Ski
pt
oMai
ncont
ent
El
sev
i
erl
ogo
Enzy
mes
Enzy
mesar
easpeci
al
i
zedcl
assofpr
ot
ei
nsr
esponsi
bl
ef
orcat
al
y
zi
ngchemi
cal
r
eact
i
ons
wi
t
hi
nt
hecel
l
andt
husar
ei
deal
dr
ugt
ar
get
s.
Fr
om:
Pedi
at
r
i
cCr
i
t
i
cal
Car
e(
Four
t
hEdi
t
i
on)
,
2011
Rel
at
edt
er
ms:
Cy
t
ochr
omeP450Gl
ucoseAmi
noAci
dsEi
cosanoi
dRecept
or
Adenosi
neTr
i
phosphat
eI
n
Vi
t
r
oPept
i
dePr
ot
ei
nDNAPr
ot
ei
nase
Vi
ewal
l
Topi
cs
Enzy
mes
Ant
oni
oBl
anco,
Gust
av
oBl
anco,
i
nMedi
cal
Bi
ochemi
st
r
y
,
2017
Summar
y
Enzy
mesar
ecat
al
y
st
st
hat
,
wi
t
hi
nt
hemi
l
dcondi
t
i
onsoft
emper
at
ur
e,
pH,
andpr
essur
eoft
he
cel
l
s,
car
r
youtchemi
cal
r
eact
i
onsatamazi
nghi
ghr
at
e.Theyar
echar
act
er
i
zedbyar
emar
kabl
e
ef
f
i
ci
encyandspeci
f
i
ci
t
y
.
Subst
r
at
esar
et
hesubst
ancesonwhi
chenzy
mesact
.
Enzy
mesar
enamedbyaddi
ngt
hesuf
f
i
xaset
ot
henameoft
hesubst
r
at
et
hatt
heymodi
f
y(
i
.
e.
,
ur
easeandt
y
r
osi
nase)
,
ort
het
y
peofr
eact
i
ont
heycat
al
y
ze(
dehy
dr
ogenase,
decar
boxy
l
ase)
.
Somehav
ear
bi
t
r
ar
ynames(
pepsi
nandt
r
y
psi
n)
.TheI
nt
er
nat
i
onal
Uni
onofBi
ochemi
st
r
yand
Mol
ecul
arBi
ol
ogyassi
gnseachenzy
meanameandanumbert
oi
dent
i
f
yt
hem.
Enzy
mesar
ecl
assi
f
i
edi
nt
osi
xcat
egor
i
esaccor
di
ngt
ot
het
y
peofr
eact
i
oncat
al
y
zed:
Oxi
dor
educt
ases,
t
r
ansf
er
ases,
hy
dr
ol
ases,
l
y
ases,
l
i
gases,
andi
somer
ases.
St
r
uct
ur
al
l
y
,
t
hev
astmaj
or
i
t
yofenzy
mesar
epr
ot
ei
ns.Al
soRNAmol
ecul
eshav
ecat
al
y
t
i
c
act
i
v
i
t
y(
r
i
bozy
mes)
.
Coenzy
mesar
esmal
l
nonpr
ot
ei
nmol
ecul
est
hatar
eassoci
at
edt
osomeenzy
mes.Many
coenzy
mesar
er
el
at
edt
ov
i
t
ami
ns.Coenzy
mesandt
hepr
ot
ei
npor
t
i
onwi
t
hcat
al
y
t
i
cact
i
v
i
t
yor
apoenzy
mef
or
mt
hehol
oenzy
me.Theapoenzy
mei
sr
esponsi
bl
ef
ort
heenzy
me’
ssubst
r
at
e
speci
f
i
ci
t
y
.Coenzy
mesunder
gochangest
ocompensat
ef
ort
het
r
ansf
or
mat
i
onsoccur
r
i
ngi
n
t
hesubst
r
at
e.
Met
al
l
oenzy
mesar
eenzy
mest
hatcont
ai
nmet
al
i
ons.
Themechani
sm ofact
i
onofenzy
mesdependsont
heabi
l
i
t
yofenzy
mest
oaccel
er
at
et
he
r
eact
i
onr
at
ebydecr
easi
ngt
heact
i
v
at
i
onener
gy
.Dur
i
ngt
hecour
seoft
her
eact
i
on,
t
heenzy
me
(
E)bi
ndst
ot
hesubst
r
at
e/
s(
S)andf
or
msat
r
ansi
entenzy
me–subst
r
at
ecompl
ex(
ES)
.Att
he
endoft
her
eact
i
on,
t
hepr
oduct
/
sar
ef
or
med,
t
heenzy
mer
emai
nsunchanged,
canbi
ndanot
her
subst
r
at
eandcanber
eusedmanyt
i
mes.
Act
i
v
esi
t
eorcat
al
y
t
i
csi
t
ei
st
hespeci
f
i
cpl
acei
nt
heenzy
mewher
et
hesubst
r
at
ebi
nds.The
st
r
uct
ur
al
compl
ement
ar
i
t
ybet
weenEandSal
l
owsanexactr
eci
pr
ocal
f
i
t
.Theenzy
meadapt
s
t
ot
hesubst
r
at
ev
i
aaconf
or
mat
i
onal
changeknownasi
nducedf
i
t
.Thepr
esencei
nt
heact
i
v
e
si
t
eofami
noaci
dst
hatbi
ndf
unct
i
onal
gr
oupsi
nt
hesubst
r
at
eensur
esadequat
el
ocat
i
onof
t
hesubst
r
at
eandf
or
mat
i
onoft
het
r
ansi
t
i
oni
nt
er
medi
ar
y
,
whi
chwi
l
l
besubj
ect
edt
ocat
al
y
si
s.
Zy
mogensorpr
oenzy
mesar
ei
nact
i
v
epr
ecur
sor
sofenzy
mes.Theyacqui
r
eact
i
v
i
t
yaf
t
er
hy
dr
ol
y
si
sofapor
t
i
onoft
hei
rmol
ecul
e.
Cel
l
ul
arl
ocat
i
onofenzy
mesv
ar
i
es,
t
hemaj
or
i
t
ybei
ngi
ndi
f
f
er
entcompar
t
ment
soft
hecel
l
,
whi
l
eot
her
sar
eext
r
acel
l
ul
ar
.
