Lewin Benjamin (ed.) Genes IX
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they are located
and
often cause
chromosomal
breaks.
The result
of the transpositions
is
there-
fore to inactivate
the
genome.
P Elements
Are
Activated
in
the GermLine
o
P
elements are activated in
the
germtine
of
P
male x M femate
crosses
because
a tissue-
specific
spticing event removes
one intron,
which
generates
the
coding sequence for
the transoosase.
.
The P
etement also
produces
a repressor
of
transposition, which is inherited
maternatty in
the
cytoptasm.
e
The
presence
of the
repressor
exptains why M male
x
P femate crosses remain
ferti[e.
Activation
of P
elements is
tissue-specific: it
occurs only in
the
germline.
P
elements are tran-
scribed,
though, in both
germline
and somatic
tissues. Tissue-specificity
is
conferred by a
change in
the splicing
pattern.
riiii:
riii:
,
: ; I
depicts the
organization of the
element
and
its
transcripts.
The
primary
tran-
script extends for 2.5 kb
or 3.0 kb, the
difference
probably
reflecting
merely
the leakiness of the
termination site. TWo
protein
products
can be
produced:
.
In somatic tissues,
only the first
two
introns are
excised, creating
a coding
region
of ORF0-ORFI
-ORF2.
Transla-
tion of this RNA yields
a
protein
of
66 kD. This
protein
is a repressor
o{
transposon activity.
.
In
germline
tissues, an
additional splic-
ing
event occurs
to remove intron 3.
This connects
all four
open
reading
frames
into an mRNA
that is translated
to
generate
a
protein
of 87 kD. This
pro-
tein
is
the transposase.
TWo types of experiment
have demonstrated
that
splicing of the third intron is needed for
transposition. First, if
the splicing
junctions
are
mutated
in vitro and the P
element is reintro-
duced into flies, its transposition
activity is abol-
ished.
Second,
if
the third intron is
deleted, so
that ORF3 is constitutively
included in the
mRNA in
all tissues, transposition
occurs
in
somatic tissues as well as the
germline.
Thus
whenever ORFI is
spliced to the
preceding
read-
ing frame, the P element
becomes active. This
is the crucial regulatory event,
and usually
it
occurs only in the
germline.
i,iiii.i{'11
.'
1,,:rrr The P etement
has
four
exons.
The first
three are spliced
together
in somatic expression;
at[ four
are sp[iced together
in
germtine
expression.
What
is responsible
for the tissue-specific
splicing? Somatic cells
contain
a
protein
that
binds to sequences
in exon
3 to
prevent
splic-
ing of the last
intron
(see
Section
26.I2, AIter'
native Splicing Involves
Differential
Use of Splice
Junctions).
The absence
of
this
protein
in
germline
cells
allows splicing
to
generate
the
mRNA that codes
for the transposase.
Ttansposition
of a
P element
requires
-I50
bp
of
terminal DNA.
The transposase
binds
to
l0
bp sequences
that
are adjacent
to the
3l bp
inverted repeats.
Transposition
occurs
by a non-
replicative
"cut
and
paste" mechanism
resem-
bling that of
Tnl0.
(It
contributes
to hybrid
dysgenesis
in two ways:
Insertion
of the
trans-
posed
element
at a
new site
may cause
muta-
tions, and
the break
that
is left at the
donor
site-see
Figure 21 .7-h'as
a deleterious
effect.)
It is interesting
that,
in a significant
propor-
tion of cases,
the break
in donor
DNA
is repaired
by using the sequence
of
the homologous
chro-
mosome. If the
homolog
has a P element,
the
presence
of
a P element
at
the donor site
may
be restored
(so
the event
resembles
the
result
of a replicative
transposition).
If the
homolog
lacks
a
P element,
repair
may
generate
a se-
quence
lacking the
P element,
thus apparently
I
I Transcription
V
Short RNA
.e*
Long RNA
irr
lntron
1 Intron 2 Intron 3
I
66
kD repressor
Germline
t.+-+:;5!igqFffi
mRNA
-
[ts2iing
frame 4'
I
T
Y
87
kD transposase
21..1,5
P Elements
Are Activated
in the Germline
545
providing
a
precise
excision
(an
unusual event
in
other transposable systems).
The dependence
of
hybrid
dysgenesis on
the
sexual orientation of a cross shows
that the
cytoplasm is important, as
well
as the
P factors
themselves. The contribution
of the cytoplasm
is described as the
cytotype; a line of flies con-
taining P
elements
has
P cytotype, whereas
a
line
of flies lacking P elements has
M cytotype.
