday ekasikaxYrk,alCaRbPBfamBl . b:uEnþ Ketone bodies EdlmanbgÁúM carboxyl esrI rlayenAkñúg QamnwgGaccUleTAkñúgxYrk,al . 6- \T§iBlrbs; INSULIN nig GLUCAGON eTAelIemtabUlIsekasika G½rm:Un Insulin RtUv)anplitedayedayekasika β énGIuLÚt× Langerhands enAkñúglMEBg . vaRtUv)anbeBa©jBIekasikaTaMgenHkñúgkareqøIytbeTAnwgkarekIneLIgnUvkMrit glucose enAkñúgQam . Insulin eFIV[fycuHkMhab;én glucose kñúgQamedaykarbgá[karpøas;bþÚrmYycMnYnenAkñúgemtabUlIs ¬taragTI 23-1¦. viFIsamBaØCageKbMputkñúgkareFIV[cuHkMrit glucose kñúgQamKWeP¶ackarsþúkTwkén glucose

TaMgCa glycogen nig Triglycerides . ehtudUecñH Insulin eFIV[skmμnUvdMeNIrkarCIvsMeyaK nig raraMg dMeNIrkarbMEbk . Insulin manGMeBIeTAelIEtekasikaTaMgenaH ¬EdleKsÁal;faCaekasikacMNuc¦ Edlman Specific insulin receptor protein enAkñúg plasma membranes . ekasikacMNucsMxan;cMeBaH insulin

KWCaekasikaeføIm ekasikaxøaj; nig ekasikasac;dMu . kMrit glucose kñúgQamFmμtamanRbEhl 10 mmol . b:uEnþ karbriePaKGahar (meal) begáInnUv kMhab; glucose enAkñúgQamya:gxøaMg nig eP¶ackarbeBa©j Insulin . karcgCabnÞab;rbs; Insulin

eTAnwgRbUetGIunEdlCaGñkTTYlenAelIPñasekasikab:Hnwg)aøs μa (plasma membrane insulin receptor

protein) begáInGRtakardwknaM glucose qøgkat;PñasnigcUleTAkñúgekasika . Insulin man\T§iBlCaeRcInRbePTeTAelIRKb;TidæPaBénemtabUlIs . ∙ Carbohydrate metabolism - Insulin eP¶ackarsMeyaK glycogen . enAkñúgeBlCamYyKña vararaMg glycogenolysis nig gluconeogenesis . lT§plsrubénskmμPaBTaMgenHkarsþúkTukén glucose EdlelIs . ∙ Protein metabolism - Insulin eP¶ackardwknaM nig kareRbIR)as;énGasIutGamIneN k¾dUcCa smahr½nkm μénGasIutGamIneNeTAkñúgRbUetGIun . ∙ Lipid metabolism - Insulin eP¶ackareRbIR)as; glucose edayCalikaxøaj (adipose cells) k¾dUcCakarsMeyaK nig karsþúkTuk Triglycerides . dUcEdleyIg)aneXIj karsþúkTukénlIBIdRtUvkar RbPB glucose nig Insulin CYykarEkécñedaykarbegáIn glucose Edlman . kñúgeBlCamYyKñaenH Insulin raraMgkarbMEbkénsþúk Triglycerides .

taragTI 23-1 Comparaison of Metabolic Effects of Insulin and Glucagon

Action Insulin Glucagon

Cellular glucose transport Increased No effect Glycogen synthesis Increased Decreased Glycogenolysis in liver Decreased Increased Glyconeogenesis Decreased Increased Amino acid uptake and protein synthesis Increased No effect Inhibition of amino acid release and protein degradation

Decrease No effect

Lipogenesis Increased No effect Lipolysis Decreased Increased Ketogenesis Decreased Increased

dUcEdlGñkGac)a:n;sμanrYcmkehIy Insulin KWCaEpñkEtmYyKt;énkarEktMrUvTaMgmUlénemta-bUlIsenAkñúgekasika . G½rm:UnTIBIr glucagon RtUv)anbeBa©jedayekasika α énGIuLÚt× Langerhans

kñúgkareqøIytbeTAnwgkarfycuHkMrit glucose kñúgQam . \T§iBlrbs; glucagon CaTUeTARbQmeTAnwg \T§iBlrbs; Insulin RtUv)ansegçbenAkñúgtaragTI 23-1 . eTaHbICavaminman\T§iBledaypÞal;eTAelI kareRbIR)as; glucose k¾b:uEnþ glucagon raraMgkarsMeyaK glycogen nig eP¶ac glycogenolysis nig gluconeogenesis . vak¾eP¶acpgEdrnUvkarbMEbkxøaj; nig Ketogenesis . \T§iBlpÞúyKña (Antigonistic effects) rbs;G½rm:UnTaMgBIrenH RtUv)aneXIjenAkñúgrUbTI 23-17 vasMxan;sMrab;karrkSaénkMrit glucose kñúgQamRKb;RKan; . enAkñúgry³eBlminTan;briePaKGahar krit glucose kñúgQamTabeP¶ackarplit glucagon EdlbegáInnUv glucose kñúgQamedaykareP¶ac karbMEbkén glycogen nig karplit glucose eday gluconeogenesis . karenHFananUvkarpþl;rYcCa eRscnUv glucose sMrab;Calika CaBiessxYrk,al . müa:gvijeTot enAeBlkMrit glucose kñúgQamx<s; xøaMgeBk Insulin RtUv)anbeBa©j . vaeP¶ackardkecjnUv glucose EdlelIsedaykarCMrujkareRbIR)as; nig bNþal[manpøÚvénkarsþúkTuk .

rUbTI23-17 segçbén\T§iBlRbQmKñaén Insulin nig Glucagon

Fin


Metabolism

Fasting (↓ Glucose)

Absorption of meal

(↑ Glucose)

Blood ↑ Glucose ↑ Amino acid ↑ Fatty acid ↑ Ketone bidies

Metabolism

↓ Insulin ↑ Glucagon ↑ Insulin

↓ Glucagon

↑ Insulin/ Glucagon ratio

↓ Insulin/ Glucagon ratio

Hydrolysis of glycogen, fat, and

protein +

Gluconeogenesis and ketogenesis

Formation of glycogen, fat, and protein

Blood ↓ Glucose ↓ Amino acid ↓ Fatty acid ↓ Ketone bidies

International University Biochemistry

by Mr. Chroeng Sopheap, Ph, phone: 012864353 or 011717361, e-mail: [email protected] 1 of 36

esckþIepþImBn§úviC¢aénm:UelKul (Introduction to Molecular Genetics)

vtßúbMNgemeron 1-KUrcnasm<½n§TUeTAén DNA nig RNA nucleotides

2-BiB’NnaBIrcnasm<½n§rbs; DNA nigeRbóbeFobvaCamYynwg RNA

3-Bnül;BIkarsMeyaK DNA (DNA replication)

4-rab;fñak;TaMgbIénm:UelKul RNA nig BiB’NnaBImuxgarrbs;BYkva 5-Bnül;BIdMeNIrkarénkarsMeyaK RAN (Transcription)

6-rab; nig Bnül;BIRbePTTaMgbIén post-transcriptional modifications rbs; eukaryotic mRNA

7-BiB’NnaBIFatusMxan;rbs; genetic code nig begáItnUv {kardwg} cMeBaHPaBsmsYnrbs;va (“feel” for its

elegance)

8-BiB’NnaBIdMeNIrkarénkarsMeyaKRbUetGIun (Translation) . 9-[niymn½y mutation nig yl;faetI mutations bgá[manCMgWmharIk nig karsøab;ekasikatamrebobdUcemþc 10-BiB’NnaBI]bkrN_EdleKeRbIkñúgkarsikSaBI DNA nig visVkm μénEhSn (genetic engineering) . 11-BiB’NnaBIdMeNIrkarén Polymerase Chain Reaction (PCR) nig BiPakSaBIkareRbId¾sMxan;éndMeNIrkar enH .




KMeragemeron 1-rcnasm<½n§rbs;nuyekøGUTId (The Structure of the Nucleotide)

k-smasPaBKImIrbs; DNA nig RNA (Chemical Composition of DNA and RNA)

x-rcnasm<½n§rbs;nuyekøGUTId (Nucleotide Structure)

2-rcnasm<½n§rbs; DNA nig RNA (The Structure of DNA and RNA)

k-rcnasm<½n§rbs; DNA : The Double Helix (DNA Structure : The Double Helix)

x-rcnasm<½n§rbs; RNA

3-karsMeyaK DNA (DNA Replication)

4-B½t’manhUrcUleTAkñúgRbB½n§CIvsa®sþ (Information Flow in Biological Systems)

k-fñak;énm:UelKul RNA (Classes of RNA Molecules)

x-karsMeyaK RNA (Transcription)

K-dMeNIrkarbnÞab;BIkarsMeyaK RNA (Post-transcriptional processing of RNA)

5-kUdénEhSn (Genetic Code)

6-karsMeyaKRbUetGIun (Protein Synthesis)

k-tYnaTIrbs; Transfer RNA (Role of Transfer RNA)

x-dMeNIrkarénkarsMeyaKRbUetGIun (The Process of Translation)

7-brivtþn_ kaMrsμIB’NsVay nig karCYsCul DNA

(Mutation, Ultraviolet Light, and DNA Repair)

k-RbePTénbrivtþn_ (The Nature of Mutations)

x-lT§plénbrivtþn_ (The Results of Mutations)

K-sarFatueFIV[manbrivtþn_ nig sarFatueFIV[manmharIk (Mutagens and Carcinogens)

X-karxUcxatedaysarkaMrsμIB’NsVay nig karCYsCul DNA

(Ultraviolet Light Damage and DNA Repair)

g-plvi)akénPaBex©aHenAkñúgkarCYsCul DNA (Consequences of Defects in DNA Repair)

8-DNA Edlbgáat; (Recombinant DNA)

k-]bkrN_eRbIenAkñúgkarsikSa DNA (Tools Used in the Study of DNA)

x-visVkm μénEhSn (Genetic Engineering)

9-POLYMERASE CHAIN REACTION (PCR)




cUrRkeLkemIlnisSitenAkñúgfñak;eronKImIrbs;Gñk . BYkeKTaMgGs;KñamanlkçN³rYmCaeRcIn ³ CMhreLIg elI/ k,almanEPñkBIr RcmuHmYy nig mat;mYy RtecokmYyenATaMgBIrxagénk,al nig GIVepSg²eTot . Gñknwgman karlM)akkñúgkarrab;BIPaBdUcKñaEdlkMNt;BIGñk nig mitþrYmfñak;rbs;GñkfaCa Homo Sapiens . enAeBlGñkBinitüedayykcitþTukdak;eTAbuKÁlTaMgLay Gñkcab;epþImkt;sMKal;PaBxusKñaCaeRcIn . B’N EPñk B’Nsk; B’NEs,k rUbragRcmuH kMBs; maDdgxøÜn ³ lkçN³TaMgGs;enH nig lkçN³eRcIneTotbgðajBI RbePT epSg²EdleFIV[EbøkBImnusSmñak;eTAmnusSmñak;eTot . sUm,IEtenAkñúgRKYsarEtmYyk¾eday EdlPaBdUcKñaGacRtUv )anbgðajeRcInEfmeTot buKÁlmñak;²manBhilkçN³mYyrbs;xøÜn (unique appearance) . CakarBit kUnePøaH dUcKñaemIleTAdUcKñaEmnETn ¬kñúgPaKeRcInéneBlevla¦ . m:UelKulEdlTTYlxusRtUvcMeBaHPaBdUcKña nig PaBxusKñaTaMgGs;enHKWCa Deoxyribonucleic acid

(DNA) . karrmUedaytwgnUvrcnasm<½n§EdleKehAfaCa Chromosomes enAkñúgéNVy:Urbs;ekasika DNA

mankUdénEhSn (genetic code) edIm,IbegáIt)anrab;Ban;énRbUetGIunxus²KñaEdlbegáIt)anCarUbeyIg . RbUetGIun TaMgenHrYmmanGg;sIumEdlTTYlxusRtUvcMeBaHkarplitnUv Pigment melanin . Melanin kan;EteRcIn

