Partial Loss of USP9X Function Leads to a Male Neurodevelopmental and Behavioral Disorder Converging on Transforming Growth Factor β Signaling.


Journal

Biological psychiatry
ISSN: 1873-2402
Titre abrégé: Biol Psychiatry
Pays: United States
ID NLM: 0213264

Informations de publication

Date de publication:
15 01 2020
Historique:
received: 27 03 2019
revised: 23 05 2019
accepted: 30 05 2019
pubmed: 25 8 2019
medline: 7 1 2021
entrez: 25 8 2019
Statut: ppublish

Résumé

The X-chromosome gene USP9X encodes a deubiquitylating enzyme that has been associated with neurodevelopmental disorders primarily in female subjects. USP9X escapes X inactivation, and in female subjects de novo heterozygous copy number loss or truncating mutations cause haploinsufficiency culminating in a recognizable syndrome with intellectual disability and signature brain and congenital abnormalities. In contrast, the involvement of USP9X in male neurodevelopmental disorders remains tentative. We used clinically recommended guidelines to collect and interrogate the pathogenicity of 44 USP9X variants associated with neurodevelopmental disorders in males. Functional studies in patient-derived cell lines and mice were used to determine mechanisms of pathology. Twelve missense variants showed strong evidence of pathogenicity. We define a characteristic phenotype of the central nervous system (white matter disturbances, thin corpus callosum, and widened ventricles); global delay with significant alteration of speech, language, and behavior; hypotonia; joint hypermobility; visual system defects; and other common congenital and dysmorphic features. Comparison of in silico and phenotypical features align additional variants of unknown significance with likely pathogenicity. In support of partial loss-of-function mechanisms, using patient-derived cell lines, we show loss of only specific USP9X substrates that regulate neurodevelopmental signaling pathways and a united defect in transforming growth factor β signaling. In addition, we find correlates of the male phenotype in Usp9x brain-specific knockout mice, and further resolve loss of hippocampal-dependent learning and memory. Our data demonstrate the involvement of USP9X variants in a distinctive neurodevelopmental and behavioral syndrome in male subjects and identify plausible mechanisms of pathogenesis centered on disrupted transforming growth factor β signaling and hippocampal function.

Sections du résumé

BACKGROUND
The X-chromosome gene USP9X encodes a deubiquitylating enzyme that has been associated with neurodevelopmental disorders primarily in female subjects. USP9X escapes X inactivation, and in female subjects de novo heterozygous copy number loss or truncating mutations cause haploinsufficiency culminating in a recognizable syndrome with intellectual disability and signature brain and congenital abnormalities. In contrast, the involvement of USP9X in male neurodevelopmental disorders remains tentative.
METHODS
We used clinically recommended guidelines to collect and interrogate the pathogenicity of 44 USP9X variants associated with neurodevelopmental disorders in males. Functional studies in patient-derived cell lines and mice were used to determine mechanisms of pathology.
RESULTS
Twelve missense variants showed strong evidence of pathogenicity. We define a characteristic phenotype of the central nervous system (white matter disturbances, thin corpus callosum, and widened ventricles); global delay with significant alteration of speech, language, and behavior; hypotonia; joint hypermobility; visual system defects; and other common congenital and dysmorphic features. Comparison of in silico and phenotypical features align additional variants of unknown significance with likely pathogenicity. In support of partial loss-of-function mechanisms, using patient-derived cell lines, we show loss of only specific USP9X substrates that regulate neurodevelopmental signaling pathways and a united defect in transforming growth factor β signaling. In addition, we find correlates of the male phenotype in Usp9x brain-specific knockout mice, and further resolve loss of hippocampal-dependent learning and memory.
CONCLUSIONS
Our data demonstrate the involvement of USP9X variants in a distinctive neurodevelopmental and behavioral syndrome in male subjects and identify plausible mechanisms of pathogenesis centered on disrupted transforming growth factor β signaling and hippocampal function.

