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Intron
From Wikipedia, the free encyclopedia
For the interferon-based drug used in viral and cancer treatments, see Intron A.


Representation of intron and exons within a simple gene containing a single intron.
An intron is any nucleotide sequence within a gene that is removed by RNA splicing while the final mature
RNA product of a gene is being generated.
[1][2]
The term intron refers to both the DNA sequence within a gene,
and the corresponding sequence in RNA transcripts.
[3]
Sequences that are joined together in the final mature
RNA after RNA splicing are exons. Introns are found in the genes of most organisms and many viruses, and
can be located in a wide range of genes, including those that generate proteins, ribosomal RNA (rRNA),
and transfer RNA(tRNA). When proteins are generated from intron-containing genes, RNA splicing takes place
as part of the RNA processing pathway that follows transcription and precedes translation.
The word intron is derived from the term intragenic region; i.e., a region inside a gene. Although introns are
sometimes called intervening sequences, the term "intervening sequence" can refer to any of several families
of internal nucleic acid sequences that are not present in the final gene product, including inteins, untranslated
sequences (UTR), and nucleotides removed by RNA editing, in addition to introns.
Contents
[hide]
1 Simple illustration of an intron and RNA splicing
o 1.1 1. Identify exon and intron sequences
o 1.2 2. Cleavage at intron-exon junctions
o 1.3 3. Link the exons together
2 Introduction
3 Classification
4 Biological functions and evolution
5 See also
6 References
7 External links
[edit]Simple illustration of an intron and RNA splicing
Introns and the process of RNA splicing can be illustrated using a text example. Consider the following
quotation from Groucho Marx:
"Outside of a dog, a book is man's best friend. Inside of a dog it's too
dark to read."
To make this text look more like a nucleic acid sequence, we'll simply
delete all punctuation and make all of the text uppercase. This will be our
"message".
OUTSIDEOFADOGABOOKISMANSBESTFRIENDINSIDEOFADOGIT
STOODARKTOREAD
If there were a single intron within this sequence, it would look
something like this
[edit]1. Identify exon and intron sequences
To remove the intron, the sequences corresponding to the intron and
exons must be precisely identified. Here, the exon sequences are
changed to bold. Note that misidentifying the boundaries between the
exons and intron by even a single letter will result in a nonsensical
message.
OUTSIDEOFADOGABOOKISMANSBEGUITUITGLSAJSAKHDLAYSI
OEYASHDKLSALKDN
KLASNDKLASKGDASJKBDNKNSKDKLSANDKNSKANDKNSAKNDK
SAKLDHSDJGJASBDKN
SANDLNSMALVJOPDHVANVLNVKLSAKNFADNGKLHKNIUHSAFLS
AFLFASFOLSANFLNK
FNKDSNGOIFSGHSKDHGKDHSFIABHFIHEHRFKHLKDHSUNSIUAN
IDAUBIOAYICMUSF
AVMASSTFRIENDINSIDEOFADOGITSTOODARKTOREAD
[edit]2. Cleavage at intron-exon
junctions
Next, the sequence must be cut precisely at the
boundaries between the exons and the intron
OUTSIDEOFADOGABOOKISMANSBE
GUITUITGLSAJSAKHDLAYSIOEYASHDKLSALKDN
KLASNDKLASKGDASJKBDNKNSKDKLSANDKNSKANDKNSAKNDK
SAKLDHSDJGJASBDKN
SANDLNSMALVJOPDHVANVLNVKLSAKNFADNGKLHKNIUHSAFLS
AFLFASFOLSANFLNK
FNKDSNGOIFSGHSKDHGKDHSFIABHFIHEHRFKHLKDHSUNSIUAN
IDAUBIOAYICMUSF
AVMAS
STFRIENDINSIDEOFADOGITSTOODARKTOREAD
[edit]3. Link
the exons
together
Last, the exons
must be joined
together to
generate the
final message.
(The excised
intron is usually
broken down to
its component
letters, which
are then
recycled.)
OUTSIDEOFADOGABOOKISMANSBESTFRIENDINSIDEOFADOGIT
STOODARKTOREAD
[edit]Intr
oductio
n
Introns
were first
discovered
in protein-
coding
genes
of adenovir
us,
[4][5]
and
were
subsequen
tly
identified
in genes
encoding
transfer
RNA and
ribosomal
RNA
genes.
Introns are
now known
to occur
within a
wide
variety of
genes
throughout
organisms
and
viruses
within all of
the
biological
kingdoms.
The fact
that genes
were split
or
interrupted
by introns
was
discovered
independe
ntly in
1977
by Phillip
Allen
Sharp and
Richard J.
Roberts,
for which
they
shared
the Nobel
Prize in
Physiology
or
Medicine i
n
1993.
[6]
Th
e
term intron
was
introduced
by
American
biochemist
Walter
Gilbert:
[7]

