Identification of a Novel Germ Line Variant Hotspot Mutant
p53-R175L in Pediatric Adrenal Cortical Carcinoma
Alina Nico West,
1,3
Raul C. Ribeiro,
4,7
Jesse Jenkins,
5
Carlos Rodriguez-Galindo,
4,7
Bonald C. Figueiredo,
8
Richard Kriwacki,
2,6
and Gerard P. Zambetti
2,3
Departments of
1
Interdisciplinary Sciences and
2
Molecular Sciences, University of Tennessee Health Science Center; Departments of
3
Biochemistry,
4
Hematology and Oncology,
5
Pathology, and
6
Structural Biology, and the
7
International Outreach Program, St. Jude
Children’s Research Hospital, Memphis, Tennessee; and
8
Department of Pediatrics, Center for Molecular Genetics and
Cancer Research in Children CEGEMPAC, Federal University of Parana ´, Curitiba PR, Brazil
Abstract
Hotspot mutations in the p53 tumor suppressor gene result in
the disruption of DNA contact points or alter the overall
structure of the protein to prevent DNA binding. When
inherited, hotspot mutants are associated with Li-Fraumeni
syndrome (LFS), a familial cancer predisposition. One of the
most common hotspot mutations occurs at codon 175,
resulting in an arginine to histidine substitution. We have
identified a novel germ line variant of the 175 mutant (Arg to
Leu; R175L) in a pediatric patient who developed adrenal
cortical carcinoma. Surprisingly, the family is not tumor prone
or associated with LFS. In vitro , the R175L mutant displayed an
attenuated tumor suppressor activity in the regulation of
transcription,colonyformation,andapoptosiswhencompared
with wild-type p53 and the R175H mutant. These findings
suggest that p53-R175L retains sufficient activity to suppress
LFS, but not adrenal cortical carcinoma. Therefore, not all
hotspot mutants are functionally equivalent and the biochem-
ical nature of the mutant may significantly influence clinical
outcome. The implications of these results for genetic
counseling are discussed. (Cancer Res 2006; 66(10): 5056-62)
Introduction
Thep53tumorsuppressorplaysaprominentroleintheprevention
of cancer by functioning as a transcription factor that induces
downstreamtargetswhichnegativelycontrolcellgrowth(e.g.,p21
Cip1
)
and viability (e.g., Puma and Noxa; refs. 1–3). It is, therefore, not
surprising that half of all human cancers have sustained inactivating
somatic mutations in p53. Several residues are more frequently
targeted than others and these are referred to as hotspot mutations
(for review, see ref. 4). Interestingly, hotspot mutations disrupt criti-
cal DNA contact points or alter the structure in such a way that
the protein no longer binds DNA in a sequence-specific manner (5).
In either case, prototypical hotspot mutants are unable to efficiently
induce target gene expression, cell cycle arrest, and cell death.
Germ line p53 hotspot mutations strongly predispose carriers to
cancer as children or young adults. Indeed, epidemiology studies
estimate that f70% of males and 100% of females who inherit a
p53 mutation will develop cancer, including tumors of the breast,
brain, soft tissue, bone, blood, and adrenal cortex (6). The
extraordinarily high occurrence of tumors associated with germ
line p53 mutations is referred to as Li-Fraumeni syndrome (LFS;
ref. 7). LFS is formally defined by a proband with a sarcoma before
the age of 45 years and a first-degree relative with any cancer under
45 years of age, and an additional first-degree or second-degree
family member with any cancer before 45 years or sarcoma at any
age (8). Subsequent studies identified a variation of LFS, referred to
as Li-Fraumeni-like (LFLS) syndrome, which includes a proband
with any childhood cancer or a sarcoma, brain tumor, or adrenal
cortical tumor before 45 years of age, and a first-degree or second-
degree relative with a LFS-type tumor and an additional first-
degree or second-degree relative with any cancer before 60 years of
age (9). A telling feature of LFS and LFLS is a child who has
developed an adrenal cortical tumor, although other mechanisms
(e.g., Beckwith-Wiedemann syndrome) are also recognized to
promote childhood adrenal cortical tumorigenesis.
Pediatric adrenal cortical tumors are extremely rare, with an
annual worldwide incidence of 0.3 to 0.4 per million children under
the age of 15 (10). Because childhood adrenal cortical tumors often
arise within LFS and LFLS families, they are usually associated with
a germ line p53 mutation (11). In many of these cases, the proband
has inherited a hotspot p53 mutation. However, childhood adrenal
cortical tumors can also occur outside the context of LFS/LFLS
and be associated with a constitutional mutant p53 allele (12). For
example, recent studies identified a group of pediatric adrenal
cortical tumor patients from southern Brazil who inherited a
mutation in exon 10, corresponding to an arginine to histidine
substitution at amino acid 337 (R337H; ref. 13). Interestingly, the
mutation occurs within the COOH-terminal oligomerization
domain and not the DNA-binding region. Although the mutation
could be tracked through multiple generations within the adrenal
cortical tumor families, there were no reports of sarcoma or
indications of an increased susceptibility to cancer in general. The
R337H mutation strongly predisposes carriers to adrenal cortical
tumors, but not to LFS or any other tumors (14). Interestingly, the
histidine substitution at this site within the oligomerization
domain alters the stability of the protein structure in a pH-
dependent manner, which presumably forms the basis for the
tumor specificity of this particular mutant (15).
The overall structure of p53 relies on arginine 175 (Arg
175
),
which is located in the L2 loop of the DNA binding domain (5).
Arg
175
mediates the interaction between the L2 and L3 loops to
maintain structural stability. Substitution of histidine for arginine
at amino acid 175 (R175H) is one of the most common somatic
mutations of p53 detected in human cancers (16, 17). The mutant
R175H DNA binding domain is unable to bind specifically to the
gadd45 promoter, is significantly less thermodynamically stable
than the wild-type domain, and is predicted to be completely
denatured at 37jC. Furthermore, full-length mutant p53 R175H
Requests for reprints: Gerard P. Zambetti, Department of Biochemistry, St. Jude
Children’s Research Hospital, 332 North Lauderdale, Memphis, TN 38105. Phone:
901-495-3429; Fax: 901-525-8025; E-mail: gerard.zambetti@stjude.org.
I2006 American Association for Cancer Research.
doi:10.1158/0008-5472.CAN-05-4580
Cancer Res 2006; 66: (10). May 15, 2006 5056 www.aacrjournals.org
Research Article
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