Identifiants

pubmed: 31443933
pii: S0006-3223(19)31479-9
doi: 10.1016/j.biopsych.2019.05.028
pmc: PMC6925349
mid: NIHMS1538027
pii:
doi:

Substances chimiques

Transforming Growth Factor beta 0
USP9X protein, human 0
Ubiquitin Thiolesterase EC 3.4.19.12
Usp9x protein, mouse EC 3.4.19.12

Types de publication

Journal Article Research Support, N.I.H., Extramural Research Support, Non-U.S. Gov't

Langues

eng

Sous-ensembles de citation

IM

Pagination

100-112

Subventions

Organisme : NHGRI NIH HHS
ID : U01 HG007672
Pays : United States
Organisme : NICHD NIH HHS
ID : U54 HD090255
Pays : United States
Organisme : Wellcome Trust
ID : WT098051
Pays : United Kingdom
Organisme : NIMH NIH HHS
ID : R01 MH107182
Pays : United States
Organisme : Wellcome Trust
Pays : United Kingdom

Investigateurs

Loren Pena (L)
Vandana Shashi (V)
Kelly Schoch (K)
Jennifer A Sullivan (JA)
Maria T Acosta (MT)
David R Adams (DR)
Aaron Aday (A)
Mercedes E Alejandro (ME)
Patrick Allard (P)
Euan A Ashley (EA)
Mahshid S Azamian (MS)
Carlos A Bacino (CA)
Guney Bademci (G)
Eva Baker (E)
Ashok Balasubramanyam (A)
Dustin Baldridge (D)
Deborah Barbouth (D)
Gabriel F Batzli (GF)
Alan H Beggs (AH)
Hugo J Bellen (HJ)
Jonathan A Bernstein (JA)
Gerard T Berry (GT)
Anna Bican (A)
David P Bick (DP)
Camille L Birch (CL)
Stephanie Bivona (S)
Carsten Bonnenmann (C)
Devon Bonner (D)
Braden E Boone (BE)
Bret L Bostwick (BL)
Lauren C Briere (LC)
Elly Brokamp (E)
Donna M Brown (DM)
Matthew Brush (M)
Elizabeth A Burke (EA)
Lindsay C Burrage (LC)
Manish J Butte (MJ)
Olveen Carrasquillo (O)
Ta Chen Peter Chang (TC)
Hsiao-Tuan Chao (HT)
Gary D Clark (GD)
Terra R Coakley (TR)
Laurel A Cobban (LA)
Joy D Cogan (JD)
F Sessions Cole (FS)
Heather A Colley (HA)
Cynthia M Cooper (CM)
Heidi Cope (H)
William J Craigen (WJ)
Precilla D'Souza (P)
Surendra Dasari (S)
Mariska Davids (M)
Jean M Davidson (JM)
Jyoti G Dayal (JG)
Esteban C Dell'Angelica (EC)
Shweta U Dhar (SU)
Naghmeh Dorrani (N)
Daniel C Dorset (DC)
Emilie D Douine (ED)
David D Draper (DD)
Annika M Dries (AM)
Laura Duncan (L)
David J Eckstein (DJ)
Lisa T Emrick (LT)
Christine M Eng (CM)
Gregory M Enns (GM)
Cecilia Esteves (C)
Tyra Estwick (T)
Liliana Fernandez (L)
Carlos Ferreira (C)
Elizabeth L Fieg (EL)
Paul G Fisher (PG)
Brent L Fogel (BL)
Irman Forghani (I)
Noah D Friedman (ND)
William A Gahl (WA)
Rena A Godfrey (RA)
Alica M Goldman (AM)
David B Goldstein (DB)
Jean-Philippe F Gourdine (JF)
Alana Grajewski (A)
Catherine A Groden (CA)
Andrea L Gropman (AL)
Melissa Haendel (M)
Rizwan Hamid (R)
Neil A Hanchard (NA)
Frances High (F)
Ingrid A Holm (IA)
Jason Hom (J)
Alden Huang (A)
Yong Huang (Y)
Rosario Isasi (R)
Fariha Jamal (F)
Yong-Hui Jiang (YH)
Jean M Johnston (JM)
Angela L Jones (AL)
Lefkothea Karaviti (L)
