1
Ge
n
e
ti
c P
r
ac
ti
c
um
Re
port
B
io
c
h
e
mi
ca
l s
e
p
a
r
a
tion of
e
y
e
pigm
e
nts
D
rosophil
a
m
e
l
a
nog
a
st
e
r
W
ith thin l
a
y
e
r
c
hrom
a
togr
a
phy
B
r
ad
for
d Haa
g
A
rizon
a S
t
a
t
e U
niv
e
rsity
B
iology
De
p
a
rtm
e
nt
Ma
r
c
h
2024
Ab
str
ac
t
D
rosophil
a Me
l
a
nog
a
st
e
r
'
s
e
y
e
pigm
e
nt involv
e
s two l
a
n
e
s
:
ommo
c
hrom
e a
n
d
pt
e
ri
d
in
e
.
T
h
e
ommo
c
hrom
e
p
a
th pro
d
u
ce
s
b
rown pigm
e
nts
,
whil
e
pt
e
ri
d
in
e
pro
d
u
ce
s r
ed
,
y
e
llow
a
n
d b
lu
e
pigm
e
nts
.
M
ut
a
tions on th
e
s
e
p
a
thw
a
ys
ca
n
c
h
a
ng
e
th
e c
olor of th
e e
y
e
pigm
e
nt
.
D
iff
e
r
e
n
ce
s in
pigm
e
nts
be
tw
ee
n
D
rosophil
a Me
l
a
nog
a
st
e
r norm
a
l
a
n
d
mut
a
nts
ca
n
be a
n
a
lyz
ed
using
c
hrom
a
togr
a
phy
.
C
hrom
a
togr
a
phy
,
e
sp
ec
i
a
lly thin l
a
y
e
r
c
hrom
a
togr
a
phy
,
s
e
p
a
r
a
t
e
s
c
olor
c
ompon
e
nts
ba
s
ed
on
d
iff
e
r
e
n
ce
s in migr
a
tion r
a
t
e
in th
e
st
a
tion
a
ry ph
a
s
e
.
T
h
e a
n
a
lysis shows th
a
t
norm
a
l f
e
m
a
l
e e
y
e
pigm
e
nts
c
ont
a
in
c
ompon
e
nts su
c
h
a
s
d
rosopt
e
rin
e a
n
d
x
a
nthopt
e
rin
e
,
whil
e
th
e
mut
a
nt
e
y
e
pigm
e
nt s
e
pi
a
only
c
ont
a
ins
b
iopt
e
rin
e a
n
d
isos
e
pi
a
p
e
rin
e
.
M
ut
a
tions
ca
us
e
th
e
mut
a
nt
e
y
e
s
e
pi
a
to turn into
da
rk
b
rown
.
C
hrom
a
togr
a
phi
c
r
e
sults
a
r
e
m
ea
sur
ed b
y
a
p
a
r
a
m
e
t
e
r
ca
ll
ed
r
e
t
e
ntion f
ac
tor
.
2
1
.
P
r
e
limin
a
ry
M
ut
a
tions
ca
n o
cc
ur on th
e
ommo
c
hrom
e a
n
d
pt
e
ri
d
in
e
p
a
ths in th
e P
igm
e
nt of
D
rosophil
a
Me
l
a
nog
a
st
e
r
,
ca
using
e
y
e c
olor
c
h
a
ng
e
s
.
T
h
e
mut
a
tion o
cc
urs
d
uring th
e
pro
ce
ss of
c
h
a
nging
DNA
to
RNA a
n
d
th
e
n
bec
om
e
s
a
prot
e
in
,
from tr
a
ns
c
ription to tr
a
nsl
a
tion
,
in
acc
or
da
n
ce
with
Ce
ntr
a
l
D
ogm
a
(
N
i
c
holl
2008
:
15
).