Mul
t
i
enzy
mesy
st
emsar
et
hosecomposedofaser
i
esofenzy
mesorenzy
mecompl
exes.Ther
e
ar
eal
somul
t
i
f
unct
i
onal
enzy
meswi
t
hsev
er
al
di
f
f
er
entcat
al
y
t
i
csi
t
esi
nt
hesamemol
ecul
e.
Enzy
meact
i
v
i
t
yi
sdet
er
mi
nedbymeasur
i
ngt
heamountofpr
oductf
or
med,
orsubst
r
at
e
consumedi
nar
eact
i
oni
nagi
v
ent
i
me.I
ni
t
i
al
v
el
oci
t
ycor
r
espondst
ot
heact
i
v
i
t
ymeasur
ed
whent
heamountofconsumedsubst
r
at
ei
sl
esst
han20%oft
het
ot
al
subst
r
at
eor
i
gi
nal
l
y
pr
esent
.OneI
Uofenzy
mecat
al
y
zest
heconv
er
si
onof1μmol
ofsubst
r
at
epersecondunder
def
i
nedcondi
t
i
onsofpHandt
emper
at
ur
e.Speci
f
i
cact
i
v
i
t
yi
st
heuni
t
sofenzy
mepermi
l
l
i
gr
am
ofpr
ot
ei
npr
esenti
nt
hesampl
e.Mol
aract
i
v
i
t
yort
ur
nov
ernumberar
et
hesubst
r
at
emol
ecul
es
conv
er
t
edi
nt
opr
oductperuni
tt
i
meperenzy
memol
ecul
e,
undercondi
t
i
onsofsubst
r
at
e
sat
ur
at
i
on.
Ther
at
eoft
heenzy
mat
i
cr
eact
i
oni
sdi
r
ect
l
ypr
opor
t
i
onal
t
ot
heamountofenzy
mer
at
epr
esent
i
nt
hesampl
e.
Al
so,
atl
ow[
S]andunderconst
antcondi
t
i
onsoft
hemedi
um,
enzy
meact
i
v
i
t
yr
api
dl
yi
ncr
eases
wi
t
ht
her
ai
sei
n[
S]
.Athi
ghersubst
r
at
el
ev
el
s,
t
heact
i
v
i
t
yi
ncr
easessl
owl
yandt
endst
or
eacha
maxi
mum.Theef
f
ectf
ol
l
owsahy
per
bol
i
cf
unct
i
on;
atl
ow[
S]t
her
eact
i
oni
sf
i
r
stor
der
;
athi
gh
[
S]t
her
eact
i
oni
szer
oor
derwi
t
hr
espectt
ot
hesubst
r
at
e.
Km orMi
chael
i
sconst
anti
st
he[
S]atwhi
cht
her
eact
i
onr
at
er
eachesav
al
ueequal
t
ohal
ft
he
maxi
mum.
Undergi
v
encondi
t
i
onsofpHandt
emper
at
ur
e,
t
heKm v
al
uei
sdi
st
i
nct
i
v
ef
oreachenzy
meand
i
susedt
ochar
act
er
i
zei
t
.Formostenzy
mes,
t
heKm v
al
uei
si
nv
er
sel
yr
el
at
edt
ot
heaf
f
i
ni
t
yof
t
heenzy
mef
ort
hesubst
r
at
e,
t
hehi
ghert
heaf
f
i
ni
t
y
,
t
hel
owert
heKm.
Temper
at
ur
eaf
f
ect
senzy
meact
i
v
i
t
y
,
i
ncr
easi
ngi
tt
or
eachapeak,
whi
chcor
r
espondst
ot
he
opt
i
mal
enzy
meact
i
v
i
t
y
.Bey
ondt
hi
smaxi
mum,
enzy
meact
i
v
i
t
yr
api
dl
ydr
ops.Theopt
i
mal
t
emper
at
ur
ef
ormostmammal
i
anenzy
mesi
sar
ound37°
C.Thei
nact
i
v
at
i
ngef
f
ectof
t
emper
at
ur
esabov
e40°
Ci
sduet
opr
ot
ei
ndenat
ur
at
i
on.
pHaf
f
ect
senzy
meact
i
v
i
t
y
,
byi
nf
l
uenci
ngt
hest
at
eofdi
ssoci
at
i
onoff
unct
i
onal
gr
oupsi
nv
ol
v
ed
i
nt
heEScompl
ex.Enzy
meshav
eanopt
i
mum pHandext
r
emev
al
uesofpHcauseenzy
me
denat
ur
at
i
on.
Enzy
mei
nhi
bi
t
or
scanbecl
assi
f
i
edas:
I
r
r
ev
er
si
bl
e,
whi
chper
manent
l
yi
nact
i
v
at
et
heenzy
me,
and
Rev
er
si
bl
e,
whi
chconsi
stoft
hef
ol
l
owi
ngi
nhi
bi
t
or
s:
Compet
i
t
i
v
e:
i
ncr
easet
heKm butnott
heVmax,
i
t
sact
i
oni
sr
ev
er
sedbyi
ncr
easi
ng[
S]
.Some
hav
est
r
uct
ur
al
si
mi
l
ar
i
t
yt
ot
hesubst
r
at
eandcompet
ewi
t
hi
tf
ort
heact
i
v
esi
t
e.
Noncompet
i
t
i
v
e:
bi
ndt
ot
heenzy
mei
nasi
t
edi
f
f
er
entt
ot
hecat
al
y
t
i
ccent
er
.Theydecr
ease
Vmax,
l
eav
eKm unaf
f
ect
ed,
andar
enoti
nf
l
uencedby[
S]
.
Ant
i
compet
i
t
i
v
e:
r
educeKm andVmax.
Enzy
mesar
esubj
ect
edt
or
egul
at
i
on,
t
oadaptt
ot
her
equi
r
ement
sofdi
f
f
er
entcel
l
s.Whent
he[
S]
i
nt
hecel
l
i
sbel
owt
heKm,
changesi
n[
S]modi
f
yt
heact
i
v
i
t
y
.Al
l
ost
er
i
cenzy
mesar
et
hose
modul
at
edbyagent
st
hatbi
ndt
ot
hem atasi
t
edi
f
f
er
entt
ot
heact
i
v
ecent
er
.Thecur
v
eofi
ni
t
i
al
v
el
oci
t
yv
er
sus[
S]f
oral
l
ost
er
i
cenzy
mesi
snothy
per
bol
i
c,
butsi
gmoi
d.Enzy
meact
i
v
i
t
yi
sal
so
changedbycov
al
entmodi
f
i
cat
i
on,
suchasphosphor
y
l
at
i
on.