Hybrid
dysgenesis occurs only when
chromo-
somes containing P factors find
themselves in
M cytotype, that is, when
the male
parent
has
P elements
and the female
parent
does not.
Cytotype shows an inheritable
cytoplasmic
e{fect; when a
cross occurs through P cytotype
(the
female
parent
has P elements),
hybrid dys-
genesis
is suppressed for
several
generations
of
crosses with M female
parents.
Thus something
in P cytotype,
which can be diluted
out over
some
generations,
suppresses hybrid
dysgenesis.
The effect
of cytotlpe is explained in molec-
ular
terms by the model
of
ii{,i.ii:ii: .:i"-:il.
It
depends on the ability
o{ the 66 kD
protein
to
repress transposition.
The
protein
is
provided
::iliriltii
;i i.:;i;
Hybrid
dysgenesis is
determined
by the interactions
between
P
etements in
the
genome
and
66 kD
repressor
in
the cytotype.
CHAPTER
2L Transposons
as a
maternal factor in the egg. In
a
P line, there
must
be sufficient
protein
to
prevent
transpo-
sition from occurring, even though the P ele-
ments are
present.
In any
cross
involving
a
P female, its
presence prevents
either synthesis
or activity of the transposase. When the female
parent
is M type, though, there is no repressor
in the
egg, and the
introduction
of a
P
element
from the male
parent
results in activity of trans-
posase
in
the
germline.
The
ability of P cytotlpe
to exert an effect through more than
one
gen-
eration
suggests
that there must
be enough
repressor
protein
in the egg, and it must be sta-
ble enough, to be
passed
on through
the adult
to be
present
in
the eggs of the next
generation.
Strains of. D. melanogaster descended from
flies
caught
in the
wild
more
than 30
years
ago
are always M. Strains descended from flies caught
in the
past
l0
years
are almost always P. Does
this
mean that the P element family
has invaded
wild
populations
of D. melanogaster in recent
years?
P
elements are
indeed
highly invasive
when introduced into a new
population;
the
source of the invading element
would have to
be another species.
Hybrid dysgenesis reduces interbreeding,
and
thus
it is
a step on the
path
to speciation.
Suppose that a dysgenic
system is created by a
transposable
element
in
some
geographic
Ioca-
tion. Another element may
create a different
system in
some other
location.
Flies in
the two
areas will be dysgenic for two
(or
possibly
more)
systems. If this renders
them intersterile and
the
populations
become
genetically
isolated,
further separation may
occur. Multiple dysgenic
systems
therefore
lead
to inability to mate-
and to speciation.
Sum
mary
Prokaryotic
and eukaryotic cells
contain a vari-
ety of transposons that mobilize
by moving
or
copying DNA
sequences. The transposon
can
be
identified
only as an
entity within
the
genome;
its
mobility does not involve
an inde-
pendent
form. The transposon
could
be selfish
DNA,
concerned only
with
perpetuating
itself
within
the resident
genome;
if
it conveys
any
selective advantage
upon the
genome,
this
must be indirect.
All transposons
have
systems
to limit the
extent of transposition,
because
unbridled
transposition is
presumably
damag-
ing,
but the molecular
mechanisms
are differ-
ent in each
case.
The archetypal
transposon
has inverted
repeats at its
termini and
generates
direct repeats
of a short sequence
at the
site of insertion.
The
simplest
types are the
bacterial
insertion
sequences
(IS),
which
consist
essentially
of the
inverted
terminal repeats
flanking
a coding
frame(s)
whose
product(s)
provide
transposi-
tion
activity. Composite
transposons
have ter-
minal modules that
consist
of
IS
elements;
one
or both
of the
IS
modules
provides
transposase
activity, and the sequences
between
them
(often
carrying
antibiotic resistance)
are treated
as
pas-
sengers.
The
generation
of target
repeats flanking
a
transposon reflects
a common feature
of trans-
position.
The target
site is cleaved
at
points
that
are staggered on each DNA
strand
by a fixed
distance
(often
five
or nine
base
pairs).
The
transposon
is in effect inserted
between
pro-
truding single-stranded
ends
generated
by the
staggered cuts. Thrget
repeats
are
generated
by
filling in
the single-stranded
regions.
IS elements,
composite
transposons, and
P elements mobilize
by
nonreplicative
transposi-
tion, in
which
the
element moves
directly from a
donor
site
to a recipient
site. A
single transposase
enzyme undertakes the reaction.
It occurs
by a
"cut
and
paste"
mechanism
in which
the transpo-
son is separated from flanking
DNA.
