{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,1,13]],"date-time":"2026-01-13T06:00:48Z","timestamp":1768284048607,"version":"3.49.0"},"reference-count":82,"publisher":"Springer Science and Business Media LLC","issue":"6","license":[{"start":{"date-parts":[[2024,6,1]],"date-time":"2024-06-01T00:00:00Z","timestamp":1717200000000},"content-version":"tdm","delay-in-days":0,"URL":"https:\/\/creativecommons.org\/licenses\/by\/4.0"},{"start":{"date-parts":[[2024,6,14]],"date-time":"2024-06-14T00:00:00Z","timestamp":1718323200000},"content-version":"vor","delay-in-days":13,"URL":"https:\/\/creativecommons.org\/licenses\/by\/4.0"}],"funder":[{"DOI":"10.13039\/100012352","name":"Universit\u00e0 degli Studi di Milano","doi-asserted-by":"crossref","id":[{"id":"10.13039\/100012352","id-type":"DOI","asserted-by":"crossref"}]}],"content-domain":{"domain":["link.springer.com"],"crossmark-restriction":false},"short-container-title":["Hum. Genet."],"published-print":{"date-parts":[[2024,6]]},"abstract":"<jats:title>Abstract<\/jats:title><jats:p>NF1 microdeletion syndrome, accounting for 5\u201311% of NF1 patients, is caused by a deletion in the <jats:italic>NF1<\/jats:italic> region and it is generally characterized by a severe phenotype. Although 70% of NF1 microdeletion patients presents the same 1.4\u00a0Mb type-I deletion, some patients may show additional clinical features. Therefore, the contribution of several pathogenic mechanisms, besides haploinsufficiency of some genes within the deletion interval, is expected and needs to be defined. We investigated an altered expression of deletion flanking genes by qPCR in patients with type-1 NF1 deletion, compared to healthy donors, possibly contributing to the clinical traits of NF1 microdeletion syndrome. In addition, the 1.4-Mb deletion leads to changes in the 3D chromatin structure in the 17q11.2 region. Specifically, this deletion alters DNA-DNA interactions in the regions flanking the breakpoints, as demonstrated by our 4C-seq analysis. This alteration likely causes position effect on the expression of deletion flanking genes.<\/jats:p><jats:p>Interestingly, 4C-seq analysis revealed that in microdeletion patients, an interaction was established between the <jats:italic>RHOT1<\/jats:italic> promoter and the <jats:italic>SLC6A4<\/jats:italic> gene, which showed increased expression. We performed NGS on putative modifier genes, and identified two \u201clikely pathogenic\u201d rare variants in RAS pathway, possibly contributing to incidental phenotypic features.<\/jats:p><jats:p>This study provides new insights into understanding the pathogenesis of NF1 microdeletion syndrome and suggests a novel pathomechanism that contributes to the expression phenotype in addition to haploinsufficiency of genes located within the deletion.This is a pivotal approach that can be applied to unravel microdeletion syndromes, improving precision medicine, prognosis and patients\u2019 follow-up.<\/jats:p>","DOI":"10.1007\/s00439-024-02683-0","type":"journal-article","created":{"date-parts":[[2024,6,14]],"date-time":"2024-06-14T13:02:05Z","timestamp":1718370125000},"page":"775-795","update-policy":"https:\/\/doi.org\/10.1007\/springer_crossmark_policy","source":"Crossref","is-referenced-by-count":3,"title":["Genetic\/epigenetic effects in NF1 microdeletion syndrome: beyond the haploinsufficiency, looking at the contribution of not deleted