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Copyright 2007, 2001, 1988, 1977 New Age International (P) Ltd., Publishers
Published by New Age International (P) Ltd., Publishers
All rights reserved.
No part of this ebook may be reproduced in any form, by photostat, microfilm,
xerography, or any other means, or incorporated into any information retrieval
system, electronic or mechanical, without the written permission of the publisher.
All inquiries should be emailed to rights@newagepublishers.com

ISBN : 978-81-224-2429-4

PUBLISHING FOR ONE WORLD

NEW AGE INTERNATIONAL (P) LIMITED, PUBLISHERS

4835/24, Ansari Road, Daryaganj, New Delhi - 110002
Visit us at www.newagepublishers.com

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Hydrocarbon chain
Polar head (hydrophilic)
Water
(a)

Water

Water

(b)

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&

!"#$%&'()*+#,&,-*

(a)
Danielli and Davson
1935

(b)
Davson and Daneilli
1943

(c)
Robertson
1961/1965

!"#$%&$(

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bilayer

Hydrophilic
area

Hydrophilic
area

lntegral
protein

Peripheral
protein

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Ribosomes

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Proteases
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Glycosidases
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Lipases phospholipases
Phosphatases

Phagosome

Proteins
Nucleic acids
Polysaccharides
Organic linked sulphates
Lipids
Organic linked phosphates

Autophagic vacuole

(b)

Early autolysosome

Early phagolysosome

Autolysosome

Phagolysosome
(digestive vacuole)

Residual body

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Complex I
NADH

Complex III

Succinate

!"#$%&$&&

Complex II

Complex 1V

U&,$%+,$-.,-$"*/0*,7"*3"3G$%#"*/0*3',/.7/#)$'/#5

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%6798:7)'7:;?
[(,)#+,&-,+31&-$#&,.'2)#O&'.)#,()#1&-03&,.01+1*&.4&-&0('8#$&.4&0(,)#.*+3#1&^.+0#3&,7-+06+1#9&!-,7
0(,)#.*+3#&+1&-&'.0.'#$&-03&+1&'-3#&(2&.4&*7$##&2-$*15&-&2($+0#&.$&2%$+'+3+0#5&-&2#0*.1#&1(/-$&-03&27.127-*#&/$.(29&:7#&2($+0#1&-$#&-3#0+0#&-03&/(-0+0#5&67+)#&2%$+'+3+0#1&-$#&*7%'+0#5&,%*.1+0#&-03
($-,+)&AU+/9&H9HKB9&=.*7&*7#&2($+0#1&-03&2%$+'+3+0#1&-$#&8-1#19
NH2

OH

PURINES
C

C
N

C
C
C

C
N

H2N

Adenine

H Guanine

PYRIMIDINES
NH2

CH
N
H
Cytosine

!"#$%&$&'

C
CH

CCH3

HN
O

CH
N
H
Thymine

HN
O

CH
CH

C
N
H
Uracil

R7"3'.%&*+,$-.,-$"*/0*2-$'#"+*%#)*2D$'3')'#"+5

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6.17(24,1.0%#;'-0"#1%":%60-@%)48
Monosaccharides
Disaccharides
(one sugar compounds) (two sugar compounds)

Polysaccharides
(many sugar compounds)
+
more

Glucose

Sucrose

Starch
+
more

Fructose

Maltose

Dextrin
+
more

Fructose

Maltose

Glycogen
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more
Cellulose

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/%.'"#b0(#6(/%#+-1"0c8#A1;)4#%'(;52#02%#:1(0%-'"#+(1/#02%#/("0#-/:(10.'0#6('"0-0;%'0"#(+#02%#.'-/.)
7(,4#.',#.66(;'0#+(1#.7(;0#('%#2.)+#(+#02%#0(0.)#,14#$%-520#(+#02%#7(,48#!#)-7%1.)#";::)4#(+#:1(0%-'#-"
'%6%"".14#0(#02%#7(,4#.',#$-02(;0#-0#'(#)-+%#-"#:(""-7)%8#90#/;"0#7%#6('"0.'0)4#";::)-%,#0(#02%#7(,4#+(1
51($02#.',#1%:.-18
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-0"#(15.'"*#+);-,"#.',#0-"";%"8#`(#0$(#:1(0%-'"#.1%#%3.60)4#.)-&%#-'#02%-1#:24"-()(5-6.)#7%2.@-(;18#E(/%
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,-++%1%'0#%'B4/%"8
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6(7.)08#`-01(5%'#-"#02%#,-"0-'5;-"2-'5#%)%/%'0#-'#02%#:1(0%-'"#7%6.;"%#-0#-"#:1%"%'0#-'#02%"%#6(/:(;',"
$2%1%."#-0#-"#.7"%'0#-'#6.17(24,1.0%"#.',#+.0"8
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(+0%'# 1%+%11%,# 0(# ."# /.61(/()%6;)%"8# E-'6%# .# ).15%# ';/7%1# (+# "-/-).1# ;'-0"*# &'($'# ."
/.61(/()%6;)%"*#.1%#X(-'%,#0(#+(1/#62.-'F)-&%#/()%6;)%"#02%4#.1%#62.1.60%1-B%,#."#7-(F:()4/%1"8

%B2768%42=1
!#6(/:)%0%#24,1()4"-"#(+#@.1-(;"#:1(0%-'"#4-%),"#.7(;0#IT#,-++%1%'0#./-'(#.6-,"*#"2($-'5#02%1%74#02.0
02%#:1(0%-'"#.1%#/.,%#;:#(+#./-'(#.6-,#";7;'-0"8#!))#02%"%#./-'(#.6-,"#.1%#"F./-'(#.6-,"*#-8%8#02%
./-'(#51(;:#-"#.00.62%,#0(#02%#"F6.17('8#d.62#./-'(#.6-,#2."#('%#:.10#$2-62#-"#"./%#+(1#.))#.',#('%
:.10# $2-62# -"# ":%6-+-6# (')4# 0(# 02.0# :.10-6;).1# ./-'(# .6-,8# A.&-'5# 02%"%# :(-'0"# -'# @-%$# 02%# 5%'%1.)
+(1/;).#(+#.'#./-'(#.6-,#-"#"&%062%,#."#+())($"<

.,,/+0122+
R

H
N

&(

OH

C
O

A2%#:.10#).7%))%,#f#@.1-%"#+(1#%.62#./-'(#.6-,<#$2%1%."#1%"0#(+#02%#/()%6;)%#-"#02%#"./%#+(1#.))8
A2%#"-/:)%"0#(+#02%#./-'(#.6-,#-"#./-'(F.6%0-6#.6-,#(1#5)46-'%8
H

H
H

N
O

Acetic acid

H
C

C
O

Amino acetic acid (Glycine)

!).'-'%*#02%#'%30#"-/:)%"0#./-'(#.6-,*#-"#./-'(F:1(:-('-6#.6-,8#A2%1%#.1%#0$(#./-'(#,%1-@.0-@%"
(+#:1(:-('-6#.6-,#.',#02%-1#+(1/.0-('#-"#7."%,#('#02%#.00.62/%'0#(+#02%#./-'(#51(;:#%-02%1#0(#"4,(1
!F6.17('#.0(/8
Propionic acid

!-amino-propionic
acid (alanine)

"-amino-propionic
acid ("-alanine)

NH2

OH

C
H

or

H H

OH

H H

OH

H NH2

OH

or

or

OH

OH

C
O

NH2 H

9'#02%#"./%#$.4#021%%#:(""-7)%#./-'(#.6-,"#6.'#7%#,%1-@%,#+1(/#7;041-6#.6-,8#D;0#02%#:1(0%-'"#.1%
6('"0-0;0%,#(')4#$-02#"F./-'(#.6-,"#.',#'(0#$-02#(02%1#./-'(#.6-,"8
9'#.))#./-'(#.6-,"*#$-02#02%#%36%:0-('#(+#5)46-'%*#02%#"F6.17('#.0(/#-"#.00.62%,#0(#+(;1#,-++%1%'0
.0(/"#(1#51(;:"8#A2%#"F6.17('#.0(/#-"#02%1%+(1%#."4//%01-68#A2%#./-'(#.6-,"#6('0.-'-'5#."4//%01-6
"F6.17('#.1%#(:0-6.))4#.60-@%#.',#6.'#%3-"0#-'#0$(#"0%1%(F-"(/%1-6#+(1/"#>=-58#I8I?8
A2%# '.0;1.))4# (66;11-'5# ('%"# 7%)('5# 0(# ZF+(1/8# D;0# "(/%# ./-'(# .6-,"# ";62# ."# .).'-'%# .1%
,%301(1(0.0(1# .)02(;52# 02%4# 7%)('5# 0(# ZF+(1/8# A2%4# .1%# ./:2(0%1-6# %)%601()40%"# 7%6.;"%# 02%4# 2.@%
7(02#./-'(*#.',#6.17(34)#51(;:"#$2-62#1%.60#."#.6-,"#-'#02%#:1%"%'6%#(+#7."%"#.',#."#7."%"#-'#02%
:1%"%'6%#(+#.6-,"8

&!

!"#$%&'()*+#,&,-*
OH
C
O

OH
C
O

NH2

!"#$%&$&

NH2

9+*'"(*1&%".'2*%1('*2'-65"./0'/1",*'/."37

$0(1&01(+2 %#*2 &.%$$)3)&%0)"#2 "32 %!)#"2 %&)*$2 %4%).%5.+2 )#2 -("0+)#$,# A2%# 6).""-+-6.0-('# (+
./-'(# .6-,"*# (70.-'%,# 74# 02%# 24,1()4"-"# (+# :1(0%-'"# -"# 5-@%'# 7%)($# .',# -0# -"# 7."%,# ;:('# 02%
6(/:("-0-('#(+#02%#"-,%#62.-'*#-8%8#fF51(;:#$2-62#@.1-%"#+1(/*#('%#./-'(#.6-,#0(#.'(02%18
()*+,%&$'

81",*'8."3('/,3'9)&"%':-%$.-$%/0'!*%1$0/-&

504+

6'()*')(0.%-,(4).0

;7 :"3&'.)/",('+"-)',*'2$,.-"*,/0'#%*$5(<("150&'/1",*'/."3(7
H

=06.",&'>=06?
@ABCDEA

OH

C
O

NH2

80/,",&'>80/?
@FBGDEA

H/0",&'>H/0?
@FBIIDEA

H3C

NH2

;(*0&$.",&'>;0.?
@KBIFDEA

H3C

H3C

O
OH
C
O

NH2

H 3C
J&$.",&'>J&$?
@KBIFDEA

OH
C

H3C

NH2

OH
C

CH3

NH2

O
OH
C
O
78,#'129

&&

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78,#'129

;;7 6"1+%*:0"#&%+"-)')63%*L60".'>EB?'#%*$5(<B63%*L6M/1",*'/."3(7
:&%",&'>:&%?
@FBGDEF

OH NH2

;;;7 :"3&'.)/",('+"-)'($02$%'/-*1(7
H

@6(-",&'>@6(?
@KBIADAEN:A
H

P&-)"*,",&'>P&-?
@CBIIDEA:

B63%*L606(",&'>B60?
@KBINDAEF

8%#",",&'>8%#?
@KBINDAEA

B"(-"3",&'>B"(?
@KBODFEA

OH
C

NH2

NH2

O
OH

C
O
OH
C
O

NH2

OH
C
O

CH3

;H7 :"3&'.)/",('+"-)'/'Q/(".'#%*$57
J6(",&'>J6(?
@KBINDAEA

NH2

OH

9)%&*,",&'>9)%?
@NBODEF

OH
C

NH2 H

NH2

NH2 OH H

NH2

OH
C

NH2

NH2

HN

CH

CH2
NH

CH

OH
C
O

OH
C
O

CHOOH

NH2

C
H
78,#'129

&)

!"#$%&'()*+#,&,-*

78,#'129

H7 :"3&'.)/",('+"-)'/'./%Q*L60'#%*$5<8."3".'/1",*'/."3(<*%'-)&"%'/1"3&(
8(5/%-".'/."3'>8(5?
@NBGDEN

C C

NH2

HOOC
8(5/%/#",&'>8(,?
@NBRDAEF

OH

C C

OH
C
O

NH2 H NH2
H H H

=0$-/1".'/."3'>=0$?
@CBODEN

OH

H C C C C
O

COOH H NH2
H H H

=0$-/1",&'>=0,?
@CBISDAEF

OH

C C C C C

NH2 H H NH2

H;7 :"3&'.)/",'+"-)'/%*1/-".'>T&,U&,&M0"V&?'#%*$57

H H

W)&,60/0/,",&'>W)&?
@OBIIDEA

OH

C C

C
O

H NH2
H H

96%*(",&'>96%?
@OBIIDEF

HO

H NH2

C
9%65-*5)/,'>9%5?
@IIBIADAEA

OH

CH2

NH2

CH
N
H
H 2C

CH2

OH

CH C

H 2C

W%*0",&'>W%*?
@CBODEA

CH COOH

N
H
HO

B63%*L65%*0",&'>B65?
@CBODEF

HC

CH2

H 2C

CH
N

OH
C
O

A2%# ./-'(# .6-,"# .1%# ,%"61-7%,# ."# 02%# 7;-),-'5# 7)(6&"# (+# :1(0%-'"*# .',# /.4# 7%# 6('@%'-%'0)4
51(;:%,#;',%1#0$(#71(.,#6.0%5(1-%"*#-8%8#%""%'0-.)#.',#'('%""%'0-.)8#A2%#%""%'0-.)#./-'(#.6-,"#.1%#0(
7%#(70.-'%,#74#02%#.'-/.)#021(;52#-0"#,-%08#A2%#'('%""%'0-.)#./-'(#.6-,"#.1%*#2($%@%1*#"4'02%"-B%,#74
02%#.'-/.)#0-"";%"8

.,,/+0122+

&*

@)/3'()&7,A2%#./-'(#.6-,"#>";7;'-0"?#.1%#X(-'%,#0(5%02%1#-'#02%#+())($-'5#/.''%18
R1

R2

NH2 C C OH + NH2 C C OH
H

H
R1

R2

NH2 C C N C C OH + H2O
H

A2%# .7(@%# 1%.60-('# "2($"# 02%# ./-'(# .6-,"# ."# )-'&%,# 021(;52# 02%# 6.17(34)# (C

OH

) # .',# ./-'(
O
51(;:"8# A2%# 6.17('# .',# '-01(5%'# 7(',# 7%0$%%'# %.62# :.-1# (+# ./-'(# .6-,"# "2($'# .7(@%# -"# .# ":%6-.)
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./-'(#.6-,*#-0#"0-))#2."#.'#./-'(#51(;:#.0#('%#%',#.',#.#6.17(34)#51(;:#.0#02%#(02%1#%',8#A2%1%+(1%#-0
6.'#"0-))#1%.60#$-02#.'(02%1#./-'(#.6-,#0(#+(1/#.#)('5%1#62.-'#>01-:%:0-,%?*#$2-62#-'#0;1'#6.'#1%.60#-'
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:%:0-,%#62.-'"8#E%@%1.)#";62#62.-'"#X(-'#0(#+(1/#/.61(/()%6;)%"#(+#:1(0%-'"8

D965>278.59@45@9>
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&"

!"#$%&'()*+#,&,-*

A2%#6(/:)%0%#./-'(#.6-,#"%C;%'6%#(+#"%@%1.)#(02%1#:1(0%-'"#-"#&'($'8#A2%"%#-'6);,%#%'B4/%"
";62#."#1-7(';6)%."%*#)4"(4/%#.',#02%#";70-)-"-'#(+#A0$3)1'?.,&?*3'"'&2,%068
A2;"*#;',%1#02%#:1-/.14#"01;60;1%#$%#2.@%#6('"-,%1%,#02%#./-'(#.6-,#"%C;%'6%8#d.62#./-'(#.6-,
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"01('5%"0#)-'&.5%"8

.>467=09<8.59@45@9>8638D965>271
9'#.,,-0-('#0(#02%#.7(@%#/%'0-('%,#:1-/.14#"01;60;1%#02%#7-()(5-6.)#.60-@-04#(+#:1(0%-'#.)"(#,%:%',"#('
02%#"%6(',.14#"01;60;1%8#\',%1#02%#"%6(',.14#"01;60;1%#(+#:1(0%-'#$%#6('"-,%1#02%#"01;60;1.)#.":%60"
)-&%#+(),-'5#(+#02%#:()4:%:0-,%#62.-'"#-'0(#.#":%6-+-6#6(-)%,#"01;60;1%#.1-"-'5#."#.#1%";)0#(+#02%#)-'&.5%
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)-'&%,#74#/%.'"#(+#24,1(5%'#7(',"#.',#,-";):2-,%#7(',"#>=-58#I8M?8

O
CH

CH2

CH

N
H
O

C H
O

CH

C
H

O
N O COOH
CH CH2 R
H
N

O CH CH2 R

CH

N
H
O

CH

CH

!"#$%&$-

CH2

N
O

CH2

NH2
H

805)/M)&0"L'>/2-&%'W"V&'/,3'T%*+,4'5)'("'",#4'IOKG4'57NS?7

,>95209<8.59@45@9>8638D965>271
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K@05>9709<8.59@45@9>8638D965>27
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.,,/+0122+

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COOH
Fe

NH2
(a)

!"#$%&$.

(b)

>/?'9&%-"/%6'*2'-)&'16*#0*Q",X'>Q?'-)&'.*$%(&'*2'-)&'5*065&5-"3&'.)/",7

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CH3 (CH2)16COOH

Stearic acid (saturated) C18H36O2

H H H H H H H H

H H H H H H H O

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H H H H H H H H H H H H H H H H H
HC (CH2)7CH2
HC (CH2)7CHOOH

Oleic acid (monounsaturated) C18H34O2

H H H H H

H H H H H H H O

H C C C C C C C C C C C C C C C C C C OH
H H H H H H H H H H H H H H H H H
Linoleic acid (polyunsaturated) C18H32O2

HC(CH2)4CH3
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CH2

OH

H OOC R

CH

OH + H OOC R

CH2 OH
H OOC R
One mol. glycerol
Three moles
of fatty acid

CH2 OOC R
CH OOC R + 3H2O
CH2 OOC R
One mol. of
triglyceride

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CH2.O.CO.C17H35

CH2OH

CH.O.CO.C17H35 + 3NaOH # CHOH + 3C17H35COONa

CH2.O.CO.C17H35
Tristerin

CH2OH
Glycerol

Sodium stearate

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@.14#(')4#-'#02%#'.0;1%#(+#('%#(+#02%#51(;:"#)-'&%,#0(#02%#:2(":2.0%#:(10-('#(+#02%#/()%6;)%8#=;102%1*

.,,/+0122+

)*

/("0#(+#02%#6(/:(;',"#.1%#(+#:2(":2(,-%"0%1"#'.0;1%8#P'%#04:%#(+#:2(":2()-:-,*#@-B8#:2(":2.0-,-6
.6-,#-"#-',-6.0%,#7%)($#$-02#.#5%'%1.)#+(1/;).8
-'"$-'%0)*)&2%&)*$,#H2(":2.0-,-6#.6-,"#.1%#6(/:(;',"#6('"-"0-'5#(+#5)46%1()*#0$(#+.004#.6-,"*
.',#.#:2(":2.0%#51(;:8
O
H 2C O C R 1
O
H 2C O C R 2
O
H2C O P OH
OH

A2%#fJ#.',#fI#-'#02%#.7(@%#+(1/;).#1%:1%"%'0#02%#1%"-,;%"#(+#02%#/()%6;)%"#(+#+.004#.6-,"8#A2%
:2(":2.0-,-6#.6-,"#"2(;),#7%#@-%$%,#."#+.0"#-'#$2-62#('%#(+#02%#+.004#.6-,"#-"#1%:).6%,#74#:2(":2(1-6
.6-,8# A2%# "01;60;1%# ";55%"0"*# 02%"%# .6-,"# 6(;),# %."-)4# 5-@%# 1-"%# 0(# 01-5)46%1-,%"# (1# 0(# :2(":2()-:-,"8
A2%# :2(":2.0-,-6# .6-,"# .1%# '(0# :1%"%'0# -'# "-5'-+-6.'0# ./(;'0"# -'# 02%# 0-"";%# %301.60"8# A2%4# :).4
-/:(10.'0#1()%#."#.60-@%#-'0%1/%,-.0%"#-'#02%#7-("4'02%"-"#(+#(02%1#)-:-,#6(/:(;',"8
H)$'3-'+&,>:2(":2.0-,4)#62()-'%"?<#Z%6-02-'"#.1%#62()-'%#%"0%1"#(+#:2(":2.0-,-6#.6-,8#A2%4#.1%#+.0"
-'#$2-62#('%#(+#02%#+.004#.6-,"#-"#1%:).6%,#74#:2(":2(1-6#.6-,#.',#02%#'-01(5%'(;"#62()-'%8#A2%4#.1%
7%"0# &'($'# .',# :1(7.7)4# 02%# /("0# 6(//('# +(1/# (+# :2(":2()-:-,"# -'# .'-/.)"8# P'# 24,1()4"-"*
)%6-02-'"#71%.&#;:#-'0(#5)46%1()*#+.004#.6-,"*#:2(":2(1-6#.6-,#.',#62()-'%8
Z%6-02-'"#.1%#"();7)%#-'#02%#+.0#"()@%'0"#%36%:0#.6%0('%#.',#74#02-"#:1(:%104#02%4#.1%#,-"0-'5;-"2%,
+1(/#02%#+.0"8#A2%"%#.1%#1%C;-1%,#+(1#02%#01.'":(10*#.',#;0-)-B.0-('#(+#(02%1#)-:-,"#%":%6-.))4#-'#02%#)-@%18
9+#02%#"4'02%"-"#(+#62()-'%*#.'#-/:(10.'0#6(/:('%'0#(+#)%6-02-'#-"#-'0%11;:0%,*#02%#"4'02%"-"#(+#)%6-02-'
.)"(# $-))# 7%# "0(::%,8# ]-02# 02%# 1%";)0*# 02%# 01.'":(10# (+# +.0# 0(# .',# +1(/# 02%# )-@%1# -"# -'0%11;:0%,8
G('"%C;%'0)4*#02%#)-:-,"#.66;/;).0%#-'#02%#)-@%1#5-@-'5#1-"%#0(#.#6(',-0-('#6.))%,#C033#,"'K)15,A2%#).6&#(+
,-%0.14#62()-'%#.)"(#)%.,"#0(#.#@.1-%04#(+#01(;7)%"#";62#."#51($02#+.-);1%*#2%/(112.5-6#&-,'%4"*#.',
")-::%,#0%',('"*#%068
O
H 2C O C R 1
O
HC O C R2
O
+

H2C O P O CH2CH2N (CH3)3

O

A2%#%'B4/%*#)%6-02-'."%#!*#:1%"%'0#-'#02%#@%'(/"#(+#"'.&%"#.00.6&"#)%6-02-'"#.',#1%/(@%"#('%#(+
02%# +.004# .6-,# 1%"-,;%"# )%.@-'5# .# :1(,;60# &'($'# ."# )4"()%6-02-'8# A2-"# :1(,;60# 2."# 02%# .7-)-04# 0(
2.%/()4B%#02%#1%,#7)((,#6(1:;"6)%"8
>)/-0"'+&, >H2(":2.0-,4)# %02.'()./-'%"?<# A2%"%# .1%# :2(":2()-:-,"# $2-62# 1%"%/7)%# )%6-02-'"# -'
/("0# :1(:%10-%"*# 7;0# ,-++%1# -'# 6('0.-'-'5# ./-'(%024)# .)6(2()# >`NIKGNIKGNIKPN?# -'# :).6%# (+
62()-'%8

)"

!"#$%&'()*+#,&,-*

O
H 2C O C R 1
O
HC O C R2
O
+

H2C C P O CH2CH2NH3
O

H2(":2.0-,4)"%1-'%#-"#.#6%:2.)-'#)-&%#6(/:(;',#.',#6('0.-'"#02%#./-'(#.6-,#"%1-'%#-'#:).6%#(+
%02.'()./-'%8
O
H 2C O C R 1
O
HC O C R2
O
+

H2C O P O CH2CH NH3

O

COOH

@-%&/-03'(#",'+%&'3%",-"#.#:2(":2()-:-,#"-/-).1#-'#"01;60;1%#0(#02%#)%6-02-'#7;0#2."#-'("-0()#-'#:).6%
(+#62()-'%8
O
H 2C O C R 1
O
HC O C R2
O
H2C O P O C
OH

OH

OH

H
C

OH

OH

OH
H

A2%"%#.1%#/.-')4#+(;',#-'#:).'0"#.',#-'#'%1@(;"#0-"";%"8
@"0&.0"%!)+&7,A2%"%# :2(":2()-:-,"# .1%# .7;',.'0# -'# 71.-'# .',# /;"6)%8# A2%4# 1%"%/7)%# )-6-02-'"
.',#6%:2.)-'"#-'#"01;60;1%#7;0#:(""%""#.'#.),%24,%#51(;:#-'#:).6%#(+#('%#(+#02%#+.004#.6-,"#-'#04:-6.)
:2(":2()-:-,#/()%6;)%8#A2%#"01;60;1%#"2($'#7%)($#1%:1%"%'0"#.#04:-6.)#:)."/.)(5%'8#A2%#:(10-('#(+
02%#+(1/;).#%'6)("%,#74#)-'%#-"#.'#%02('()./-'%#1%"-,;%8

.,,/+0122+

)#

H2C O CH = CH R1
O
HC O C R2
O
+

H2C O P O CH2 CH2 NH3

O
Structure of plasmalogen (phosphatidal ethanolamine)

6/-'+!%&'+), "'/'(&7, A2%"%# .1%# /(1%# 6(/:)-6.0%,# :2(":2.0-,%"# 6('0.-'-'5# ":2-'5("-'%*# .# )('5F
62.-'#./-'(#.)6(2()#-'#:).6%#(+#5)46%1()8
6/-'+!%.#)"'+&,>":2-'5("-'%#:2(":2.0-,%"?<#A2%"%#:2(":2()-:-,"#6('0.-'#":2-'5("-'%#.',#.#+.004
.6-,#.00.62%,#-'#02%#+(1/#(+#./-,%#)-'&.5%#0(#02%#'-01(5%'#(+#6.17('#I*#.',#:2(":2(1462()-'%#.00.62%,
0(# 02%# 0%1/-'.)# 6.17('# .0(/# (+# ":2-'5("-'%8# P'# 24,1()4"-"# 02%4# 4-%),# +.004# .6-,"*# :2(":2(1-6# .6-,*
62()-'%# .',# ":2-'5("-'%8# A2%# ":2-'5(/4%)-'"# .1%# +(;',# -'# ).15%# C;.'0-0-%"# -'# 02%# '%1@%# 0-"";%*
:.10-6;).1)4#-'#02%#/4%)-'#"2%.02#(+#02%#'%1@%8
7./&".)-)*$,#G%1%71("-,%"#.',#5.'5)(-"-,%"#.1%#02%#62-%+#5)46()-:-,"8
CH3(CH2)12CH=CH

CH3(CH2)12CH=CH

HO CH

HO CH
O

H2N CH
CH2OH

R C N CH
H

CH2 O P O CH2
Sphingosine

Sphingomyelins O

CH2N (CH3)3

A2%"%# .1%# :.10-6;).1)4# .7;',.'0# -'# 02%# 71.-'# .',# -'# 02%# /4%)-'# "2%.02# (+# '%1@%"8# A2%1%# .1%# 02%
)-:-,"#6('0.-'-'5#6.17(24,1.0%#1.,-6.)"#.',#'-01(5%'#-'#02%#/()%6;)%8#9'#"01;60;1%#02%"%#.1%#1%).0%,#0(
":2-'5(/4%)-'"*#7;0#6('0.-'#.#2%3("%#-'"0%.,#(+#:2(":2(1462()-'%8#P'#24,1()4"-"#02%4#5-@%#1-"%#0(
+.004#.6-,"*#":2-'5("-'%#.',#5.).60("%8

!"P QE,%GEF.
9'#02%#4%.1#J^JJ#=;'&*#.#H()-"2#7-(62%/-"0*#6(-'%,#02%#0%1/#b@-0./-'%c#0(#,%"-5'.0%#02%#.'0-7%1-7%1+.60(18#9'#02%#0%1/#@-0./-'%#K'30, ";55%"0"#02%#%""%'0-.)#'.0;1%#(+#02%#+.60(1*#.',#0.'+), -',-6.0%"#02%
62%/-6.)# "01;60;1%8# Z.0%1# 02%# "./%# 0%1/# $."# ;"%,# +(1# "%@%1.)# ;'&'($'# ,-%0.14# +.60(1"8# `(0# .))# 02%
&'($'# +.60(1"# 2.@%# 02%# ./-'%# "01;60;1%8# A2%1%+(1%# -'# J^J^# 02%# )."0# b%c# +1(/# 02%# @-0./-'%# $."
,-"6('0-';%,#+1(/#;"%#02%1%74#1%/(@-'5#02%#-/:)-6.0-('#(+#62%/-6.)#"01;60;1%8
_-0./-'"# 6.'# 7%# ,%"61-7%,# ."# .66%""(14# +((,# +.60(1"# $2-62# .1%# %""%'0-.)# +(1# "(/%# /%0.7()-6
1%.60-('"#$-02-'#02%#6%))#.',#$2-62#/;"0#7%#:1(@-,%,#-'#02%#,-%0#-'#/-';0%#./(;'0"8#9'#/.'4#.'-/.)"

