Nathan Young, PhD

Nathan Young, PhD

Nathan Young, PhD

Associate Professor

Basic Science

Trauma & Problem Fractures

Orthopaedic Trauma Institute - Young Lab

2550 23rd Street
San Francisco CA 94110

628-206-8812

Orthopaedic Trauma Institute at Zuckerberg San Francisco General - Trauma & Problem Fractures

2540 23rd Street,
San Francisco CA 94110

4154762124
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Publications

Hassan MG, Kaler H, Zhang B, Cox TC, Young N, Jheon AH. Effects of Multi-Generational Soft Diet Consumption on Mouse Craniofacial Morphology. Front Physiol. 2020; 11:783. PMID: 32754047

Lee ECS, Roach NT, Clouthier AL, Bicknell RT, Bey MJ, Young NM, Rainbow MJ. 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. PMID: 32580097

Zaid MB, Young NM, Pedoia V, Feeley BT, Ma CB, Lansdown DA. Radiographic shoulder parameters and their relationship to outcomes following rotator cuff repair: a systematic review. Shoulder Elbow. 2021 Aug; 13(4):371-379. PMID: 34394734

Hallgrímsson B, Katz DC, Aponte JD, Larson JR, Devine J, Gonzalez PN, Young NM, Roseman CC, Marcucio RS. Integration and the Developmental Genetics of Allometry. Integr Comp Biol. 2019 11 01; 59(5):1369-1381. PMID: 31199435

Zaid MB, Young NM, Pedoia V, Feeley BT, Ma CB, Lansdown DA. 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. PMID: 31353303

Ralph Marcucio, Diane Hu, Benedikt Hallgrimsson, Nathan Young. Emergent Properties of Facial Morphogenesis Regulated by Fgf Signaling. The FASEB Journal. 2019 Apr 1; 33(S1):774.18-774.18. PMID:

Benedikt Hallgrimsson, Jose David Aponte, David C Katz, Jordan J Bannister, Rebecca M Green, Nathan M Young, James M Cheverud, Nils Daniel Forkert, Ralph S Marcucio. Integration and the genetics of variation in facial shape. The FASEB Journal. 2019 Apr 1; 33(S1):330.2-330.2. PMID:

Heather A Richbourg, Nathan Young, Risa Takenaka, Ralph Marcucio. Quantifying the Genotype to Phenotype Map in Developing Mice. The FASEB Journal. 2018 Apr 1; 32:lb529-lb529. PMID:

Alexis Lainoff, Nathan Young, Benedikt Hallgrimsson, Ralph Marcucio. Untangling Sources of Phenotypic Variation Characterizing the Craniofacial Disease Holoprosencephaly. The FASEB Journal. 2018 Apr 1; 32:776.11-776.11. PMID:

Young NM, Linde-Medina M, Fondon JW, Hallgrímsson B, Marcucio RS. Craniofacial diversification in the domestic pigeon and the evolution of the avian skull. Nat Ecol Evol. 2017 Mar 13; 1(4):95. PMID: 28812673

Nathan M. Young, Terence D. Capellini. Out on a Limb. Developmental Approaches to Human Evolution. 2016 Jan 29; 101-137. PMID:

Young NM, Capellini TD, Roach NT, Alemseged Z. 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. PMID: 26676585

Hallgrimsson B, Percival CJ, Green R, Young NM, Mio W, Marcucio R. Morphometrics, 3D Imaging, and Craniofacial Development. Curr Top Dev Biol. 2015; 115:561-97. PMID: 26589938

Marcucio R, Hallgrimsson B, Young NM. Facial Morphogenesis: Physical and Molecular Interactions Between the Brain and the Face. Curr Top Dev Biol. 2015; 115:299-320. PMID: 26589930

Hu D, Young NM, Xu Q, Jamniczky H, Green RM, Mio W, Marcucio RS, Hallgrimsson B. Signals from the brain induce variation in avian facial shape. Dev Dyn. 2015 Sep; 244(9):1133-1143. PMID: 25903813