genes"],"prefix":"10.1007","volume":"143","author":[{"ORCID":"https:\/\/orcid.org\/0000-0002-4558-8376","authenticated-orcid":false,"given":"Viviana","family":"Tritto","sequence":"first","affiliation":[],"role":[{"role":"author","vocabulary":"crossref"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-6597-3714","authenticated-orcid":false,"given":"Paola","family":"Bettinaglio","sequence":"additional","affiliation":[],"role":[{"role":"author","vocabulary":"crossref"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-4040-4654","authenticated-orcid":false,"given":"Eleonora","family":"Mangano","sequence":"additional","affiliation":[],"role":[{"role":"author","vocabulary":"crossref"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-6819-0846","authenticated-orcid":false,"given":"Claudia","family":"Cesaretti","sequence":"additional","affiliation":[],"role":[{"role":"author","vocabulary":"crossref"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-0699-5323","authenticated-orcid":false,"given":"Federica","family":"Marasca","sequence":"additional","affiliation":[],"role":[{"role":"author","vocabulary":"crossref"}]},{"ORCID":"https:\/\/orcid.org\/0000-0003-0595-1553","authenticated-orcid":false,"given":"Chiara","family":"Castronovo","sequence":"additional","affiliation":[],"role":[{"role":"author","vocabulary":"crossref"}]},{"ORCID":"https:\/\/orcid.org\/0000-0001-6996-4033","authenticated-orcid":false,"given":"Roberta","family":"Bordoni","sequence":"additional","affiliation":[],"role":[{"role":"author","vocabulary":"crossref"}]},{"ORCID":"https:\/\/orcid.org\/0000-0003-3025-9657","authenticated-orcid":false,"given":"Cristina","family":"Battaglia","sequence":"additional","affiliation":[],"role":[{"role":"author","vocabulary":"crossref"}]},{"ORCID":"https:\/\/orcid.org\/0000-0003-4830-6204","authenticated-orcid":false,"given":"Veronica","family":"Saletti","sequence":"additional","affiliation":[],"role":[{"role":"author","vocabulary":"crossref"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-0602-5335","authenticated-orcid":false,"given":"Valeria","family":"Ranzani","sequence":"additional","affiliation":[],"role":[{"role":"author","vocabulary":"crossref"}]},{"ORCID":"https:\/\/orcid.org\/0000-0003-0527-9234","authenticated-orcid":false,"given":"Beatrice","family":"Bodega","sequence":"additional","affiliation":[],"role":[{"role":"author","vocabulary":"crossref"}]},{"ORCID":"https:\/\/orcid.org\/0000-0003-4744-9377","authenticated-orcid":false,"given":"Marica","family":"Eoli","sequence":"additional","affiliation":[],"role":[{"role":"author","vocabulary":"crossref"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-8625-3089","authenticated-orcid":false,"given":"Federica","family":"Natacci","sequence":"additional","affiliation":[],"role":[{"role":"author","vocabulary":"crossref"}]},{"ORCID":"https:\/\/orcid.org\/0000-0001-6481-8467","authenticated-orcid":false,"given":"Paola","family":"Riva","sequence":"additional","affiliation":[],"role":[{"role":"author","vocabulary":"crossref"}]}],"member":"297","published-online":{"date-parts":[[2024,6,14]]},"reference":[{"key":"2683_CR1","doi-asserted-by":"publisher","first-page":"1167","DOI":"10.1002\/ajmg.a.35847","volume":"161","author":"IE Amarillo","year":"2013","unstructured":"Amarillo IE, Dipple KM, Quintero-Rivera F (2013) Familial microdeletion of 17q24.3 Upstream of SOX9 is Associated with isolated Pierre Robin sequence due to position effect. Am J Med Genet Part A 161:1167\u20131172. https:\/\/doi.org\/10.1002\/ajmg.a.35847","journal-title":"Am J Med Genet Part