EdlxøÜneyIgdak;kmμviFIedayEhSnedIm,IeFIV B’Nrbs;sk; EPñk nig Es,kkan;EtexμA . RbUetGIunepSgeTotKWCa RbUetGIunrcnasm<½n§ . EhSnrbs; α-keratin EdlbegáItsk;kMNt;faetIsk;Twkrlk Rtg; b¤ rYj . rab;Ban;EhSn pÞúkB½t’manénEhSncMeBaHrab;Ban;RbUetGIunEdlbBa¢akarbegáItrUbrbs;eyIg nig CMenOmYycMnYn ¬\riyabf¦ . lkçN³EhSnRtUv)anbBa¢ÚnBICMnan;mYyeTA[CMnan;bnÞab; . enAeBlEs<m (sperm) rYmCamYynwgGUvul (ovum) karbgákMeNIt (conceptus) RtUv)anbegáIteLIgBImYysMrab;Kt;énRkUm:UhSÚmmþay (maternal chromo-

some) nig mYysMrab;Kt;énRkUm:hSÚmjativgS . enAeBlBgEdlbgákeNItenH (fertilized egg) bMEbkxøÜn ekasikakUnnimYy²nwgTTYl)anvtßúcMLg (a copy) mYyénRkUm:UhSÚmnimYy²rbs;RkUm:UhSÚmTaMgBIrenH . EhSnEdl enAelIRkUm:UhSÚmTaMgenHnwgkMNt;TisedAkarGPivDÆTarkkñúgépÞ (fetal development) BIekasikaEdlbgákMeNIt enaHeTACaTarkeTIbnwgekIt (Newborn) rYmCamYynwglkçN³TaMgGs;EdleyIgTTYlsÁal;faCamnusS . enAkñúgCMBUkenHeyIgnwgRsavRCavBIrcnasm<½n§rbs; DNA nig ehtukarN_énm:UelKulEdlbkERbnUv B½t’manénEhSnrbs;EhSnmYyeTACarcnasm<½n§rbs;RbUetGIun . 1-rcnasm<½n§rbs;nuyekøGUTId (The Structure of the Nucleotide)

k-smasPaBKImIrbs; DNA nig RNA (Chemical Composition of DNA and RNA)

DNA RtUv)anrkeXIjeday Friedrich Meischer enAkñúgqñaM 1869 . b:uEnþ vaminRtUv)aneKTTYlsÁal; faCaB½t’manénEhSnrhUtdl;qñaM 1950 . enAkñúgqñaM 1953 James Watson nig Francis Crick )ane)aHBum< GtßbTEdlBiB’NnaBIrcnasm<½n§rbs;m:UelKul DNA . cab;BImanGtßbTenaH nig karsikSaCaeRcInepSgeTot eyIg )ansÁal;fa deoxyribonucleic acid (DNA) nig ribonucleic acid (RNA) KWCa polymer Evgénnuy-



ekøGUTId (Nucleotide) . nuyekøGUTIdnimYy²ekIteLIgBI ÉktabI ³ sárEdlman 5GatUmkae)an )asGasUt (nitrogenous base) nig manpgEdrnUvbgÁúMhVÚssVatCYnkalmYy BIr b¤ bI . )asGasUtKWCarcnasm<½n§ExSbiTcMruH (heterocyclic ring structure) EdlExSemmanGatUmkae)an nig GasUt . vamancMNat;fñak;BIrén)asGasUt . )asGasUtCa Purine KWCaExSbiTEdlman 6 GatUmCab;CamYynwgExSbiTEdlman 5GatUm . )asGasUtCa Pyrimidine manExSbiTEtmYyEdlman 6 GatUm . rcnasm<½n§enHRtUvbgðajenAkñúgrUbTI 24-1 .

rUbTI 24-1

sárEdlman5GatUmkae)anenAkñúg RNA KWCa Ribose nig sárEdlenAkñúg DNA KWCa 2’-

deoxyribose. PaBxusKñaEtmYyKt;rvagsárTaMgBIrenHKWGvtþmanénbgÁúM Hydroxyl enAelIkae)an 2’ rbs; 2’-

deoxyribose . Purine enAkñúgTaMg DNA nig RNA KWCa Adenine nig Guanine . TaMg DNA nig RNA

man)as Pyrimidine Ca Cytosine b:uEnþ)asTI 4 KW Thymine manEtenAkñúg DNA nig Uracil manenAkñúgEt RNA . smasPaBKImIrbs; DNA nig RNA RtUv)ansegçbenAkñúgtaragTI 24-1 . epSgBIenHeTot nuyekøGUTIdEdlbegáIt DNA nig RNA manbgÁúMhVÚsVat . nuyekøGUTIdGacmanbgÁúM-hVÚsVatmYy (mono) BIr (di) nig bI (tri) .



taragTI 24-1

Chemical Composition of Nucleic Acids

DNA RNA Sugar 2’-Deoxyribose Ribose Purine Nitrogenous Bases Adenine (A)

Guanine (G) Adenine (A) Guanine (G)

Pyrimidine Nitrogenous Bases Cytosine (C) Thymine (T)

Cytosine (C) Uracil (U)

x-rcnasm<½n§rbs;nuyekøGUTId (Nucleotide Structure)

nuyekøGUTIdRtUv)anplitedaykarrYmKñaénsár )asGasUt nig y:agehacNas;k¾bgÁúMhVÚsVatmYyEdr dUcEdl)anbgðajenAkñúgrUbTI 24-2 . GaRs½yedayrcnasm<½n§FMenHmanm:UelKulExSbiTBIr sár nig )as eyIgRtUv EtmanrebobgayRsYlmYyedIm,IBiB’NnaBIGatUmrbs;ExSbiTnimYy² . cMeBaHehtuplenH GatUménExSbiTrbs;sár RtUv)aneKkMNt;edayRBIm (prime “ ’ ” ) edIm,IeFIV[eXIjxusKñaBIGatUménExSbiTrbs;)as . sm<½n§kUva:Lg;rvag sár nig bgÁúMhVÚsVatKWCasm<½n§hVÚsVÚeGEsÞr (phosphoester linkage) EdlekIteLIgedaysarRbtikmμdkm:UelKul TwkmYyrvag 5’-OH rbs;sár nig ─OH rbs;bgÁúMhVÚsVat . sm<½n§rvag)as nig sárKWCasm<½n§ β-N-glycosidic

nig vaP¢ab;eTAnwg 1’-Carbon rbs;sár nig GatUmGasUtrbs;)asGasUt (N-9 rbs; purine nig N-1 rbs; pyrimidine) . edIm,IehAeQμaHnuyekøGUTId cab;epþImedaysamBaØCamYynwgeQ μaHrbs;)asGasUt nig GnuvtþtamviFan samBaØxageRkam ³ dkBaküxagcug “-ine” ecj nig CMnYsvaeday “-osine” cMeBaH purine nig “idine”cMeBaH pyri-

midine . Uracil KWCakarelIkElgmYycMeBaHviFanenH . enAkñúgkrNIenH “-acil” RtUv)anCMnYseday “-

idine” edaybegáIt)anCaeQμaH UUridine . nuyekøGUTIdrYmCamYynwgsárCa Ribose KWCa Ribonucleotides nig nuyekøGUTIdEdlmansár 2’-deo-

xyribose KWCa Deoxyribonucleotides . cMeBaH deoxyribonucleotide mYymanbuBVbT “deoxy-” dak; enAxagmuxeQμaH)asGasUtEdlEkrYc . minmanbuBVbTRtUvkarsMrab; ribonucleotides b¤ cMeBaH Thymindine

EdleKeXIjmanvaenAEtkñúg DNA . bEnßmbuBVbTedIm,IbgðajBIcMnYnbgÁúMhVÚsVatEdlRtUv)anP¢ab; . monophosphate manbgÁúMhVÚsVatmYy diphosphate manbgÁúMhVÚsVatBIr nig triphosphate manbgÁúMhVÚsVatbI . GaRs½yedayeQμaHeBjeljrbs;nuyekøGUTIdeFIV[mankareTIsETg GkSrkat;y:agRsYlRtUv)aneKeRbICa TUeTA . GkSrkat;RtUv)ansegçbeTAkñúgtaragTI 24-2 .



tarragTI 24-2

Names and Abbreviations of Ribonucleotides and Deoxyribonucleotides Containing Adenine

Nucleotides Abbreviation Deoxyadenosine monophosphate Deoxyadenosine diphosphate Deoxyadenosine triphosphate Adenosine monophosphate Adenosine diphosphate Adenosine triphosphate

dAMP dADP dATP AMP ADP ATP

rUbTI 24-2

2-rcnasm<½n§rbs; DNA nig RNA (The Structure of DNA and RNA)

ExSEtmYyKt;rbs; DNA (single strand of DNA) KWCa polymers rbs;nuyekøGUTIdEdlcgP¢ab;Kña BImYyeTAmYyedaysm<½n§ 3’-5’phosphodiester . ExSemrbs; polymer RtUv)aneKehAfaCa Sugar-

phosphate backbone GaRs½yedayvaekIteLIgGMBIxñatqøas;Kñaén sugar 2’-deoxyribose nig bgÁúM phosphate enAkñúgsm<½n§ phosphodiester . )asGasUtRtUv)ancgeTAnwgsárnimYy²edaysm<½n§ N-glycosidic

¬rUbTI 24-3¦ .



rUbTI 24-3

k-rcnasm<½n§rbs; DNA : The Double Helix (DNA Structure : The Double Helix)

James Watson nig Francis Crick KWCaGñkTImYyEdlBiB’NnaBIrcnasm<½n§ 3-D rbs; DNA enAkñúg qñaM 1953 . BYkKat;RsavRCaveXIjrcnasm<½n§edaykarsg;nUvKMrU (model) edaysMGageTAelIlT§plrbs;Gñk epSgeTot . Irwin Chargaff )ansegáteXIjfacMnYnrbs; Adenine enAkñúgm:UelKul DNA NamYyes μIeTAnwg cMnYn Thymine . dUcKñaenHEdr Kat;)aneXIjfacMnYnrbs; Cytosine nig Guanine k¾es μIKñaEdr . karsikSa edaykareFIV[)aøtkaMrsμ IGiucrbs; Rosalind Franklin nig Maurice Wilkens )anbgðajBIcMgaydUc²KñaCaeRcIn EdlbBa¢ak;BIlkçN³rbs; DNA : 0.34 nm, 3.4 nig 2 nm ¬cUrBinitüemIlrcnasm<½n§rbs; DNA kñúgrUbTI 24-4 edIm,IBinitüemIlBIsar³sMxan;énkarvas;enH¦ . rYmCamYynwgB½t’manenH Watson nig Crick )ansnñidæanfa DNA KWCa double helix énExSBIrrbs; DNA evjcUlKña . vamansar³sMxan;kñúgkarRbdUcrcnasm<½n§rbs; double helix eTAnwgCeNþIrrages<ór (spiral staircase) . sugar-phosphate backbones rbs;ExSTaMgBIr én DNA rMuCMuvijBIxageRkAén helix dUcCabgáan;édCeNþIr enAelICeNþIrrages<ór (spiral staircase) . )asGasUt lUtcUleTAEpñkkNþaledaybegáItCamMuCamYynwgG½kSrbs; helix . GñkGacRsm½yfa)asGasUtbegáIt)anCakaM rbs;CeNþIr . rcnasm<½n§rbs;m:UelKulRbNitenHRtUv)anbgðajenAkñúgrUbTI 24-4 .



rUbTI 24-4

ExSTaMgBIrrbs; DNA RtUv)ancgP¢ab;Kñaedaysm<½n§GIuRdUEsnrvag)asGasUtenAEpñkkNþalrbs; helix. Adenine begáIt)anCasm<½n§GIuRdUEsnBIrCamYynwg Thymine nig Cytosine begáIt)anCasm<½n§GIuRdUEsnbICamYy nwg Guanine ¬rUbTI 24-4 nig TI 24-5¦ . BYkvaTaMgGs;enHRtUv)aneKehAfaCa KU)as (base pairs) . ExSTaMg

BIrrbs; DNA KWCaExSbMeBjKña (complemetary strands) GaRs½yedayQutrbs;)as (the sequence of

bases) enAelIExS DNA mYykMNt;edaysV½yRbvtþinUvQutrbs;)asenAelIExS DNA mYyeTot . enAeBlvaman Adenine enAelIExSmYyCaTUeTAvaman Thymine enAelITItaMgdUcKñaenAelIExSQmKña .