"T
he
no
tio
n
of
th
e
cis
tro
n
[...
]
m
us
t
be
re
pl
ac
ed
by
th
at
of
a
tra
ns
cri
pti
on
un
it
co
nt
ai
ni
ng
re
gi
on
s
w
hi
ch
wil
l
be
lo
st
fro
m
th
e
m
at
ur
e
m
es
se
ng
er
-
w
hi
ch
I
su
gg
es
t
w
e
ca
ll
int
ro
ns
(fo
r
int
ra
ge
ni
c
re
gi
on
s)
-
alt
er
na
tin
g
wit
h
re
gi
on
s
w
hi
ch
wil
l
be
ex
pr
es
se
d -
ex
on
s."
(G
ilb
ert
19
78
)
The
frequency
of introns
within
different
genomes
is
observed
to vary
widely
across the
spectrum
of
biological
organisms.
For
example,
introns are
extremely
common
within the
nuclear
genome of
higher
vertebrates
(e.g.
humans
and mice),
where
protein-
coding
genes
almost
always
contain
multiple
introns,
while
introns are
rare within
the nuclear
genes of
some
eukaryotic
microorgan
isms,
[8]
for
example b
aker's/bre
wer's
yeast (Sac
charomyce
s
cerevisiae)
. In
contrast,
the mitoch
ondrial
genomes o
f
vertebrates
are entirely
devoid of
introns,
while those
of
eukaryotic
microorgan
isms may
contain
many
introns.
Introns are
well known
in bacterial
and
archaeal
genes, but
occur more
rarely than
in most
eukaryotic
genomes.


Simple
illustrati
on of an
unsplice
d
mRNA
precurs
or, with
two
introns
and
three
exons
(top).
After
the
introns
have
been
remove
d via
splicing,
the
mature
mRNA
sequen
ce is
ready
for
translati
on
(bottom
).
[edit]Cla
ssificati
on
Splicing of
all intron-
containing
RNA
molecules
is
superficiall
y similar,
as
described
above.
However,
different
types of
introns
were
identified
through
the
examinatio
n of intron
structure
by DNA
sequence
analysis,
together
with
genetic
and
biochemic
al analysis
of RNA
splicing
reactions.
At least
four
distinct
classes of
introns
have been
identified.
[1]

Intron
s in
nuclea
r
protei
n-
coding
genes
that
are
remov
ed
by spli
ceoso
mes
Intron
s in
nuclea
r and
archa
eal
transf
er
RNA
genes
that
are
remov
ed by
protei
ns
(tRNA
splicin
g
enzym
es)
Self-
splicin
g grou
p I
introns
that
are
remov
ed
by RN
A
cataly
sis.
Self-
splicin
g grou
p II
introns
that
are
remov
ed
by RN
A
cataly
sis
Group III
introns are
proposed
to be a fifth
family, but
little is
known
about the
biochemic
al
apparatus
that
mediates
their
splicing.
They
appear to
be related
to group II
introns,
and
possibly to
spliceosom
al introns.
[9]