Emily G Kelley (EG)
David M Koeller (DM)
Isaac S Kohane (IS)
Jennefer N Kohler (JN)
Deborah Krakow (D)
Donna M Krasnewich (DM)
Susan Korrick (S)
Mary Koziura (M)
Joel B Krier (JB)
Jennifer E Kyle (JE)
Seema R Lalani (SR)
Byron Lam (B)
Brendan C Lanpher (BC)
Ian R Lanza (IR)
C Christopher Lau (CC)
Jozef Lazar (J)
Kimberly LeBlanc (K)
Brendan H Lee (BH)
Hane Lee (H)
Roy Levitt (R)
Shawn E Levy (SE)
Richard A Lewis (RA)
Sharyn A Lincoln (SA)
Pengfei Liu (P)
Xue Zhong Liu (XZ)
Sandra K Loo (SK)
Joseph Loscalzo (J)
Richard L Maas (RL)
Ellen F Macnamara (EF)
Calum A MacRae (CA)
Valerie V Maduro (VV)
Marta M Majcherska (MM)
May Christine V Malicdan (MCV)
Laura A Mamounas (LA)
Teri A Manolio (TA)
Thomas C Markello (TC)
Ronit Marom (R)
Martin G Martin (MG)
Julian A Martínez-Agosto (JA)
Shruti Marwaha (S)
Thomas May (T)
Jacob McCauley (J)
Allyn McConkie-Rosell (A)
Colleen E McCormack (CE)
Alexa T McCray (AT)
Jason D Merker (JD)
Thomas O Metz (TO)
Matthew Might (M)
Eva Morava-Kozicz (E)
Paolo M Moretti (PM)
Marie Morimoto (M)
John J Mulvihill (JJ)
David R Murdock (DR)
Avi Nath (A)
Stan F Nelson (SF)
J Scott Newberry (JS)
John H Newman (JH)
Sarah K Nicholas (SK)
Donna Novacic (D)
Devin Oglesbee (D)
James P Orengo (JP)
Stephen Pak (S)
J Carl Pallais (JC)
Christina Gs Palmer (CG)
Jeanette C Papp (JC)
Neil H Parker (NH)
John A Phillips (JA)
Jennifer E Posey (JE)
John H Postlethwait (JH)
Lorraine Potocki (L)
Barbara N Pusey (BN)
Genecee Renteri (G)
Chloe M Reuter (CM)
Lynette Rives (L)
Amy K Robertson (AK)
Lance H Rodan (LH)
Jill A Rosenfeld (JA)
Robb K Rowley (RK)
Ralph Sacco (R)
Jacinda B Sampson (JB)
Susan L Samson (SL)
Mario Saporta (M)
Judy Schaechter (J)
Timothy Schedl (T)
Daryl A Scott (DA)
Lisa Shakachite (L)
Prashant Sharma (P)
Kathleen Shields (K)
Jimann Shin (J)
Rebecca Signer (R)
Catherine H Sillari (CH)
Edwin K Silverman (EK)
Janet S Sinsheimer (JS)
Kevin S Smith (KS)
Lilianna Solnica-Krezel (L)
Rebecca C Spillmann (RC)
Joan M Stoler (JM)
Nicholas Stong (N)
David A Sweetser (DA)
Cecelia P Tamburro (CP)
Queenie K-G Tan (QK)
Mustafa Tekin (M)
Fred Telischi (F)
Willa Thorson (W)
Cynthia J Tifft (CJ)
Camilo Toro (C)
Alyssa A Tran (AA)
Tiina K Urv (TK)
Tiphanie P Vogel (TP)
Daryl M Waggott (DM)
Colleen E Wahl (CE)
Nicole M Walley (NM)
Chris A Walsh (CA)
Melissa Walker (M)
Jennifer Wambach (J)
Jijun Wan (J)
Lee-Kai Wang (LK)
Michael F Wangler (MF)
Patricia A Ward (PA)
Katrina M Waters (KM)
Bobbie-Jo M Webb-Robertson (BM)
Daniel Wegner (D)
Monte Westerfield (M)
Matthew T Wheeler (MT)
Anastasia L Wise (AL)
Lynne A Wolfe (LA)
Jeremy D Woods (JD)
Elizabeth A Worthey (EA)
Shinya Yamamoto (S)
John Yang (J)
Amanda J Yoon (AJ)
Guoyun Yu (G)
Diane B Zastrow (DB)
Chunli Zhao (C)
Stephan Zuchner (S)
William Gahl (W)