)$

!"#$%&'()*+#,&,-*

02%4#6.''(0#7%#"4'02%"-B%,#74#02%#7(,48#G%10.-'#.'-/.)"#,(#"4'02%"-B%#.#+%$#@-0./-'"#-'#@%14#/-';0%
C;.'0-0-%"# 7;0# 02%4# +.))# "2(10# (+# 02%# 1%C;-1%/%'0"8# 9'# .'4# 6."%*# 02%# .'-/.)"# "2(;),# :1(6;1%# 02%/
021(;52#+((,8#_-0./-'#,%+-6-%'64#1%";)0"#-'#"%@%1.)#,-"%."%"#62.1.60%1-"0-6#(+#%.62#@-0./-'#,%+-6-%'648
_-0./-'"#.1%#.#":%6-.)#51(;:#(+#(15.'-6#";7"0.'6%"#$2-62#.1%#62%/-6.))4#;'1%).0%,8#A2%4#/.4#7%
(15.'-6# .6-,"*# ./-'%"*# ./-'(# .6-,"*# %"0%1"*# .)6(2()"*# "0%1(-,"*# %068# A2%"%# @-0./-'"# .1%# 01.,-0-('.))4
,-@-,%,#-'0(#0$(#";751(;:"#('#02%#7."-"#(+#02%-1#"();7-)-04#:1(:%10-%"*#";62#."#$.0%1#"();7)%#@-0./-'"
.',#+.0#"();7)%#@-0./-'"8

R05>9S16/@:/>8Q250B271
_-0./-'"#G#.',#D#6(/:)%3#.1%#$.0%1#"();7)%#";7"0.'6%"8
%$&"(5)&2%&)*2 "(24)0%!)#2&,#!"6(17-6#.6-,#-"#.#2%3("%#,%1-@.0-@%#.',#-"#:1(:%1)4#6).""-+-%,#."
.# 6.17(24,1.0%8# 90# -"# .# $2-0%# 614"0.))-'%# ";7"0.'6%# 2-52)4# "();7)%# -'# $.0%18# 90# -"# %."-)4# (3-,-B%,# -'
"();0-('#.',#6((&-'5#,%"01(4"#-08#A2%#@-0./-'#-"#2-52)4#;'"0.7)%#-'#.)&.)-'%#"();0-('"8
A2-"# @-0./-'# -"# 1%C;-1%,# 74# :1-/.0%"# .',# 02%# 5;-'%.# :-58# !))# (02%1# @%10%71.0%"# .',# "(/%
-'@%10%71.0%"# ."# $%))# ."# :).'0"# .',# /("0# /-61(F(15.'-"/"# 6.'# "4'02%"-B%# ."6(17-6# .6-,# +1(/
6.17(24,1.0%"8#90#-"#.'#%""%'0-.)#,-%0.14#+.60(1*#02%#,%+-6-%'64#(+#$2-62#6.;"%"#:.-'+;)#,-"%."%#(+#02%
X(-'0"#.',#5;/"#6.))%,#&$?1K#5
O=C
HO C

HO C
HC
HO C H
CH2 OH
LAscorbic acid
(reduced form)

O=C
O=C

O=C
HC
CHOH
CH2 OH
LDehydroascorbic acid
(oxidized form)

N4,1(34:1()-'%#-"#.'#;';";.)#./-'(#.6-,#+(;',#-'#02%#6()).5%'#.',#-"#%""%'0-.)#+(1#/.-'0%'.'6%
(+#'(1/.)#0-"";%"8#90#-"#+(1/%,#74#24,1(34).0-('#(+#:1()-'%#.',#02-"#:1(6%""#1%C;-1%"#@-0./-'#G8#9'#02%
.7"%'6%#(+#02-"#@-0./-'#02%#24,1(34:1()-'%#:1(,;60-('#-"#@%14#/;62#1%,;6%,8#A2%1%+(1%*#02%#6()).5%'
+(1/%,#,;1-'5#."6(17-6#.6-,#,%+-6-%'64*#6('0.-'"#'%5)-5-7)%#./(;'0#(+#24,1(34:1()-'%#.',#02-"#71-'5"
.7(;0#"01;60;1.)#.7'(1/.)-0-%"#(7"%1@%,#-'#"6(17;0-6#0-"";%"8
_-0./-'#G#-"#6('0.-'%,#-'#+1%"2#@%5%0.7)%"#.',#+1;-0"8
4)0%!)#2527("1-,#_-0./-'#D#51(;:#6('0.-'"#.7(;0#0$%)@%#&'($'#@-0./-'"8#d.62#('%#:%1+(1/"
":%6-+-6#+;'60-('8#A2%4#.1%#6())%60-@%)4#6.))%,#@-0./-'#D#6(/:)%3#.',#.1%#5%'%1.))4#+(;',#0(5%02%18
E-'0.'+),%1,K'30.'+,A7,A2-"#@-0./-'#-"#6('0.-'%,#-'#4%."0*#/-)&*#%55*#:%."#.',#7%.'"#2%'6%#1-62)4
,-"01-7;0%,# -'# :).'0"*# .'-/.)"# .',# 6%10.-'# /-61((15.'-"/"8# !# ,%+-6-%'64# (+# 02-"# @-0./-'# 6.;"%"# .
"%1-(;"#,-"%."%#&'($'#."#*)1'*)1'5,90"#"4/:0(/"#.1%#-'+).//.0-('#(+#'%1@%"*#/;"6;).1#$%.&'%""#.',
:.1.)4"-"#(+#)-/7"8

.,,/+0122+

NH2

C1

H3C

CH3

(CH2)2 OH
S

CH2 N

N
N

)%

Thiamine

A2-./-'%# +;'60-('"# ."# :.10# (+# 02%# 6(%'B4/%# 6(6.17(34)."%# $2-62# -"# 02%# 6(%'B4/%# +(1# :41;@-6
,%6.17(34)."%#.',#+(1#"%@%1.)#(02%1#%'B4/%"#$2-62#.1%#'%6%"".14#+(1#02%#(3-,.0-('#(+#:41;@-6#.6-,#0(
.6%04)KG(!#.',#"Ka%0(5);0.1-6#.6-,#0(#";66-'4) K G(!#>"%%#G2.:0%1#Q?8
L'*%C"0K'+,%1,K'30.'+,AM7,A2-"#@-0./-'#-"#$-,%)4#,-"01-7;0%,#.',#02%#1-62#"(;16%"#.1%#)-@%1*#4%."0*
$2%.0#5%1/*#/-)&*#%55"*#.',#51%%'#)%.+4#@%5%0.7)%"8#90"#,%+-6-%'64#6.;"%"#61.6&"#-'#02%#6(1'%1"#(+#02%
/(;02# .',# ,%1/.0-0-"# (+# 02%# +.6%8# A2%# %4%"# 7%6(/%# -'+)./%,# .',# 02%1%# -"# ,-/'%""# -'# @-"-('8# A2-"
@-0./-'# -"# .# :.10# (+# 0$(# 6(%'B4/%"*# +).@-'# /('(';6)%(-,%# >=L`?# .',# +).@-'# .,%'-'%# ,-';6)%(0-,%
>=!Y?#$2-62#.1%#)-'&%,#0(#:1(0%-'"#."#+).@(:1(0%-'"8
OH OH OH
CH2 C
CH3

CH3

C CH2OH

H H H
N O
NH

O
Riboflavin

N'$%3'+'$, 0$'(# %1, +'0$'+7, `-6(0-'-6# .6-,# -"# .'# %""%'0-.)# ,-%0.14# 1%C;-1%/%'08# A14:0(:2.'# -"# -0"
:1%6;1"(1# $2-62# -"# 6('@%10%,# -'0(# '-6(0-'-6# .6-,# 021(;52# .# "%1-%"# (+# 7-(62%/-6.)# 1%.60-('"8# 90"
,%+-6-%'64#-'#,-%0#6.;"%"#.#,-"%."%#:%)).51.#$2-62#-"#62.1.60%1-B%,#74#,%1/.0-0-"#,-.112(%.#.',#'%1@(;"
,-"(1,%1"8#A2-"#@-0./-'#>'-.6-'?#-"#.#6(/:('%'0#(+#6(%'B4/%"#)-&%#YH`#.',#AH`#$2-62#0.&%#:.10#-'
(3-,.0-('F1%,;60-('#(+#6.17(24,1.0%"*#+.0"*#:1(0%-'"#.',#';6)%-6#.6-,#/%0.7()-"/8
A2-"#@-0./-'#-"#$-,%":1%.,*#7;0#02%#62-%+#"(;16%"#.1%#/%.0*#)-@%1*#4%."0*#7%.'"#.',#$2%.0#5%1/8
O
C
OH
N
Nicotinic acid

O'30.'+,AP7,A2-"#@-0./-'#(66;1"#-'#021%%#+(1/"[#:41-,(3-'%*#:41-,(3.)#.',#:41(,(3./-'%8#90#-"#.'
%""%'0-.)# 6(/:('%'0# (+# 02%# 6(%'B4/%# :41-,(3.)# :2(":2.0%# $2-62# -"# .'# -/:(10.'0# 6(+.60(1# -'# 02%
01.'"/-'.0-('#.',#,%6.17(34).0-('#(+#.))#'.0;1.))4#(66;11-'5#./-'(#.6-,"8#A2%#,-%0.14#1%C;-1%/%'0#(+
@-0./-'# DO[# -'# 2;/.'# 7%-'5"# -"# '(0# 4%0# +;))4# %"0.7)-"2%,8# 90# -"# "4'02%"-B%,# 74# 51%%'# :).'0"# .',# .
';/7%1#(+#/-61((15.'-"/"8#A2%#@-0./-'#-"#+(;',#-'#/.'4#+((,"#)-&%#)-@%1*#:(1&*#&-,'%4*#4%."0*#%55F
4()&*#51.-'"#.',#@.1-(;"#"%%,"8
O
HO

CH2OH
CH2OH

CH3 N
Pyridoxine

HO
CH3

CH2 NH2
CH2OH

N
Pyridoxamine

C
OH
CH3

CH2OH

N
Pyridoxal

*'

!"#$%&'()*+#,&,-*

@0+3%3-)+'$,0$'(7,A2%#1%C;-1%/%'0#(+#02-"#@-0./-'#-'#2;/.'#7%-'5"#2."#'(0#7%%'#%"0.7)-"2%,#"(
+.1*# .)02(;52# -0# -"# %""%'0-.)# +(1# /.'4# (02%1# .'-/.)"8# H.'0(02%'-6# .6-,# -"# %""%'0-.))4# .# :(10-('# (+# 02%
6(%'B4/%8#!#/()%6;)%#$2-62#2."#.#&%4#1()%#-'#02%#/%0.7()-"/#(+#+.0"*#6.17(24,1.0%"#.',#./-'(#.6-,"8
90"#1()%#2."#7%%'#+;))4#,%0%1/-'%,#-'#1.0"#$2%1%#02%#).6&#(+#02-"#@-0./-'#-',;6%#1%0.1,.0-('#(+#51($02*
1%:1(,;60-('#-/:.-1/%'0#.',#51%4-'5#(+#7).6&#2.-18#A2%#1-62#"(;16%"#(+#02-"#@-0./-'#.1%#)-@%1*#4%."0*
%55"#.',#1(4.)#X%))48#A2-"#-"#"4'02%"-B%,#74#51%%'#:).'0"#.',#"(/%#/-61((15.'-"/"8
H CH3 OH O H H H
HOC C

C N C C COOH

H CH3 H

H H

Pantothenic acid

A'%3'+7,D-(0-'#1%C;-1%/%'0"#-'#/.'#2.@%#'(0#7%%'#,%0%1/-'%,#"(#+.1*#7;0#-0"#-/:(10.'6%#-'#(02%1
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Glutamic
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Folic Acid (Folacin)

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Lipoic acid (oxidized form)

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Cu

Zn

A red

B ox

A ox

B red

or

++

Zn

Cu

(Ox: oxidized)
(red: reduced)

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-+"+2(+85
ATP

!G = 10 kcal/mole

ADP + Pi

!G = +10 kcal/mole

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NH2
N
N
CH
Adenine
O

CH2

OH

OH

H
D-Ribose

Adenosine Triphosphate (ATP)

!"#$%&$'

!"#$%"$#&'()'*+,-

Uncatalysed reaction
+

H Catalysed reaction
Enzyme catalysed reaction

Energy

Urea
H2O

CO2 + 2NH3
+ 13800 calories

Reaction coordinate

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-+./$-+#+@*-+,+")#(,2""#2,0/&*(#03#+&9),+(5
E3# 2# '"/10(+# ,0"+1/"+# $(# (/:`+1*+8# *0# 2# (+-$+(# 03# +&9),+A12*2")9+8# -+21*$0&(?# *7+# ,0"+1/"+# $(
:-0;+&#80<&#<$*7#,$&$,/,#+@4+&8$*/-+#03#+&+-')5#I0,+*$,+(#*7+#+&+-')#-+./$-+8#30-#*7+#-+21*$0&#$(
4-+(+&*#$&#*7+#;$&+*$1#30-,#$&#*7+#-+21*$0&#$*(+"3?#:/*#,0-+#3-+./+&*")?#*7+#+&9),+#21*(#$&#10,:$&2*$0&
<$*7#!6C#-+./$-$&'#"0<#21*$%2*$0&#+&+-')5#D0*7#()&*7+*$1#2&8#8+'-282*$%+#42*7<2)(#-+./$-+#*7+#/(+#03
!6C5#E&#()&*7+*$1#-+21*$0&(#*7+#!6C#,0"+1/"+#10,:$&+(#<$*7#*7+#+&9),+#2*#0&+#03#*7+#21*$%+#($*+(#2&8
*7+#(/:(*-2*+#10,:$&+(#<$*7#0*7+-#($*+(5#67$(#12&#:+#(70<&#2(K
I/:(*-2*+#W#+&9),+(#W#!6C#"#!6C#M#+&9),+#M#(/:(*-2*+#"
!]C#W#C$#W#+&9),+#W#4-08/1*
I$&1+#*7+#-+21*$0&#$(#+&9),+#,+8$2*+8?#$*#12&#:+#($,4"$3$+8#*7/(K
ATP
A
(Substrate)

ADP + Pi
B
(Product)

Enzyme

E&# 8+'-282*$0&# -+21*$0&(?# *7+# *+-,$&2"# 470(472*+# '-0/4# 03# !6C# X!# M# C# M# C# M# CY# $(# /(/2"")
*-2&(3+--+8#*0#*7+#(/:(*-2*+#<7$17#'+*(#470(40-)"2*+85#67$(#,2)#:+#(70<&#2(K
ATP
A

ADP

Enzyme

B
(Phosphorylated
substrate)

I$&1+# !6C# $(# /(+8# :0*7#$&# ()&*7+*$1# 2&8# 8+'-282*$%+# -+21*$0&(?# $*(# -+'+&+-2*$0&# $(# +((+&*$2"# 30+33$1$+&*# 3/&1*$0&$&'# 03# *7+# :$0"0'$12"# ()(*+,(# XZ$'5# [5[Y5# \+'+&+-2*$0&# 03# !6C# $(# 40(($:"+# $&# *<0
<2)(K# X$Y# :)# *-2&(3+-# 03# 2# 470(472*+# '-0/4# 3-0,# 2# 7$'7A+&+-')# ,0"+1/"+# X(/:(*-2*+A"+%+"
470(470-)"2*$0&Y#2&8#X$$Y#:)#+"+1*-0&#*-2&(40-*#()(*+,5

.#,&,-#/%&'01#2%3#,"+
ADP
+
Pi

Oxidation of glucose

#'

Muscle contraction
Nerve conduction

Fatty acid oxidation

Glucose absorption

Oxidation of amino acids

Energy
requiring
reaction

Energy
yielding

Protein synthesis

Photosynthesis

ATP

!"#$%&$&

*+,'%1%4&'7/'&/&#819"#3/5)&#':#(%&55&5-

!":

&8%63+%&(;)'64,+%&(1)'3,+%&(-

O&+# 03# *7+# ,2`0-# 1"2((+(# 03# -+21*$0&(# $(# *7+# 0@$82*$0&A-+8/1*$0&# -+21*$0&# $&# <7$17# +"+1*-0&(# 2-+
*-2&(3+--+8#3-0,#0&+#2*0,#*0#2&0*7+-5#N+#72%+#2"-+28)#&0*+8#*72*#0@$82*$0&#$&%0"%+(#"0((#03#+"+1*-0&(
<7+-+2(#-+8/1*$0&#$&%0"%+(#'2$&#03#+"+1*-0&(5#67$(#,2)#:+#(70<&#2(K
Oxidation
Fe+++ + e

Fe++
Reduction

E&#(0,+#03#*7+#-+80@#-+21*$0&(?#*7+#%2"/+(#03#!:#2-+#%+-)#7$'7?#7+&1+#*7+)#,2)#(+-%+#2(#!6C
-+'+&+-2*$0&# -+21*$0&(5# E&# 2""# -+80@# -+21*$0&(?# 2# -+8/1$&'# 2'+&*# X+"+1*-0&# 80&0-Y# 2&8# 2&# 0@$8$9$&'
2'+&*# X+"+1*-0&# 211+4*0-Y# 2-+# 4-+(+&*5# ]/-$&'# *7+# -+21*$0&?# -+8/1$&'# 2'+&*# '+*(# 0@$8$9+8# 2&8# *7+
0@$8$9$&'# 2'+&*# '+*(# -+8/1+8# 2&8# *7$(# $(# 8+4+&8+&*# /40&# *7+# 2:$"$*)# 03# -+8/1$&'# 2'+&*# *0# 3/-&$(7
+"+1*-0&(# 2&8# *7+# *+&8+&1)# 03# 0@$8$9$&'# 2'+&*# *0# 211+4*# *7+,5# 67$(# -+"2*$%+# 2:$"$*)# *0# 80&2*+# 2&8
211+4*#+"+1*-0&(#$(#12""+8#$!#&),4&*!"*'355#E*#$(#-+4-+(+&*+8#:)#*7+#30""0<$&'#+./2*$0&K
7+#L#7&#W#
<7+-+
7+
7&
=
;
"

L
L
L
L
L

=;
X0@$82&*Y
#"&
">
X-+8/1*2&*Y

*7+#-+80@#40*+&*$2"
*7+#(*2&82-8#+,3#03#*7+#()(*+,
'2(#10&(*2&*#$&#`0/"+(T,0"+T8+'-++
2:(0"/*+#*+,4+-2*/-+
%2"+&1+#03#*7+#$0&#0-#&/,:+-#03#+"+1*-0&#+./$%2"+&*(

#(

!"#$%&'()*+#,&,-*

> L Z2-282)#XaQ?^SS#10/"0,:(Y
"& L "0'2-$*7,#*0#*7+#:2(+#+
N7+&#*7+#10&1+&*-2*$0&#03#0@$82&*#2&8#-+8/1*2&*#2-+#+./2"?#E&#X0@$82&*T-+8/1*2&*Y#:+10,+(#S#2&8
7+#L#7&5#67+#-+80@#40*+&*$2"#%2"/+(#2(#1$*+8#2:0%+#2-+#244"$12:"+#<7+&#*7+#4U#%2"/+#$(#S?#0*7+-<$(+#$*
<0/"8#%2-)#<$*7#*7+#172&'+#$&#4U5#67+#7&,03#*7+#7)8-0'+&#2*#4U#b5S#$(#M#S5cVS#F#X-+80@#40*+&*$2"#03
7)8-0'+&#2*#4U#S#$(#S5SSSY5

!"< +='1,/+&,=)&9'1-/-+'9
67+# *-2&(3+-# 03# +"+1*-0&(# 011/-(# 0&# 2&# 2*0,$1# "+%+"5# N7+*7+-# *7+# 2*0,# (70/"8# 21*# 2(# 2# -+8/1$&'# 00@$8$9$&'#2'+&*#8+4+&8(#0&#*7+#(*-/1*/-+#03#*7+#,0"+1/"+5#67+#1)*017-0,+(#2-+#2#'-0/4#03#10,4"+@
*)4+(#03#,0"+1/"+(#:+"0&'$&'#*0#*7+#1"2((#03#40-47)-$&(5#67+)#72%+#2&#2*0,#03#$-0&#XZ+Y#7+"8#$&#*7+
40-47)-$&# -$&'# (*-/1*/-+5# !&# $,40-*2&*# 4-04+-*)# 03# *7+# 1)*017-0,+(# $(# *7+$-# 2:$"$*)# *0# /&8+-'0
-+%+-($:"+#0@$82*$0&#$&%0"%$&'#2#172&'+#$&#*7+#%2"+&1)#03#$-0&5
!#%2-$+*)#03#1)*017-0,+(#72%+#:++&#$8+&*$3$+8#3-0,#4"2&*(#2&8#2&$,2"(5#E&#,2,,2"$2&#1+""(?#3$%+
*)4+(#03#1)*017-0,+(#72%+#:++&#$8+&*$3$+8K#>)*017-0,+(#:?#1P?#1?#2#2&8#2[5#67+(+#72%+#$,40-*2&*#-0"+(
$&#*7+#0@$82*$0&#-+21*$0&(5#E&#:$0"0'$12"#()(*+,(?#-+21*$0&(#$&%0"%$&'#*7+#-+,0%2"#03#7)8-0'+&#2*0,
011/-# ,0(*# 3-+./+&*")# 2&8# *7$(# 7)8-0'+&# 2*0,# $(# *-2&(3+--+8# *0# *7+# 7)8-0'+&# 211+4*0-5# d!]
X&$10*$&2,$8+# 28+&$&+# 8$&/1"+0*$8+Y# $(# *7+# 10,,0&# 7)8-0'+&# 211+4*0-# <7$17# 21*(# 2(# 2&# 0@$8$9$&'
2'+&*5#E&#*7+#10&%+-($0&#03#4)-/%2*+#*0#>OV#2&8#21+*2*+?#-0"+#03#d!]#12&#:+#$&8$12*+8K
COOH
NAD
C

NADH + H

CO2 + CoA
C
CH3
(Acetyl CoA)

CH3
(Pyruvate)

E&# *7+# 2:0%+# -+21*$0&?# *7+# 12-:0@)"# '-0/4# $(# "0(*# :)# *7+# 4)-/%2*+# 2&8# *7+# -+,2$&$&'# VA12-:0&
40-*$0&#$(#`0$&+8#*0#10+&9),+#!5#N7+&#d!]#211+4*(#2#7)8-0'+&?#$*#2"(0#$&%0"%+(#$0&$1#:0&8$&'#*0#2
(+10&8#7)8-0'+&?#'$%$&'#-$(+#*0#4-08/1*#d!]U#W#UWK
d!]#W#4)-/%2*+#"#d!]U#W#UW#W#V+M#W#>OV#W#21+*2*+
\+'+&+-2*$0&#03#!6C#$(#8+4+&8+&*#/40&#-+21*$0&(#$&%0"%$&'#-+,0%2"#03#7)8-0'+&5#67+#0@$8$9$&'
2'+&*# d!]# 72(# 2&# 70# 035/!, &.# M S5[VS?# <7+-+2(# *7+# -+8/1$&'# 2'+&*# 4)-/%2*+# 72(# 2&# B0# %2"/+# 03# M
S5bSS5#67+-+#2-+#,2&)#10,40/&8(#$&#:$0"0'$12"#()(*+,(#<7$17#72%+#,0-+#40($*$%+#-+80@#40*+&*$2"(
*72&#d!]#2&8#d!]U5#N7+&#d!]#-+21*(#<$*7#(/17#10,40/&8(?#$*#(+-%+(#2(#2#-+8/1$&'#2'+&*#$&#$*(
d!]U# W# UW# 30-,5# d!]U# X-+8/1+8# 30-,Y# +&*+-(# $&*0# 2# (+-$+(# 03# -+21*$0&(# $&%0"%$&'# *-2&(3+-# 03
+"+1*-0&(5# E&# *7+# 10/-(+# 03# -+21*$0&(# '-+2*# 8+2"# 03# +&+-')# $(# '+&+-2*+85# I+-$+(# 03# (/17# -+21*$0&(# 2-+
12""+8#+"+1*-0&#*-2&(40-*#()(*+,5#E&#*7+#(17+,+#03#-+21*$0&(?#0@)'+&#$(#*7+#3$&2"#7)8-0'+&#211+4*0XZ$'5#[5cY5

#!