Ggát;p©itrbs; double helix KW 2.0 nm . karenHRtUv)ankMNt;edayTMhMrbs;KUr)as purine-pyrimi-

dine . helix bMeBjmYyCuMral; 10 KU)as . mYyCMueBjmanRbEvg 3.4 nm . ehtudUecñHKU)asnimYy²BnøÚt helix

eTAmux)anRbEvg 0.34 nm . TidæPaBsMxan;cugeRkaybMputmYyeTotrbs; DNA double helix KWfaExSTaMgBIrmanTisedApÞúyKña (antiparallel strands) dUcCa]TahrN_enAxageRkamenH ³ 5’ P─S─P─S─P─S─P─S─P─S─P─S─OH 3’ │ │ │ │ │ │ A T G C G A : : : : : : T A C G C T │ │ │ │ │ │ 3’OH─S─P─S─P─S─P─S─P─S─ P─S─P 5’

m:üagvijeTot ExSTaMgBIrrbs; helix rt;edayb®Ba©as;TisedAKña ¬rUbTI 24-4¦ . enAeBlEtExSTaMgBIr RtUvEt antiparallel eTGac[KU)asbegáIt)anCasm<½n§GIuRdUEsnEdlcgExSTaMgBIr[enACab;Kña .

rUbTI 24-5

x-rcnasm<½n§rbs; RNA

Sagra-phosphate backbone rbs; RNA man ribonucleotide k¾RtUv)ancgP¢ab;Kñatamry³sm<½n§ 3’-5’ phosphodiester . sm<½n§ phosphodiester TaMgenHdUceTAnwgsm<½n§ phosphodiester enAkñúg DNA

Edr . b:uEnþ m:UelKul RNA xusBIm:UelKul DNA eTAtamlkçN³RKwHbIy:ag .



m:UelKul RNA CaFmμtamanExSEtmYyKt; (single-strand)

ExSem sugar-phosphate rbs; RNA man ribonucleotides EdlcgP¢ab;Kñatamry³sm<½n§ 3’-5’

phosphodiester . ehtudUecñH sugar ribose RtUv)aneKeXIjenATIkEnøgrbs; 2’-deoxyribose . )asGasUt Uracil (U) CMnYs Thymine (T)

eTaHbICam:UelKul RNA manExSEtmYyKt;k¾eday b:uEnþ KU)asrvag Uracil nig Adenine nig rvag Guanine nig Cytosine GacekItmaneLIg . eyIgnwgbgðajBIsar³sMxan;énlkçN³enH enAeBleyIgBInitüBI rebobEdlm:UelKul RNA Cab;Tak;TgenAkñúgkarsMEdg[eXIjnUvB½t’manénEhSnenAkñúg DNA . 3-karsMeyaK DNA (DNA Replication)

DNA RtUvEt)ansMeyaK (replicate) muneBlekasikabMEbkxøÜn (divide) dUecñH ekasikakUnnimYy² (daughter cells) TTYl)anvtßúcMLgénEhSnnimYy² . ekasikaEdl)at;bg;nUvEhSnsMxan; (critical gene) nwg RtUvsøab; dUcCabuKÁlEdlmanCMgWEhSn (genetic disease) PaBex©aHenAkñúgEhSnsMxan; (a defect in an

important gene) Gacsøab;munGayu . ehtudUecñH vamansar³sMxan;EdldMeNIrkarénkarsMeyaK DNA (DNA

replication) begáItnUvvtßúcMLgedaycMTisedAdac;xat (absolutely accurate copy) BIB½t’manEhSndMbUg (original genetic information) . RbsinebImankMhusRtUv)anekIteLIgcMeBaHEhSnsMxan; lT§plGacCa brivtþn_EdlbNþal[søab; (lethal mutation) .

rUbTI 24-6



CMhanTImYyenAkñúgkarsMeyaK DNA KWkarEjkecjBIKñaénExSTaMgBIrrbs; DNA . RbUetGIunEdleFIVkar garenHedaykarpþac;sm<½n§GIuRdUEsnrvagKU)as . bnÞab;BI Gg;sIum DNA polymerase {Gan} parental strand k¾ GacRtUv)aneKehAfaCa vtßúKMrU (template) nig sMrYlkareFIV polymerization énExSkUntamviFanbMeBjKña . m:UelKul Deoxyribonucleotide triphosphate RtUv)aneRbICavtßúnaMmux (precursor) sMrab;karsMeyaK DNA. b:uEnþ bgÁúMhVÚsVatBIrcueRkayRtUv)anpþac;ecjenAkñúgdMeNIrkarenH . karpþac;enHbeBa©jnUvfamBlEdlRtUv kareday DNA polymerase edIm,IbegáItCasm<½n§ phosphoester rvag 3’-OH rbs; 2’-deoxyribose nig 5’-phosphoryl group rbs; deoxyribonucleotide monophosphate RtUv)anbEnßeTAkñúgExS DNA . GaRs½yeday DNA polymerase {Gan} parental strand nig begáItnUvExS DNA kUnEdlman lkçN³bMeBjKña m:UelKul DNA f μ InimYy²man parental strand mYy nig daughter strand EdleTIbsMeyaK )anmYy . rebobénkarsMeyaK DNA enHRtUv)aneKehAfa {karrsMeyaKrkSaTukBak;kNþal} (semi-

conservative replication) ¬rUbTI 24-6¦. P½sþútagcMeBaHynþkarénkarsMeyaK DNA enHRtUv)anpþl;edaykarBiesaFn_EdlerobcMeday Matthew

Meselson nig Franklin Stahl enAkñúgqñaM 1958 . enAkñúgeBlenaH rebobénkarsMeyaKBIrRtUv)aneKelIkyk mkBicarNar ³ rebobénkarsMeyaKrkSaTukBak;kNþalEdleTIbEt)anBiB’NnaBIxagelI nig rebobrkSaTuk (conservative model) . rebobTI 2 enH)anbBa¢ak;fa double-strand parental DNA molecule RtUv)an rkSaTukedayminmankarb:HBal; nig double-strand daughter DNA molecule RtUv)anbegáIt . Meselson

nig Stahl bNþúH Escherichia coli cell enAkñúgfñalbNþúHEdl 15NH4+ KWCaRbPBGasUtEtmYyKt; . cUrrMlwk

faRKb isotopes énFatuKImImYybgðajBIlkçN³KImIdUcKña . ehtudUecñH 15NH4+ nwgbMeBjmuxgartamrebobdUcKña

eTAnwg 14NH4+ Fmμta . ehtudUecñH RKb; DNA rbs;ekasikaman nonradioactive, heavy isotope enH .

ekasikaTaMgenHmanEt 15NH4+-labeled DNA bnÞab;mkRtUv)aneKbEnßmeTAelIfñalEdlsMbUrEtGIusUtUbGasUt

14NH4+ nig GnuBaØat;[vaduHmYyvdþénkarbMEbkekasika . enAeBlEdlm:UelKul DNA kUnRtUv)aneKEjkecj

nig viPaK eKeXIjfa DNA nimYy²ekIteLIgBIExSmYyén “heavy” DNA EdlCa parental strand nig ExSmYy eTotBI “light” DNA EdlCa daughter strand fμ I . bnÞab;BIvdþTI 2 énkarbMEbkekasikaenAkñúgfñalbNþúHEdl man 14NH4

+ mYyPaKBIr (1/2) isolated DNA minman 15N nig mYyPaKBIr (1/2) manl,ay 50¼50 én DNA Edlman 14N nig 15N ¬rUbTI 24-6¦ . Tinñn½yTaMgenHpþl;karKaMRTy:agxøaMgcMeBaHkarsMeyaKrkSaTuk-Bak;kNþal (semiconservative replication) edaykarbgðajfa parental strand rbs;m:UelKul DNA eRbI CavtßúKMrUsMrab;karsMeyaKén daughter strand . dUecñH m:UelKul DNA EdlsMeyaKf μ Iman parental strand

mYy nig daughter strand EdleTIbsMeyaKf μ ImYy . GaRs½yedayvamansar³sMxan;kñúgkarbegáItvtßúcMLgcMTisedAmYyBI parental DNA vamansar³xøaMg Nas;kñúgkareCosvagPaBmankMhuskñúgdMeNIrkarsMeyaK . bEnámBIelIkarsMrYlkarsMeyaKén DNA strand f μ I DNA polymerase GacemIlEk (proofread) strand EdleTIbsMeyaKf μ I . RbsinebInuyekøGUTIdxus (wrong



nucleotide) RtUv)anbEnßmeTAelI DNA EdllUtlas; vaRtUv)andkecj nig CMnYnedaynuyekøGUTIdRtwmRtUvvij. enAkñúgviFIenH vtßúcMLgBitR)akdrbs; parental DNA RtUv)anFana . RKb;B½t’manEhSnrbs;)ak;etrIdUcCa E. coli man circular piece of DNA EtmYyKt;EdlekIteLIgBI bIlannuekøGUTId nig RtUv)aneKehAfaCaRkUm:UsUm (chromosome) . DNA replication enAkñúg E. coli

cab;epþImBIQutEtmYyKt;enA circular chromosome EdleKsÁal;faCa Replication origin . karsMeyaK (replication) ekIteLIgedayTisedABIrpÞúyKñakñúgel,ÓnRbEhl 500 nuyekøGUTIdf μIral;vinaTI ¡ cMNucEdl deoxyribonucleotides RtUv)anbEnßmedIm,IbegáItnUv daughter strand RtUv)aneKsÁal;faCa Replication

fork ¬rUbTI 24-7¦ . enATIenHEdl DNA RtUv)anebIkedIm,IGnuBaØati[mankarcgnUvRbUetGIunepSg² nig Gg;sIum EdlTTYlxusRtUvcMeBaH DNA replication . GaRs½yeday karsMeyaK DNA ekIteLIgedayTisedABIrpÞúyKña vaRtUvman replication fork BIrEdleFIVclnakñúgTisedApÞúyKña . dUecñH Circular DNA molecules manragdUc GkSr theta (θ) .

rUbTI 24-7



4-karhUrcUlB½t’maneTAkñúgRbB½n§CIvsa®sþ (Information Flow in Biological Systems)

Central dogma ¬siT§anþkNþal¦rbs;CIvviTüaénm:UelKulEføgfaenAkñúgekasika karhUrénB½t’manBn§ú (genetic information) EdlmanenAkñúg DNA KWCapøÚvEdlmanTisedAEtmYy (one-way street) EdldwknaM ecjBI DNA eTA RAN eTARbUetGIun . dMeNIrkarenHEdl single strand of DNA eRbICavtßúKMrUsMrab; karsMeyaKm:UelKul RNA RtUv)aneKehAfaCa Transcription . Baküfa Transcription manRsLayBI BaküLataMg transcribere nig mann½yedaysamBaØfa {eFIVkarftcMLg} . ehtudUecñH enAkñúgdMeNIrkarenH Epñk mYyénB½t’manenAkñúg DNA RtUv)anftcMLgeTAkñúg RNA strand . dMeNIrkarEdlsar (message) RtUv)an bMElgeTACaRbUetGIunRtUv)aneKehAfaCa Translation . mindUceTAnwg Transcription dMeNIrkarén Translation Cab;Tak;TgeTAnwgkarbMElgB½t’manBIPasamYyeTACaPasamYyepSgeTot . enAkñúgkrNIenH B½t’manBn§úenAkñúg linear sequence rbs;nuyekøGUTIdRtUv)anbkERbeTACaRbUetGIun EdlCa linear sequence of

amino acids . karsMEdk[eXIjnUvB½t’manEdlmanenAkñúg DNA KWCamUldæanRKwHedIm,IeFIVkarGPivDÆ begáIt nig rkSanUvRKb;srIragÁ . k-fñak;énm:UelKul RNA (Classes of RNA Molecules)

cMNat;rbs;m:UelKul RNA bIRtUv)anbegáIteLIgeday Transcription : messenger RNA, transfer

