1
1M
CB
140
11/
29/0
6
The
gene
tics
of h
eter
ochr
omat
in
in m
etaz
oa
3M
CB
140
11/
29/0
64
MC
B 1
40 1
1/29
/06
Her
man
n Jo
seph
Mul
ler
1946
Nob
el P
rize
in M
edic
ine:
"for t
he d
isco
very
of t
he p
rodu
ctio
n of
m
utat
ions
by
mea
ns o
f X-r
ay ir
radi
atio
n"
2
5M
CB
140
11/
29/0
6
Whi
te w
ild-ty
pe
Whi
tem
utan
t
The
true
mea
ning
of "
red
eye
redu
ctio
n":
6M
CB
140
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29/0
612
.14
7M
CB
140
11/
29/0
612
.14
8M
CB
140
11/
29/0
6
Gen
e be
havi
or c
an c
hang
e de
pend
ing
on
whe
re o
n th
e ch
rom
osom
e th
e ge
ne li
es.
= “p
ositi
on e
ffect
”(ba
ris
the
mos
t com
mon
ly u
sed
exam
ple)
“Pos
ition
effe
ct v
arie
gatio
n”(P
EV
): ce
ll-to
-cel
l va
riabi
lity
of e
xpre
ssio
n of
a g
ene
that
has
bee
n re
loca
ted
to a
new
pos
ition
in th
e ge
nom
e.
Epi
gene
tic p
heno
men
on:
Sta
ble
chan
ge in
exp
ress
ion
with
out c
hang
e in
seq
uenc
e!
3
9M
CB
140
11/
29/0
6
Enh
ance
r of P
EV
Sup
pres
sor o
f PE
V
10M
CB
140
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29/0
6
2 ge
nes:
Su(
var)
2-5
Su(
var)
3-9
11M
CB
140
11/
29/0
612
MC
B 1
40 1
1/29
/06
4
13M
CB
140
11/
29/0
6
HP
1(S
arah
Elg
in)
(het
eroc
hrom
atin
pro
tein
1)
Iden
tifie
d in
a B
IOC
HE
MIC
AL
sche
me
to d
isco
ver p
rote
ins
that
are
ass
ocia
ted
with
he
tero
chro
mat
in.
14M
CB
140
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29/0
6
15M
CB
140
11/
29/0
6
HP
1 =
Su(
var)2
-5
Con
serv
ed in
hum
ans
and
in m
ice
(bot
h in
term
s of
seq
uenc
e an
d in
tranu
clea
rlo
catio
n!).
bioc
hem
istry
gene
tics
Why
doe
s H
P1
go to
pla
ces
that
HP
1 go
es to
?
16M
CB
140
11/
29/0
6
“Bio
chem
ical
epi
stas
is”
(T. J
enuw
ein)
Ove
rexp
ress
ion
of m
ouse
Su(
var)
3-9
lead
s to
a
MA
SS
IVE
redi
strib
utio
n of
HP
1 in
the
nucl
eus
of m
ouse
cel
ls.
5
17M
CB
140
11/
29/0
6
Who
wou
ld h
ave
thun
k it?
NC
BI:
Su(
var)
3-9
cont
ains
a d
omai
n (th
e S
ET
dom
ain)
that
is s
omew
hat s
imila
r to,
ah
em, R
UB
ISC
O m
ethy
ltran
sfer
ase.
Su(
var)3
-9 is
a H
ISTO
NE
met
hyltr
ansf
eras
e.
18M
CB
140
11/
29/0
6
His
tone
met
hyla
tion
19M
CB
140
11/
29/0
6
Cal
ling
Dav
id D
ucho
vny
and
Gill
ian
And
erso
n
•S
u(va
r)3-9
was
giv
en th
is n
ame
beca
use
it w
as th
e 9t
hge
ne is
olat
ed o
n th
e 3r
d
chro
mos
ome
in a
scr
een
for S
u(va
r)s.
•It
met
hyla
tes
lysi
ne 9
in h
isto
ne H
3.
This
was
dis
cove
red
18 y
ears
afte
r it w
as
nam
ed.
20M
CB
140
11/
29/0
6
And
fina
lly
•H
P1
pref
eren
tially
BIN
DS
his
tone
H3
met
hyla
ted
on ly
sine
9.
