Editing David C. Page

{{other people|David Page}}
{{Infobox scientist
| name              = David Page
| birth_name        = David C. Page
| image             = <!--(filename only, i.e. without "File:" prefix)-->
| birth_date        = <!--{{birth date |YYYY|MM|DD}}--> {{birth year and age|1956}}
| birth_place       = [[Harrisburg, Pennsylvania]], U.S.
| fields            = [[Genetics]]
| workplaces        = {{plainlist|
* [[Whitehead Institute]]
* [[Massachusetts Institute of Technology]]}}
| education         = {{plainlist|
* [[Swarthmore College]] ([[B. A.|BA]])
* [[Harvard University]] ([[M. D.|MD]])}}
| doctoral_advisor  = 
| doctoral_students = [[Bruce Lahn]]
| known_for         = 
| awards            = {{Plainlist|
* [[MacArthur Fellows Program|MacArthur Prize Fellowship]]
* [[Searle Scholars Program|Searle Scholar Award]]}}
| spouse            = <!--(or | spouses = )-->
| children          = 
| website           = {{URL|http://wi.mit.edu/people/faculty/page}}
}}
'''David C. Page''' (born 1956) is an American biologist who is a professor at the [[Massachusetts Institute of Technology]] (MIT). He is the director of the [[Whitehead Institute]] and a [[Howard Hughes Medical Institute]] (HHMI) investigator.<ref>{{Cite web|url=https://www.hhmi.org/scientists/david-c-page|title=David C. Page|website=HHMI.org}}</ref>  He is best known for his work on mapping the [[Y-chromosome]] and on its evolution in mammals and expression during development.

==Education and early life==
Page was born in [[Harrisburg, Pennsylvania]], in 1956 and grew up in the rural outskirts of [[Pennsylvania Dutch Country|Pennsylvania Dutch country]].<ref name=":5">{{Cite journal|title = Profile of David C. Page|journal = Proceedings of the National Academy of Sciences of the United States of America|date = 2006|issn = 0027-8424|pmc = 1413862|pmid = 16481618|pages = 2471–2473|volume = 103|issue = 8|doi = 10.1073/pnas.0600615103|first = Bijal|last = Trivedi|bibcode = 2006PNAS..103.2471T|doi-access = free}}</ref><ref name=":0">{{Cite web|title = Why, Oh Y? {{!}} The Scientist Magazine|url = http://www.the-scientist.com/?articles.view/articleNo/41689/title/Why--Oh-Y-/|website = The Scientist|access-date = 2016-02-04}}</ref><ref>{{Cite journal|url = http://d3fnt9hd6tfohz.cloudfront.net/assets/issues/MURJ29_full.pdf|title = MURJ Spotlight: Professor David C. Page, Director of the Whitehead Institute|last1 = Jagetia|first1 = Riya|date = Spring 2015|journal = MIT Undergraduate Research Journal|access-date = 4 February 2016|last2 = Berg|first2 = Elizabeth|last3 = Kim|first3 = Jae Hyun|last4 = Jones|first4 = Brianna|url-status = dead|archive-url = https://web.archive.org/web/20160223095046/http://d3fnt9hd6tfohz.cloudfront.net/assets/issues/MURJ29_full.pdf|archive-date = 23 February 2016}}</ref> The first of his family to go to college, Page attended [[Swarthmore College]], where he graduated with a [[B.A.]] with highest honors in chemistry in 1978.<ref name=":0" /><ref name=":1">{{Cite web|title = DAVID PAGE LABORATORY|url = http://pagelab.wi.mit.edu/|website = pagelab.wi.mit.edu|access-date = 2016-02-04}}</ref> During his final year at Swarthmore, Page attended class just one day a week and spent the rest of his time researching [[chromatin]] structure in the laboratory of molecular biologist Robert Simpson at the [[National Institutes of Health]].<ref name=":0" /><ref name=":1" /> In 1978, Page enrolled at [[Harvard Medical School]] and the [[Harvard–MIT Division of Health Sciences and Technology|Harvard-MIT Health Sciences Program]], where he worked in the laboratories of [[David Botstein]] at MIT and Raymond White at the [[University of Massachusetts Medical School]].<ref name=":1" /> In White's lab, Page worked on a project to develop a [[Genetic linkage|genetic linkage map]] of the human genome that would become a precursor to the [[Human Genome Project]].<ref name=":5" /> The work relied on locating [[restriction fragment length polymorphism]]s (RFLP). The first RFLP that Page found was from a site of homology between the [[X chromosome]] and Y chromosome, a coincidence that would set the direction of his subsequent career.<ref name=":5" />