Smith FJ, Percival CJ, Young NM, Hu D, Schneider RA, Marcucio RS, Hallgrimsson B. Divergence of craniofacial developmental trajectories among avian embryos. Dev Dyn. 2015 Sep; 244(9):1158-1167. PMID: 25703037

Young NM. Macroevolutionary diversity of amniote limb proportions predicted by developmental interactions. J Exp Zool B Mol Dev Evol. 2013 Nov; 320(7):420-7. PMID: 23913472

Smith F, Hu D, Young NM, Lainoff AJ, Jamniczky HA, Maltepe E, Hallgrimsson B, Marcucio RS. The effect of hypoxia on facial shape variation and disease phenotypes in chicken embryos. Dis Model Mech. 2013 Jul; 6(4):915-24. PMID: 23592613

Nathan Young, Diane Hu, Alexis Lainoff, Benedikt Hallgrímsson, Ralph Marcucio. Embryonic origins of novelty and constraint in the amniote upper jaw. The FASEB Journal. 2013 Apr 1; 27:319.3-319.3. PMID:

Xin Li, Stephen Tropp, Nathan Young, Benedikt Hallgrimsson, Ralph Marcucio. 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. PMID:

Benedikt Hallgrimsson, Heather A. Jamniczky, Neus Martínez-Abadías, Nathan M. Young, Ralph S. Marcucio. The Phenogenomics of Craniofacial Shape. The FASEB Journal. 2012 Apr 1; 26:337.4-337.4. PMID:

Ralph Marcucio, Nathan Young, Diane Hu, Benedikt Hallgrimsson. Tissue Interactions that Regulate Facial Morphogenesis. The FASEB Journal. 2012 Apr 1; 26:337.2-337.2. PMID:

Chong HJ, Young NM, Hu D, Jeong J, McMahon AP, Hallgrimsson B, Marcucio RS. Signaling by SHH rescues facial defects following blockade in the brain. Dev Dyn. 2012 Feb; 241(2):247-56. PMID: 22275045

Marcucio RS, Young NM, Hu D, Hallgrimsson B. Mechanisms that underlie co-variation of the brain and face. Genesis. 2011 Apr; 49(4):177-89. PMID: 21381182

Young NM, Wagner GP, Hallgrímsson B. Development and the evolvability of human limbs. Proc Natl Acad Sci U S A. 2010 Feb 23; 107(8):3400-5. PMID: 20133636

Brugmann SA, Powder KE, Young NM, Goodnough LH, Hahn SM, James AW, Helms JA, Lovett M. 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. PMID: 20015954

Nathan M. Young, Diane Hu, H. Jonathan Chong, Benedikt Hallgrímsson, Wei Liu, Ralph Marcucio. 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. PMID:

Nathan M. Young, Benedikt Hallgrímsson, Theodore Garland. 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. PMID:

Boughner JC, Wat S, Diewert VM, Young NM, Browder LW, Hallgrímsson B. Short-faced mice and developmental interactions between the brain and the face. J Anat. 2008 Dec; 213(6):646-62. PMID: 19094181

Young NM. 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. PMID: 18324623

M. E. Steiper, N. M. Young. 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. PMID:

Benedikt Hallgrímsson, Miriam Leah Zelditch, Trish E. Parsons, Erika Kristensen, Nathan M. Young, Steven K. Boyd. Morphometrics and Biological Anthropology in the Postgenomic Age. Biological Anthropology of the Human Skeleton. 2008 Mar 21; 207-235. PMID:

Katherine Elizabeth Willmore, Nathan M. Young, Joan T. Richtsmeier. Phenotypic Variability: Its Components, Measurement and Underlying Developmental Processes. Evolutionary Biology. 2007 Dec 1; 34(3-4):99-120. PMID:

Deepak M. Gupta, Nathan M. Young, Derrick C. Wan, Lisa J. Brunet, Richard M. Harland, Jill A. Helms, Michael T. Longaker. Geometric morphometric analysis of craniofacial deformity in the noggin mutant. Journal of the American College of Surgeons. 2007 Sep 1; 205(3):s60. PMID:

Young NM, Wat S, Diewert VM, Browder LW, Hallgrímsson B. Comparative morphometrics of embryonic facial morphogenesis: implications for cleft-lip etiology. Anat Rec (Hoboken). 2007 Jan; 290(1):123-39. PMID: 17441205

Hallgrimsson B, Lieberman DE, Young NM, Parsons T, Wat S. Evolution of covariance in the mammalian skull. Novartis Found Symp. 2007; 284:164-85; discussion 185-90. PMID: 17710853

Young NM. Function, ontogeny and canalization of shape variance in the primate scapula. J Anat. 2006 Nov; 209(5):623-36. PMID: 17062020

Steiper ME, Young NM. Primate molecular divergence dates. Mol Phylogenet Evol. 2006 Nov; 41(2):384-94. PMID: 16815047

Willmore KE, Zelditch ML, Young N, Ah-Seng A, Lozanoff S, Hallgrímsson B. Canalization and developmental stability in the Brachyrrhine mouse. J Anat. 2006 Mar; 208(3):361-72. PMID: 16533318

Young NM, Hallgrímsson B. Serial homology and the evolution of mammalian limb covariation structure. Evolution. 2005 Dec; 59(12):2691-704. PMID: 16526515

Nathan M. Young, Benedikt Hallgrmsson. SERIAL HOMOLOGY AND THE EVOLUTION OF MAMMALIAN LIMB COVARIATION STRUCTURE. Evolution. 2005 Jan 1; 59(12):2691-2704. PMID:

Steiper ME, Young NM, Sukarna TY. 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. PMID: 15572456

Young N. Modularity and integration in the hominoid scapula. J Exp Zool B Mol Dev Evol. 2004 May 15; 302(3):226-40. PMID: 15211684

Young NM, MacLatchy L. The phylogenetic position of Morotopithecus. J Hum Evol. 2004 Feb; 46(2):163-84. PMID: 14871561

Young NM. Integrating "Evo" and "Devo": The Limb as Model Structure. Integr Comp Biol. 2017 12 01; 57(6):1293-1302. PMID: 28992180

Young NM, Marcucio RS, Hallgrímsson B, Richbourg HA, Green R. 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. PMID:

Zuzana Vavrušová, Diane Hu, Wei Liu, Ralph S. Marcucio, Benedikt Hallgrímsson, Nathan M. Young. Developmental Morphospace of Embryonic Brain and Face Growth in the Chicken (Gallus gallus). The FASEB Journal. 2022 May 13; 36(S1). PMID:

Scott A. Williams, Thomas Cody Prang, Gabrielle A. Russo, Nathan M. Young, Daniel L. Gebo. African apes and the evolutionary history of orthogrady and bipedalism. American Journal of Biological Anthropology. 2023 Jan 13. PMID:

Hallgrímsson B, Jamniczky HA, Young NM, Rolian C, Schmidt-Ott U, Marcucio RS. The generation of variation and the developmental basis for evolutionary novelty. J Exp Zool B Mol Dev Evol. 2012 Sep; 318(6):501-17. PMID: 22649039

Young NM, Chong HJ, Hu D, Hallgrímsson B, Marcucio RS. Quantitative analyses link modulation of sonic hedgehog signaling to continuous variation in facial growth and shape. Development. 2010 Oct; 137(20):3405-9. PMID: 20826528

Hu D, Young NM, Li X, Xu Y, Hallgrímsson B, Marcucio RS. 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. PMID: 25605783

Young NM, Hu D, Lainoff AJ, Smith FJ, Diaz R, Tucker AS, Trainor PA, Schneider RA, Hallgrímsson B, Marcucio RS. Embryonic bauplans and the developmental origins of facial diversity and constraint. Development. 2014 Mar; 141(5):1059-63. PMID: 24550113

Li X, Young NM, Tropp S, Hu D, Xu Y, Hallgrímsson B, Marcucio RS. 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. PMID: 23906837

