Robin Allshire

Allshire grew up in the fishing village of Howth, Co Dublin 1960–1978.{{citation needed|date=September 2018}} His parents were Arthur Gordon Allshire (1925-2012) who was a Pharmacist and Freda Margaret (née Schmutz; 1933–2014).{{cite news |url=https://notices.irishtimes.com/death/allshire-arthur/21440606 |newspaper=The Irish Times|title=ALLSHIRE, Arthur : Death notice - Irish Times Family Notices}}{{cite news |url=https://notices.irishtimes.com/death/allshire-freda-margaret/39263256 |newspaper=The Irish Times|title=ALLSHIRE, Freda Margaret : Death notice - Irish Times Family Notices}} He was awarded his Bachelor of Arts degree in Genetics by Trinity College Dublin, in 1981{{citation needed|date=September 2018}} where he was motivated by the inspirational teaching of David McConnell and colleagues at the Dept of Genetics to undertake post-graduate studies.{{cite web|url=http://allshire.biology.ed.ac.uk|title=EPIGENETICS AND SPECIALIZED CHROMATIN}} He subsequently joined the [[Medical Research Council (United Kingdom)|

Medical Research Council]] (MRC) Mammalian Genome Unit at the University of Edinburgh where he obtained his PhD in 1985{{cite thesis|degree=PhD|publisher=University of Edinburgh|hdl=1842/11176|title=Construction and analysis of vectors based on bovine papilloma virus|first= Robin Campbell|last=Allshire|year=1985|id={{EThOS|uk.bl.ethos.355979}}|oclc=606010479}} {{free access}} under the guidance of Chris Bostock and Edwin Southern investigating the use of bovine papillomavirus as a chassis for mammalian artificial chromosome construction.