Const
i
t
ut
i
v
eenzy
mesar
et
hosewhosel
ev
el
sr
emai
nconst
antt
hr
oughoutt
hel
i
f
eoft
hecel
l
.
I
nduci
bl
eenzy
mes,
ar
et
hosewhosesy
nt
hesi
si
sact
i
v
at
edasr
equi
r
ed.
I
sozy
mesar
edi
f
f
er
entpr
ot
ei
nst
hathav
et
hesameenzy
meact
i
v
i
t
y
.
Vi
ewchapt
er
Pur
chasebook
Enzy
mes
Ger
al
dLi
t
wackPh.
D.
,
i
nHumanBi
ochemi
st
r
y
,
2018
Coenzy
mes
Someenzy
mesar
eact
i
v
ewi
t
houtcoenzy
mes.Howev
ermany
,
r
equi
r
eacoenzy
met
obeact
i
v
e.
Anenzy
met
hati
si
nact
i
v
ei
nt
heabsenceofi
t
scoenzy
mei
scal
l
edanapoenzy
me.I
nt
he
pr
esenceofi
t
scoenzy
met
opr
oducet
heact
i
v
ef
or
m oft
heenzy
me,
i
ti
scal
l
edahol
oenzy
me:
apoenzy
me+coenzy
me⇌hol
oenzy
me
Al
t
houghsomeenzy
mescont
ai
nacoenzy
met
hati
st
i
ght
l
ybound,
ot
her
smaycont
ai
na
coenzy
met
hati
sr
eadi
l
ydi
ssoci
abl
e.I
nt
hel
at
t
ercase,
t
hecoenzy
mecanbeconsi
der
edasa
r
eact
antorsubst
r
at
e.Thus,
i
nt
hel
act
at
edehy
dr
ogenaser
eact
i
on,
f
orexampl
e,
py
r
uv
at
eand
t
hecoenzy
meNADHneedt
obeaddedt
ot
heenzy
meand,
ki
net
i
cal
l
y
,
t
hi
swoul
dbeconsi
der
ed
t
obeat
wosubst
r
at
er
eact
i
on:
py
r
uv
at
e+NADH⇌l
act
at
e+NAD+
I
nadoubl
er
eci
pr
ocal
pl
oti
nwhi
ch1/
v
el
oci
t
y(
y
axi
s)i
spl
ot
t
edagai
nst1/
[
py
r
uv
at
e]hol
di
ngt
he
concent
r
at
i
onofNADHhi
gh,
t
hepl
otwoul
dgi
v
et
heKm f
orpy
r
uv
at
e.Asi
mi
l
arexper
i
menti
n
whi
ch[
py
r
uv
at
e]wasatasat
ur
at
i
ngl
ev
el
and[
NADH]wasv
ar
i
ed,
t
her
eci
pr
ocal
pl
otwoul
dgi
v
e
t
heKm f
orNADH.
I
ngener
al
,
coenzy
mesar
ev
i
t
ami
nsorder
i
v
at
i
v
esofv
i
t
ami
ns.Theyar
el
i
st
edi
nTabl
e5.
4.
Tabl
e5.
4.TheVi
t
ami
ns,
Thei
rCoenzy
mes,
andThei
rChemi
cal
Funct
i
ons
Vi
t
ami
n
Coenzy
me
React
i
onCat
al
y
zed HumanDef
i
ci
encyDi
sease
Wat
er
Sol
ubl
eVi
t
ami
ns
Ni
aci
n(
ni
aci
nat
e)
NAD+,
NADP'Oxi
dat
i
on
Pel
l
agr
a
NADH,
NADPHReduct
i
on
Ri
bof
l
av
i
n(
v
i
t
ami
nB2)
FADH2,
FMNH2
FAD,
FMN
Oxi
dat
i
on
Ski
ni
nf
l
ammat
i
on
Reduct
i
on
Thi
ami
ne(
v
i
t
ami
nB1)
Thi
ami
nepy
r
ophosphat
e(
TPP)
Li
poi
caci
d(
l
i
poat
e) Li
poat
e
Oxi
dat
i
on
Twocar
bont
r
ansf
er Ber
i
ber
i
–
Di
hy
dr
ol
i
poat
eReduct
i
on
Pant
ot
heni
caci
d(
pant
ot
henat
e)
Bi
ot
i
n(
v
i
t
ami
nH)
Bi
ot
i
n Car
box
y
l
at
i
on–
Py
r
i
doxi
ne(
v
i
t
ami
nB6)
Tr
ansami
nat
i
on
Coenzy
meA(
CoASH)
Acy
l
t
r
ansf
er –
Py
r
i
doxal
phosphat
e(
Pl
.
P) Decar
boxy
l
at
i
on
Anemi
a
Racemi
zat
i
on
Cα–Cβbondcl
eav
age
α,
βEl
i
mi
nat
i
on
βSubst
i
t
ut
i
on
Vi
t
ami
nB12 Coenzy
meB12
Fol
i
caci
d(
f
ol
at
e)
I
somer
i
zat
i
onPer
ni
ci
ousanemi
a
Tet
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ahy
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of
ol
at
e(
THF)
Ascor
bi
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d(
v
i
t
ami
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–
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bont
r
ansf
er Megal
obl
ast
i
canemi
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Scur
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Wat
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nsol
ubl
e(
Li
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dSol
ubl
e)Vi
t
ami
ns
Vi
t
ami
nA
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–
–
Vi
t
ami
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cket
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Vi
t
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t
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Vi
t
ami
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y
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at
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on
–
Thi
st
abl
ei
sr
epr
oducedf
r
om
ht
t
p:
/
/
wps.
pur
shal
l
.
com/
wps/
medi
a/
obj
ect
s/
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nst
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uct
or
Resour
ces/
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_
25/
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xt
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mages/
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TB01.
JPG.
Repr
oducedf
r
om G.Li
t
wack,
HumanBi
ochemi
st
r
yandDi
sease,
Academi
cPr
ess/
El
sev
i
er
,
Tabl
e
31,
page119,
2008
Vi
ewchapt
er
Pur
chasebook
Enzy
mes
Rei
nhar
dRenneber
g,
.
.
.Vany
aLor
och,
i
nBi
ot
echnol
ogyf
orBegi
nner
s(
SecondEdi
t
i
on)
,
2017
2.
3TheRol
eofCof
act
or
si
nCompl
exEnzy
mes
Notal
l
enzy
mesconsi
stexcl
usi
v
el
yofpr
ot
ei
n,
asdoesl
y
sozy
me.Manyi
ncl
udeaddi
t
i
onal
chemi
cal
component
sorcof
act
or
swhi
chser
v
east
ool
s.Suchenzy
mesar
eknownasqual
i
f
i
ed
enzy
mesandhav
emor
ecompl
i
cat
edr
eact
i
onmechani
sms.
Cof
act
or
scanconsi
stofoneormor
ei
nor
gani
ci
ons(
suchasFe3+,
Mg2+,
Mn2+,
orZn2+)or
mor
ecompl
exor
gani
cmol
ecul
es,
knownascoenzy
mes.Someenzy
mesr
equi
r
ebot
ht
y
pesof
cof
act
or
s.
Coenzy
mesar
eor
gani
ccompoundst
hatbi
ndt
ot
heact
i
v
esi
t
eofenzy
mesorneari
t
.They
modi
f
yt
hest
r
uct
ur
eoft
hesubst
r
at
eormov
eel
ect
r
ons,
pr
ot
ons,
andchemi
cal
gr
oupsbackand
f
or
t
hbet
weenenzy
meandsubst
r
at
e,
negot
i
at
i
ngconsi
der
abl
edi
st
anceswi
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hi
nt
hegi
ant
enzy
memol
ecul
e.Whenusedup,
t
heysepar
at
ef
r
om t
hemol
ecul
e.
Manycoenzy
mesar
eder
i
v
edf
r
om v
i
t
ami
npr
ecur
sor
s,
whi
chexpl
ai
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equi
r
ea
const
antl
owl
ev
el
suppl
yofcer
t
ai
nv
i
t
ami
ns.Oneoft
hemostessent
i
al
coenzy
mes,
NAD+
(
ni
cot
i
nami
deadeni
nedi
nucl
eot
i
de)
,
i
sder
i
v
edf
r
om ni
aci
n.Mostwat
er
sol
ubl
ev
i
t
ami
nsoft
he
v
i
t
ami
nBgr
oupactascoenzy
mepr
ecur
sor
sv
er
ymuchl
i
keni
aci
n.
Ot
t
oHei
nr
i
chWar
bur
g(
1883–1970,
Fi
g.2.
3)di
scov
er
edt
her
espi
r
at
or
yenzy
mecy
t
ochr
ome
oxi
dase(
Fi
g.1.
14)andNAD.Hi
sdi
scov
er
yandsubsequentst
r
uct
ur
al
anal
y
si
swasoneoft
he
shi
ni
nghour
sofmoder
nbi
ochemi
st
r
y
.I
nt
heabsenceofni
aci
ni
nt
hedi
et
,
cer
t
ai
nenzy
mes(
e.
g.
,
dehy
dr
ogenases)cannotwor
kef
f
ect
i
v
el
yi
nt
hebody
.Theaf
f
ect
edhumanwi
l
l
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el
oppel
l
agr
a,
adi
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i
t
ami
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aci
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i
ci
ency
.Ot
t
oWar
bur
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el
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i
cal
t
est
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et
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i
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educedNADHatawav
el
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hof340nm (
t
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zedNAD+
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i
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hi
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el
engt
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.I
twasnowpossi
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et
omeasur
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i
al
enzy
me
r
eact
i
ons,
suchast
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ect
i
onofgl
ucoseusi
nggl
ucosedehy
dr
ogenase(
seeChapt
er
:
Anal
y
t
i
cal
Bi
ot
echnol
ogyandt
heHumanGenome)
.
Fi
gur
e2.
3.Ot
t
oHei
nr
i
chWar
bur
g(
1883–1970)di
scov
er
edt
hecof
act
orni
cot
i
nami
deadeni
ne
di
nucl
eot
i
de(
NAD)andr
espi
r
at
or
yenzy
mescont
ai
ni
ngi
r
on,
suchascy
t
ochr
omeoxi
dase(
see
Fi
g.1.
14)
.Hewasawar
dedt
heNobel
Pr
i
zei
n1941.
Nowaday
s,
v
i
t
ami
nsl
i
keB2(
r
i
bof
l
av
i
n)
,
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n)
,
andC(
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bi
caci
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e
pr
oducedbyt
het
onusi
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ot
echnol
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cal
met
hods(
seeChapt
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:
Whi
t
eBi
ot
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:
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l
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asSy
nt
het
i
cFact
or
i
es)
.
Cof
act
or
st
hatar
ecov
al
ent
l
ybondedt
ot
heenzy
mear
ecal
l
edpr
ost
het
i
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oups.Fl
av
i
nadeni
ne
di
nucl
eot
i
deact
sasapr
ost
het
i
cgr
oupf
orGOD.Per
oxi
daseandcy
t
ochr
omeP450cont
ai
na
hemegr
oup,
asf
oundi
nmy
ogl
obi
nandhemogl
obi
n.Thehemegr
oupi
t
sel
fconsi
st
sofa
por
phy
r
i
nr
i
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ncor
por
at
i
ngani
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oni
oni
ni
t
scent
er
.
Coenzy
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r
ast
,
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yl
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and,
j
ustl
i
kesubst
r
at
es,
t
heyunder
gochanges
i
nt
hebi
ndi
ngpr
ocessandar
eusedup.Unl
i
kesubst
r
at
es,
howev
er
,
t
heybi
ndt
oawhol
ehostof
enzy
mes(
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g.
,
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l
yal
l
dehy
dr
ogenases)andar
er
egener
at
edand
r
ecy
cl
edi
nsi
det
hecel
l
s(
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i
on2.
13)
.Enzy
mest
hatbi
ndt
ot
hesamecoenzy
meusual
l
y
r
esembl
eeachot
heri
nt
hei
rchemi
cal
mechani
sms.
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l
ewer
ef
er
r
edt
ot
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t
ool
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hepr
ot
ei
nsect
i
onoft
heenzy
mei
st
he
“
cr
af
t
sman”usi
ngt
heset
ool
s,
whoi
sr
esponsi
bl
ef
ort
hei
ref
f
ect
i
v
eness.Asal
way
s,
cr
af
t
smen
andt
ool
sr
el
yoneachot
hert
oachi
ev
et
hebestpossi
bl
er
esul
t
.