Page finished his [[M.D.]] degree in the spring of 1984 and started his own lab as the first Whitehead Fellow at the [[Whitehead Institute|Whitehead Institute for Biomedical Research]] researching the genetics of [[XX male syndrome]], or de la Chapelle Syndrome.<ref name=":5" /><ref name=":1" /> After Page won the [[MacArthur Fellows Program|MacArthur "Genius Grant"]] in 1986, Page was promoted to the faculty of the Whitehead Institute and the [[MIT Department of Biology]] in 1988.<ref name=":1" /><ref>{{Cite journal|url = http://bulletin.swarthmore.edu/bulletin-issue-archive/wp-content/archived_issues_pdf/Bulletin_1999_09.pdf|title = Save the males: David Page '78 reveals the evolutionary roots of sex and gender|last = Cruzan Morton|first = Carol|date = September 1999|journal = Swarthmore Bulletin|access-date = 4 February 2016}}</ref> In 1990, Page was named a Howard Hughes Medical Institute Investigator, and in 2005 he was named as director of the Whitehead Institute.<ref name=":1" />

== Research and career==
Page has worked in several areas of [[genetics]].<ref name=gs>{{Google scholar id}}</ref><ref>{{Cite news|url=https://www.nytimes.com/2009/09/15/science/15chrom.html|title=New Clues to Sex Anomalies in How Y Chromosomes Are Copied|first=Nicholas|last=Wade|work=The New York Times |date=September 14, 2009|via=NYTimes.com}}</ref>

===Mapping the Y chromosome===
In his work on de la Chapelle Syndrome in 1986, Page collaborated with the geneticist who originally identified the first XX male, [[Albert de la Chapelle]], and geneticist [[Jean Weissenbach]] to show that XX males carry a small piece of the Y chromosome.<ref name=":5" /><ref>{{Cite journal|title = Chromosome Y-specific DNA in related human XX males|journal = Nature|date = 1985-05-16|issn = 0028-0836|pmid = 2987697|pages = 224–226|volume = 315|issue = 6016|first1 = D. C.|last1 = Page|first2 = A.|last2 = de la Chapelle|first3 = J.|last3 = Weissenbach|doi=10.1038/315224a0|bibcode = 1985Natur.315..224P|s2cid = 4344130}}</ref><ref>{{Cite journal|title = Chromosome Y-specific DNA is transferred to the short arm of X chromosome in human XX males|journal = Science|date = 1986-08-15|issn = 0036-8075|pmid = 3738510|pages = 786–788|volume = 233|issue = 4765|first1 = M.|last1 = Andersson|first2 = D. C.|last2 = Page|first3 = A.|last3 = de la Chapelle|doi=10.1126/science.3738510|bibcode = 1986Sci...233..786A|s2cid = 32456133}}</ref>

In the following year, he reported that the gene [[ZFY]] induced the development of the [[testes]], a finding which received a great deal of media attention since it putatively resolving a decade-long search for the sex-determining gene.<ref name=":5" /><ref name=":6">{{Cite journal|title = The sex-determining region of the human Y chromosome encodes a finger protein|journal = Cell|date = 1987-12-24|issn = 0092-8674|pmid = 3690661|pages = 1091–1104|volume = 51|issue = 6|first1 = D. C.|last1 = Page|first2 = R.|last2 = Mosher|first3 = E. M.|last3 = Simpson|first4 = E. M.|last4 = Fisher|first5 = G.|last5 = Mardon|first6 = J.|last6 = Pollack|first7 = B.|last7 = McGillivray|first8 = A.|last8 = de la Chapelle|first9 = L. G.|last9 = Brown|doi=10.1016/0092-8674(87)90595-2|s2cid = 7454260}}</ref> In 1989, a British team of scientists led by [[Peter Goodfellow (geneticist)|Peter Goodfellow]] and [[Robin Lovell-Badge]] began to report that the testis-determining gene was not ZFY, which led Page to review his data. Page found that he had misinterpreted his data because one of the XY females in his study had a second deletion at the site which proved to be the location of the real sex-determining gene. Launching a second round of media attention, [[Nature (journal)|Nature]] published his findings together with a paper from the British group that identified the sex-determining gene, which they termed [[Testis determining factor|SRY]].<ref name=":5" /><ref name=":6" /><ref>{{Cite journal|title = A gene from the human sex-determining region encodes a protein with homology to a conserved DNA-binding motif|journal = Nature|date = 1990-07-19|issn = 0028-0836|pmid = 1695712|pages = 240–244|volume = 346|issue = 6281|doi = 10.1038/346240a0|first1 = A. H.|last1 = Sinclair|first2 = P.|last2 = Berta|first3 = M. S.|last3 = Palmer|first4 = J. R.|last4 = Hawkins|first5 = B. L.|last5 = Griffiths|first6 = M. J.|last6 = Smith|first7 = J. W.|last7 = Foster|first8 = A. M.|last8 = Frischauf|first9 = R.|last9 = Lovell-Badge|bibcode = 1990Natur.346..240S|s2cid = 4364032|url = http://rcin.org.pl/Content/66315|url-access = subscription}}</ref>