T
his mut
a
tion
ca
n o
cc
ur in g
e
n
e
s th
a
t r
e
gul
a
t
e D
rosophil
a
Me
l
a
nog
a
st
e
r
'
s
e
y
e
pigm
e
nts
,
support th
e bead
l
e
hypoth
e
sis
a
n
d
t
a
tum th
a
t
eac
h g
e
n
e e
n
c
o
de
s
on
e
sp
ec
ifi
c e
nzym
e
or prot
e
in
(
Ra
v
e
n
e
t
a
l
2001
:
296
).
T
o t
e
st th
e
hypoth
e
sis
,
Bead
l
e a
n
d Ta
tum isol
a
t
e
th
e M
ut
a
nt
Ne
urospor
a
,
whi
c
h grows
b
oth
in
a
rtifi
c
i
a
l m
ed
ium
a
n
d d
isinown
UV
to in
c
r
ea
s
e
mut
a
tions
(
Ra
v
e
n
e
t
a
l
2001
:
296
).
T
h
e
r
e
sults
show
ed a c
h
a
ng
e
in
a
mino
ac
i
d
s
,
proving th
a
t th
e
m
e
t
ab
olism of th
e
org
a
nism
de
p
e
n
d
s on th
e
e
nzym
e c
o
ded b
y th
e
g
e
n
e
.
T
his hypoth
e
sis is known
a
s
O
n
e Ge
n
e
-
O
n
e E
nzym
e
(
Pa
ss
a
rg
e
2001
:
17
).
A
lthough
eac
h g
e
n
e
pro
d
u
ce
s prot
e
in
,
not
a
ll prot
e
ins
a
r
e e
nzym
e
s
.
Ke
r
a
tin
a
n
d
hormon
e
s
a
r
e e
x
a
mpl
e
s of non
-
e
nzym
e
prot
e
ins
(
Ca
mp
be
ll
e
t
a
l
2008
:
373
).
Beca
us
e
m
a
ny prot
e
ins
c
onsist of
s
e
v
e
r
a
l polyp
e
pti
de c
h
a
ins
,
eac
h polyp
e
pti
de
is
c
o
ded b
y
a
p
a
rti
c
ul
a
r g
e
n
e
,
th
e
hypoth
e
sis of
O
n
e
Ge
n
e
-
O
n
e E
nzym
e
is
c
h
a
ng
ed
to
O
n
e Ge
n
e
-
O
n
e P
olyp
e
pti
de
(
Ra
v
e
n
e
t
a
l
2001
:
297
).
Bead
l
e a
n
d Ta
tum formul
a
t
e
th
e
four
ba
si
c
prin
c
ipl
e
s of th
e
hypoth
e
sis of on
e
g
e
n
e
-
on
e
polyp
e
pti
de
: (
1
)
g
e
n
e
s
a
r
e
v
e
ry import
a
nt in
b
io
c
h
e
mi
ca
l pro
ce
ss
e
s
,
beca
us
e eac
h g
e
n
e
r
e
gul
a
t
e
s
th
e
pro
ce
ss
. (
2
)
B
io
c
h
e
mi
ca
l pro
ce
ss
e
s
a
r
e
gr
ad
u
a
l
. (
3
)
Eac
h
b
io
c
h
e
mi
ca
l r
eac
tion is gov
e
rn
ed b
y
3
ce
rt
a
in g
e
n
e
s
,
m
ea
ning th
a
t on
e
g
e
n
e
pro
d
u
ce
s
a
sp
ec
i
a
l prot
e
in
. (
4
)
M
ut
a
tions in g
e
n
e
s
ca
n
c
h
a
ng
e ce
ll
ab
iliti
e
s to
ca
rry out
b
io
c
h
e
mi
ca
l r
eac
tions
(
S
uryo
2008
:
285
).
T
h
e
thr
ee
m
a
in pro
ce
ss
e
s in th
e ce
ll inform
a
tion p
a
th
a
r
e
r
e
pli
ca
tion
(
d
ou
b
ling
DNA
),
tr
a
ns
c
ription
(
DNA c
h
a
ng
e
s to
RNA
),
a
n
d
tr
a
nsl
a
tion
(
c
h
a
ng
e
s in
RNA
into prot
e
in
).