Vi
ewchapt
er
Pur
chasebook
Const
i
t
uent
soft
heHumanBody
Tsugi
kazuKomoda,
Toshi
y
uki
Mat
sunaga,
i
nBi
ochemi
st
r
yf
orMedi
cal
Pr
of
essi
onal
s,
2015
Cl
assi
f
i
cat
i
onofEnzy
mesi
nCat
al
y
si
s
Enzy
mescanbecl
assi
f
i
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i
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l
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ot
hedi
f
f
er
encebet
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eact
i
onand
subst
r
at
especi
f
i
ci
t
y
,
andt
hemechani
sm ofact
i
on.Theenzy
mecode(
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cl
assi
f
i
cat
i
on.Not
at
i
onaccor
di
ngt
oECnumber
,
i
.
e.ECX.
X.
X.
X,
i
sshownasf
ol
l
ows.Thef
i
r
st
ECnumbercl
assi
f
i
est
heenzy
mer
eact
i
onmechani
sm i
nt
osi
xgr
oups,
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y
oxi
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i
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on,
t
r
ansi
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i
on,
hy
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ol
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i
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nt
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s(
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i
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ht
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ni
t
i
al
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assi
f
i
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e:
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EC1.
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X.
Xr
educt
ase
•
EC2.
X.
X.
Xt
r
ansf
er
ase
•
EC3.
X.
X.
Xhy
dr
ol
ase
•
EC4.
X.
X.
Xl
y
ase
•
EC5.
X.
X.
Xi
somer
ase
•
EC6.
X.
X.
Xl
i
gase.
Al
t
houghst
andar
dsofcl
assi
f
i
cat
i
ondi
f
f
eri
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oup,
t
heyar
esubdi
v
i
dedbyt
hedi
f
f
er
ence
bet
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mer
eact
i
onandsubst
r
at
especi
f
i
ci
t
y
.Thesenumber
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eassi
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ot
heent
i
r
e
enzy
me,
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er3000enzy
mer
eact
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.Mor
eov
er
,
enzy
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eav
ar
i
et
yofact
i
v
i
t
i
es;
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orexampl
e,
ATPasecat
al
y
zest
hehy
dr
ol
y
si
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h
pr
ot
ei
nsandATP.Somet
i
mes,
t
hesubst
r
at
et
hatt
heenzy
memet
abol
i
zesi
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t
t
edf
r
om t
he
sy
st
emi
cnami
ng,
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ul
esasasy
st
emi
cname.Forexampl
e,
t
hesy
st
emi
c
nameofEC1.
1.
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1i
sal
cohol
dehy
dr
ogenase.Ther
ear
eal
somanyenzy
mesnamedi
n
accor
dancewi
t
ht
henami
ngconv
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on,
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y
mer
ase.
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t
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megener
al
l
yconsi
st
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ot
ei
n,
af
ewenzy
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ai
nnonpr
ot
ei
n
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d.Ther
i
bozy
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scov
er
edbyThomasCechandot
her
si
n
1986i
sacat
al
y
stmadeofRNA,
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chact
soni
t
sel
fandcl
eav
esRNA.
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equi
r
eot
hermol
ecul
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unct
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,
et
c.
)
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me,
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ot
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t
i
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t
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act
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,
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i
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i
v
i
t
y
,
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oenzy
me,
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ot
ei
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t
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v
i
t
y
.
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gani
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henonpr
ot
ei
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st
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eact
i
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nt
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i
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ecent
er
i
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l
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me.Si
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al
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ement
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nt
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i
v
ef
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me,
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ost
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oup.Al
t
hought
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erf
r
om t
y
pi
cal
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ost
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oups,
t
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l
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at
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om t
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meandbeconsumeddur
i
nganenzy
mer
eact
i
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t
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r
at
e
l
i
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e,
si
ncet
hecy
t
ochr
omeP450(
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mei
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ent
l
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t
ht
he
hemei
r
on,
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at
ef
r
om t
heCYPenzy
me.Ther
ef
or
e,
t
hi
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et
yi
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l
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me.Al
t
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i
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al
ent
l
yt
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me,
l
i
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at
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r
om t
heenzy
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et
y
,
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i
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ef
or
e,
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i
t
er
i
a
def
i
ni
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ost
het
i
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oupsar
enotst
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i
ct
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Anenzy
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i
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ot
ei
nchai
ns(
pept
i
dechai
n)
.Wheni
tconsi
st
s
oft
woormor
epept
i
dechai
ns,
eachpept
i
dechai
ni
scal
l
edasubuni
t
.
Vi
ewchapt
er
Pur
chasebook
Toxi
col
ogy
P.
D.Fel
gat
e,
i
nEncy
cl
opedi
aofFor
ensi
cSci
ences(
SecondEdi
t
i
on)
,
2013
Cl
onedEnzy
meDonorI
mmunoassay
CEDI
Ai
samor
er
ecenthomogeneousi
mmunoassayt
hatusest
hebi
ndi
ngofanant
i
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i
v
i
t
yofgenet
i
cal
l
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r
agment
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om Escher
i
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abel
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i
v
ef
r
agment
s,
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heenzy
me
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or(
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ai
nsasmal
l
por
t
i
onofenzy
memi
ssi
ngf
r
om
t
hel
ar
gerEAf
r
agment
.Ant
i
bodi
est
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ndt
ot
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ent
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ot
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r
agment
pr
ev
entt
her
eassoci
at
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v
i
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y
.Ast
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ugi
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agmentdecr
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i
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yduet
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zes
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edβgal
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he
enzy
meact
i
v
i
t
ycanbemeasur
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r
ophot
omet
r
i
cal
l
y
.
Vi
ewchapt
er
Pur
chasebook
TheNer
v
ousSy
st
emsofNonHumanPr
i
mat
es
D.
S.St
ol
zenber
g,
i
nEv
ol
ut
i
onofNer
v
ousSy
st
ems(
SecondEdi
t
i
on)
,
2017
3.
24.
2.
3.
1.
2.