A"},{"key":"2683_CR2","doi-asserted-by":"publisher","first-page":"68","DOI":"10.1038\/nature15393","volume":"526","author":"A Auton","year":"2015","unstructured":"Auton A, Abecasis GR, Altshuler DM et al (2015) A global reference for human genetic variation. Nature 526:68\u201374. https:\/\/doi.org\/10.1038\/nature15393","journal-title":"Nature"},{"key":"2683_CR3","doi-asserted-by":"publisher","first-page":"931","DOI":"10.1038\/s41431-023-01377-x","volume":"31","author":"P Bettinaglio","year":"2023","unstructured":"Bettinaglio P, Mangano E, Tritto V et al (2023) New insights into the molecular basis of spinal neurofibromatosis type 1. Eur J Hum Genet 31:931\u2013938. https:\/\/doi.org\/10.1038\/s41431-023-01377-x","journal-title":"Eur J Hum Genet"},{"key":"2683_CR4","doi-asserted-by":"publisher","first-page":"2114","DOI":"10.1093\/bioinformatics\/btu170","volume":"30","author":"AM Bolger","year":"2014","unstructured":"Bolger AM, Lohse M, Usadel B (2014) Trimmomatic: a flexible trimmer for Illumina sequence data. Bioinformatics 30:2114\u20132120. https:\/\/doi.org\/10.1093\/bioinformatics\/btu170","journal-title":"Bioinformatics"},{"key":"2683_CR5","doi-asserted-by":"publisher","first-page":"e157","DOI":"10.1371\/journal.pbio.0030157","volume":"3","author":"A Bolzer","year":"2005","unstructured":"Bolzer A, Kreth G, Solovei I et al (2005) Three-dimensional maps of all chromosomes in human male fibroblast nuclei and prometaphase rosettes. PLoS Biol 3:e157. https:\/\/doi.org\/10.1371\/journal.pbio.0030157","journal-title":"PLoS Biol"},{"key":"2683_CR6","doi-asserted-by":"publisher","first-page":"i85","DOI":"10.1093\/noajnl\/vdz040","volume":"2","author":"ADA Costa","year":"2020","unstructured":"Costa ADA, Gutmann DH (2020) Brain tumors in neurofibromatosis type 1. Neuro-Oncology Adv 2:i85\u2013i97. https:\/\/doi.org\/10.1093\/noajnl\/vdz040","journal-title":"Neuro-Oncology Adv"},{"key":"2683_CR7","doi-asserted-by":"publisher","first-page":"386","DOI":"10.1101\/gr.277187.122","volume":"33","author":"D Dang","year":"2023","unstructured":"Dang D, Zhang S-W, Duan R, Zhang S (2023) Defining the separation landscape of topological domains for decoding consensus domain organization of the 3D genome. Genome Res 33:386\u2013400. https:\/\/doi.org\/10.1101\/gr.277187.122","journal-title":"Genome Res"},{"key":"2683_CR8","doi-asserted-by":"publisher","first-page":"1288","DOI":"10.1086\/374821","volume":"72","author":"T De Raedt","year":"2003","unstructured":"De Raedt T, Brems H, Wolkenstein P et al (2003) Elevated risk for MPNST in NF1 microdeletion patients. Am J Hum Genet 72:1288\u20131292. https:\/\/doi.org\/10.1086\/374821","journal-title":"Am J Hum Genet"},{"key":"2683_CR9","doi-asserted-by":"publisher","first-page":"429","DOI":"10.1111\/cge.13150","volume":"93","author":"C Deltas","year":"2018","unstructured":"Deltas C (2018) Digenic inheritance and genetic modifiers. Clin Genet 93:429\u2013438. https:\/\/doi.org\/10.1111\/cge.13150","journal-title":"Clin Genet"},{"key":"2683_CR10","doi-asserted-by":"publisher","first-page":"491","DOI":"10.1038\/ng.806","volume":"43","author":"MA DePristo","year":"2011","unstructured":"DePristo MA, Banks E, Poplin R et al (2011) A framework for variation discovery and genotyping using next-generation DNA sequencing data. Nat Genet 43:491\u2013498. https:\/\/doi.org\/10.1038\/ng.806","journal-title":"Nat Genet"},{"key":"2683_CR11","doi-asserted-by":"publisher","first-page":"635","DOI":"10.1038\/ng.2963","volume":"46","author":"PS