.#,&,-#/%&'01#2%3#,"+
Reduced
substrate
A-H2
2e

ATP
FADH2

NAD
2e
NADH2

quinone
2e

2e

FAD

ADP + P

!"#$%&$)

2 Fe

ATP

2+

cyt. b 2e
3+

hydroquinone

2 Fe

Coenzyme
Q

ADP + P

ATP
2+

3+

2 Fe

2 Fe

cyt. c 2e

cyt. a 2e

2+

3+

2 Fe

2 Fe

3+

2 Fe

cyt. a3
2+

2 Fe

H 2O
2e
1/2O2

ADP + P

.4&%"#(/'"#3/5:(#"'515"&2-

B"+1*-0&#*-2&(40-*#()(*+,#$&#8$33+-+&*#0-'2&$(,(#,2)#8$33+-#$&#1+-*2$&#(*+4(5#U0<+%+-?#*7+#,2`010,40&+&*(#03#*7+#()(*+,#2-+K
X2Y 2&#+&9),+_d!]#X&$10*$&2,$8+#28+&$&+#8$&/1"+0*$8+Y
X:Y 2#3"2%04-0*+$&
X1Y 10+&9),+#e
X8Y 1)*017-0,+#10,40/&8(

!"> +='1253?&0)&+'%(E&#:$0"0'$12"#()(*+,(?#,2&)#,+*2:0"$*+(#/&8+-'0#0@$82*$0&#:)#*7+#"0((#03#7)8-0'+&#*7-0/'7#12*2")9+8
-+21*$0&(5#67+#(4+1$3$1#+&9),+(#$&%0"%+8#2-+#;&0<&#2(##!+-#$&%!"31!1#2&8#*7+#-+21*$0&#12&#4-01++8
<$*7#*7+#2$8#03#2#7)8-0'+&#211+4*0-5#]+7)8-0'+&2*$0&#03#2#(/:(*-2*+#8+4+&8(#/40&#*7+#2%2$"2:$"$*)#03
d!]W# 0-# d!]CW5# 67+# 0@$8$9+8# 30-,(# 03# *7+(+# +&9),+(# 2-+# 4-+(+&*# $&# *7+# ()(*+,# $&# ,$&/*+
./2&*$*$+(?#2&8#$&#0-8+-#*0#,2$&*2$&#2#10&(*2&*#(/44")?#d!]U#2&8#d!]CU#2-+#-+0@$8$9+8#*0#d!]
2&8#d!]C5#67+#,+172&$(,#*0#-+0@$8$9+#*7+(+#+&9),+(#$(#3/-&$(7+8#:)#1+-*2$&#.530&4$&*!'"15#67+(+
2-+#4-0*+$&(#2&8#21*#2(#10+&9),+(5#67+)#10&*2$&#2#4-0(*7+*$1#'-0/4#ZGd#X3"2%$&#,0&0&/1"+0*$8+Y#0Z!]#X3"2%$&#28+&$&+#8$&/1"+0*$8+Y5#67+(+#10+&9),+(#12*2")9+#0@$82*$0&A-+8/1*$0&#-+21*$0&(K
+2H
FPH2

FP
2H

6<0#10A+&9),+(#03#*7$(#'-0/4?#d!]UV#2&8#d!]CUV#-+0@$8$9+#d!]U#2&8#d!]CU5#67$(#12&
:+#(70<&#2(K
NADH + H

NAD

FP

FPH2

NADPH + H

NADP

FP

FPH2

#)

!"#$%&'()*+#,&,-*

67+#3"2%04-0*+$&#+&9),+(#21*#/40&#1+-*2$&#,+*2:0"$*+(#*72*#3/&1*$0&#2(#(/:(*-2*+(5#Z"2%04-0*+$&(
*2;+#7)8-0'+&#3-0,#d!]#2&8#*7+-+:)#'+*#-+8/1+85#67+#(2,+#3"2%04-0*+$&#,0"+1/"+#$(#-+0@$8$9+8#*0
:+10,+#21*$%+#2'2$&#*0#42-*$1$42*+#$&#3/-*7+-#-+21*$0&(5#67$(#12&#:+#(70<&#2(K
AH2

NAD

A
(Substrate)

NADH
+
+
H

FP-H2

FP
(Flavoprotein)

\+0@$82*$0&#03#3"2%04-0*+$&(#$(#2110,4"$(7+8#:)#1)*017-0,+#()(*+,5

!"@ 6'=/6)&*'(3+%&(
67+#$&$*$2"#(*+4#03#*7+#+"+1*-0&#*-2&(40-*#()(*+,#$(#*7+#0@$82*$0&#03#*7+#0-'2&$1#(/:(*-2*+#:)#d!]?#2&8
$&#*/-&#d!]#'+*(#-+8/1+8#*0#d!]U5#E*#,2)#:+#(70<&#2(#/&8+-K
G+*2:0"$*+#W#d!]#"#0@$8$9+8#,+*2:0"$*+#W#d!]U#W#UW
E&#*7+#2:0%+#-+21*$0&?#*7+#0@$8$9$&'#2'+&*#d!]#$(#-+8/1+8#*0#d!]U#:)#211+4*$&'#*<0#+"+1*-0&(
2&8# 0&+# 7)8-0'+&?# *7+# 0*7+-, 7)8-0'+&# 2*0,# $(# -+"+2(+8# 2(# 7)8-0'+&# $0&# XUWY5# I/17# -+21*$0&(
$&%0"%$&'# "0((# 03# 7)8-0'+&# 2&8# *<0# +"+1*-0&(# :)# *7+# ,0"+1/"+# *0# :+# 0@$8$9+8# 2-+# 12""+8
8+7)8-0'+&2*$0&(5# 67+# +&9),+(# <7$17# 12*2")9+# (/17# -+21*$0&(# 2-+# 12""+8##!+-#$&%!"31!15# I4+1$3$1
8+7)8-0'+&2(+(#2-+#-+./$-+8#30-#8$33+-+&*#0-'2&$1#,0"+1/"+(5
67+#d!]U#*7/(#30-,+8#$(#-+0@$8$9+8#*0#d!]#:)#2#3"2%04-0*+$&#+&9),+5#Z"2%$&#3/&1*$0&(#2(#2
10+&9),+5#67+#d!]U#$(#0@$8$9+8#*0#d!]#2&8#3"2%$&#211+4*(#*<0#+"+1*-0&(?#2#7)8-0'+&#2*0,#2&8#*7+
UW#3-0,#*7+#(0"/*$0&5
UW#W#d!]U#W#Z"2%$&#!#Z"2%$&#UV#W#d!]
67/(#3"2%$&#$(#-+8/1+8?#2&8#3/-*7+-#/&8+-'0+(#-+0@$82*$0&#:)#10+&9),+#?5#67+#-+21*$0&#$&%0"%+(
0@$82*$0&# 03# -+8/1+8# 3"2%$&# :)# *7+# *-2&(3+-# 03# *<0# +"+1*-0&(# 2&8# *<0# 7)8-0'+&(# *0# 2# ,0"+1/"+# 03
10+&9),+#?5
Flavin H2

Flavin
Coenzyme Q

E&# *7+# &+@*# -+21*$0&(?# -+0@$82*$0&# 03# 10+&9),+# ?# *2;+(# 4"21+# :)# 1)*017-0,+(# <7$17# 21*# 2(
+"+1*-0&# 12--$+-(5# 67+-+# 2-+# 2# &/,:+-# 03# (/17# 10,40/&8(# <7$17# 011/-# $&# *7+# ,$*0170&8-$2# 03# *7+
1+""(?#2&8#8$33+-#3-0,#+217#0*7+-#0&#*7+#:2($(#03#*7+$-#(*-/1*/-+#<$*7#-+(4+1*#*0#*7+#4-0*+$&#,0"+1/"+5
O<$&'#*0#*7+$-#(*-/1*/-2"#8$33+-+&1+(?#*7+)#2"(0#8$33+-#<$*7#-+(4+1*#*0#*7+$-#2:$"$*)#*0#211+4*#0-#80&2*+
+"+1*-0&(5# 67+)# -+21*# $&# 2# 42-*$1/"2-# 0-8+-?# 1)*017-0,+# @# "# 1)*017-0,+# (# "# 1)*017-0,+# 3# "

.#,&,-#/%&'01#2%3#,"+

#*

1)*017-0,+#3[5#>)*017-0,+#@#211+4*(#2&#+"+1*-0&#3-0,#10+&9),+#?#2&8#*7/(#'+*(#-+8/1+85#I$&1+#0&+
+"+1*-0&# $(# 211+4*+8# :)# 0&+# ,0"+1/"+# 03# 1)*017-0,+# @?# *<0# ,0"+1/"+(# 2-+# &+1+((2-)# *0# -+0@$8$9+
10+&9),+#?5#67+#7)8-0'+&(#03#*7+#-+8/1+8#10+&9),+#?#2-+#&0*##211+4*+8#:)#1)*017-0,+(#2&8#*7+)
2-+#-+"+2(+8#2(#7)8-0'+&#$0&(#$&#*7+#,+8$/,5
!&# +"+1*-0&# *7/(# 211+4*+8# :)# 1)*017-0,+# @# $(# &+@*# 42((+8# 0&# *0# (?# *7+&# *0# 3?# 2&8# 3$&2"")# *0
1)*017-0,+# 3[# <$*7# 2"*+-&2*+# 0@$82*$0&# 2&8# -+8/1*$0&# 03# *7+# $-0&# 2*0,# 03# *7+# 1)*017-0,+(5
>)*017-0,+#3[# $(# 10,,0&")# ;&0<&# 2(# 1)*017-0,+# 0@$82(+# <7$17# $(# 1242:"+# 03# /&8+-'0$&'# 8$-+1*
0@$82*$0&# :)# ,0"+1/"2-# 0@)'+&5# >)*017-0,+# 0@$82(+# $(# 2&# $,40-*2&*# +&9),+# (70<$&'# 2# (4+1$3$1
:+72%$0/-5#E&#*7+#4-+(+&1+#03#0@)'+&#$*#<0-;(#+33$1$+&*")#2&8#10&*$&/0/(")#*-2&(3+-(#+"+1*-0&(#$&#0&+
8$-+1*$0&?#*72*#$(#*0#0@)'+&5#E&#*7+#2:(+&1+#03#0@)'+&?#$*#:+10,+(#$&21*$%+5#67+#(+./+&1+#03#-+21*$0&(
$(#(70<&#$&#Z$'5#[5c5

!"A

'(')*/1)'5'3-'13(61&8%63+%?'10=&-0=&)/53+%&(

E3#0&+#,0"+#03#'"/10(+#$(#10,4"+*+")#0@$8$9+8?#QRQ#;12"#03#+&+-')#$(#-+"+2(+85#67$(#12&#:+#(70<&#*7/(K
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ATP

ADP

(1 mole)
glucose
(2 moles)
3-phosphoglyceric
acid
(2 moles)
2-phosphoglyceric
acid

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ATP
glucose 6-phosphate

2ATP
H2O

2ADP

ADP

(2 moles)
3-phosphoglycoraldehyde

fructose 1, 6-diphosphate

(2 moles)
1, 3-diphosphoglyceric
acid
2ADP
(2 moles)
phosphoenol pyruvate

2NAD.H2

2NAD

2P

ATP
(2 moles)
pyruvate

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##

Pyruvic acid
NAD
NADH2

3ATP

3ATP

NADH2

CoA

CO2
Acetyl CoA
CoA
2
Oxaloacetic
Citric acid
acid
10

NAD

Malic acid
H2O

FADH2

NAD

Fumaric acid

2ATP

Isocitric
acid

CITRIC ACID CYCLE

NADPH2
4
Oxalosuccinic acid

3ATP

FAD
Succinic acid
CoA

CO2
"-ketoglutaric
acid

7
CO2 6
CoA
GTP GDP
Succinyl
NAD
acid
Pi H2O
NADH2
1ATP
3ATP

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1+5&.,5&-#%/)&#$%&#/.$*)/$*+"#&"&(74#8:;>#$+#(&/.$#$+7&$%&(#/$#/"4+"&#1+1&"$:#!"#/#-5+6#(&/.$*+"#+"54
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8H*7:#;:M>:
Enzyme lowers energy barrier
so that EA! < EA

Heat raises energy

content of reactants
so that EA!! < EA

Reactants
C6H12O6 + 6O2

#G
Energy content
of reactants

U|
|V
||
WU
||
|
V|
||
|W

EA
(without
heat or
enzyme)

U|
V| EA" (with heat)
W
U|
V| EA! (with enzyme)
W

Products
6CO2 + 6H2O

Heating
This difference equals
(EA EA! ) since heating
raises the energy content
of the products as well
as that of the reactants

Progress of reaction

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/1*"+#/.*2-#8H*7:#;:;>T
Catalytic residues

Binding residues

Active site

Active site
Structural
residues

Enzyme

Non essential
residues

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Q6 IA28#-,(#(9&:/.$/&$,%,123#&.-$'+<'+,%.'-&'"&4'1#$)1,+
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U6 G#+'A.5,(#(9&%/#2&)(#&;PIP&,(&'A.5,-%

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V&D#%'&8+')<&*#%:##-&,4.-'&,-5&D#%'&,$.5
P6 D.-,(#(9&$,%,123#&%/#&%+,-("#+&'"&,&GIR&+,5.$,1

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Q6 7$21%+,-("#+,(#(9& $,%,123#& %/#& %+,-("#+& '"& ,$21X,$#%21

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P6 Y12$'(.5,(#(9&$,%,123#&812$'(.5.$&*'-5(
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&"341&# .+".&"$(/$*+"# /"2# *".(&/-*"7# -,0-$(/$&
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Enzyme concentration
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9:
P$# '+*"$# =# $%&# -,0-$(/$&# .+".&"$(/$*+"# *-# 1,.%
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1/V

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bit

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bito

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*$(-#"-$-,"+&/+#0"+%&.35+J-+$'';+#0"2"+$2"+$%&1#+:O+-$#12$''3+&,,122(-=+$*(-&+$,(.)+?0(,0+/$''+1-."2
#?&+,$#"=&2(")>+"))"-#($'+$-.+-&-P"))"-#($'+G@$%'"+Q5NH5
!"#$%& '() !""#$%&'()'$*)+,$-#""#$%&'()./&$,).0&*"
!""#$%&'(

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123#,$&$#
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@0"+ 92&,"))+ &/, -.!(&"!+&$", #$A")+ 9'$,"+ (-+ #0"+ '(7"2;+ A(.-"3+ $-.+ (-#")#(-$'+ *1,&)$;+ $'#0&1=0+ 12"$
/&2*$#(&-+()+,&-/(-".+#&+#0"+'(7"2+&-'35+J-+#0()+92&,"))+$**&-($+()+'(%"2$#".+(-+#0"+(-#")#(-"+$-.+#0"
A(.-"3;+$-.+=&")+(-#&+,(2,1'$#(&-+(-+#0"+/&2*+&/,='1#$*(-"5+R"23+'(##'"+$*&1-#+&/+$**&-($+()+/&1-.+(#0"+ )3)#"*(,+ ,(2,1'$#(&-5+ @0"+ $**&-($+ 92&.1,".+ $)+ $+ 2")1'#+ &/+ ."$*(-$#(&-+ &/+ $*(-&+ $,(.)+ ()
,&-7"2#".+(-#&+12"$+(-+#0"+'(7"2+?0(,0+()+#0"-+"<,2"#".+&1#5
NH2
R. CH

2H

R. C

NH
H2O

O
R. C

+ NH3

COOH

COOA

COOH

amino acid

imino acid

keto acid

/0!"'!(&"!+&$", (-7&'7")+ (-#"2,&-7"2)(&-+ &/+ $+ 9$(2+ &/+ $*(-&+ $,(.)+ $-.+ $+ 9$(2+ &/+ A"#&+ $,(.)
,$#$'34".+%3+#2$-)$*(-$)")+&2+$*(-&P#2$-)/"2$)")5+@0"+2"$,#(&-)+$2"+2"7"2)(%'"5
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+,(*$'-./0)(&'&10
R.CH (NH2) COOCH

R.CO.COOH
(keto acid)

*+,(&'()

NH3-to urea cycle

!-ketoglutarate

Glutamate
Transamination

13'$"'/&$'%&,$)3#'0%&,$E

CH3

CH3
CH

Transmination

NH2

CO +NH3

COOH

COOH
pyruvic acid

alanine

I)9$2#(,+ $,(.+ ()+ ,&-7"2#".+ #&+ &<$'&$,"#(,+ $,(.;+ ?0('"+ ='1#$*(,+ $,(.+ ()+ ,0$-=".+ #&+ $'90$
A"#&='1#$2(,+$,(.5
COOH

COOH
H 2O

CH2

CH2

CH2

NAD
NH2

CH
COOH

glutamic acid

*+,(&'(-

NADH
+H

CH2 + NH3

CO
COOH
!-Ketoglutaric acid

B,$F#3"&,$),G)@(8%'/&0)'0&*)%,)!-H#%,@(8%'3&0)'0&*)D5)@(8%'/'%#)*#25*3,@#$'"#E

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$**&-(1*+ )$'#)+ G$**&-(1*+ )$'#)+ $2"+ "<,2"#".+ (-+ *"#$%&'(,+ $,(.&)()H5+ D&?"7"2;+ *1,0+ $*&1-#+ &/
$**&-($+ ()+ "<,2"#".+ (-+ #0"+ /&2*+ &/+ 12"$5+ @0"+ ,&-7"2)(&-+ &/+ $**&-($+ (-#&+ 12"$+ #$A")+ 9'$,"+ (-+ #0"
'(7"25+@0"+/&2*$#(&-+&/+12"$+()+$+,&*9'"<+92&,"))+$-.+#$A")+9'$,"+7($+&2-(#0(-"+,3,'"+$)+92&9&)".+%3
S2"%)5+J-+#0"+),0"*"+&/+12"$+/&2*$#(&-+$2=(-(-"+()+03.2&'34".+%3+$-+"-43*"+$2=(-$)"+#&+3("'.+&-"
*&'",1'"+&/+12"$+$-.+&-"+&/+&2-(#0(-"5+@0"+."#$(')+&/+#0"+92&,"))+$2"+."),2(%".+(-+80$9#"2+NT5

/"9 2(4!%,&17-!')&*
B",$2%&<3'$#(&-+ ()+ $+ 92&,"))+ (-+ ?0(,0+ ,"2#$(-+ $*(-")+ $2"+ /&2*".+ %3+ #0"+ 2"*&7$'+ &/+ 8U:+ /2&*+ #0"
8UUD+G,$2%&<3'(,+=2&19H+&/+$*(-&+$,(.)5+I*(-")+$2"+903)(&'&=(,$''3+(*9&2#$-#5+B",$2%&<3'$#(&-+()
,$#$'34".+ %3+ $*(-&+ $,(.+ .",$2%&<3'$)")+ (-+ #0"+ 92")"-,"+ &/+ ,&"-43*"+ 932(.&<$'+ 90&)90$#"5+ @0"
2")1'#(-=+$*(-");+/&2+"<$*9'";+0()#(.(-"+3("'.)+0()#$*(-";+#32&)(-"+#32$*(-";+$-.+)"2(-"+3("'.)+"#0$-&'
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I'#0&1=0+ ='3,(-"+ ()+ #0"+ )(*9'")#+ $*(-&+ $,(.;+ (#+ ()+ $+ 92",12)&2+ &/+ #0"+ 2(-=+ )3)#"*)+ (-+ 912(-"+ $-.
9&29032(-)5+ V'3,(-"+ ,$-+ %"+ ,&-7"2#".+ (-#&+ )"2(-"+ ?0"-+ (#+ ,&*%(-")+ ?(#0+ /&2*$'."03."+ GD8DUH;+ $
)(-='"+,$2%&-+,&*9&1-.5+V'3,(-";+)"2(-";+$'$-(-"+$-.+='1,&)"+$2"+(-#"2,&-7"2#(%'"+GW(=5+M5XH5
Glycine
+
formaldehyde

Serine

Pyruvic
acid

3-Phosphoglycerate

Phosphoserine
transamination
3-Phosphohydroxy
pyruvic acid

Transamination
Alanine

*+,(&'(.

:$%#30,$F#3"&,$),G)@(50&$#I)"#3&$#)'$*)'('$&$#)J"02#/'%&0KE

F&*"+$*(-&+$,(.)+0$7"+#2"*"-.&1)+903)(&'&=(,$'+(*9&2#$-,"5+L0"-3'$'$-(-"+$-.+#32&)(-"+/$''+(#0()+,$#"=&23+?0(,0+0$7"+$2&*$#(,+-1,'"(5+@0")"+#?&+$*(-&+$,(.)+/&2*+#0"+92",12)&2)+&/+#0"+0&2*&-")
#032&<(-";+-&2$.2"-$'(-"+$-.+$.2"-$'(-"5+L0"-3'$'$-(-"+,$-+=(7"+2()"+#&+#32&)(-"+(-+#0"+%&.3;+%1#+#0()
()+$-+(22"7"2)(%'"+2"$,#(&-5+B"/",#(7"+*"#$%&'()*+&/+#0")"+$*(-&+$,(.)+,$1)")+$+.()"$)"+!%*!1+$"20.!
?0(,0+()+.1"+#&+$-+(-%&2-+="-"+"22&25+J-+#0()+.()"$)"+#0"+12(-"+#12-)+%'$,A+?0"-+"<9&)".+#&+$(25+I+="-"
*1#$#(&-+,$1)")+#0"+$%)"-,"+&/+3$($4."+&'!+.,$564."!'.,/2&*+#0"+,"'');+?(#0+#0"+2")1'#;+0&*&="-#()(,
$,(.+ $,,1*1'$#")+ (-+ #0"+ ,"'')+ $-.+ $99"$2)+ (-+ #0"+ 12(-"5+ J-+ $-&#0"2+ .()"$)";+ 13."6%7.+$"20&!), #0"
,&-7"2)(&-+ &/+ 90"-3'$'$-(-"+ #&+ #32&)(-"+ ()+ %'&,A".;+ ,&-)"E1"-#'3+ #0"+ 9(=*"-#+ *"'$-(-+ ()+ -&#
92&.1,".5+80('.2"-+."/(,("-#+(-+90"-3'$'$-(-"+)1//"2+/2&*+*"-#$'+(*%",('(#3+G/&2+*&2"+."#$(')+2"/"2+#&
,0$9#"2+&-+L03)(&'&=(,$'+V"-"#(,)H5
Kidney
Arginine

Glyco-cyamine
Glycine

Ornithine
(Liver)

In muscle
Creatinine
Creatine - P
H 2O + P 1

*+,(&'(/

L,3/'%&,$),G)03#'%&$#)G3,/)'3@&$&$#)&$)%2#)(&F#3)J"02#/'%&0KE

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82"$#(-"+()+*"#03'+=1$-(.(-"P$,"#(,+$,(.;+$-.+(#)+/&2*1'$+()>
YD:
Z
8 [ YD
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82"$#(-"+()+/&1-.+$%1-.$-#'3+(-+*1),'";+?0(,0+()+92&%$%'3+#0"+)(#"+&/+(#)+/&2*$#(&-5+J#+()+,&*9&)".
&/+$*(-&+$,(.)+='3,(-";+$2=(-(-"+$-.+*"#0(&-(-"5
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92&7(.")+"-"2=3+/&2+*1),'"+*"#$%&'()*;+/&2+2")3-#0")()+&/+$."-3'+932&90&)90$#";+)199'(")+"-"2=3+/&2
*1),'"+,&-#2$,#(&-5+82"$#(-"+$,,"9#)+90&)90$#")+.12(-=+='3,&'3)()5+60"-+,&*%(-".+?(#0+90&)90&2(,
$,(.;+,2"$#(-"+)"27")+$)+$+%1//"25
82"$#(-(-"+()+$-+$-03.2(."+&/+,2"$#(-"5+J#)+)#21,#12$'+/&2*1'$+()>
YD
Z
8 [ YD
Z
Y \ 8DX
Z
8D:
Z
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82"$#(-(-"

82"$#(-(-"+()+$-+"<,2"#&23+92&.1,#+$-.+)&'"'3+."2(7".+/2&*+,2"$#(-"+GW(=5+M5TH5+J#+()+/&1-.+(-+#0"
*1),'")+?0"2"+(#+()+)3-#0")(4".+$-.+"<,2"#".+(-+#0"+12(-"5+I%&1#+N5O+#&+N5Q+=*+&/+,2"$#(-(-"+9"2+.$3+()
"<,2"#".+ (-+ #0"+ 12(-"+ $-.+ (#+ ()+ (-."9"-."-#+ &/+ #0"+ 92&#"(-+ (-#$A"5+ I/#"2+ 0"$73+ *1),1'$2+ "<"2,()"
,2"$#(-(-"+&1#91#+()+#"*9&2$2('3+$,,"'"2$#".+?0(,0+)&&-+)#&9)+.12(-=+#0"+2",&7"23+9"2(&.5

/"/ .6-$86%#+('!,&-).+
F1'9012+()+$+,&-)#(#1"-#+&/+)1'9012+,&-#$(-(-=+$*(-&+$,(.)+,3)#"(-";+,3)#(-"+$-.+*"#0(&-(-"5+J#+()+$')&
/&1-.+ (-+ ='3,&92&#"(-)+ $)+ *1,&(#(-P)1'9012(,+ $,(.+ (-+ *1,(-";+ )1'90&'(9(.)+ (-+ -"27&1)+ #())1";+ &2+ $)
(-&2=$-(,+ )1'90$#")5+ F1'9012+ ()+ 92")"-#+ (-+ #())1"+ 92&#"(-);+ (-+ 0$(2);+ 0&2-)+ $-.+ /"$#0"2);+ (-+ *1,(-+ $)
*1,&(#(-)1'9012(,+$,(.;+(-+)&*"+='3,&92&#"(-)+&/+#"-.&-);+,&2-"$+$-.+,&--",#(7"+#())1")5
F*$''+$*&1-#)+&/+)1'9012P,&-#$(-(-=+$*(-&+$,(.)+$2"+1#('(4".+/&2+#0"+)3-#0")()+&/+#0"+(-)1'(-5+@0"
#2(9"9#(."+ ='1#$#0(&-"+ $-.+ #0"+ "\ *"2,$9#&"#03'$*(-";+ $+ ,&-)#(#1"-#+ &/+ ,&"-43*"+ I5+ J*9&2#$-#
9$#0?$3)+&/+)1'9012+*"#$%&'()*+$2"+)0&?-+(-+W(=5+M5Q5

!"#$%&'()*

"#'

" - mercaptoethylamine

Glutathione

CO2

Cystine

Cysteine

Pyruvate
Urea

NH3
H 2S

Cysteine
sulphuric acid

Thiosulphate

Cysteic
acid
CO2

H2SO3

+ Pyruvate

*+,(&'(0

H2SO4

Taurine

Thiosulphate
H2SO4

Mercapturic
acid

+ Cholic acid
Taurocholic
acid

98(;283)/#%'D,(&"/)J"02#/'%&0KE

!$C&2(#3+ &/+ )1'9012P,&-#$(-(-=+ $*(-&+ $,(.)+ $2"+ ,$#$%&'(4".+ (-+ #0"+ '(7"2+ 92&.1,(-=+ 12"$;+ $-.
)1'9012+()+&<(.(4".+$)+)1'9012(,+$,(.+#&+%"+"<,2"#".+$)+)1'90$#")+(-+#0"+12(-"5+@0"+)1'9012+,&*9&1-.)
$2"+1)"/1'+(-+#0"+/&''&?(-=+?$3)>
GNH F3-#0")()+&/+#())1"+92&#"(-)+'(A"+0$(2;+/"$#0"2);+"#,5
G:H F3-#0")()+&/+='1#$#0(&-";+#$12(-";+(-)1'(-;+)1'90$#(.");+"#,5
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90"-&'5
GTH W&2*$#(&-+&/+0"9$2(-5
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/"<

+('!,&-).+#&3#*64-(&$%&'()*

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,3#&9'$)*5+80"*(,$''3+#0"3+$2"+,&*9&)".+&/+)(*9'"+92&#"(-)+'(A"+92&#$*(-");+92&'$*(-")+&2+0()#&-")
,&-C1=$#".+?(#0+-1,'"(,+$,(.)+$-.+$2"+2(,0+(-+%$)(,+$*(-&+$,(.)5+@0"3+$2"+*$."+19+&/+%$)(,+%1('.(-=
%'&,A)+ ,$''".+ -1,'"&#(.")+ ,&-#$(-(-=+ 912(-"+ $-.+ 932(*(.(-"+ %$)");+ 9"-#&)"+ )1=$2)+ $-.+ 90&)90&2(,
$,(.5+@0"+912(-")+$2"+,&-7"2#".+#&+12(,+$,(.+$-.+932(*(.(-")+$2"+&<(.(4".+#&+92&.1,"+8U:+$-.+YDX5
@0"+ 9"-#&)"+ )1=$2)+ $2"+ (-+ #0"+ /&2*+ &/+ ."&<32(%&)"+ &2+ 2(%&)"+ 2")1'#(-=+ (-+ ."&<32(%&)"+ -1,'"(,+ $,(.
GBYIH+&2+2(%&)"+-1,'"(,+$,(.+G]YIH+2")9",#(7"'35+@0"+912(-"+%$)")+,&*92()"+$."-(-"+$-.+=1$-(-";
$-.+932(*(.(-")+,&-)()#+&/+#03*(-"+$-.+,3#&)(-"5+I''+#0")"+/&12+%$)")+$2"+92")"-#+(-+BYI;+?0"2"$)+(]YI+#03*(-"+()+2"9'$,".+?(#0+12$,('5

"#!