2TheDy
nami
cI
nt
er
pl
ayofHi
st
oneAcet
y
l
t
r
ansf
er
aseEnzy
mesandHi
st
one
Deacet
y
l
aseEnzy
mesi
nRegul
at
i
ngGeneExpr
essi
on
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mesant
agoni
zeeachot
her
.Toel
uci
dat
eexact
l
yhowt
hei
nt
er
pl
aybet
ween
t
heset
woenzy
mesaf
f
ect
sgeneexpr
essi
on,
Wangetal
.(
2009)usedaChI
Psequenci
ng
t
echni
quet
oi
dent
i
f
yt
hel
ocat
i
onofsev
er
al
HATandHDACenzy
mest
hr
oughoutchr
omat
i
n.Not
sur
pr
i
si
ngl
yt
heyf
oundt
hatHATswer
er
ecr
ui
t
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oact
i
v
egenepr
omot
er
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t
i
v
el
y
cor
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el
at
edwi
t
hbot
hhi
st
oneacet
y
l
at
i
onandgenet
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anscr
i
pt
i
on.Unexpect
edl
y
,
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er
,
t
hey
f
oundt
hatt
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st
r
i
but
i
onofHDACenzy
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r
equent
l
yov
er
l
appedwi
t
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er
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ui
t
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oact
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v
egenepr
omot
er
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pi
cal
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yabsenti
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l
encedgenes.
Thesedat
asuggestt
hatmostHDACsf
unct
i
ont
or
esetchr
omat
i
ni
nact
i
v
egenepr
omot
er
sby
r
emov
i
ngt
heacet
y
l
at
i
ont
hatcont
r
i
but
edt
ot
r
anscr
i
pt
i
oni
ni
t
i
at
i
on.Thecoi
nci
dentr
ecr
ui
t
ment
oft
hesest
opandgosi
gnal
sconf
er
st
i
ghtt
empor
al
cont
r
ol
ont
r
anscr
i
pt
i
oni
nr
esponset
oacel
l
sur
f
acesi
gnal
byar
api
dr
epr
essi
onoft
r
anscr
i
pt
i
onwhent
hesi
gnal
hassubsi
ded.Ther
ef
or
e
act
i
v
egenepr
omot
er
scy
cl
ebet
weenacet
y
l
at
edanddeacet
y
l
at
edst
at
es.
Vi
ewchapt
er
Pur
chasebook
Pr
oduct
i
on,
Pur
i
f
i
cat
i
on,
andAppl
i
cat
i
onofMi
cr
obi
al
Enzy
mes
Ani
l
KumarPat
el
,
.
.
.AshokPandey
,
i
nBi
ot
echnol
ogyofMi
cr
obi
al
Enzy
mes,
2017
2.
1I
nt
r
oduct
i
on
Enzy
mesar
et
heact
i
v
epr
ot
ei
ns(
exceptRNAse)t
hatcancat
al
y
zebi
ochemi
cal
r
eact
i
ons.These
ar
ebi
omol
ecul
esr
equi
r
edf
orbot
hsy
nt
hesesaswel
l
asbr
eakdownr
eact
i
onsbyl
i
v
i
ng
or
gani
sms.Al
l
l
i
v
i
ngor
gani
smsar
ebui
l
tandmai
nt
ai
nedbyt
heseenzy
mes,
whi
char
et
r
ul
y
t
er
medasbi
ol
ogi
cal
cat
al
y
st
shav
i
ngt
hecapabi
l
i
t
yt
oconv
er
taspeci
f
i
ccompound(
as
subst
r
at
e)i
nt
opr
oduct
sathi
gherr
eact
i
onr
at
es.Li
kechemi
cal
cat
al
y
st
s,
enzy
mesi
ncr
easet
he
r
eact
i
onr
at
ebyl
ower
i
ngi
t
sact
i
v
at
i
onener
gy(
Ea)
,
hence,
pr
oduct
sar
ef
or
medf
ast
erand
r
eact
i
onsr
eacht
hei
requi
l
i
br
i
um st
at
emor
er
api
dl
y
.Ther
at
esofmostenzy
mat
i
cr
eact
i
onsar
e
mi
l
l
i
onsoft
i
mesf
ast
ert
hant
hoseoft
heuncat
al
y
zedr
eact
i
ons.Theycanper
f
or
m conv
er
si
ons
i
nmi
nut
esorev
eni
nsecondswhi
chot
her
wi
semayt
akehundr
edsofy
ear
s(
Dal
by
,
2003;
Ot
t
en
andQuax,
2005)
.Enzy
mesar
eknownt
ocat
al
y
zeabout4000bi
ochemi
cal
r
eact
i
onsi
nl
i
v
i
ng
bei
ngs(
Bai
r
och,
2000)
.Forexampl
e,
l
act
asei
sagl
y
cosi
dehy
dr
ol
aset
hati
sabl
et
ohy
dr
ol
y
ze
l
act
ose(
mi
l
ksugar
)i
nt
oi
t
sconst
i
t
uentgal
act
oseandgl
ucosemonomer
s.I
ti
spr
oducedby
v
ar
i
ousmi
cr
oor
gani
smsandal
soi
nt
hesmal
l
i
nt
est
i
neofhumansandot
hermammal
shel
pi
ng
t
odi
gestmi
l
kcompl
et
el
y
.Enzy
mesar
eal
soenant
i
osel
ect
i
v
ecat
al
y
st
s,
whi
chcanbeused
ei
t
heri
nt
hesepar
at
i
onofenant
i
omer
sf
r
om ar
acemi
cmi
xt
ur
eori
nt
hesy
nt
hesi
sofchi
r
al
compounds.
Humansr
ecogni
zedt
hei
mpor
t
anceofenzy
mest
housandsofy
ear
sago;
cl
ar
i
f
i
cat
i
onand
f
i
l
t
r
at
i
onofwi
nesandbeerbei
ngt
heear
l
i
estexampl
esoft
heappl
i
cat
i
onofi
ndust
r
i
al
enz
y
mes.