Despite a belief among geneticists that the Y chromosome contained few genes other than the sex-determining gene, Page continued to map the Y chromosome. He had already published [[Deletion mapping|DNA-based deletion maps]] of the Y chromosome in 1986,<ref>{{Cite journal|title = A deletion map of the human Y chromosome based on DNA hybridization.|journal = American Journal of Human Genetics|date = 1986-02-01|issn = 0002-9297|pmc = 1684749|pmid = 3004206|pages = 109–124|volume = 38|issue = 2|first1 = G|last1 = Vergnaud|first2 = D C|last2 = Page|first3 = M C|last3 = Simmler|first4 = L|last4 = Brown|first5 = F|last5 = Rouyer|first6 = B|last6 = Noel|first7 = D|last7 = Botstein|first8 = A|last8 = de la Chapelle|first9 = J|last9 = Weissenbach}}</ref> and went on to develop comprehensive clone-based physical maps of the chromosome in 1992<ref>{{Cite journal|url = http://pagelab.wi.mit.edu/pdf/1992%20-%20The%20human%20Y%20chromosome%20-%20a%2043-interval%20map%20based%20on%20naturally%20occurring%20deletions.pdf|title = The Human Y Chromosome: A 43-Interval Map Based on Naturally Occurring Deletions|last1 = Vollrath|first1 = Douglas|date = 2 October 1992|journal = Science|doi = 10.1126/science.1439769|pmid = 1439769|access-date = 4 February 2016|last2 = Foote|first2 = Simon|last3 = Hilton|first3 = Adrienne|last4 = Brown|first4 = Laura G.|last5 = Beer-Romero|first5 = Peggy|last6 = Bogan|first6 = Jonathan|last7 = Page|first7 = David C.|issue = 5079|volume = 258|pages = 52–59|bibcode = 1992Sci...258...52V}}</ref><ref>{{Cite journal|url = http://pagelab.wi.mit.edu/pdf/1992%20-%20The%20human%20Y%20chromosome%20-%20overlapping%20DNA%20clones%20spanning%20the%20euchromatic%20region.pdf|title = The Human Y Chromosome: Spanning the Euchromatic Region|last1 = Foote|first1 = Simon|date = 2 October 1992|journal = Science|doi = 10.1126/science.1359640|pmid = 1359640|access-date = 4 February 2016|last2 = Vollrath|first2 = Douglas|last3 = Hilton|first3 = Adrienne|last4 = Page|first4 = David C.|issue = 5079|volume = 258|pages = 60–66}}</ref> and systematic catalogs of Y-linked genes in 1997.<ref>{{Cite journal|url = http://pagelab.wi.mit.edu/pdf/1997%20-%20Functional%20coherence%20of%20the%20human%20Y%20chromosome.pdf|title = Functional Coherence of the Human Y Chromosome|last1 = Lahn|first1 = Bruce|date = 24 October 1997|journal = Science|doi = 10.1126/science.278.5338.675|pmid = 9381176|access-date = 4 February 2016|last2 = Page|first2 = David C.|issue = 5338|volume = 278|pages = 675–680|bibcode = 1997Sci...278..675L}}</ref> Page collaborated with a team at the [[McDonnell Genome Institute|Genome Institute]] at [[Washington University in St. Louis|Washington University]] to make a complete map of the Y chromosome, which they achieved in 2003.<ref name=":3">{{Cite journal |last1 = Skaletsky|first1 = H.|last2 = Kuroda-Kawaguchi|first2 = T.|last3 = Minx|first3 = P. J.|last4 = Cordum|first4 = H. S.|last5 = Hillier|first5 = L.|last6 = Brown|first6 = L. G.|last7 = Repping|first7 = S.|last8 = Pyntikova|first8 = T.|last9 = Ali|first9 = J.|last10 = Bieri|doi = 10.1038/nature01722|first10 = T.|last11 = Chinwalla|first11 = A.|last12 = Delehaunty|first12 = A.|last13 = Delehaunty|first13 = K.|last14 = Du|first14 = H.|last15 = Fewell|first15 = G.|last16 = Fulton|first16 = L.|last17 = Fulton|first17 = R.|last18 = Graves|first18 = T.|last19 = Hou|first19 = S. F.|last20 = Latrielle|first20 = P.|last21 = Leonard|first21 = S.|last22 = Mardis|first22 = E.|last23 = Maupin|first23 = R.|last24 = McPherson|first24 = J.|last25 = Miner|first25 = T.|last26 = Nash|first26 = W.|last27 = Nguyen|first27 = C.|last28 = Ozersky|first28 = P.|last29 = Pepin|first29 = K.|last30 = Rock|first30 = S.|title = The male-specific region of the human Y chromosome is a mosaic of discrete sequence classes|journal = Nature|volume = 423|issue = 6942|pages = 825–837|year = 2003|pmid = 12815422|bibcode = 2003Natur.423..825S|doi-access = free}}</ref> To do so, Page and his colleagues developed a new sequencing technique, single-haplotype iterative mapping and sequencing (SHIMS), since mammalian sex chromosomes contain too many repetitive sequences to be sequenced by conventional approaches.<ref name=":0" /> The development of SHIMS allowed Page to identify long palindromic sequences on the long arm of the Y chromosome, which he would go on to show made the Y chromosome vulnerable to the deletions that cause spermatogenic failure (an inability to produce sperm).<ref>{{Cite journal|title = Recombination between palindromes P5 and P1 on the human Y chromosome causes massive deletions and spermatogenic failure|journal = American Journal of Human Genetics|date = 2002-10-01|issn = 0002-9297|pmc = 419997|pmid = 12297986|pages = 906–922|volume = 71|issue = 4|doi = 10.1086/342928|first1 = Sjoerd|last1 = Repping|first2 = Helen|last2 = Skaletsky|first3 = Julian|last3 = Lange|first4 = Sherman|last4 = Silber|first5 = Fulco|last5 = Van Der Veen|first6 = Robert D.|last6 = Oates|first7 = David C.|last7 = Page|first8 = Steve|last8 = Rozen}}</ref> In 2012, Page characterized the most common genetic cause of spermatogenic failure, the deletion of the AZFc region of the Y chromosome.<ref name=":4">{{Cite web|url=http://wi.mit.edu/people/faculty|title=Whitehead Institute - Faculty|website=wi.mit.edu|date=22 May 2024 }}</ref><ref>{{Cite journal|title = AZFc deletions and spermatogenic failure: a population-based survey of 20,000 Y chromosomes|journal = American Journal of Human Genetics|date = 2012-11-02|issn = 1537-6605|pmc = 3487143|pmid = 23103232|pages = 890–896|volume = 91|issue = 5|doi = 10.1016/j.ajhg.2012.09.003|first1 = Steven G.|last1 = Rozen|first2 = Janet D.|last2 = Marszalek|first3 = Kathryn|last3 = Irenze|first4 = Helen|last4 = Skaletsky|first5 = Laura G.|last5 = Brown|first6 = Robert D.|last6 = Oates|first7 = Sherman J.|last7 = Silber|first8 = Kristin|last8 = Ardlie|first9 = David C.|last9 = Page}}</ref> The lab also found that aberrant crossing over within the Y chromosome's palindromes underlies a wide range of disorders of sexual differentiation, including Turner syndrome.<ref name=":4" />

