Rare deleterious mutations of HNRNP genes result in shared neurodevelopmental disorders.


Journal

Genome medicine
ISSN: 1756-994X
Titre abrégé: Genome Med
Pays: England
ID NLM: 101475844

Informations de publication

Date de publication:
19 04 2021
Historique:
received: 04 09 2020
accepted: 16 03 2021
entrez: 20 4 2021
pubmed: 21 4 2021
medline: 19 1 2022
Statut: epublish

Résumé

With the increasing number of genomic sequencing studies, hundreds of genes have been implicated in neurodevelopmental disorders (NDDs). The rate of gene discovery far outpaces our understanding of genotype-phenotype correlations, with clinical characterization remaining a bottleneck for understanding NDDs. Most disease-associated Mendelian genes are members of gene families, and we hypothesize that those with related molecular function share clinical presentations. We tested our hypothesis by considering gene families that have multiple members with an enrichment of de novo variants among NDDs, as determined by previous meta-analyses. One of these gene families is the heterogeneous nuclear ribonucleoproteins (hnRNPs), which has 33 members, five of which have been recently identified as NDD genes (HNRNPK, HNRNPU, HNRNPH1, HNRNPH2, and HNRNPR) and two of which have significant enrichment in our previous meta-analysis of probands with NDDs (HNRNPU and SYNCRIP). Utilizing protein homology, mutation analyses, gene expression analyses, and phenotypic characterization, we provide evidence for variation in 12 HNRNP genes as candidates for NDDs. Seven are potentially novel while the remaining genes in the family likely do not significantly contribute to NDD risk. We report 119 new NDD cases (64 de novo variants) through sequencing and international collaborations and combined with published clinical case reports. We consider 235 cases with gene-disruptive single-nucleotide variants or indels and 15 cases with small copy number variants. Three hnRNP-encoding genes reach nominal or exome-wide significance for de novo variant enrichment, while nine are candidates for pathogenic mutations. Comparison of HNRNP gene expression shows a pattern consistent with a role in cerebral cortical development with enriched expression among radial glial progenitors. Clinical assessment of probands (n = 188-221) expands the phenotypes associated with HNRNP rare variants, and phenotypes associated with variation in the HNRNP genes distinguishes them as a subgroup of NDDs. Overall, our novel approach of exploiting gene families in NDDs identifies new HNRNP-related disorders, expands the phenotypes of known HNRNP-related disorders, strongly implicates disruption of the hnRNPs as a whole in NDDs, and supports that NDD subtypes likely have shared molecular pathogenesis. To date, this is the first study to identify novel genetic disorders based on the presence of disorders in related genes. We also perform the first phenotypic analyses focusing on related genes. Finally, we show that radial glial expression of these genes is likely critical during neurodevelopment. This is important for diagnostics, as well as developing strategies to best study these genes for the development of therapeutics.