RNA nig ribosomal RNA . 1. Messenger RNA (mRNA) dwknaMB½t’manBn§úsMrab;RbUetGIunBI DNA eTA[ ribosomes . vaenH KWCa complementary RNA copy énBn§úenAelI DNA . 2. Ribosomal RNA (rRNA) smasPaBénrcnasm<n§ nig muxgarrbs; ribosomes EdlKWCa {evTika} (platform) EdlkarsMeyaKRbUetGIunekIteLIgenATIenaH . vamanm:UelKul rRNA bIRbePTenAkñúg ribosome )ak;etrI nig bYnRbePTenAkñúg ribosomes rbs;BYk eukaryotes . 3. Transfer RNA (tRNA) bkERbkUdBn§úrbs; mRNA eTACa primary sequence of amino

acids enAkñúgRbUetGIun . bEnßmBIrcnasm<½n§TI 1 m:UelKul tRNA manragrcnasm<½n§TI 2 ragdUcsøwkeQImanbI snøwk EdlbNþalmkBIkarcgsm<½n§GIuRdUEsnénKU)as (A─U nig G─C) nig rcnasm<n§TI 3 EdlmanragCaGkSr L ¬rUbTI 24-8¦ . Qut CCA RtUv)aneKXIjmanenAnwgcug 3’én tRNA . bgÁúM 3’─OH rbs;nuyekøGUTIdcug eK (termial nucleotide) Adenosine GacRtUv)anP¢ab;tamry³sm<½n§kUva:Lg;CamYynwgGasIutGamIneN . nuyekøGUTId 3 EdlenAEpñk)aténcugsnøwk tRNA begáIt)anCa Anticodon . eyIgnwgBiPakSaBisþarenAkñúgEpñk 24-6 RtIKuNén)at (triplet of base ) begáItsm<½n§GIuRdUEsnCamYynwg Codon ¬QutbMeBjKñaén)as¦ enAelI m:UelKul mRNA enAelIépÞrbs; ribosome kñúgry³eBlénkarsMeyaKRbUetGIun . karcgsm<½n§GIuRdUEsnrvag codon nig anticodon enHnaMmknUvGasIutGamIneNRtwmRtUveTAnwgTIkEnøg (site) énkarsMeyaKRbUetGIunenAkñúg TItaMgsmRsbkñúgkarbegáItExS peptide ¬rUbTI 24-9¦ .



rUbTI 24-8

rUbTI 24-9

x-karsMeyaK RNA (Transcription)

Transcription EdlRtUv)anbgðajenAkñúgrUbTI 24-10 RtUv)ansMrYledayGg;sIum RNA

polymerase . dMeNIrkarenHekIteLIgCabIdMNak;kar . dMNak;karTI 1 RtUv)aneKehAfaCa Iniation Cab;Tak; TgeTAnwgkarP¢ab; RNA polymerase eTAnwg specific nucleotide squence EdleKehAfaCa Promoter

enAkEnøgcab;epþImén gene . Gnþrkm μrbs; RNA polymerase enHCamYynwg specific promoter DNA

sequence GnuBaØati[ RNA polymerase sÁal;cMNucab;epþImsMrab;eFIV transcription kEnøgcab;epþIménEhSn (the beginning of gene) . vak¾kMNt;EdrnUvGIVEdl DNA strand nwgRtUv)anbkERbBnül; (transcribe) .



mindUceTAnwg DNA replication , Transcription begáIt complementary copy BIExSmYykñúgcMeNamExS TaMgBIrbs; DNA . dUcEdlva)anP¢ab;eTAnwg DNA , RNA polymerase EjkExSTaMgBIrbs; DNA dUecñHva Gac {Gan} Qut)asrbs; DNA . dMNak;karTI 2 karBnøÚtExS[Evg (elongation) cab;epþImenAeBl RNA polymerase {Gan} DNA

templet strand nig sMrYl polymerization énkarftcMLg complemetary RNA . CamYynwgral;CMhanénkarsMrYl RNA polymerase bBa¢Ún ribonucleotide eTA[cugénExS RNA EdlduHeLIg nig sMrYlkarbegáItnUv sm<½n§ 3’-5’phosphodiester rvag 5’phosphoryl group rbs; ribonucleotide

Edlnwgmkdl; CamYynig 3’hydroxyl group rbs; ribonucleotide éncugExSEdlkMBugduH . RbtikmμenHRtUv)anbgðajenAkñúg rUbTI 24-11 . dMNak;karTI 3 rbs; transcription KWCa karbBa©b; (termination) . RNA polymrase CYbnwg Qut bBa©b; (termination sequence) enAcugénEhSn nig beBa©jnUvm:UelKul RNA EdlbegáIt)anfμ I mYy .

rUbTI 24-10

K-karrcnabnÞab;BIkarsMeyaK RNA (Post-transcriptional processing of RNA)

enAkñúg)ak;etrI EdlCaBYk prokaryotes karbBa©b; (termination) beBa©jnUv mRNA eBjelj (mature mRNA) sMrab;karsMeyaKRbUetGIun (translation) . CakarBit GaRs½yedayBYk prokaryotes

minmanPñaséNVyU:EdlEjk DNA nig sIutU)aøs;[dac;BIKña karsMeyaKRbUetGIuncab;epþImyUrmuneBl mRNA

RtUv)anbBa©b; . cMeBaH Eucaryotes karsMeyaK RNA (transcription) begáItnUv primary transcript Edl RtUvEtTTYlrgnUvnUvkarEklMGy:ageRcInbnÞab;BIkarsMeyaKmuneBlvaRtUv)andwknaMecjeTAeRkAéNVy:UsMrab;eFIV karsMeyaKRbUetGIunenAkñúgsIutU)aøs; .



rUbTI 24-11

Eukaryotic primary transcripts TTYlrgnUvkarEklMGrbIbnÞab;BIkarsMeyaK RNA . vaTaMgenaHKWCa karbEnßmén 5’cap structure nig 3’poly(A) tail nig RNA splicing . cMeBaHkarEkrlMGrTI 1 Cap structure RtUv)anbEnßmedaysarGg;sIumeTAcug 5’rbs; primary

transcript . Cap structure ¬rUbTI 24-12¦ man 7-methyl-guanosine P¢ab;eTAnwgcug 5’rbs; RNA

edaysars<an 5’-5’trphosphate . Nucleotides BIrdMbUgrbs; mRNA k¾RtUvTTYlnUvkarP¢ab;bgÁúMemTIlEdr (methylated) . Cap structure RtUvkarsMrab;karsMeyaKRbUetGIunEdlmanRbsiTi§PaBrbs; final mature

mRNA .



rUbTI 24-12

karEklMGrTI 2 KWCakarbEnßmGg;sIumén poly(A) tail eTAelIcug 3’én transcript . Poly(A)

polymerase eRbI ATP nig sMrYl polymerization tamlMdab;BI 100-200 adenosine nucleotides enAelI cug 3’rbs; RNA . Pol(A) tail karBarcug 3’rbs; mRNA BIkarbMEbkedayGg;sIum nig ehtudUecñHry³eBl rs;enArbs; mRNA . karEklMGrTI 3 RNA splicing Cab;Tak;TgeTAnwgkardkecjénEpñkrbs; primary transcript Edl minEmnCakardak;kUdRbUetGIun (protein coding) . EhSnrbs;)ak;etrIbnþCab; RKb;QutnuyekøGUTIdrbs;EhSnRtUv )aneXIjenAkñúg mRNA . b:uEnþ karsikSaBIrcnasm<½n§rbs;EhSnrbs; Eukaryotes bgðajPaBxusKñaén PaBTak;Taj (fascinating difference) . Eukaryotic genes minbnþCab;KñaeT vaman Extra DNA

sequences enAkñúgEhSnTaMgenHEdlminmandak;kUd (encode) [ amino acid sequence mYyNasMrab; RbUetGIuneT . QutTaMgenHRtUv)aneKehAfaCa Intervening Sequence of Introns . Primary transcript

manTaMg Introns nig QutEdlkUd[RbUetGIun EdleKehAfaCa Exons . vtþmanrbs; Introns enAkñúg mRNA

nwgeFIV[minGacmandMeNIrkarénkarsMeyaKRbUetGIunedIm,sMeyaKnUvRbUetGIunRtwmRtUvmYy . ehtudUecñH BYkva (introns) RtUv)andkecj EdlkardkenHRtUv)aneFIVedaydMeNIrkar RNA splicing . enAeBlGñkRsm½y RNA splicing RtUvEteTogTat;xøaMgNas; . RbsinebIvadkeRcIneBk b¤ ticeBk RNA RtUv)andkecj mRNA nwgminGacGnuvtþnUvkUdRtwmRtUvsMrab;RbUetGIunEdr . ehtudUecñH vaman {sBaØa} (signals) enAkñúg DNA edIm,IkMNt;RBMEdnrbs; Introns . CaTUeTA sequence GpU eKeXIjmanenA intron’s

5’boundary nig sequence ApG eXIjenA 3’boundary .



karTTYlsÁal;énRBMEdndMN (splice boundaries) nig kareFIV[manesßrPaBénbNþúMEdltrYc (splicing

complex) RtUvkarCMnYyBIPaKl¥itEdleKehAfaCa spliceosomes . Spliceosomes ekIteLIgBIRbePTmYyén small nuclear ribonucleoproteins (snRNPs Ganfa “snurps”) . snRNP man small RNA rYmCa mYynwgRbUetGIun . RNA components rbs; snRNPs bMeBjKña (complementary) eTAnwgQutepSg²Edl Cab;Tak;TgeTAnwg splicing .

rUbTI 24-13

edaysarkarcgsm<½n§GIuRdUEsneTAnwgRBMEdndMN (splice boundary) b¤ intron sequences snRPNs

sÁal; nig dak;rYmKñanUvQutEdlCab;Tak;TgenAkñúgRbtikm μkat;t (splicing reactions) . mYykñúgcMeNamEhSnTI 1 rbs; eukaryote bgðajfaman introns KWCaEhSnsMrab; β-subunit rbs; Hb

mnusSeBjv½y ¬rUbTI 24-13¦ . enAelI DNA EhSnsMrab; β-hemoglobin manRbEvg 1200 nucleotides

b:uEnþmanEt 438 nucleotides EdlmanpÞúkB½t’manEhSnsMrab;RbUetGIun . QutEdlenAsl;RtUv)aneKeXIjenAkñúg introns BIrEdlman 116 nucleotides nig 646 nucleotides EdlRtUv)andkecjeday splicing muneBlman karsMeyaKRbUetGIun . vaRtUvmankarcab;GarmμN_fa intron FM EvgCag final β-hemoglobin mRNA ! cMeBaH EhSnEdlRtUv)aneKsikSa Introns RtUv)aneKCYbRbTHeXIjmancenøaHTMhMBI 50-20,000 nucleotides KitCaRbEvg nig GacmaneRcInenABaseBjénEhSn . ehtudUecñH Typical human gene GacmanBI 10-30 dg EvgCa Final mRNA . 5-kUdénEhSn (Genetic Code)

mRNA manpÞúk gentic code sMrab;RbUetGIun . b:uEnþ GIVeTAEdlCaRbePTénkUdenH ? enAkñúgqñaM 1954 George Gamow )anelIkeLIgfa GaRs½yedayvamanEtbYn {GkSr} enAkñúg DNA alphabet (A, T, G nig