Nuclear
pre-mRNA
introns
(spliceoso
mal
introns)
are
characteriz
ed by
specific
intron
sequences
located at
the
boundaries
between
introns and
exons.
[10]
T
hese
sequences
are
recognized
by
spliceosom
al RNA
molecules
when the
splicing
reactions
are
initiated.
[11]
In
addition,
they
contain a
branch
point, a
particular
nucleotide
sequence
near the 3'
end of the
intron that
becomes
covalently
linked to
the 5' end
of the
intron
during the
splicing
process,
generating
a branched
(lariat)
intron.
Apart from
these three
short
conserved
elements,
nuclear
pre-mRNA
intron
sequences
are highly
variable.
Nuclear
pre-mRNA
introns are
often much
longer than
their
surroundin
g exons.
Group I
and group
II introns
are found
in genes
encoding
proteins
(messenge
r
RNA), tran
sfer
RNA and ri
bosomal
RNA in a
very wide
range of
living
organisms.
,
[12][13]
Follo
wing
transcriptio
n into
RNA,
group I
and group
II introns
also make
extensive
internal
interaction
s that allow
them to
fold into a
specific,
complex th
ree-
dimension
al
architectur
e. These
complex
architectur
es allow
some
group I
and group
II introns to
be self-
splicing,
that is, the
intron-
containing
RNA
molecule
can
rearrange
its own
covalent
structure
so as to
precisely
remove the
intron and
link the
exons
together in
the correct
order. In
some
cases,
particular
intron-
binding
proteins
are
involved in
splicing,
acting in
such a way
that they
assist the
intron in
folding into
the three-
dimension
al structure
that is
necessary
for self-
splicing
activity.
Group I
and group
II introns
are
distinguish
ed by
different
sets of
internal
conserved
sequences
and folded
structures,
and by the
fact that
splicing of
RNA
molecules
containing
group II
introns
generates
branched
introns
(like those
of
spliceosom
al RNAs),
while
group I
introns use
a non-
encoded
guanosine
nucleotide
(typically
GTP) to
initiate
splicing,
adding it
on to the
5'-end of
the
excised
intron.
Transfer
RNA
introns that
depend
upon
proteins for
removal
occur at a
specific
location
within the
anticodon
loop of
unspliced
tRNA
precursors,
and are
removed
by a tRNA
splicing
endonucle
ase. The
exons are
then linked
together by
a second
protein, the
tRNA
splicing
ligase.
[14]
N
ote that
self-
splicing
introns are
also
sometimes
found
within
tRNA
genes.
[15]

[edit]Biol
ogical
functio
ns and
evoluti
on
As a first
approximat
ion, it is
possible to
view
introns as
unimportan
t
sequences
whose only
function is
to be
removed
from an
unspliced
precursor
RNA in
order to
generate
the
functional
mRNA,
rRNA or
tRNA
product.
However, it
is now
well-
establishe
d that
some
introns
themselve
s encode
specific
proteins or
can be
further
processed
after
splicing to
generate n
oncoding
RNAmolec
ules.
[16]
Alt
ernative
splicing is
widely
used to
generate
multiple
proteins
from a
single
gene.
Furthermor
e, some
introns
represent
mobile
genetic
elements a
nd may be
regarded
as
examples
of selfish
DNA.
[17]

The
biological
origins of
introns are
obscure.
After the
initial
discovery
of introns
in protein-
coding
genes of
the
eukaryotic
nucleus,
there was
significant
debate as
to whether
introns in
modern-
day
organisms
were
inherited
from a
common
ancient
ancestor
(termed
the introns-
early
hypothesis
), or
whether
they
appeared
in genes
rather
recently in
the
evolutionar
y process
(termed
the introns-
late
hypothesis
). Another
theory is
that
the spliceo
some and
the intron-
exon
structure of
genes is a
relic of
the RNA
world (the
introns-first
hypothesis
).
[18]
There
is still
considerab
le debate
about the
extent to
which of
these
hypothese
s is most
correct.
The
popular
consensus
at the
moment is
that introns
arose
within the
eukaryote
lineage as
selfish
elements.
Early
studies of
genomic
DNA
sequences
from a
wide range
of
organisms
show that
the intron-
exon
structure of
homologou
s genes in
different
organisms
can vary
widely.
[19]