Dhandapany","year":"2014","unstructured":"Dhandapany PS, Razzaque MA, Muthusami U et al (2014) RAF1 mutations in childhood-onset dilated cardiomyopathy. Nat Genet 46:635\u2013639. https:\/\/doi.org\/10.1038\/ng.2963","journal-title":"Nat Genet"},{"key":"2683_CR12","doi-asserted-by":"publisher","first-page":"1147","DOI":"10.1038\/nsmb.2669","volume":"20","author":"L Di Croce","year":"2013","unstructured":"Di Croce L, Helin K (2013) Transcriptional regulation by polycomb group proteins. Nat Struct Mol Biol 20:1147\u20131155. https:\/\/doi.org\/10.1038\/nsmb.2669","journal-title":"Nat Struct Mol Biol"},{"key":"2683_CR13","doi-asserted-by":"publisher","first-page":"963","DOI":"10.1038\/ng2083","volume":"39","author":"J Douglas","year":"2007","unstructured":"Douglas J, Cilliers D, Coleman K et al (2007) Mutations in RNF135, a gene within the NF1 microdeletion region, cause phenotypic abnormalities including overgrowth. Nat Genet 39:963\u2013965. https:\/\/doi.org\/10.1038\/ng2083","journal-title":"Nat Genet"},{"key":"2683_CR14","doi-asserted-by":"publisher","first-page":"4384","DOI":"10.1038\/s41467-018-06908-5","volume":"9","author":"F Fang","year":"2018","unstructured":"Fang F, Xia N, Angulo B et al (2018) A distinct isoform of ZNF207 controls self-renewal and pluripotency of human embryonic stem cells. Nat Commun 9:4384. https:\/\/doi.org\/10.1038\/s41467-018-06908-5","journal-title":"Nat Commun"},{"key":"2683_CR16","doi-asserted-by":"publisher","first-page":"1329","DOI":"10.1007\/s00439-017-1832-5","volume":"136","author":"L Ferrari","year":"2017","unstructured":"Ferrari L, Scuvera G, Tucci A et al (2017) Identification of an atypical microdeletion generating the RNF135-SUZ12 chimeric gene and causing a position effect in an NF1 patient with overgrowth. Hum Genet 136:1329\u20131339. https:\/\/doi.org\/10.1007\/s00439-017-1832-5","journal-title":"Hum Genet"},{"key":"2683_CR15","doi-asserted-by":"publisher","first-page":"1432","DOI":"10.1038\/s41431-020-0658-0","volume":"28","author":"L Ferrari","year":"2020","unstructured":"Ferrari L, Mangano E, Bonati MT et al (2020) Digenic inheritance of subclinical variants in Noonan Syndrome patients: an alternative pathogenic model? Eur J Hum Genet 28:1432\u20131445. https:\/\/doi.org\/10.1038\/s41431-020-0658-0","journal-title":"Eur J Hum Genet"},{"key":"2683_CR17","doi-asserted-by":"publisher","first-page":"1502","DOI":"10.1016\/j.cell.2019.01.020","volume":"176","author":"EH Finn","year":"2019","unstructured":"Finn EH, Pegoraro G, Brand\u00e3o HB et al (2019) Extensive heterogeneity and intrinsic variation in spatial Genome Organization. Cell 176:1502\u20131515e10. https:\/\/doi.org\/10.1016\/j.cell.2019.01.020","journal-title":"Cell"},{"key":"2683_CR18","doi-asserted-by":"publisher","first-page":"524","DOI":"10.1016\/j.ajhg.2009.03.010","volume":"84","author":"HV Firth","year":"2009","unstructured":"Firth HV, Richards SM, Bevan AP et al (2009) DECIPHER: database of chromosomal imbalance and phenotype in humans using Ensembl resources. Am J Hum Genet 84:524\u2013533. https:\/\/doi.org\/10.1016\/j.ajhg.2009.03.010","journal-title":"Am J Hum Genet"},{"key":"2683_CR19","doi-asserted-by":"publisher","first-page":"6495","DOI":"10.1074\/jbc.M208609200","volume":"278","author":"\u00c5 Fransson","year":"2003","unstructured":"Fransson \u00c5, Ruusala A, Aspenstr\u00f6m P (2003) Atypical rho GTPases have roles in mitochondrial homeostasis and apoptosis. J Biol Chem 278:6495\u20136502. https:\/\/doi.org\/10.1074\/jbc.M208609200","journal-title":"J