Enzy
meshav
ebeenusedi
nbr
ewi
ng,
baki
ng,
andal
cohol
pr
oduct
i
onsi
ncepr
ehi
st
or
i
ct
i
mes;
howev
er
,
t
heydi
dnotcal
l
t
hem enzy
mes.Oneoft
heear
l
i
estwr
i
t
t
enr
ef
er
encest
oenzy
mesi
s
f
oundi
nHomer
’
sGr
eekepi
cpoemsdat
i
ngf
r
om about800BC,
wher
ei
twasment
i
onedt
hat
enzy
meswer
eusedi
nt
hepr
oduct
i
onofcheese.TheJapanesehav
eal
sousednat
ur
al
l
y
occur
r
i
ngenzy
mesi
nt
hepr
oduct
i
onoff
er
ment
edpr
oduct
sl
i
kesake,
Japaneseschnapps
br
ewedf
r
om r
i
ce,
f
ormor
et
hanat
housandy
ear
s.Someenzy
meshav
ebeendesi
gnedby
nat
ur
et
of
or
m compl
exmol
ecul
esf
r
om si
mpl
eroneswhi
l
eot
her
shav
ebeendesi
gnedf
or
br
eaki
ngupcompl
exmol
ecul
esi
nt
osi
mpl
erones,
al
soaf
ewmodi
f
ymol
ecul
es.These
r
eact
i
onsi
nv
ol
v
et
hemaki
ngandbr
eaki
ngoft
hechemi
cal
bondsi
nt
hecomponent
s.Owi
ngt
o
t
hei
r“
speci
f
i
ci
t
y
,
”apr
oper
t
yofanenz
y
met
hatal
l
owsi
tt
or
ecogni
zeapar
t
i
cul
arsubst
r
at
et
hat
t
heyar
edesi
gnedt
ot
ar
get
,
t
heyar
eusef
ul
f
ori
ndust
r
i
al
pr
ocessesandar
ecapabl
eof
cat
al
y
zi
ngt
her
eact
i
onbet
weenpar
t
i
cul
archemi
cal
sev
eni
ft
heyar
epr
esenti
nmi
xt
ur
eswi
t
h
manychemi
cal
s.Theseenzy
mesar
eenv
i
r
onment
al
l
ysaf
e,
nat
ur
al
,
andar
eappl
i
edv
er
ysaf
el
y
i
nf
oodandev
enphar
maceut
i
cal
i
ndust
r
i
es.St
i
l
l
,
enzy
mesar
epr
ot
ei
ns,
whi
chl
i
keanypr
ot
ei
n
cancauseandhav
ecausedi
nt
hepastal
l
er
gi
cr
eact
i
ons,
hence,
pr
ot
ect
i
v
emeasur
esar
e
necessar
yi
nt
hei
rpr
oduct
i
onandappl
i
cat
i
ons.
Enzy
met
echnol
ogyi
sanev
erev
ol
v
i
ngbr
anchof“
Sci
enceandTechnol
ogy
.
”Wi
t
ht
he
i
nt
er
v
ent
i
onandi
nf
l
uenceofBi
ot
echnol
ogyandBi
oi
nf
or
mat
i
cs,
cont
i
nuousl
ynov
el
ori
mpr
ov
ed
appl
i
cat
i
onsofenzy
mesar
eemer
gi
ng.Wi
t
hnov
el
appl
i
cat
i
ons,
t
heneedf
orenzy
meswi
t
h
i
mpr
ov
edpr
oper
t
i
esar
eal
soemer
gi
ngsi
mul
t
aneousl
y
.Dev
el
opmentofcommer
ci
al
enzy
mesi
s
aspeci
al
i
zedbusi
nesswhi
chi
susual
l
yunder
t
akenbycompani
espossessi
nghi
ghski
l
l
si
n:
●
Scr
eeni
ngf
ornewandi
mpr
ov
edenzy
mes
●
Sel
ect
i
onofmi
cr
oor
gani
smsandst
r
ai
ni
mpr
ov
ementf
orqual
i
t
at
i
v
eandquant
i
t
at
i
v
e
i
mpr
ov
ement
●
Fer
ment
at
i
onf
orenzy
mepr
oduct
i
on
●
Lar
gescal
eenzy
mepur
i
f
i
cat
i
ons
●
For
mul
at
i
onofenzy
mesf
orsal
e
Enzy
met
echnol
ogyof
f
er
si
ndust
r
i
esandconsumer
sanoppor
t
uni
t
yt
or
epl
acepr
ocessesusi
ng
aggr
essi
v
echemi
cal
swi
t
hmi
l
dandenv
i
r
onment
al
l
yf
r
i
endl
yenzy
mepr
ocesses.About3000
enzy
mesar
eknownofwhi
chonl
y150–170ar
ebei
ngexpl
oi
t
edi
ndust
r
i
al
l
y
.Atpr
esentonl
y5%
ofchemi
cal
pr
oduct
sar
epr
oducedt
hr
oughabi
ol
ogi
cal
r
out
ei
nt
hi
sgr
eener
a.Howev
er
,
economi
cal
l
yf
easi
bl
eandecof
r
i
endl
yenzy
mat
i
cpr
ocessesar
eemer
gi
ngasal
t
er
nat
i
v
est
o
phy
si
cochemi
cal
andmechani
cal
pr
ocesses.Basedont
hedi
f
f
er
entappl
i
cat
i
onsect
or
s,
i
ndust
r
i
al
enzy
mescanbecl
assi
f
i
edas:
(
1)Enzy
mesi
nt
hef
oodi
ndust
r
y
,
(
2)Enzy
mesf
or
pr
ocessi
ngai
ds,
(
3)Enzy
mesasi
ndust
r
i
al
bi
ocat
al
y
st
,
(
4)Enzy
mesi
ngenet
i
cengi
neer
i
ng,
and
(
5)Enzy
mesi
ncosmet
i
cs.
Today
,
enzy
mesar
eenv
i
sagedast
hebr
eadandbut
t
erofbi
ot
echnol
ogybecauset
heyar
et
he
mai
nt
ool
sf
orsev
er
al
bi
ot
echnol
ogi
cal
t
echni
ques(
gener
est
r
i
ct
i
on,
l
i
gat
i
on,
andcl
oni
ng,
et
c.
)
,
bi
opr
ocesses(
f
er
ment
at
i
onandcel
l
cul
t
ur
e)
,
andi
nanal
y
t
i
csi
nhumanandani
mal
t
her
apyas
medi
ci
nesorasdr
ugt
ar
get
s.Fur
t
her
mor
et
heyf
i
ndappl
i
cat
i
onsi
nsev
er
al
ot
heri
ndust
r
i
es,
suchasf
oodandf
eed,
t
ext
i
l
es,
ef
f
l
uentandwast
et
r
eat
ment
,
paper
,
t
anner
y
,
baki
ng,
br
ewi
ng,
dai
r
y
,
phar
maceut
i
cal
s,
conf
ect
i
onar
y
,
et
c.(
Pandeyetal
.
,
2006)
.