Cleavage of
the transposon
ends, nicking
of the target site,
and connection of the transposon
ends to the
stag-
gered
nicks all occur in
a
nucleoprotein
complex
containing the transposase.
Loss
of the transpo-
son
from
the donor creates a
double-strand break
whose
fate is not
clear. In the case
of
Tn I0,
trans-
position
becomes
possible
immediately
after DNA
replication, when sites recognized
by t}i'e dam
methylation system
are transiently hemimeth-
ylated.
This imposes
a demand for
the existence
of two copies of the donor
site, which may enhance
the cell's chances for survival.
The TnA family
of transposons mobilize
by
replicative
transposition. After
the transposon
at the donor site becomes connected
to the tar-
get
site, replication
generates
a cointegrate mol-
ecule that
has
two copies of the
transposon. A
resolution reaction
that involves recombina-
tion between two
particular
sites then frees
the
two
copies of
the
transposon, so that one
remains
at the donor site and one appears
at
the target site. TWo enzymes coded
by the trans-
poson
are
required: Ttansposase
recognizes the
ends of the
transposon
and connects them to
the target site, and resolvase
provides
a site-
specific recombination function.
Phage Mu
undergoes replicative transpo-
sition by the same mechanism as TnA. It also can
use
its
cointegrate
intermediate
to transpose by
a
nonreplicative mechanism. The difference
between this reaction and the
nonreplicative
transposition
of
IS
elements
is that the cleavage
events occur in a different order.
The
best characterized
transposons
in
plants
are the controlling elements
of maize, which
fall into
several
families. Each
family contains
a
single type of autonomous
element
that is
analogous to bacterial transposons
in
its
ability
to mobilize. A family also contains
many dif-
ferent
nonautonomous
elements that
are
derived
by
mutations
(usually
deletions) of the
autonomous element.
The nonautonomous
ele-
ments lack the ability to
transpose, but display
transposition
activity
and other
abilities of the
autonomous element when
an autonomous
element is
present
to
provide
the necessary
tr ans
-
acting
f
unctions.
In addition to the
direct consequences
of
insertion and excision, the
maize elements
may
also control the activities
of
genes
at or near the
sites where
they
are inserted;
this control
may
be subject to developmental
regulation. Maize
elements
inserted into
genes
may be excised
from the transcripts,
which explains
why they
do not
simply
impede
gene
activity. Control
of
target
gene
expression
involves
a variety of
molecular
effects,
including
activation by
pro-
vision of an enhancer
and suppression
by inter-
ference with
posttranscriptional events.
Tiansposition of
maize elements
(in partic-
ular Ac)
is
nonreplicative,
and
probably
requires
only a
single
transposase
enzyme coded
by the
element.
Transposition occurs
preferentially
after replication of
the element.
It is
likely
that
there are mechanisms
to limit
the frequency of
transposition. Advantageous
rearrangements
of the
maize
genome
may
have been connected
with the
presence
of the
elements.
P elements
in D. melanogaster
are
responsi-
ble for hybrid dysgenesis,
which
could be a
fore-
runner
of
speciation.
A cross between
a male
carrying
P
elements
and
a female
lacking them
generates
hybrids
that are sterile.
A P element
has four open
reading
frames, which
are sepa-
rated
by introns. Splicing
of
the first three
ORFs
generates
a
66 kD repressor
and occurs
in all
cells. Splicing of
all four
ORFs to
generate
the
87 kD transposase
occurs only
in the
germline
by a tissue-specific
splicing
event.
P elements
mobilize when exposed
to cytoplasm
lacking
the
repressor. The burst
of transposition
events
inactivates the
genome
by
random
insertions.
Only a complete P element
can
generate
trans-
posase,
but
defective elements
can
be mobilized
intrans bv the enzvme.
21.16
Summary
547
References
@
Introduction
Reviews
Campbell, A.
(
I98l
).
Evolutionary
significance of
accessory DNA
elements
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35,
55-83
Finnegan, D.
J.
(1985).
Transposable
elements
in
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Cytol
9), 281-)26.
@
Insenron
sequences
nre
srmpre
Transoosition
Modules
Reviews
Berg, D. E. and Howe, M.,
eds.
(I989).
Mobile
DNA
Washington, DC: American
Society for Micro-
biology
Press.
Calos, M. and Miller, J. H.
(1980).
Transposable
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Cell
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Craig, N. L.
(1997).
Target
site selection
in
transpo-
silio}l. Annu Rey. Biochem.
66,
437 47 4.
Galas, D. J. and
Chandlea
M.
