UOA01-13: The Paternal Age Effect: His Clock is Ticking
Submitting Institution
University of OxfordUnit of Assessment
Clinical MedicineSummary Impact Type
PoliticalResearch Subject Area(s)
Biological Sciences: Genetics
Medical and Health Sciences: Neurosciences, Oncology and Carcinogenesis
Summary of the impact
Research from the University of Oxford's Clinical Genetics Laboratory
initiated the introduction of
an upper age limit of 40 years for sperm donors in the UK and
internationally and led to increased
public awareness of the effect of paternal age in the transmission of
inherited disease. Oxford
researchers, led by Professor Andrew Wilkie, were the first to describe
the exclusively paternal
transmission of de novo mutations, in a rare craniofacial disorder
called Apert Syndrome; they also
showed that the accumulation of such mutations leads to a disproportionate
risk of disease
transmission with age. By showing that the frequency of mutations
increases with paternal age,
this research contributed to important changes in clinical practice
relating to sperm donation. This
has also had a significant cultural impact, as the research and its
clinical outcomes have
challenged public perceptions of paternal age.
Underpinning research
Since 1993, the University of Oxford's Clinical Genetics Laboratory,
directed by Professor Andrew
Wilkie, has been collaborating closely with plastic surgeons at the Oxford
Craniofacial Unit to
research the genetic basis of rare craniofacial disorders. In 1995 this
group discovered that
mutations in the FGFR2 gene were responsible for Apert syndrome: a
rare craniofacial disorder1.
Since previous studies had revealed a positive correlation between
parental age and Apert
syndrome2, the group set out to investigate the origin of FGFR2
mutations.
In a pivotal study of 57 Apert syndrome families published in 19963,
the researchers revealed that
without exception the FGFR2 mutation was inherited from the
father. Furthermore, they found that
an increase in paternal age was associated with an increased likelihood of
inheriting an FGFR2
mutation3. Further work by the Clinical Genetics Laboratory
revealed that while FGFR2 mutations
can be detrimental to an embryo, they are advantageous to the cells that
generate sperm4. These
findings showed that sperm-generating cells with an FGFR2 mutation
could outcompete their non-mutant
counterparts in the testes, leading to an increased proportion of sperm
carrying the FGFR2
mutation4. This research established the general principle that
deleterious mutations can have
competitive advantage, and the group went on to show a similar paternal
age-effect mechanism for
mutations in other genes, responsible for testicular tumors and other
craniofacial disorders5. The
Oxford researchers have argued that these findings will have broad
implications for the origins of
many inherited genetic diseases4-5. They recently showed how
spermatogonial selection combines
with mutation rates to cause an increased burden of HRas mutations in the
sperm of healthy older
men6.
The research has shown, unambiguously, that paternal age is a risk factor
for the occurrence of
mutations, and has identified a mechanism by which this risk can be
amplified with age, causing
serious and unexpected disorders in offspring.
References to the research
1. Wilkie, A. O. et al. Apert syndrome results from localized
mutations of FGFR2 and is allelic
with Crouzon syndrome. Nat. Genet. 9, 165-172 (1995)
doi:10.1038/ng0295-165. Primary
paper describing the FGFR2 gene mutation in Apert Syndrome.
2. Risch, N., Reich, E. W., Wishnick, M. M. & McCarthy, J. G.
Spontaneous mutation and
parental age in humans. Am. J. Hum. Genet. 41, 218-248
(1987). Inconclusive statistical
analysis study of parental age and the incidence of new mutations.
3. Moloney, D. M. et al. Exclusive paternal origin of new
mutations in Apert syndrome. Nat.
Genet. 13, 48-53 (1996) doi:10.1038/ng0596-48. Study
showing that an increase in
paternal age is associated with an increased likelihood of inheriting
an FGFR2
mutation.
4. Goriely, A., McVean, G. A. T., Röjmyr, M., Ingemarsson, B. &
Wilkie, A. O. M. Evidence for
selective advantage of pathogenic FGFR2 mutations in the male germ line. Science
301,
643-646 (2003) doi: 10.1126/science.1085710. The first paper to
describe the accurate
measurement of any individual mutation in human sperm.
5. Goriely, A. et al. Activating mutations in FGFR3 and HRAS
reveal a shared genetic origin for
congenital disorders and testicular tumors. Nat. Genet. 41,
1247-1252 (2009) doi:
10.1038/ng.470. Paper outlining the implications for the
inheritance of
craniosynostosis syndromes and other disease-causing mutations in
sperm.
6. Giannoulatou E, McVean G, Taylor IB, McGowan SJ, Maher GJ, Iqbal Z,
Pfeifer SP, Turner
I, Burkitt-Wright EMM, Shorto J, Itani A, Turner K, Gregory L, Buck D,
Rajpert-De Meyts E,
Looijenga LHJ, Kerr B, Wilkie AOM* & Goriely A* (2013). Contributions
of intrinsic mutation
rate and selfish selection to levels of de novo HRAS mutations in the
paternal germline.
Proc Natl Acad Sci USA in press. Shows how selfish spermatogonial
selection
combines with mutation rate to cause the increased burden of mutations
in the sperm
of healthy older men.
This research was funded by the Wellcome Trust, with support from the
Government of Malaysia,
who funded a DPhil post.
Details of the impact
Research carried out by the University of Oxford's Clinical Genetics
Laboratory has directly
influenced changes in sperm donor guidelines in the UK and abroad. The
research has also
received significant media attention, contributing to improved social
awareness of paternal age
effect mutations, as well as public awareness of genetic diagnostic
testing.