T
his
c
on
ce
pt is
ca
ll
ed Ce
ntr
a
l
D
ogm
a b
y
F
r
a
n
c
is
C
ri
c
k in
1958
(
P
i
e
r
ce e
t
a
l
2005
:
281
).
T
his inform
a
tion
ca
n
c
h
a
ng
e
c
h
a
ng
e
s through mut
a
tions
,
whi
c
h
a
r
e c
h
a
ng
e
s in th
e c
omposition of
DNA
nu
c
l
e
oti
de
s
spont
a
n
e
ously or
d
u
e
to mut
a
g
e
n
(
S
olomon
e
t
a
l
2005
:
260
;
R
ittn
e
r
e
t
a
l
2004
:
237
).
C
h
a
ng
e
s in g
e
n
e
s
ca
n
be de
t
ec
t
ed
using
c
hrom
a
togr
a
phy
,
m
e
tho
d
s th
a
t s
e
p
a
r
a
t
e a
n
d
i
de
ntify
c
ompon
e
nts with simil
a
r
c
h
e
mi
ca
l
a
n
d
physi
ca
l prop
e
rti
e
s
.
C
hrom
a
togr
a
phy fun
c
tions
ba
s
ed
on
mol
ec
ul
a
r int
e
r
ac
tions through st
a
tion
a
ry ph
a
s
e
s th
a
t
a
r
e
influ
e
n
ced b
y th
e
motion ph
a
s
e
,
wh
e
r
e
solv
e
nts
ac
t
a
s th
e
motion ph
a
s
e
.
M
ol
ec
ul
e
s th
a
t mov
e be
tw
ee
n s
e
p
a
r
a
t
e
ph
a
s
e
s
a
r
e ba
s
ed
on th
e
d
istin
c
tion of th
e
ro
a
r
,
with th
e
pol
a
r
c
ompon
e
nt mor
e ea
sily
d
issolv
ed
in th
e
ph
a
s
e
of th
e
pol
a
r
motion
a
n
d
vi
ce
v
e
rs
a
.
T
h
e
r
e a
r
e
v
a
rious typ
e
s of
c
hrom
a
togr
a
phy
ba
s
ed
on th
e
st
a
tion
a
ry ph
a
s
e
,
n
a
m
e
ly
:
-
Ad
sorption
c
hrom
a
togr
a
phy
:
us
e
solution or g
a
s
a
s
a
motion ph
a
s
e ab
sor
bed b
y
a
soli
d
st
a
tion
a
ry
ph
a
s
e
.
-
Pa
rtition
c
hrom
a
togr
a
phy
:
s
e
p
a
r
a
t
e
su
b
st
a
n
ce
s
ba
s
ed
on solu
b
ility in th
e
motion
a
n
d
st
a
tion
a
ry
ph
a
s
e
in th
e
form of solution
.
-
I
on
e
x
c
h
a
ng
e
r
c
hrom
a
togr
a
phy
:
using ions
a
s
a
st
a
tion
a
ry ph
a
s
e a
n
d
solution
a
s
a
motion ph
a
s
e
,
s
e
p
a
r
a
ting su
b
st
a
n
ce
s
ba
s
ed
on
e
l
ec
trost
a
ti
c c
h
a
rg
e
s
.
-
Pe
rm
ea
s
a
tion
c
hrom
a
togr
a
phy
:
us
e
porous g
e
l to s
e
p
a
r
a
t
e c
ompon
e
nts
ba
s
ed
on siz
e
,
without
4
int
e
r
ac
tion with th
e
motion ph
a
s
e
.
-
A
ffinity
c
hrom
a
togr
a
phy
:
s
e
p
a
r
a
t
e ba
s
ed
on sp
ec
ifi
c
int
e
r
ac
tions
be
tw
ee
n st
a
tion
a
ry ph
a
s
e
s
a
n
d
motion ph
a
s
e
s through ion or p
H c
h
a
ng
e
s
.