Hallgrímsson B, Jamniczky H, Young NM, Rolian C, Parsons TE, Boughner JC, Marcucio RS. Deciphering the Palimpsest: Studying the Relationship Between Morphological Integration and Phenotypic Covariation. Evol Biol. 2009 Dec; 36(4):355-376. PMID: 23293400

Young NM, Sherathiya K, Gutierrez L, Nguyen E, Bekmezian S, Huang JC, Hallgrímsson B, Lee JS, Marcucio RS. Facial surface morphology predicts variation in internal skeletal shape. Am J Orthod Dentofacial Orthop. 2016 Apr; 149(4):501-8. PMID: 27021454

Young NM, Capellini TD, Roach NT, Alemseged Z. 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. PMID: 26862164

Marchini M, Hu D, Lo Vercio L, Young NM, Forkert ND, Hallgrímsson B, Marcucio R. Wnt Signaling Drives Correlated Changes in Facial Morphology and Brain Shape. Front Cell Dev Biol. 2021; 9:644099. PMID: 33855022

Benedikt Hallgrímsson, J. David Aponte, Marta Vidal-Garcia, Heather Richbourg, Rebecca Green, Nathan M. Young, James M. Cheverud, Anne L. Calof, Arthur D. Lander, Ralph S. Marcucio. The Developmental Basis for Evolvability. Evolvability. 2023 Jun 27; 171-198. PMID:

Marta Linde-Medina, John Fondon, Ralph Marcucio, Benedikt Hallgrímsson, Nathan M. Young. Evolvability and the Diversification of the Craniofacial Skeleton in the Domesticated Pigeon ( Columba livia ). The FASEB Journal. 2016 Apr 1; 30(S1). PMID:

Richbourg HA, Vidal-García M, Brakora KA, Devine J, Takenaka R, Young NM, Gong SG, Neves A, Hallgrímsson B, Marcucio RS. Dosage-dependent effects of FGFR2W290R mutation on craniofacial shape and cellular dynamics of the basicranial synchondroses. Anat Rec (Hoboken). 2024 Feb 26. PMID: 38409943

Lee ECS, Young NM, Rainbow MJ. A comparative approach for characterizing the relationship among morphology, range-of-motion and locomotor behaviour in the primate shoulder. Proc Biol Sci. 2023 Oct 25; 290(2009):20231446. PMID: 37848066

Musa B Zaid, Nathan M Young, Valentina Pedoia, Brian T Feeley, C Benjamin Ma, Drew A Lansdown. Radiographic shoulder parameters and their relationship to outcomes following rotator cuff repair: a systematic review. Shoulder & Elbow. 2020 Jan 10; 175857321989598. PMID:

Young NM, Capellini TD, Roach NT, Alemseged Z. 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. PMID: 26351685

Young NM, Winslow B, Takkellapati S, Kavanagh K. Shared rules of development predict patterns of evolution in vertebrate segmentation. Nat Commun. 2015 Apr 01; 6:6690. PMID: 25827599

Young NM. A reassessment of living hominoid postcranial variability: implications for ape evolution. J Hum Evol. 2003 Dec; 45(6):441-64. PMID: 14643673

Grants & Awards

A Predictive Developmental Morphospace Model of Cleft Lip
2020-09-03 - 2021-08-31
NIH R56DE029124
Role: Principal Investigator

The developmental genetic basis for evolutionary variation in the hominin shoulder
2015-07-01 - 2018-06-30
NSF BCS-1518596
Role: Co-Investigator

A Molecular Investigation of Growth and Shape Variation in Cleft-Lip Etiology
2007-09-01 - 2009-08-31
NIH F32DE018596
Role: Principal Investigator

Education

	Postdoctoral Studies	University of California, San Francisco	2007
	Postdoctoral Studies	Stanford University	2006
	Postdoctoral Studies	University of Calgary	2004
	Ph.D.	Harvard University	2003
	B.A.	New College of Florida	1995

Videos

Science Speakeasy | Nathan Young | On the Shoulders of Apes

Bowl Science: Brains or Brawn?

About Nathan Young, PhD

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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