Informations de copyright

Copyright © 2019 Society of Biological Psychiatry. All rights reserved.

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Auteurs

Brett V Johnson (BV)

University of Adelaide and Robinson Research Institute, Adelaide, Australia.

Raman Kumar (R)

University of Adelaide and Robinson Research Institute, Adelaide, Australia.

Sabrina Oishi (S)

School of Biomedical Sciences, The University of Queensland, Brisbane, Australia.

Suzy Alexander (S)

Queensland Brain Institute, The University of Queensland, Brisbane, Australia; Queensland Centre for Mental Health Research, Wacol, Queensland, Australia.

Maria Kasherman (M)

School of Biomedical Sciences, The University of Queensland, Brisbane, Australia; Griffith Institute for Drug Discovery, Griffith University, Brisbane, Australia.

Michelle Sanchez Vega (MS)

Queensland Brain Institute, The University of Queensland, Brisbane, Australia.

Atma Ivancevic (A)

University of Adelaide and Robinson Research Institute, Adelaide, Australia; BioFrontiers Institute, University of Colorado Boulder, Boulder, Colorado.

Alison Gardner (A)

University of Adelaide and Robinson Research Institute, Adelaide, Australia.

Deepti Domingo (D)

University of Adelaide and Robinson Research Institute, Adelaide, Australia.

Mark Corbett (M)

University of Adelaide and Robinson Research Institute, Adelaide, Australia.

Euan Parnell (E)

Department of Physiology, Northwestern University Feinberg School of Medicine, Chicago, Illinois.

Sehyoun Yoon (S)

Department of Physiology, Northwestern University Feinberg School of Medicine, Chicago, Illinois.

Tracey Oh (T)

Department of Medical Genetics, British Columbia Women's Hospital and University of British Columbia, Vancouver, British Columbia, Canada.

Matthew Lines (M)

Children's Hospital of Eastern Ontario, Ottawa, Ontario, Canada.

Henrietta Lefroy (H)

Oxford Centre for Genomic Medicine, Oxford University Hospitals National Health Services Foundation Trust, Oxford, United Kingdom.

Usha Kini (U)

Oxford Centre for Genomic Medicine, Oxford University Hospitals National Health Services Foundation Trust, Oxford, United Kingdom.

Margot Van Allen (M)

Department of Medical Genetics, University of British Columbia, Vancouver, British Columbia, Canada.

Sabine Grønborg (S)

Center for Rare Diseases, Department of Pediatrics and Department of Clinical Genetics, University Hospital Copenhagen, Copenhagen, Denmark.

Sandra Mercier (S)

Service de Génétique Médicale, Centre Hospitalier Universitaire Nantes and l'Institut du Thorax, Institut National de la Santé et de la Recherche Médicale, Centre National de la Recherche Scientifique, Université de Nantes, Nantes, France.

Sébastien Küry (S)

Service de Génétique Médicale, Centre Hospitalier Universitaire Nantes and l'Institut du Thorax, Institut National de la Santé et de la Recherche Médicale, Centre National de la Recherche Scientifique, Université de Nantes, Nantes, France.

Stéphane Bézieau (S)

Service de Génétique Médicale, Centre Hospitalier Universitaire Nantes and l'Institut du Thorax, Institut National de la Santé et de la Recherche Médicale, Centre National de la Recherche Scientifique, Université de Nantes, Nantes, France.

Laurent Pasquier (L)

Service de Génétique Clinique, Centre de Référence Déficiences Intellectuelles de Causes Rares, Centre Hospitalier Universitaire Hôpital Sud, Rennes, France.

