Nathan Young, PhD

Title(s)Associate Professor, Orthopaedic Surgery
SchoolSchool of Medicine
Address1001 Potrero Ave
San Francisco CA 94110
Phone628-206-5366
ORCID ORCID Icon0000-0002-3437-7493 Additional info
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    Collapse Biography 
    Collapse Education and Training
    University of California, San FranciscoPostdoctoral Studies2007Orthopaedic Surgery
    Stanford UniversityPostdoctoral Studies2006Plastic Surgery
    University of CalgaryPostdoctoral Studies2004Cell Biology & Anatomy
    Harvard UniversityPh.D.2003Biological Anthropology
    New College of FloridaB.A.1995Anthropology

    Collapse Overview 
    Collapse Overview
    My research utilizes the concepts of variation, functional compromise, and historical constraint as fundamental explanatory principles to address basic biomedical questions through the lens of evolution. My lab combines classical embryology in a range of model and non-model systems with modern genetic tools and advanced methods for quantifying and comparing phenotypes at a range of scales. This approach yields significant insights into not only the processes that contribute to evolutionary diversity among species, but also how individual differences ranging from normal to disease states are generated. My research therefore investigates both normal mechanisms of development as well as the etiology of structural birth defects, and is relevant to longstanding goals of providing personalized and predictive medicine.
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    A Predictive Developmental Morphospace Model of Cleft Lip
    NIH R56DE029124Sep 3, 2020 - Aug 31, 2021
    Role: Principal Investigator
    The developmental genetic basis for evolutionary variation in the hominin shoulder
    NSF BCS-1518596Jul 1, 2015 - Jun 30, 2018
    Role: Co-Investigator
    A Molecular Investigation of Growth and Shape Variation in Cleft-Lip Etiology
    NIH F32DE018596Sep 1, 2007 - Aug 31, 2009
    Role: Principal Investigator

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    Collapse Publications
    Publications listed below are automatically derived from MEDLINE/PubMed and other sources, which might result in incorrect or missing publications. Researchers can login to make corrections and additions, or contact us for help. to make corrections and additions.
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    Altmetrics Details PMC Citations indicate the number of times the publication was cited by articles in PubMed Central, and the Altmetric score represents citations in news articles and social media. (Note that publications are often cited in additional ways that are not shown here.) Fields are based on how the National Library of Medicine (NLM) classifies the publication's journal and might not represent the specific topic of the publication. Translation tags are based on the publication type and the MeSH terms NLM assigns to the publication. Some publications (especially newer ones and publications not in PubMed) might not yet be assigned Field or Translation tags.) Click a Field or Translation tag to filter the publications.
    1. Cellular Dynamics and the Developmental Basis for Craniofacial Variation in Human Evolution and Disease. Evolutionary Cell Processes in Primates Volume 1: Bones, Brains, and Muscle (J. Richtsmeier & K. Pitirri, editors). 2021; 227-244. Young NM, Marcucio RS, Hallgrímsson B, Richbourg HA, Green R.
    2. Wnt Signaling Drives Correlated Changes in Facial Morphology and Brain Shape. Front Cell Dev Biol. 2021; 9:644099. Marchini M, Hu D, Lo Vercio L, Young NM, Forkert ND, Hallgrímsson B, Marcucio R. PMID: 33855022; PMCID: PMC8039397.
      View in: PubMed   Mentions: 2  
    3. Effects of Multi-Generational Soft Diet Consumption on Mouse Craniofacial Morphology. Front Physiol. 2020; 11:783. Hassan MG, Kaler H, Zhang B, Cox TC, Young N, Jheon AH. PMID: 32754047; PMCID: PMC7367031.
      View in: PubMed   Mentions: 3  
    4. Three-dimensional scapular morphology is associated with rotator cuff tears and alters the abduction moment arm of the supraspinatus. Clin Biomech (Bristol, Avon). 2020 08; 78:105091. Lee ECS, Roach NT, Clouthier AL, Bicknell RT, Bey MJ, Young NM, Rainbow MJ. PMID: 32580097; PMCID: PMC8161464.
      View in: PubMed   Mentions: 1     Fields:    Translation:Humans
    5. Radiographic shoulder parameters and their relationship to outcomes following rotator cuff repair: a systematic review. Shoulder Elbow. 2021 Aug; 13(4):371-379. Zaid MB, Young NM, Pedoia V, Feeley BT, Ma CB, Lansdown DA. PMID: 34394734; PMCID: PMC8355656.