Biol Chem"},{"key":"2683_CR20","doi-asserted-by":"publisher","first-page":"133","DOI":"10.2147\/TACG.S18675","volume":"7","author":"S Gallati","year":"2014","unstructured":"Gallati S (2014) Disease-modifying genes and monogenic disorders: experience in cystic fibrosis. Appl Clin Genet 7:133\u2013146. https:\/\/doi.org\/10.2147\/TACG.S18675","journal-title":"Appl Clin Genet"},{"key":"2683_CR21","doi-asserted-by":"publisher","first-page":"69","DOI":"10.1007\/s00439-004-1101-2","volume":"115","author":"C Gervasini","year":"2004","unstructured":"Gervasini C, Orzan F, Bentivegna A et al (2004) Evidence for non-homologous end joining and non-allelic homologous recombination in atypical NF1 microdeletions. Hum Genet 115:69\u201380. https:\/\/doi.org\/10.1007\/s00439-004-1101-2","journal-title":"Hum Genet"},{"key":"2683_CR22","doi-asserted-by":"publisher","first-page":"273","DOI":"10.1016\/j.ygeno.2004.10.014","volume":"85","author":"C Gervasini","year":"2005","unstructured":"Gervasini C, Venturin M, Orzan F et al (2005) Uncommon alu-mediated NF1 microdeletion with a breakpoint inside the NF1 gene. Genomics 85:273\u2013279. https:\/\/doi.org\/10.1016\/j.ygeno.2004.10.014","journal-title":"Genomics"},{"key":"2683_CR23","doi-asserted-by":"publisher","first-page":"e79973","DOI":"10.1371\/journal.pone.0079973","volume":"8","author":"N Gheldof","year":"2013","unstructured":"Gheldof N, Witwicki RM, Migliavacca E et al (2013) Structural variation-associated expression changes are paralleled by chromatin architecture modifications. PLoS ONE 8:e79973. https:\/\/doi.org\/10.1371\/journal.pone.0079973","journal-title":"PLoS ONE"},{"key":"2683_CR24","doi-asserted-by":"publisher","first-page":"D514","DOI":"10.1093\/nar\/gki033","volume":"33","author":"A Hamosh","year":"2004","unstructured":"Hamosh A (2004) Online mendelian inheritance in man (OMIM), a knowledgebase of human genes and genetic disorders. Nucleic Acids Res 33:D514\u2013D517. https:\/\/doi.org\/10.1093\/nar\/gki033","journal-title":"Nucleic Acids Res"},{"key":"2683_CR25","doi-asserted-by":"publisher","first-page":"101","DOI":"10.1097\/MCD.0000000000000153","volume":"26","author":"VL Hartill","year":"2017","unstructured":"Hartill VL, Dillon MW, Warren DJ, Blyth M (2017) RAF1-associated Noonan syndrome presenting antenatally with an abnormality of skull shape, subdural haematoma and associated with novel cerebral malformations. Clin Dysmorphol 26:101\u2013106. https:\/\/doi.org\/10.1097\/MCD.0000000000000153","journal-title":"Clin Dysmorphol"},{"key":"2683_CR26","doi-asserted-by":"publisher","first-page":"1807","DOI":"10.1002\/cncr.23648","volume":"113","author":"RT Jensen","year":"2008","unstructured":"Jensen RT, Berna MJ, Bingham DB, Norton JA (2008) Inherited pancreatic endocrine tumor syndromes: advances in molecular pathogenesis, diagnosis, management, and controversies. Cancer 113:1807\u20131843. https:\/\/doi.org\/10.1002\/cncr.23648","journal-title":"Cancer"},{"key":"2683_CR27","doi-asserted-by":"publisher","first-page":"268","DOI":"10.1016\/j.devcel.2013.12.013","volume":"28","author":"H Jiang","year":"2014","unstructured":"Jiang H, He X, Wang S et al (2014) A microtubule-associated zinc finger protein, BuGZ, regulates mitotic chromosome alignment by ensuring Bub3 stability and kinetochore targeting. Dev Cell 28:268\u2013281. https:\/\/doi.org\/10.1016\/j.devcel.2013.12.013","journal-title":"Dev Cell"},{"key":"2683_CR28","doi-asserted-by":"publisher","first-page":"434","DOI":"10.1038\/s41586-020-2308-7","volume":"581","author":"KJ Karczewski","year":"2020","unstructured":"Karczewski KJ, Francioli LC, Tiao G et al (2020) The mutational constraint spectrum quantified from variation in 141,456 humans. Nature 581:434\u2013443. https:\/\/doi.org\/10.1038\/s41586-020-2308-7","journal-title":"Nature"},{"key":"2683_CR29","doi-asserted-by":"publisher","first-page":"410","DOI":"10.1086\/423624","volume":"75","author":"H Kehrer-Sawatzki","year":"2004","unstructured":"Kehrer-Sawatzki H, Kluwe L, Sandig C et al (2004) High frequency of mosaicism among patients with neurofibromatosis type 1 (NF1) with microdeletions caused by somatic recombination of the JJAZ1 gene. Am J Hum Genet 75:410\u2013423. https:\/\/doi.org\/10.1086\/423624","journal-title":"Am J Hum Genet"},{"key":"2683_CR30","doi-asserted-by":"publisher","first-page":"349","DOI":"10.1007\/s00439-017-1766-y","volume":"136","author":"H Kehrer-Sawatzki","year":"2017","unstructured":"Kehrer-Sawatzki H, Mautner V-F, Cooper DN (2017) Emerging genotype\u2013phenotype relationships in patients with large NF1 deletions. Hum Genet 136:349\u2013376. https:\/\/doi.org\/10.1007\/s00439-017-1766-y","journal-title":"Hum Genet"},{"key":"2683_CR31","doi-asserted-by":"publisher","first-page":"187","DOI":"10.1001\/archderm.139.2.187","volume":"139","author":"K Khosrotehrani","year":"2003","unstructured":"Khosrotehrani K, Bastuji-Garin S, Zeller J et al (2003) Clinical risk factors for mortality in patients with neurofibromatosis 1: a cohort study of 378 patients. Arch Dermatol 139:187\u2013191. https:\/\/doi.org\/10.1001\/archderm.139.2.187","journal-title":"Arch Dermatol"},{"key":"2683_CR32","doi-asserted-by":"publisher","first-page":"1611","DOI":"10.1093\/hmg\/7.10.1611","volume":"7","author":"DJ Kleinjan","year":"1998","unstructured":"Kleinjan DJ, van Heyningen V (1998) Position effect in human genetic disease. Hum Mol Genet 7:1611\u20131618. https:\/\/doi.org\/10.1093\/hmg\/7.10.1611","journal-title":"Hum Mol Genet"},{"key":"2683_CR33","doi-asserted-by":"publisher","first-page":"17","DOI":"10.1016\/j.ymeth.2019.07.014","volume":"170","author":"PHL Krijger","year":"2020","unstructured":"Krijger PHL, Geeven G, Bianchi V et al (2020) 4\u00a0C-seq from beginning to end: a detailed protocol for sample preparation and data analysis. Methods 170:17\u201332. https:\/\/doi.org\/10.1016\/j.ymeth.2019.07.014","journal-title":"Methods"},{"key":"2683_CR34","doi-asserted-by":"publisher","first-page":"D1062","DOI":"10.1093\/nar\/gkx1153","volume":"46","author":"MJ Landrum","year":"2018","unstructured":"Landrum MJ, Lee JM, Benson M et al (2018) ClinVar: improving access to variant interpretations and supporting evidence. Nucleic Acids Res 46:D1062\u2013D1067. https:\/\/doi.org\/10.1093\/nar\/gkx1153","journal-title":"Nucleic Acids Res"},{"key":"2683_CR35","doi-asserted-by":"publisher","first-page":"736","DOI":"10.1016\/j.stem.2019.03.004","volume":"24","author":"M Laugsch","year":"2019","unstructured":"Laugsch M, Bartusel M, Rehimi R et al (2019) Modeling the pathological Long-Range Regulatory effects of Human Structural Variation with patient-specific hiPSCs. Cell Stem Cell 24:736\u2013752e12. https:\/\/doi.org\/10.1016\/j.stem.2019.03.004","journal-title":"Cell Stem Cell"},{"key":"2683_CR36","doi-asserted-by":"publisher","first-page":"285","DOI":"10.1038\/nature19057","volume":"536","author":"M Lek","year":"2016","unstructured":"Lek M, Karczewski KJ, Minikel EV et al (2016) Analysis of protein-coding genetic