C) nig GaRs½edayvaman 20 GasIutGamIneN genetic code RtUvEtmanBaküEdlekIteLIgy:agehacBI 3 GkSr EdlRtUvykBIkñúgcMeNam 4 én DNA alphabet . etIKat;eFIVkarsnñidæanrebobdUcemþc ? Kat;)anBicarNafakUd EdlBakümanBIrGkSr (code of two-letter words) ekIteLIgBIkarrYmKñaNamYyBIGkSrcMnYnbYnman {s½BÞsa®sþ} énBakücMnYnEt 16 Kt; (42) . m:üagvijeTot vamanEt 16 rebobKt;EdlxusKñakñúgkartMerob A, T, C nig G

rYmKñaedIm,IbegáItCa)asmYy (AA, AT, AC, AG, TT, TA, TA, etc) . karenHvaminRKb;RKan;kñúgkardak;kUd [)anRKb;cMeBaHGasiIutGamIneNTaMg 20 eT . kUdEdlmanbYnGkSrpþl;nUvBakücMnYn 256 (44) eRcInq¶ayBItMrUv kar . b:uEnþ kUdEdlmanbIGkSrpþl;s½BÞsa®sþcMnYn 64 Bakü (43) RKb;RKan;kñúgkarpþl;kUdcMeBaHGasIutGamIneN 20 b:uEnþminelIsxøaMgeBkeT . karBiesaFn_mYyes‘rId¾l¥bgðajfa Gamow BitCaRtUv edaykarbgðajfa genetic code KWCa triple

code . brivtþn_ (mutations) RtUv)aneKbgðajenAkñúg DNA rbs; Bacterial virus . brivtþn_enHRtUv)anbBa©Úl ¬b¤ lubecjBI¦ nuyekøGUTId mYy BIr b¤ bIeTAkñúgEhSn . bnÞab;mkGñkRsavRCavBinitüRbUetGIunEdldak;kUdeday EhSnenaH . enAeBlnuyekøGUTId mYy b¤ BIr RtUv)anbBa©Úl minmanRbUetGIunRtUv)anbegáIteT . b:uEnþenAeBl)asTI 3 RtUv)anbBa©Úl Gtßn½y (sense) rbs; mRNA RtUv)anCYsCul (restore) nig RbUetGIunRtUv)anplit . GñkGac Rsm½ykarBiesaFn_enHedaykareRbIXøaEdlekIteLIgBIBaküEdlmanbIGkSr ]TahrN_ ³ THE CAT RAN OUT

etImanGIVekIteLIgcMeBaHGtßn½yrbs;XøaRbsinebIeyIgbBa©ÚlGkSrmYy ? THE FCA TRA NOU T

karGanénXøaenHRtUv)anxUcxat nig Xøa\LÚvenHElgmann½yeTotehIy . etI \LÚvenHeyIgGacCYsCul XøaenHedaykarbBa©ÚlGkSrTI 2 Edrb¤eT ? THE FAC ATR ANO UT

eT \LÚvenHeyIgminGacCYsCulGtßn½yrbs;XøaenH)aneT . Caf μ ImþgeTot eyIg)aneFIV[xUcxaténkar GanXøa b:uEnþGaRs½yedaykUdrbs;eyIgmanBaküEdlmanbIGkSr XøaenAEtminmanGtßn½y . RbsinebIeyIg bBa©Úl GkSrTI 3 vanwgCYsCulkarGanenH ³ THE FAT CAT RAN OU T

tamkarBit edaysarkarbBa©ÚlGkSrbIfμ IeTot eyIg)anCYsCulnUvGtßn½yrbs;saredaysarkarCYsCul karGan (reading frame) . karenHvaKWCarebobc,s;las;Edlsarrbs; mRNA RtUv)anbkRsay . RkumnimYy²EdlmanbInuyekøGUTIdenAkñúgQutrbs; mRNA RtUv)aneKehAfaCa codon nig kUdnimYy²sMrab;Et GasIutGamIneNmYyKt; RbsinebIQutenHRtUv)ankat;pþac; b¤ pøas;bþÚr vaGacpøas;bþÚrsmasPaBrbs;GasIutGamIn-eNrbs;RbUetGIun b¤ bNþal[karpliténRbUetGIunminekIteLIgTal;EtesaH . dUcEdleyIg)ankt;sMKal; genetic code EdlmanbIGkSrmanBakücMnYn 64 EdleKehAfaCa Codon

b:uEnþvamanEt 20 GasIutGamIneN . ehtudUecñH vaman codons 44 eRcInCageKRtUvkaredIm,IbBa¢ak;RKb; GasIutGamIneNenAkñúgRbUetGIun . Codons bIkñúgcMeNam codons TaMgGs; (UAA, UAG nig UGA) bBa¢ak;BI sBaØaénkarbBa©b; (termination signals) cMeBaHdMeNIrkarsMeyaKRbUetGIun (translation) . b:uEnþ karenHenAEt



Tuk[eyIgnUv 41 codons eTot . etIGIVeTACamuxgarrbs; “extra” code words ? Crick )anesñIeLIgfa genetic code KWCa degenerate code . Baküfa degenerate RtUv)aneKeRbIedIm,Ibgðajfa triplet codons

xusKñaGaceRbICa code words sMrab;GasIutGamIneNdUcKña . Complete genetic code RtUv)anbgðajenAkñúgrUbTI 24-14 . eyIgGaceFIVkarsegátCaeRcInGMBI genetic code . TI 1 Methionine nig Tryptophan KWCaGasIutGamIneNEdlman codon EtmYyKt; . GasIutGamIneNTaMgGs;epSgeToty:agehack¾man codons BIrEdrsMrab;GasIutGamIneNnimYy² nig serine nig leucine man codons 6 sMrab;vanimYy² . genetic code k¾GacFn;nwgbrivtþn_bnþicbnþÜcEdr (mutation-

resistant . cMeBaHGasIutGamIneNTaMgenHEdlman multiple codons Cajwkjab;)asBIxagedImdUcKña nig dUecñH nig bBa¢ak;GtþsBaØaNGasIutGamIneN nig manEtTItaMgTI 3 ERbRbYlxusKña . brivtþn_ (mutation) ¬karpøas;bþÚr enAkñúgQut)as¦ enAkñúgTItaMgTI 3 Cajwkjab;minman\T§iBleTAelIGasIutGamIneNEdlRtUv)aneKsmahr½nkm μ cUleTAkñúgRbUetGIuneT .

rUbTI 24-14



6-karsMeyaKRbUetGIun (Protein Synthesis)

dMeNIrkarénkarsMeyaKRbUetGIunRtUv)aneKehAfaCa Translation . vaCab;Tak;TgeTAnwgkarbkERb B½t’manEhSnBIQutrbs;nuyekøGUTIdeTACaQutGasIutGamIneNenAkñúgrcnasm<½n§TImYyrbs;RbUetGIun . karsMeyaK RbUetGIun (Translation) RtUv)anGnuvtþenAelI Ribosomes EdlCabNþúMén ribosomal RNA (rRNA) CamYy nwgRbUetGIun . Ribosome nimYy²ekIteLIgBIBIr subunits : a small and a large subunit ¬rUbTI 24-15a¦. enAkñúg eukaryotic cells, small ribosomal subunit man rRNA mYym:UelKul nig ribosomal proteins

33 xus²Kña nig large ribosomal subunit man rRNA bIm:UelKul nig RbEhl 49 énRbUetGIunepSg² . karsMeyaKRbUetGIunCab;Tak;TgeTAnwgGMeBIekIteLIgdMNalKñarbs; ribosomes CaeRcInenAelIm:UelKul mRNA EtmYyKt; . bNþúMTaMgenHén ribosomes CaeRcInenAtambeNþay mRNA EtmYyKt;enHRtUv)aneKsÁal; faCa Polyribosomes b¤ Polysomes ¬rUbTI 24-15b¦ . Ribosome nimYy²sMeyaKvtßúcMLgmYyén m:UelKulRbUetGIunEdldak;kUdeday mRNA . ehtudUecñH vtßúcMLgCaeRcInénRbUetGIunRtUv)anekIteLIgdMNalKña .

rUbTI 24-15

k-tYnaTIrbs; Transfer RNA (Role of Transfer RNA)

Codons rbs; mRNA RtUvEt)anGan RbsinebI genetic message nwgRtUv)anbkERbeTACaRbUetGIun . m:UelKulEdlbkRsayB½t’mankUdenAkñúg mRNA eTACarcnasm<½n§TImYyrbs;RbUetGIunKWCa transfer RNA

(tRNA) . edIm,IbkRsay genetic message eTACa primary sequence of protein, tRNA RtUvEtbMeBj muxgaredayesμaHsμRKBIr .



muxgarTI 1 tRNA RtUvEtP¢ab;CamYynwgGasIutGamIneNmYytamry³sm<½n§kUva:Lg; nig GasIutGamIneN BiessEtmYyKt; (only one, specific amino acid) . vamany:agehac tRNA mYysMrab;GasIutGamIneN nimYy² . dUcEdl)anbgðajenAkñúgrUbTI 24-8 RKb;m:UelKul tRNA manQut CCA enAcug 3’rbs;BYkva . enH KWCakEnøgEdlGasIutGamIneNnwgRtUvP¢ab;tamsm<½nÐkUv:aLg;eTAnwgm:UelKul tRNA . tRNA nimYy²RtUv)anTTYl sÁal;tamlkçN³Biesseday active site rbs;Gg;sIumEdleKehAvaCa Aminoacyl tRNA synthetase . Gg;sIumenHk¾TTYlsÁal;GasIutGamIneNRtwmRtUv nig cgGasIutGamIneNedaysm<½n§kUva:Lg;eTAnwgcug 3’rbs; tRNA . rUbTI 24-16a bgðajBIkarTTYlsÁal;GasIutGamIneN Methionine nig tRNA rbs;vaedaysar Methionyl tRNA synthetase . rcnsm<½n§EdlTTYl)anRtUv)aneKehAfaCa Aminoacyl tRNA enAkñúg krNIenH Methionyl tRNA . enAkñúgrUbTI 24-16b RbtikmμEdlbNþal[mankarP¢ab;énbgÁúM aminoacyl eTA nwg tRNA EdlRtUv)anbgðaj . GasIuGamIneNEdlcgP¢ab;eTAnwg tRNA nwgRtUvbBa¢ÚnBI tRNA eTA[ExS polypeptide enAkñúgry³eBlénkarsMeyaKRbUetGIun . muxgarTI 2 tRNA RtUvEtGacsÁal; Codon RtwmRtUvenAelI mRNA RtUvkarGasIutGamIneNenaH . karenHRtUv)ansMrYltamry³QutEdlmanbI)as (sequence of three bases) EdleKehAfaCa anticodon

EdlsßitenAnwg)atrbs; tRNA cloverleaf ¬emIlrUbTI 24-8¦ . Anticodon sequence sMrab; tRNA

nimYy²enHRtUvbMeBjeTAnwg codon EdlmanenAelI mRNA EdlbBa¢ak;BIlkçN³Biessrbs;GasIutGamIneN mYy . dUcEdlGñk)aneXIjenAkñúgrUbTI 24-9 sm<½nGIuRdUEsnénPaBbMeBjKña anticodon-codon nwgnaMmknUv GasIutGamIneNRtwmRtUveTAnwgTIkEnøgénkarsMeyaKRbUetGIun .

rUbTI 24-16

x-dMeNIrkarénkarsMeyaKRbUetGIun (The Process of Translation)

1. karpþÜcepþIm (Initiation)

dMNak;karTImYyénkarsMeyaKRbUetGIunKW karpþÜcepþIm (initiation) . RbUetGIunEdleKehAfaCa Initiation factors RtUvkaredIm,IsMrYlkarebegáItnUv translation complex EdlekIteLIgBIm:UelKul mRNA,

small ribosomal subunits nig large ribosomal subunits nig initiator tRNA . initiator tRNA enH sÁal; codon AUG nig dwknaM methionine EdlCaGasIutGamIneN.



Ribosome manTIkEnøgBIsMrab;cgP¢ab;CamYynwgm:UelKul tRNA . kEnøgTI 1 eKehAfaCa Peptidy

tRNA binding site (P-site) cgP¢ab; peptidyl tRNA edaybegáInnUvkarcg peptide eTAnwgm:UelKul tRNA . kEnøgTI 2 eKehAfaCa Aminoacyl tRNA binding site (A-site) cab; aminoacyl tRNA

edaydwkGasIutGamIneNbnÞab;eTAbEnßmelIExS peptide . m:UelKul tRNA nimYy² cgtamry³sm<½n§GIuRdUEsn eTAnwg mRNA eday codon-anticodon complematary . complex TaMgmUlRtUv)aneFIV[manesßrPaBEfm eTotedaykarBitEdlfa mRNA k¾RtUv)ancgP¢ab;eTAnwg Ribosome Edr . rUbPaB 24-17 a bgðajBIes‘rI énRBwtþikarN_EdlbNþal[mankarbegáItnUv initiation complex . initiator methionyl tRNA TTYlbnÞúk enA P-site enAkñúgbNþúMenH .