In 1985 Allshire joined Nicholas Hastie's research group at the MRC Human Genetics Unit, Edinburgh (formerly MRC Clinical and Population Cytogentics Unit) as a postdoctoral researcher where he discovered that mammalian telomeres are composed of simple repetitive sequences similar to those of unicellular eukaryotes{{cite journal |doi=10.1038/332656a0 |pmid=2833706 |title=Telomeric repeat from T. Thermophila cross hybridizes with human telomeres |journal=Nature |volume=332 |issue=6165 |pages=656–9 |year=1988 |last1=Allshire |first1=Robin C |last2=Gosden |first2=John R |last3=Cross |first3=Sally H |last4=Cranston |first4=Gwen |last5=Rout |first5=Derek |last6=Sugawara |first6=Neal |last7=Szostak |first7=Jack W |last8=Fantes |first8=Peter A |last9=Hastie |first9=Nicholas D |bibcode=1988Natur.332..656A |s2cid=4352376 }} and that telomere length in blood cells shorten with age and are further eroded in cancerous cells.{{cite journal |doi=10.1038/346866a0 |pmid=2392154 |title=Telomere reduction in human colorectal carcinoma and with ageing |journal=Nature |volume=346 |issue=6287 |pages=866–8 |year=1990 |last1=Hastie |first1=Nicholas D |last2=Dempster |first2=Maureen |last3=Dunlop |first3=Malcolm G |last4=Thompson |first4=Alastair M |last5=Green |first5=Daryll K |last6=Allshire |first6=Robin C |bibcode=1990Natur.346..866H |s2cid=4258451 }} This work resulted from following the fate of fission yeast (Schizosaccharomyces pombe) telomeres after introdroducing fission yeast chromosomes into mouse cell in collaboration with Peter Fantes.{{cite journal |pmid=3475186 |year=1987 |last1=Allshire |first1=R. C |title=A fission yeast chromosome can replicate autonomously in mouse cells |journal=Cell |volume=50 |issue=3 |pages=391–403 |last2=Cranston |first2=G |last3=Gosden |first3=J. R |last4=Maule |first4=J. C |last5=Hastie |first5=N. D |last6=Fantes |first6=P. A |doi=10.1016/0092-8674(87)90493-4 |s2cid=2193386 }} In 1989 he took a position as an independent visiting scientist at Cold Spring Harbor Laboratory (CSHL) for 18 months before joining the MRC Human Genetics Unit as a junior group leader. While at CSHL he decided to switch his focus to investigating chromosomal elements in the genetically tractable fission yeast.{{cite journal |pmid=2349217 |pmc=54043 |year=1990 |last1=Allshire |first1=R. C |title=Introduction of large linear minichromosomes into Schizosaccharomyces pombe by an improved transformation procedure |journal=Proceedings of the National Academy of Sciences of the United States of America |volume=87 |issue=11 |pages=4043–7 |doi=10.1073/pnas.87.11.4043 |bibcode=1990PNAS...87.4043A |doi-access=free }} At the MRC HGU, Edinburgh (1990 - 2002), and subsequently at the Wellome Trust Centre for Cell Biology, University of Edinburgh (2002–present), he discovered that genes are silenced when placed within fission yeast centromeres{{cite journal |pmid=8287474 |year=1994 |last1=Allshire |first1=R. C |title=Position effect variegation at fission yeast centromeres |journal=Cell |volume=76 |issue=1 |pages=157–69 |last2=Javerzat |first2=J. P |last3=Redhead |first3=N. J |last4=Cranston |first4=G |doi=10.1016/0092-8674(94)90180-5 |s2cid=42369473 }}{{cite journal |pmid=7851795 |year=1995 |last1=Allshire |first1=R. C |title=Mutations derepressing silent centromeric domains in fission yeast disrupt chromosome segregation |journal=Genes & Development |volume=9 |issue=2 |pages=218–33 |last2=Nimmo |first2=E. R |last3=Ekwall |first3=K |last4=Javerzat |first4=J. P |last5=Cranston |first5=G |doi=10.1101/gad.9.2.218 |doi-access=free }} and telomeres,{{cite journal |pmid=8070408 |pmc=395293 |year=1994 |last1=Nimmo |first1=E. R |title=Telomere-associated chromosome breakage in fission yeast results in variegated expression of adjacent genes |journal=The EMBO Journal |volume=13 |issue=16 |pages=3801–11 |last2=Cranston |first2=G |last3=Allshire |first3=R. C |doi=10.1002/j.1460-2075.1994.tb06691.x }} and then utilised this gene silencing to gain fundamental insights into the processes of chromosome segregation,{{cite journal |doi=10.1038/33941 |pmid=9572142 |title=Defective meiosis in telomere-silencing mutants of Schizosaccharomyces pombe |journal=Nature |volume=392 |issue=6678 |pages=825–8 |year=1998 |last1=Nimmo |first1=Elaine R |last2=Pidoux |first2=Alison L |last3=Perry |first3=Paul E |last4=Allshire |first4=Robin C |bibcode=1998Natur.392..825N |s2cid=4412433 }}{{cite journal |pmid=11069763 |year=2000 |last1=Pidoux |first1=A. L |title=Live analysis of lagging chromosomes during anaphase and their effect on spindle elongation rate in fission