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Nucleoprotein

Protein

Nucleic acid
(DNA and RNA)
Nucleotides

Nucleosides

Bases
(Purines, Pyrimidines)

*+,(&'('

Phosphoric acid

Pentose sugars
(Deoxyribose, Ribose)

>5*3,(5"&"),G)$80(#,;3,%#&$")J"02#/'%&0KE

Y1,'"&#(.")+ $2"+ -1,'"&)(."+ 90&)90$#")5+ Y1,'"&)(.")+ $2"+ /&2*".+ ?0"-+ #0"+ 90&)90&2(,+ $,(.
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F"7"2$'+-1,'"&#(.")+/1-,#(&-+$)+,&"-43*")+?0(,0+$2"+."2(7$#(7")+&/+QP$."-3'(,+$,(.5+F&*"+&/+#0"
(*9&2#$-#+ ,&"-43*"+ -1,'"&#(.")+ ?0(,0+ #$A"+ 9$2#+ (-+ #0"+ *"#$%&'()*+ $2">+ -(,&#(-$*(."+ $."-(-"
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2D?FGH=>?@ABCD=B#!B=E
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#())1")+?0"2"+#0"+,"''+92&'(/"2$#(&-+()+*$<(*1*+$-.+2$9(.5

4IJI>?C=KL#?M#*ABCD=B#!B=E
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$,#(&-+&/+92&#"$)")5+@0"+)9",(/(,+"-43*");+."&<32(%&-1,'"$)"+$-.+2(%&-1,'"$)"+%2"$A+#0"+BYI+$-.

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2")9",#(7"'35+ `3+ )9",(/(,+ "-43*");+ $."-$)"+ $-.+ =1$-$)";+ $."-(-"+ $-.+ =1$-(-"+ $2"+ ,&-7"2#".+ #&
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Fat + Glucose + Urea

Glucocortin

Insulin
Glucocortin

Ox
ida
tio
n

Blood
glucose

Lowered by
glucocortin

Fats

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by insulin

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Glucogen
(1 , 4 and 1 , 6 glycosyl
units)

UDP

Pi

Branching enzyme
1,4 Glycosyl
units
Phosphorylase
Glucose
S

Glycogen
primer
Uridine
diphosphate
glucose (UDPG)

Debranching
enzyme

Glycogen
synthetase

Cyclic AMP

S
Insulin

Glucose
from
debranching
enzyme

Glucagon
cpinephrine

Touronic
pathway
UDPG Pyrophosphorylase
PPi

Uridine
triphosphate
(UTP)

Glucose 1 phosphate

Mg

++

Phosphoglucomultase
Glucose 6 phosphate

H2O
S Simulation
I Inhibition

Glucose-6
phosphate

ADP
Mg++ Glucokinase

Pi

ATP
Glucose

*+,(&'(3

4(50,@#$#"&"E

to
HMP
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!"#$%&'()*

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*1),'"+ ='3,&="-+ ()+ /$2+ =2"$#"2+ (-+ $*&1-#+ #0$-+ #0"+ '(7"2+ ='3,&="-5+ f(7"2+ ()+ #0"+ )#&2$="+ &2=$-+ /&2
='3,&="-;+?0"2"$)+*1),'"+='3,&="-+$,#)+$)+$+)&12,"+&/+"-"2=3+.12(-=+,&-#2$,#(&-5+F#$27$#(&-+.&")+-&#
$//",#+#0"+*1),'"+='3,&="-5+F3-#0")()+&/+='3,&="-+(-+#0"+*1),'"+#$A")+9'$,"+(-+#0"+)$*"+*$--"2+$)+(,$)"+&/+'(7"25+!1),'"+,$--&#+,&-7"2#+='3,&="-+#&+='1,&)"+)(-,"+#0"+)9",(/(,+"-43*"+2"E1(2".+/&2+(#)
,&-7"2)(&-+()+$%)"-#+(-+#0"+*1),'"5+D&?"7"2;+='3,&="-+()+%2&A"-+.&?-+#&+'$,#(,+$,(.+(-+#0"+*1),'"
/2&*+?0"2"+(#+()+,$22(".+#&+#0"+'(7"2+#02&1=0+#0"+%'&&.)#2"$*+#&+%"+,&-7"2#".+(-#&+='3,&="-5

/"09 +('!,&-).+#&3#&'8(%#.6P!%.
V'1,&)";+ =$'$,#&)"+ $-.+ /21,#&)"+ $2"+ )1=$2)+ &/+ =2"$#+ *"#$%&'(,+ (*9&2#$-,"5+ @0")"+ )1=$2)+ "-#"2+ #0"
*"#$%&'(,+ 9$#0?$3)+ $/#"2+ %"(-=+ 90&)90&23'$#".5+ L0&)90&23'$#(&-+ &/+ ='1,&)"+ 0$)+ $'2"$.3+ %""."),2(%".+(-+,&--",#(&-+?(#0+='3,&'3)()5+@0"+)&12,"+&/+=$'$,#&)"+()+*$(-'3+#0"+'$,#&)"+,&-#"-#+&/+#0"
*('A5+V$'$,#&)"+()+/(2)#+90&)90&23'$#".+(-+#0"+92")"-,"+&/+$+)9",(/(,+"-43*"+4!%!*+$7&"!'.)$-.+#0"-+(#
2"$,#)+ ?(#0+ 12(.(-"+ .(90&)90&=$'$,#&)"+ #&+ /&2*+ 12(.(-"+ .(90&)90&='1,&)"+ ?0(,0+ *$3+ 9$2#(,(9$#"+ (='3,&="-+)3-#0")()5+f$2="+E1$-#(#(")+&/+=$'$,#&)"+(-+%'&&.+$2"+A-&?-+#&+%"+#&<(,+$-.+,$1)"+$+.()"$)"
4!%!*+$'.(&!9
W21,#&)"+()+&%#$(-".+/2&*+)1,2&)"+$-.+/21(#)+$-.+()+2"$.('3+,&-7"2#".+(-#&+='1,&)"+&2+='3,&="-+(#0"+'(7"25+J#+()+90&)90&23'$#".+(-+#0"+92")"-,"+&/+/21,#&A(-$)"+(-#&+/21,#&)"PNP90&)90$#"5+@0"+)"E1"-,"
&/+"7"-#)+()+)0&?-+%"'&?+GW(=5+M5NNH5
U,,$)(&-$''3+ ='3,"2$'."03."+ ()+ &<(.(4".+ #&+ ='3,"2&'5+ V'3,"2$'."03."+ ,$-+ $')&+ %"+ 2".1,".+ #&
='3,"2$'."03."PXP90&)90$#"+G#2(&)"H;+$-.+#?&+*&'",1'")+&/+#2(&)"+,$-+%"+,&-7"2#".+#&+/21,#&)"PN;+MP
.(90&)90$#"5

4"#3!'#+('!,&-).+
W$#)+$-.+'(9(.)+$2"+(*9&2#$-#+,&-)#(#1"-#)+&/+#0"+92&#&9'$)*5+@0"3+*$3+%"+92")"-#+(-+#0"+/&&.)+&2+*$3
%"+ /&2*".+ (-+ #0"+ %&.35+ 6"+ 0$7"+ ."),2(%".+ #0"+ ,0("/+ ,'$))")+ &/+ /$#)+ $-.+ '(9(.)+ (-+ 80$9#"2+ :5+ @0"

!"#$%&'()*

""%

D fructose
)#21,#12$'+/$#)+$2"+7"23+,&*9'"<+,&*9&1-.);+?0"2"$)+2")"27"+/$#)
&2+."9&#+/$#)+$2"+#0"+&2.(-$23+/$#)+?0(,0+*$A"+19+#0"+%1'A+&/+#0"
Fructokinase
%&.35+B"9&#+/$#)+$2"+(-+#0"+/&2*+&/+-"1#2$'+#2(='3,"2(.")+?0(,0+,$ATP
%"+ 03.2&'34".+ #&+ /&2*+ *&-&='3,"2(.")+ $-.+ /2""+ /$##3+ $,(.)5
@2(='3,"2(.")+ *$3+ %"+ 2")3-#0")(4".+ /2&*+ #0"+ /$##3+ $,(.)+ (-+ #0"
Fructose 1 phosphate
,"'')+&/+(-#")#(-$'+*1,&)$5
J-+#0"+*"#$%&'()*+&/+/$#;+#02""+*$C&2+92&,"))")+$2"+(-7&'7".>
Aldolase
GNH !&%('(4$#(&-+ &/+ /$#+ /2&*+ )#&2$="+ /$#+ ."9&#)+ &/+ #0"+ %&.3; Dihydroxyacetone
?0(,0+'$#"2+#$A"+9$2#+(-+,$#$%&'()*5
D Glyceraldehyde
G:H I%)&29#(&-+&/+.(=")#".+/$#)5
+
GXH F3-#0")()+&/+/$#)+(-+#0"+'(7"2;+/2&*+(-#")#(-$'+*1),&)$+$-.
NAD
Aldehyde
#0"+$.(9&)"+#())1"+,$2%&03.2$#"+$-.+92&#"(-+)&12,")5
dehydrogenase
!&%('"+ /$#+ ()+ (-+ #0"+ /&2*+ &/+ *(-1#"+ &('3+ .2&9'"#)+ A-&?-+ $)
+
NAD+H
*36%$(&*0$"',?0(,0+#2$7"'+(-+#0"+%'&&.)#2"$*5+803'&*(,2&-)+$2"
D Glycerate
*$."+ 19+ &/+ -"1#2$'+ /$#);+ $-.+ $')&+ ,&-)()#+ &/+ 90&)90&'(9(.)+ G/$#
9'1)+90&)90$#"H;+,0&'")#"2&'+$-.+,0&'")#"2&'+")#"2)+&/+/$##3+$,(.)5
*+,(&'()) <#%'D,(&"/),G)G380%,"#E
@0"+ ,03'&*(,2&-)+ $2"+ $%)&2%".+ %3+ #0"+ '(7"2+ ?0"2"+ #0"3+ $2"
03.2&'34".+ =(7(-=+ 2()"+ #&+ /2""+ ='3,"2&')+ $-.+ /$##3+ $,(.)5+ !&%('(4$#(&-+ &/+ /$#)+ ,$-+ %"+ ,&-7"-("-#'3
&%)"27".+(-+,$)"+&/+)#$27(-=+$-(*$')5+I/#"2+)#$27$#(&-+/&2+$+)0&2#+9"2(&.;+#0"+='3,&="-+2")"27"+&/+#0"
'(7"2+()+."9'"#".5+F(-,"+-&+*&2"+&/+,$2%&03.2$#")+$2"+)3-#0")(4".+G"<,"9#+%3+?$3+&/+='1,&-"&="-")()H;
#0"+'(7"2+.&")+-&#+2","(7"+(#)+,$2%&03.2$#"+)199'35+e-."2+)1,0+,&-.(#(&-);+'$2=";+$*&1-#)+&/+/$#)+$2"
#2$-)9&2#".+#&+#0"+'(7"2+?0(,0+#$A"+9$2#+(-+*"#$%&'()*5

/"0: %&-(#&3#-)S(%#)*#3!'#+('!,&-).+
f(7"2+0$)+$+A"3+2&'"+(-+#0"+*"#$%&'()*+&/+/$#)5+J#+0$)+%""-+92&7".+%"3&-.+.&1%#+#0$#+(-+,&-.(#(&-)+&/
,$2%&03.2$#"+."9'"#(&-;+*&)#+&/+#0"+/$#+&/+#0"+%&.3+()+*&%('(4".+#&+#0"+'(7"2+#&+92&7(."+$-+$'#"2-$#(7"
)&12,"+ &/+ "-"2=35+ f(7"2+ ()+ -&#+ -&2*$''3+ $-+ $,,1*1'$#&2+ &/+ /$#)+ $)+ (#+ ()+ /&2+ ,$2%&03.2$#")5+ @0"+ /$#
,&-#"-#+(-+#0"+'(7"2+()+*$(-#$(-".+1-(/&2*'3+,&-)#$-#+%"#?""-+X+$-.+_+9"2+,"-#5+@01)+"<,"))+&/+/$#
."9&)(#)+ ()+ #2$-)/&2*".+ %3+ #0"+ '(7"2+ (-#&+ 1)"/1'+ )1%)#$-,")+ #02&1=0+ 7$2(&1)+ (-#"2,&-7"2)(&-)
GW(=5+M5N:H5
`")(.")+(-#"2,&-7"2)(&-+&/+92&#"(-);+/$#)+$-.+,$2%&03.2$#");+?0(,0+=&")+&-+(-+#0"+'(7"2;+(#+()+$')&
2")9&-)(%'"+ /&2+ #2$-)/&2*$#(&-)+ &/+ '(9(.)+ (-#&+ 90&)90&'(9(.)+ $-.+ ,0&'")#"2&';+ .")$#12$#(&-+ &/+ /$##3
$,(.);+ &<(.$#(&-+ &/+ /$##3+ $,(.);+ "#,5+ @0"+ )'1==()0+ /1-,#(&-+ &/+ #0"+ '(7"2+ 2")1'#)+ (-+ )"7"2$'+ *"#$%&'(,
.()&2."2);+?0(,0+*$3+%"+-&2*$''3+.1"+#&+#0"+"//",#+&/+,"2#$(-+9&()&-);+/$#P2(,0+.("#;+92&#"(-P9&&2+.("#;
."/(,("-,3+&/+7(#$*(-)+$-.+$+0&)#+&/+&#0"2+,$1)")5+@0"2"/&2";+?"+,$-+)$3+#0$#+'(7"2+()+#0"+92".&*(-$-#
&2=$-+?0(,0+*$(-#$(-)+#0"+0"$'#03+)#$#"+&/+$-+(-.(7(.1$'5

""&

+,(*$'-./0)(&'&10

Galactose

ATP
Galactokinase

Galactose 1 phosphate

Uridine diphosphate
glactose

Uridine diphosphate
glucose
Glycogen
phosphorylase
glucose 1 phosphate

Glucose

Fructose

*+,(&'()-

ATP
Glucokinase
ATP
Fructokinase

Glucose 6 phosphate

Glucose
phosphate

Fructose 6 phosphate

:$%#30,$F#3"&,$),G)0'3D,25*3'%#")&$)%2#)(&F#3E

/"0; &1)2!')&*#&3#3!''7#!4)2.
W$#);+")9",($''3+#2(='3,"2(.");+$2"+03.2&'3)".+#&+#0"(2+,&-)#(#1"-#)+/$##3+$,(.)+$-.+='3,"2&'+%"/&2"+#0"3
92&,"".+ #&+ ,$#$%&'(,+ 9$#0?$35+ !1,0+ &/+ #0"+ /$#+ 03.2&'3)()+ #$A")+ 9'$,"+ (-+ $.(9&)"+ #())1";+ ?0"2"+ /2""
/$##3+ $,(.)+ GWWIH+ $2"+ 92&.1,".+ #&+ %"+ ,$22(".+ (-#&+ #0"+ 9'$)*$+ GW(=5+ M5NXH5+ @0"+ WWI+ $,(.)+ 2"$,0+ #0"
#())1")+G'(7"2;+A(.-"3;+0"$2#;+*1),'";+#")#();+%2$(-;+$-.+$.(9&)"+#())1"H+?0"2"+&<(.$#(&-+#$A")+9'$,"5+@0"
'&-=+ ,0$(-+ /$##3+ $,(.)+ $2"+ )3)#"*$#(,$''3+ %2&A"-+ .&?-+ #&+ :P,$2%&-+ 1-(#)+ (-+ #0"+ /&2*+ &/+ g$,#(7"
$,"#$#")h5+ @0"+ $,"#$#"+ $-.+ #0"+ '&-=+ ,0$(-+ /$##3+ $,(.)+ $2"+ *"#$%&'()".+ #02&1=0+ $+ ,&**&-+ 9$#0?$3;
2"E1(2(-=+I@L+$-.+8&I+"-43*"+GW(=5+M5NTH5
CH3

CH3
CHOH
CH2

Oxidation
Reduction

COOH
"-Hydroxybutyrate

CO
CH2

CH3
Decarboxylation

CO

+ CO2

CH3

COOH
Acetoacetate

Acetone

@0"+$,"#3'+8&I+,$-+"(#0"2+,&*%(-"+?(#0+&<$'&$,"#$#"+%"/&2"+"-#"2(-=+#0"+,(#2(,+$,(.+,3,'";+&2+(#
*$3+%"+.(2",#'3+&<(.()".+#&+$,"#&$,"#$#";+#0"+/(2)#+A"#&-,+%&.35+I,"#&$,"#$#"+*$3+/12#0"2+%2"$A.&?#&+"P03.2&<3%1#32$#"+$-.+$,"#&-";+?0(,0+$,,1*1'$#"+(-+#0"+'(7"2+(-+)*$''+$*&1-#)5+@0"+'(7"2+#())1"
,$--&#+&<(.()"+$,"#&$,"#$#"5+I,"#&-"+$2()")+/2&*+$,"#&$,"#$#"+%3+)9&-#$-"&1)+.",$2%&<3'$#(&-5

!"#$%&'()*

""'

Carbohydrates

Fats

Glycerol

Triose phosphate

Fatty acid

Pyruvate

Acetate
Oxidized to
AcetylCoA

Oxaloa
c

Acetylations
Acetoacetate

CO2

etic acid

CO2 + H2O
Citrate

Acetone
" Hydroxybutyrate

*+,(&'().

902#/'%&0)3#;3#"#$%'%&,$),G)/#%'D,(&"/),G)G'%")'$*)0'3D,25*3'%#"E

ur

n,

io

at

CH3CO ~ CoA
Throlysis

at

RCO ~ CoA

es

Citric
acid
cycle

(CoA derivative)
RCH2CH2COCoA
(active and acid)

1 ATP
CoA

(Sat. fatty acid)

R CH2 CH2 COOH
(Acetyl CoA)

5
(Unsaturated fatty acid)

RCO CH2CO ~ CoA

sis
oly

RCH OH CH2CO ~ CoA

(Hydroxy fatty acid CoA)

dr

Hy

Dehydrogenation

RCH CH CO ~ CoA
Enzymes
(1) Thiokinase
(2) Acyldehdrogenase
(3) Enol hydrase
(4) " hydroxy acyldehydrogenase
(5) Thiolase

*+,(&'()/

<#%'D,(&"/),G)G'%%5)'0&*"E

@0"+1#('()$#(&-+&/+='3,"2&'+()+."9"-."-#+&-+#0"+$,#(7$#(-=+"-43*";+4%6*.0$%,7&"!'.),?0(,0+()+/&1-.
(-+)1//(,("-#+$*&1-#)+(-+'(7"2;+A(.-"3;+(-#")#(-";+%2&?-+$.(9&)"+#())1"+$-.+'$,#$#(-=+*$**$23+='$-.)5

/"0/ !TR1)2!')&*#&3#3!''7#!4)2.
W$##3+ $,(.)+ $2"+ *$(-'3+ &<(.()".+ %3+ $+ 92&,"))+ ,$''".+"P&<(.$#(&-;+ $+ ),0"*"+ 92&9&)".+ %3+ S-&&9+ (NcOT5+ D"+ 9&)#1'$#".+ #0$#+ )(-,"+ *$C&2(#3+ &/+ -"1#2$'+ /$#)+ ,&-#$(-+ "7"-+ -1*%"2+ &/+ ,$2%&-+ $#&*);
"P&<(.$#(&-+()+#0"+*&2"+'(A"'3+92&,"))+(-+?0(,0+#0"+/$##3+$,(.+*&'",1'"+()+%2&A"-+.&?-+(-+$+)#"9?()"
*$--"2;+2"*&7(-=+:P,$2%&-+$#&*)+/2&*+#0"+,$2%&<3'+"-.+(-+"$,0+)#"9;+3("'.(-=+$,"#$#"+"E1(7$'"-#)5

""!

+,(*$'-./0)(&'&10
O
Fatty acid

R R CH2 CH2 COH

CoA SH

ATP
++

Thiokinase

Mg

1
AMP + PPi
Acyl CoA
O
(Active fatty acid) R CH CH C~S CoA
2
2
Fp(Flavoprotein)
Acyl CoA
2~P
Dehydrogenase
Fp H2
H 2O
2
O Respiratory chain
A " Unsaturated R CH = CH C~S CoA
Acyl CoA
Enoyl
H2O
hydrase
3
O
OH
"hydroxy
RCHCH2C~S CoA
acyl CoA
3~ P
"hydroxyacyl
NAD
++
CoA dehydrogenase
NADH+H
H 2O
4
Respiratory chain
O
O
"Ketoacyl
RCCH2C~S CoA
CoA
Thiolase
" Ketothiolase

CoA SH
5

O
O
RC~SCoA + CH3C~S CoA
AcylCoA
AcetylCoA
Citric
acid
cycle
2CO2

*+,(&'()0

"-,?&*'%&,$E

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='1,&)"+ $-.+ &#0"2+ ,&*9'"<+ ,$2%&03.2$#")+ $)+ 9"2+ -"".)+ &/+ #0"+ $-(*$'5+ F&*"+ (*9&2#$-#+ )#"9)+ &/+"P
&<(.$#(&-+),0"*"+$2"+)0&?-+(-+W(=5+M5NQ5
J#+*1)#+%"+%&2-"+(-+*(-.+#0$#+/$##3+$,(.+&<(.$#(&-+#$A")+9'$,"+(-+#0"+*(#&,0&-.2($;+%1#+%"/&2"+WI
"-#"2)+ #0"+ *(#&,0&-.2($+ (#+ 0$)+ #&+ %"+ *$."+ 2"$.3+ /&2+ &<(.$#(&-+ 2"$,#(&-)5+ @0"+ WI+ (-+ #0"+ ,3#&)&'+ ()
$,#(7$#".+ %3+ $+ *&'",1'"+ &/+ I@L+ (-+ #0"+ 92")"-,"+ &/+ $,3',&"-43*"+ IG8&IFDH5+ @0"+ 2"$,#(&-+ &,,12)
"(#0"2+ (-+ #0"+ "-.&9'$)*(,+ 2"#(,1'1*+ &2+ $#+ #0"+ &1#"2+ *(#2&,0&-.2($'+ *"*%2$-";+ 2")1'#(-=+ (-+ #0"
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*(#&,0&-.2($+?(#0+#0"+0"'9+&/+*!0"&+&".),$+,$22("2+*&'",1'"5+@0()+2"$,#(&-+()+,$#$'3)".+%3+$-+"-43*";
$,3'+ 8&I+ #2$-)/"2$)"5+ U-,"+ #0"+ /$##3+ $,3'+ 8&I+ "-#"2)+ #0"+ *(#&,0&-.2($'+ *$#2(<;+ #0"2"+ /&''&?)+ #0"
2"*&7$'+&/+:+03.2&="-+$#&*)+/2&*+#0"+!,$-.+"+,$2%&-);+,$#$'3)".+%3+$+."03.2&="-$)";+2")1'#(-=+(-

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&/+0(=0+90&)90$#"+%&-.+"-"2=3+Gc__j:XTO+$+NOOH+&/+#0"+#&#$'+"-"2=3+&/+,&*%1)#(&-+&/+#0"+/$##3+$,(.5