rUbTI 24-17



2. kareFIVlUtRbEvgExS (Chain elongation)

dMNak;karTI 2 én translation KW chain elongation . karenHekIteLIgCabIdMNak;EdrekItsareLIj vijrhUtdl;karsMeyaKRbUetGIun)anbBa©b; . eyIgcUleTAkñúgGMeBIenHbnÞab;BI tetrapeptide RtUv)anekItrYcehIy nig peptidyl tRNA ftcMLgnUv P-site ¬rUbTI 24-17b¦ . RBwtþkrN_dMbUgKWCakarcgén aminoacyl-tRNA molecule eTAnwg A-site EdlTMenr . bnÞab;mk karbegáItsm<½n§ peptide ekIteLIg . karenHRtUv)ansMrYledayGg;sIummYyenAelI ribosome EdleKehAfaCa Peptidyl transferase . \T\LÚv peptide chain RtUvpøas;eTA[ tRNA EdlftcMLg A-site . cugbB©ab; tRNA enAkñúg P-site Føak;eTAq¶ay nig ribosome pøas;bþÚrTItaMg dUecñH codon bnÞab;enAelI mRNA ftcMLg A-site . clnarbs; ribosome enHRtUv)aneKehAfaCa Translation . dMeNIrkarenHpøas;bþÚr peptidyl

tRNA f μ IBI A-site eTA P-site . Chain elongation én translation RtUvkarkareFIV hydrolysis én GTP

eTACa GDP nig Pi . Elogation factors CaeRcInk¾Cab;Tak;TgenAkñúgdMeNIrkarenHEdr . 3. karBa©b; (Termination)

dMNak;cugeRkaybMputén translation KWCa Termination . vaman Termination codons bI (UAA, UAG, nig UGA) Edlminman tRNA molecules RtUvKña (no corresponding tRNA

molecules). enAeBl “stop” codon mYykñúgcMeNamBYkvaRtUvCYb Translation RtUv)anbBa©b; . Release

factor cgP¢ab;eTAnwg A-site TMenr . Peptidyl transferase EdlsMrYlBImunmknUvkarbegáItsm<½n§ peptide eFIV hydrolyze sm<½n§ ester rvag peptidyl tRNA nig amino acid cugeRkayrbs;RbUetGIunsMeyaKehIyf μ I² ¬rUbTI 24-17c¦ . enAnwgcMNucenH tRNA ¬peptide EdleTIbsMeyaKfμ I¦ nig ribosomal subunits BIrRtUv)an beBa©j . Peptide EdlRtUv)anbeBa©jbnÞab;BI translation minmansar³sMxan;caM)ac;enAkñúgTMrg;bMeBjmuxgar rbs;vaeT . enAkñúgkrNImYycMnYn peptide enHRtUv)ankat;edayGg;sIum proteolytic muneBlvaGacbMeBaj muxgar)an . karsMeyaKGg;sIuménRbdab;rMlayGahareRbIyuT§sa®sþenH . CYnkalRbUetGIunenHRtUv)anrYmCamYy peptides epSgeTotedIm,IbegáItCaRbUetGIunEdlGacbMeBjmuxgar (functional protein) dUcCakñúgkrNI hemoglobin . Cellular enzymes bEnßmbgÁúM carbohydrates b¤ lipids eTAelIRbUetGIunmYycMnYnCaBiess RbUetGIunEdlcugrbs;vabBa©b;enAelIépÞekasika . karERbRbYlcugeRkayenHmanlkçN³BiesscMeBaHRbUetGIun BiessmYy nig dUcCaQuténRbUetGIunxøÜnvaRtUv)ankMNt;TisedAeday cellular genetic information . 7-brivtþn_ kaMrsμIB’NsVay nig karCYsCul DNA

(Mutation, Ultraviolet Light, and DNA Repair)

k-RbePTénbrivtþn_ (The Nature of Mutations)

karpøas;bþÚrGacekIteLIgenAkñúgQut nucleotides rbs; DNA molecule . karpøas;bþÚrénEhSnrebob hñwgRtUv)aneKehAfaCa Mutation . Mutations KWCaGacelcecjBIkMhusEdleFIVeLIgeday DNA



polymerase enAkñúgry³eBlén DNA replication . BYkvak¾GacbNþalmkBIGMeBIénCatiKImIEdleKehAfaCa Mutagens EdleFIV[xUcxat DNA . Mutations GacRtUv)aneKeFIVcMNat;fñak;eTAtamRbePTénkarpøas;bþÚrEdlekIteLIgenAkñúg DNA . karCMnYsén Nucleotide EtmYyKtcMeBaH nucleotide mYyRtUv)aneKehAfaCa point mutation ³ ATGGACTTC Normal DNA sequence ATGCACTTC Point mutation

CYnkal nucleotide mYy b¤ sUm,IEtEtEpñkeRcInén DNA RtUv)an)at; . karenHRtUv)aneKehAfaCa deletion mutation : ATGGACTTC Normal DNA sequence ATGTTC Deletion mutation

mþgmáal nucleotide mYy b¤ eRcInRtUv)anbEnßmeTAelI DNA sequence . karenHRtUv)aneKehAfaCa insertion mutation : ATGGACTTC Normal DNA ATGCTCGCTTC Insertion mutation

x-lT§plénbrivtþn_ (The Results of Mutations)

brivtþn_ (Mutations) mYycMnYnKWCa brivtþn_es¶óms¶at; (silent mutation) dUecñH BYkvaminbNþal[man karpøas;bþÚrenAkñúgRbUetGIuneT . b:uEnþCajwkjab; brivtþn_man\T§iBlGviC¢maneTAelIsuxPaBrbs;srIragÁ . \T§iBl rbs;brivtþn_GaRs½yeTAnwgfaetI genetic code sMrab;RbUetGIunxUcxaty:agdUcemþc . cUrBicarNaelI codons BIr sMrab; Glutamic acid : GAA nig GAG . brivtþn_enAnwgcMNuc (point mutation) EdleFIV[xUc nucleotide

TI 3 én GGA eTACa GAG nwgenAEtbNþal[mansmahr½Nkmμén glutamic acid enAkñúgTItaMgRtwmRtUv rbs;RbUetGIun . dUcKñaenHEdr karpøas;bþÚrBI GAG eTACa GAA k¾sßitenAes¶óms¶at;Edr . vaman human genetic diseases RbmaN 4000 CMgWEdlbNþal;mkBIbrivtþn_ . karenHekIteLIg GaRs½yedaybrivtþn_enAkñúg DNAEdlpøas;bþÚr codon nig bNþal[mankareFIVsmahr½Nkmμén amino acid

xuscUeTAkñúgRbetGIun . karenHbgá[RbUetGIunminGacbMeBjmuxgar b¤ bMeBjmuxgarmin)anRtwmRtUv . cUrBicarNaBI human genetic disease KW sickle cell anemia . enAkñúg normal β-chain rbs; hemoglobin amino acid cMnYn 6 KWCa glutamic acid . enAkñúg β-chain rbs; sickle cell , amino acids

TaMg 6 enaHKWCa Valine . etIkarCMnYs amino acid enHekIteLIgrebobdUcemþc ? cMeLIyCab;Tak;TgeTAnwgkar Binitüén codons sMrab; glutaminc acid nig valine ³ Glutamic acid : GAA b¤ GAG

Valine : GUG, GUC, GUA b¤ GUU

Point mutation BI A→U enAelI nucleotide TI 2 pøas;bþÚr codons mYycMnYnsMrab; glutaminc

acid eTACa codons sMrab; valine ³ GAA → GUA GAG → GUG Glutaminc acid Valine codon codon



Mutation enAkñúg codon EtmYyKt;enHbNþal[maNkarpøas;bþÚrQut amino acid enATItaMgTI 6 enA kñúg β-chain én hemoglobin rbs;mnusSBI glutamic acid eTACa valine . lT§plénkarpøas;bþÚrhakdUcCa tictYcKWCa sickle cell anemia enAkñúgbuKÁlEdlTTYlmrtknUvvtßúcMLgBIrén mutant gene . K-sarFatueFIV[manbrivtþn_ nig sarFatueFIV[manmharIk (Mutagens and Carcinogens)

CatiKImIEdlbgá[mankarpøas;bþÚrenAkñúgQut DNA (DNA sequence) RtUv)aneKehAfaCa Mutagens . Cajwkjab; mutagens k¾GacCa Carcinogens Edr ¬CatiKImIEdlbgá[manCMgWmharIk¦ . PaKeRcInénCMgWmharIkbNþalmkBI mutations enAkñúgekasikaFmμtaEtmYyKt; . mutations TaMgenHbNþal[ mankar)at;bg;nUvkarRtYtBinitükarlUtlas;Fmμta edaybgá[manekasikaminFmμtaEbksayPay (prolife-

rate). RbsinebIkarlUtlas;enaHminRtUv)anRtYtBinitü b¤ RtUv)anbMpøaj vanwgbNþal[søab;buKÁlEdlman CMgWenH. eyIgRtUv)anb:HeTAnwg carcinogens CaeRcInenAkñúgry³eBlénkarrs;enArbs;eyIg . CYnkaleyIgb:H eTAnwg carcinogen edayécdnü b:uEnþenAkñúgkrNImYycMnYnvaRtUv)anb:HedaykareRCIserIs . vamanRbmaN smasFatuKImI 3000 muxenAkñúgEpSg)arI nig sarFatuCaeRcInCa mutagens EdlmansukþanuBl . CalT§pl mnusSEdlCk;)arIman»kasnwgekItCMgWmharIksYteRcInCagGñkEdlminCk;)arI . X-karxUcxatedaysarkaMrsμIB’NsVay nig karCYsCul DNA

(Ultraviolet Light Damage and DNA Repair)

BnøWB’NsVay (UV) KWCaPñak;mYyepSgeTotEdlbgákarxUcxatcMeBaH DNA . karRsUbénBnøWB’NsVay eday DNA bgá[man adjacent pyrimidine bases eTAcgP¢ab;Kñatamsm<½n§kUv:aLg; . plitpl ¬rUbTI 24-18¦ RtUv)aneKehAfaCa Pyrimidine dimer . CalT§pl karbegáItCa Pyrimidine dimer vaminmankarcg sm<½n§GIuRdUEsnrvag pyrimidine molecules TaMgenH nig complementary bases enAelI DNA strand epSg eTot . karBnøatén DNA enHminGaceFIV replication b¤ transcription )aneT ¡ )ak;etrIdUcCa Escherichia coli manynþkarxus²KñabYnkñúgkarCYsCulkarxUcxatedaykaMrs μ IB’NsVay. b:uEnþ sUm,IEtdMeNIrkarCYsCulGaceFIV[mankMhusk¾eday . Mutations ekIteLIgenAeBlRbB½n§CYsCul karxUcxatedaykaMrs μIB’NsVayeFIV[mankMhus nig bgá[mankarpøas;bþÚrenAkñúgQut nucleotide rbs; DNA . enAkñúgevC¢sa®sþ Rbtikmμ pyrimidine dimerization RtUv)aneKeRbIedIm,ICaRbeyaCn_enAkñúgmnÞIreBTü EdlenATIenaH germicidal (UV) light RtUv)aneKeRbIedIm,IsMlab;)ak;etrIenAkñúgxül; nig enAelIépÞCMuvij dUcCa bnÞb;sMrab;vHkat;CaedIm,I . karsøab;rbs;ekasikaenHRtUv)anbgáedaykarbegáIt pyrimidine dimer enAkñúgTMhM eRcIn . RbB½n§CYsCulrbs;)ak;etrIRtUv)anrMCylxøaMg nig ekasikasøab; . GBa©wg RbePTdUcKñaénkarbegáIt pyrimidine dimer GacekIteLIgcMeBaHekasikamnusSEdr . kardak; [enAxageRkAb:HnwgBnøWRBHGaTitüral;éf¶edIm,ITTYl)anEs,kmanB’NdaMEdg (fashionable tan) eFIV[Es,kb:H eTAkaMrsμ IB’NsVayy:ageRcIn . karenHeFIV[xUcEs,kedaykarbegáItnUv Pyrimidine dimers CaeRcIn . kardak; eTAnwgkaMrs μ IB’NsVaykMritx<s;BIBnøWRBHGaTitü b¤ k¾kareFIV[Es,keLIgB’NenAkNþalval)anCab;Tak;TgeTAnwg eRKaHfñak;énCMgWmharIEs,kenAkñúgcMeNamRbCaCnTaMgenaH .



g-plvi)akénPaBex©aHenAkñúgkarCYsCul DNA (Consequences of Defects in DNA Repair)

RbB½n§CYsCulrbs;mnusScMeBaH pyrimidine dimers s μ úRKs μajxøaMgNas; EdlRtUvkary:agehacGg;sIum 5 . CMhanTI 1 kñúgkarCYsCul pyrimidine dimer KWkarkat; sugar-phosphate backbone rbs; DNA Edl enAEk,rTIkEnøgxUcxat . Gg;sIumEdleFIVkarkat; sugar-phosphate backbone enHRtUv)aneKehAfaCa repair

endonuclease . RbsinebIEhSnEdlftcMLgGg;sIumenHmanPaBex©aH (defective) pyrimidine dimer

minGacCYsCul)aneT . karekIneLIgén mutations rYmCamYynwgkarfycuHRBmKñaRbsiT§PaBénynþkarCYsCul DNA rbs;ekasikabNþal[mankarekIneLIgnUv incident of cancer . ]TahrN_ mutation én repair

endonuclease gene nig enAkñúg genes énpøÚvCYsCulepSgeTot (repair pathway) bNþal[man genetic

skin disorder EdleKehAfa xeroderma pigmentosum . RbCaCnEdlTTYlrgBI xeroderma

pigmentosum rYsy:agxøaMgeTAnwgkaMrs μ IB’NsVayénBnøWRBHGaTitü nig ekIt multiple skin cancer CaFmμta munGayu 20 qñaM . 8-DNA Edlbgáat; (Recombinant DNA)

k-]bkrN_eRbIenAkñúgkarsikSa DNA (Tools Used in the Study of DNA)

