Hum Genet (1985) 69:44-46
© Springer-Verlag 1985
Linkage and recombination
between fragile X-linked mental retardation and the factor IX gene
Stephen T. Warren 1, Thomas W. Glover 2, Richard L. Davidson 1, and Pudur Jagadeeswaran a
1Center for Genetics, University of Illinois College of Medicine, Chicago, IL 60612, USA
2The Genetics Center of the Southwest Biomedical Research Institute, Tempe, AZ 85281, USA
Summary. Linkage analysis on a family with fragile X-linked
mental retardation was performed using a Taq 1 restriction
fragment length polymorphism detected by a cloned human
coagulation factor IX eDNA. Two affected brothers in this
sibship were found to have different factor IX RFLP alleles,
indicating a recombinational event occurred between the two
genes. Our data therefore indicate that the gene responsible
for fragile X-linked mental retardation is not as tightly linked
to the factor IX gene as the previously published data may
suggest.
Introduction
A major cause of genetically determined mental retardation is
fragile X-linked mental retardation [fra(X)-linked MR]. This
condition results in moderate to profound mental retardation
in hemizygous males and is associated with a fragile site near
the vicinity of band Xq28 (Jacobs et al. 1980; Turner and
Jacobs 1983). Since carrier detection and prenatal diagnosis of
this condition are associated with technical and penetrance
problems (Glover 1983; Sherman et al. 1984), close linkage of
a marker gene to fra(X)-linked MR would be of considerable
benefit.
The human coagulation factor IX gene recently has been
cloned and mapped to the region from Xq26 to Xq28 by
several laboratories, including our own, and a common Taq I
restriction fragment length polymorphism (RFLP) has been
identified (Kurachi and Davie 1982; Choo et al. 1982; Jaye et
al. 1983; Camerino et al. 1984; Jagadeeswaran et al. 1984).
Camerino et al. (1983) recently reported linkage between the
fra(X)-linked MR gene and the factor IX Taq I RFLP. Out of
17 informative meioses in two families, recombination be-
tween these genes was not observed, resulting in an estimate
of the genetic distance as less than 12 centimorgans (cM).
However, in the absence of recombination, it was not clear
how tightly linked these genes are.
We have carried out linkage analyses with a factor IX
eDNA probe in families with fra(X)-linked MR and have
identified a recombinant individual. Thus, our data indicate
that the fra(X)-linked MR and factor IX genes are not as close
as the data of Camerino et al. (1983) might suggest.
Offprint requests to: P. Jagadeeswaran, Center for Genetics, Univer-
sity of Illinois College of Medicine, Chicago, IL 60612, USA
Materials and methods
Cytogenetic analyses. Heparinized blood was cultured by con-
ventional whole blood microculture techniques for 96h. To
induce expression of the fra(X) site, cells were grown in either
RPMI 1640 medium with 5% fetal bovine serum (FBS) but
without folate (Irvine Scientific), or in folate containing RPMI
1640 medium with 10% FBS to which 0.1uM 5-fluorodeoxy-
uridine (FUdR) was added at the beginning of the culture
period (Glover 1981).
Epstein-Barr virus (EBV) transformed lymphoblastoid cell
lines were established from some family members by the
method of Jacobs et al. (1982). For fra(X) expression, lym-
phoblastoid cells were cultured in RPMI 1640 medium with
10% FBS. FUdR (0.1uM) was added 24h prior to chromo-
some harvest. For all cultures, air dried slides were stained
with Giemsa or Q-banded and at least 50 cells scored for fra-
gile X expression.
DNA isolation and Southern blot analyses. High molecular
weight DNA was isolated from either lymphocytes or lympho-
blastoid cells and subjected to Southern blot analyses (South-
ern 1975), using as probe a 1.0kb Hpa II fragment, containing
the factor IX eDNA insert (Jagadeeswaran et al. 1984).
Lod score calculations. The relative probability of linkage and
lod scores were calculated at various recombination intervals
from 0 to 50% (Morton 1955; Connea!ly and Rivas 1980).
Results and discussion
Figure 1 shows the pedigree of the family described in this
report. This family, of English/Irish descent, was ascertained
through individuals III-2 and III-3, who were diagnosed as
having fra(X)-linked MR on the basis of profound psycho-
motor retardation and expression of the fragile site at Xq28.
The fragile site was expressed, with FUdR induction, in both
lymphocytes and lymphoblastoid cells from both individuals
(III-2, 22-25% fra(X) expression; III-3, 14-23% fra(X)
expression). A phenotypically normal brother, III-4, did not
express the fra(X) in 100 lymphocyte metaphases examined.
Two female siblings were mentally normal. However, one
sister (III-5) was determined to be a carrier of fra(X)-linked
MR on the basis of expression of the fra(X) site (10-27%
fra(X) expression in lymphocytes and lymphoblastoid cells).