[CANCER RESEARCH 56. 1241-1243. March 15. I996|
Advances in Brief
Germline and Somatic Mutations in an Oncogene: RET Mutations in Inherited
Medullary Thyroid Carcinoma1
Debbie J. Marsh, Scott D. Andrew, Charis Eng, Diana L. Learoyd, Amanda G. Capes, Ruth Pojer,
Ann-Louise Richardson, Carol Houghton, Lois M. Mulligan, Bruce A. J. Ponder, and Bruce G. Robinson2
Koi/ing Institute of Medical Research ¡D.J. M.. S. D. A.. D. L L, A. C. C. R. P.. A-L R.. B. G. R.¡and Department of Endocrinology [D. L. L. B. G. R.¡,Royal North Shore
Hospital, St Leonards. N.S.W. 2065, Australia; University of Sydney, Sydney, N.S.W., Australia ¡D.J. M.. D. L. L, A. G. C., B. G. R.¡:Department of Medicine, Dana-Farber
Cancer Institute, Harvard Medical School, Boston, Massachusetts [C. E.I; Cancer Research Campaign Human Cancer Genetics Research Group, University of Cambridge,
Addi'nbrooke's Hospital, Bo\ 23X, Cambridge CB2 2QQ, England 1C. £., C. H., B. A, J. P.]; and Departments of Paediatrics and Pathologv, Queen's University, Kingston,
Ontario. K7L 3N6 Canada ¡L.M. M.¡
Abstract
Inherited cancer syndromes predispose an individual to the develop
ment of specific tumors. Somatic and germline mutations in the same
tumor suppressor gene, as described in Knudson's two-mutation model,
are well recognized. Inherited mutations in the RET proto-oncogene,
which encodes a receptor tyrosine kinase, predispose individuals to the
multiple endocrine neoplasia type 2 (MEN 2) cancer syndromes. The
major component tumor of these syndromes is medullary thyroid carci
noma (MTC). To date, somatic mutations in RET have not been identified
in tumors from individuals with MEN 2, although they have been well
documented in sporadic MEN 2-related tumors. We have identified,
among 16 MEN 2 cases with well-defined RET germline mutations, a
somatic missense mutation at codon 918 of RET in 3 of 15 MTCs and in
a sample with hyperplastic C-cells (the presumed precursor to hereditary
MTC). We suggest that the presence of a somatic mutation, in addition to
the preexisting germline mutation in hereditary MTCs, may contribute to
tumorigenesis in vivo.
Introduction
MTC1 is a malignancy of the thyroid C-cells and is the character
istic component tumor of the inherited cancer syndromes MEN types
2A and 2B and FMTC. Germline mutations in the RET proto-onco
gene, which encodes a receptor tyrosine kinase, cause MEN 2A, MEN
2B, and FMTC (1-9). The majority of germline mutations that have
been found in MEN 2A and FMTC families lie within exons 10 and
11 of RET at codons 609, 611,618, 620, and 634 and result in the
substitution of a cysteine by an alternative amino acid (1, 2). A single
mutation at codon 918 causing the substitution of a methionine (ATG)
with a threonine (ACG) is associated with MEN 2B and sporadic
MTC (3-5, 10-12). Somatic mutations in RET codons 768 and 883
have also been described in sporadic MTC at very low frequencies
(7, 8, 13). Studies using transient transfection and other methods have
shown that RET codon 634 missense mutations result in increased
receptor dimerization independent of ligand binding, and the codon
Received 12/13/95; accepted 1/31/96.
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1This work has been supported by a number of grants and scholarships: Mary Jo Reeve
Fellowship (to D. L. L.): National Health and Medical Research Council biomedicai
scholarship (to A. G. C.); core and program grants from the Cancer Research Campaign
(CRC), CRC Dana-Farber Cancer Institute Fellowship. Lawrence and Susan Marx Inves-
tigatorship in Cancer Genetics in the Division of Cancer Epidemiology and Control,
Dana-Farber Cancer Institute, and Young Scientist Award from the Markey Charitable
Trust (to C. E.); Medical Research Council of Canada (to L. M. M.); and the CRC Gibb
Life Fellowship (to B. A. J. P.).
2 To whom requests for reprints should be addressed, at Kolling Institute of Medical
Research, Royal North Shore Hospital, St. Leonards, N.S.W. 2065. Australia. Phone: +61
2 9926 7267: Fax: +612 9926 8523.
The abbreviations used are: MTC, medullary thyroid carcinoma; MEN, multiple
endocrine neoplasia; FMTC. familial MTC; CCH. C-cell hyperplasia.
918 mutation leads to altered substrate specificity (6, 9, 14). Both
mutations increase receptor tyrosine kinase activity (6, 9).
Inherited cancer syndromes are generally caused by germline mu
tations in tumor suppressor genes (15). Activating mutations in the
RET proto-oncogene in the germline of MEN 2 and FMTC individuals
represent an exception to this rule. According to Knudson's two-
mutation model originally invoked for tumor suppressor genes (16), a
primary mutation in a tumor suppressor gene occurs in the germline,
with a secondary mutation occurring in somatic tissue within the same
gene on the other alÃ-eleleading to tumorigenesis (17, 18). It has been
thought that this double mutation is not necessary in the case of
oncogenes since a single activating mutation in RET has been shown
to transform cells (6, 9). However, although the RET germline muta
tion is obviously present from birth in an affected individual, the
factors that determine how CCH, the presumed precursor to hereditary
MTC (19, 20), or MTC develop in C-cells are yet to be elucidated. We
hypothesized that the presence of a somatic mutation, in addition to
the preexisting germline mutation in hereditary MTCs, may contribute
to tumorigenesis in vivo.
Materials and Methods
Patients. Thyroid tissue, from 16 members of MEN 2A or FMTC families,
in which germline RET mutations had been identified previously (21-23), was
studied. Patient 3. aged 11 years, was identified as having a high risk of
inheriting the MEN 2A phenotype by linkage analysis, prior to the advent of
direct mutation detection, and thyroidectomy was performed (24). CCH was
diagnosed in the thyroid specimen from this patient. All other patients were
adult and had MTCs that varied in extent from 8 mm (patient 1) to widespread
metastatic disease in the remaining patients, with the exception of patients
6 and 16.
Mutation Detection. DNA was extracted from 5 paraffin-embedded and
11 frozen thyroid tissue specimens by methods described previously (25,
26). Sequencing of PCR products was performed using the Cyclist DNA
Sequencing Kit (Stratagene, La Jolla, CA). Exon 16 was amplified using
PCR and the primers CRT5G and CRT5H (3, 27). PCR products from this
primary amplification were used as templates in a secondary PCR, which
used a modified forward primer, to generate fragments specifically de
signed to be cleaved by the restriction endonuclease Rsal (New England
Biolabs, Inc., Beverly, MA) only in the presence of the codon 918ATG —¿Â»
ACG mutation (28). Ten MTCs, whose codon 918 mutation status has been
reported previously as negative (patients 6-8 and 10-16, Table 1; Ref. 12),
were also re-analyzed with this sensitive primer-mediated technique. Sam
ples were screened for mutations at codons 768 and 883 (7, 8, 13) by PCR
amplification with CRT4E and CRT4F (exon 13), and CRT17B and
CRT17G or CRT17S and CRT17A (exon 15), respectively (27), and
digestion with the restriction endonuclease Alul (New England Biolabs,
Inc.). The presence of each of the mutations causes loss of an Alul site. All
results were independently replicated in separate laboratories.
1241
Research.
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