Gary Ruvkun

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Gary Ruvkun
Gary Ruvkun, 2024 Nobel Prize Laureate in Medicine.jpg
Ruvkun in 2024
Born (1952-03-26) March 26, 1952 (age 72) [1]
Berkeley, California, U.S. [2]
Alma mater University of California, Berkeley (BA)
Harvard University (PhD)
Awards
Scientific career
Institutions University of California, Berkeley
Harvard University
Massachusetts Institute of Technology
Massachusetts General Hospital
Thesis The molecular genetic analysis of symbiotic nitrogen fixation (NIF) genes from rhizobium meliloti  (1982)
Doctoral advisor Frederick Ausubel
Website ruvkun.hms.harvard.edu

Gary Bruce Ruvkun (born March 26, 1952) is an American molecular biologist and Nobel laureate at Massachusetts General Hospital and professor of genetics at Harvard Medical School in Boston. [3]

Contents

Ruvkun discovered the mechanism by which lin-4 , the first microRNA (miRNA) discovered by Victor Ambros, regulates the translation of target messenger RNAs via imperfect base-pairing to those targets, and discovered the second miRNA, let-7 , and that it is conserved across animal phylogeny, including in humans. These miRNA discoveries revealed a new world of RNA regulation at an unprecedented small size scale, and the mechanism of that regulation. Ruvkun also discovered many features of insulin-like signaling in the regulation of aging and metabolism.

He was elected a Member of the American Philosophical Society in 2019. Ruvkun was awarded the 2024 Nobel Prize in Physiology or Medicine for the discovery of microRNA and its role in post-transcriptional gene regulation. [4]

Early life and education

Ruvkun was born into a Jewish family, the son of Samuel and Dora (née Gurevich) Ruvkun. [5]

Ruvkun received a Bachelor of Arts (BA) with a major in biophysics from the University of California, Berkeley in 1973. He received a Doctor of Philosophy (PhD) in biophysics from Harvard University in 1982. [6] He conducted his doctoral studies in the laboratory of Frederick M. Ausubel, where he investigated bacterial nitrogen fixation genes. Ruvkun completed postdoctoral research with Robert Horvitz at the Massachusetts Institute of Technology (MIT) and Walter Gilbert of Harvard. [7]

Research

miRNA lin-4

Ruvkun's research revealed that the miRNA lin-4, a 22 nucleotide regulatory RNA discovered in 1992 by Victor Ambros' lab, regulates its target mRNA lin-14 by forming imperfect RNA duplexes to down-regulate translation. The first indication that the key regulatory element of the lin-14 gene recognized by the lin-4 gene product was in the lin-14 3’ untranslated region came from the analysis of lin-14 gain-of-function mutations which showed that they are deletions of conserved elements in the lin-14 3’ untranslated region. Deletion of these elements relieves the normal late stage-specific repression of LIN-14 protein production, and lin-4 is necessary for that repression by the normal lin-14 3' untranslated region. [8] [9] In a key breakthrough, the Ambros lab discovered that lin-4 encodes a very small RNA product, defining the 22 nucleotide miRNAs. When Ambros and Ruvkun compared the sequence of the lin-4 miRNA and the lin-14 3’ untranslated region, they discovered that the lin-4 RNA base pairs with conserved bulges and loops to the 3’ untranslated region of the lin-14 target mRNA, and that the lin-14 gain of function mutations delete these lin-14 complementary sites to relieve the normal repression of translation by lin-4. In addition, they showed that the lin-14 3' untranslated region could confer this lin-4-dependent translational repression on unrelated mRNAs by creating chimeric mRNAs that were lin-4-responsive. In 1993, Ruvkun reported in the journal Cell on the regulation of lin-14 by lin-4. [10] In the same issue of Cell, Victor Ambros described the regulatory product of lin-4 as a small RNA. [11] These papers revealed a new world of RNA regulation at an unprecedented small size scale, and the mechanism of that regulation. [12] [13] Together, this research is now recognized as the first description of microRNAs and the mechanism by which partially base-paired miRNA::mRNA duplexes inhibit translation. [14]