Sections du résumé

BACKGROUND
With the increasing number of genomic sequencing studies, hundreds of genes have been implicated in neurodevelopmental disorders (NDDs). The rate of gene discovery far outpaces our understanding of genotype-phenotype correlations, with clinical characterization remaining a bottleneck for understanding NDDs. Most disease-associated Mendelian genes are members of gene families, and we hypothesize that those with related molecular function share clinical presentations.
METHODS
We tested our hypothesis by considering gene families that have multiple members with an enrichment of de novo variants among NDDs, as determined by previous meta-analyses. One of these gene families is the heterogeneous nuclear ribonucleoproteins (hnRNPs), which has 33 members, five of which have been recently identified as NDD genes (HNRNPK, HNRNPU, HNRNPH1, HNRNPH2, and HNRNPR) and two of which have significant enrichment in our previous meta-analysis of probands with NDDs (HNRNPU and SYNCRIP). Utilizing protein homology, mutation analyses, gene expression analyses, and phenotypic characterization, we provide evidence for variation in 12 HNRNP genes as candidates for NDDs. Seven are potentially novel while the remaining genes in the family likely do not significantly contribute to NDD risk.
RESULTS
We report 119 new NDD cases (64 de novo variants) through sequencing and international collaborations and combined with published clinical case reports. We consider 235 cases with gene-disruptive single-nucleotide variants or indels and 15 cases with small copy number variants. Three hnRNP-encoding genes reach nominal or exome-wide significance for de novo variant enrichment, while nine are candidates for pathogenic mutations. Comparison of HNRNP gene expression shows a pattern consistent with a role in cerebral cortical development with enriched expression among radial glial progenitors. Clinical assessment of probands (n = 188-221) expands the phenotypes associated with HNRNP rare variants, and phenotypes associated with variation in the HNRNP genes distinguishes them as a subgroup of NDDs.
CONCLUSIONS
Overall, our novel approach of exploiting gene families in NDDs identifies new HNRNP-related disorders, expands the phenotypes of known HNRNP-related disorders, strongly implicates disruption of the hnRNPs as a whole in NDDs, and supports that NDD subtypes likely have shared molecular pathogenesis. To date, this is the first study to identify novel genetic disorders based on the presence of disorders in related genes. We also perform the first phenotypic analyses focusing on related genes. Finally, we show that radial glial expression of these genes is likely critical during neurodevelopment. This is important for diagnostics, as well as developing strategies to best study these genes for the development of therapeutics.

Identifiants

pubmed: 33874999
doi: 10.1186/s13073-021-00870-6
pii: 10.1186/s13073-021-00870-6
pmc: PMC8056596
doi:

Substances chimiques

Heterogeneous-Nuclear Ribonucleoproteins 0

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

63

Subventions

Organisme : NIEHS NIH HHS
ID : P30 ES010126
Pays : United States
Organisme : NHGRI NIH HHS
ID : T32 HG000035
Pays : United States
Organisme : NINDS NIH HHS
ID : K08 NS092898
Pays : United States
Organisme : Wellcome Trust
Pays : United Kingdom
Organisme : NIMH NIH HHS
ID : R01 MH101221
Pays : United States
Organisme : Howard Hughes Medical Institute
Pays : United States
Organisme : Department of Health
Pays : United Kingdom