T
hin l
a
y
e
r
c
hrom
a
togr
a
phy
(
TLC
)
a
n
d
high
-
p
e
rform
a
n
ce
thin l
a
y
e
r
c
hrom
a
togr
a
phy
(
HTLC
)
a
r
e
c
hrom
a
togr
a
phi
c
v
a
ri
a
nts th
a
t us
e a
thin l
a
y
e
r of sili
ca
g
e
l
,
a
lumin
a
,
or pl
a
sti
c
on th
e
su
b
str
a
t
e a
s
a
st
a
tion
a
ry ph
a
s
e
.
T
h
e
ph
a
s
e
of th
e
mov
e
m
e
nt is solv
e
nt
,
a
n
d
s
e
p
a
r
a
tion o
cc
urs
ba
s
ed
on solu
b
ility
d
iff
e
r
e
n
ce
s
.
T
h
e c
hrom
a
togr
a
phi
c
m
e
tho
d
is us
ed
to
a
n
a
lyz
e c
ompoun
d
s su
c
h
a
s pigm
e
nts
,
for
e
x
a
mpl
e e
y
e
pigm
e
nts on
D
rosophil
a Me
l
a
nog
a
st
e
r
.
T
his pigm
e
nt in
c
lu
de
s pt
e
ri
d
in
e a
n
d
ommo
c
hrom
e
,
whi
c
h
a
ff
ec
ts
e
y
e c
olor
.
Da
m
a
g
e
to this pigm
e
nt syst
e
m
ca
n
c
h
a
ng
e
th
e c
olor of th
e e
y
e
.
T
h
e
pigm
e
nt will
mov
e
to
a ce
rt
a
in position on th
e c
hrom
a
togr
a
m
,
ca
ll
ed
th
e RF
(
Re
t
e
ntion
Fac
tor
)
v
a
lu
e
,
whi
c
h is
ca
l
c
ul
a
t
ed a
s th
e
r
a
tio of
c
ompon
e
nt
d
ispl
ace
m
e
nt
d
ist
a
n
ce
to solv
e
nt
d
ist
a
n
ce
.
T
hin l
a
y
e
r
c
hrom
a
togr
a
phy pr
ac
ti
ce
s
a
im to un
de
rst
a
n
d
th
e
t
ec
hniqu
e
of s
e
p
a
r
a
tion of pigm
e
nts in
D
rosophil
a Me
l
a
nog
a
st
e
r
,
th
e
working prin
c
ipl
e
of
c
hrom
a
togr
a
phy
,
a
n
d
th
e e
ff
ec
t of mut
a
tion in
ph
e
notyp
e
s
.
2
.
T
h
e
m
e
tho
d
ology
T
ools us
ed
in thin l
a
y
e
r
c
hrom
a
togr
a
phy pr
ac
ti
ce
s to s
e
p
a
r
a
t
e
th
e d
rosophil
a
m
e
l
a
nog
a
st
e
r
e
y
e
pigm
e
nt in
c
lu
d
ing s
c
rip
e
rs
,
b
ottl
e
sp
ec
im
e
ns
,
e
th
e
riz
e
r
b
ottl
e
s
,
b
ottl
e
s of j
a
m
,
n
eed
l
e
son
de
,
tw
ee
z
e
rs
,
pip
e
tt
e
s
,
p
e
tri
d
ish
e
s
,
ca
r
b
on p
a
p
e
r
,
G
lov
e
s
,
UV
tr
a
nsmitt
e
rs
,
UV
prot
ec
tiv
e
gl
a
ss
e
s
,
5
st
a
tion
e
ry
,
a
n
d
rul
e
rs
.
T
h
e
m
a
t
e
ri
a
l us
ed
in
c
lu
de
s
a
norm
a
l f
e
m
a
l
e d
rosophil
a a
n
d
s
e
pi
a
mut
a
nt
,
nh
4
oh solution
,
n
-
propyl
a
l
c
ohol solution
,
sili
ca
p
a
p
e
r
(
TLC
),
a
n
d d
i
e
thyl
e
th
e
r
.