More
recent
studies of
entire euka
ryotic geno
mes have
now shown
that the
lengths
and
density
(introns/ge
ne) of
introns
varies
considerab
ly between
related
species.
For
example,
while the
human
genome
contains
an average
of 8.4
introns/gen
e (139,418
in the
genome),
the
unicellular
fungus Enc
ephalitozo
on
cuniculi co
ntains only
0.0075
introns/gen
e (15
introns in
the
genome).
[2
0]
Since
eukaryotes
arose from
a common
ancestor
(Common
descent),
there must
have been
extensive
gain and/or
loss of
introns
during
evolutionar
y
time.
[21][22]

This
process is
thought to
be subject
to
selection,
with a
tendency
towards
intron gain
in larger
species
due to their
smaller
population
sizes, and
the
converse
in smaller
(particularl
y
unicellular)
species.
[23]

Biological
factors
also
influence
which
genes in a
genome
lose or
accumulat
e
introns.
[24][2
5][26]

Alternative
splicing of
introns
within a
gene acts
to
introduce
greater
variability
of protein
sequences
translated
from a
single
gene,
allowing
multiple
related
proteins to
be
generated
from a
single
gene and a
single
precursor
mRNA
transcript.
The control
of
alternative
RNA
splicing is
performed
by
complex
network of
signaling
molecules
that
respond to
a wide
range of
intracellula
r and
extracellul
ar signals.
Introns
contain
several
short
sequences
that are
important
for efficient
splicing,
such as
acceptor
and donor
sites at
either end
of the
intron as
well as a
branch
point site,
which are
required
for proper
splicing by
the spliceo
some.
Some
introns are
known to
enhance
the
expression
of the gene
that they
are
contained
in by a
process
known
as intron-
mediated
enhancem
ent (IME).
[edit]See
also
Structure:
Exon
mRNA
Eukar
yotic
chrom
osome
fine
structu
re
Small
t
intron
Splicing:
Altern
ative
splicin
g
Minor
splice
osome
Function
Micro
RNA
Others:
Intein
Interru
pted
gene
Nonco
ding
DNA
Nonco
ding
RNA
Selfish
DNA
Twintr
on
[edit]Ref
erence
s
1. ^
a

b
A
lb
er
ts,
Br
uc
e
(2
0
0
8)
.
M
ol
ec
ul
ar
bi
ol
o
gy
of
th
e
ce
ll.
N
e
w
Y
or
k:
G
ar
la
n
d
S
ci
e
nc
e.
I
S
B
N
0-
8
1
5
3-
4
1
0
5-
9.
2. ^
St
ry
er
,
L
u
b
er
t;
B
er
g,
Je
re
m
y
M
ar
k;
T
y
m
oc
zk
o,
Jo
h
n
L.
(2
0
0
7)
.
Bi
oc
h
e
mi
st
ry
.
S
a
n
Fr
a
nc
is
co
:
W
.H
.
Fr
e
e
m
a
n.
I
S
B
N
0-
7
1
6
7-
6
7
6
6-
X.
3. ^
Ki
n
ni
b
ur
g
h,
Al
a
n;
m
er
tz,
j.
a
n
d
R
os
s,
J.
(J
ul
y
1
9
7
8)
. "
T
h
e
pr
ec
ur
so
r
of
m
o
us
e
-
gl
o
bi
n
m
es
se
n
g
er
R
N
A
co
nt
ai
ns
tw
o
int
er
ve
ni
n
g
R
N
A
se
q
u
e
nc
es
".
C
ell
1
4
(3
):
6
8
1

