First published online March 24, 2005
doi: 10.1242/10.1242/dev.01775
Development 132, 1923-1934 (2005)
Published by The Company of Biologists 2005
Analysis of xbx genes in C. elegans
Evgeni Efimenko1,
Kerry Bubb2,
Ho Yi Mak4,
Ted Holzman3,
Michel R. Leroux5,
Gary Ruvkun4,
James H. Thomas2 and
Peter Swoboda1,*
1 Karolinska Institute, Department of Biosciences and Södertörn
University College, Section of Natural Sciences, S-14189 Huddinge,
Sweden
2 Department of Genome Sciences, University of Washington, Seattle, WA 98195,
USA
3 Department of Microbiology, University of Washington, Seattle, WA 98195,
USA
4 Department of Molecular Biology, Massachusetts General Hospital and Department
of Genetics, Harvard Medical School, Boston, MA 02114, USA
5 Department of Molecular Biology and Biochemistry, Simon Fraser University,
Burnaby, British Columbia V5A 1S6, Canada

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Fig. 1. Computational search for the X-box motif. (A) Schematic of the X-box search
algorithm. The algorithm finds all matches for a defined motif consensus and
cross-matches them against a list of predicted genes or available ESTs. The
search space upstream of predicted genes was set to 1000 bp. (B) The number of
matches obtained with different consensus sequences (top to bottom: relaxed,
average, refined) searching the C. elegans genome. (C) The `relaxed'
X-box consensus (RYYNYY WW RRNRAC), used for initial searches, generates the
largest number of matches that spread equally within promoter regions. (D) The
`refined' X-box consensus (GTHNYY AT RRNAAC), obtained on the basis of in vivo
expression analysis of xbx genes, shows a significant concentration
of matches in the region of around 100 bp upstream of the ATG.
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Fig. 2. Functional repertoire of xbx genes. Functional repertoire of
xbx genes and some representative members from different molecular
groups, including possible components of cilium structure and transport
machinery, transcription factors and receptors. The diagram represents data
from the `average' consensus list of xbx genes/candidates (758
members) (see Table S1 in supplementary material), including experimentally
proven genes.
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Fig. 3. Newly discovered X-box genes. (A) xbx-2::gfp is expressed in most
or all CSN. (B) xbx-3::gfp is expressed in amphid and phasmid (not
shown) neurons. (C) xbx-4::gfp is expressed in a subset of amphid
neurons. (D) xbx-5::gfp is expressed in a subset of amphid neurons.
(E) xbx-6::gfp is expressed in many different types of cells,
including the phasmid neurons PHA and PHB. (F) xbx-7::gfp is
expressed in a subset of amphid and interlabial neurons. (G)
tub-1::gfp is expressed in most or all CSN. (H) Schematic diagram of
CSN positioning in the C. elegans hermaphrodite (after
Collet et al., 1998 ).
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Fig. 4. An example of retrograde movement for XBX-2::GFP particles in a phasmid
cilium (see also Movie 1 in supplementary material). The ciliary transition
zone is marked with an asterisk. The arrowhead indicates the initial position
of the moving particle at time zero (t=0). The arrow indicates the position of
the moving particle at different time points (t=0.5 and 1 seconds). Scale
bars: 1 µm.
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Fig. 5. Properties of the X-box motif in C. elegans. (A) The X-box
consensus sequence obtained from in vivo expression analysis. Nucleotides
marked in red are strongly conserved and important for the proper function of
the motif. (B) Proposed scheme for the difference in expression patterns
observed for xbx genes. Depending on the nucleotide composition of
the X-box, DAF-19 can bind to the motif either in homodimer (driving
expression in many, most or all CSN) or in heterodimer form, interacting
together with some cell-specific factor (driving expression in a subset of
CSN).
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Fig. 6. Development of a `ciliary module' in C. elegans. DAF-19 regulates
the development of the module, which includes genes for the cilium structure
and transport machinery, receptors and other factors. The activation of this
module leads to the formation of functional ciliated endings during
specification of sensory neurons in the worm.
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© The Company of Biologists Ltd 2005