P
r
ac
ti
ca
l st
e
ps
be
gin with pr
e
p
a
ring sili
ca
p
a
p
e
r
,
giving
1
c
m lin
e
from th
e e
n
d
of using
a
p
e
n
c
il
,
a
n
d
m
a
rking two points with
a d
ist
a
n
ce
of
2
c
m
.
A
voi
d
tou
c
hing p
a
p
e
r
b
y h
a
n
d
to pr
e
v
e
nt
c
ont
a
min
a
tion
.
D
rosophil
a Me
l
a
nog
a
st
e
r is f
e
m
a
l
e
norm
a
l isol
a
t
ed
,
s
eda
t
ed
in
d
i
e
thyl
e
th
e
r
,
a
n
d
th
e
e
y
e
s
a
r
e
t
a
k
e
n with
a
son
de
n
eed
l
e
.
E
y
e
s
a
r
e
pl
aced
on
c
hrom
a
togr
a
phi
c
p
a
p
e
r
a
n
d
wr
e
stl
ed
with
a
gl
a
ss short
e
r
.
T
h
e
s
a
m
e
pro
ce
ss w
a
s r
e
p
ea
t
ed
for mut
a
nt s
e
pi
a
.
S
ili
ca
p
a
p
e
r is ins
e
rt
ed
into
a b
ottl
e
of j
a
m
c
ont
a
ining
a
mixtur
e
of
NH
4
OH a
n
d N
-
P
ropil
a
l
c
ohol un
de
r th
e
sour
c
los
e
t
,
th
e
n
c
ov
e
r
ed
with
ca
r
b
on p
a
p
e
r
.
T
h
e
p
a
p
e
r w
a
s illumin
a
t
ed b
y
UV
for
75
minut
e
s
,
th
e
n t
a
k
e
n with tw
ee
z
e
rs
a
n
d
o
b
s
e
rv
ed
un
de
r
UV
using th
e
glov
e
s
a
n
d
prot
ec
tiv
e
gl
a
ss
e
s
.
D
rosophil
a Me
l
a
nog
a
st
e
r
'
s
e
y
e
pigm
e
nt
c
ompon
e
nt looks lik
e a c
olor on
c
hrom
a
togr
a
phi
c
p
a
p
e
r
a
n
d
is m
ea
sur
ed
from th
e
lin
e
to
ca
l
c
ul
a
t
e
th
e RF
v
a
lu
e
.
3
.
Re
sults
a
n
d D
is
c
ussion
T
h
e
s
e
p
a
r
a
tion of
D
rosophil
a Me
l
a
nog
e
r
'
s
e
y
e
pigm
e
nt is
d
on
e
using thin l
a
y
e
r
c
hrom
a
togr
a
phy
,
with
a
1
-
2
c
m lin
ed
lin
e
from th
e e
n
d
to f
ac
ilit
a
t
e RF
m
ea
sur
e
m
e
nt
a
n
d da
t
a
v
a
li
d
ity
.
T
h
e d
ist
a
n
ce be
tw
ee
n points in th
e
pl
a
t
e a
lso
a
voi
d
s mixing pigm
e
nts
.
T
h
e
pigm
e
nt must
be c
rush
ed
to
be c
l
ea
rly visi
b
l
e
un
de
r
UV
(
Occ
i
de
nt
a
l
C
oll
e
g
e
2012
:
2
).
6
T
h
e
solv
e
nt us
ed
is
NH
4
OH a
n
d N
-
P
ropil
a
l
c
ohol
.
NH
4
OH b
in
d
s th
e c
ompon
e
nt in th
e
st
a
tion
a
ry ph
a
s
e
,
whil
e
th
e N
-
P
ropil
a
l
c
ohol
ea
sily
e
v
a
por
a
t
e
s
,
a
llows pigm
e
nt
c
ompon
e
nts to
be
dede
m
be
r
ed
to
c
hrom
a
togr
a
phi
c
pl
a
t
e
s
(
M
orris
2012
:
6
).