yeast |journal=Journal of Cell Science |volume=113 Pt 23 |pages=4177–91 |last2=Uzawa |first2=S |last3=Perry |first3=P. E |last4=Cande |first4=W. Z |last5=Allshire |first5=R. C |issue=23 |doi=10.1242/jcs.113.23.4177 }}{{cite journal |doi=10.1126/science.1064027 |pmid=11598266 |title=Requirement of Heterochromatin for Cohesion at Centromeres |journal=Science |volume=294 |issue=5551 |pages=2539–42 |year=2001 |last1=Bernard |first1=P |last2=Maure |first2=J. F |last3=Partridge |first3=J. F |last4=Genier |first4=S |last5=Javerzat |first5=J. P |last6=Allshire |first6=R. C |bibcode=2001Sci...294.2539B |s2cid=31166180 }} and heterochromatin{{cite journal |pmid=7660126 |year=1995 |last1=Ekwall |first1=K |title=The chromodomain protein Swi6: A key component at fission yeast centromeres |journal=Science |volume=269 |issue=5229 |pages=1429–31 |last2=Javerzat |first2=J. P |last3=Lorentz |first3=A |last4=Schmidt |first4=H |last5=Cranston |first5=G |last6=Allshire |first6=R |doi=10.1126/science.7660126 |bibcode=1995Sci...269.1429E |s2cid=38678389 }}{{cite journal |pmid=9428524 |year=1997 |last1=Ekwall |first1=K |title=Transient inhibition of histone deacetylation alters the structural and functional imprint at fission yeast centromeres |journal=Cell |volume=91 |issue=7 |pages=1021–32 |last2=Olsson |first2=T |last3=Turner |first3=B. M |last4=Cranston |first4=G |last5=Allshire |first5=R. C |doi=10.1016/S0092-8674(00)80492-4 |doi-access=free }}{{cite journal |doi=10.1038/35065138 |pmid=11242054 |title=Selective recognition of methylated lysine 9 on histone H3 by the HP1 chromo domain |journal=Nature |volume=410 |issue=6824 |pages=120–4 |year=2001 |last1=Bannister |first1=Andrew J |last2=Zegerman |first2=Philip |last3=Partridge |first3=Janet F |last4=Miska |first4=Eric A |last5=Thomas |first5=Jean O |last6=Allshire |first6=Robin C |last7=Kouzarides |first7=Tony |bibcode=2001Natur.410..120B |s2cid=4334447 }}{{cite journal |pmid=12361567 |year=2002 |last1=Partridge |first1=J. F |title=Cis-acting DNA from fission yeast centromeres mediates histone H3 methylation and recruitment of silencing factors and cohesin to an ectopic site |journal=Current Biology |volume=12 |issue=19 |pages=1652–60 |last2=Scott |first2=K. S |last3=Bannister |first3=A. J |last4=Kouzarides |first4=T |last5=Allshire |first5=R. C |doi=10.1016/S0960-9822(02)01177-6 |doi-access=free }} and kinetochore CENP-A chromatin{{cite journal |doi=10.1083/jcb.200212110 |pmid=12719471 |pmc=2172903 |title=Sim4 |journal=The Journal of Cell Biology |volume=161 |issue=2 |pages=295–307 |year=2003 |last1=Pidoux |first1=Alison L |last2=Richardson |first2=William |last3=Allshire |first3=Robin C }}{{cite journal |pmid=17677001 |pmc=1934396 |year=2007 |last1=Castillo |first1=A. G |title=Plasticity of fission yeast CENP-A chromatin driven by relative levels of histone H3 and H4 |journal=PLOS Genetics |volume=3 |issue=7 |pages=e121 |last2=Mellone |first2=B. G |last3=Partridge |first3=J. F |last4=Richardson |first4=W |last5=Hamilton |first5=G. L |last6=Allshire |first6=R. C |last7=Pidoux |first7=A. L |doi=10.1371/journal.pgen.0030121 |doi-access=free }}{{cite journal |doi=10.1016/j.molcel.2007.10.010 |pmid=18158900 |pmc=2193228 |title=A NASP (N1/N2)-Related Protein, Sim3, Binds CENP-A and is Required for Its Deposition at Fission Yeast Centromeres |journal=Molecular Cell |volume=28 |issue=6 |pages=1029–44 |year=2007 |last1=Dunleavy |first1=Elaine M |last2=Pidoux |first2=Alison L |last3=Monet |first3=Marie |last4=Bonilla |first4=Carolina |last5=Richardson |first5=William |last6=Hamilton |first6=Georgina L |last7=Ekwall |first7=Karl |last8=McLaughlin |first8=Paul J |last9=Allshire |first9=Robin C }}{{cite journal |doi=10.1016/j.molcel.2009.01.019 |pmid=19217404 |pmc=2697330 |title=Fission Yeast Scm3: A CENP-A Receptor Required for Integrity of Subkinetochore Chromatin |journal=Molecular Cell |volume=33 |issue=3 |pages=299–311 |year=2009 |last1=Pidoux |first1=Alison L |last2=Choi |first2=Eun Shik |last3=Abbott |first3=Johanna K.R |last4=Liu |first4=Xingkun |last5=Kagansky |first5=Alexander |last6=Castillo |first6=Araceli G |last7=Hamilton |first7=Georgina L |last8=Richardson |first8=William |last9=Rappsilber |first9=Juri |last10=He |first10=Xiangwei |last11=Allshire |first11=Robin C }}{{cite journal |doi=10.1016/j.cell.2009.06.010 |pmid=19563746 |pmc=4397584 |title=Common Ancestry of the CENP-A Chaperones Scm3 and HJURP |journal=Cell |volume=137 |issue=7 |pages=1173–4 |year=2009 |last1=Sanchez-Pulido |first1=Luis |last2=Pidoux |first2=Alison L |last3=Ponting |first3=Chris P |last4=Allshire |first4=Robin C }}{{cite journal |doi=10.1098/rsob.140043 |pmid=24789708 |pmc=4043117 |title=Eic1 links Mis18 with the CCAN/Mis6/Ctf19 complex to promote CENP-A assembly |journal=Open Biology |volume=4 |issue=4 |pages=140043 |year=2014 |last1=Subramanian |first1=L |last2=Toda |first2=N. R. T |last3=Rappsilber |first3=J |last4=Allshire |first4=R. C }} establishment{{cite journal |doi=10.1126/science.1150944 |pmid=18174443 |pmc=2586718 |title=Heterochromatin and RNAi Are Required to Establish CENP-A Chromatin at Centromeres |journal=Science |volume=319 |issue=5859 |pages=94–7 |year=2008 |last1=Folco |first1=H. D |last2=Pidoux |first2=A. L |last3=Urano |first3=T |last4=Allshire |first4=R. C |bibcode=2008Sci...319...94F }}{{cite journal |doi=10.1126/science.1172026 |pmid=19556509 |pmc=2949999 |title=Synthetic Heterochromatin Bypasses RNAi and Centromeric Repeats to Establish Functional Centromeres |journal=Science |volume=324 |issue=5935 |pages=1716–9 |year=2009 |last1=Kagansky |first1=A |last2=Folco |first2=H. D |last3=Almeida |first3=R |last4=Pidoux |first4=A. L |last5=Boukaba |first5=A |last6=Simmer |first6=F |last7=Urano |first7=T |last8=Hamilton |first8=G. L |last9=Allshire |first9=R. C |bibcode=2009Sci...324.1716K }} and maintenance.{{cite journal |pmid=10766735 |pmc=316498 |year=2000 |last1=Partridge |first1=J. F |title=Distinct protein interaction domains and protein spreading in a complex centromere |journal=Genes & Development |volume=14 |issue=7 |pages=783–91 |last2=Borgstrøm |first2=B |last3=Allshire |first3=R. C |doi=10.1101/gad.14.7.783 }}{{cite journal |doi=10.1038/sj.emboj.7601892 |pmid=17948055 |pmc=2048757 |title=The JmjC domain protein Epe1 prevents unregulated assembly and disassembly of heterochromatin |journal=The EMBO Journal |volume=26 |issue=22 |pages=4670–82 |year=2007 |last1=Trewick |first1=Sarah C |last2=Minc |first2=Elsa |last3=Antonelli |first3=Richard |last4=Urano |first4=Takeshi |last5=Allshire |first5=Robin C }}{{cite journal |doi=10.1038/emboj.2013.72 |pmid=23572080 |pmc=3642681 |title=Distinct roles for Sir2 and RNAi in centromeric heterochromatin nucleation, spreading and maintenance |journal=The EMBO Journal |volume=32 |issue=9 |pages=1250–64 |year=2013 |last1=Buscaino |first1=Alessia |last2=Lejeune |first2=Erwan |last3=Audergon |first3=Pauline |last4=Hamilton |first4=Georgina |last5=Pidoux |first5=Alison |last6=Allshire |first6=Robin C }} He is particularly interested in the epigenetic mechanisms that allow the persistence of specialised chromatin domains through multiple cell divisions and meiosis.{{cite journal |doi=10.1126/science.1260638 |pmid=25838386 |pmc=4397586 |title=Restricted epigenetic inheritance of H3K9 methylation |journal=Science |volume=348 |issue=6230 |pages=132–5 |year=2015 |last1=Audergon |first1=P. N. C. B |last2=Catania |first2=S |last3=Kagansky |first3=A |last4=Tong |first4=P |last5=Shukla |first5=M |last6=Pidoux |first6=A. L |last7=Allshire |first7=R. C |bibcode=2015Sci...348..132A }} He has investigated how RNA interference (RNAi) mediates heterochromatin formation{{cite journal |doi=10.1038/embor.2009.273 |pmid=20062003 |pmc=2828748 |title=Hairpin RNA induces secondary small interfering RNA synthesis and silencing in trans in fission yeast |journal=EMBO Reports |volume=11 |issue=2 |pages=112–8 |year=2010 |last1=Simmer |first1=Femke |last2=Buscaino |first2=Alessia |last3=Kos-Braun |first3=Isabelle C |last4=Kagansky |first4=Alexander |last5=Boukaba |first5=Abdelhalim |last6=Urano |first6=Takeshi |last7=Kerr |first7=Alastair R W |last8=Allshire |first8=Robin C }}{{cite journal |doi=10.1016/j.cell.2010.01.038 |pmid=20211136 |pmc=2875855 |title=Stc1: A Critical Link between RNAi and Chromatin Modification Required for Heterochromatin Integrity |journal=Cell |volume=140 |issue=5 |pages=666–77 |year=2010 |last1=Bayne |first1=Elizabeth H |last2=White |first2=Sharon A |last3=Kagansky |first3=Alexander |last4=Bijos |first4=Dominika A |last5=Sanchez-Pulido |first5=Luis |last6=Hoe |first6=Kwang-Lae |last7=Kim |first7=Dong-Uk |last8=Park |first8=Han-Oh |last9=Ponting |first9=Chris P |last10=Rappsilber |first10=Juri |last11=Allshire |first11=Robin C }}{{cite journal |doi=10.1371/journal.pgen.1002499 |pmid=22319459 |pmc=3271066 |title=Raf1 is a DCAF for the Rik1 DDB1-Like Protein and Has Separable Roles in siRNA Generation and Chromatin Modification |journal=PLOS Genetics |volume=8 |issue=2 |pages=e1002499 |year=2012 |last1=Buscaino |first1=Alessia |last2=White |first2=Sharon A |last3=Houston |first3=Douglas R |last4=Lejeune |first4=Erwan |last5=Simmer |first5=Femke |last6=De Lima Alves |first6=Flavia |last7=Diyora |first7=Piyush T |last8=Urano |first8=Takeshi |last9=Bayne |first9=Elizabeth H |last10=Rappsilber |first10=Juri |last11=Allshire |first11=Robin C |doi-access=free }} and shown that splicing factors contribute to heterochromatin integrity via siRNA generation and RNAi.