6
9
3.
d
oi:
1
0.
1
0
1
6/
0
0
9
2-
8
6
7
4(
7
8)
9
0
2
5
1-
9.
P
M
ID
6
8
8
3
8
8.
4. ^
C
h
o
w
L
T,
G
eli
n
as
R
E,
Br
ok
er
T
R,
R
o
b
er
ts
R
J
(S
e
pt
e
m
b
er
1
9
7
7)
.
"A
n
a
m
az
in
g
se
q
u
e
nc
e
ar
ra
n
g
e
m
e
nt
at
th
e
5'
e
n
ds
of
a
d
e
n
ov
ir
us
2
m
es
se
n
g
er
R
N
A"
.
C
ell
1
2
(1
):
1

8.
d
oi:
1
0.
1
0
1
6/
0
0
9
2-
8
6
7
4(
7
7)
9
0
1
8
0-
5.
P
M
ID
9
0
2
3
1
0.
5. ^
B
er
g
et
S
M
,
M
o
or
e
C,
S
h
ar
p
P
A
(A
u
g
us
t
1
9
7
7)
. "
S
pli
ce
d
se
g
m
e
nt
s
at
th
e
5'
te
r
mi
n
us
of
a
d
e
n
ov
ir
us
2
lat
e
m
R
N
A"
.P
ro
c.
N
atl
.
A
ca
d.
S
ci.
U.
S.
A.
7
4
(8
):
3
1
7
1

5.
d
oi:
1
0.
1
0
7
3/
p
n
as
.7
4.
8.
3
1
7
1.
P
M
C
4
3
1
4
8
2.
P
M
ID
2
6
9
3
8
0.
6. ^
ht
tp
://
w
w
w.
n
o
b
el
pr
iz
e.
or
g/
n
o
b
el
_
pr
iz
es
/
m
e
di
ci
n
e/l
a
ur
e
at
es
/1
9
9
3/
pr
es
s.
ht
ml
7. ^
Gi
lb
er
t,
W
alt
er
(
1
9
7
8)
.
"
W
hy
g
e
n
es
in
pi
ec
es
".
N
at
ur
e
2
7
1(
5
6
4
5)
:
5
0
1

5
0
1.
d
oi:
1
0.
1
0
3
8/
2
7
1
5
0
1
a
0.
P
M
ID
6
2
2
1
8
5.
8. ^
St
aji
ch
J
E,
Di
et
ric
h
F
S,
R
oy
S
W
(2
0
0
7)
. "
C
o
m
p
ar
ati
ve
g
e
n
o
mi
c
a
n
al
ys
is
of
fu
n
g
al
g
e
n
o
m
es
re
ve
al
s
int
ro
n-
ric
h
a
nc
es
to
rs
".
G
e
n
o
m
e
Bi
ol.
8
(
1
0)
:
R
2
2
3.
d
oi:
1
0.
1
1
8
6/
g
b-
2
0
0
7-
8-
1
0-
r2
2
3.
P
M
C
2
2
4
6
2
9
7.
P
M
ID
1
7
9
4
9
4
8
8.
9. ^
C
o
p
er
tin
o
D
W
,
H
all
ic
k
R
B
(D
ec
e
m
b
er
1
9
9
3)
.
"
G
ro
u
p
II
a
n
d
gr
o
u
p
III
int
ro
ns
of
tw
int
ro
ns
:
p
ot
e
nti
al
re
lat
io
ns
hi
ps
wi
th
n
uc
le
ar
pr
e-
m
R
N
A
int
ro
ns
".
Tr
e
n
ds
Bi
oc
h
e
m
.
S
ci.
1
8
(1
2)
:
4
6
7

7
1.
P
M
ID
8
1
0
8
8
5
9.
10. ^
P
a
d
g
et
t
R
A,
G
ra
b
o
w
sk
i
P
J,
K
o
n
ar
sk
a
M
M
,
S
eil
er
S,
S
h
ar
p
P
A
(1
9
8
6)
.
"S
pli
ci
n
g
of
m
es
se
n
g
er
R
N
A
pr
ec
ur
so
rs
".
A
n
n
u.
R
ev
.
Bi
oc
h
e
m
.
5
5:
1
1
1
9