•Th
at’s
why
Su(
var)3
-9 d
eter
min
es
loca
lizat
ion
of H
P1
to h
eter
ochr
omat
in (i
t m
ethy
late
s hi
ston
es in
het
eroc
hrom
atin
).•
At l
east
in fi
ssio
n ye
ast,
and
perh
aps
in
wor
ms,
this
has
to d
o w
ith R
NA
i. ☺
6
21M
CB
140
11/
29/0
622
MC
B 1
40 1
1/29
/06
23M
CB
140
11/
29/0
6
HP1
HP1
24M
CB
140
11/
29/0
6
HP1
HP1HP1
HP1
HP1
HP1
HP1
HP1
==
=
7
25M
CB
140
11/
29/0
6
Rem
embr
ance
of t
hing
s pa
st:
chro
mat
in a
s an
epi
gene
tic v
ehic
le
26M
CB
140
11/
29/0
6
Hom
olog
y(o
rthol
ogs
of h
eter
ocho
mat
in p
rote
ins
in fi
ssio
n ye
ast,
inse
cts,
and
hum
ans)
27M
CB
140
11/
29/0
6
Ana
logy
Fiss
ion
yeas
t, fli
es, m
amm
als.
Bud
ding
yea
st.
28M
CB
140
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29/0
6
Nat
ure,
Oct
ober
10,
200
2
The
poly
com
bgr
oup
prot
ein
EZH
2is
invo
lved
in p
rogr
essi
on o
f pr
osta
te c
ance
rV
aram
bally
et a
l.
Pros
tate
can
cer i
s a
lead
ing
caus
e of
can
cer-r
elat
ed d
eath
in m
ales
and
is
seco
nd o
nly
to lu
ng c
ance
r. A
lthou
gh e
ffect
ive
surg
ical
and
radi
atio
n tre
atm
ents
exi
st fo
r clin
ical
ly lo
caliz
ed p
rost
ate
canc
er, m
etas
tatic
pro
stat
e ca
ncer
rem
ains
ess
entia
lly in
cura
ble.
Her
e w
e sh
ow, t
hrou
gh g
ene
expr
essi
on p
rofil
ing,
that
the
poly
com
b gr
oup
prot
ein
enha
ncer
of z
este
hom
olog
2 (E
ZH2)
is o
vere
xpre
ssed
in h
orm
one-
refra
ctor
y, m
etas
tatic
pr
osta
te c
ance
r. …
Dys
regu
late
d ex
pres
sion
of E
ZH2
may
be
invo
lved
in th
e pr
ogre
ssio
n of
pro
stat
e ca
ncer
, as
wel
l as
bein
g a
mar
ker t
hat d
istin
guis
hes
indo
lent
pro
stat
e ca
ncer
from
thos
e at
risk
of l
etha
l pro
gres
sion
.
8
29
From
egg
to e
mbr
yo
?
30
31M
CB
140
11/
29/0
6
Hom
eotic
mut
atio
ns (W
. Bat
eson
)
“…N
ot th
at th
ere
has
mer
ely
been
a c
hang
e, b
ut th
at
som
ethi
ng h
as b
een
chan
ged
into
the
liken
ess
of s
omet
hing
el
se.”
Gen
etic
s
Alle
le
Het
eroz
ygou
s
Hom
ozyg
ous
32M
CB
140
11/
29/0
6
wt
ante
nnap
edia
9
33M
CB
140
11/
29/0
6
34
The
segm
enta
tion
hier
arch
y
35M
CB
140
11/
29/0
6
“Do
you
have
any
idea
who
I th
ink
I am
?!!”
1.S
egm
ent i
dent
ity is
det
erm
ined
by
trans
crip
tion
fact
ors.
2.Th
ey a
ct o
n ta
rget
gen
es o
nly
trans
ient
ly. T
hen
they
go
away
, and
the
activ
ity o
f the
ir ta
rget
s is
m
aint
aine
d by
larg
e co
mpl
exes
: Pol
ycom
b re
pres
ses
gene
s, a
nd T
ritho
rax
activ
ates
them
.3.
Nob
ody
knew
how
Pol
ycom
b an
d Tr
ithor
ax d
o th
is.