Martine Raynaud (M)

Centre Hospitalier Régional Universitaire de Tours, Service de Génétique, Unité Nixte de Recherche 1253, iBrain, Université de Tours, Institut National de la Santé et de la Recherche Médicale, Tours, France.

Alexandra Afenjar (A)

Groupe de Recherche Clinique No. 19, ConCer-LD, Département de Génétique, Assistance Publique-Hôpitaux de Paris, Hôpital Armand Trousseau, Centres de Référence Maladies Rares des Déficits Intellectuels de Causes Rares, Paris, France.

Thierry Billette de Villemeur (T)

Sorbonne Université, Groupe de Recherche Clinique No. 19, ConCer-LD, Neuropédiatrie, Centres de Référence Maladies Rares Neurogénétique, Institut National de la Santé et de la Recherche Médicale, Assistance Publique-Hôpitaux de Paris, Hôpital Armand Trousseau, Paris, France.

Boris Keren (B)

Hôpital de la Pitié-Salpêtrière, Département de Génétique, Paris, France.

Julie Désir (J)

Erasme University Hospital, Université Libre de Bruxelles, Brussels, Belgium.

Lionel Van Maldergem (L)

Centre de Génétique Humaine, Université de Franche-Comté, Besançon, France.

Martina Marangoni (M)

Erasme University Hospital, Université Libre de Bruxelles, Brussels, Belgium.

Nicola Dikow (N)

Institute of Human Genetics, Heidelberg University, Heidelberg, Germany.

David A Koolen (DA)

Department of Human Genetics, Radboud University Medical Center, Nijmegen, The Netherlands.

Peter M VanHasselt (PM)

Department of Metabolic Diseases, University Medical Center Utrecht, Utrecht, The Netherlands.

Marjan Weiss (M)

Department of Clinical Genetics, Vrije Universiteit University Medical Center, Amsterdam, The Netherlands.

Petra Zwijnenburg (P)

Medical Genetics Unit, Hospital Pediátrico, Centro Hospitalar e Universitário de Coimbra, Coimbra, Portugal.

Joaquim Sa (J)

Medical Genetics Unit, Hospital Pediátrico, Centro Hospitalar e Universitário de Coimbra, Coimbra, Portugal.

Claudia Falcao Reis (CF)

Medical Genetics Unit, Hospital Pediátrico, Centro Hospitalar e Universitário de Coimbra, Coimbra, Portugal.

Carlos López-Otín (C)

Departamento de Bioquímica y Biología Molecular, Instituto Universitário de Oncología del Principado de Asturias, Universidad de Oviedo, Oviedo, Spain; Centro de Investigación Biomédica en Red de Cáncer, Spain.

Olaya Santiago-Fernández (O)

Departamento de Bioquímica y Biología Molecular, Instituto Universitário de Oncología del Principado de Asturias, Universidad de Oviedo, Oviedo, Spain.

Alberto Fernández-Jaén (A)

Unidad de Neurología Infantil, Hospital Universitário Quirón Madrid, Madrid, Spain.

Anita Rauch (A)

Institute of Medical Genetics, University of Zurich, Schlieren, Switzerland.

Katharina Steindl (K)

Institute of Medical Genetics, University of Zurich, Schlieren, Switzerland.

Pascal Joset (P)

Institute of Medical Genetics, University of Zurich, Schlieren, Switzerland.

Amy Goldstein (A)

Children's Hospital of Philadelphia, Philadelphia, Pennsylvania.

Suneeta Madan-Khetarpal (S)

Children's Hospital of Pittsburgh, Pittsburgh, Pennsylvania.

Elena Infante (E)

Children's Hospital of Pittsburgh, Pittsburgh, Pennsylvania.

Elaine Zackai (E)

Children's Hospital of Philadelphia, Philadelphia, Pennsylvania.

Carey Mcdougall (C)

Children's Hospital of Philadelphia, Philadelphia, Pennsylvania.