microRNA, let-7

In 2000, the Ruvkun lab reported the identification of second C. elegans microRNA, let-7, which like the first microRNA regulates translation of the target gene, in this case lin-41, via imperfect base pairing to the 3’ untranslated region of that mRNA. [15] [16] This was an indication that miRNA regulation via 3’ UTR complementarity may be a common feature, and that there were likely to be more microRNAs. The generality of microRNA regulation to other animals was established by the Ruvkun lab later in 2000, when they reported that the sequence and regulation of the let-7 microRNA is conserved across animal phylogeny, including in humans. [17]

miRNAs and siRNAs

When siRNAs of the same 21-22 nucleotide size as lin-4 and let-7 were discovered in 1999 by Hamilton and Baulcombe in plants, [18] the fields of RNAi and miRNAs suddenly converged. It seemed likely that the similarly sized miRNAs and siRNAs would use similar mechanisms. In a collaborative effort, the Mello and Ruvkun labs showed that the first known components of RNA interference and their paralogs, Dicer and the PIWI proteins, are used by both miRNAs and siRNAs. [19] Ruvkun's lab in 2003 identified many more miRNAs, [20] [21] identified miRNAs from mammalian neurons, [22] and in 2007 discovered many new protein-cofactors for miRNA function. [23] [24] [25]

C. elegans metabolism and longevity

Ruvkun's laboratory has also discovered that an insulin-like signaling pathway controls C. elegans metabolism and longevity. Klass [26] Johnson [27] and Kenyon [28] showed that the developmental arrest program mediated by mutations in age-1 and daf-2 increase C. elegans longevity. The Ruvkun lab established that these genes constitute an insulin like receptor and a downstream phosphatidylinositol kinase that couple to the daf-16 gene product, a highly conserved Forkhead transcription factor. [29] Homologues of these genes have now been implicated in regulation of human aging. [30] These findings are also important for diabetes, since the mammalian orthologs of daf-16 (referred to as FOXO transcription factors) are also regulated by insulin. [31] The Ruvkun lab has used full genome RNAi libraries to discover genes that regulate aging and metabolism. Many of these genes are broadly conserved in animal phylogeny and could be targeted in diabetes drug development. [32]

SETG: The Search for Extraterrestrial Genomes

The Ruvkun lab in collaboration with Maria Zuber at MIT, Chris Carr (now at Georgia Tech), and Michael Finney (now a San Francisco biotech entrepreneur) has been developing protocols and instruments that can amplify and sequence DNA and RNA to search for life on another planet that is ancestrally related to the Tree of Life on Earth. [33] The Search for Extraterrestrial Genomes, or SETG, project has been developing a small instrument that can determine DNA sequences on Mars (or any other planetary body), and send the information in those DNA sequence files to Earth for comparison to life on Earth. [34]

Innate immune surveillance

In 2012, Ruvkun made an original contribution to the field of immunology with the publication of a featured paper in the journal Cell describing an elegant mechanism for innate immune surveillance in animals that relies on the monitoring of core cellular functions in the host, which are often sabotaged by microbial toxins during the course of infection. [35]

Microbial life beyond the Solar System

In 2019, Ruvkun, together with Chris Carr, Mike Finney and Maria Zuber, [36] presented the argument that the appearance of sophisticated microbial life on Earth soon after it cooled, and the recent discoveries of Hot Jupiters and disruptive planetary migrations in exoplanet systems favors the spread of DNA-based microbial life across the galaxy. The SETG project is working to have NASA send a DNA sequencer to Mars to search for life there in the hope that evidence will be uncovered that life did not arise originally on Earth, but elsewhere in the universe. [37]

Published articles and recognition

As of 2018, Ruvkun has published about 150 scientific articles. Ruvkun has received numerous awards for his contributions to medical science, for his contributions to the aging field [38] and to the discovery of microRNAs. [39] He is a recipient of the Lasker Award for Basic Medical Research, [40] the Gairdner Foundation International Award, and the Benjamin Franklin Medal in Life Science. [41] Ruvkun was elected as a member of the National Academy of Sciences in 2008. [42]

Awards

Ruvkun received Gruber Prize in Genetics alongside Victor Ambros in 2014. Genetics laureates.jpg
Ruvkun received Gruber Prize in Genetics alongside Victor Ambros in 2014.