T
h
e
solv
e
nt is stor
ed
in
a ca
r
b
on p
a
p
e
r
-
pl
a
t
ed b
ottl
e
to prot
ec
t from
UV
r
a
ys th
a
t
ca
n int
e
rf
e
r
e
with pigm
e
nt o
b
s
e
rv
a
tions
.
C
hrom
a
togr
a
phi
c
pl
a
t
e
is l
e
ft for
75
minut
e
s in th
e ac
i
d
i
c
sp
ace
for m
a
ximum s
e
p
a
r
a
tion
(
M
orris
2012
:
7
).
C
hrom
a
togr
a
phi
c
r
e
sults
a
r
e
o
b
s
e
rv
ed
un
de
r
UV
r
a
ys
beca
us
e
pigm
e
nts
a
r
e
not s
ee
n
without th
e
h
e
lp of
UV
.
UV
r
a
ys
a
r
e
n
eeded beca
us
e
light
e
n
e
rgy s
ee
ms not in
acc
or
da
n
ce
with
pigm
e
nt
e
n
e
rgy
.
UV
o
b
s
e
rv
a
tions
a
lso r
e
quir
e
glov
e
s to prot
ec
t from
UV e
xposur
e
th
a
t
ca
n
ca
us
e
mut
a
tions
(
SAMAL
2010
:
1
).
T
hin l
a
y
e
r
c
hrom
a
togr
a
phy m
a
k
e
s it
ea
si
e
r
a
n
d
impr
ac
t th
e
s
e
p
a
r
a
tion of mol
ec
ul
e
s
.
T
his
m
e
tho
d
is mor
e e
ff
ec
tiv
e
th
a
n p
a
p
e
r
c
hrom
a
togr
a
phy
,
with th
e be
n
e
fits of
a
simpl
e
o
b
s
e
rv
a
tion
pro
ced
ur
e
,
goo
d
s
e
p
a
r
a
tion sp
eed
,
high s
e
nsitivity
,
a
n
d
th
e ab
ility to us
e
org
a
ni
c
s
a
mpl
e
s
(
E
ugin
e
2011
:
2
).
D
iff
e
r
e
n
ce
s in th
e
int
e
nsity of pigm
e
nts in f
e
m
a
l
e a
n
d
norm
a
l
d
rosophil
a
m
e
l
a
nog
a
st
e
r
P
igm
e
nts on
D
rosophil
a Me
l
a
nog
a
st
e
r f
e
m
a
l
e a
n
d
norm
a
l m
a
l
e
show
d
iff
e
r
e
n
ce
s
.
Fe
m
a
l
e
s
h
a
v
e b
right
e
r pigm
e
nts th
a
n m
a
l
e
s
(
Sc
ott
1996
:
1086
).
T
his is
beca
us
e c
hromosom
e X
in f
e
m
a
l
e
s
b
rings g
e
n
e
s th
a
t
a
ff
ec
t pigm
e
nt synth
e
sis
,
so f
e
m
a
l
e
s th
a
t h
a
v
e
two
c
hromosom
e
s x shows high
e
r
pigm
e
nt int
e
nsity th
a
n m
a
l
e
s th
a
t only h
a
v
e
on
e c
hromosom
e X
(
Sc
ott
1996
:
1086
).
7
T
h
e
r
e
sults of th
e
pigm
e
nt o
b
s
e
rv
a
tions in f
e
m
a
l
e
s
Ob
s
e
rv
a
tions from our group
(
4
D
)
show
ed
th
a
t
D
rosophil
a N
orm
a
l f
e
m
a
l
e Me
l
a
nog
a
st
e
r
f
ea
tur
ed
thr
ee
s
e
qu
e
nti
a
l pigm
e
nts
:
y
e
llow
c
h
ee
rful with
a
v
a
lu
e
of
RF
0
.
5
,
gr
ee
nish y
e
llow with
a
v
a
lu
e
of
RF
0
.
76
,
a
n
d b
luish gr
ee
n with
a
v
a
lu
e
of
RF
0
.
83
.