Vinodh Narayanan (V)

Center for Rare Childhood Disorders, Translational Genomics Research Institute, Phoenix, Arizona.

Keri Ramsey (K)

Center for Rare Childhood Disorders, Translational Genomics Research Institute, Phoenix, Arizona.

Saadet Mercimek-Andrews (S)

Division of Clinical and Metabolic Genetics, Department of Pediatrics, University of Toronto, The Hospital for Sick Children, Toronto, Ontario, Canada.

Loren Pena (L)

Division of Human Genetics, Cincinnati Children's Hospital; Department of Pediatrics, University of Cincinnati College of Medicine, Cincinnati, Ohio.

Vandana Shashi (V)

Department of Pediatrics, Division of Medical Genetics, Duke University Medical Center, Durham, North Carolina.

Kelly Schoch (K)

Department of Pediatrics, Division of Medical Genetics, Duke University Medical Center, Durham, North Carolina.

Jennifer A Sullivan (JA)

Department of Pediatrics, Division of Medical Genetics, Duke University Medical Center, Durham, North Carolina.

Filippo Pinto E Vairo (F)

Department of Clinical Genomics, Mayo Clinic, Rochester, Minnesota; Center for Individualized Medicine, Mayo Clinic, Rochester, Minnesota.

Pavel N Pichurin (PN)

Department of Clinical Genomics, Mayo Clinic, Rochester, Minnesota.

Sarah A Ewing (SA)

Department of Clinical Genomics, Mayo Clinic, Rochester, Minnesota.

Sarah S Barnett (SS)

Department of Laboratory Medicine and Pathology, Mayo Clinic, Rochester, Minnesota.

Eric W Klee (EW)

Department of Clinical Genomics, Mayo Clinic, Rochester, Minnesota.

M Scott Perry (MS)

Jane and John Justin Neuroscience Center, Cook Children's Medical Center, Fort Worth, Texas.

Mary Kay Koenig (MK)

Department of Pediatrics, University of Texas Medical School at Houston, Houston, Texas.

Catherine E Keegan (CE)

Division of Genetics, Department of Pediatrics, University of Michigan, Ann Arbor, Michigan.

Jane L Schuette (JL)

Division of Genetics, Department of Pediatrics, University of Michigan, Ann Arbor, Michigan.

Stephanie Asher (S)

Translational Medicine & Human Genetics, Hospital of the University of Pennsylvania, Philadelphia, Pennsylvania.

Yezmin Perilla-Young (Y)

Division of Pediatric Genetics and Metabolism, University of North Carolina, Chapel Hill, North Carolina.

Laurie D Smith (LD)

Division of Pediatric Genetics and Metabolism, University of North Carolina, Chapel Hill, North Carolina.

Jill A Rosenfeld (JA)

Baylor College of Medicine, Houston, Texas.

Elizabeth Bhoj (E)

Children's Hospital of Philadelphia, Philadelphia, Pennsylvania.

Paige Kaplan (P)

Children's Hospital of Philadelphia, Philadelphia, Pennsylvania.

Dong Li (D)

Children's Hospital of Philadelphia, Philadelphia, Pennsylvania.

Renske Oegema (R)

Department of Genetics, University Medical Center Utrecht, Utrecht, The Netherlands.

Ellen van Binsbergen (E)

Department of Genetics, University Medical Center Utrecht, Utrecht, The Netherlands.

Bert van der Zwaag (B)

Department of Genetics, University Medical Center Utrecht, Utrecht, The Netherlands.

Marie Falkenberg Smeland (MF)

Department of Medical Genetics, University Hospital of North Norway, Tromsø, Norway.

Ioana Cutcutache (I)

Translational Medicine, UCB Pharma, Slough, United Kingdom.

Matthew Page (M)

Translational Medicine, UCB Pharma, Braine-l'Alleud, Belgium.

Martin Armstrong (M)

Translational Medicine, UCB Pharma, Braine-l'Alleud, Belgium.

Angela E Lin (AE)

Medical Genetics Unit, Mass General Hospital for Children, Boston, Massachusetts.