GñkviTüasa®sþCajwkjab;RtUv)aneKecaTsYrfaehtuGIV)aneKsikSaRbFanbTEdlTMngCaminmansar³sMxan; dUcCa bacterial DNA replication . ehtuplmYyd¾l¥KWfakarsikSarebobhñwgCajwkjab;pþl;nUvkaryl;c,as;-las;eTAkñúgkargarén human genetic systems . TIBIr KWfakarRsavRCavCajwkjab;plitnUv]bkrN_Edl GnuBaØati[elatcUleTAkñúgbec©kviTüaf μ I . \LÚv karenHvaBitxøaMgCagenHenAkñúgkarGPivDÆ recombinant DNA

technology . PaKeRcInénbec©keTs nig ]bkrN_ (tools) EdleKeRbIenAkñúg recombinant DNA studies

RtUv)anbegáIt nig RsavRCaveXIjenAkñúgkarsikSamUldæan (basic studies) eTAelI bacterial DNA

replication nig gene expression . karTaMgenHrYmbBa©ÚlGg;sIumCaeRcInEdlsMrYlRbtikmμrbs; DNA

molecules , gel electrophoresis, cloning vectors nig hybridization techniques . 1. Restriction Enzymes

Restriction enzymes KWCa bacterial enzymes Edl {kat;} sugar-phosphate backbone rbs; DNA molecules enAnwg specific nucleotide sequence . Gg;sIumTI 1 kñúgcMeNamGg;sIumTaMgenHEdlRtUv )aneKbnSúT§ nig sikSaRtUv)aneKehAfa EcoR1 . eQμaHmanRsLayBIeQμaH genus nig species rbs;)ak;etrI EdlGg;sIumenHRtUv)ancMraj;ecjmk enAkñúgkrNIenHKW E. coli . xageRkamenHKWCa specific nucleotide

sequence EdlRtUv)anTTYlsÁal;eday EcoliR1 . 5’................................. GAATTC.................................3’ 3’................................. CTTAAG.................................5’

enAeBl EcoR1 kat; DNA enATIkEnøgenH vakat;enAkñúgrebobbBaäit (staggered fashion) . eday manlkçN³Biess vakat;rvag G nig A TImYyenAelI strands TaMgBIr . karkat;begáIt)anCa DNA BIrkMNat; Edlmanrcnasm<½n§dUcxageRkam ³



5’................................. G AATTC.................................3’ 3’................................. CTTAA G..............................5’

cugbBaäitTaMgenH (staggered termini) RtUv)aneKehAfaCa Sticky ends GaRs½yedayBYkvaGacrYmKña CamYynwgcugmYyepSgeTotedaykarsm<½n§GIuRdUEsn . karenHKWCalkçN³rbs; DNA fragments EdlbegáIt eday restriction enzymes Edlvamansar³sMxan;xøaMgNas;cMeBaH gene cloning . ]TahrN_én restriction enzymes epSgeTot nig specific recognition sequences

rbs;BYkvaRtUv)anerobrab;enAkñúgtaragTI 24-3 . TIkEnøgenAelI sugar-phosphate backbone EdlRtUv)an kat;edayGg;sIumTaMgenHRtUv)anbgðajedaysBaØa “/” . Gg;sIumTaMgenHRtUv)aneKeRbIedIm,Ikat; DNA molecules FM²[eTACakMNat;tUc² (smaller

fragments) énTMhMBiess (specific size) . GaRs½yeday restriction enzyme CaTUeTAkat;enATIkEnøgdUcKña DNA EdlTTYl)anBIbuKÁlBiessmYybegáItnUv reproductive set of DNA fragments . karenHsmRsb sMrab;karsikSa b¤ karcMLg DNA BIRbPBNamYy (cloning of DNA) .

taragTI 24-18

2. Agarose Gel Electrophoresis

meFüa)aymYyénkarsikSaBI DNA fragmentsEdlbegáIteday restriction enzyme digestion KW Agarose gel electrophoresis . Digested DNA sample RtUv)andak;eTAkñúg sample well enAkñúg gel

nig crnþGKÁisnIRtUv)aneKtP¢ab; . GaRs½ybnÞúkGviC¢manén Phosphoryl groups enAkñúg sugar-phosphate

backbone , DNA fragments eFIVclnatamry³ gel ecjBI negative electrode (cathode) eq<aHeTA positive electrode (anode) . Smaller DNA fragments eFIVclnaelOnCag larger DNA fragments

nig CalT§pl DNA fragments RtUvray)a:yCacugbBa©b;enABaseBj gel eyageTAtamTMhMrbs;va . TMhMrbs; fragment nimYy²GacRtUv)ankMNt;edaykareRbóbeFobCamYynwgKMrUénkarpøas;TItaMg (migration pattern)



rbs; DNA frag,ments EdleKsÁal;TMhM . Agarose gel Edlbgðaj DNA bands EdlmanTMhMxus²KñaRtUv )anbgðajenAkñúgrUbTI 24-19 .

rUbTI 24-19

3. Hybridization ¬karbgáat;BUC¦ Agarose gel electrophoresis GnuBaØati[kMNt;BITMhMén DNA fragment . b:uEnþenAkñúg recombinant DNA research vak¾mansar³sMxan;kñúgkarkMNt;GtþsBaØaNfaetI gene NaRtUvGnuvtþeday particular DNA fragments . Hybridization KWCabec©keTsmYyRtUv)aneKeRbIedImI,kMNt;vtþmanrbs; gene enAelI particular

DNA fragment . bec©keTsenHsMGageTAelIkarBitEdlfa complementary DNA sequences nwgcg sm<½n§GIuRdUEsn b¤ bgáat;BUC (hybridize) CamYynwg DNA sequence mYyepSgeTot . vaminmanbBaðaeT faetI complementary single-stranded sequence elcecjBI DNA molecule b¤ BI DNA molecules ecjBI srIragÁBIRbePTxusKña . RbsinebI DNA sequences bMeBjKña (complementary) BYkvanwgbgáat;BUC (hybridize) CamYynwgsrIragÁmYyepSgeTot . CakaBit sUm,IEt RNA GacRtUv)aneKeRbIkñúgkareFIV hybridization studies . RNA Gacbgáat;CamYy nwg DNA molecules b¤ eTAnwg RNA molecules epSgeTot . Cafμ ImþgeTot lkçN³vinicä½yEtmYyKt;cMeBaH hybridization KWvtþmanén complementary nucleic acid sequences enAelI strands TaMgBIr . bec©keTsmYy EdleKehAfaCa Southern Blooting Cab;Tak;TgeTAnwg hybrization én DNA

fragments EdlTTYl)anBI agarose gel ¬rUbTI 24-20¦ . CMhanTI 1 énnitiviFIKWCakarbMEbk DNA eday sar restriction enzymes bnÞab;mk fragments TaMgenHRtUv)anEjk[ecjBIKñaeTAtamTMhMeday agarose

gel electrophoresis . bnÞab; DNA fragments RtUv)anbBa¢ÚnedaykareFIV[eLIgCaB’N (bloting) enAelI



special membrane filter . DNA molecules enAelI filter RtUv)an {rlay} eTACa single DNA strand

dUecñHBYkvaRtUv)anerobcMrYcCaeRscsMrab; hybridization . kartMrUvbnÞab;mkeTotcMeBaHnitiviFIenHKWCaRsavRCavrkEdlnwgkNt;rknUv large gene sequence Edl eKcgrk . sarFatusMrab;RsavRCav (probe) KWCa DNA b¤ RNA EdlRtUvBMBak;pøakedayviTüúskmμ EdlGacRtUv )aneKeRbIedIm,IkMNt;rk specific gene . enAeBlEdl probe RtUv)aneFIVeLIg vaRtUv)aneKdak;enAkñúg sUluysüúg[b:HeTAnwg membrane filter . Probe nwg hybridize EtCamYynwg DNA fragments TaMgenaH enAelI filter Edlman complentary sequence CamYynwgva edIm,IkMNt;rkfaetI DNA bans NaenAelI gelRtUv)anTTYlrgnUv hybridization, filter RtUv)aneKdak; X-ray film mYydMu . bnÞab;BIkardak;[b:HnwgBnøW nig karbegáIt (exposure and development) X-ray film

nwgbgðajCa black band b¤ bands EdlmanenAkEnøgNamYyEdl hybridaztion )anekIteLIg . enAkñúgrebob enH GñkviTüasa®sþGackNt;GtþsBaØaNfaetI DNA fragment mYyNaEdlman gene EdleKcgrk .

rUbTI 24-20



4. DNA cloning Vectors

DNA cloning experiments rYmbec©kviTüaTaMgenHCamYynwgl,icmYyb¤BIrbEnßmeTot edIm,Idkyk vtßúftcMLgEtmYyKt;én gene nig bnÞab;mkbegáItrab;ekadiénvtßúftcMLg (billions of copies) . edIm,IbegáIt vtßúftcMLgcMruHénEhSn (multiple copies of gene) vaGacRtUv)aneKP¢ab;CamYynwg cloning vectors . Cloning vectors KWCaEpñkén DNA Edlman replication origin rbs;vapÞal; dUecñHvaGaceFIV replication

enAkñúg host cell Cajwkjab; E. coli eRbICa host cell Edl vector man cloned DNA RtUv)aneFIV replication edayeRcIn . vamanBIRbePTsMxan;én cloning vectors . TImYyKWCa bacterial virus b¤ phage vectors . BYkva TaMgenHKWCa bacterial viruses EdlCaTUeTARtUv)anbMElgedIm,IGnuBaØati[mankarbEnßmén cloned DNA

fragments . Viruses TaMgenHminRtwmEtman origin of replication b:uEnþk¾man genes epSg²eTotTaMgGs; EdlRtUvkarsMrab;eFIV replication [)an 100-200 copies én virus ¬nig cloned fragment¦ kñúgmYy infected cell . TIBIrKWCa vector EdleKeRbIedayjwkjab;KWCa Plasmid vector . Plasmids KWCa extra piece rbs; circular DNA EdleKeXIjmanenAkñúgRbePTPaKeRcInbMputénBYk)ak;etrI . BYkvaman replication origin

rbs;BYkva b:uEnþKWCa replication enzymes rbs; host cell EdlTTYlxusRtUvkñúgkareFIV replication rbs; plasmids . Plasmids enAkñúgFmμCatiCajwkjab;man genes sMrab; antibiotic resistance . ehtudUecñH ekasika Edlman plasmids Fn; (resistant) eTAnwgGMeBImYy b¤ eRcInrbs; antibiotics . Plasmids RtUv)aneKeRbICa cloning vectors k¾RtUv)anbMElgedIm,ITTYlyk)annUv cloned DNA fragments nig Cajwkjab;man antibiotic resistance genes Edlmansar³sMxan;enAkñúgekasikaEdlman plasmid .

rUbTI 24-21



mYyén plasmid rebobhñwgKW pBR322 . karKUsbøg;rbs; restriction enzyme én pBR322 RtUv)an bgðajenAkñúgrUbTI 24-21 . Plasmids enHman antibiotic resistance genes BIr ³ mYysMrab; Ampicillin

nig mYyeTotsMrab; Tetracycline . enAkñúg antibiotic resistance genes man restriction enzyme sites

CaeRcInEdlsmrmüsMrab;kareFIV cloning . x-visVkm μénEhSn (Genetic Engineering)

PaKeRcInén]bkrN_ (tools) EdlRtUvkarcMeBaH cloning experiment RtUvEteRbIrYmKña karsMerccitþ bnÞab;RtUv)aneFIVeLIgKW gene mYyNanwgRtUvftcMLg (clone) . ]TahrN_ EdleyIgnwgeRbIKWkarftcMLgén β-

globin genes cMeBaH Hemoglobin Fmμta nig sickle cell hemoglobin . DNA Edl)anmkbuKÁlman normal hemoglobin RtUv)anbMEbkeday restriction enzyme RbEhlCa BamHI . vaenHKWCa Target

