Xing-Wang Deng
xingwang.deng@
yale.edu
Wolfgang Dubiel
Ning Wei, Kay Hofmann
Kirsten Mundt
John Colicelli
Jun-ya Kato
Michael Naumann
Daniel Segal
Michael Seeger
Michael Glickman
Daniel A. Chamovitz
Anthony Carr
and if the transposase displaces polymer subunits that are
nearby on the DNA, transposition will occur most fre-
quently at distant sites. Daniel Voytas (Iowa State
University, Ames, USA) reported that the Sir 3 and 4
genes, involved in silencing at the telomeres, direct yeast
Ty5 to silent chromatin. And Nancy Craig (Howard
Hughes Medical Institute/Johns Hopkins University,
Baltimore, USA) reported that the TnsE pathway of Tn7 is
biased to insert near double strand breaks and at replication
termination sites. We imagine this is part of a strategy for
Tn7 to get itself transferred onto a conjugating plasmid.
Several speakers discussed host suppression of transpos-
ition. Among them, Ronald Plasterk (Netherlands Cancer
Institute, Amsterdam, The Netherlands) spoke on suppres-
sion of transposition in the germ-line of Bristol strains of
Caenorhabditis elegans. Surprisingly, the suppression path-
way overlaps with the RNAi pathway, a mysterious
process by which exogenous double-stranded RNA blocks
expression of the proteins it encodes. Silke Jensen (Institut
Gustave Roussy, Villejuif, France) reported on suppression
of I elements in Drosophila. Interestingly, transgenes con-
taining fragments of I, whether in the sense or in the anti-
sense direction, suppress invasion by a new I element. Joan
Curcio (Wadsworth Center, Albany, USA) reported on
inhibition of both Ty transposition and invasive growth by
the yeast protein Fus3.
Zooming in from evolution to mechanism, several ses-
sions focused on the assembly of protein–DNA com-
plexes. Assembly is coupled to activation, to ensure that
recombination occurs at the proper time and place and on
the proper substrates. The crystal structure of Cre bound
to a Lox Holliday junction, presented by Greg Van Duyne
(University of Pennsylvania School of Medicine,
Philadelphia, USA), explained how, of the four DNA mol-
ecules in a Holliday junction, only two at a time are acti-
vated to be cleaved. Phoebe Rice (University of Chicago,
USA) presented the structure of Flp recombinase, confirm-
ing biochemical predictions that its catalytic tyrosine,
unlike that of closely related Cre, is donated in trans; the
tyrosine reaches into the active site of the neighboring sub-
unit, to cleave the DNA bound by the neighbor. Similarly,
William Reznikoff (University of Wisconsin, Madison,
USA) reported that Tn5 transposase donates catalytic
acidic residues (the DDE motif) in trans. This mechanism
helps ensure that two distant DNA sites move together.
Transposons have evolved means to recognize them-
selves, to find a good insertion site, and to coordinate the
cutting and pasting of their two ends: a complex series of
tasks. And yet they must work with the tools they carry on
their backs and the proteins they co-opt from their hosts.
These elements are streamlined, and we see in them the
beauty both of complexity and of simplicity.
Outlook MEETING REPORTS Non-homologous recombination
TIG May 2000, volume 16, No. 5
202
T
he COP9 signalosome was first identified in
Arabidopsis thaliana as an essential regulator of light
signal transduction
1
. Subsequently, the COP9 signalosome
was identified in animal systems, suggesting that this com-
plex had a general role in developmental regulation
2,3
.
Genetic analysis in Drosophila indicated that this complex
is essential for animal development
4
. This role might
include the control of the cell cycle and regulation of
MAP-kinase signaling
3,5–10
. The biochemical purification
of the COP9 signalosome from cauliflower
11
, mammals
9,16
and Arabidopsis
12
indicated that this complex comprises
eight core subunits. The complex does not exist in
Saccharomyces cerevisiae but appears to be present in
Schizosaccharomyces pombe
5
. The COP9 signalosome is
similar, both in size and composition, to two other regula-
tory complexes: the lid of the 19S proteasome regulatory
particle and the eukaryotic translational regulatory com-
plex eIF3 (Refs 13, 14). Most subunits from all three
complexes contain one of two structural motifs: the
PCI/PINT domain and the MPN domain
14,15
. Reports have
suggested interactions between these complexes and/or
their subunits
3
.
Because the genes encoding these subunits were iso-
lated either through a variety of unrelated genetic screens,
or by the biochemical purification of the complex from
various organisms, the original names for the COP9 sig-
nalosome subunits are largely unrelated; in most cases,
the names do not imply that the protein product is a
COP9 signalosome subunit. The fact that many of the
subunits have several names adds to the confusion. In
order to clarify this situation, a unified subunit nomencla-
ture was agreed upon that conveys the fact that these pro-
teins function cooperatively within a specific complex,
and that will be applicable for all organisms. The sub-
units from human, mouse and Drosophila have previ-
ously been numbered in decreasing size according to their
Unified nomenclature for the COP9
signalosome and its subunits: an
essential regulator of development
Outlook LETTERS
0168-9525/00/$ – see front matter © 2000 Elsevier Science Ltd. All rights reserved. PII: S0168-9525(00)01982-X