Changes to Practice Guidelines
Oxford University's 1996 discovery of the paternal age effect on mutations
in Apert syndrome was
a major contributor to the 1999 decision of the British Andrology Society
to introduce an upper age
limit of 40 years for sperm donors7. The University of Oxford's
1996 paper3 was one of two studies
from the same year that were cited as primary evidence for imposing the
age limit to prevent the
transmission of somatic mutations (the other describing an association of
increase in parental age
with cases of dyskinetic cerebral palsy). In a review of sperm donor
guidance in 20048, the UK
Human Fertilisation and Embryology Authority confirmed the British
Andrology Society's
recommendations for an upper age limit, again citing the group's 1996
paper3.
This upper age limit for sperm donors was upheld in the British Andrology
Society's 2008
guidance9. The Practice Committee of the American Society for
Reproductive Medicine, and the
Practice Committee of the Society for Assisted Reproductive Technology
have since introduced an
upper age limit for sperm donors10.
Public Awareness of Paternal Age Effect
In October 2009, the Clinical Genetics Laboratory's description of
selective advantage due to
deleterious mutations in the testes5 received wide coverage in
national newspapers. Articles
featuring commentary from Professor Wilkie were published in The Times11,
The Telegraph12, and
the Daily Mail Online13. These articles and the associated
research have increased public
consciousness and understanding of the parental age effect. The impact
this research has had on
society is illustrated in a recent feature article on late fatherhood,
published in the Daily Mail14. The
article received over 70 comments from readers14.
Sources to corroborate the impact
- British Andrology Society. British Andrology Society guidelines for
the screening of semen
donors for donor insemination (1999). Hum. Reprod. 14,
1823-1826 (1999) doi:
10.1093/humrep/14.7.1823. Guidelines recommending the upper age
limit of 40 years
for all sperm donors within the UK. Guidelines directly cite the
Moloney et al. (1996)
paper from Oxford as key evidence to support the upper age limit of
40 years.
- Human Fertilisation & Embryology Authority: Seed Review: (SCAG/ELC
(06/04) 02 — ANNEX
A)(2004, June 17). (Accessed 2013), Available from
http://www.hfea.gov.uk/docs/ELC_Annex_A_Audit_June04.pdf.
The Human Fertilisation and Embryology Authority sperm donor
guidance review,
confirming recommendations for an upper age limit of 40 years for
sperm donors.
This audit directly cites Moloney, et al (1996) as key evidence.
- Association of Biomedical Andrologists; Association of Clinical
Embryologists; British
Andrology Society; British Fertility Society; Royal College of
Obstetricians and
Gynaecologists UK guidelines for the medical and laboratory screening of
sperm, egg and
embryo donors (2008) Human Fertility,11:4,201 — 210
(2008) doi:
10.1080/14647270802563816. Also available from
http://www.britishandrology.org.uk/BAS/Policy/New%202008%20Donor%20Guidelines.pdf
(Accessed 2013) British Andrology Society guidelines
confirming the upper age limit
of 40 for all sperm donors within the UK.
- Practice Committee of the American Society for Reproductive Medicine
and the Practice
Committee of the Society for Assisted Reproductive Technology.
Recommendations for
gamete and embryo donation: a committee opinion Fertil Steril.
Jan;99(1):47-62 (2013). doi:
10.1016/j.fertnstert.2012.09.037. Also available from
http://www.asrm.org/uploadedFiles/ASRM_Content/News_and_Publications/Practice_Guidelines/Guidelines_and_Minimum_Standards/2008_Guidelines_for_gamete(1).pdf
(Accessed
2013) Guidelines from the Practice Committee of the American
Society for
Reproductive Medicine, and the Practice Committee of the Society for
Assisted
Reproductive Technology recommending an upper age limit of 40 for
all sperm
donors within the US.
- Scientists discover link between older dads and genetic diseases — The
Times (October 26th
2009) (Accessed 2013). Available from
http://www.thetimes.co.uk/tto/science/genetics/article1844016.ece
Article reporting findings from Goriely et al (2009) paper
featuring commentary from
Professor Andrew Wilkie.
- Older fathers linked to genetic disease due to testicular tumours — The
Telegraph (October
26th 2009) (Accessed 2013). Available
from
http://www.telegraph.co.uk/health/healthnews/6435802/Older-fathers-linked-to-genetic-
disease-due-to-testicular-tumours.html Article reporting
findings from Goriely et al
(2009) paper featuring commentary from Professor Andrew Wilkie.
- Why older fathers are more likely to have children with genetic
disorders — Mail Online
(October 26th 2009) (Accessed 2013) Available from
http://www.dailymail.co.uk/health/article-1223025/Why-older-fathers-likely-children-genetic-disorders.html
Article reporting findings from Goriely et al (2009) paper
featuring
commentary from Professor Andrew Wilkie.
- We all know late-life motherhood poses risks for babies. But worrying
new research reveals
how having an older father can damage a child's health too. — Mail
Online (February 21st
2013) (Accessed 2013) Available from http://www.dailymail.co.uk/femail/article-
2282033/We-know-late-life-motherhood-poses-risks-babies-But-worrying-new-research-
reveals-having-older-father-damage-childs-health.html Article
reviewing the risks of late
fatherhood, quoting Professor Wilkie and his work.