Investigateurs

John Acampado (J)
Andrea J Ace (AJ)
Alpha Amatya (A)
Irina Astrovskaya (I)
Asif Bashar (A)
Elizabeth Brooks (E)
Martin E Butler (ME)
Lindsey A Cartner (LA)
Wubin Chin (W)
Wendy K Chung (WK)
Amy M Daniels (AM)
Pamela Feliciano (P)
Chris Fleisch (C)
Swami Ganesan (S)
William Jensen (W)
Alex E Lash (AE)
Richard Marini (R)
Vincent J Myers (VJ)
Eirene O'Connor (E)
Chris Rigby (C)
Beverly E Robertson (BE)
Neelay Shah (N)
Swapnil Shah (S)
Emily Singer (E)
Lee Anne G Snyder (LAG)
Alexandra N Stephens (AN)
Jennifer Tjernagel (J)
Brianna M Vernoia (BM)
Natalia Volfovsky (N)
Loran Casey White (LC)
Alexander Hsieh (A)
Yufeng Shen (Y)
Xueya Zhou (X)
Tychele N Turner (TN)
Ethan Bahl (E)
Taylor R Thomas (TR)
Leo Brueggeman (L)
Tanner Koomar (T)
Jacob J Michaelson (JJ)
Brian J O'Roak (BJ)
Rebecca A Barnard (RA)
Richard A Gibbs (RA)
Donna Muzny (D)
Aniko Sabo (A)
Kelli L Baalman Ahmed (KL)
Evan E Eichler (EE)
Matthew Siegel (M)
Leonard Abbeduto (L)
David G Amaral (DG)
Brittani A Hilscher (BA)
Deana Li (D)
Kaitlin Smith (K)
Samantha Thompson (S)
Charles Albright (C)
Eric M Butter (EM)
Sara Eldred (S)
Nathan Hanna (N)
Mark Jones (M)
Daniel Lee Coury (DL)
Jessica Scherr (J)
Taylor Pifher (T)
Erin Roby (E)
Brandy Dennis (B)
Lorrin Higgins (L)
Melissa Brown (M)
Michael Alessandri (M)
Anibal Gutierrez (A)
Melissa N Hale (MN)
Lynette M Herbert (LM)
Hoa Lam Schneider (HL)
Giancarla David (G)
Robert D Annett (RD)
Dustin E Sarver (DE)
Ivette Arriaga (I)
Alexies Camba (A)
Amanda C Gulsrud (AC)
Monica Haley (M)
James T McCracken (JT)
Sophia Sandhu (S)
Maira Tafolla (M)
Wha S Yang (WS)
Laura A Carpenter (LA)
Catherine C Bradley (CC)
Frampton Gwynette (F)
Patricia Manning (P)
Rebecca Shaffer (R)
Carrie Thomas (C)
Raphael A Bernier (RA)
Emily A Fox (EA)
Jennifer A Gerdts (JA)
Micah Pepper (M)
Theodore Ho (T)
Daniel Cho (D)
Joseph Piven (J)
Holly Lechniak (H)
Latha V Soorya (LV)
Rachel Gordon (R)
Allison Wainer (A)
Lisa Yeh (L)
Cesar Ochoa-Lubinoff (C)
Nicole Russo (N)
Elizabeth Berry-Kravis (E)
Stephanie Booker (S)
Craig A Erickson (CA)
Lisa M Prock (LM)
Katherine G Pawlowski (KG)
Emily T Matthews (ET)
Stephanie J Brewster (SJ)
Margaret A Hojlo (MA)
Evi Abada (E)
Elena Lamarche (E)
Tianyun Wang (T)
Shwetha C Murali (SC)
William T Harvey (WT)
Hannah E Kaplan (HE)
Karen L Pierce (KL)
Lindsey DeMarco (L)
Susannah Horner (S)
Juhi Pandey (J)
Samantha Plate (S)
Mustafa Sahin (M)
Katherine D Riley (KD)
Erin Carmody (E)
Julia Constantini (J)
Amy Esler (A)
Ali Fatemi (A)
Hanna Hutter (H)
Rebecca J Landa (RJ)
Alexander P McKenzie (AP)
Jason Neely (J)
Vini Singh (V)
Bonnie Van Metre (B)
Ericka L Wodka (EL)
Eric J Fombonne (EJ)
Lark Y Huang-Storms (LY)
Lillian D Pacheco (LD)
Sarah A Mastel (SA)
Leigh A Coppola (LA)
Sunday Francis (S)
Andrea Jarrett (A)
Suma Jacob (S)
Natasha Lillie (N)
Jaclyn Gunderson (J)
Dalia Istephanous (D)
Laura Simon (L)
Ori Wasserberg (O)
Angela L Rachubinski (AL)
Cordelia R Rosenberg (CR)
Stephen M Kanne (SM)
Amanda D Shocklee (AD)
Nicole Takahashi (N)
Shelby L Bridwell (SL)
Rebecca L Klimczac (RL)
Melissa A Mahurin (MA)
Hannah E Cotrell (HE)
Cortaiga A Grant (CA)
Samantha G Hunter (SG)
Christa Lese Martin (CL)
Cora M Taylor (CM)
Lauren K Walsh (LK)
Katherine A Dent (KA)
Andrew Mason (A)
Anthony Sziklay (A)
Christopher J Smith (CJ)

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GenBank Overview. Available from: https://www.ncbi.nlm.nih.gov/genbank/

Auteurs

Madelyn A Gillentine (MA)

Department of Genome Sciences, University of Washington School of Medicine, 3720 15th Ave NE S413A, Box 355065, Seattle, WA, 981095-5065, USA.