      View in: PubMed   Mentions:
    6. Integration and the Developmental Genetics of Allometry. Integr Comp Biol. 2019 11 01; 59(5):1369-1381. Hallgrímsson B, Katz DC, Aponte JD, Larson JR, Devine J, Gonzalez PN, Young NM, Roseman CC, Marcucio RS. PMID: 31199435; PMCID: PMC6934422.
      View in: PubMed   Mentions: 12     Fields:    Translation:HumansAnimals
    7. Anatomic shoulder parameters and their relationship to the presence of degenerative rotator cuff tears and glenohumeral osteoarthritis: a systematic review and meta-analysis. J Shoulder Elbow Surg. 2019 Dec; 28(12):2457-2466. Zaid MB, Young NM, Pedoia V, Feeley BT, Ma CB, Lansdown DA. PMID: 31353303.
      View in: PubMed   Mentions: 7     Fields:    Translation:Humans
    8. Emergent Properties of Facial Morphogenesis Regulated by Fgf Signaling. The FASEB Journal. 2019 Apr 1; 33(S1):774.18-774.18. Marcucio MR, Hu HD, Hallgrimsson HB, Young YN. .
      View in: Publisher Site   Mentions:
    9. Integration and the genetics of variation in facial shape. The FASEB Journal. 2019 Apr 1; 33(S1):330.2-330.2. Hallgrimsson HB, Aponte AJ, Katz KD, Bannister BJ, Green GR, Young YN, Cheverud CJ, Forkert FN, Marcucio MR. .
      View in: Publisher Site   Mentions:
    10. Untangling Sources of Phenotypic Variation Characterizing the Craniofacial Disease Holoprosencephaly. The FASEB Journal. 2018 Apr 1; 32:776.11-776.11. Lainoff LA, Young YN, Hallgrimsson HB, Marcucio MR. .
      View in: Publisher Site   Mentions:
    11. Quantifying the Genotype to Phenotype Map in Developing Mice. The FASEB Journal. 2018 Apr 1; 32:lb529-lb529. Richbourg RH, Young YN, Takenaka TR, Marcucio MR. .
      View in: Publisher Site   Mentions:
    12. Integrating "Evo" and "Devo": The Limb as Model Structure. Integr Comp Biol. 2017 12 01; 57(6):1293-1302. Young NM. PMID: 28992180.
      View in: PubMed   Mentions:    Fields:    Translation:Animals
    13. Craniofacial diversification in the domestic pigeon and the evolution of the avian skull. Nat Ecol Evol. 2017 Mar 13; 1(4):95. Young NM, Linde-Medina M, Fondon JW, Hallgrímsson B, Marcucio RS. PMID: 28812673; PMCID: PMC5559897.
      View in: PubMed   Mentions: 15     Fields:    
    14. Facial surface morphology predicts variation in internal skeletal shape. Am J Orthod Dentofacial Orthop. 2016 Apr; 149(4):501-8. Young NM, Sherathiya K, Gutierrez L, Nguyen E, Bekmezian S, Huang JC, Hallgrímsson B, Lee JS, Marcucio RS. PMID: 27021454; PMCID: PMC4813327.
      View in: PubMed   Mentions: 10     Fields:    Translation:Humans
    15. Reply to Almécija: A new direction for reconstructing our last common ancestor with chimpanzees. Proc Natl Acad Sci U S A. 2016 Feb 23; 113(8):E945. Young NM, Capellini TD, Roach NT, Alemseged Z. PMID: 26862164; PMCID: PMC4776488.
      View in: PubMed   Mentions:    Fields:    Translation:HumansAnimals
    16. Out on a Limb. Developmental Approaches to Human Evolution. 2016 Jan 29; 101-137. Young YN, Capellini CT. .
      View in: Publisher Site   Mentions:
    17. Reply to Melillo: Woranso-Mille is consistent with an australopithecine shoulder intermediate between African apes and Homo. Proc Natl Acad Sci U S A. 2015 Dec 29; 112(52):E7160. Young NM, Capellini TD, Roach NT, Alemseged Z. PMID: 26676585; PMCID: PMC4702962.
      View in: PubMed   Mentions: 2     Fields:    Translation:HumansAnimals
    18. Morphometrics, 3D Imaging, and Craniofacial Development. Curr Top Dev Biol. 2015; 115:561-97. Hallgrimsson B, Percival CJ, Green R, Young NM, Mio W, Marcucio R. PMID: 26589938; PMCID: PMC5299999.