Fe
m
a
l
e
mut
a
nt s
e
pi
a
only shows
b
luish
gr
ee
n with
RF
v
a
lu
e
of
0
.
89
.
P
igm
e
nt
de
t
a
ils in norm
a
l f
e
m
a
l
e
s
a
n
d
mut
a
nts
D
rosophil
a Me
l
a
nog
a
st
e
r
N
orm
a
l f
e
m
a
l
e
s show
ed
th
e P
igm
e
nt
c
ompon
e
nt
a
s follows
:
RF
0
.
5
for
D
rosopt
e
rin
e
(
y
e
llow
c
h
ee
r
e
n
a
n
),
RF
0
.
76
for
SEPIAPTERINE
(
gr
ee
nish y
e
llow
),
a
n
d RF
0
.
83
for
Xa
nthopt
e
rin
e
(
gr
ee
n
b
luish
).
Fe
m
a
l
e
mut
a
nt s
e
pi
a
with
RF
0
.
89
only shows x
a
nthopt
e
rin
e
(
c
h
ad
sp
ace
:
1
).
Ob
s
e
rv
a
tions
a
t norm
a
l m
a
l
e a
n
d
mut
a
nt
D
rosophil
a Me
l
a
nog
a
st
e
r
Ma
l
e N
orm
a
l from th
e
5
D
group shows th
e P
igm
e
nt
c
ompon
e
nt
a
s
follows
:
RF
0
.
09
for
D
rosopt
e
rin
e
(
OREN
),
RF
0
.
021
for isox
a
nthopt
e
rin
e
(
gr
ee
nish
b
lu
e
),
RF
0
.
52
for
ALEPTERINE
(
Ye
llow
),
RF
0
.
78
for
a
fr
a
nti
c
in
e
(
OREN
),
a
n
d RF
0
.
84
for
I
sos
e
pi
aP
t
e
rin
e
(
Ye
llow
).
T
h
e
8
W
hit
e M
ut
a
nt
Ma
l
e
in th
e
1
C
group
d
i
d
not show pigm
e
nts
a
n
d
look whit
e
.
Sca
rl
e
t
Ma
l
e M
ut
a
nt
from
G
roup
5
D
shows
:
RF
0
.
15
for
D
rosopt
e
rin
e
(
O
r
e
n
),
RF
0
.
50
for
Xa
nthopt
e
rin
e
(
gr
ee
n
),
RF
0
.
73
for
SEPIAPTERINE
(
Ye
llow
),
a
n
d RF
0
.
78
for
SEPIAPTERINE
(
Ye
llow
) (
C
h
ad
sp
ace
:
1
).
C
omp
a
rison with th
e
lit
e
r
a
tur
e
T
h
e
r
e
sults of th
e
o
b
s
e
rv
a
tion of th
e N
orm
a
l
D
rosophil
a Me
l
a
nog
a
st
e
r
E
y
e P
igm
e
nt show
ed
l
e
ss th
a
n s
e
v
e
n pigm
e
nts
,
d
iff
e
r
e
nt from th
e
lit
e
r
a
tur
e
th
a
t m
e
ntions s
e
v
e
n
P
t
e
ri
d
in
e P
igm
e
nts
(
Occ
i
de
nt
a
l
C
oll
e
g
e
2009
:
7
).
Se
pi
a
mut
a
nts show four pigm
e
nts
,
c
ontr
a
ry to th
e
lit
e
r
a
tur
e
(
C
h
ad
sp
ace
:
1
).
W
hit
e
mut
a
nts
d
o not show pigm
e
nts
,
acc
or
d
ing to th
e L
it
e
r
a
tur
e
(
Occ
i
de
nt
a
l
C
oll
e
g
e
2009
:
7
).
Sca
rl
e
t mut
a
nts h
a
v
e
four of th
e e
xp
ec
t
ed
s
e
v
e
n pt
e
ri
d
in
e
pigm
e
nts
,
a
n
d d
o not
h
a
v
e B
rown pigm
e
nts
d
u
e
to mut
a
tions on th
e
ommo
c
hrom
e
p
a
th
,
d
iff
e
r
e
nt from th
e L
it
e
r
a
tur
e
(
Occ
i
de
nt
a
l
C
oll
e
g
e
2009
:
6
).