===Evolution of the Y chromosome===
With the development of more detailed maps of the Y chromosome, in the mid-1990s Page began to find genetic evidence confirming the theory that both the X and Y chromosomes had evolved from [[autosome]]s, beginning with the 1996 discovery that a family of genes called [[DAZ associated protein 1|DAZ]] (deleted in azoospermia) had been transposed from an autosome to the Y chromosome.<ref name=":0" /><ref>{{Cite journal|title = The DAZ gene cluster on the human Y chromosome arose from an autosomal gene that was transposed, repeatedly amplified and pruned|journal = Nature Genetics|date = 1996-11-01|issn = 1061-4036|pmid = 8896558|pages = 292–299|volume = 14|issue = 3|doi = 10.1038/ng1196-292|first1 = R.|last1 = Saxena|first2 = L. G.|last2 = Brown|first3 = T.|last3 = Hawkins|first4 = R. K.|last4 = Alagappan|first5 = H.|last5 = Skaletsky|first6 = M. P.|last6 = Reeve|first7 = R.|last7 = Reijo|first8 = S.|last8 = Rozen|first9 = M. B.|last9 = Dinulos|s2cid = 34964224}}</ref> In 1999, Page and his then-graduate student [[Bruce Lahn]] showed that the X and Y chromosomes had diverged in four steps, beginning 200-300 million years ago.<ref>{{Cite journal|title = Four evolutionary strata on the human X chromosome|journal = Science|date = 1999-10-29|issn = 0036-8075|pmid = 10542153|pages = 964–967|volume = 286|issue = 5441|first1 = B. T.|last1 = Lahn|first2 = D. C.|last2 = Page|doi=10.1126/science.286.5441.964}}</ref> Later cross-species comparisons would show that while ancestral genes on the Y chromosome initially underwent rapid decay,<ref name=":3" /><ref>{{Cite journal|title = Convergent evolution of chicken Z and human X chromosomes by expansion and gene acquisition|journal = Nature|date = 2010-07-29|issn = 1476-4687|pmc = 2943333|pmid = 20622855|pages = 612–616|volume = 466|issue = 7306|doi = 10.1038/nature09172|first1 = Daniel W.|last1 = Bellott|first2 = Helen|last2 = Skaletsky|first3 = Tatyana|last3 = Pyntikova|first4 = Elaine R.|last4 = Mardis|first5 = Tina|last5 = Graves|first6 = Colin|last6 = Kremitzki|first7 = Laura G.|last7 = Brown|first8 = Steve|last8 = Rozen|first9 = Wesley C.|last9 = Warren|bibcode = 2010Natur.466..612B|url = http://dspace.mit.edu/bitstream/1721.1/66578/1/050610_Submitted.pdf}}</ref> the remaining genes have remained stable for the last 25 million years,<ref>{{Cite journal|title = Strict evolutionary conservation followed rapid gene loss on human and rhesus Y chromosomes|journal = Nature|date = 2012-03-01|issn = 0028-0836|pmc = 3292678|pmid = 22367542|pages = 82–86|volume = 483|issue = 7387|doi = 10.1038/nature10843|language = en|first1 = Jennifer F.|last1 = Hughes|first2 = Helen|last2 = Skaletsky|first3 = Laura G.|last3 = Brown|first4 = Tatyana|last4 = Pyntikova|first5 = Tina|last5 = Graves|first6 = Robert S.|last6 = Fulton|first7 = Shannon|last7 = Dugan|first8 = Yan|last8 = Ding|first9 = Christian J.|last9 = Buhay|bibcode = 2012Natur.483...82H}}</ref> overturning the long-held view that the Y chromosome was going extinct.<ref name=":0" /> In a 2014 study, Page concluded that the conserved genes on the Y chromosome played an important role in male viability, since they were dosage-dependent genes with similar but not identical counterparts on the X chromosome that all have regulatory roles in transcription, translation, and protein stability. Because these genes are expressed throughout the body, Page further concluded that these genes give rise to differences in the biochemistry of male and female tissues.<ref>{{Cite journal|title = Mammalian Y chromosomes retain widely expressed dosage-sensitive regulators|journal = Nature|date = 2014-04-24|issn = 0028-0836|pmc = 4139287|pmid = 24759411|pages = 494–499|volume = 508|issue = 7497|doi = 10.1038/nature13206|language = en|first1 = Daniel W.|last1 = Bellott|first2 = Jennifer F.|last2 = Hughes|first3 = Helen|last3 = Skaletsky|first4 = Laura G.|last4 = Brown|first5 = Tatyana|last5 = Pyntikova|first6 = Ting-Jan|last6 = Cho|first7 = Natalia|last7 = Koutseva|first8 = Sara|last8 = Zaghlul|first9 = Tina|last9 = Graves|bibcode = 2014Natur.508..494B}}</ref>