&F=EIJ=?@#?M#6@KIJAHIJDE#3IJJG#!B=EK
`&.3+'(9(.)+$2"+2(,0+(-+1-)$#12$#".+/$##3+$,(.)+$-.+#0")"+$2"+&<(.()".+*&2"+)'&?'35+F&*"+"<$*9'")+$2">
9$'*(#&'"(,+$,(.+GNM>NH;+&'"(,+$,(.+GN_>NH;+'(-&'"(,+$,(.+GN_>:H;+'(-&'"-(,+$,(.+GN_>XH+$-.+$2$,0(.&-(,
$,(.+G:O>TH5+I''+.&1%'"+%&-.)+(-+-$#12$''3+&,,122(-=+1-)$#12$#".+/$##3+$,(.)+$2"+(-+*&'P,&-/(=12$#(&-5
L$'*(#&'"(,+$-.+&'"(,+$,(.)+$2"+-&#+"))"-#($'+$)+#0"3+,$-+%"+/&2*".+(-+#0"+%&.3;+%1#+#0"+2"*$(-(-=+#02""
$,(.)+,&*"+1-."2+#0"+.''."+&!%,=!++6,!*&-',,$#"=&23+$-.+0$7"+#&+%"+)199'(".+(-+#0"+.("#5+U<(.$#(&-+&/
1-)$#12$#".+/$##3+$,(.)+92&,"".)+#0"+1)1$'+"P&<(.$#(7"+9$#0?$3+1-#('+#0"+.&1%'"+%&-.+()+2"$,0".5+@0"
.&1%'"+%&-.+(-+*&';,&-/(=12$#(&-+()+-&#+71'-"2$%'"+#&+"-43*(,+$##$,A+1-'"))+(#+()+()&*"2()".+#&++0!"'P
,&-/(=12$#(&-5+ L&'31-)$#12$#".+ /$##3+ $,(.);+ )1,0+ $)+ '(-&'"(,;+ $2$,0(.&-(,+ "#,5+ $2"+ *&2"+ ,&*9'"<+ $-.
2"E1(2"+ $..(#(&-$'+ "-43*"+ /&2+ &<(.$#(&-5+ @0"3+ $2"+ -&2*$''3+ /&1-.+ $)+ )#21,#12$'+ ,&*9&-"-#)+ ($))&,($#(&-+ ?(#0+ ,0&'")#"2&'+ $-.+ 90&)90&'(9(.)+ G"5=5+ *"*%2$-")+ $-.+ 2"92&.1,#(7"+ &2=$-)H5+ J*$**$');+$2$,0(.&-(,+$-.+)&*"+2"'$#".+8P:O+/$##3+$,(.)+$2"+A-&?-+#&+=(7"+2()"+#&+1-(E1"+,&*9&1-.)
'(A"+10$'+!4%!"-&"',?0(,0+0$7"+0&2*&-"P'(A"+$,#(7(#35

"")

+,(*$'-./0)(&'&10

/"0< +('!,&-).+#&3#P-74(%&U-"+ &/+ #0"+ 03.2&'3)()+ 92&.1,#)+ &/+ #2(='3,"2(.")+ ()+ ='3,"2&'+ ?0(,0+ ()+ *"#$%&'()".+ &2+ 1#('()".+ (&2=$-)j#())1")+?0"2"+)9",(/(,+"-43*"+='3,"2&'+A(-$)"+()+$%1-.$-#'3+92")"-#5+8"2#$(-+&2=$-)+)1,0+$)
'(7"2;+A(.-"3;+(-#")#(-$'+*1,&)$+$-.+'$,#$#(-=+*$**$23+='$-.)+$2"+2(,0+(-+#0"+"-43*";+?0('"+*1),'")
$-.+$.(9&)"+#())1"+,&-#$(-+7"23+'(##'"+$,#(7(#35+V'3,"2&'+()+92".&*(-$-#'3+,&-7"2#".+(-#&+,$2%&03.2$#"
#02&1=0+='3,"2&'+90&)90$#";+/&2*".+%3+$+)9",(/(,+='3,"2&'+A(-$)"+$#+#0"+"<9$-)"+&/+I@L5
CH2OH

CH2OH

CHOH + ATP

CHOH + ADP

CH2OH

CH2OP

Glycerol

Glycerol phosphate
(triose phosphate)

V'3,"2&'+90&)90$#"+()+#0"-+&<(.()".+#&+#2(&)"90&)90$#"+%3+$+='3,"2&'+90&)90$#"+."03.2&="-$)"
?0(,0+1'#(*$#"'3+/&2*)+='3,&="-+#02&1=0+='3,&="-")()5+@2(&)"90&)90$#"+*$3+%";+0&?"7"2;+&<(.()".
#&+93217(,+$,(.+%3+?$3+&/+='3,&'3)()+GW(=5+M5NMH5+J-+.($%"#(,+&2+90'&220(4(-()".+$-(*$');+='3,"2&'+()
,&-7"2#".+$'*&)#+E1$-#(#$#(7"'3+#&+='1,&)"5

*+,(&'()'

L'%#),G)@(50#3,()/#%'D,(&"/E

/"0O .7*'8(.).#&3#P-74(%)2(.#!*2#3!''7#!4)2.
J#+ 0$)+ %""-+ A-&?-+ /&2+ $+ '&-=+ #(*"+ #0$#+ /$#)+ $2"+ )3-#0")()".+ /2&*+ *"#$%&'(#")+ )1,0+ $)+ $,"#$#"+ $-.
$,"#&$,"#$#"5+W$#)+$2"+$')&+)3-#0")()".+/2&*+92&#"(-+$-.+,$2%&03.2$#"+)&12,")5+I''+-$#12$''3+&,,122(-=
/$##3+$,(.)+9&))"))+"7"-+-1*%"2+&/+,$2%&-+$#&*)K+#01)+(#+()+'&=(,$'+#0$#+/$##3+$,(.)+*1)#+%"+)3-#0")()".
/2&*+ :P,$2%&-+ /2$=*"-#)5+ J#+ 0$)+ %""-+ )0&?-+ #0$#+ #0"+ )#$2#(-=+ *$#"2($'+ ()+ $,"#3'P8&I+ ?0(,0+ ,$-+ %"
."2(7".+/2&*+93217$#"5+I'#"2-$#(7"'3;+93217$#"+,$-+$')&+%"+."2(7".+/2&*+/2"";+$,"#$#"+?0"-+(#+2"$,#)
?(#0+,&"-43*"+I+$-.+I@L5

.G@JNDK=K#?M#PCGBDH=EDK
@2(='3,"2(.")+$2"+/&2*".+%3+2"$,#(&-)+%"#?""-+$,3'P8&I+,&*9&1-.)+$-.+%P='3,"2&90&)90$#";+?0(,0
()+ /&2*".+ %3+ )9",(/(,+ ='3,"2&'+ A(-$)"5+ @0"2"+ $2"+ &#0"2+ "-43*")+ ?0(,0+ ,$#$'3)"+ #0"+ /&2*$#(&-+ &/
*&-&i$-.+.(90&)90$#(.(,+$,(.)+$#+#0"+"<9"-)"+&/+/$##3+$,3'+8&I+."2(7$#(7")>

!"#$%&'()*

""*

CH2 OR1
CHOR2
CH2 OP

OH
O
OH

Phosphatidic acid

J-+ #0"+ -"<#+ )#$="+ #0"+ 90&)90$#"+ =2&19+ ()+ 2"*&7".+ %3+ $+ )9",(/(,+ 90&)90$#$)"+ /&''&?".+ %3
2"9'$,"*"-#+%3+$+#0(2.+/$##3+$,3'+2")(.1">

CHOR2
OH
O
CH2OP
OH
Phosphatidic acid

CH2 OR1

CH2 OR1

CH2 OR1
+H2O

CHOR2
CH2OH
Diglyceride

+ Acyl CoA

CHOR2
CH2OR3
Triglyceride

.G@JNDK=K#?M#3IJJG#!B=EK
W$##3+ $,(.+ &<(.$#(&-+ &,,12)+ (-+ #0"+ *(#&,0&-.2($;+ %1#+ /$##3+ $,(.+ )3-#0")()+ #$A")+ 9'$,"+ -&#+ &-'3+ (*(#&,0&-.2($+%1#+$')&+(-+*(#&,0&-.2($P/2""+)3)#"*)5+@0"+9$#0?$3+/&2+)3-#0")()+()+-&#+"<$,#'3+2"7"2)$'
&/+"P&<(.$#(&-+),0"*";+%1#+(#+(-7&'7");+)&*"+*&.(/(,$#(&-)5+e-."2+$-$"2&%(,+,&-.(#(&-);+*(#&,0&-.2($
,$#$'3)"+#0"+(-,&29&2$#(&-+&/+$,"#3'P8&I+1-(#)+(-#&+'&-=+,0$(-+/$##3+$,(.)+G7(45+)#"$2(,+$,(.;+9$'*(#(,
$,(.H;+2"E1(2(-=+I@L;+YIBD+$-.+YIBLD5+F3-#0")()+(-+"<#2$P*(#&,0&-.2($'+)3)#"*;+")9",($''3+(-+#0"
'(7"2;+ %2$(-;+ A(.-"3+ "#,5;+ $,"#3'P8&I+ 1-(#)+ $2"+ (-,&29&2$#".+ (-#&+ /$##3+ $,(.);+ ,$#$'3)".+ %3+ ,3#&)&'(,
"-43*")+$-.+,&/$,#&2)+)1,0+$)+I@L;+YIBLD+$-.+!=:d+&2+!-:d+(&-)5+@0()+)3)#"*+()+."9"-."-#+&8U:+)199'(".+%3+%(,$2%&-$#"5+80$(-+"'&-=$#(&-+1)1$''3+#$A")+9'$,"+(-+#0"+*(,2&)&*")5

/"0Q +('!,&-).+#&3#$8&.$8&-)$)2.
L0&)90&'(9(.)+$2"+/&1-.+(-+$''+,"'')+$-.+$2"+)3-#0")()".+"(#0"2+/2&*+90&)90$#(.(,+$,(.+&2+90&)90$#(.3'
,0&'(-"5+ f",(#0(-+ ()+ #0"+ *&)#+ (*9&2#$-#+ 90&)90&'(9(.+ (-+ #0"+ %&.3+ $-.+ ()+ )3-#0")()".+ (-+ #0"+ '(7"25
F3-#0")()+&/+90&)90&'(9(.)+/2&*+/$#)+(-7&'7")+*&%('()$#(&-+&/+/$#)+(-+$-.+&1#+&/+#0"+,"'')5+D"-,"+#0"
$,#1$'+ )(#")+ $2"+ '(7"2;+ (-#")#(-"+ $-.+ A(.-"35+ L0&)90&'(9(.)+ $2"+ '$2="'3+ /&1-.+ (-+ ,&*%(-$#(&-+ ?(#0
92&#"(-)+$-.+$2"+#2$-)9&2#".+(-+#0"+%'&&.+(-+#0"+/&2*+&/+92&#"(-+,&*9'"<")5
@0"+*&)#+,&**&-+$*&-=+90&)90&'(9(.)+$2"+'",(#0(-)+$-.+,"90$'(-);+?0('"+?0"2"+$2"+&#0"2)+(?0(,0+%$)")+$2"+2"9'$,".+%3+)"2(-"+&2+(-&)(#&'5+8$#$%&'()*+&/+'",(#0(-+()+$,,&*9'()0".+(-+#0"+/&''&?(-=
*$--"2+GW(=5+M5NaH>
F90(-=&*3"'(-)+ $2"+ 90&)90&'(9(.)+ ,&-#$(-(-=+ $+ /$##3+ $,(.;+ 90&)90&2(,+ $,(.;+ ,0&'(-";+ $-.+ $
,&*9'"<+&/+$*(-&+$',&0&';+)90(-=&)(-";+%1#+$2"+."7&(.+&/+='3,"2&'5+I%-&2*$'+E1$-#(#(#(")+&/+90&)90&P
$-.+ )90(-=&'(9(.)+ (-+ ,"2#$(-+ #())1");+ ")9",($''3+ (-+ #0"+ -"27&1)+ )3)#"*;+ ,$1)"+ .()"$)")+ '(A"
)90(-=&'(9(.&)");+$-.+."*3'"(-$#(-=+.()"$)")+#0$#+$2"+(-0"2(#".5

/"5R +('!,&-).+#&3#48&-(.'(%&80&'")#"2&'+&,,12)+(-+7$2(&1)+%&.3+#())1")+$-.+(-+#0"+9'$)*$+"(#0"2+(-+#0"+/2""+/&2*+&2+)#&2$="+/&2*+$)
'&-=+,0$(-+/$##3+$,(.+")#"2)+%"'&-=(-=+#&+#0"+,'$))+&/+)#"2&')5+J#+()+$')&+$-+(*9&2#$-#+,&*9&-"-#+&/+#0"

"$#

+,(*$'-./0)(&'&10
Lecithin
RCOOH

Phospholipase A

Lysolecithin
RCOOH

Lysophospholipase

Glyceryephosphoryl choline
Glycerylphosphoryl
choline esterase

! - Glycerophosphate + Choline

*+,(&'()1

B'%'D,(&"/),G)(#0&%2&$E

*"*%2$-"+)3)#"*+&/+#0"+,"''+$-.+()+#0"+92",12)&2+&/+)#"2&(.+0&2*&-");+"<,2"#".+/2&*+#0"+%&.3+(-+#0"
/&2*+&/+%('"+$,(.)+G)$'#)H5+@0"+,0&'")#"2&'+,&-#"-#+&/+#0"+%'&&.+2$-=")+/2&*+NQO+\+:QO+*=jNOO+*'5

.?AHBDK#?M#4N?CDKJDH?C#=@#JND#,?EG
60('"+*$C&2+9$2#+&/+#0"+%&.3+,0&'")#"2&'+G$%&1#+N+=j.$3H+()+)3-#0")(4".+(-+#0"+%&.3;+$+)*$''+9&2#(&-+()
G$%&1#+O5X+=j.$3H+92&7(.".+%3+7$2(&1)+/&&.)+(-#$A"+)1,0+$)+"==+3&'A;+*"$#;+'(7"2;+$-.+%2$(-+$-.+(#)
19#$A"+ ()+ #02&1=0+ '&?+ ."-)(#3+ '(9&92&#"(-)+ GfBf)H5+ F3-#0")()+ &,,12)+ /2&*+ $,"#3'P8&I+ G#?&P,$2%&1-(#)H+92&.1,".+%3+%"#$+&<(.$#(&-+&/+/$##3+$,(.)+(-+#0"+'(7"2;+(-#")#(-"+$-.+$'*&)#+$''+,"'')5+I%&1#+aOO
*=j.+()+)3-#0")(4".+%3+$+?"''+2"=1'$#".+*",0$-()*5+W2""+,0&'")#"2&'+()+2"*&7".+/2&*+#0"+%&.3+%3
9'$)*$+0(=0P."-)(#3+'(9&92&#"(-)+GDBf)H5

.G@JNDK=K#?M#4N?CDKJDH?C
I'#0&1=0+,0&'")#"2&'+()+)3-#0")(4".+%3+*$-3+#())1")+)1,0+$)+$.2"-$'+,&2#"<;+)A(-;+(-#")#(-"+#")#")+"#,5
'(7"2+()+,&-)(."2".+#&+%"+#0"+*$(-+)(#"5+@?&+*&'",1'")+&/+$,"#3'P8&I+,&-."-)"+#&+/&2*+$,"#&$,"#3'P
8&I;+?0(,0+$=$(-+2"$,#)+?(#0+$-&#0"2+*&'",1'"+&/+$,"#3'P8&I+#&+/&2*+XP03.2&<3PXP*"#03'='1#$23'P
8&I+GD!VP8&IH;+'"$.(-=+#&+#0"+)3-#0")()+&/+*"7$'&-$#"+GW(=5+M5N_H5+J#+*1)#+%"+-&#".+#0$#+$''+,$2%&$#&*)+ &/+ ,0&'")#"2&'+ &2(=(-$#"+ /2&*+ $,"#3'P8&I+ "-43*"5+ !"7$'&-$#"+ ()+ #0"+ ,21,($'+ ,&*9&1-.+ $-.
#02&1=0+$+)"2(")+&/+2"$,#(&-);+=(7")+2()"+#&+,0&'")#"2&'5

'HI@KV?HJ#I@E#(FBHDJ=?@#?M#4N?CDKJDH?C
80&'")#"2&';+(-+$))&,($#(&-+?(#0+&#0"2+'(9(.);+()+$%)&2%".+(-+#0"+(-#")#(-"+$-.+#0"2"$/#"2+(-,&29&2$#".
(-#&+,03'&*(,2&-)+$-.+7"23+'&?P."-)(#3+'(9&92&#"(-)+GRfBf)H5+I+=2"$#"2+9$2#+&/+,0&'")#"2&'+G$%&1#
_OkH+(-+#0"+'3*90+1-."2=&")+")#"2(/(,$#(&-+?(#0+'&-=P,0$(-+/$##3+$,(.)+$-.+#2$-)9&2#".+$)+'(9&92&#"(-)
(-+#0"+9'$)*$5+D(=0")#+92&9&2#(&-+&/+,0&'")#"2&'+()+/&1-.+(-+#0"+fBf5+W2""+,0&'")#"2&'+(-+#0"+9'$)*$+()

!"#$%&'()*

"$"

2 CH3COSCoA
(Acetyl CoA)
CoA
CH3COCH2COSCoA
(Acetoacetyl CoA)
CH3CoSCoA
OH

CoA
CH3CCH2CoSCoA
CH2COOH
(3-Hydroxy-3-methylgutaryl CoA)
2NADPH + H+
2NADP+
OH

CoA HMG CoA reductase

CH3CCH2CH2OH
CH2COOH
(Mevalonic Acid)

To synthesis of cholesterol

*+,(&'()2

9%#;")&$)%2#)"5$%2#"&"),G)/#F'(,$&0)'0&*I)%2#);3#083",3),G)02,(#"%#3,()"5$%2#"&")J"%#;")$,%)"2,M$KE

"E1('(%2$#".+ ?(#0+ #0"+ ,0&'")#"2&'+ ")#"2)+ $-.+ $%&1#+ 0$'/+ &/+ #0"+ /2""+ ,0&'")#"2&'+ ()+ + "'(*(-$#".+ %3
"<,2"#(&-+(-+#0"+/$",");+?0('"+2"*$(-."2+()+"<,2"#".+$)+-"1#2$'+)#"2&(.5
@0"+'(7"2+,&-#$(-)+$+9&&'+&/+1-")#"2(/(".+,0&'")#"2&'+/2&*+?0"2"+(#+()+#2$-)9&2#".+#&+#0"+(-#")#(-"
$-.+#0"-+%$,A+#&+'(7"2+#02&1=0+,03'&*(,2&-)5+80&'")#"2&'+()+"<,2"#".+/2&*+#0"+%&.3+$)+,0&'")#"2&'+&2
%('"+$,(.)5+A$10$'+!"$%+()+#0"+*$(-+)#"2&'+)",2"#".+(-+#0"+/$",");+$+92&.1,#+&/+,0&'")#"2&';+$2()(-=+/2&*
#0"+(-#")#(-$'+%$,#"2($+$,#(7(#35+!1,0+&/+#0"+,0&'")#"2&'+)",2"#".+(-+#0"+%('"+()+"(#0"2+2"$%)&2%".+&2+="#)
."9&)(#".+(-+#0"+$2#"2(")+(-+#0"+/&2*+&/+")#"2)+&2+*$3+9$2#(,(9$#"+(-+)#"2&(.+)3-#0")()5

/"50 W('&P(*(.).
B12(-=+ ,&-.(#(&-)+ &/+ )#$27$#(&-+ $-.+ .($%"#");+ &<(.$#(&-+ &/+ /2""+ /$##3+ $,(.)+ ()+ "-0$-,".+ (-+ #0"
*(#&,0&-.2($5+ J-+ 7"2#"%2$#");+ #0"+ $,"#3'P8&I+ &2(=(-$#(-=+ /2&*+ #0"+ "P&<(.$#(&-+ ,$-+ '"$7"+ #0"
*(#&,0&-.2(&-+ (-+ #0"+ /&2*+ &/+ ,(#2$#"5+ D&?"7"2;+ $+ *$C&2+ 9$2#+ &/+ #0()+ $,"#3'P8&I+ ()+ 1)".+ /&2+ #0"
)3-#0")()+ &/+ 7.+$"., #$-&.'+ (-+ #0"+ '(7"2+ G7.+$'&'H5+ @0"2"+ $2"+ #02""+ #39")+ &/+ A"#&-"+ %&.(")>+ $,"#&-";
$,"#&$,"#(,+ $,(.;+ $-.+ XP03.2&<3%1#32(,+ $,(.;+ ?0(,0+ $2"+ ="-"2$#".+ #02&1=0+ $+ 92&,"))+ ,$''".
7.+$4.".'&';+#$A(-=+9'$,"+(-+#0"+*(#&,0&-.2($5

3?HLIJ=?@#?M#WDJ?@D#,?E=DK
f&-=P,0$(-+ /$##3+ $,(.+ $2"+ #2$-)9&2#".+ #&+ #0"+ (--"2+ *(#&,0&-.2($'+ *"*%2$-"+ #02&1=0+ ,$2-(#(-"
."2(7$#(7")5+"P&<(.$#(&-+&/+/$##3+$,(.)+#$A")+9'$,"+(-+#0"+*(#&,0&-.2(&-;+2"'"$)(-=+$,"#3'P8&I;+?(#0

"$$

+,(*$'-./0)(&'&10

#0"+,&-)"E1"-#+92&.1,#(&-+&/+'$2="+$*&1-#+&/+I@L5+@0"+(-(#($'+)#"9)+$2"+)(*('$2+#&+#0&)"+&/+,0&'")#"2&'
)3-#0")()+GW(=5+M5NcH5+@0"+XP03.2&<3PXP*"#03'='1#$23'P8&I+/&2*".+()+)9'(#+(-#&+$,"#3'P8&I+$-.+/2""
$,"#&$,"#(,+$,(.5
@0"+ $,"#&$,"#(,+ $,(.+ ()+ "(#0"2+ )9&-#$-"&1)'3+ .",$2%&<3'$#".+ #&+ $,"#&-"+ &2+ 2".1,".+ #&+ XP
03.2&<3%1#32(,+$,(.>
CH3COCH2COOH
(Acetoacetic acid)

CO2 + CH3COCH3
(Acetone)

CH3COCH2COOH + NADH + H

CH3CHOHCH2COOH + NAD
(3-Hydroxybutyric acid)

f&-=P,0$(-+/$##3+$,(.)+G"7"-+-1*%"2H+92&.1,"+$,"#3'P8&I+1-(#)+#02&1=0+%"#$+&<(.$#(&-;+%1#+&..
,0$(-+/$##3+$,(.)+92&.1,"+$,"#3'P8&I+$-.+92&9(&-3'P8&I;+?0(,0+()+='1,&="-(,5+L"2&<()&*")+&<(.(4"
'&-=P,0$(-+/$##3+$,(.)5

WDJ?@D#,?E=DK#.DHXD#IK#(@DHUG#.?AHBD
I,"#&$,"#(,+$,(.+$-.+XP03.2&<3%1#32(,+$,(.+$2"+92&.1,".+(-+'$2="+$*&1-#+$-.+$2"+1#('(4".+$)+"-"2=3
)&12,"+ %3+ 7$2(&1)+ #())1")+ )1,0+ $)+ *1),'");+ A(.-"3+ $-.+ #0"+ %2$(-+ G"<#2$0"9$#(,+ #())1")H5+ I,"#&$,"#(,
$,(.+.(//1)")+/2""'3+$,2&))+#0"+,"''+*"*%2$-")+$-.+,$--&#+%"+2"$,#(7$#".+1-'"))+#0"3+2"$,0+#0"+,3#&)&'
?0"2"+#0"3+,$-+9$2#(,(9$#"+(-+,0&'")#"2&'+)3-#0")()+$)+0$99"-)+?(#0+$,"#&$,"#$#"5
I,"#&$,"#$#"+()+2"$,#(7$#".+(-+"<#2$0"9$#(,+#())1")>
I,"#&$,"#$#"+d+F1,,(-3'P8&I+#+ F1,,(-(,+ $,(.+ d+ I,"#&$,"#3'P8&I5+ I,"#&$,"#3'P8&I+ )9'(#)+ %3
#0(&'3)()+(-#&+#?&+*&'",1'")+&/+$,"#3'P8&I+?0(,0+"-#"2)+#0"+S2"%)+,3,'"5+S"#&="-")()+()+$-+"<#2"*"'3
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(2 Acetyl CoA)
CoA
CH3COCH2COSCoA
(Acetoacetyl CoA)
H2O + Acetyl CoA
CoA
CH2COOH
CH3COH
CH2COSCoA
(3-Hydroxy-3-methylglutaryl CoA)
Acetyl CoA
CH3COCH2COOH
(Acetoacetate)
NANDH + H
NAD
OH
CH3CCH2COOH
H
(3-Hydroxybutyrate)

*+,(&'()3

CO2

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CH3COCH3
(Acetone)

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follicle

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Muscles of the mucosa
Submucosa

Lymphatic
plexus

Circular muscle

Auerbach!s
plexus

Longitudinal muscle
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(a)

Surface coat

Microvilli

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

Lumen

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Apoferritin
Fe

+++

Plasma

Apoferritin
Ferritin

+++

+++

++

Fe

++

Ligand
Fe Complex
Fe

*+,(&'()4

+++

++

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} Transferrin
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Fe

++

Low molecular
Weight ligands

++

Fe

Fe chelate

Fe Complex
Macromolecules

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Solution-1
1
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1
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Solution-2

1
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#$$+")'%5)%04()A.5)-%28,"+-%$.39-%(6$%6$8,"6$+%.5%$4"*")(.$%4(6'$%8,()."."$2%5#%$;3$22%2(4.9%?56%./"2-%./$
7/(4$2%2$$*%.5%0522$22%*56$%057$6#,4%A"+)$12%./()%./52$%06$2$).%")%/,*()%&$")'29

!"#$%&'

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!"#$#%"&'$()*)+,'-*./('$.#(&'$$*0(1$'.)'()*)+,'-*./(.2,,#2-0('$$("-0"3"02'$(&*$$.('-0(4#,)(&$#.*0
&#)1',5)*-5.6( 78*( )*)+,'-*( ".( '.9))*5,"&/( 8"%8$9( 3".&#2.( '-0( 09-')"&( "-( -'52,*( 5#( 1,#3"0*
.*$*&5"3*(1*,)*'+"$"59(#:"-%(5#(58*(1,*.*-&*(#4("#-;&8'--*$.('-0("#-(12)1.6(<-(1$'-5./("5($"*.("-5*,-'$
5#(58*(&*$$(:'$$('-0(*-&$#.*.(58*(&595#1$'.)(#4(58*(&*$$6(!*."0*.(5,'-.1#,5(#4(&*$$2$',()'5*,"'$./("#-./
:'5*,('-0()'&,#)#$*&2$*./(58*()*)+,'-*.('$.#(42-&5"#-("-(5,'-.)*)+,'-*(."%-'$"-%('-0(&*$$;5#;&*$$
"-5*,'&5"#-.6(=*4"&"*-&"*.("-(&*$$()*)+,'-*(,*.2$5("-('(3',"*59(#4(0".*'.*./(:8"&8(:"$$(+*(0*'$5(:"58"&8'15*,(>?(#-(189."#$#%"&'$(0".#,0*,.6