      View in: PubMed   Mentions: 21     Fields:    Translation:HumansAnimals
    19. Facial Morphogenesis: Physical and Molecular Interactions Between the Brain and the Face. Curr Top Dev Biol. 2015; 115:299-320. Marcucio R, Hallgrimsson B, Young NM. PMID: 26589930; PMCID: PMC6936956.
      View in: PubMed   Mentions: 33     Fields:    Translation:HumansAnimalsCells
    20. Fossil hominin shoulders support an African ape-like last common ancestor of humans and chimpanzees. Proc Natl Acad Sci U S A. 2015 Sep 22; 112(38):11829-34. Young NM, Capellini TD, Roach NT, Alemseged Z. PMID: 26351685; PMCID: PMC4586873.
      View in: PubMed   Mentions: 14     Fields:    Translation:HumansAnimals
    21. Signals from the brain induce variation in avian facial shape. Dev Dyn. 2015 Sep; 244(9):1133-1143. Hu D, Young NM, Xu Q, Jamniczky H, Green RM, Mio W, Marcucio RS, Hallgrimsson B. PMID: 25903813; PMCID: PMC4618284.
      View in: PubMed   Mentions: 31     Fields:    
    22. Divergence of craniofacial developmental trajectories among avian embryos. Dev Dyn. 2015 Sep; 244(9):1158-1167. Smith FJ, Percival CJ, Young NM, Hu D, Schneider RA, Marcucio RS, Hallgrimsson B. PMID: 25703037; PMCID: PMC4544654.
      View in: PubMed   Mentions: 18     Fields:    
    23. Shared rules of development predict patterns of evolution in vertebrate segmentation. Nat Commun. 2015 Apr 01; 6:6690. Young NM, Winslow B, Takkellapati S, Kavanagh K. PMID: 25827599.
      View in: PubMed   Mentions: 9     Fields:    Translation:HumansAnimals
    24. A dynamic Shh expression pattern, regulated by SHH and BMP signaling, coordinates fusion of primordia in the amniote face. Development. 2015 Feb 01; 142(3):567-74. Hu D, Young NM, Li X, Xu Y, Hallgrímsson B, Marcucio RS. PMID: 25605783; PMCID: PMC4302993.
      View in: PubMed   Mentions: 28     Fields:    Translation:AnimalsCells
    25. Embryonic bauplans and the developmental origins of facial diversity and constraint. Development. 2014 Mar; 141(5):1059-63. Young NM, Hu D, Lainoff AJ, Smith FJ, Diaz R, Tucker AS, Trainor PA, Schneider RA, Hallgrímsson B, Marcucio RS. PMID: 24550113; PMCID: PMC3929406.
      View in: PubMed   Mentions: 57     Fields:    Translation:HumansAnimals
    26. Macroevolutionary diversity of amniote limb proportions predicted by developmental interactions. J Exp Zool B Mol Dev Evol. 2013 Nov; 320(7):420-7. Young NM. PMID: 23913472.
      View in: PubMed   Mentions: 3     Fields:    Translation:Animals
    27. Quantification of shape and cell polarity reveals a novel mechanism underlying malformations resulting from related FGF mutations during facial morphogenesis. Hum Mol Genet. 2013 Dec 20; 22(25):5160-72. Li X, Young NM, Tropp S, Hu D, Xu Y, Hallgrímsson B, Marcucio RS. PMID: 23906837; PMCID: PMC3842176.
      View in: PubMed   Mentions: 15     Fields:    Translation:HumansAnimalsCells
    28. The effect of hypoxia on facial shape variation and disease phenotypes in chicken embryos. Dis Model Mech. 2013 Jul; 6(4):915-24. Smith F, Hu D, Young NM, Lainoff AJ, Jamniczky HA, Maltepe E, Hallgrimsson B, Marcucio RS. PMID: 23592613; PMCID: PMC3701211.
      View in: PubMed   Mentions: 10     Fields:    Translation:Animals
    29. Embryonic origins of novelty and constraint in the amniote upper jaw. The FASEB Journal. 2013 Apr 1; 27:319.3-319.3. Young YN, Hu HD, Lainoff LA, Hallgrímsson HB, Marcucio MR. .
      View in: Publisher Site   Mentions:
    30. The generation of variation and the developmental basis for evolutionary novelty. J Exp Zool B Mol Dev Evol. 2012 Sep; 318(6):501-17. Hallgrímsson B, Jamniczky HA, Young NM, Rolian C, Schmidt-Ott U, Marcucio RS. PMID: 22649039; PMCID: PMC3648206.