In super-resolution studies of the sex chromosomes, Page has found evidence of an evolutionary "arms race" between the X and Y chromosomes for transmission to the next generation. In one study, Page found that human X and mouse Y chromosomes have converged, independently acquiring and amplifying gene families expressed in testicular germ cells.<ref>{{Cite journal|title = Independent specialization of the human and mouse X chromosomes for the male germ line|journal = Nature Genetics|date = 2013-09-01|issn = 1061-4036|pmc = 3758364|pmid = 23872635|pages = 1083–1087|volume = 45|issue = 9|doi = 10.1038/ng.2705|language = en|first1 = Jacob L.|last1 = Mueller|first2 = Helen|last2 = Skaletsky|first3 = Laura G.|last3 = Brown|first4 = Sara|last4 = Zaghlul|first5 = Susan|last5 = Rock|first6 = Tina|last6 = Graves|first7 = Katherine|last7 = Auger|first8 = Wesley C.|last8 = Warren|first9 = Richard K.|last9 = Wilson}}</ref> Another study found that the mouse Y chromosome had acquired and massively amplified genes homologous to the testis-expressed gene families on the mouse X chromosome.<ref>{{Cite journal|title = Sequencing the mouse Y chromosome reveals convergent gene acquisition and amplification on both sex chromosomes|journal = Cell|date = 2014-11-06|issn = 1097-4172|pmc = 4260969|pmid = 25417157|pages = 800–813|volume = 159|issue = 4|doi = 10.1016/j.cell.2014.09.052|first1 = Y. Q. Shirleen|last1 = Soh|first2 = Jessica|last2 = Alföldi|first3 = Tatyana|last3 = Pyntikova|first4 = Laura G.|last4 = Brown|first5 = Tina|last5 = Graves|first6 = Patrick J.|last6 = Minx|first7 = Robert S.|last7 = Fulton|first8 = Colin|last8 = Kremitzki|first9 = Natalia|last9 = Koutseva}}</ref>