5
0.
d
oi:
1
0.
1
1
4
6/
a
n
n
ur
ev
.bi
.5
5.
0
7
0
1
8
6.
0
0
5
3
5
1.
P
M
ID
2
9
4
3
2
1
7.
11. ^
G
ut
hr
ie
C,
P
at
te
rs
o
n
B
(1
9
8
8)
.
"S
pli
ce
os
o
m
al
sn
R
N
A
s"
.A
n
n
u.
R
ev
.
G
e
n
et
.
2
2:
3
8
7

4
1
9.
d
oi:
1
0.
1
1
4
6/
a
n
n
ur
ev
.g
e.
2
2.
1
2
0
1
8
8.
0
0
2
1
3
1.
P
M
ID
2
9
7
7
0
8
8.
12. ^
C
ec
h
T
R
(1
9
9
0)
.
"S
elf
-
sp
lic
in
g
of
gr
o
u
p
I
int
ro
ns
".
A
n
n
u.
R
ev
.
Bi
oc
h
e
m
.
5
9:
5
4
3

6
8.
d
oi:
1
0.
1
1
4
6/
a
n
n
ur
ev
.bi
.5
9.
0
7
0
1
9
0.
0
0
2
5
5
1.
P
M
ID
2
1
9
7
9
8
3.
13. ^
Mi
ch
el
F,
F
er
at
JL
(1
9
9
5)
.
"S
tr
uc
tu
re
a
n
d
ac
tiv
iti
es
of
gr
o
u
p
II
int
ro
ns
".
A
n
n
u.
R
ev
.
Bi
oc
h
e
m
.
6
4:
4
3
5

6
1.
d
oi:
1
0.
1
1
4
6/
a
n
n
ur
ev
.bi
.6
4.
0
7
0
1
9
5.
0
0
2
2
5
1.
P
M
ID
7
5
7
4
4
8
9.
14. ^
G
re
er
C
L,
P
e
e
bl
es
C
L,
G
e
g
e
n
h
ei
m
er
P,
A
b
el
so
n
J
(F
e
br
u
ar
y
1
9
8
3)
.
"
M
ec
h
a
ni
s
m
of
ac
tio
n
of
a
ye
as
t
R
N
A
lig
as
e
in
tR
N
A
sp
lic
in
g"
.
C
ell
3
2
(2
):
5
3
7

4
6.
d
oi:
1
0.
1
0
1
6/
0
0
9
2-
8
6
7
4(
8
3)
9
0
4
7
3-
7.
P
M
ID
6
2
9
7
7
9
8.
15. ^
R
ei
n
h
ol
d-
H
ur
ek
B,
S
h
u
b
D
A
(
M
ay
1
9
9
2)
.
"S
elf
-
sp
lic
in
g
int
ro
ns
in
tR
N
A
g
e
n
es
of
wi
d
el
y
di
ve
rg
e
nt
b
ac
te
ri
a"
.
N
at
ur
e
3
5
7
(6
3
7
4)
:
1
7
3

6.
d
oi:
1
0.
1
0
3
8/
3
5
7
1
7
3
a
0.
P
M
ID
1
5
7
9
1
6
9.
16. ^
R
e
ar
ic
k
D,
Pr
ak
as
h
A,
M
c
S
w
e
e
ny
A,
S
h
e
p
ar
d
S
S,
F
e
d
or
ov
a
L,
F
e
d
or
ov
A
(
M
ar
ch
2
0
1
1)
. "
Cr
iti
ca
l
as
so
ci
ati
o
n
of
nc
R
N
A
wi
th
int
ro
ns
".
N
uc
lei
c
A
ci
ds
R
es
.
3
9
(6
):
2
3
5
7

6
6.
d
oi:
1
0.
1
0
9
3/
n
ar
/g
kq
1
0
8
0.
P
M
C
3
0
6
4
7
7
2.
P
M
ID
2
1
0
7
1
3
9
6.
17. ^
L
a
m
b
o
wi
tz
A
M
,
B
elf
or
t
M
(1
9
9
3)
.
"I
nt
ro
ns
as
m
o
bil
e
g
e
n
eti
c
el
e
m
e
nt
s"
.
A
n
n
u.
R
ev
.
Bi
oc
h
e
m
.
6
2:
5
8
7