See also

Related Research Articles

<i>Caenorhabditis elegans</i> Free-living species of nematode

Caenorhabditis elegans is a free-living transparent nematode about 1 mm in length that lives in temperate soil environments. It is the type species of its genus. The name is a blend of the Greek caeno- (recent), rhabditis (rod-like) and Latin elegans (elegant). In 1900, Maupas initially named it Rhabditides elegans. Osche placed it in the subgenus Caenorhabditis in 1952, and in 1955, Dougherty raised Caenorhabditis to the status of genus.

microRNA Small non-coding ribonucleic acid molecule

Micro ribonucleic acid are small, single-stranded, non-coding RNA molecules containing 21–23 nucleotides. Found in plants, animals, and even some viruses, miRNAs are involved in RNA silencing and post-transcriptional regulation of gene expression. miRNAs base-pair to complementary sequences in messenger RNA (mRNA) molecules, then silence said mRNA molecules by one or more of the following processes:

Howard Robert Horvitz ForMemRS NAS AAA&S APS NAM is an American biologist whose research on the nematode worm Caenorhabditis elegans was awarded the 2002 Nobel Prize in Physiology or Medicine, together with Sydney Brenner and John E. Sulston, whose "seminal discoveries concerning the genetic regulation of organ development and programmed cell death" were "important for medical research and have shed new light on the pathogenesis of many diseases".

<span class="mw-page-title-main">Andrew Fire</span> American biologist and professor of pathology and genetics

Andrew Zachary Fire is an American biologist and professor of pathology and of genetics at the Stanford University School of Medicine. He was awarded the 2006 Nobel Prize in Physiology or Medicine, along with Craig C. Mello, for the discovery of RNA interference (RNAi). This research was conducted at the Carnegie Institution of Washington and published in 1998.

The Let-7 microRNA precursor gives rise to let-7, a microRNA (miRNA) involved in control of stem-cell division and differentiation. let-7, short for "lethal-7", was discovered along with the miRNA lin-4 in a study of developmental timing in C. elegans, making these miRNAs the first ever discovered. let-7 was later identified in humans as the first human miRNA, and is highly conserved across many species. Dysregulation of let-7 contributes to cancer development in humans by preventing differentiation of cells, leaving them stuck in a stem-cell like state. let-7 is therefore classified as a tumor suppressor.

lin-4 microRNA precursor

In molecular biology lin-4 is a microRNA (miRNA) that was identified from a study of developmental timing in the nematode Caenorhabditis elegans. It was the first to be discovered of the miRNAs, a class of non-coding RNAs involved in gene regulation. miRNAs are transcribed as ~70 nucleotide precursors and subsequently processed by the Dicer enzyme to give a 21 nucleotide product. The extents of the hairpin precursors are not generally known and are estimated based on hairpin prediction. The products are thought to have regulatory roles through complete or partial complementarity to mRNA. The lin-4 gene has been found to lie within a 4.11kb intron of a separate host gene.

mir-10 microRNA precursor family Short non-coding RNA gene

The mir-10 microRNA precursor is a short non-coding RNA gene involved in gene regulation. It is part of an RNA gene family which contains mir-10, mir-51, mir-57, mir-99 and mir-100. mir-10, mir-99 and mir-100 have now been predicted or experimentally confirmed in a wide range of species. miR-51 and miR-57 have currently only been identified in the nematode Caenorhabditis elegans.

<span class="mw-page-title-main">David Baulcombe</span> British plant scientist and geneticist

Sir David Charles Baulcombe is a British plant scientist and geneticist. As of October 2024 he was Head of Group, Gene Expression, in the Department of Plant Sciences at the University of Cambridge, and the Edward Penley Abraham Royal Society Research Professor and Regius Professor of Botany Emeritus at Cambridge. He held the Regius botany chair in that department from 2007 to 2020.

<span class="mw-page-title-main">LIN28</span>

Lin-28 homolog A is a protein that in humans is encoded by the LIN28 gene.

<span class="mw-page-title-main">Victor Ambros</span> American developmental biologist (born 1953)

Victor R. Ambros is an American developmental biologist and Nobel Laureate who discovered the first known microRNA (miRNA). He is a professor at the University of Massachusetts Medical School. He completed both his undergraduate and doctoral studies at the Massachusetts Institute of Technology. Ambros received the Nobel Prize in Physiology or Medicine in 2024 for his research on microRNA.

mir-48 microRNA is a microRNA which is found in nematodes, in which it controls developmental timing. It acts in the heterochronic pathway, where it controls the timing of cell fate decisions in the vulva and hypodermis during larval development.