Tianyun Wang (T)

Department of Genome Sciences, University of Washington School of Medicine, 3720 15th Ave NE S413A, Box 355065, Seattle, WA, 981095-5065, USA.

Kendra Hoekzema (K)

Department of Genome Sciences, University of Washington School of Medicine, 3720 15th Ave NE S413A, Box 355065, Seattle, WA, 981095-5065, USA.

Jill Rosenfeld (J)

Baylor Genetics Laboratories, Houston, TX, USA.
Department of Molecular & Human Genetics, Baylor College of Medicine, Houston, TX, USA.

Pengfei Liu (P)

Baylor Genetics Laboratories, Houston, TX, USA.

Hui Guo (H)

Department of Genome Sciences, University of Washington School of Medicine, 3720 15th Ave NE S413A, Box 355065, Seattle, WA, 981095-5065, USA.
Center for Medical Genetics and Hunan Key Laboratory of Medical Genetics, School of Life Sciences, Central South University, Changsha, Hunan, China.

Chang N Kim (CN)

Department of Anatomy, University of California, San Francisco, CA, USA.
Department of Psychiatry, University of California, San Francisco, CA, USA.
Weill Institute for Neurosciences, University of California at San Francisco, San Francisco, CA, USA.
The Eli and Edythe Broad Center of Regeneration Medicine and Stem Cell Research, University of California, San Francisco, CA, USA.

Bert B A De Vries (BBA)

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

Lisenka E L M Vissers (LELM)

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

Magnus Nordenskjold (M)

Department of Molecular Medicine and Surgery, Center for Molecular Medicine, Karolinska Institutet, Stockholm, Sweden.
Department of Clinical Genetics, Karolinska University Hospital, Stockholm, Sweden.

Malin Kvarnung (M)

Department of Molecular Medicine and Surgery, Center for Molecular Medicine, Karolinska Institutet, Stockholm, Sweden.
Department of Clinical Genetics, Karolinska University Hospital, Stockholm, Sweden.

Anna Lindstrand (A)

Department of Molecular Medicine and Surgery, Center for Molecular Medicine, Karolinska Institutet, Stockholm, Sweden.
Department of Clinical Genetics, Karolinska University Hospital, Stockholm, Sweden.

Ann Nordgren (A)

Department of Molecular Medicine and Surgery, Center for Molecular Medicine, Karolinska Institutet, Stockholm, Sweden.
Department of Clinical Genetics, Karolinska University Hospital, Stockholm, Sweden.

Jozef Gecz (J)

School of Medicine and the Robinson Research Institute, the University of Adelaide at the Women's and Children's Hospital, Adelaide, South Australia, Australia.
Genetics and Molecular Pathology, SA Pathology, Adelaide, South Australia, Australia.
South Australian Health and Medical Research Institute, Adelaide, South Australia, Australia.

Maria Iascone (M)

Laboratorio di Genetica Medica - ASST Papa Giovanni XXIII, Bergamo, Italy.

Anna Cereda (A)

Department of Pediatrics, ASST Papa Giovanni XXIII, Bergamo, Italy.

Agnese Scatigno (A)

Department of Pediatrics, ASST Papa Giovanni XXIII, Bergamo, Italy.

Silvia Maitz (S)

Genetic Unit, Department of Pediatrics, Fondazione MBBM S. Gerardo Hospital, Monza, Italy.

Ginevra Zanni (G)

Unit of Neuromuscular and Neurodegenerative Disorders, Department Neurosciences, Bambino Gesù Children's Hospital, IRCCS, 00146, Rome, Italy.

Enrico Bertini (E)

Unit of Neuromuscular and Neurodegenerative Disorders, Department Neurosciences, Bambino Gesù Children's Hospital, IRCCS, 00146, Rome, Italy.

Christiane Zweier (C)

Institute of Human Genetics, Friedrich-Alexander-Universität Erlangen-Nürnberg (FAU), Erlangen, Germany.

Sarah Schuhmann (S)

Institute of Human Genetics, Friedrich-Alexander-Universität Erlangen-Nürnberg (FAU), Erlangen, Germany.