C
on
c
lusion
C
hrom
a
togr
a
phy is
a
m
e
tho
d
for s
e
p
a
r
a
ting
c
h
e
mi
ca
l
c
ompon
e
nts
ba
s
ed
on th
e
migr
a
tion r
a
t
e
in th
e
st
a
tion
a
ry ph
a
s
e
or sil
e
nt ph
a
s
e
.
T
hin l
a
y
e
r
c
hrom
a
togr
a
phy
ca
n
be
us
ed
for i
de
ntifi
ca
tion of
D
rosophil
a Me
l
a
nog
a
st
e
r
'
s
e
y
e
pigm
e
nt
.
Se
p
a
r
a
t
e
pigm
e
nts
acc
or
d
ing to th
e
ir
c
h
e
mi
ca
l
c
h
a
r
ac
t
e
risti
c
s
.
T
h
e
pigm
e
nt on
D
rosophil
a Me
l
a
nog
a
st
e
r norm
a
lly
c
om
e
s from th
e
pt
e
ri
d
in
e a
n
d
9
ommo
c
hrom
e
p
a
th
,
pro
d
u
ce
s
a
r
ed c
olor
,
whil
e
th
e
mut
a
nt s
e
pi
a
shows
da
m
a
g
e
to th
e
pt
e
ri
d
in
e
p
a
th
,
pro
d
u
ce
s
da
rk
c
ho
c
ol
a
t
e
.
T
h
e d
iff
e
r
e
n
ce
in r
e
sults o
b
s
e
rv
ed c
omp
a
r
ed
to th
e
lit
e
r
a
tur
e
m
a
y
be ca
us
ed b
y
e
rrors in th
e
insul
a
tion of pigm
e
nts
,
in
c
lu
d
ing
c
ont
a
min
a
tion
a
n
d e
xposur
e
to th
e
pr
ac
ti
ca
lity
.
Re
f
e
r
e
n
ce
s
Ca
mp
be
ll
,
N
.
A
.,
J
.
B
.
Reece
,
L
.
A
.
U
rr
a
y
,
M
.
L
.
Ca
in
,
P
.
V
.
M
inorsky
,
R
.
B Jac
kson
&
S
.
A
.
Wa
ss
e
rm
a
n
.
2008
.
B
iology
.
8
th
ed
.
Pea
rson
Ed
u
ca
tion
I
n
c
.,
Sa
n
F
r
a
n
c
is
c
o
:
1465
p
.
C
h
ad
sp
ace
.
Se
p
a
r
a
tion of
D
rosophil
a
E
y
e
pigm
e
nt with
c
hrom
a
togr
a
phy
.
1
pp
.
http
://
c
h
ad
sp
ace
.
unwis
d
om
.
org
/
wp
-
c
ont
e
nt
/
uplo
ad
s
/
2013
/
06
/
l
ab
-
15
-
s
e
p
a
r
a
tion
-
of
-
d
rosophil
a
-
e
y
e
-
pigm
e
nts
-
with
-
c
hrom
a
togr
a
phy
.
p
d
f
.
A
pril
4
,
2015
PK
.
14
.
15
C
l
a
rk
,
J
im
.
2007
.
T
hin l
a
y
e
r
c
hrom
a
togr
a
phy
.
1
pp
.
http
://
www
.
c
h
e
mgui
de
.
c
o
.
uk
/
a
n
a
lysis
/
c
hrom
a
togr
a
phy
/
thinl
a
y
e
r
.
html
.
A
pril
3
PK
.
11
.
37
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ug
e
n
e
,
B
.
M
.
2011
.
E
xp
e
rim
e
nt
6
:
T
hin
La
y
e
r
C
hrom
a
togr
a
phy
.
6
p
.
http
://
www
.
acade
mi
a
.
ed
u
/
2112360
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