6
2
2.
d
oi:
1
0.
1
1
4
6/
a
n
n
ur
ev
.bi
.6
2.
0
7
0
1
9
3.
0
0
3
1
0
3.
P
M
ID
8
3
5
2
5
9
7.
18. ^
P
e
n
ny
D,
H
o
e
p
p
n
er
M
P,
P
o
ol
e
A
M
,
Je
ff
ar
es
D
C
(N
ov
e
m
b
er
2
0
0
9)
.
"A
n
ov
er
vi
e
w
of
th
e
int
ro
ns
-
fir
st
th
e
or
y"
. J
o
ur
n
al
of
M
ol
ec
ul
ar
E
vo
lut
io
n
6
9
(5
):
5
2
7

4
0.
d
oi:
1
0.
1
0
0
7/
s0
0
2
3
9-
0
0
9-
9
2
7
9-
5.
P
M
ID
1
9
7
7
7
1
4
9.
19. ^
R
o
dr
g
u
ez
-
Tr
ell
es
F,
T
ar
r
o
R,
A
ya
la
F
J
(2
0
0
6)
.
"
O
ri
gi
ns
a
n
d
ev
ol
uti
o
n
of
sp
lic
e
os
o
m
al
int
ro
ns
".
A
n
n
u.
R
ev
.
G
e
n
et
4
0:
4
7

7
6.
d
oi:
1
0.
1
1
4
6/
a
n
n
ur
ev
.g
e
n
et
.4
0.
1
1
0
4
0
5.
0
9
0
6
2
5.
P
M
ID
1
7
0
9
4
7
3
7.
20. ^
M
o
ur
ie
r
T,
Je
ff
ar
es
D
C
(
M
ay
2
0
0
3)
.
"E
uk
ar
yo
tic
int
ro
n
lo
ss
".
S
ci
e
nc
e
3
0
0
(5
6
2
4)
:
1
3
9
3

1
3
9
3.
d
oi:
1
0.
1
1
2
6/
sc
ie
nc
e.
1
0
8
0
5
5
9.
P
M
ID
1
2
7
7
5
8
3
2.
21. ^
R
oy
S
W
,
Gi
lb
er
t
W
(
M
ar
ch
2
0
0
6)
.
"T
h
e
ev
ol
uti
o
n
of
sp
lic
e
os
o
m
al
int
ro
ns
:
p
at
te
rn
s,
p
uz
zl
es
a
n
d
pr
o
gr
es
s"
.N
at
ur
e
R
ev
ie
w
s
G
e
n
eti
cs
7
(
3)
:
2
1
1

2
1.
d
oi:
1
0.
1
0
3
8/
nr
g
1
8
0
7.
P
M
ID
1
6
4
8
5
0
2
0.
22. ^
d
e
S
o
uz
a
S
J
(J
ul
y
2
0
0
3)
.
"T
h
e
e
m
er
g
e
nc
e
of
a
sy
nt
h
eti
c
th
e
or
y
of
int
ro
n
ev
ol
uti
o
n"
.
G
e
n
eti
ca
1
1
8
(2

3)
:
1
1
7

2
1.
d
oi:
1
0.
1
0
2
3/
A:
1
0
2
4
1
9
3
3
2
3
3
9
7.
P
M
ID
1
2
8
6
8
6
0
2.
23. ^
Ly
nc
h
M
(A
pr
il
2
0
0
2)
. "
In
tr
o
n
ev
ol
uti
o
n
as
a
p
o
p
ul
ati
o
n-
g
e
n
eti
c
pr
oc
es
s"
.
Pr
oc
e
e
di
n
gs
of
th
e
N
ati
o
n
al
A
ca
d
e
m
y
of
S
ci
e
nc
es
9
9
(9
):
6
1
1
8