Antje Wiesener (A)

Institute of Human Genetics, Friedrich-Alexander-Universität Erlangen-Nürnberg (FAU), Erlangen, Germany.

Micah Pepper (M)

Center on Human Development and Disability, University of Washington, Seattle, WA, USA.
Seattle Children's Autism Center, Seattle, WA, USA.

Heena Panjwani (H)

Center on Human Development and Disability, University of Washington, Seattle, WA, USA.
Seattle Children's Autism Center, Seattle, WA, USA.

Erin Torti (E)

GeneDX, Gaithersburg, MD, USA.

Farida Abid (F)

Department of Pediatrics-Neurology, Baylor College of Medicine, Houston, TX, USA.
Texas Children's Hospital, Houston, TX, USA.

Irina Anselm (I)

Department of Neurology, Boston Children's Hospital, Harvard Medical School, Boston, MA, USA.

Siddharth Srivastava (S)

Department of Neurology, Boston Children's Hospital, Harvard Medical School, Boston, MA, USA.

Paldeep Atwal (P)

The Atwal Clinic: Genomic & Personalized Medicine, Jacksonville, FL, USA.

Carlos A Bacino (CA)

Department of Molecular & Human Genetics, Baylor College of Medicine, Houston, TX, USA.

Gifty Bhat (G)

Department of Pediatrics, Section of Genetics, University of Illinois at Chicago, Chicago, IL, USA.

Katherine Cobian (K)

Department of Pediatrics, Section of Genetics, University of Illinois at Chicago, Chicago, IL, USA.

Lynne M Bird (LM)

Department of Pediatrics, University of California San Diego, San Diego, CA, USA.
Genetics/Dysmorphology, Rady Children's Hospital San Diego, San Diego, CA, USA.

Jennifer Friedman (J)

Department of Pediatrics, University of California San Diego, San Diego, CA, USA.
Rady Children's Institute for Genomic Medicine, San Diego, CA, USA.
Department of Neurosciences, University of California San Diego, San Diego, CA, USA.

Meredith S Wright (MS)

Department of Pediatrics, University of California San Diego, San Diego, CA, USA.
Rady Children's Institute for Genomic Medicine, San Diego, CA, USA.

Bert Callewaert (B)

Department of Biomolecular Medicine, Ghent University Hospital, Ghent, Belgium.

Florence Petit (F)

Clinique de Génétique, Hôpital Jeanne de Flandre, Bâtiment Modulaire, CHU, 59037, Lille Cedex, France.

Sophie Mathieu (S)

Sorbonne Universités, Centre de Référence déficiences intellectuelles de causes rares, département de génétique et embryologie médicale, Hôpital Trousseau, AP-HP, Paris, France.

Alexandra Afenjar (A)

Sorbonne Universités, Centre de Référence déficiences intellectuelles de causes rares, département de génétique et embryologie médicale, Hôpital Trousseau, AP-HP, Paris, France.

Celenie K Christensen (CK)

Department of Pediatrics, Indiana University School of Medicine, Indianapolis, IN, USA.

Kerry M White (KM)

Department of Medical and Molecular Genetics, IU Health, Indianapolis, IN, USA.

Orly Elpeleg (O)

Department of Genetics, Hadassah, Hebrew University Medical Center, Jerusalem, Israel.

Itai Berger (I)

Pediatric Neurology, Assuta-Ashdod University Hospital, Ashdod, Israel.
Health Sciences, Ben-Gurion University of the Negev, Beersheba, Israel.

Edward J Espineli (EJ)

Department of Pediatrics-Neurology, Baylor College of Medicine, Houston, TX, USA.
Texas Children's Hospital, Houston, TX, USA.

Christina Fagerberg (C)

Department of Clinical Genetics, Odense University Hospital, Odense, Denmark.

Charlotte Brasch-Andersen (C)

Department of Clinical Genetics, Odense University Hospital, Odense, Denmark.