      View in: PubMed   Mentions: 16     Fields:    Translation:Animals
    31. A Novel Gene Crispld2 may Contribute to Facial Dysmophology in a Chicken Model of Crouzon's Syndrome. The FASEB Journal. 2012 Apr 1; 26:907.15-907.15. Li LX, Tropp TS, Young YN, Hallgrimsson HB, Marcucio MR. .
      View in: Publisher Site   Mentions:
    32. Tissue Interactions that Regulate Facial Morphogenesis. The FASEB Journal. 2012 Apr 1; 26:337.2-337.2. Marcucio MR, Young YN, Hu HD, Hallgrimsson HB. .
      View in: Publisher Site   Mentions:
    33. The Phenogenomics of Craniofacial Shape. The FASEB Journal. 2012 Apr 1; 26:337.4-337.4. Hallgrimsson HB, Jamniczky JH, Martínez-Abadías MN, Young YN, Marcucio MR. .
      View in: Publisher Site   Mentions:
    34. Signaling by SHH rescues facial defects following blockade in the brain. Dev Dyn. 2012 Feb; 241(2):247-56. Chong HJ, Young NM, Hu D, Jeong J, McMahon AP, Hallgrimsson B, Marcucio RS. PMID: 22275045; PMCID: PMC3547623.
      View in: PubMed   Mentions: 30     Fields:    Translation:AnimalsCells
    35. Mechanisms that underlie co-variation of the brain and face. Genesis. 2011 Apr; 49(4):177-89. Marcucio RS, Young NM, Hu D, Hallgrimsson B. PMID: 21381182; PMCID: PMC3086711.
      View in: PubMed   Mentions: 68     Fields:    Translation:HumansAnimalsCells
    36. Quantitative analyses link modulation of sonic hedgehog signaling to continuous variation in facial growth and shape. Development. 2010 Oct; 137(20):3405-9. Young NM, Chong HJ, Hu D, Hallgrímsson B, Marcucio RS. PMID: 20826528; PMCID: PMC2947484.
      View in: PubMed   Mentions: 73     Fields:    Translation:AnimalsCells
    37. Development and the evolvability of human limbs. Proc Natl Acad Sci U S A. 2010 Feb 23; 107(8):3400-5. Young NM, Wagner GP, Hallgrímsson B. PMID: 20133636; PMCID: PMC2840520.
      View in: PubMed   Mentions: 41     Fields:    Translation:HumansAnimals
    38. Comparative gene expression analysis of avian embryonic facial structures reveals new candidates for human craniofacial disorders. Hum Mol Genet. 2010 Mar 01; 19(5):920-30. Brugmann SA, Powder KE, Young NM, Goodnough LH, Hahn SM, James AW, Helms JA, Lovett M. PMID: 20015954; PMCID: PMC2816616.
      View in: PubMed   Mentions: 32     Fields:    Translation:HumansAnimals
    39. Deciphering the Palimpsest: Studying the Relationship Between Morphological Integration and Phenotypic Covariation. Evol Biol. 2009 Dec; 36(4):355-376. Hallgrímsson B, Jamniczky H, Young NM, Rolian C, Parsons TE, Boughner JC, Marcucio RS. PMID: 23293400; PMCID: PMC3537827.
      View in: PubMed   Mentions: 124  
    40. The relationship between variable SHH signaling and the severity of structural defects in the face and brain. The FASEB Journal. 2009 Apr 1; 23:180.5-180.5. Young YN, Hu HD, Chong CH, Hallgrímsson HB, Liu LW, Marcucio MR. .
      View in: Publisher Site   Mentions:
    41. Epigenetic Effects on Integration of Limb Lengths in a Mouse Model: Selective Breeding for High Voluntary Locomotor Activity. Evolutionary Biology. 2009 Mar 1; 36(1):88. Young YN, Hallgrímsson HB, Garland GT. .
      View in: Publisher Site   Mentions:
    42. Short-faced mice and developmental interactions between the brain and the face. J Anat. 2008 Dec; 213(6):646-62. Boughner JC, Wat S, Diewert VM, Young NM, Browder LW, Hallgrímsson B. PMID: 19094181; PMCID: PMC2666134.
      View in: PubMed   Mentions: 31     Fields:    Translation:Animals
    43. Timing primate evolution: Lessons from the discordance between molecular and paleontological estimates. Evolutionary Anthropology Issues News and Reviews. 2008 Jul 1; 17(4):179-188. Steiper SM, Young YN. .
