Development 112, 207-219 (1991)
Printed in Great Britain © The Company of Biologists Limited 1991
207
A chicken caudal homologue, CHox-cad, is expressed in the epiblast with
posterior localization and in the early endodermal lineage
AYALA FRUMKIN
1
, ZEHAVA RANGINI
2
, ADI BEN-YEHUDA
1
, YOSEF GRUENBAUM
2
and
ABRAHAM FAINSOD
1
*
1
Department of Cellular Biochemistry, Hebrew Umversity-Hadassah Medical School, Em Kerem, Jerusalem 91010, Israel
2Department of Genetics, Hebrew University, Gtvat Ram, Jerusalem 91904, Israel
* Author for correspondence
Summary
CHox-cad is a chicken homeobox gene whose homeo-
domain is homologous to the Drosophila caudal and the
murine Cdxl genes. Based on sequence analysis of a
2.5 kb CHox-cad cDNA clone, we deduced that the
primary translation product consists of 248 ami no acids.
Comparison between the cDNA and genomic clones
revealed the presence of an intron within the CHox-cad
homeodomain between amino acids 44 and 45. The onset
of CHox-cad transcription correlates temporarily with
the beginning of gastrulation. During primitive streak
stages CHox-cad exhibits a caudally localized pattern of
expression restricted to the epiblast and the primitive
streak. At these stages, CHox-cad transcripts can also be
detected in the definitive endoderm cells. Later in
embryogenesis CHox-cad is expressed in the epithelial
lining of the embryonic gut and yolk sac. After four days
of chicken development, no CHox-cad transcripts could
be detected. The early CHox-cad posterior expression in
the germ layer undergoing gastrulation and its continu-
ous expression in the early endodermal lineage raise the
possibility of CHox-cad involvement in the establishment
of the definitive endoderm.
Key words: homeobox, caudal, chicken embryogenesis,
cloning, in situ hybridization, endoderm, CHox-cad, cell
lineage.
Introduction
Numerous homeobox-containing genes (homeobox
genes) have been cloned from vertebrate genomes and,
in several instances, the same or closely related genes
have been cloned from different organisms (Scott et al.
1989; Graham et al. 1989; Duboule and Dolle", 1989).
The interest in this vertebrate gene family arose as a
result of the discovery of the homeobox sequence in
Drosophila developmental genes (see Gehring, 1987a,/?
for reviews), which was subsequently shown to have
been conserved during evolution, in animals including
vertebrates (McGinnis et al. 1984). The large number of
vertebrate homeobox genes cloned to date are found in
one of four major Hox clusters which, in mice and
humans, contain 6-10 homeobox genes (Acampora et
al. 1989; Graham et al. 1989; Duboule and Dolle\ 1989;
Nj0lstad et al 1990; Fritz et al. 1989; Wedden et al.
1989). The organization of vertebrate homeobox genes
in clusters is reminiscent of Drosophila homeobox gene
organization. Comparison of sequence and genomic
organization of the clusters in mouse and man led to the
suggestion that the vertebrate Hox clusters arose as a
result of duplication of an ancestral gene complex (Hart
et al. 1987; Kappen et al. 1989; Schughart et al. 1989).
The similarity between the Drosophila and vertebrate
homeobox gene clusters extends beyond the arrange-
ment of homeobox sequences in a gene complex.
Comparison of vertebrate genes in the clusters to the
Drosophila genes in the Antennapedia and Bithorax
complexes revealed a colinear arrangement of cognate
genes (Acampora et al. 1989; Duboule and Doll6, 1989;
Graham et al. 1989). Furthermore, as in Drosophila,
the order of the genes along the cluster represents the
anteroposterior order of their rostral boundary of
expression (Gaunt et al. 1988; Graham et al. 1989;
Duboule and Dolle", 1989).
The high degree of homology between the vertebrate
homeobox sequences and their Drosophila counter-
parts has resulted in the organization of the genes into
families based on their homeodomain sequences (Scott
et al. 1989). In some instances, a homeobox gene family
contains several sequences from the same organism due
to the duplication of the Hox complexes. The homology
between cognate genes is not restricted to the homeo-
box region but it extends throughout the genes