Lars Kjærsgaard Hansen (LK)

H C Andersen Chilldrens Hospital, Odense University Hospital, Odense, Denmark.

Timothy Feyma (T)

Gillette Children's Specialty Healthcare, Saint Paul, MN, USA.

Susan Hughes (S)

Division of Clinical Genetics, Children's Mercy Kansas City, Kansas City, MO, USA.
The University of Missouri-Kansas City, School of Medicine, Kansas City, MO, USA.

Isabelle Thiffault (I)

The University of Missouri-Kansas City, School of Medicine, Kansas City, MO, USA.
Children's Mercy Kansas City, Center for Pediatric Genomic Medicine, Kansas City, MO, USA.

Bonnie Sullivan (B)

Division of Clinical Genetics, Children's Mercy Kansas City, Kansas City, MO, USA.

Shuang Yan (S)

Division of Clinical Genetics, Children's Mercy Kansas City, Kansas City, MO, USA.

Kory Keller (K)

Oregon Health & Science University, Corvallis, OR, USA.

Boris Keren (B)

Department of Genetics, Hópital Pitié-Salpêtrière, Paris, France.

Cyril Mignot (C)

Department of Genetics, Hópital Pitié-Salpêtrière, Paris, France.

Frank Kooy (F)

Department of Medical Genetics, University of Antwerp, Antwerp, Belgium.

Marije Meuwissen (M)

Department of Medical Genetics, University of Antwerp, Antwerp, Belgium.

Alice Basinger (A)

Genetics Department, Cook Children's Hospital, Fort Worth, TX, USA.

Mary Kukolich (M)

Genetics Department, Cook Children's Hospital, Fort Worth, TX, USA.

Meredith Philips (M)

Genetics Department, Cook Children's Hospital, Fort Worth, TX, USA.

Lucia Ortega (L)

Genetics Department, Cook Children's Hospital, Fort Worth, TX, USA.

Margaret Drummond-Borg (M)

Genetics Department, Cook Children's Hospital, Fort Worth, TX, USA.

Mathilde Lauridsen (M)

Department of Clinical Genetics, Odense University Hospital, Odense, Denmark.

Kristina Sorensen (K)

Department of Clinical Genetics, Odense University Hospital, Odense, Denmark.

Anna Lehman (A)

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

Elena Lopez-Rangel (E)

Department of Medical Genetics, University of British Columbia, Vancouver, Canada.
Division of Developmental Pediatrics, Department of Pediatrics, BC Children's Hospital, University of British Columbia, Vancouver, BC, Canada.
Sunny Hill Health Centre for Children, Vancouver, BC, Canada.

Paul Levy (P)

Department of Pediatrics, The Children's Hospital at Montefiore, Bronx, NY, USA.

Davor Lessel (D)

Institute of Human Genetics, University Medical Center Hamburg-Eppendorf, Hamburg, Germany.

Timothy Lotze (T)

Department of Pediatrics-Neurology, Baylor College of Medicine, Houston, TX, USA.

Suneeta Madan-Khetarpal (S)

Department of Human Genetics, University of Pittsburgh, Pittsburgh, PA, USA.
UPMC Children's Hospital of Pittsburgh, Pittsburgh, PA, USA.

Jessica Sebastian (J)

Department of Human Genetics, University of Pittsburgh, Pittsburgh, PA, USA.

Jodie Vento (J)

Department of Human Genetics, University of Pittsburgh, Pittsburgh, PA, USA.

Divya Vats (D)

Kaiser Permanente Southern California, Los Angeles, CA, USA.

L Manace Benman (LM)

The Permanente Medical Group, Oakland, CA, USA.

Shane Mckee (S)

Northern Ireland Regional Genetics Service, Belfast City Hospital, Belfast, UK.

Ghayda M Mirzaa (GM)

Center for Integrative Brain Research, Seattle Children's Research Institute, Seattle, WA, USA.
Department of Pediatrics, University of Washington, Seattle, WA, USA.
Brotman Baty Institute for Precision Medicine, Seattle, WA, USA.