Marcie A Steeves (MA)

Medical Genetics Unit, Mass General Hospital for Children, Boston, Massachusetts.

Nicolette den Hollander (ND)

Department of Clinical Genetics, Leiden University Medical Center, Leiden, The Netherlands.

Mariëtte J V Hoffer (MJV)

Department of Clinical Genetics, Leiden University Medical Center, Leiden, The Netherlands.

Margot R F Reijnders (MRF)

Department of Clinical Genetics, Maastricht University Medical Center, Maastricht, The Netherlands.

Serwet Demirdas (S)

Department of Clinical Genetics, Erasmus University Medical Center, Rotterdam, The Netherlands.

Daniel C Koboldt (DC)

Nationwide Children's Hospital, Columbus, Ohio.

Dennis Bartholomew (D)

Nationwide Children's Hospital, Columbus, Ohio.

Theresa Mihalic Mosher (TM)

Nationwide Children's Hospital, Columbus, Ohio.

Scott E Hickey (SE)

Department of Pediatrics, The Ohio State University College of Medicine, Columbus, Ohio.

Christine Shieh (C)

David Geffen School of Medicine, University of California-Los Angeles, California.

Pedro A Sanchez-Lara (PA)

Department of Pediatrics, Cedars-Sinai Medical Center, Los Angeles, California.

John M Graham (JM)

Department of Pediatrics, Cedars-Sinai Medical Center, Los Angeles, California.

Kamer Tezcan (K)

Department of Genetics, Kaiser Permanente, Sacramento, California.

G B Schaefer (GB)

Section of Genetics and Metabolism, University of Arkansas for Medical Sciences, Little Rock, Arkansas.

Noelle R Danylchuk (NR)

Department of Genetic Counseling, University of Arkansas for Medical Sciences, Little Rock, Arkansas.

Alexander Asamoah (A)

Department of Pediatrics, University of Louisville School of Medicine, Louisville, Kentucky.

Kelly E Jackson (KE)

Department of Pediatrics, University of Louisville School of Medicine, Louisville, Kentucky.

Naomi Yachelevich (N)

Clinical Genetics Services, Department of Pediatrics, New York University School of Medicine, New York, New York.

Margaret Au (M)

Department of Pediatrics, Cedars-Sinai Medical Center, Los Angeles, California.

Luis A Pérez-Jurado (LA)

University of Adelaide and Robinson Research Institute, Adelaide, Australia; Women's and Children's Hospital, Adelaide, Australia; South Australian Health and Medical Research Institute, Adelaide, South Australia; Hospital del Mar Research Institute, Network Research Centre for Rare Diseases and Universitat Pompeu Fabra, Barcelona, Spain.

Tjitske Kleefstra (T)

Department of Human Genetics, Radboud University Medical Center, Nijmegen, The Netherlands.

Peter Penzes (P)

Department of Physiology, Northwestern University Feinberg School of Medicine, Chicago, Illinois.

Stephen A Wood (SA)

Griffith Institute for Drug Discovery, Griffith University, Brisbane, Australia.

Thomas Burne (T)

Queensland Brain Institute, The University of Queensland, Brisbane, Australia; Queensland Centre for Mental Health Research, Wacol, Queensland, Australia.

Tyler Mark Pierson (TM)

Department of Pediatrics, Cedars-Sinai Medical Center, Los Angeles, California; Department of Neurology and the Board of Governors Regenerative Medicine Institute, Cedars-Sinai Medical Center, Los Angeles, California.

Michael Piper (M)

School of Biomedical Sciences, The University of Queensland, Brisbane, Australia; Queensland Brain Institute, The University of Queensland, Brisbane, Australia.

Jozef Gécz (J)

University of Adelaide and Robinson Research Institute, Adelaide, Australia; South Australian Health and Medical Research Institute, Adelaide, South Australia. Electronic address: Jozef.Gecz@adelaide.edu.au.

Lachlan A Jolly (LA)

University of Adelaide and Robinson Research Institute, Adelaide, Australia. Electronic address: Lachlan.Jolly@adelaide.edu.au.

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