Theenzy
mesut
i
l
i
zedt
odayar
eal
sof
oundi
nani
mal
s(
pepsi
n,
t
r
y
psi
n,
pancr
at
i
n,
andchi
mosi
n)
andpl
ant
s(
papai
n,
br
omel
ai
n,
andf
i
ci
n)
,
butmostoft
hem ar
eofmi
cr
obi
al
or
i
gi
n,
suchas
gl
ucoamy
l
ase,
αamy
l
ase,
pect
i
nases,
et
c.Theadv
ant
ageofusi
ngmi
cr
obesf
orenzy
me
pr
oduct
i
oni
st
hei
rhi
ghergr
owi
ngabi
l
i
t
i
es,
hi
gherpr
oduct
i
v
i
t
y
,
andt
hei
reasi
ergenet
i
c
mani
pul
at
i
onf
orenhancedenzy
mepr
oduct
i
on,
et
c.Enzy
mespr
oducedf
r
om mi
cr
obi
al
or
i
gi
ns
ar
et
er
medasmi
cr
obi
al
enzy
mes.Mi
cr
obesar
emai
nl
yexpl
oi
t
edi
ni
ndust
r
i
esf
orenzy
me
pr
oduct
i
on.Mor
eov
er
,
mi
cr
obi
al
enzy
mesar
esuppl
i
ed,
wel
l
st
andar
di
zed,
andmar
ket
edby
sev
er
al
compet
i
ngcompani
eswor
l
dwi
de.Dependi
ngont
het
y
peofpr
ocess,
enzy
mescanbe
usedi
nsol
ubl
ef
or
m(
ani
mal
pr
ot
easesandl
i
pasesi
nt
anner
y
)andi
ni
mmobi
l
i
zedf
or
m
(
i
somer
i
zat
i
onofgl
ucoset
of
r
uct
osebygl
ucosei
somer
ase)
.
Vi
ewchapt
er
Pur
chasebook
I
ndust
r
i
al
Appl
i
cat
i
onsofAsy
mmet
r
i
cSy
nt
hesi
s
R.
A.Shel
don,
i
nCompr
ehensi
v
eChi
r
al
i
t
y
,
2012
9.
15.
3.
3Scopeoft
heCLEATechnol
ogy
TheCLEAt
echnol
ogyhasbr
oadscopeandhasbeenappl
i
edt
oani
ncr
easi
ngl
ywi
desel
ect
i
on
ofhy
dr
ol
ases,
oxi
dor
educt
ases,
andl
y
ases.Asnot
edear
l
i
er
,
addi
t
i
onal
pr
ot
ei
nscanbe
i
ncor
por
at
edi
nt
oCLEAsbycopr
eci
pi
t
at
i
onandcr
ossl
i
nki
ng.Ext
r
apol
at
i
onoft
hi
snot
i
onl
eads
t
ot
hei
deaofpr
epar
i
ngmul
t
i
pur
poseCLEAsf
r
om cr
udeenzy
meext
r
act
sconsi
st
i
ngofmul
t
i
pl
e
enzy
mes.Forexampl
e,
Dal
al
etal
.
33pr
epar
edaCLEAf
r
om apor
ci
nepancr
eat
i
cacet
one
powderext
r
actcont
ai
ni
ngl
i
pase,
phosphol
i
paseA2,
andαamy
l
aseenzy
mes.Al
l
t
hr
eeenzy
me
act
i
v
i
t
i
eswer
ecompl
et
el
yr
et
ai
nedi
nt
heCLEAandt
hel
at
t
erwasr
ecy
cl
edt
hr
eet
i
meswi
t
hout
appr
eci
abl
el
ossofact
i
v
i
t
y
.Weuset
het
er
m mul
t
i
CLEAt
odescr
i
besuchCLEAst
hatcanbe
usedf
orper
f
or
mi
ngdi
f
f
er
entbi
ot
r
ansf
or
mat
i
onsi
ndependent
l
y
.Howev
er
,
weenv
i
sagedt
he
del
i
ber
at
ecoi
mmobi
l
i
zat
i
onoft
woormor
eenzy
mesi
nasi
ngl
eCLEAf
ort
hesol
epur
poseof
per
f
or
mi
ngt
woormor
ebi
ot
r
ansf
or
mat
i
onsi
namul
t
i
enzy
mecascadepr
ocess.Wehav
egi
v
en
t
henamecombi
CLEAst
osuchcat
al
y
st
sandexampl
esofcascadepr
ocessesper
f
or
medwi
t
h
suchcombi
CLEAswi
l
l
bedi
scussedl
at
er
.
Vi
ewchapt
er
Pur
chasebook
Enzy
mesandEnzy
meRegul
at
i
on
N.
V.Bhagav
an,
ChungEunHa,
i
nEssent
i
al
sofMedi
cal
Bi
ochemi
st
r
y
,
2011
Nomencl
at
ur
e
Enzy
mesar
egener
al
l
ynamedf
ort
hesubst
r
at
eorchemi
cal
gr
ouponwhi
cht
heyact
,
andt
he
namet
akest
hesuf
f
i
xase.Thus,
t
heenzy
met
hathy
dr
ol
y
zesur
eai
snamedur
ease.Exampl
es
ofexcept
i
onst
ot
hi
st
er
mi
nol
ogyar
et
r
y
psi
n,
pepsi
n,
andpapai
n,
whi
char
et
r
i
v
i
al
names.A
sy
st
emat
i
cnomencl
at
ur
ef
orenzy
meshasbeendev
el
opedbyt
heEnzy
meCommi
ssi
onoft
he
I
nt
er
nat
i
onal
Uni
onofBi
ochemi
st
r
y
.Thi
ssy
st
em pr
ov
i
desar
at
i
onal
andpr
act
i
cal
basi
sf
or
i
dent
i
f
i
cat
i
onofal
l
enzy
mescur
r
ent
l
yknown,
aswel
l
asf
ornewenzy
mes.Thesy
st
emat
i
cname
descr
i
best
hesubst
r
at
e,
t
henat
ur
eoft
her
eact
i
oncat
al
y
zed,
andot
herchar
act
er
i
st
i
cs(
see
Suppl
ement
ar
yMat
er
i
al
sont
hewebsi
t
ef
ort
hecompl
et
el
i
st
i
ngofenzy
menomencl
at
ur
e
desi
gnat
i
ons)
.
Vi
ewchapt
er
Pur
chasebook
ENZYMES|
Enzy
mesi
nPhy
si
ol
ogi
cal
Sampl
es
N.Ki
ba,
i
nEncy
cl
opedi
aofAnal
y
t
i
cal
Sci
ence(
SecondEdi
t
i
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