===The genetics of germ cells===
Page used the mouse as a model to study the genetics of [[Meiosis|meiotic]] initiation, showing that [[retinoic acid]] (RA) is the key factor which induces meiosis, as well as identifying several important genes crucial to the meiotic initiation pathway, including Stra8 and [[DAZL]].<ref>{{Cite journal|title = In germ cells of mouse embryonic ovaries, the decision to enter meiosis precedes premeiotic DNA replication|journal = Nature Genetics|date = 2006-12-01|issn = 1061-4036|pmid = 17115059|pages = 1430–1434|volume = 38|issue = 12|doi = 10.1038/ng1919|first1 = Andrew E.|last1 = Baltus|first2 = Douglas B.|last2 = Menke|first3 = Yueh-Chiang|last3 = Hu|first4 = Mary L.|last4 = Goodheart|first5 = Anne E.|last5 = Carpenter|first6 = Dirk G.|last6 = de Rooij|first7 = David C.|last7 = Page|s2cid = 17258424}}</ref><ref>{{Cite journal|title = Germ Cell-Intrinsic and -Extrinsic Factors Govern Meiotic Initiation in Mouse Embryos|journal = Science|date = 2008-12-12|issn = 0036-8075|pmid = 19074348|pages = 1685–1687|volume = 322|issue = 5908|doi = 10.1126/science.1166340|language = en|first1 = Yanfeng|last1 = Lin|first2 = Mark E.|last2 = Gill|first3 = Jana|last3 = Koubova|first4 = David C.|last4 = Page|bibcode = 2008Sci...322.1685L|s2cid = 11261341}}</ref><ref>{{Cite journal|title = Retinoic acid regulates sex-specific timing of meiotic initiation in mice|journal = Proceedings of the National Academy of Sciences of the United States of America|date = 2006-02-21|issn = 0027-8424|pmc = 1413806|pmid = 16461896|pages = 2474–2479|volume = 103|issue = 8|doi = 10.1073/pnas.0510813103|language = en|first1 = Jana|last1 = Koubova|first2 = Douglas B.|last2 = Menke|first3 = Qing|last3 = Zhou|first4 = Blanche|last4 = Capel|first5 = Michael D.|last5 = Griswold|first6 = David C.|last6 = Page|bibcode = 2006PNAS..103.2474K|doi-access = free}}</ref> Page further discovered that the differentiation germ cells into [[gametocyte]]s ([[oocyte]]s in females or [[spermatocyte]]s in males) does not depend on meiotic initiation, as commonly thought, showing that germ cells deficient in Stra8, a gene that activates the meiotic pathway, are still capable of growth and differentiation.<ref>{{Cite journal|title = Periodic retinoic acid-STRA8 signaling intersects with periodic germ-cell competencies to regulate spermatogenesis|journal = Proceedings of the National Academy of Sciences of the United States of America|date = 2015-05-05|issn = 1091-6490|pmc = 4426408|pmid = 25902548|pages = E2347–2356|volume = 112|issue = 18|doi = 10.1073/pnas.1505683112|first1 = Tsutomu|last1 = Endo|first2 = Katherine A.|last2 = Romer|first3 = Ericka L.|last3 = Anderson|first4 = Andrew E.|last4 = Baltus|first5 = Dirk G.|last5 = de Rooij|first6 = David C.|last6 = Page|bibcode = 2015PNAS..112E2347E|doi-access = free}}</ref><ref>{{Cite journal|title = Oocyte differentiation is genetically dissociable from meiosis in mice|journal = Nature Genetics|date = 2013-08-01|issn = 1061-4036|pmc = 3747777|pmid = 23770609|pages = 877–883|volume = 45|issue = 8|doi = 10.1038/ng.2672|language = en|first1 = Gregoriy A.|last1 = Dokshin|first2 = Andrew E.|last2 = Baltus|first3 = John J.|last3 = Eppig|first4 = David C.|last4 = Page}}</ref>