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@*)+,'-*.( ',*( 4#2-0( 5#( &#-.".5.( #4( )'"-$9( $"1"0./( 1,#5*"-./( .#)*( &',+#890,'5*./( '-0( :'5*,6( 78*
1,#5*"-(5#($"1"0(,'5"#(3',"*.(%,*'5$96(A#,(*B')1$*/(58*("--*,()"5#&8#-0,"'$()*)+,'-*(8'.(CDE(1,#5*"-/
:8"$*(58*()9*$"-()*)+,'-*(#4(-*2,#-.(1#..*..(#-$9(FGE(1,#5*"-6(78*(0"44*,*-&*("-(1,#5*"-;$"1"0(,'5"#
'&&#2-5.(4#,(58*(.1*&"4"&(42-&5"#-.(58*(#,%'-*$$*.(8'3*(5#(1*,4#,)6

045678+9:+'8;<74=8+*>?>@A
H"1"0.("-(58*()*)+,'-*(',*('(&#)1$*B()"B52,*(#4(&8#$*.5*,#$('-0(4'559('&"0./()'"-$9("-(58*(4#,)(#4
%$9&*,"0*.( '-0( 18#.18#$"1"0.6( I8#$*.5*,#$( ".( :"0*$9( 0".5,"+25*0( "-( 58*( )*)+,'-*/( 58#2%8
18#.18#$"1"0.(1,*0#)"-'5*6(!*&'2.*(#4("5.(8"%8(18#.18#$"1"0(&#-5*-5/(58*()9*$"-(#4(58*(-*,3*(&*$$.
&'-(*$*&5,"&'$$9("-.2$'5*(58*(&*$$(4,#)("5.(*-3"#,-)*-56(78*($"1"0(&#)1#."5"#-(3',"*.(')#-%(0"44*,*-5
)*)+,'-*.6

1>B8+5C?8A+9:+-D9A?D9E>?>@A+196=@+>=+'8;<74=8A
A"3*( ")1#,5'-5( 591*.( #4( 18#.18#$"1"0.( "-&$20*( 18#.18'5"0"&( '&"0/( $*&"58"-/( 18#.18'5"09$( "-#."5#$/
18#.18'5"09$( .*,"-*/( '-0( 18#.18'5"09$( *58'-#$')"-*( JA"%6( K6FL6( 78#2%8( '$$( )*)+,'-*.( &#-5'"-( '

!"#$%&'"()*+,-./.0+

"#$

Phosphatidyl choline
CH2 OCOR
CH2 OCOR
2

R COO

C
CH2

R COOCH
OH

CH2

O
O

OH

OCH2 CH2 N(CH3)3

Phosphatidyl aminoethanol
1

3 sn-Phosphatidic acid

CH2 OCOR
2

R COO CH
Phosphatidyl inositol
1
CH2 OCOR

CH2

R COO CH
CH2

O
O

P
OH

OH

OH

OH

OH

O
O

O
H

OCH2 CH2 NH33

Phosphatidyl serine (Cephalin)

CH2 OCOR

OH

R COO CH
CH2

O
O

P
OH

!"#$%&$'

OCH2 CH2 N(CH3)3

COO

!"#$%#&'$%()*&'+,-)./)#$.0#$.1%#%203)0$.4%*5)$62,.#$%1%()&*2)$62,.#$.7%(),-5%.*08

.2+.5'-5"'$( 1,#1#,5"#-( #4( 18#.18#$"1"0./( 1,*0#)"-'-5( +*"-%( 18#.18#%$9&*,"0*.( 8'3"-%( '( %$9&*,#$
+'&M+#-*6(N$$()*)+,'-*(18#.18#$"1"0.(',*(!"#$%#!&$%'/(8'3"-%(+#58(890,#18"$"&('-0(890,#18#+"&
1#,5"#-.( JA"%6( K6>L6( O58*,( $"1"0.( "-&$20*( &8#$*.5*,#$/( %$9&#$"1"0./( 18#.18'5"09$&8#$"-*( '-0
.18"-%#)9*$"-6(P18"-%#)9*$"-($'&M.('(%$9&*,#$(+'&M+#-*/(+25(Q2"5*(&#))#-("-(1$'.)'()*)+,'-*.6
<-.5*'0( #4( '( %$9&*,#$( +'&M+#-*( "5( &#-5'"-.( .18"-%#."-*/( '-( ')"-#( '$&#8#$( :"58( '( $#-%( 2-.'52,'5*0
890,#&',+#-(&8'"-6(N(4'559('&9$(."0*(&8'"-(".($"-M*0(5#(58*(')"-#(%,#21(#4(.18"-%#."-*(+9('-(')"0*
+#-0( 5#( '( &*,')"0*6( 78*( 5*,)"-'$( 890,#B9$( %,#21( #4( .18"-%#."-*( ".( *.5*,"4"*0( 5#( 18#.18#&8#$"-*/
)'M"-%(58*(890,#18#+"&(8*'0(#4(.18"-%#)9*$"-(.")"$',(5#(58'5(#4(18#.18'5"09$&8#$"-*6
N()'R#,(0"44*,*-&*(')#-%(18#.18#$"1"0.(&#-&*,-.(58*(&8',%*(&',,"*0(+9(58*(1#$',(8*'0(%,#21.('5
-*25,'$(1S6(P#)*(18#.18#%$9&*,"0*.(J18#.18'5"09$&8#$"-*/(18#.18'5"09$(*58'-#$')"-*L(8'3*(-#(-*5
*$*&5,"&(&8',%*T(#58*,.(J&',0"#$"1"-/(18#.18'5"09$.*,"-*L(8'3*('(-*5(-*%'5"3*(&8',%*6(N(4*:(,',*($"1"0.
&',,9('(-*5(1#."5"3*(&8',%*('5(-*25,'$(1S6(U#-*58*$*../(58*(1#$',(8*'0(%,#21.("-('$$(18#.18#$"1"0.(&'1'&M(5#%*58*,("-5#(58*(&8','&5*,".5"&(+"$'9*,(.5,2&52,*6(P18"-%#)9*$"-('-0(%$9&#$"1"0.(',*(.")"$',(".8'1*('-0(&'-(4#,)(+"$'9*,(:"58(58*)6

$D9E8A5879E+4=+(;?9754=5+$9=A5>568=5+9:+'8;<74=8+*>?>@A
I8#$*.5*,#$('-("5.(0*,"3'5"3*.(&#-.5"525*('-#58*,(")1#,5'-5(&$'..(#4()*)+,'-*($"1"0./(58*(.5*,#"0.6(78*
+'."&( .5,2&52,*( #4( 58*( .5*,#"0.( ".( 58*( 4#2,;,"-%( 890,#&',+#-( JA"%6( K6?L6( I8#$*.5*,#$( ".( 58*( )'R#,

"#%

1'-#&/()*+,-./.0+
CH2N (CH2)3
Hydrophilic Head

CH2
Special region

Phosphate

O P

O
CH2CH CH2

Fatty acid tail

OC

CH2 CH2
CH2 CH2
CH2 CH
CH2 CH
CH2

CH2

CH2
CH2

Hydrophobic Tail

CH3

!"#$%&$(

CH2
CH2
CH3

9',+('+,-)./)#$.0#$.1%#%20)#,-0-*')%*)'$-)#1&0"&)"-"7,&*08

CH2

CH3 CH2
CH2
CH
CH3
CH2
CH
CH2
CH2
C
CH2
CH
CH2 3 CH
CH
CH2
C
CH

CH
HO

C
CH2

CH3
CH
CH3

CH2
CH

Cholesterol (full structure)

!"#$%&$)

9',+('+,-)./)($.1-0'-,.1)4$.0-)2-,%:&'%:-0)&,-)#,-0-*')%*)'$-)"-"7,&*-8

&#-.5"52*-5( J'( .5*,#"0L( #4( '-")'$( 5"..2*.T( #58*,( .5*,#"0.( 1$'9( ")1#,5'-5( ,#$*.( "-( 1$'-5.6( N$58#2%8
&8#$*.5*,#$(".(*-5",*$9('(890,#&',+#-("-(&#)1#."5"#-/("5(".(')18"1'58"&(+*&'2.*("5(&#-5'"-.('(890,#B9$
%,#21(58'5("-5*,'&5.(:"58(:'5*,6(I8#$*.5*,#$(".(*.1*&"'$$9('+2-0'-5("-(1$'.)'()*)+,'-*(#4()'))'$"'&*$$./(+25(4#2-0(5#('+.*-5(4,#)()#.5(1,#M',9#5"&(&*$$.6

!"#$%&'"()*+,-./.0+

"#&

-7958>=A+478+4EA9+:96=@+>=+'8;<74=8A
V,#5*"-.(4#,)(58*(+2$M(#4(&*$$()*)+,'-*/(*B".5"-%('.(%(&)*+!,(#,(#)+%#$)+!,(1,#5*"-.6

(=58F74E+'8;<74=8+-7958>=A
@'-9(#4(58*(1,#5*"-.('..#&"'5*0(:"58(58*(1$'.)'()*)+,'-*(',*(5"%85$9(+#2-0(5#("5/(:8"$*(.#)*(',*
'55'&8*0(5#($"1"0.("-(58*(+"$'9*,6(<-(#58*,.;58*(&+!(-")".+!()/#+0&)%(-/(58*(1#$91*15"0*(&8'"-('&52'$$9
5,'3*,.*.(58*($"1"0(+"$'9*,6(N$$(W;1,#5*"-(&#21$*0(,*&*15#,.(J*6%6(,*&*15#,.(#4(1*15"0*(8#,)#-*./('-0
#0#2,.L/(*'&8(.1'-(58*(1$'.)'()*)+,'-*(.*3*-(5")*.(JA"%6(K6XL6(<-('$$(58*.*(&'.*./(58*(1,#5*"-(:"58(58*
$"1"0(+"$'9*,(&#-.".5.(1,")',"$9(#4(890,#18#+"&(')"-#('&"0.6(78*.*(',*(2.2'$$9(',,'-%*0("-('-('$18'
8*$"B( .#( 58'5( 58*( 1#$',( YI( Z( O( '-0( YUS( %,#21.( '5( 58*( 1*15"0*( +#-0.( &'-( "-5*,'&5( :"58( *'&8( #58*,
,'58*,(58'-(:"58(58*",(890,#18#+"&(.2,,#2-0"-%.6(78#.*(1#,5"#-.(#4(58*(1#$91*15"0*(58'5(1,#R*&5(#25
4,#)(58*(+"$'9*,(5*-0(5#(8'3*('(8"%8(1*,&*-5'%*(#4(890,#18"$"&(')"-#('&"0.6(A2,58*,)#,*/(58#.*(58'5
1,#R*&5( "-5#( 58*( 'Q2*#2.( .2,,#2-0"-%.( #4( 58*( &*$$( ',*( 2.2'$$9( %$9&#1,#5*"-./( :"58( )'-9( 890,#18"$"&
.2%',( ,*."02*.( '55'&8*0( 5#( 58*( 1',5( #4( 58*( 1#$91*15"0*( *B1#.*0( '5( 58*( .2,4'&*( #4( 58*( &*$$6( P#)*
5,'-.)*)+,'-*(1,#5*"-.(58'5(.1'-(58*(+"$'9*,(.*3*,'$(5")*.(4#,)('(890,#18"$"&(&8'--*$(58,#2%8(:8"&8
&*,5'"-("#-.('-0()#$*&2$*.(&'-(*-5*,(#,($*'3*(58*(&*$$(JA"%6(K6[L6
NH2
LIGAND

Plasma membrane
Cytoplasm

IV

III
II

V
I

VI
VII
T

"
GDP

COOH

!"#$%&$*

!);)#,.'-%*<(.+#1-2),-(-#'.,8)=')(.*0%0'0)./)0-:-*)',&*0"-"7,&*-)#,.'-%*0)4%'$)><'-,"%*&1)#,.?-('%*5)'.)'$-@'-,%.,8

-87>?D874E+'8;<74=8+-7958>=A
V*,"18*,'$()*)+,'-*(1,#5*"-.(',*(58#.*(58'5('08*,*(#-$9($##.*$9(5#(58*()*)+,'-*(:"58(:8"&8(58*9(',*
'..#&"'5*06(78*.*()#$*&2$*.(0#(-#5(.1'-(58*($"1"0(+"$'9*,(&#,*(#4(58*()*)+,'-*/(+25('55'&8("-0",*&5$9/
591"&'$$9( +9( +"-0"-%( 5#( "-5*%,'$( )*)+,'-*( 1,#5*"-.6( 78*,*4#,*/( 58*( .#( &'$$*0( ,*%2$'5#,9( 1,#5*".2+2-"5.( #4( )'-9( "#-( &8'--*$.( '-0( 5,'-.)*)+,'-*( ,*&*15#,./( 4#,( *B')1$*/( )'9( +*( 0*4"-*0( '.
1*,"18*,'$( )*)+,'-*( 1,#5*"-.6( 78*.*( 1,#5*"-./( "-( &#-5,'.5( 5#( "-5*%,'$( )*)+,'-*( 1,#5*"-./( 5*-0( 5#

"#'

1'-#&/()*+,-./.0+

Peripheral
membrane protein

Transmembrane
protein

Phospholipid
bilayer

Integral
membrane protein

Peripheral
membrane
protein

!"#$%&$0

Q.#.5,&#$%(&1)&,,&*5-"-*')./)#,.'-%*0)%*)'$-)#1&0"&)"-"7,&*-8

&#$$*&5( "-( 58*( :'5*,;.#$2+$*( 4,'&5"#-( 02,"-%( 1,#5*"-( 12,"4"&'5"#-6( 78*.*( ',*( )#,*( $##.*$9( '..#&"'5*0
:"58( 58*( )*)+,'-*6( 78*9( ',*( 2.2'$$9( '55'&8*0( -#-&#3'$*-5$9( 5#( 58*( 1,#5,20"-%( 1#,5"#-.( #4( "-5*%,'$
)*)+,'-*(1,#5*"-.6(@*)+,'-*(1,#5*"-.(',*(#45*-(,*.5,"&5*0("-(58*",()#3*)*-5.(JA"%6(K6[L6

$47<9DC@7458A+478+)EA9+-78A8=5+>=+5D8+'8;<74=8
I',+#890,'5*.("-(58*(1$'.)'()*)+,'-*(',*(1,*.*-5("-(58*(4#,)(#4(&#3'$*-5$9;$"-M*0()#$*&2$*.(:"58
1,#5*"-.('-0($"1"0.6(78*,*(5:#(591*.\(*,1'0#+0&)%(-('-0(*,1'0,%#%2-6(78*(&#))#-(.2%',.('..#&"'5*0
:"58(58*(1,#5*"-.(',*(=;%$''2&#.*/(=;%'&5#.*/(=;)'--#.*(*5&6/(:8"&8(',*(#$"%#.'&&8',"0*(&#)1$*B*.6
!*."0*.( .")1$*( .2%',./( .2%',( 0*,"3'5"3*.( .2&8( '.( U;'&*59$;=;%$2&#.')"-*( '-0( U;'&*59$;-*2,')"-"&
'&"0(J."'$"&('&"0L(',*('$.#(1,*.*-56(N$)#.5('$$(1,#5*"-.(1,*.*-5(#-(58*(#25*,(.2,4'&*(#4(58*()*)+,'-*
8'3*(-#5(%#5(&',+#890,'5*(&#)1#-*-5/('.(:"$$(+*(.**-(4,#)(58*(7'+$*(K6F6
+,-./%&$' A&,%&'%.*)%*)B,.'-%*C)D%#%2)&*2)E&,7.$62,&'-)E."#.0%'%.*)./)9."-)F-"7,&*-0
!"#$%&'"

(%)*"+',-

.+/+0,-

1&%$)230%&*",-

F6-1%*
L+"&*)-,6'$,.(6'-

GH
MJ

IJ
MK

K
H

F%'.($.*2,%&1)%**-,)"-"7,&*-

IJ

NM

4#)"$&)#1&0"&)"-"7,&*-

PM

MN

!"G '&'23)0&+)3$%(/&$/H3&
@*)+,'-*( 8'.( '( 09-')"&( .5,2&52,*6( N$58#2%8( 58*,*( ',*( $',%*( 3',"'5"#-.( "-( 58*( &#)1#."5"#-( #4
)*)+,'-*/(58*(+'."&(.5,2&52,'$(2-"5(#4(3",52'$$9('$$(+"#$#%"&'$()*)+,'-*.(".(58*(#$0-#$0,%#%2/.%,!1)+6
78".(+"$'9*,(".('(.8**5;$"M*(.5,2&52,*(&#)1#.*0(#4(5:#($'9*,.(#4(18#.18#$"1"0()#$*&2$*.(:8#.*(1#$',
8*'0( %,#21.( 4'&*( 58*( .2,,#2-0"-%( :'5*,( '-0( 58*",( 4'559( '&"0( &8'"-.( 4#,)( '( &#-5"-2#2.( 890,#18#+"&
"-5*,"#,('-0(?(-)(58"&M6

!"#$%&'"()*+,-./.0+
Exterior

Oligosaccharide

Glycoprotein

"#(

Peripheral protein

Glycolipid

Intergal
protein

Hydrophobic core

Fatty acyl
tails

Hydrophilic
Protein
Integral protein

Hydrophilic polar
head

Peripheral proteins

!"#$%&$1

Phospholipid

Cytosol

Phospholipid bilayer

Leaflets

R%1-&/1-')"-2-1)./)'$-)"-"7,&*-)0$.4%*5)&,,&*5-"-*')./)#,.'-%*0)&*2)'$-%,)0+5&,)(."#.*-*'08

]'&8(18#.18#$"1"0($'9*,("-(58".($')*$$',(.5,2&52,*(".(&'$$*0('(,)!3,)&(JA"%6(K6DL6(78*()'R#,(0,"3"-%
4#,&*(4#,(58*(4#,)'5"#-(#4(18#.18#$"1"0(+"$'9*,.(".(58*(890,#18#+"&("-5*,'&5"#-(+*5:**-(58*(4'559('&9$
&8'"-.(#4(%$9&#$"1"0('-0(18#.18#$"1"0()#$*&2$*.6(^'-(0*,(:''$.("-5*,'&5"#-.(')#-%(58*(890,#&',+#&8'"-.( 4'3#2,( &$#.*( 1'&M"-%( #4( 58*.*( 890,#18#+"&( 5'"$.6( S90,#%*-( +#-0"-%( '-0( *$*&5,#.5'5"&
"-5*,'&5"#-.(+*5:**-(58*(1#$',(8*'0(%,#21.('-0(:'5*,()#$*&2$*.('$.#(.5'+"$"_*(58*(+"$'9*,6(@"&*$$*.(',*
%*-*,'$$9( -#5( 4#,)*0( +9( 18#.18#$"1"0.( "-( 'Q2*#2.( .#$25"#-./( ."-&*( 58*( 4'559( '&9$( &8'"-.( ".18"-%#)9*$"-./( %$9&#$"1"0./( '-0( '$$( 18#.18#$%9&*,"0*.( ',*( 5##( $',%*( 5#( 4"5( "-5#( 58*( "-5*,"#,( #4( '
)"&*$$*(JA"%6(K6CL6
Water

(a)

!"#$%&$2

Water

(b)

!"#$%#&'$%()#$.0#$.1%#%2)".1-(+1-0)%*)'$-)&S+-.+0)#$&0-3)T&U)/.,"&'%.*)./)"%(-11-0)%*)4$%($)#.1&,)$-&20)&,2%,-('-2)'.4&,20)4&'-,V)T7U)/.,"&'%.*)./)7%1%#%2)1&6-,8

"##

1'-#&/()*+,-./.0+

1E6>@I;9A4>J+'9@8E+9:+-E4A;4+'8;<74=8
78*( 3,4%25"0-!%'/ "02),( #4( 58*( 1$'.)'( )*)+,'-*( ".( 58*( 2-"3*,.'$$9( '&&*15*0( )#0*$/( :8"&8( :'.
1,#1#.*0(+9(P"-%*,('-0(U"&#$.#-("-(FKC>6(78*()#0*$(.2%%*.5.(58'5(58*($"1"0.(4#,)('(3".&#2./(5:#;
0")*."#-'$( 4$2"0( )'5,"B( "-5#( :8"&8( 1,#5*"-.( ',*( "-.*,5*0( '-0( "-5*%,'5*0( )#,*( #,( $*..( 0**1$96( 78*
1*,"18*,'$(1,#5*"-.(',*(.21*,4"&"'$$9($#&'5*0('-0()'-9(#4(58*)(42-&5"#-('.(*-_9)*./(:8"$*(58*("-5*%,'$
1,#5*"-.(',*('..#&"'5*0(:"58($"1"0.('-0(1*-*5,'5*("-5#(58*("-5*,"#,(#4(58*()*)+,'-*('$#-%(:"58(4'559
'&"0(."0*(&8'"-.(JK6GL6(78*9(',*(5"%85$9(+#2-0(5#(58*($"1"0.('-0(&#-.5"525*(42-&5"#-'$(1,#5*"-.(58'5(',*
-#5( *'."$9( .*1','+$*6( N$$( )*)+,'-*;+#2-0( *-_9)*.( '-0( &',,"*,.( ',*( "-&$20*0( "-( 58".( &'5*%#,96
V*,"18*,'$( 1,#5*"-.( 8'3*( '( $##.*( '44"-"59( '-0( &'-( +*( *'."$9( 0".1$'&*06( P2&8( '( )*)+,'-*( ".( )#,*
09-')"&('-0(*B1$'"-.(58*("-5,"&'5*(.*)";1*,)*'+$*(18*-#)*-#-6

Peripheral protein

Glycolipid

Lipid layer

!"#$%&$3

Integral protein

W1+%2<".0&%()".2-1)./)9%*5-,8

,7>8=545>9=+9:+-7958>=A+>=+5D8+'8;<74=8
O,"*-5'5"#-( #4( 58*( 1,#5*"-.( ".( ")1#,5'-5( "-( 58*( 42-&5"#-"-%( #4( 58*( )*)+,'-*.6( 78*( U;5*,)"-2.( ".
'$:'9.('5(58*(."0*(58'5(".(52,-*0(':'9(4,#)(58*(&95#.#$6(`.2'$$9(58*(890,#&',+#-(,*."02*.(%*5('55'&8*0
5#(58*(U;5*,)"-2.6(791"&'$()*)+,'-*(1,#5*"-.(&'-(58*,*4#,*(+*(&#2-5*0(')#-%(58*(%$9&#1,#5*"-.6(78*
I;5*,)"-2.(#4(1,#5*"-.(&'-(*"58*,($"*('5(58*(#25."0*/(:"58"-(58*($"1"0($'9*,(#,(1,#5,20*("-5#(58*(&95#.#$/
"6*6(58*(."5*(#4(1,#5*"-(.9-58*.".(JA"%6(K6KL6

!"#$%&'"()*+,-./.0+

"#!

NH3
Extracellular
domain

+ +

O-linked
oligosaccharides

Lipid
bilayer

Central
hydrophobic
alpha helix

+
Cytosolic
hydrophilic
domain

!"#$%&$&

COO

=*'-5,&1)"-"7,&*-)#,.'-%*0)0$.4%*5)'4.)"-"7,&*-<0#&**%*5),-5%.*0C)%8-8)516(.#$.,%*08

(=58F74E+-7958>=A+2>=@+)AC;;857>J4EEC+59+5D8+*>?>@+2>E4C87
]3*,9("-5*%,'$()*)+,'-*(1,#5*"-(8'.('(."-%$*/(.1*&"4"&(#,"*-5'5"#-(:"58(,*.1*&5(5#(58*(&95#.#$"&('-0
*B#1$'.)"&(4'&*.(#4(58*()*)+,'-*6(@#$*&2$*.(#4("-5*%,'$()*)+,'-*(1,#5*"-/(.2&8('.(*,1'0#$0+%(/($"*
"-(58*(.')*(0",*&5"#-/(1,#3"0"-%('.9))*5,9(5#(58*(5:#()*)+,'-*(4'&*.6(<-(&#-5,'.5(5#(18#.18#$"1"0./
1,#5*"-.(8'3*(-*3*,(+**-(#+.*,3*0(5#(4$"1;4$#1('&,#..('()*)+,'-*/(+*&'2.*(.2&8()#3*)*-5(:#2$0(+*
*-*,%*5"&'$$9(2-4'3#2,'+$*('-0(:#2$0(,*Q2",*('(5,'-."*-5()#3*)*-5(#4(890,#18"$"&(')"-#('&"0.('-0
.2%',( ,*."02*.( 58,#2%8( 58*( 890,#18#+"&( "-5*,"#,( #4( 58*( )*)+,'-*6( 78*( '.9))*5,9( #4( )*)+,'-*
1,#5*"-.(".(*.5'+$".8*0(02,"-%(58*",(+"#.9-58*.".('-0()'"-5'"-*0(58,#2%8#25(1,#5*"-.a($"4*5")*6
@*)+,'-*( '.9))*5,9( ".( )#.5( #+3"#2.( "-( 58*( &'.*( #4( )*)+,'-*( %$9&#1,#5*"-.6( <-( 58*( 1$'.)'
)*)+,'-*/( '$$( 58*( O;'-0( U;$"-M*0( #$"%#.'&&8',"0*.( #4( %$9&#1,#5*"-.( '-0( #$"%#.'&&8',"0*.( "%$9&#$"1"0.(',*(#-(58*(*B#1$'.)"&(.2,4'&*6(<-(58*(*-0#1$'.)"&(,*5"&2$2)/(58*9(',*(4#2-0(#-(58*("-5*,"#,
#,($2)*-'$()*)+,'-*(.2,4'&*6

*>?>@+$9;?9A>5>9=+9:+K>::878=5+'8;<74=8+*84:E85A
N$$( 58*( 1$'.)'( )*)+,'-*.( 8'3*( 0"44*,*-5( $"1"0( &#)1#."5"#-( "-( 58*( 5:#( $*'4$*5.6( <-( 58*( 82)'*,958,#&95*/('$$(58*(%$9&#$"1"0.('-0('$)#.5('$$(#4(.18"-%#)9*$"-('-0(18#.18'5"09$(&8#$"-*(',*(4#2-0("58*(*B#1$'.)"&($*'4$*56(<-(&#-5,'.5/(58*(&95#.#$"&($*'4$*5(&#-5'"-.($"1"0.(:"58(-*25,'$(#,(-*%'5"3*(1#$',
8*'0(%,#21./(.2&8('.(18#.18'5"09$*58'-#$')"-*('-0(18#.18'5"09$.*,"-*6(78*(,*$'5"3*('+2-0'-&*(#4('
1',5"&2$',( 18#.18#$"1"0( "-( 58*( 5:#( $*'4$*5.( #4( '( )*)+,'-*( &'-( +*( 0*5*,)"-*0( #-( 58*( +'.".( #4( "5.
.2.&*15"+"$"59(5#(890,#$9.".(+9(18#.18#$"1'.*.(:8*-('00*0(5#(58*(&*$$(*B5*,"#,/(+*&'2.*(18#.18#$"1'.*.
',*(2-'+$*(5#(1*-*5,'5*(58*(&95#.#$"&(4'&*(JA"%6(K6FbL6

'9B8;8=5+9:+'9E8J6E8A+>A+/L9I@>;8=A>9=4E
<-(+#58(-'52,'$()*)+,'-*.('-0(12,*(18#.18#$"1"0(+"$'9*,./(58*,)'$()#5"#-(1*,)"5.(18#.18#$"1"0('-0
%$9&#$"1"0( )#$*&2$*.( 5#( ,#5'5*( 4,**$9( ',#2-0( 58*",( $#-%( 'B".( '-0( 5#( 0"442.*( $'5*,'$$9( :"58"-( 58*
)*)+,'-*($*'4$*56(P"-*(58*.*()#3*)*-5.(',*($'5*,'$(#,(,#5'5"#-'$/(58*(4'559('&9$(&8'"-.(,*)'"-("-(58*
890,#18#+"&("-5*,"#,(#4(58*()*)+,'-*(JA"%6(K6FFL6

"!)