36M
CB
140
11/
29/0
6
How
Pol
ycom
b an
d Tr
ithor
ax w
ork
10
37M
CB
140
11/
29/0
6
extra
sex
com
bs
enha
ncer
of z
este
38M
CB
140
11/
29/0
6
E(z
) doe
s it
Post
ed S
epte
mbe
r 13,
200
2 –
CEL
L im
med
iate
ear
ly p
ublic
atio
n
Cze
rmin
, B.,
Mel
fi, R
., M
cCab
e, D
., S
eitz
, V.,
Imho
f, A
., an
d P
irrot
ta, V
. D
roso
phila
Enh
ance
r of Z
este
/ES
C c
ompl
exes
hav
e a
hist
one
H3
met
hyltr
ansf
eras
eac
tivity
that
mar
ks c
hrom
osom
al P
olyc
omb
site
s.
Cel
l. P
ublis
hed
onlin
e S
epte
mbe
r 13,
200
2.
10.1
016/
S009
2867
4020
0975
3
Mül
ler,
J., H
art,
C.M
., Fr
anci
s, N
.J.,
Var
gas,
M.L
., S
engu
pta,
A.,
Wild
, B
., M
iller,
E.L
., O
'Con
nor,
M.B
., K
ings
ton,
R.E
., an
d S
imon
, J.A
. H
isto
ne m
ethy
ltran
sfer
ase
activ
ity o
f a D
roso
phila
Pol
ycom
b gr
oup
repr
esso
r com
plex
. Cel
l. P
ublis
hed
onlin
e S
epte
mbe
r 13,
200
2.
10.1
016/
S009
2867
4020
0976
5
39M
CB
140
11/
29/0
640
MC
B 1
40 1
1/29
/06
“Influ
entia
l ide
as a
re a
lway
s si
mpl
e. S
ince
nat
ural
ph
enom
ena
need
not
be
sim
ple,
we
mas
ter t
hem
, if
at a
ll, b
y fo
rmul
atin
g si
mpl
e id
eas
and
expl
orin
g th
eir
limita
tions
.”
Al H
ersh
ey
11
41M
CB
140
11/
29/0
6
Reg
ulat
ion
of g
enes
occ
urs
via
the
inte
ract
ion
of tr
ans-
actin
g fa
ctor
s (p
rote
ins)
with
cis
-act
ing
sequ
ence
s ne
ar th
e ge
nes
them
selv
es.
+st
imul
us
+
42M
CB
140
11/
29/0
6
4344
Bic
oid
is th
e an
terio
r m
orph
ogen
12
45M
CB
140
11/
29/0
646
MC
B 1
40 1
1/29
/06
Wha
t dem
ocra
cy, I
mea
n, g
ene
regu
latio
n, is
real
ly li
ke•
Tran
s-ac
ting
fact
ors
do n
ot d
istri
bute
in th
e nu
cleu
s ba
sed
on th
e pr
imar
y se
quen
ce o
f the
gen
ome:
som
e fa
ctor
s fa
il to
bin
d m
ost g
enes
that
hav
e se
quen
ces
wai
ting
for t
hem
, and
oth
er fa
ctor
s bi
nd a
larg
e nu
mbe
r of
gen
es th
at d
o N
OT
have
seq
uenc
es fo
r the
m•
Eve
n w
hen
a fa
ctor
bin
ds n
ext t
o a
gene
, man
y tim
es,
noth
ing
happ
ens;
the
sam
e fa
ctor
bou
nd to
two
diffe
rent
ge
nes
can
exer
t dia
met
rical
ly o
ppos
ite e
ffect
s•
Mos
t gen
es in
the
hum
an g
enom
e ar
e un
der
cons
ider
able
regu
lato
ry in
fluen
ce fr
om e
ntiti
es o
ther
than
“s
impl
e” tr
ans-
actin
g fa
ctor
s; th
ese
entit
ies
incl
ude
nonc
odin
g R
NA
and
mod
ified
his
tone
s
47M
CB
140
11/
29/0
6
Boy
er a
nd Y
oung
C
ell S
ept.
23, 2
005
Fisc
hle,
Wan
g, A
llis
CO
CB
200
3
Dav
id A
llis: “
the
hist
one
code
”
13
51M
CB
140
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29/0
6
1963
-200
020
00 -
…
Hen
ry e
t al.
(11/
1/20
03) G
enes
Dev
. 17:
264
8.
Gen
etic
info
rmat
ion
Lac
oper
ator
gaat
tgtg
agcg
gata
acaa
ttt
14
Gen
etic
info
rmat
ion
Gen
etic
info
rmat
ion
-
+?