===Awards and honors===
His honors and awards include:
* 1986 [[MacArthur Fellows Program|MacArthur Prize Fellowship]]<ref>{{Cite web|title = David C. Page — MacArthur Foundation|url = https://www.macfound.org/fellows/282/|website = www.macfound.org|access-date = 2016-02-04}}</ref>
* 1989 [[Searle Scholars Program|Searle Scholar Award]]<ref>{{Cite web|title = Searle Scholars Program : David C. Page (1989)|url = http://www.searlescholars.net/person/313|website = www.searlescholars.net|access-date = 2016-02-04}}</ref>
* 1997 Francis Amory Prize, [[American Academy of Arts and Sciences]] (for genetic studies of mammalian sex determination; prize shared with Peter Goodfellow and Robin Lovell-Badge)
* 2003 [[Curt Stern Award]], [[American Society of Human Genetics]] (for outstanding scientific achievement in human genetics)<ref>{{Cite journal|title = On Low Expectations Exceeded; or, The Genomic Salvation of the Y Chromosome|journal = American Journal of Human Genetics|date = 2004-03-01|issn = 0002-9297|pmc = 1182254|pmid = 15053010|pages = 399–402|volume = 74|issue = 3|first = David C.|last = Page|doi=10.1086/382659}}</ref>
* 2005 Member, [[National Academy of Sciences]]<ref name=":5"/>
* 2007 Fellow, [[American Association for the Advancement of Science]]<ref name=":1" />
* 2008 Member, [[National Academy of Medicine]]<ref>{{Cite web|title = MIT Office of the Provost, Institutional Research|url = http://web.mit.edu/ir/pop/awards/iom.html|website = web.mit.edu|access-date = 2016-02-04}}</ref>
* 2011 Fellow, [[American Academy of Arts and Sciences]] (Class Speaker, Biological Sciences)<ref>{{Cite web|title = Press Releases - American Academy of Arts & Sciences|url = https://www.amacad.org/content/news/pressReleases.aspx?pr=133|website = www.amacad.org|access-date = 2016-02-04}}</ref>

==References==
{{Reflist}}

{{Authority control}}

{{DEFAULTSORT:Page, David C.}}
[[Category:Living people]]
[[Category:Swarthmore College alumni]]
[[Category:Harvard Medical School alumni]]
[[Category:Massachusetts Institute of Technology faculty]]
[[Category:MacArthur Fellows]]
[[Category:Members of the United States National Academy of Sciences]]
[[Category:Whitehead Institute faculty]]
[[Category:Howard Hughes Medical Investigators]]
[[Category:21st-century American biologists]]
[[Category:1956 births]]
[[Category:Scientists from Pennsylvania]]
[[Category:People from Harrisburg, Pennsylvania]]
[[Category:Members of the National Academy of Medicine]]