2
3.
d
oi:
1
0.
1
0
7
3/
p
n
as
.0
9
2
5
9
5
6
9
9.
P
M
C
1
2
2
9
1
2.
P
M
ID
1
1
9
8
3
9
0
4.
24. ^
Je
ff
ar
es
D
C,
M
o
ur
ie
r
T,
P
e
n
ny
D
(J
a
n
u
ar
y
2
0
0
6)
.
"T
h
e
bi
ol
o
gy
of
int
ro
n
g
ai
n
a
n
d
lo
ss
".
Tr
e
n
ds
in
G
e
n
eti
cs
2
2
(1
):
1
6

2
2.
d
oi:
1
0.
1
0
1
6/j
.ti
g.
2
0
0
5.
1
0.
0
0
6.
P
M
ID
1
6
2
9
0
2
5
0.
25. ^
Je
ff
ar
es
D
C,
P
e
nk
et
t
C
J,
B

hl
er
J
(A
u
g
us
t
2
0
0
8)
.
"
R
a
pi
dl
y
re
g
ul
at
e
d
g
e
n
es
ar
e
int
ro
n
p
o
or
".
Tr
e
n
ds
in
G
e
n
eti
cs
2
4
(8
):
3
7
5

8.
d
oi:
1
0.
1
0
1
6/j
.ti
g.
2
0
0
8.
0
5.
0
0
6.
P
M
ID
1
8
5
8
6
3
4
8.
26. ^
C
as
till
o-
D
av
is
CI
,
M
ek
h
e
d
ov
S
L,
H
ar
tl
D
L,
K
o
o
ni
n
E
V,
K
o
n
dr
as
h
ov
F
A
(A
u
g
us
t
2
0
0
2)
.
"S
el
ec
tio
n
fo
r
sh
or
t
int
ro
ns
in
hi
g
hl
y
ex
pr
es
se
d
g
e
n
es
".
N
at
ur
e
G
e
n
eti
cs
3
1
(4
):
4
1
5

8.
d
oi:
1
0.
1
0
3
8/
n
g
9
4
0.
P
M
ID
1
2
1
3
4
1
5
0.
[edit]Ext
ernal
links

Look up intron in
Wiktionary, the free
dictionary.
A
search
engine
for
exon/i
ntron
seque
nces
define
d by
NCBI
Bruce
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s,
Alexa
nder
Johns
on,
Julian
Lewis,
Martin
Raff,
Keith
Robert
s, and
Peter
Walter
Molec
ular
Biolog
y of
the
Cell,
2007,
ISBN
978-0-
8153-
4105-
5.
Fourth
edition
is
availa
ble
online
throug
h the
NCBI
Books
helf: li
nk
Jerem
y M
Berg,
John L
Tymoc
zko,
and
Lubert
Stryer,
Bioch
emistr
y 5th
edition
,
2002,
W H
Freem
an.
Availa
ble
online
throug
h the
NCBI
Books
helf: li
nk
Intron
finding
tool
for
plant
geno
mic
seque
nces
Exon-
intron
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c
maker
[hide]
V

T

E
Post-transcriptional modification


Nuclear
Precursor mRNA 5' cap formation Polyadenylation (CPSF, CstF, PAP, PAB2, CFI, CFII) Poly(A)-binding protein
RNA splicing: intron/exon snRNP spliceosome (minor spliceosome, U1) alternative splicing pre-mRNA processing
factor (PLRG1, PRPF3, PRPF4, PRPF4B, PRPF6, PRPF8, PRPF18, PRPF19, PRPF31, PRPF38A,PRPF38B, PRPF39, PRPF40A, PRPF40B)
RNA editing Polyuridylation


Cytosolic
5' cap methylation
Messenger RNA decapping: DCP1A DCP1B DCP2 DCPS EDC3 EDC4

see also disorders of transcription and post transcriptional modification
B bsyn: dna (repl, cycl, reco, repr) tscr (fact, tcrg, nucl, rnat, rept, ptts) tltn (risu, pttl, nexn) dnab, rnab/runp stru (domn, 1, 2, 3, 4)
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