DNA . Vector DNA RtUvEt)anbMEBkedayGg;sIumdUcKña ¬rUbTI 24-22¦ . enAkñúg]TahrN_rbs;eyIg eyIg nwgeRbI plasmid vector pBR322 .

rUbTI 24-22

Digested vector nig target DNA RtUv)anlaycUlKñaeRkamlkçxNÐEdlCMruj[ BamHI sticky

ends rbs; target nig vector DNA edIm,IrYmKña (hybridize) CamYyKñaBImYyeTAmYy . GaRs½yeday sticky

ends RtUv)ancgP¢ab;Kñaedaysm<½n§GIuRdUEsnBIrb¤bI eyIgRtUvEtP¢ab; target nig vector DNA tamry³sm<½n§ kUv:aLg;muneBldak;BYkvaeTAkñúg bacterial cell . karenHRtUv)aneKGnuvtþedaygayRsYledaykareRbIGg;sIum



DNA ligase . DNA ligase KWCaGg;sIummYyepSgeTotEdlRtUv)aneKrkeXIjenAkñúgry³eBlkarsikSamUldæan én DNA replication nig DNA repair . vasMrYlkarbegáItnUvsm<½n§ phosphoester rvagEpñkTaMgBIrrbs; DNA . \LÚvdl;eBleyIgbBa©Úl recombinant DNA molecules eTAkñúg bacterial cell . meFüa)aymYy énkarbBa©ÚlenHKWCakarbMElg (transformation) dMeNIrkarEdl E. coli cells TTYlrgRbRBwtþkmμKImIeRkam lkçxNÐEdlmanTMenareTArkkarbBa©Úl DNA cUleTAkñúg cells . ekasikaénl,ayEdl)anbMElg (the cells of

transformation mixture) RtUv)aneFIVbnÞHenAelI solid nutient agar medium mYy . ekasikanimYy²nwg lUtlas;Ca colony énekasikaenAelI agar medium enH . KYr[GP½BV karbMElgenHminmanRbsiT§PaBxøaMgeT nig PaKeRcInénekasikaenAkñúgl,ayEdl)anbMElgminmanpÞúk plasmids eT . BYkvaTaMgenHeKminman karcab;GarmμN_eT etIRtUveFIVdUcemþcedIm,Idkvaecj ? enAeBlvaRtUv)andkecj CamYynwg pBR322 vagayRsYl xøaMgNas; . RbsinebIekasikaRtUv)anCabnÞHenAelI agar medium Edlman ampicillin RKb;ekasikaEdlmin TTYlyk plasmid nwgRtUv)ansMlab;eday antibiotic . karbgáat;BUC (hybridization) GacRtUv)aneKeRbIedIm,IkMNt;rk vtßúcMLg (clones) EdlmanpÞúk β-

globin gene . vtßúsÞg;viTüúskmμ (Radioctive probe) RtUv)aneKerobcMedaykarcMraj; β-globin mRNA

ekasikaRKab;QamRkhm . ekasikaTaMgenHman nucleic acid tictYcCag globin mRNA . β-globin

mRNA RtUv)anbMBak;pøakviTüúskmμ nig RtUv)aneKeRbICavtßúsÞg;edIm,IkMNt;rk colonies TaMgenaHEdlman desired cloned DNA . vtßúcMLg (replica) én experimental plate RtUv)aneFIVeLIgedaykarsgát; steril velveteen fabric

eTAelIépÞ . Velveteen Edlmanekasika)anBI original colonies RtUvsgát;eTAelIépÞén fresh agar plate

edaybegáIt)anCa precise replica . membrane filter RtUv)aneKdak;eTAelI colonies enAelI agar plate . ekasikaBI colony nimYy²nwgRtUv)anbBa¢ÚneTA membrane filter . ekasikaTaMgenH RtUv)anbMEbkedayfñm² edaykareFIVRbtikmμKImICamYynwg detergents nig Gg;sIum nig filter RtUv)anTTYlrgRbtikmμKImI dUecñH released

DNA RtUv)anP¢ab;eTAnwg membrane . enAeBlbgáat;BUC (hybridization) RtUv)anGnuvtþenAelI filters

TaMgenH vtßúsÞg;viTüúskmμnwgP¢ab; (hybridize) EtCamYynwg complementary sequences én β-globin

gene. enAeBl membrane filter RtUv)anbHeTAnwgkaMrs μ I X sñam (spot) nwgelcecjeLIgenA developed

film EtenATIkEnøgrbs; colony EdlmanpÞúk desired clone . edaykarRtLb;eTA original plate eyIgGac eRCIserIsekasikaBI colony TaMgenaH nig bNþúHekasikaTaMgenHsMrab;karsikSaeTAGnaKt ¬rUbTI 24-23¦ nitiviFIdUcKñaGacRtUv)aneKeRbIedIm,IftcMLg (clone) β-chain gene rbs; sickle cell hemoglobin. bnÞab;mkBYkvaTaMgBIrGacRtUv)aneKsikSa nig eRbóbeFobedIm,IkMNt;BIFmμCatién genetic defect . ]TahrN_EdlRsYlenHeFIV[vaeXIjfaRKb; gene cloning RsYlxøaMgNas; nig mins μ úRKs μaj . karenH RtUv)anbgðajeRcInCagq¶ayBIkarBit . visVkrEhSn (Genetic engineers) RtUvEtCMnHelI]bs½KÁTMagLaykñúgkar ftcMLg eukaryotic genes énplRbeyaCn_evC¢sa®sþBiessNamYy . ]bs½KÁmYykñúgcMeNam]bs½KÁTImYy



EdlCYbRbTHKWCavtþmanrbs; introns enAkñúg eukaryotic genes . )ak;etrIEdlRtUv)aneRbIsMrab;karftcMLg minman enzymatic machinery edIm,I dk introns ecj (splice out) . Molecular biologist rkeXIjfa DNA copy én eukaryotic mRNA GacRtUv)aneFIVeLIgedaykareRbI enzyme reverse transcriptase

BIGMbUrvirusEdleKehAfa Retroviruses . DNA copy én mRNA rebobhñwgmannUvRKb; protein-coding

sequences rbs; gene b:uEnþminman intron sequence eT . dUecñH )ak;etrIGac transcribe nig translate nUv cloned DNA nig begáItnUvplitplEdlmantMélsMrab;kareRbIenAkñúgevC¢sa®sþ nig kargarGnuvtþepSgeTot .

rUbTI 24-23

karenHKWCabBaðabec©keTsEtmYyKt;kñúgcMeNambBaðabec©keTsCaeRcInEdlRtUv)anCMnHeday karbegáIt EdleFIV[q¶l; (amazing development) enAkñúg recombinant DNA technology . esckþIsegçbb:uEnþ KYr[cab;GarmμN_rbs;plitplsMxan;evC¢sa®sþénvisVkmμEhSnRtUv)anbgðajenAkñúgtaragTI 24-4 .



eTaHbICavDÆnPaBya:gxøaMgRtUv)anekIteLIgenAkñúgkarGnuvtþvisVkmμénEhSncMeBaHbBaðevC¢sa®sþsMxan;² k¾eday b:uEnþPaKeRcInénplitplTaMgenHbgðajRtwmEtkarBüa)al b:uEnþmineFIV[CasHes,IyeT . karsubinþ cugeRkaysMrab;GnaKtKWedIm,IeFIVGnþraKmn_[)anedaydac;xat edIm,IdkecjnUvEhSnminl¥TaMgGs;ecjBImnusS . kardak;CMnYs mutant gene enAkñúg fertilized egg CamYynwg normal gene KWCadMeNaHRsaymYycMeBaH eKaledAenH . b:uEnþkarBiténsubinþenaHcMeBaHmnusSKWeRcInqñaMNas; RbEhlmYyTsSvtS . kñúgxN³enaHEdr molecular biologists bnþeFIVkargareq<aHeTAeKaledAenaH edaykarsikSarcnasm<½n§Fm μta nig expression én DNA nig edaykareFIVkargaredIm,IbegáItnUvRbB½n§beBa©jEdlmansuvtßiPaBmYysMrab;karcak;bBa©Úlén DNA f μI cUleTAkñúgekasika dUcCa fragile human fertilized egg . 9-POLYMERASE CHAIN REACTION (PCR)

)ak;etrIEdleKcMraj;BI hot spring enA Yellowstone National Park pþl;nUvKnøwHsMxan;cMeBaH molecular tool cMeBaHkarsikSaBI DNA . Polymerase Chain Reaction (PCR) GnuBaØati[GñkviTüa-sa®sþbegáItnUvcMnYncugbBa©b;énEhSnNamYyEdlmanRbeyaCn_ nig Bacterium Thermus aquaticus begáIt DNA polymerase EdlFn;nwgkMedA (Taq polymerase) EdlGnuBaØati[dMeNIrkarRbRBwtþeTA . Human genome manRbEhlbIBan;lanén base pairs rbs; DNA . b:uEnþ ]bmafaGñkcab;GarmμN_ kñúgkasikSaEtEhSnEtmYyKt; RbEhlEhSnEdlTTYlxusRtUvcMeBaH muscular dystrophy b¤ cystic

fibrosis. vahak;dUcCakarBinitüemIlm¢úlenAkñúgKMnrcMebIg . edayeRbI PCR GñkviTüasa®sþGacbegáItrab;lanén vtßúcMLgrbs;EhSnEdlGñkcgsikSa kñúgxN³EdlmineGIeBIcMeBaH 100,000-200,000 énEhSnepSgeTotenAelI human chromosomes . PaBGa’fkM)aMgcMeBaH specificity enHKWkarsMeyaKén DNA primer EpñkxøIén single-stranded DNA

Edlnwgbgáat;edaymanlkçN³BiesseTAnwgkarcab;epþIménEhSnBiessNamYy . DNA polymerase RtUvkar primer mYysMrab;karpþÜcepþIménkarsMeyaK DNA GaRs½yedayBYkvamanGMeBIedaykarbEnßm nucleotides f μ I eTAelI 3’─OH rbs; nucleotide cugeRkayrbs; primer . edIm,IeFIV PCR cMnYntictYcén DNA RtUv)anlayCamYynwg Taq polymerase, the primer nig 4

DNA nucleotide triphosphates . l,ayenHbnÞab;mkRtUv)andak;enAkñúg]bkrN_EdleKehAfaCa Thermocycler . sItuNðPaBenAkñúg Thermocycler RtUv)anekIneLIgBI 94-96oC kñúgry³eBlCeRcInnaTI edIm,IEjk strands TaMgBIrrbs; DNA . bnÞab;sItuNðPaBRtUvTMlak;cuH[mkenARtwm 50-56oC edIm,I[GnuBaØati primer eFIVkarbgáat; (hybridize) CamYynwg target DNA . cugbBa©b;sItuNðPaBRtUv)aneFIV[eLIgdl; 72oC

edIm,IGnuBaØati[ Tag polymerase edIm,ImanGMeBI edayGan template DNA strand nig eFIV polymerization

)anCa daughter strand edayBnøÚtBI primer . enAcugbBa©b;énCMhanenH cMnYnénEhSn)anekIneLIgeTVr ¬rUbTI 24-24¦ . \LÚvCMhanTaMgbIRtUv)aneFIVsareLIgvij . CamYynwgvdþnimYy²cMnYnénEhSnRtUv)anekIneLIgeTVr . CaRTwsþI bnÞab;BI 30 vdþ GñkmanmYyBan;landgén DNA eRcInCacMnYnEdlGñkcab;epþIm¡



PCR GacRtUv)aneKeRbIedIm,IRsavRCavrkEhSnEdlTTYlxusRtUvcMeBaH muscular dystrophy . vak¾GacRtUv)aneKeRbIedIm,IeFIVeraKvinicä½yCMgW . cMeBaHehtuplenH vaGacRtUv)aneRbIedIm,IBRgIkcMnYntictYcén HIV

enAkñúgQamrbs;GñkmanCMgWeGds_ . vak¾GacRtUv)aneKeRbIedayGñkviTüasa®sþnitievC¢viTüa (forensic scientist)

edIm,IBRgIkcMnYn DNA BIb¤ssk; b¤ dMNk;QamtUcmYyenAkEnøgXatkmμ . rUbTI 24-24

Fin

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