{{cite journal |doi=10.1126/science.1164029 |pmid=18948543 |pmc=2585287 |title=Splicing Factors Facilitate RNAi-Directed Silencing in Fission Yeast |journal=Science |volume=322 |issue=5901 |pages=602–6 |year=2008 |last1=Bayne |first1=E. H |last2=Portoso |first2=M |last3=Kagansky |first3=A |last4=Kos-Braun |first4=I. C |last5=Urano |first5=T |last6=Ekwall |first6=K |last7=Alves |first7=F |last8=Rappsilber |first8=J |last9=Allshire |first9=R. C |bibcode=2008Sci...322..602B }}{{cite journal |doi=10.1186/s13059-014-0481-4 |pmid=25274039 |pmc=4210515 |title=A systematic genetic screen identifies new factors influencing centromeric heterochromatin integrity in fission yeast |journal=Genome Biology |volume=15 |issue=10 |pages=481 |year=2014 |last1=Bayne |first1=Elizabeth H |last2=Bijos |first2=Dominika A |last3=White |first3=Sharon A |last4=Alves |first4=Flavia de Lima |last5=Rappsilber |first5=Juri |last6=Allshire |first6=Robin C |doi-access=free }} He has provided insight into how transcription and resulting non-coding RNA might influence the assembly of specialised CENP-A chromatin{{cite journal |doi=10.1074/jbc.M111.228510 |pmid=21531710 |pmc=3123123 |title=Identification of Noncoding Transcripts from within CENP-A Chromatin at Fission Yeast Centromeres |journal=Journal of Biological Chemistry |volume=286 |issue=26 |pages=23600–7 |year=2011 |last1=Choi |first1=Eun Shik |last2=Strålfors |first2=Annelie |last3=Castillo |first3=Araceli G |last4=Durand-Dubief |first4=Mickaël |last5=Ekwall |first5=Karl |last6=Allshire |first6=Robin C |doi-access=free }}{{cite journal |doi=10.1371/journal.pgen.1002985 |pmid=23028377 |pmc=3447972 |title=Factors That Promote H3 Chromatin Integrity during Transcription Prevent Promiscuous Deposition of CENP-ACnp1 in Fission Yeast |journal=PLOS Genetics |volume=8 |issue=9 |pages=e1002985 |year=2012 |last1=Choi |first1=Eun Shik |last2=Strålfors |first2=Annelie |last3=Catania |first3=Sandra |last4=Castillo |first4=Araceli G |last5=Svensson |first5=J. Peter |last6=Pidoux |first6=Alison L |last7=Ekwall |first7=Karl |last8=Allshire |first8=Robin C |doi-access=free }}{{cite journal |doi=10.1371/journal.pgen.1004986 |pmid=25738810 |pmc=4349457 |title=Sequence Features and Transcriptional Stalling within Centromere DNA Promote Establishment of CENP-A Chromatin |journal=PLOS Genetics |volume=11 |issue=3 |pages=e1004986 |year=2015 |last1=Catania |first1=Sandra |last2=Pidoux |first2=Alison L |last3=Allshire |first3=Robin C |doi-access=free }}{{cite journal |doi=10.1016/j.cub.2018.10.049 |pmid=30503616 |pmc=6303189 |title=Centromere DNA Destabilizes H3 Nucleosomes to Promote CENP-A Deposition during the Cell Cycle |journal=Current Biology |volume=28 |issue=24 |pages=3924–3936 |year=2018 |last1=Shukla M |first1=Manu |last2=Allshire |first2=Robin C }} and demonstrated that some acts of lncRNA transcription are responsive to environmental stimuli and regulate neighbouring genes by transcriptional interference.{{cite journal |doi=10.1038/ncomms6576 |pmid=25428589 |pmc=4255232 |title=Long non-coding RNA-mediated transcriptional interference of a permease gene confers drug tolerance in fission yeast |journal=Nature Communications |volume=5 |pages=5576 |year=2014 |last1=Ard |first1=Ryan |last2=Tong |first2=Pin |last3=Allshire |first3=Robin C |bibcode=2014NatCo...5.5576A }}{{cite journal |doi=10.1093/nar/gkw801 |pmid=27613421 |pmc=5159543 |title=Transcription-coupled changes to chromatin underpin gene silencing by transcriptional interference |journal=Nucleic Acids Research |volume=44 |issue=22 |pages=10619–10630 |year=2016 |last1=Ard |first1=Ryan |last2=Allshire |first2=Robin C }} Recently using fission yeast his team discovered an epigenetic mechanism that allows fungi to develop resistance to antifungal drugs without alterations to their DNA.{{cite journal |last1=Torres-Garcia |first1=Sito |title=Epigenetic gene silencing by heterochromatin primes fungal resistance |journal=Nature |year=2020 |volume=585 |issue=7825|pages=453–458 |doi=10.1038/s41586-020-2706-x |pmid=32908306 |url= |pmc=7116710 |bibcode=2020Natur.585..453T }} This finding is important for understanding how pathogenic fungi become resistant to the limited number of available antifungal agents in both clinical and agricultural arenas.