MicroRNA sequencing (miRNA-seq), a type of RNA-Seq, is the use of next-generation sequencing or massively parallel high-throughput DNA sequencing to sequence microRNAs, also called miRNAs. miRNA-seq differs from other forms of RNA-seq in that input material is often enriched for small RNAs. miRNA-seq allows researchers to examine tissue-specific expression patterns, disease associations, and isoforms of miRNAs, and to discover previously uncharacterized miRNAs. Evidence that dysregulated miRNAs play a role in diseases such as cancer has positioned miRNA-seq to potentially become an important tool in the future for diagnostics and prognostics as costs continue to decrease. Like other miRNA profiling technologies, miRNA-Seq has both advantages and disadvantages.

In molecular biology mir-84 microRNA is a short RNA molecule. MicroRNAs function to regulate the expression levels of other genes by several mechanisms.

In molecular biology mir-241 microRNA is a short RNA molecule. MicroRNAs function to regulate the expression levels of other genes by several mechanisms.

<span class="mw-page-title-main">Julie Ahringer</span> American geneticist

Julie Ann Ahringer is an American/British Professor of Genetics and Genomics, Director of the Gurdon Institute and a member of the Department of Genetics at the University of Cambridge. She leads a research lab investigating the control of gene expression.

In bioinformatics, TargetScan is a web server that predicts biological targets of microRNAs (miRNAs) by searching for the presence of sites that match the seed region of each miRNA. For many species, other types of sites, known as 3'-compensatory sites are also identified. These miRNA target predictions are regularly updated and improved by the laboratory of David Bartel in conjunction with the Whitehead Institute Bioinformatics and Research Computing Group.

NamiRNAs are a type of miRNAs present in the nucleus, which can activate gene expression by binding to the enhancer, and therefore were named nuclear activating miRNAs (NamiRNAs), such as miR-24-1 and miR-26. These miRNAs loci are enriched with epigenetic markers that display enhancer activity like histone H3K27ac, P300/CBP, and DNaseI high-sensitivity loci. These NamiRNAs are able to activate the related enhancers and co-work with them to up-regulate the expression of neighboring genes. NamiRNAs are able to promote global gene transcription by binding their targeted enhancers in whole genome level.

Iva Susan Greenwald is an American biologist who is Professor of Cell and Molecular Biology at Columbia University. She studies cell-cell interactions and cell fate specification in C. elegans. She is particularly interested in LIN-12/Notch proteins, which is the receptor of one of the major signalling systems that determines the fate of cells.

Rosalind 'Candy' Lee is a biomedical scientist, best known for her breakthrough paper on the discovery of microRNA which was published in 1993. In 2002, Lee was joint recipient of the Newcomb Cleveland Prize, for the best paper published in the journal Science that year. In 2024, Lee's 1993 paper was cited as the seminal discovery for which the Nobel Prize in physiology or medicine was awarded that year, to co-author Victor Ambros, her husband.

LIN-14 is a nuclear protein that plays a crucial role in regulating developmental timing in the nematode worm Caenorhabditis elegans. It functions as a heterochronic gene, controlling the timing of developmental events during larval development. LIN-14 protein levels are high at the beginning of the first larval stage (L1) and then rapidly decline, which is essential for the transition from early to late cell fates. LIN-14 is a BEN domain transcription factor, capable of binding DNA and directly regulating gene expression. The protein's activity is tightly regulated by lin-4, a microRNA which inhibits LIN-14 protein synthesis through complementary base pairing with sequences in the lin-14 mRNA 3' untranslated region.