1'-#&/()*+,-./.0+

50
40
30
20
10
0
10
20
30
20
10

% of
total
phospholipid

!"#$%&$'4

Exoplasmic

Phosphatidyl choline
Phosphatidyl ethanol amine
Phosphatidyl serine

Cytoplasmic leaflet

9($-"&'%()0$.4%*5)1%#%2)(."#.0%'%.*)%*)-,6'$,.(6'-)"-"7,&*-8)Q$-),-1&'%:-)&7+*2&*(-)./)#$.0#$.1%#%20)%*
-&($)1-&/1-')&((.+*'0)/.,)'$-%,)0#-(%/%()/+*('%.*08

(a)

!"#$%&$''

Total phospholipids
Sphingomyelin

(b)

F-"7,&*-)1%#%20)&,-)(.*0'&*'16)".7%1-3)T&U),.'&'%.*&1)".:-"-*'V)T7U)1&'-,&1)".:-"-*'8

78*(18#.18#$"1"0.("-(12,*($"1"0(+"$'9*,.(0#(-#5(4$"1;4$#1(#,()"%,'5*(4,#)(#-*($*'4$*5(5#(58*(#58*,6
S#:*3*,/( "-( .#)*( -'52,'$( )*)+,'-*./( #&&'."#-'$( )#3*)*-5( ".( #+.*,3*0( #:"-%( 5#( &'5'$9.".( +9
)*)+,'-*( 1,#5*"-.6( ]-*,%*5"&'$$9/( .2&8( )#3*)*-5.( ',*( *B5,*)*$9( 2-4'3#2,'+$*/( +*&'2.*( 58*( 1#$',
8*'0(#4(18#.18#$"1"0()2.5(+*(5,'-.1#,5*0(58,#2%8(58*(890,#18#+"&("-5*,"#,(#4(58*()*)+,'-*6

)75>:>J>4E+'8;<74=8A+J4=+<8+197;8@+<C+'8JD4=>J4E+K>A?87A>9=+9:
-D9A?D9E>?>@A
7:#(.9.5*).(#4(12,*(18#.18#$"1"0(+"$'9*,(',*(,%#0-0")-('-0(1$'-',(+"$'9*,.6(H"1#.#)*.(',*(.18*,"&'$
3*."&$*.( 215#( $( !( "-( 0"')*5*,( &#-.".5"-%( #4( '( 18#.18#$"1"0( +"$'9*,( 58'5( *-&$#.*.( '( &*-5,'$( 'Q2*#2.
&#)1',5)*-5/(4#,)*0(+9()*&8'-"&'$$9(0".1*,."-%(18#.18#$"1"0.("-(:'5*,6(V$'-',(+"$'9*,.(',*(4#,)*0
'&,#..('(8#$*("-('(1',5"5"#-(58'5(.*1','5*.(5:#('Q2*#2.(.#$25"#-.6(c8*-('(.2.1*-."#-(#4($"1#.#)*.(#,
'(1$'-',(+"$'9*,(&#)1#.*0(#4('(."-%$*(591*(#4(18#.18#$"1"0(".(8*'5*0/("5(2-0*,%#*.('-('+,215(&8'-%*("189."&'$(1,#1*,5"*.(#3*,('(-',,#:(5*)1*,'52,*(,'-%*6(78".(18'.*(5,'-."5"#-(".(02*(5#("-&,*'.*0()#5"#'+#25(58*(I;I(+#-0.(#4(58*(4'559('&9$(&8'"-./(:8"&8(1'..(4,#)('(8"%8$9(#,0*,*0/(%*$;$"M*(.5'5*(5#('
)#,*()#+"$*(4$2"0(.5'5*6(=2,"-%(%*$;5#;4$2"0(5,'-."5"#-/('(,*$'5"3*$9($',%*(')#2-5(#4(8*'5(".('+.#,+*0
#3*,('(-',,#:(5*)1*,'52,*(,'-%*(58'5(".(58*()*$5"-%(5*)1*,'52,*(#4(58*(+"$'9*,6(H"1"0.(:"58(.8#,5(&8'"#,(2-.'52,'5*0(4'559('&9$(&8'"-.(2-0*,%#(18'.*(5,'-."5"#-('5($#:*,(5*)1*,'52,*.(58'-($"1"0.(:"58($#-%
#,(.'52,'5*0(&8'"-.6

!"#$%&'"()*+,-./.0+

!"M

"!"

'&'23)0&+/3)0.-,3/+1H0$/(,0.

78*(1$'.)'()*)+,'-*(42-&5"#-.('.('(+',,"*,(+*5:**-(58*(&*$$('-0("5.(*B5,'&*$$2$',(*-3",#-)*-5(+25
*-.2,*.(5,'-.1#,5(#4(*..*-5"'$()#$*&2$*./(.2&8('.(%$2&#.*/(')"-#('&"0./($"1"0./('-0("#-.(*5&6/("-5#(58*
&*$$('-0('$$#:.(:'.5*.(5#($*'3*(58*(&*$$6

'8;<74=8+>A+.8E8J5>B8EC+-87;84<E8
P*$*&5"3*(1*,)*'+"$"59(#4(58*(1$'.)'()*)+,'-*('$$#:.(58*(&*$$(5#()'"-5'"-('(&#-.5'-5(*-3",#-)*-5("58*("-5*,"#,6(P")"$',$9/(#,%'-*$$*.(:"58"-(58*(&*$$(#45*-(8'3*('(0"44*,*-5("-5*,-'$(*-3",#-)*-5(4,#)(58'5
#4( 58*( .2,,#2-0"-%( &95#.#$/( '( 0"44*,*-5( )'"-5'"-*0( +9( 58*( #,%'-*$$*,( )*)+,'-*.6( A#,( *B')1$*/( 58*
$9.#.#)*.( :"58"-( '-( '-")'$( &*$$( ',*( "-3#$3*0( "-( 58*( 0"%*.5"3*( '-0( .&'3*-%"-%( ,#$*./( :8*,*( 58*
&#-&*-5,'5"#-(#4(1,#5#-.(".(Fbb;Fbbb(5")*.(58'5(#4(58*(&95#.#$6(78".(1,#5#-(%,'0"*-5(".()'"-5'"-*0(+9
1,#5*"-.("-(58*(#,%'-*$$*()*)+,'-*6

-87;84<>E>5C+9:+)75>:>J>4E+'8;<74=8A
N-( ',5"4"&"'$( )*)+,'-*( &#)1#.*0( #4( 12,*( 18#.18#$"1"0( #,( #4( 18#.18#$"1"0( '-0( &8#$*.5*,#$( ".
1*,)*'+$*(5#(%'.*./(.2&8('.(IO>('-0(O>('-0(.)'$$()#$*&2$*./(.2&8('.(*58'-#$6(78*.*()#$*&2$*.(&'&,#..( &*$$( )*)+,'-*.( 2-'"0*0( +9( 5,'-.1#,5( 1,#5*"-.6( N+.#$25*$9( -#( )*5'+#$"&( *-*,%9( ".( *B1*-0*0
+*&'2.*( )#3*)*-5( ".( 4,#)( 8"%8( 5#( $#:( &#-&*-5,'5"#-( #4( 58*( )#$*&2$*./( 0#:-( 58*( &#-&*-5,'5"#%,'0"*-56
<-(&#-5,'.5/('(12,*(18#.18#$"1"0()*)+,'-*(".(#-$9(.$"%85$9(1*,)*'+$*(5#(:'5*,('-0(".(*..*-5"'$$9
")1*,)*'+$*( 5#( )#.5( :'5*,( .#$2+$*( )#$*&2$*./( .2&8( '.( 890,#%*-/( .#0"2)/( &'$&"2)( '-0( 1#5'.."2)6
V,#5*"-.(1$'9(")1#,5'-5(,#$*("-(58*(5,'-.1#,5(#4(.2&8()#$*&2$*.('-0("#-.('&,#..('$$(&*$$2$',()*)+,'-*.
+*&'2.*(0"44*,*-5(&*$$(591*.(,*Q2",*(0"44*,*-5(&#)1#."5"#-(#4(58*.*($#:()#$*&2$',(:*"%85(&#)1#2-0./
."-&*(58*(1$'.)'()*)+,'-*(#4(*'&8(&*$$(591*(&#-5'"-.('(.1*&"4"&(.*5(#4(5,'-.1#,5(1#,5*"-.(58'5(-*%#5"'5*
#-$9( &*,5'"-( "#-.( #,( )#$*&2$*.( 5#( &,#../( '.( 0#*.(58*( )*)+,'-*( .2,,#2-0"-%( *'&8( 591*( #4( #,%'-*$$*
JA"%6(K6F>L6

.9;8+/74=A?975+-7958>=A+16=J5>9=+4A+(9=IJD4==8EA
P#)*()*)+,'-*;$#&'5*0(1,#5*"-.(42-&5"#-('.("#-;&8'--*$.(5#(5,'-.1#,5(:'5*,(#,(.1*&"4"&(591*.(#4("#-.
0#:-(58*(&#-&*-5,'5"#-(%,'0"*-56(78*9(4#,)('(1,#5*"-;$"-*0(1'..'%*:'9('&,#..(58*()*)+,'-*(58,#2%8
:8"&8()2$5"1$*(:'5*,()#$*&2$*.(#,("#-.()#3*(.")2$5'-*#2.$9("-('(."-%$*(4"$*('5('(3*,9(,'1"0(,'5*(JFbGd
.*&#-0L6( A#,( *B')1$*/( 58*( 1$'.)'( )*)+,'-*( #4( '$$( '-")'$( &*$$.( ".( ,"&8( "-( ef( '-0( "5.( )#3*)*-5
0#:-8"$$(58,#2%8('$:'9.;#1*-(&8'--*$.(%*-*,'5*.('-(*$*&5,"&'$(1#5*-5"'$('&,#..(58*()*)+,'-*6(@'-9
#58*,(591*.(#4(&8'--*$(1,#5*"-.(',*(2.2'$$9(&$#.*0/('-0(#1*-(#-$9("-(,*.1#-.*(5#(.1*&"4"&(."%-'$.(JA"%6
K6F?L6
N-#58*,( &$'..( #4( )*)+,'-*( 1,#5*"-./( &'$$*0( &+!(-#0+&)+-/( )#3*( '( :"0*( 3',"*59( #4( "#-.( '-0
)#$*&2$*.('&,#..(58*()*)+,'-*6(78*.*(5,'-.1#,5*,.(+"-0(#-$9(#-*(#,('(4*:(.2+.5,'5*()#$*&2$*.('5('
5")*/(2-0*,%#('(&#-4#,)'5"#-'$(&8'-%*(5#(5,'-.1#,5()#$*&2$*.('&,#..(58*()*)+,'-*6(I#-4#,)'5"#-'$
&8'-%*( #4( 1,#5*"-.( ,*Q2",*.( *-*,%9( 4#,( )#3*)*-5/( 8*-&*( 58*",( )#3*)*-5( ".( .$#:/( '+#25( Fb>YFbX
)#$*&2$*.(1*,(.*&#-06

"!*

1'-#&/()*+,-./.0+

Gases

CO2, N2
O2

Urea
ethanol
H2O
Glucose
+

2+

K , Mg
Ca , HCO3
HPO4
2+

Amino acids
ATP
GlucoseP

!"#$%&$'(

F&?.,)'6#-0)./)',&*0#.,')#,.'-%*08)X*-)'6#-)(.+#1-0)!QB)$62,.160%0)/.,)+#<$%11)".:-"-*')./)%.*0V)'$-).'$-,)'4.
'6#-0C)4$%($)&,-)*.')!QB&0-0C)',&*0#.,')%.*0)2.4*<$%118
Ion
channel

Channel

Closed

Outside

Cystosol
ATP
ATP pump

!"#$%&$')

Open
ADP+Pi
Closed

Q,&*0#.,')#,.'-%*0)(&*)7-)(1&00%/%-2)%*'.):&,%.+0)'6#-0V)($&**-1)#,.'-%*0)',&*0#.,')4&'-,).,)0#-(%/%()%.*0)2.4*
'$-)(.*(-*',&'%.*)5,&2%-*'0).,)-1-(',%(&1)5,&2%-*'0)2.4*<$%118)=.*0).,)4&'-,)".1-(+1-0)".:-)0%*51-)/%1-)&')&
,&#%2),&'-8

78,**(591*.(#4(5,'-.1#,5*,()#$*&2$*.(8'3*(+**-("0*-5"4"*0/(:8"&8("-&$20*(4(%#0+&)+-/(!(&%#0+&)+-/
'-0( -1"#0+&)+-6( `-"1#,5*,.( 5,'-.1#,5( #-*( )#$*&2$*( '5( '( 5")*( 0#:-( 58*( &#&*-5,'5"#-( %,'0"*-5( J*6%6
%$2&#.*('-0(')"-#('&"0.L6(78*('-5"1#,5*,.('-0(.9)1#,5*,.(&'5'$9_*()#3*)*-5(#4(#-*(591*(#4("#-(#,
)#$*&2$*( '%'"-.5( 58*( &#-&*-5,'5"#-( %,'0"*-5/( &#21$*0( 5#( 58*( )#3*)*-5( #4( 0"44*,*-5( "#-( #,( )#$*&2$*
JA"%6(K6FXL6(78*.*(',*(#45*-(,*4*,,*0(5#('.('&5"3*(5,'-.1#,5*,.(+25(:"58#25(890,#$9.".(#4(N7V6

!"#$%&'"()*+,-./.0+

Uniporter

!"#$%&$'*

Symporter

"!$

Antiporter

Q,&*0#.,') #,.'-%*0) &,-) -%'$-,) 06"#.,'-,C) +*%#.,'-,) .,) &*'%#.,'-,) '6#-C) *&"-2) &((.,2%*5) '.) '$-) 2%,-('%.*&1
".:-"-*')./)".1-(+1-08

!"N '&$%)0(.'.+1,3+/3)0.-,3/+,1
')/&3()*.+)$3,..+'&'23)0&.
P*3*,'$(5,'-.1#,5()*&8'-".)(8'3*(+**-(1,#1#.*06

-4AA>B8+K>::6A>9=
V'.."3*(0"442."#-(".()#3*)*-5(#4('()#$*&2$*(4,#)(58*('Q2*#2.(.#$25"#-("-5#(58*(890,#18#+"&("-5*,"#,
#4(58*(18#.18#$"1"0(+"$'9*,6(78*(890,#18#+"&"59(#4('(.2+.5'-&*(".()*'.2,*0(+9(58*(#!+&%&%0(('0)33%'%)(&
6/( 58*( *Q2"$"+,"2)( &#-.5'-5( 4#,( "5.( 1',5"5"#-( +*5:**-( #"$( '-0( :'5*,6( P"-&*( 58*( &#)1#."5"#-( #4( 58*
"-5*,"#,( #4( 58*( 18#.18#$"1"0( +"$'9*,( ,*.*)+$*.( 58'5( #4( #"$/( 58*( 1',5"5"#-( &#*44"&*-5( #4( '( .2+.5'-&*
)#3"-%('&,#..('(+"$'9*,(*Q2'$.(58*(,'5"#(#4("5.(&#-&*-5,'5"#-(R2.5("-."0*(58*(890,#18#+"&(&#,*(#4(58*
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6(Z(I)dI'Q
78*(1',5"5"#-(&#*44"&"*-5(".('()*'.2,*(#4(58*(,*$'5"3*('44"-"59(#4('(.2+.5'-&*(4#,($"1"0(3*,.2.(:'5*,6
O-&*('()#$*&2$*()#3*.("-5#(58*(890,#18#+"&("-5*,"#,(#4('(+"$'9*,/("5(0"442.*.('&,#..("5T(4"-'$$9/(58*
)#$*&2$*()#3*.(4,#)(58*(+"$'9*,("-5#(58*('Q2*#2.()*0"2)(#-(58*(#58*,(."0*(#4(58*()*)+,'-*6(78*
890,#18#+"&(&#,*(#4('(591"&'$(&*$$()*)+,'-*(".(Fbb;Fbbb(5")*.()#,*(3".&#2.(58'-(:'5*,/(8*-&*(58*
0"442."#-(,'5*(#4('$$(.2+.5'-&*.('&,#..(18#.18#$"1"0()*)+,'-*(".()2&8(.$#:*,(58'-(58*(0"442."#-(,'5*
#4(58*(.')*()#$*&2$*("-(:'5*,6(782./()#3*)*-5('&,#..(58*(890,#18#+"&(1#,5"#-(#4('()*)+,'-*(".(58*
,'5*;$")"5"-%(.5*1("-(0"442."#-6

OE6J9A8+/74=A?975+>A+H=>?97587I$454ECP8@
^*,9(4*:()#$*&2$*.(*-5*,(#,($*'3*(&*$$./(#,(&,#..(#,%'-*$$*()*)+,'-*.(:"58#25(58*('"0(#4(1,#5*"-.6
7,'-.1#,5(1,#5*"-.(4,*Q2*-5$9('&&*$*,'5*(5,'-.1#,5(#4()#$*&2$*.(.2&8('.(:'5*,/('-0(2,*'(58'5(&'-(4,**$9
0"442.*(58,#2%8(18#.18#$"1"0(+"$'9*,.6(<5(".(*..*-5"'$(5#(2-0*,.5'-0(58*(1,#1*,5"*.(#4(3',"#2.(M"-0.(#4
)*)+,'-*(1,#5*"-.('-0(58*",(,#$*.("-(#,%'-".)"&(189."#$#%96(78".(8'.(+**-(1#.."+$*(:"58(58*(8*$1(#4
$"1#.#)*.(J',5"4"&"'$()*)+,'-*L6(H"1#.#)*.(8'3"-%('(."-%$*(591*(#4(5,'-.1#,5(1,#5*"-(&'-(+*(2.*0(5#
*B')"-*( 1,#1*,5"*.( #4( 5,'-.1#,5( 1,#5*"-( JA"%6( K6F[L6( N( -#-;"#-"&( 0*5*,%*-5/( .2&8( '.( #&59%$2&#."0*/
.#$2+"$"_*.(58*("-5*%,'$(1,#5*"-.(#4('-(*,958#,#&95*()*)+,'-*6(78*(5,'-.1#,5(1,#5*"-/('(2-"1#,5*,/(&'+*(12,"4"*0('-0(58*-("-&#,1#,'5*0("-5#($"1#.#)*.()'0*(#4(12,*(18#.18#$"1"0.6

"!%

1'-#&/()*+,-./.0+
Glucose transport
protein

Transport protein
with detergent
molecules
Disrupt
membrane

ATP
Glucose
ADP

Phospholipid

Liposome

!"#$%&$'0

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0.1+7%1%Z-0)'$-)%*'-5,&1)#,.'-%*0)./)-,6'$,.(6'-)"-"7,&*-8)!)+*%#.,'-,)'6#-)./)51+(.0-)',&*0#.,')#,.'-%*)(&*)7%*(.,#.,&'-2)%*'.)'$-)1%#.0."-)"&2-)./)#+,-)#$.0#$.1%#%208

`-"1#,5*,.(&'5'$9_*()#3*)*-5(#4(#-*(%$2&#.*()#$*&2$*('5('(5")*(0#:-('(&#-&*-5,'5"#-(%,'0"*-56
P")"$',( 5#( *-_9)*./( 2-"1#,5*,.( '&&*$*,'5*( '( ,*'&5"#-( 58'5( ".( 58*,)#09-')"&'$$9( 4'3#2,*0/( '-0( 58*
)#3*)*-5( #4( '( .2+.5'-&*( '&,#..( 58*( )*)+,'-*( 0#:-( 58*( &#-&*-5,'5"#-( %,'0"*-5( :"$$( 8'3*( '
-*%'5"3*("W6(78".(591*(#4()#3*)*-5(".(,*4*,,*0(5#('.(3!'%,%&!&)2/2%334-%0(6
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F6 78*(,'5*(#4(2-"1#,5(5,'-.1#,5(".(Q2"5*(8"%86
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?6 `-"1#,5(5,'-.1#,5(#&&2,.(3"'('($")"5*0(-2)+*,(#4(5,'-.1#,5*,(1,#5*"-./(,'58*,(58*-(58,#2%8#25
18#.18#$"1"0(+"$'9*,6(I#-.*Q2*-5$9/(58*,*(".('()'B")2)(5,'-.1#,5(,'5*/(:8"&8(".('&8"*3*0(#-$9
:8*-(58*(&#-&*-5,'5"#-(%,'0"*-5('&,#..(58*()*)+,'-*(".(3*,9($',%*6

/L9+'9@8EA+:97+/74=A?975
78*(4",.5()#0*$(".(58*('!++%)+/"02),/("-(:8"&8(58*(5,'-.1#,5*,(1,#5*"-(+"-0.(58*()#$*&2$*.(5#()#3*('5
#-*(4'&*/()#3*.(58,#2%8(58*()*)+,'-*/('-0(,*$*'.*.(58*()#$*&2$*('5(58*(#58*,(4'&*(JA"%6(K6FDL6
78*( .*&#-0( )#0*$( *-3".'%*.( 2.*( #4( 5##( )2&8( *-*,%96( A#,( *B')1$*/( 58*( '-5"+"#5"&( 3'$"-#)9&""-&,*'.*.(58*(5,'-.1#,5(#4(ef("#-.('&,#..(+"#$#%"&'$()*)+,'-*.(+9(4#,)"-%('(.18*,*(',#2-0(*'&8(ef
"#-(JA"%6(K6FCL6(78*(890,#18#+"&(')"-#('&"0(."0*(&8'"-.(#4(58*('-5"+"#5"&($"*(#-(58*(#25*,(.2,4'&*/('-0
."B( #,( *"%85( #B9%*-( '5#).( #-( 58*( "-."0*( &##,0"-'5*$9( +#2-0( 5#( 58*( ef6( 78*( 890,#18#+"&( *B5*,"#,

!"#$%&'"()*+,-./.0+

"!&

Membrane
Exterior

Interior
Unloading
of sugar

Sugar
molecules

Carrier
returned

!"#$%&$'1

E&,,%-,<"-2%&'-2)',&*0#.,')./)0+5&,)".1-(+1-08

O2

Outer surface
hydrophobic
amino acids
Four O2
interacting
+

with K
Valinomycin

!"#$%&$'2

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)'M*.( 58*( ef( &',,"*,( &#)1$*B( .#$2+$*( "-( 58*( $"1"0( "-5*,"#,( #4( 58*( )*)+,'-*.6( <5( ".( +*$*3*0( 58'5
)*)+,'-*(5,'-.1#,5*,.(2-0*,%#(&#-4#,)'5"#-'$(&8'-%*.(58'5(1*,)"5(+#2-0("#-.(#,()#$*&2$*.(5#(1'..
58,#2%8(58*()*)+,'-*6

/74=A?975+9:+OE6J9A8
78*(%$2&#.*(5,'-.1#,5*,('$5*,-'5*.(+*5:**-(5:#(&#-4#,)'5"#-'$(.5'5*.\
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%$2&#.*(4,#)(#25."0*(2-0*,%#*.('(&#-4#,)'5"#-'$(&8'-%*(.#(58'5(58*(#25:',0(4'&"-%(."5*(".("-'&5"3'5*0
'-0(58*(+#2-0(%$2&#.*()#3*.(58,#2%8(58*(1,#5*"-('-0(+*&#)*.('55'&8*0(5#(58*(-*:$9;4#,)*0("-:',0
4'&"-%( ."5*6( 78*( %$2&#.*( ".( ,*$*'.*0( "-5#( 58*( &*$$( "-5*,"#,/( 58*( 5,'-.1#,5*,( 2-0*,%#*.( 58*( ,*3*,.*
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#25:',0;4'&"-%(%$2&#.*(+"-0"-%(."5*(JA"%6(K6FGL6

"!'