(1989).
Bacterial inser-
tion sequence. In Berg, D. E
and Howe, M.,
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109-I62.
I(leckner,
N.( 1977). Ttanslocatable
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Cell
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I(leckner,
N.
(1981).
Transposable
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Annu Rev.
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Resea
rc h
Grindley, N. D.
(1978).ISl
insertion
generates
duplication of a 9 bp sequence
at
its
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Cell
13,
419-426.
Johnsrud, L
,
Calos,
M.
P., and Miller,
J.
H.
(1978t.
The
transposon Tn9
generates
a
9
bp
repeated
sequence during integration.
Cell 15,
t209-12r9.
t@
Composite Transposons
Have IS Modutes
KCVIEW
I(eckner,
N.
(1989).
Tnl0 transposon. In Berg,
D. E.
and Howe, M. M., eds., Mobile
DN,4. Washing-
ton, DC: American
Society for Microbiology
Press,
pp.
227-268.
Transposition
0ccurs
by
Both
Repticative
and
Nonrepticative
Mechanisms
Craig, N. L.
(19971.
Target site
selection in transpo-
silion
Annu Rev. Biochem
66,4)7-474.
Grindley,
N.
D.
and Reed, R. R.
(1985).
Transposi-
tional recombination
in
prokaryotes
Annu.
Rev. Biochem
54, 86)-896.
Haren,
L., Ton-Hoang,
B., and
Chandler, M.
(19991.
Integrating
DNA:
transposases and
retroviral
integrases.
Annu Rev. Microbiol.
5),
245-28t.
Scott, J. R. and
Churchward, G.
G.
(1995).
Con-
jugative
transposition. Annw.
Rev. Immunol 49,
367-J97.
CHAPTER 21 Transoosons
@
Reviews
Common
Intermediates for Transposition
Reviews
Mizuuchi, K.
(1992).
Transpositional
recombina-
tion: mechanistic insights from studies
of
Mu
and other elements.
Annu. Rev.
Biochem. 61.
l0I l-I051.
Pato,
M. L.
(
1989). Bacteriophage
mu. In Berg, D. E.
and
Howe, M.,
eds.,
Mobile DNA.
Washington,
DC:
American
Society
for Microbiology
Press,
pp.23-52.
Resea rch
Aldaz, H., Schuster, E., and Baker, T. A.
(1996).
The interwoven architecture
of the
Mu
trans-
posase
couples DNA synthesis
to catalysis. Cel/
85,257-269.
Savilahti,
H.
and
Mizuuchi, K.
(1996).
Mu trans-
positional
recombination: donor DNA
cleav-
age and strand transfer in trans
by the Mu
transpose. Cell 85, 27 l-280.
Nonrepticative Transposition
Proceeds
by Breakage and Reunion
Resea rch
Bender,
J. and
I{eckner,
N.
(I986).
Genetic
evi-
dence that
Tnl0
transposes by a nonreplica-
tive
mechanism
Cell
45,801-815.
Bolland, S. and Ifleckner, N.
(1996).
The
three
chemical steps of Tn I 0/IS I 0 transposition
involve repeated utilization
of a single active
site.
cell
84,223-2)3.
Davies, D. R.,
Goryshin,
L Y., Reznikoff,
W.
S.,
and Rayment, I.
(2000).
Three-dimensional
structure of the Tn5 synaptic
complex trans-
position
intermediate.
Science
289,
77
-85.
Haniford, D. B., Benjamin,
H. W, and I(leckner,
N.
(
I99 I
)
. I(netic and structural
analysis of a
cleaved donor intermediate
and a strand
transfer
intermediate
in Tnl0
transposition.
Cell 64, 17l-179.
I(ennedy,
A. I(., Guhathakurta, A., I(leckner,
N.,
and Haniford, D. B.
(1998).
Tnl0
transposi-
tion
via
a DNA hairpin intermediate.
Cell 95,
125-tj4.
TnA Transposition
Requires Transposase
and Resotvase
Review
Sherratt, D.
(1989).
Tn3
and
related
transposabie
elements:
site-specific recombination
and
transposition.
In Berg, D. E.
and Howe, M.
M.,
eds., Mobile DN,4.
Washington,
DC: American
Society for Microbiology
Press,
pp.
163-l
84.
Resea rc h
Droge, P. et al.
(1990).
The two functional
domains
of
gamma
delta resolvase
act on the
same
recombination
site: implications for
the mech-
anism
of strand exchange.
Proc. Natl Acad.
Sci.
usA 87, 5336-5340.
548
619
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