References

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  12. ↑ Ruvkun, G; Wightman, B; Bürglin, T; Arasu, P (1991). "Dominant gain-of-function mutations that lead to misregulation of the C. Elegans heterochronic gene lin-14, and the evolutionary implications of dominant mutations in pattern-formation genes". Development. Supplement. 1: 47–54. PMID   1742500.
  13. ↑ Ruvkun, G.; Ambros, V.; Coulson, A.; Waterston, R.; Sulston, J.; Horvitz, H. R. (1989). "Molecular Genetics of the Caenorhabditis Elegans Heterochronic Gene Lin-14". Genetics. 121 (3): 501–516. doi:10.1093/genetics/121.3.501. PMC   1203636 . PMID   2565854.
  14. ↑ Ruvkun, G.; Wightman, B.; Ha, I. (2004). "The 20 years it took to recognize the importance of tiny RNAs". Cell. 116 (2 Suppl): S93–S96, 2 S96 following S96. doi: 10.1016/S0092-8674(04)00034-0 . PMID   15055593. S2CID   17490257.
  15. ↑ Reinhart, B. J.; Slack, F. J.; Basson, M.; Pasquinelli, A. E.; Bettinger, J. C.; Rougvie, A. E.; Horvitz, H. R.; Ruvkun, G. (2000). "The 21-nucleotide let-7 RNA regulates developmental timing in Caenorhabditis elegans". Nature. 403 (6772): 901–906. Bibcode:2000Natur.403..901R. doi:10.1038/35002607. PMID   10706289. S2CID   4384503.
  16. ↑ Slack, F. J.; Basson, M.; Liu, Z.; Ambros, V.; Horvitz, H. R.; Ruvkun, G. (2000). "The lin-41 RBCC gene acts in the C. Elegans heterochronic pathway between the let-7 regulatory RNA and the LIN-29 transcription factor". Molecular Cell. 5 (4): 659–669. doi: 10.1016/S1097-2765(00)80245-2 . PMID   10882102.
  17. ↑ Pasquinelli, A. E.; Reinhart, B. J.; Slack, F.; Martindale, M. Q.; Kuroda, M. I.; Maller, B.; Hayward, D. C.; Ball, E. E.; Degnan, B.; Müller, B.; Spring, P.; Srinivasan, J. R.; Fishman, A.; Finnerty, M.; Corbo, J.; Levine, J.; Leahy, M.; Davidson, P.; Ruvkun, E. (2000). "Conservation of the sequence and temporal expression of let-7 heterochronic regulatory RNA". Nature. 408 (6808): 86–89. Bibcode:2000Natur.408...86P. doi:10.1038/35040556. PMID   11081512. S2CID   4401732.
  18. ↑ Hamilton, A. J.; Baulcombe, D. C. (1999). "A species of small antisense RNA in posttranscriptional gene silencing in plants". Science. 286 (5441): 950–952. doi:10.1126/science.286.5441.950. PMID   10542148.
  19. ↑ Grishok, A.; Pasquinelli, A. E.; Conte, D.; Li, N.; Parrish, S.; Ha, I.; Baillie, D. L.; Fire, A.; Ruvkun, G.; Mello, C. C. (2001). "Genes and mechanisms related to RNA interference regulate expression of the small temporal RNAs that control C. Elegans developmental timing". Cell. 106 (1): 23–34. doi: 10.1016/S0092-8674(01)00431-7 . PMID   11461699. S2CID   6649604.
  20. ↑ Grad, Y.; Aach, J.; Hayes, G. D.; Reinhart, B. J.; Church, G. M.; Ruvkun, G.; Kim, J. (2003). "Computational and experimental identification of C. Elegans microRNAs". Molecular Cell. 11 (5): 1253–1263. doi: 10.1016/S1097-2765(03)00153-9 . PMID   12769849.
  21. ↑ Parry, D.; Xu, J.; Ruvkun, G. (2007). "A whole-genome RNAi Screen for C. Elegans miRNA pathway genes". Current Biology. 17 (23): 2013–2022. Bibcode:2007CBio...17.2013P. doi:10.1016/j.cub.2007.10.058. PMC   2211719 . PMID   18023351.
  22. ↑ Kim, J.; Krichevsky, A.; Grad, Y.; Hayes, G.; Kosik, K.; Church, G.; Ruvkun, G. (2004). "Identification of many microRNAs that copurify with polyribosomes in mammalian neurons". Proceedings of the National Academy of Sciences of the United States of America. 101 (1): 360–365. Bibcode:2004PNAS..101..360K. doi: 10.1073/pnas.2333854100 . PMC   314190 . PMID   14691248.
  23. ↑ Hayes, G.; Frand, A.; Ruvkun, G. (2006). "The mir-84 and let-7 paralogous microRNA genes of Caenorhabditis elegans direct the cessation of molting via the conserved nuclear hormone receptors NHR-23 and NHR-25". Development. 133 (23): 4631–4641. doi: 10.1242/dev.02655 . PMID   17065234.