Allshire was elected a Fellow of the Royal Society of Edinburgh in 2005,{{cite web|url=http://www.icg-13.org/navigation/show_navigation?nid=6002&t=17295|title=biography|publisher= ICG|access-date=4 December 2018}} a Fellow of the Royal Society (FRS) in 2011 {{cite web|url=https://royalsociety.org/people/robin-allshire-10985/|website=royalsociety.org|author=Anon|year=2011|title=Professor Robin Allshire FRS}} One or more of the preceding sentences incorporates text from the royalsociety.org website where: {{blockquote|“All text published under the heading 'Biography' on Fellow profile pages is available under Creative Commons Attribution 4.0 International License.” --{{Webarchive|url=https://web.archive.org/web/20161111170346/https://royalsociety.org/about-us/terms-conditions-policies/|title=Royal Society Terms, conditions and policies|date=2016-11-11}}}} and a Fellow of the Academy of Medical Sciences (FMedSci) in 2020.{{cite web |url=https://acmedsci.ac.uk/fellows/fellows-directory/ordinary-fellows/fellow/Professor-Robin-Allshire-0028315 |website=Academy of Medical Sciences|title=Professor Robin Allshire | the Academy of Medical Sciences}}

• 2013 Genetics Society Medal{{cite web|url=http://www.genetics.org.uk/medals-and-prizes/genetics-society-medals-and-lectures/genetics-society-medal/genetics-society-medal-2013/|title=Genetics Society Medal 2013|publisher= Genetics Society|access-date=4 December 2018}}

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Category:20th-century British biologists

Category:21st-century British biologists

Category:1960 births

Category:Wellcome Trust Principal Research Fellows

Category:Fellows of the Royal Society of Edinburgh

Category:Fellows of the Royal Society of Biology

Category:Living people

Category:Alumni of the University of Edinburgh

Category:Alumni of Trinity College Dublin