1'-#&/()*+,-./.0+

Glucose

Plasma
membrane

!"#$%&$'3

Binding
site

Exterior

Inward
binding site

Cytosol

Glucose

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5,'-.1#,5*,( 1,#5*"-.T( .9)1#,5*,( '-0( &*,5'"-( '-5"1#,5*,( &#;5,'-.1#,5( "#-.( 5#%*58*,( :"58( .1*&"4"&( .)'$$
)#$*&2$*.6(<#-;&8'--*$./("#-(12)1.('-0(.#)*('-5"1#,5*,.(5,'-.1#,5(#-$9("#-./(+25(58*(,'5*('-0(*B5*-5
#4("#-(5,'-.1#,5(".("-4$2*-&*0(+9(58*("#-(&#-&*-5,'5"#-.(#-(58*(5:#(."0*.(#4(58*()*)+,'-*('.(:*$$('.
+9('-(*$*&5,"&(1#5*-5"'$(58'5(*B".5.('&,#..(58*()*)+,'-*6

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78*(&95#.#$"&(1S(".(-*',(-*25,'$(J1S(C6bL('-0(58*(&95#.$#"&(ef("#-(&#-&*-5,'5"#-(".('$:'9.(8"%8*,(58'5
58*(U'f("#-.6(<-(+#58("-3*,5*+,'5*.('-0(3*,5*+,'5*./(ef("#-(&#-&*-5,'5"#-(".(>bYXb(5")*.(8"%8*,("-(58*
&*$$(58'-("-(58*(+$##0/(:8"$*(U'f("#-(&#-&*-5,'5"#-(".('$:'9.($#:*,6(S#:*3*,/(58*(&#-&*-5,'5"#-(#4
4,**(I'ff("#-.("-(58*(&95#.#$(".(%*-*,'$$9($*..(58'-(F()"&,#)#$',/('+#25('(F/bbb(5")*.($#:*,(58'-(58*
+$##06
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)#3*)*-5.( #4( 58*.*( "#-( 58,#2%8( 58*( &8'--*$.( &,*'5*( '( 0"44*,*-&*( "-( *$*&5,"&( 1#5*-5"'$( +*5:**-( 58*
"-."0*( '-0( #25."0*( #4( &*$$6( 78*( )'%-"520*( #4( 58".( 1#5*-5"'$( ".( YCb( )^( :"58( ,*.1*&5( 5#( "-."0*6( 78*
1$'.)'()*)+,'-*(".('-(*$*&5,"&'$(0*3"&*(&'$$*0(&'1'&"5#,6(78*("#-"&(%,'0"*-5.('-0(*$*&5,"&(1#5*-5"'$
',*( ,*.1#-."+$*( 5#( 0,"3*( )'-9( +"#$#%"&'$( 1,#&*..*./( 8*-&*( #1*-"-%( '-0( &$#."-%( #4( "#-( &8'--*$.( ',*
*..*-5"'$(5#(&#-02&5"#-(#4(*$*&5,"&'$(")12$.*.6

QR+$D4==8EA+O8=87458+&E8J57>J+-958=5>4E
78*(0".5,"+25"#-(#4(ef/(U'f('-0(I$ Y("#-.(".('$)#.5(.")"$',("-('-")'$(&*$$('-0("5.(*-3",#-)*-5('-0("4(58*
)*)+,'-*(".(")1*,)*'+$*(5#('$$("#-./(-#("#-.(:"$$(4$#:('&,#..(58*()*)+,'-*('-0(58*,*(:"$$(+*(-#
1#5*-5"'$(0"44*,*-&*6(N.(-#5*0('+#3*/(58*()*)+,'-*(1#5*-5"'$('&,#..(58*(1$'.)'()*)+,'-*(".('+#25
YCb( )^( 02*( 5#( -*%'5"3*( &8',%*( #-( 58*( &95#.#$"&( 4'&*6( 78*( )*)+,'-*( &#-5'"-.( )'-9( #1*-( ef
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58*("-."0*(5#(#25:',0/($*'3"-%('-(*B&*..(#4(-*%'5"3*(&8',%*(#-(58*("-."0*('-0(1#."5"3*(&8',%*(#-(58*
#25."0*6

!"#$%&'"()*+,-./.0+

"!(

78*(U'f(Y(ef(N7V'.*("#-(12)1()#3*.(ef("#-.("-5#(58*(&95#.#$(4,#)(58*(*B5,'&*$$2$',()*0"2)/
%*-*,'5"-%( ef( &#-&*-5,'5"#-( %,'0"*-56( @#3*)*-5( #4( ef( "#-.( 58,#2%8( 1#5'.."2)( &8'--*$.( 4,#)( 58*
&95#.#$( 0#:-( 58*( &#-&*-5,'5"#-( %,'0"*-5( %*-*,'5*.( 58*( "-."0*( -*%'5"3*( )*)+,'-*( 1#5*-5"'$6( 78*
1#5'.."2)(&8'--*$.(8'3*(-#:(+**-(&$#-*0('-0(.*Q2*-&*06

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78*( '&5"3*( 5,'-.1#,5( #4( )#$*&2$*.( '-0( "#-.( ,*Q2",*.( '-( "-125( #4( )*5'+#$"&( *-*,%9/( :8"&8( &'-( +*
0*,"3*0(*"58*,(4,#)(0",*&5(&#21$"-%(5#(58*(890,#$9.".(#4(N7V(#,(+9(&#21$"-%(5#(58*()#3*)*-5(#4('-("#0#:-("5.(&#-&*-5,'5"#-(%,'0"*-56

)J5>B8+(9=+/74=A?975
N7V;1#:*,*0(12)1.(5,'-.1#,5("#-.('%'"-.5(58*",(&#-&*-5,'5"#-(%,'0"*-5.6(A#,("-.5'-&*/(58*(U'f(Y(ef
N7V'.*(12)1.(ef("-5#(58*(&*$$('-0(U'f(#25:',0./(*.5'+$".8"-%('(8"%8(&95#.#$"&(&#-&*-5,'5"#-(#4(ef
58'5(".(*..*-5"'$(4#,(%*-*,'5"#-(#4(-*%'5"3*(1#5*-5"'$('&,#..(58*(1$'.)'()*)+,'-*6(78*(I'ff(N7V'.*
12)1.( I'ff( #25( #4( 58*( &95#.#$( "-5#( 58*( *B5,'&*$$2$',( )*0"2)( #,( "-5#( 58*( "-5,'&*$$2$',( #,%'-*$$*.( "#,0*,(5#()'"-5'"-(&#-&*-5,'5"#-(#4(I'ff("-5#(58*(&95#.#$()2&8($#:*,(58'-("-(58*(*B5,'&*$$2$',()*0"2)6
78*(12)1(0*,"3*.("5.(*-*,%9(4,#)(58*(890,#$9.".(#4(N7V(JA"%6(K6FKL6

ADP + Pi
ATP
Ca

2+
+

Na
2+

Membrane

Ca

K ATPase
pump
Inhibited by
2, 4 diitrophenol

Cell

!"#$%&$'&

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%*'.)'$-)(-118)Q$-)E&N\)])!QB&0-)#+"#0).+')E&N\)./)'$-)(6'.0.18)Q$-)#+"#0)(&*)7-)%*$%7%'-2)76)NC)M<2%*%',.#$-*.18

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"-8"+"5*0(+9(>/(X;0"-"5,#18*-#$6(78*("#-(&#-&*-5,'5"#-("-."0*(58*(&*$$(%,'02'$$9('11,#'&8*0(58'5(#4(58*
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/D788+$E4AA8A+9:+(9=+-6;?A
78,**(&$'..*.(#4(N7V;1#:*,*0(12)1.(8'3*(+**-(,*&#%-"_*0/(&'$$*0(V/(^/('-0(A6(I$'..(V(12)1.(',*
.")1$*.5('-0(&#)1#.*0(#4(4#2,(5,'-.)*)+,'-*(.2+2-"5./(.2&8('.(5:#('$18'('-0(5:#(+*5'(1#$91*15"0*.
JA"%6(K6>bL6(78*($',%*,('$18'(.2+2-"5(".(18#.18#,9$'5*0(02,"-%(5,'-.1#,5(#4("#-.6(U'f(Y(ef(N7V'.*('-0
I'ff(N7V'.*(12)1.(',*("-&$20*0("-(58".(&'5*%#,96

"!#

1'-#&/()*+,-./.0+
Exterior

!
"

Cytosol
ATP-binding
site

P-Class pump

!"#$%&$(4

c c c

!
"
ATP-binding
region
F- and V-class pump

ATP-binding
regions

ABC superfamily

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4$%1-)W)&*2)A<(1&00)#+"#0)&,-)/.+*2)%*)"%'.($.*2,%&)&*2)7&('-,%&1)#1&0"&)"-"7,&*-08

78*(^('-0(A(591*.(#4(12)1./(58#2%8(2-,*$'5*0(5#(&$'..(V(12)1./(5,'-.1#,5(#-$9(1,#5#-.6(A;&$'..
12)1(&#-5'"-(58,**(591*.(#4(5,'-.)*)+,'-*(1,#5*"-.('-0(^;&$'..(12)1.(&#-5'"-('5$*'.5(5:#(M"-0.(#4
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$9.#.#)*.('-0(#58*,(3*."&$*.("-('-")'$(&*$$.(+9(2."-%(*-*,%9(4,#)(N7V(890,#$9.".(5#(12)1(1,#5#-.
4,#)(58*(&95#.#$(5#(58*(*B#1$'.)"&(4'&*(#4(58*()*)+,'-*6(A;&$'..(12)1.(',*(4#2-0("-()"5#&8#-0,"'/
&8$#,#1$'.5.('-0("-(58*(1$'.)'()*)+,'-*(#4(+'&5*,"'6

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5#( '$$#:( %$2&#.*( 5#( )#3*( #25( #4( 58*( &*$$( +9( 4'&"$"5'5*0( 0"442."#-( 58,#2%8( '( 5,'-.1#,5*,6( 78*( $#:
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!"S 2H*Q+/3)0.-,3/+.T./&'.
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:8"&8( #58*,:".*( &'--#5( 0"442.*( 58,#2%8( 58*( )*)+,'-*( +',,"*,6( P2&8( )#$*&2$*.( "-&$20*( 1,#5*"-./

!"#$%&'"()*+,-./.0+

"!!

Epithelial cell
Gut
lumen

High Na

Low Na

High K

Blood

Carrier
Glucose

Transporter
of glucose

ATP
+

Na
Glucose

Na
K

Glucose
ADP + Pi

Na K ATPase

H 2O
H 2O

!"#$%&$('

F-($&*%0")./)51+(.0-)',&*0#.,')%*)%*'-0'%*&1)-#%'$-1%&1)(-1108

1#$9.'&&8',"0*./(1#$91*15"0*.('-0(1#$9-2&$*#5"0*.(*5&6(78".(".(0#-*(+9('(1,#&*..(&'$$*0()(20'1&0-%-6
P")"$',$9(1',5"&2$'5*()'55*,/(.*&,*5#,9()#$*&2$*.($"M*.(8#,)#-*./('-0(1,#5*"-.(&'-(+*(5,'-.1#,5*0(#25
#4(58*(&*$$(+9('(1,#&*..(&'$$*0()70'1&0-%-6(!#58(58*(1,#&*..*.("-3#$3*(58*(4#,)'5"#-(#4(3*."&$*.6

&=@9JC59A>A
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]B5,'&*$$2$',( )'&,#)#$*&2$*.( ',*( *-%2$4*0( '-0( 5,'-.1#,5*0( 5#( 58*( &*$$( "-5*,"#,( :8*,*( "5( ".
1,#%,*.."3*$9(*-&$#.*0("-('(.)'$$(1#,5"#-(#4(58*(1$'.)'()*)+,'-*6(78,**(0".5"-&5(591*.(#4(*-0#&95#5"&
1,#&*..*.( 8'3*( +**-( #+.*,3*0/( :8"&8( "-&$20*( #$!*0'1&0-%-/( #%(0'1&0-%-( '-0( +)')#&0+5")2%!&)2
*-0#&95#.".6

-D4F9JC59A>A
V8'%#&95#.".( ".( '( )'R#,( )*&8'-".)( :8*,*( 1',5"&2$'5*( )'55*,( ".( *-%2$4*0( :"58#25( 4$2"0/( .2&8( '.
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Hyperthermy

Core temperature

Homeothermy

Metabolic heat
production

Metabolic heat production

!"#$%&'$)

Hyper
thermy

t4

t5

t6

Heat death

t3

Lethal temp

Lethal temp

External temperature

Neutral zone

t2

Critical temp

t1

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Blood

Leucocytes

Plasma

Prothrombin
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Corpuscles

Tissues
Erythrocytes
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+ Ca

Platelets
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Stage II

Plasma
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(Enzyme)

Thromboplastin

Prothrombin

++

Ca

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influence
Stage III

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External
carotid

Left common caroti

Carotid body

Carotid sinus
Innominate

Chemoreceptors
aortic arch
Pulmonary
artery
Aortic
sinus

Pulmonary
veins

Aorta
Left auricle

Bicuspid
valve

Semilunar
valves
Right auricle

Tricuspid valve

Left ventricle
Right ventricle

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insects (other than diptera)

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pH 7.4

H2O (87.5%) Obligatory

Urea (40%)

pH
7.4

+
+

NH3

Na

pH 6.0
Distal tubule

REABSORPTION
Creatinine, Urea
H2O, Glucose
+
aa, Na , Uric acid
+
++
++
K , Mg , Ca

P O 4 , Cl , HCO 3
Na+(70%) Obligatory

Filtrate

Ascending limb
Descending
limb

Creatinine

SO4 , Hippuric acid

Proximal rnd

SECRETION

K
+
Na (12%)
H2O (Facultative)
Urea

ADH
Acts here
ADH

H2O ADH Controlled

Urine

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H 2O
300

300
300

300
300

300

300
300

Cortex

300 300
Outer zone of medulla 400
600

300

100

300
400
H2O

300

400

600

600

Nacl
600

Urea

800 800
1000
1200

H 2O

600

Inner zone of medulla

1000

Nacl
800
1000
1200

600

Urea

800
H2O

Na
600

Na

H 2O

1000
1200

800
1200 1200

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)($

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Blood

Proximal tubule cells

CO2
H 2O

Tubular filtrate

CO2
H 2O

CO2
H2CO3 H2O

Carbonic annydrase

!"#$%&'$&(

HCO3
Na

H 2O

HCO3

Na

H2CO3

HCO3

Na

>(;%,%:#'%(&)(*)092.(+"&)%(&/)%&)'0")1.(G%$#,)'-;-,"7

!(,*")#$((#-./$%-*1$)&#.*1"#$%&#%&$-"*%-&4#.1#)7&#3%*9.,$(#)'-'(&#$14#)7&#%&,$.1.1;#$%&#$-"*%-&4
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4.")$(#)'-'(&5
(pH 7.4)
Blood
CO2
H 2O

Distal tubule cells

Tubular filtrate

CO2

Formed at glomerulus
+

(pH 7.4)
+

H
HCO3
Na

2 Na + HPO4

H2CO3
H 2O
Carbonic annydrase

HCO3
+

Na

Na +

Na + H + HPO4 pH 6.0
To urine

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./0123#,"

Blood

Distal tubule cells

CO2
H 2O

Deamination of
amino acids
CO2
H 2O

)('

Tubular filtrate
+

NH3

Carbonic
anhydrase

H2CO3
H
HCO3

NH3

NH4

HCO3
Na

!"#$%&'$&'

Na

Na

Cl

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!"#$%&' ()*+#,&,-*

Millivolts

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Rheobase

Msec
Chronaxie

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Millivolts

0
Spike potential
20

40
EPSP

Threshold potential

60
Resting potential

6
Milliseconds

!"#$%&'$1

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10

12

./01/' ()*+#,&,-*

#!)

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Action
Potential

+ 40
+ 20
0
20
40
60
80

Resting potential

1
Axon

Na
K

+
+

Membrane potential
(millivolts)

Na
+ 40
+ 20
0
20
40
60
80

action

Na

2
K

+
+

Membrane potential
(millivolts)

Na
+ 40
+ 20
0
20
40
60
80

Minimum refractory period

Na

3
K

Na
K

Na

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56 mV

Internal
potential

20

Sodium
conductance
10

20

Potassium
conductance

10

0
0

10

Time (in sec)

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./01/' ()*+#,&,-*

#$!

Ena

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30

115 mV
20

10
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12 mV

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0

2
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(b)

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Increase in Na
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##"

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Cilia

Dendrite

Receptor cell
Sustentaclar
cell
Endoplasmic
reticulum

Basement
membrane
Axons

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Pore

Epithelium

Receptor cell

Taste bud

Nerve fibres

Basement
membrane

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!"#$%&'()"*+,#-$.$
Alpha motoneuron

##'

Extrafusal
fibre

Spindle
receptor
neuron

Intrafusal
fibres

Gamma
motoneuron

Capsule of
muscle spindle
Tendon organ
receptor neuron

Tendon
organ

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Posterior
semicircular canal

Edolymphatic duct

Lateral
semicircular

Utricle
Saccule

Cochlea

Ampulla
Fenestra ovalis
Cochlear duct

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Semicircular
canal

Utricle
Vetibulocochlear
nerve

Ossicles in
the middle ear

Vestibular
branch
Cochlear
branch
Saccule

Cochlear

External
auditory meatus

Eustachian tube

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Temporal
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Perilymph
Vestbular
membrane

Scala vestbuli

Endolymph
Scala
media

Cochlear
nerve

Tectorial
membrane
Basilar
membrane

Spiral
ganglion
cells

Organ of
corti

Scala tympani

Perilymph

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Posterior chamber
Anterior chamber

Retina
Cornea
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Choroid

Lens
Iris

Optic tract

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Outer plexiform
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layer

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Optic nerve
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Ciliary filaments

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Corneal
lens
Pigment
cells

Iris pigment
cells
Retinal
pigment cells

Crystalline
cone
Retinal
pigment cells
Rhabdom
Retinal cells

Basement membrane
(a)

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(b)

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Sensory
input
(receptor)

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Sensory
Pathway

Modular
(brain or
spinal cord)

Motor
Pathway

Output
(effector)

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Dorsal root

Dorsal horn

Skin
Spinal ganglion

Afferent
nerve

Internuncial
neuron

h i
g

Spinal nerve
Afferent nerve

Ventral
root

Ventral
horn

Commissural
neuron

Muscle

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#%*

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PARASYMPATHETIC

Constricts pupil

SYMPATHETIC

Ganglion

Vagus nerve
Stimulates flow of saliva

Slows heart beat

Constricts bronchi

Stimulates peristalsis
and secretion

Stimulates synthesis of glycogen

Contracts bladder

Medulla oblongala

Dilates papil

Inhibits flow of saliva

Accelerates heat beat

Dilates bronchi
Solar plexus

Inhibits peristelsis
and secretion
Secretion of
addrenalime and neradrenaline
stimulates conversion of glycogen
to glucose

Inhibits bladder contraction

Chain of sympathetic ganglia

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Sheath cell

Postsynaptic Membrane
Nucleus

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mv
Action potential

+45

45

90

gm

Contraction

0
Latent period

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Relaxation

10

15

20

Milliseconds

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Shortening speed
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3.0

2.0

1.0

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10

20
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30

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.//012,3'43-%"+

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.//012,3'43-%"+

&*!

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Neurosecretory cells in
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Supra optic
hypophysial tract

Third ventricle

Optic chiasma
Hypophysial portal veins
Posterior lobe

Arterial
supply

Hypophysial
artery
Primary plexus
in median eminence

Venous
drainage
Anterior lobe

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1.

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CH2

CH

#!%

COOH

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Thyroxine (tetraiodothyronine)
2.
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CH2

CH

COOH

NH2
3, 5, 3!-triiodothyronine
3.
HO

CH2

CH

COOH

NH2
3, 3, 5!-triiodothyronine
4.
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CH2

CH

COOH

NH2
3, 3!-diiodothyronine

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Serum

Mono-iodotyrosine (MIT)

Di-iodotyrosine (DIT)

Tri-iodothyronine (T3)

Tetra-iodothyroxine (T4)

Thyroglobulin
(Storage form)

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!"#$"%&'()*+,'&-."%

#)!

Cholesterol
NADPH
Pregnenolone

Dehydroepiandrosterone

NADPH

NADPH
Progesterone
NADPH

NADP

11 desoxycorticosterone
NADPH
NADPH
Aldosterone

17 OH progesterone

Corticosterone 17 OH desoxycorticosterone
NADPH
("B")
("S")
Compound
Cortisol
("F")
Adrenesterone

*+,)&'()'4

Z-$25(&0.2-2)$@),$%&-,-'7)0$%"$(.2)@%$"),0$7.2&.%$7>

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/%.$&'(0123."'"+2
CH2OH
HO

CH2OH

C=O
OH

C=O
OH

Cortisol

Cortisone

CH2OH

C=O
HO

C=O

O
Deoxycorticosterone

*+,)&'()''

O=C

CH2OH

Aldosterone

?$%&-,$2&.%$-+2>

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OH

CH3
N
H
HO

Epinephrine
OH
OH

NH2

HO
OH

*+,)&'()'.

Norepinephrine

T.+477'%5)0$%"$(.2>

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#**

Neurosecretory cells
produce brain hormone

*+,)&'()'0

Corpora allata

Prothoracic gland
stimulated

Juvenile hormone
(JH)

Ecdysone released

Chromosome
puffs

Chromosome
puffs

Puffs in giant
chromosomes

RNA synthesis
accelerated

RNA synthesis
accelerated

RNA synthesis
accelerated

Protein
synthesis

Protein synthesis

Protein synthesis
accelerated

Larva

Pupa

Adult moth

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./01,2345#,"

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Membrane
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Liquor
folliculi

Zona pellucida
Ovum
Theca
interna

Corona
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Theca
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CH3

CH3

Testosterone

O
CH3

ANDROGENS
CH3

Androstenedione
447
O

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CH3
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CH3

CH3

PROGESTERONE

Estrone

Estradiol
HO

HO
ESTROGENS

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+
30 S

3!

5 !A U G
mRNA

+ IF3

GTP
IFI
IF2

A
AUG

5!

f-met
5!

50S

3!

IF3

AUG

G D P + Pi 5O S
IF I, IF 2 Sub unit

+ G T P, IFI
IF 2, IF3

IV

III

!"#$%&'$0

II

fmet tRNA

f-met

AUG
+ IF3

I#"</$'+&'#2/'+&+#+"#+%&'%1'@*%#/+&'$=&#2/$+$9

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50 S ribosome
Site of peptidyl transferase
Peptidyl site (p)
fmet tRNA
5 'end

arg
A

f met
P

Amino acyl tRNA

mRNA

U A C G AU
AUG CGA

3 'end
AUGAUC GAA UCU CUG
Codons

30 S ribosome
GTP
f me

Paptidyl transferase
reaction

t ar
g

U A C G AU
AUG CGA

5 'end

3 'end
AUCGUC GAA UCU CUG

Direction of ribosome
movement
Peptidyl site
Amino acyl site(A)

Translocation
reaction
GDP + Pi

Out going
empty
tRNA
U C
A
5 'end

f me

met

arg

U C
A

AC U
AUG CGA AUG

Incoming
amino acyl tRNA

3 'end
AUCGAA UCU CUG

ile
f met
5 'end
AUG

arg
P

G
UA

met
A

G A U U AC
CGA AUG

3 'end
AUCGAA UCU CUG

e
id
pt
pe
ly
po rg
g
a
in
w t
ro
G f me
ch
n
arg

ile

ai

me

5 'end
AUG

!"#$%&'$1

CGA AUG

AUC

glu
P

ser
A

C U U A GA
GAA UCU CUG

:2/'$/L3/&,/'%1'$#/@$'+&'#2/'<*%;#2'%1'"'@%0=@/@#+(/',2"+&'%&'"'*+4%$%./9

3 'end

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Galactosidase activity

Nitrogen source
absent

Nitrogen source
present

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4
Hours

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#&!

0.%1*'+!"#$%&'&(#

R
Repressor

R
Repressor

Repressor

!"#$%&'$)

Operator
gene
Switch
close

Operation
Structural genes
O

Repression

Product
R

O
A
B
C
Switch
open
Induction
a
b
c
Product
R
+
!-galactoside
O
A
B
C
Switch
closes
!-galactosides
broken down
Product
R

mRNA
Proteins
(enzyme)

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Protein

CH

CH

CH
Melanin

CHNH2
COOH
Phenylalanine

Phenylpyruvic
oligohrenia

Gene pp
CH2

CH2

CH2

CHNH2

CHNH

COOH
Tyrosine

COOH
Jopa

Albinism
Gene CC

OH

CH2

CH2

CH2

C=O

C=O

C=O

COOH
COOH
Phenylpyruvic acid Phenylpyruvic acid
(in foods)
(excreted by
phenylpyruvics)

COOH
Phydroxyphenylpyruvic acid
2,5 dihydroxyphenylpyruvic acid
OH

CH2 COOH
OH
2,5 dihydroxyphenylacetic acid
homogentisic acid
(excreted by alcaptonurics)
alcaptonuria
O
geneaa
CH3 C CH COOH
Acetoacetic acid
CO2 + H2O

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Phenylalaline

Phenylpyruvic acid

Tyrosine

Dihydroxyphenylalanine
(dopa)

Diidotyrosine

Dopachrome

(2 mols)

Indole quinone

Thyroxine

Melanine
molecule

Alanine

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@" H(/+'()'*$+'52(L+
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@" 933'()'52(L+

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@" M$+')(6,5*%(/'()'&%*6.33%/+')6(,'(6/%*$%/+
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@" M$+6+'%1'*((',.&$'5&*%L%*B'()'$(,(7+/*%1%&'5&%0'(D%051+
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@" H(/+'()'*$+'52(L+
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@" H(/+'()'*$+'52(L+
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@" 933'*$+'52(L+
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@" H(/+'()'*$+'52(L+
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9" Y+3-1'%/'+3%,%/5*%/7'$B06(7+/'%(/1'%/'*$+')(6,'()'HYNI'%(/1
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>" Q5&$'5,%/('5&%0'%1'500+0'%/'%*1'-6(-+6'-35&+'*('5'76(4%/7'-+-*%0+'&$5%/'*$6(.7$'*$+'505-*(6
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@" 9'7%L+/'&(0(/'5/0'%*1'5/*%&(0(/',.1*'$5L+'%0+/*%&53'251+'1+\.+/&+1'%/'(60+6')(6'-6(-+6
251+'-5%6%/7
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9" G.1&3+'73B&(7+/'&5//(*'2+'&(/L+6*+0'*('73.&(1+`T`-$(1-$5*+
;" G.1&3+'35&S1'73.&(1+`T`-$(1-$5*51+'5&*%L%*B
>" G.1&3+&(/*5%/1'/('73.&(S%/51+
@" G.1&3+'35&S1'73.&(%1(,+651+'5&*%L%*B
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9" M6B-1%/'%1'5/'+D(-+-*%051+'*$5*'3%2+65*+1')6++'5,%/('5&%01
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@" M6B-1%/'&5/'5&*'51'5/'5&*%L5*(6')(6'533'=B,(7+/1'()'-5/&6+5*%&'-6(*+51+1

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@" E/L(3L+1'5'/.,2+6'()',%/(6'251+1'511(&%5*+0'4%*$'&$5%/'%/%*%5*%(/
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9" ;'3B,-$(&B*+1'*$5*'$5L+'2++/'*65/1)(6,+0'%/*('-351,5'&+331
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@" H(/+'()'*$+'52(L+
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@" E1'/(*')(./0'%/'-5*%+/*1'1.))+6%/7')6(,'S+*(1%1
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@" H(/+'()'*$+'52(L+
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@" 9/'%/&6+51+0'6+3+51+'()'%/1.3%/
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@" H(/+'()'*$+'52(L+
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@" ?+6%/+'0+$B065*51+
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@" M6B-*(-$5/
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@" 9'6+1+6L+'1(.6&+'()'+/+67B'*('6+-3+/%1$'9M<'%/'*$+',.1&3+
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@" U3.*5,%/51+
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9" U3B&(7+/'1*(6+1'%/'3%L+6
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@" E/50+\.5*+'-$+/B3535/%/+'%/'*$+'0%+*
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9" M65/1S+*(351+
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@" U3.&(1+`T`-$(1-$5*+'0+$B06(7+/51+
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9" C5%3.6+'*('75%/'4+%7$*
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@" d+0.&%/7'1.756'%/'.6%/+')(33(4%/7',%3S
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9" E/&(66+&*'1+&(/056B'1*6.&*.6+
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9" ;(./0'*('Y2'51'&5625,%/('76(.-1
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@" 91'>:A'751'%/'*$+'-351,5
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9" E*'%1'5&*%L5*+0'2B'2%3+'153*1'1('*$5*'%*'&5/'&3+5L+'*6B-1%/(7+/'-6(0.&%/7'5&*%L+'*6B-1%/'4$%&$
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@" E*'%1'/(*'5'=B,(7+/'5/0'*$+6+)(6+'*$+'(/3B'-6(*+51+'*$5*'%1')(./0'%/'5/'5&*%L+')(6,'5*'*$+
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