      View in: Publisher Site   Mentions:
    44. A comparison of the ontogeny of shape variation in the anthropoid scapula: functional and phylogenetic signal. Am J Phys Anthropol. 2008 Jul; 136(3):247-64. Young NM. PMID: 18324623.
      View in: PubMed   Mentions: 12     Fields:    Translation:Animals
    45. Morphometrics and Biological Anthropology in the Postgenomic Age. Biological Anthropology of the Human Skeleton. 2008 Mar 21; 207-235. Hallgrímsson HB, Zelditch ZM, Parsons PT, Kristensen KE, Young YN, Boyd BS. .
      View in: Publisher Site   Mentions:
    46. Phenotypic Variability: Its Components, Measurement and Underlying Developmental Processes. Evolutionary Biology. 2007 Dec 1; 34(3-4):99-120. Willmore WK, Young YN, Richtsmeier RJ. .
      View in: Publisher Site   Mentions:
    47. Geometric morphometric analysis of craniofacial deformity in the noggin mutant. Journal of the American College of Surgeons. 2007 Sep 1; 205(3):s60. Gupta GD, Young YN, Wan WD, Brunet BL, Harland HR, Helms HJ, Longaker LM. .
      View in: Publisher Site   Mentions:
    48. Evolution of covariance in the mammalian skull. Novartis Found Symp. 2007; 284:164-85; discussion 185-90. Hallgrimsson B, Lieberman DE, Young NM, Parsons T, Wat S. PMID: 17710853.
      View in: PubMed   Mentions: 17     Fields:    Translation:Animals
    49. Comparative morphometrics of embryonic facial morphogenesis: implications for cleft-lip etiology. Anat Rec (Hoboken). 2007 Jan; 290(1):123-39. Young NM, Wat S, Diewert VM, Browder LW, Hallgrímsson B. PMID: 17441205.
      View in: PubMed   Mentions: 22     Fields:    Translation:Animals
    50. Function, ontogeny and canalization of shape variance in the primate scapula. J Anat. 2006 Nov; 209(5):623-36. Young NM. PMID: 17062020; PMCID: PMC2100339.
      View in: PubMed   Mentions: 27     Fields:    Translation:Animals
    51. Primate molecular divergence dates. Mol Phylogenet Evol. 2006 Nov; 41(2):384-94. Steiper ME, Young NM. PMID: 16815047.
      View in: PubMed   Mentions: 130     Fields:    Translation:HumansAnimalsCells
    52. Canalization and developmental stability in the Brachyrrhine mouse. J Anat. 2006 Mar; 208(3):361-72. Willmore KE, Zelditch ML, Young N, Ah-Seng A, Lozanoff S, Hallgrímsson B. PMID: 16533318; PMCID: PMC2100242.
      View in: PubMed   Mentions: 12     Fields:    Translation:Animals
    53. Serial homology and the evolution of mammalian limb covariation structure. Evolution. 2005 Dec; 59(12):2691-704. Young NM, Hallgrímsson B. PMID: 16526515.
      View in: PubMed   Mentions: 65     Fields:    Translation:Animals
    54. SERIAL HOMOLOGY AND THE EVOLUTION OF MAMMALIAN LIMB COVARIATION STRUCTURE. Evolution. 2005 Jan 1; 59(12):2691-2704. Young YN, Hallgrmsson HB. .
      View in: Publisher Site   Mentions:
    55. Genomic data support the hominoid slowdown and an Early Oligocene estimate for the hominoid-cercopithecoid divergence. Proc Natl Acad Sci U S A. 2004 Dec 07; 101(49):17021-6. Steiper ME, Young NM, Sukarna TY. PMID: 15572456; PMCID: PMC535389.
      View in: PubMed   Mentions: 40     Fields:    Translation:HumansAnimals
    56. Modularity and integration in the hominoid scapula. J Exp Zool B Mol Dev Evol. 2004 May 15; 302(3):226-40. Young N. PMID: 15211684.
      View in: PubMed   Mentions: 8     Fields:    Translation:Animals
    57. The phylogenetic position of Morotopithecus. J Hum Evol. 2004 Feb; 46(2):163-84. Young NM, MacLatchy L. PMID: 14871561.
      View in: PubMed   Mentions: 13     Fields:    Translation:HumansAnimals
    58. A reassessment of living hominoid postcranial variability: implications for ape evolution. J Hum Evol. 2003 Dec; 45(6):441-64. Young NM. PMID: 14643673.
      View in: PubMed   Mentions: 13     Fields:    Translation:HumansAnimals
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