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  25. ↑ Pierce, M.; Weston, M.; Fritzsch, B.; Gabel, H.; Ruvkun, G.; Soukup, G. (2008). "MicroRNA-183 family conservation and ciliated neurosensory organ expression". Evolution & Development. 10 (1): 106–113. doi:10.1111/j.1525-142X.2007.00217.x. PMC   2637451 . PMID   18184361.
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  29. ↑ Lee, Siu Sylvia; Kennedy, Scott; Tolonen, Andrew C.; Ruvkun, Gary (April 25, 2003). "DAF-16 Target Genes That Control C. elegans Life-Span and Metabolism". Science. 300 (5619): 644–647. Bibcode:2003Sci...300..644L. doi:10.1126/science.1083614. PMID   12690206.
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  32. ↑ "Gary Ruvkun, Ph.D. | Mass General Research Institute". Mass General Research Institute. Retrieved October 9, 2024.
  33. ↑ Ruvkun, Gary; Finney, Michael; Zuber, Maria T.; Carr, Chris; Church, George M.; Gilbert, Walter; Quake, Stephen; Mayer, William F. "SETG, a Search for Extraterrestrial Genomes: An in situ PCR Detector for Life on Mars Ancestrally Related to Life on Earth" (PDF). Archived (PDF) from the original on October 9, 2024. Retrieved October 9, 2024.
  34. ↑ "Overview ‹ Search for Extra-Terrestrial Genomes (SETG) — MIT Media Lab". MIT Media Lab. Archived from the original on November 30, 2023. Retrieved October 9, 2024.
  35. ↑ Melo, Justine A.; Ruvkun, Gary (April 13, 2012). "Inactivation of conserved C. elegans genes engages pathogen- and xenobiotic-associated defenses". Cell. 149 (2): 452–466. doi:10.1016/j.cell.2012.02.050. ISSN   1097-4172. PMC   3613046 . PMID   22500807.
  36. ↑ Ruvkun, Gary (April 17, 2019). "YouTube Video (24:32) – Breakthrough Discuss 2019 – What is True for E. coli on Earth Will Be True for Life on Proxima Centauri b". University of Berkeley . Archived from the original on October 9, 2024. Retrieved July 9, 2019.
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  38. ↑ "Dan David Prize 10th Anniversary 2011 Laureates Announced: The Coen Brothers – for Cinema; Marcus Feldman – for Evolution; Cynthia Kenyon and Gary Ruvkun – for Ageing". www.newswire.ca. Archived from the original on April 21, 2018. Retrieved April 25, 2018.
  39. ↑ "Gary Ruvkun" Archived May 12, 2008, at the Wayback Machine – The Gairdner Foundation (Retrieved on May 25, 2008)
  40. ↑ "Gary Ruvkun" Archived July 16, 2010, at the Wayback Machine – The Lasker Foundation (Retrieved on September 15, 2008)
  41. ↑ "Franklin Award". Archived from the original on May 15, 2008. Retrieved December 14, 2021.
  42. 1 2 "Gary Ruvkun – NAS". National Academy of Sciences. Retrieved October 9, 2024.
  43. ↑ "Rosenstiel Award Winners". Brandeis University. Archived from the original on August 4, 2017. Retrieved October 7, 2024.
  44. ↑ MGH Executive Committee on Research. "Warren Triennial Prize" . Retrieved October 9, 2024.
  45. ↑ "Gary Ruvkun - Gairdner Foundation Award Winner". The Gairdner Foundation. October 7, 2024. Retrieved October 9, 2024.
  46. 1 2 3 4 Morrison, Mike (October 7, 2024). "Mass General Hospital researcher Gary Ruvkun honored with 2024 Nobel Prize". Massachusetts General Hospital. Retrieved October 9, 2024.
  47. ↑ "2010 - 2001 Awardees". Columbia University Irving Medical Center. November 11, 2022. Retrieved October 9, 2024.
  48. ↑ "Gary B. Ruvkun | American Academy of Arts and Sciences". American Academy of Arts & Sciences. October 9, 2024. Retrieved October 9, 2024.
  49. ↑ "Past Laureates". Massry Prize. Retrieved October 9, 2024.
  50. 1 2 3 "Center for Computational and Integrative Biology" . Retrieved October 9, 2024.
  51. ↑ "Victor Ambros awarded 2016 March of Dimes prize for co-discovery of MicroRNAs". University of Massachusetts Medical School. May 3, 2016. Archived from the original on July 22, 2024. Retrieved September 9, 2016.
  52. ↑ "ScholarGPS Profile: Gary Ruvkun". Archived from the original on October 9, 2024. Retrieved October 9, 2024.
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