Candace Muss (C)

Al Dupont Hospital for Children, Wilmington, DE, USA.

John Pappas (J)

NYU Grossman School of Medicine, Department of Pediatrics, Clinical Genetic Services, New York, NY, USA.

Hilde Peeters (H)

Center for Human Genetics, KU Leuven and Leuven Autism Research (LAuRes), Leuven, Belgium.

Corrado Romano (C)

Oasi Research Institute-IRCCS, Troina, Italy.

Maurizio Elia (M)

Oasi Research Institute-IRCCS, Troina, Italy.

Ornella Galesi (O)

Oasi Research Institute-IRCCS, Troina, Italy.

Marleen E H Simon (MEH)

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

Koen L I van Gassen (KLI)

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

Kara Simpson (K)

Rare Disease Institute, Children's National Health System, Washington, DC, USA.

Robert Stratton (R)

Department of Genetics, Driscoll Children's Hospital, Corpus Christi, TX, USA.

Sabeen Syed (S)

Department of Pediatric Gastroenterology, Driscoll Children's Hospital, Corpus Christi, TX, USA.

Julien Thevenon (J)

Àrea de Genètica Clínica i Molecular, Hospital Vall d'Hebrón, Barcelona, Spain.

Irene Valenzuela Palafoll (IV)

Centre de référence Anomalies du développement, CHU Grenoble-Alpes, Grenoble, France.

Antonio Vitobello (A)

UF Innovation en Diagnostic Génomique des Maladies Rares, FHU-TRANSLAD, CHU Dijon Bourgogne and INSERM UMR1231 GAD, Université de Bourgogne Franche-Comté, F-21000, Dijon, France.
INSERM UMR 1231 Génétique des Anomalies du Développement, Université Bourgogne Franche-Comté, Dijon, France.

Marie Bournez (M)

Centre de Référence Maladies Rares « déficience intellectuelle », Centre de Génétique, FHU-TRANSLAD, CHU Dijon Bourgogne, Dijon, France.
Centre de Référence Maladies Rares « Anomalies du Développement et Syndromes malformatifs »​ Université Bourgogne Franche-Comté, Dijon, France.

Laurence Faivre (L)

INSERM UMR 1231 Génétique des Anomalies du Développement, Université Bourgogne Franche-Comté, Dijon, France.
Centre de Référence Maladies Rares « Anomalies du Développement et Syndromes malformatifs »​ Université Bourgogne Franche-Comté, Dijon, France.

Kun Xia (K)

Center for Medical Genetics and Hunan Key Laboratory of Medical Genetics, School of Life Sciences, Central South University, Changsha, Hunan, China.

Rachel K Earl (RK)

Center on Human Development and Disability, University of Washington, Seattle, WA, USA.
Seattle Children's Autism Center, Seattle, WA, USA.
Department of Psychiatry and Behavioral Sciences, University of Washington, Seattle, WA, USA.

Tomasz Nowakowski (T)

Department of Anatomy, University of California, San Francisco, CA, USA.
Department of Psychiatry, University of California, San Francisco, CA, USA.
Weill Institute for Neurosciences, University of California at San Francisco, San Francisco, CA, USA.
The Eli and Edythe Broad Center of Regeneration Medicine and Stem Cell Research, University of California, San Francisco, CA, USA.

Raphael A Bernier (RA)

Center on Human Development and Disability, University of Washington, Seattle, WA, USA.
Seattle Children's Autism Center, Seattle, WA, USA.
Department of Psychiatry and Behavioral Sciences, University of Washington, Seattle, WA, USA.

Evan E Eichler (EE)

Department of Genome Sciences, University of Washington School of Medicine, 3720 15th Ave NE S413A, Box 355065, Seattle, WA, 981095-5065, USA. eee@gs.washington.edu.
Howard Hughes Medical Institute, University of Washington, Seattle, WA, USA. eee@gs.washington.edu.

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