variation in 60,706 humans. Nature 536:285\u2013291. https:\/\/doi.org\/10.1038\/nature19057","journal-title":"Nature"},{"key":"2683_CR37","unstructured":"Li H (2013) Aligning sequence reads, clone sequences and assembly contigs with BWA-MEM. arXiv Prepr arXiv"},{"key":"2683_CR38","doi-asserted-by":"publisher","first-page":"2078","DOI":"10.1093\/bioinformatics\/btp352","volume":"25","author":"H Li","year":"2009","unstructured":"Li H, Handsaker B, Wysoker A et al (2009) The sequence Alignment\/Map format and SAMtools. Bioinformatics 25:2078\u20132079. https:\/\/doi.org\/10.1093\/bioinformatics\/btp352","journal-title":"Bioinformatics"},{"key":"2683_CR39","doi-asserted-by":"publisher","first-page":"1343","DOI":"10.1038\/s41467-020-15198-9","volume":"11","author":"M Li","year":"2020","unstructured":"Li M, Nishio S-Y, Naruse C et al (2020) Digenic inheritance of mutations in EPHA2 and SLC26A4 in Pendred syndrome. Nat Commun 11:1343. https:\/\/doi.org\/10.1038\/s41467-020-15198-9","journal-title":"Nat Commun"},{"key":"2683_CR40","doi-asserted-by":"publisher","first-page":"3146","DOI":"10.1093\/ndt\/gfn208","volume":"23","author":"M L\u00f6wik","year":"2008","unstructured":"L\u00f6wik M, Levtchenko E, Westra D et al (2008) Bigenic heterozygosity and the development of steroid-resistant focal segmental glomerulosclerosis. Nephrol Dial Transplant off Publ Eur Dial Transpl Assoc -. Eur Ren Assoc 23:3146\u20133151. https:\/\/doi.org\/10.1093\/ndt\/gfn208","journal-title":"Eur Ren Assoc"},{"key":"2683_CR41","doi-asserted-by":"publisher","first-page":"31","DOI":"10.4049\/jimmunol.1402639","volume":"195","author":"BA Lubeck","year":"2015","unstructured":"Lubeck BA, Lapinski PE, Oliver JA et al (2015) Cutting Edge: Codeletion of the ras GTPase-Activating proteins (RasGAPs) Neurofibromin 1 and p120 RasGAP in T cells results in the development of T cell Acute Lymphoblastic Leukemia. J Immunol 195:31\u201335. https:\/\/doi.org\/10.4049\/jimmunol.1402639","journal-title":"J Immunol"},{"key":"2683_CR42","doi-asserted-by":"publisher","first-page":"1012","DOI":"10.1016\/j.cell.2015.04.004","volume":"161","author":"DG Lupi\u00e1\u00f1ez","year":"2015","unstructured":"Lupi\u00e1\u00f1ez DG, Kraft K, Heinrich V et al (2015) Disruptions of topological chromatin domains cause pathogenic rewiring of gene-enhancer interactions. Cell 161:1012\u20131025. https:\/\/doi.org\/10.1016\/j.cell.2015.04.004","journal-title":"Cell"},{"key":"2683_CR43","doi-asserted-by":"publisher","first-page":"823","DOI":"10.1126\/science.1215040","volume":"335","author":"DG MacArthur","year":"2012","unstructured":"MacArthur DG, Balasubramanian S, Frankish A et al (2012) A systematic survey of loss-of-function variants in human protein-coding genes. Sci (80-) 335:823\u2013828. https:\/\/doi.org\/10.1126\/science.1215040","journal-title":"Sci (80-)"},{"key":"2683_CR44","doi-asserted-by":"publisher","first-page":"623","DOI":"10.1136\/jmg.2009.075937","volume":"47","author":"V-F Mautner","year":"2010","unstructured":"Mautner V-F, Kluwe L, Friedrich RE et al (2010) Clinical characterisation of 29 neurofibromatosis type-1 patients with molecularly ascertained 1.4\u00a0mb type-1 NF1 deletions. J Med Genet 47:623\u2013630. https:\/\/doi.org\/10.1136\/jmg.2009.075937","journal-title":"J Med Genet"},{"key":"2683_CR45","doi-asserted-by":"publisher","first-page":"803","DOI":"10.1007\/s10815-020-01736-3","volume":"37","author":"ML Mebrek","year":"2020","unstructured":"Mebrek ML, Cl\u00e8de S, de Chalus A et al (2020) Simple FISH-based evaluation of spermatic nuclear architecture shows an abnormal chromosomal organization in balanced chromosomal rearrangement carriers. J Assist Reprod Genet 37:803\u2013809. https:\/\/doi.org\/10.1007\/s10815-020-01736-3","journal-title":"J Assist Reprod Genet"},{"key":"2683_CR46","doi-asserted-by":"publisher","first-page":"e08","DOI":"10.1136\/jmg.2005.034256","volume":"43","author":"KA Mensink","year":"2005","unstructured":"Mensink KA (2005) Connective tissue dysplasia in five new patients with NF1 microdeletions: further expansion of phenotype and review of the literature. J Med Genet 43:e08\u2013e08. https:\/\/doi.org\/10.1136\/jmg.2005.034256","journal-title":"J Med Genet"},{"key":"2683_CR47","doi-asserted-by":"publisher","first-page":"213","DOI":"10.1002\/humu.21418","volume":"32","author":"L Messiaen","year":"2011","unstructured":"Messiaen L, Vogt J, Bengesser K et al (2011) Mosaic type-1 NF1 microdeletions as a cause of both generalized and segmental neurofibromatosis type-1 (NF1). Hum Mutat 32:213\u2013219. https:\/\/doi.org\/10.1002\/humu.21418","journal-title":"Hum Mutat"},{"key":"2683_CR48","doi-asserted-by":"publisher","first-page":"177","DOI":"10.1016\/j.hoc.2017.11.003","volume":"32","author":"S Mettananda","year":"2018","unstructured":"Mettananda S, Higgs DR (2018) Molecular basis and genetic modifiers of Thalassemia. Hematol Oncol Clin North Am 32:177\u2013191. https:\/\/doi.org\/10.1016\/j.hoc.2017.11.003","journal-title":"Hematol Oncol Clin North Am"},{"key":"2683_CR49","doi-asserted-by":"publisher","first-page":"4284","DOI":"10.1093\/nar\/gkq1224","volume":"39","author":"M Minczuk","year":"2011","unstructured":"Minczuk M, He J, Duch AM et al (2011) TEFM (c17orf42) is necessary for transcription of human mtDNA. Nucleic Acids Res 39:4284\u20134299. https:\/\/doi.org\/10.1093\/nar\/gkq1224","journal-title":"Nucleic Acids Res"},{"key":"2683_CR50","doi-asserted-by":"publisher","first-page":"1412","DOI":"10.1128\/MCB.01177-13","volume":"34","author":"G Morlino","year":"2014","unstructured":"Morlino G, Barreiro O, Baixauli F et al (2014) Miro-1 Links Mitochondria and Microtubule Dynein Motors to control Lymphocyte Migration and Polarity. Mol Cell Biol 34:1412\u20131426. https:\/\/doi.org\/10.1128\/MCB.01177-13","journal-title":"Mol Cell Biol"},{"key":"2683_CR51","doi-asserted-by":"publisher","first-page":"E1506","DOI":"10.1002\/humu.21271","volume":"31","author":"E Pasmant","year":"2010","unstructured":"Pasmant E, Sabbagh A, Spurlock G et al (2010) NF1 microdeletions in neurofibromatosis type 1: from genotype to phenotype. Hum Mutat 31:E1506\u2013E1518. https:\/\/doi.org\/10.1002\/humu.21271","journal-title":"Hum Mutat"},{"key":"2683_CR52","doi-asserted-by":"publisher","first-page":"483","DOI":"10.1136\/jmedgenet-2012-100978","volume":"49","author":"E Pasmant","year":"2012","unstructured":"Pasmant E, Vidaud M, Vidaud D, Wolkenstein P (2012) Neurofibromatosis type 1: from genotype to phenotype. J Med Genet 49:483\u2013489. https:\/\/doi.org\/10.1136\/jmedgenet-2012-100978","journal-title":"J Med Genet"},{"key":"2683_CR53","doi-asserted-by":"publisher","first-page":"841","DOI":"10.1093\/bioinformatics\/btq033","volume":"26","author":"AR Quinlan","year":"2010","unstructured":"Quinlan AR, Hall IM (2010) BEDTools: a flexible suite of utilities for comparing genomic features. Bioinformatics 26:841\u2013842. https:\/\/doi.org\/10.1093\/bioinformatics\/btq033","journal-title":"Bioinformatics"},{"key":"2683_CR54","doi-asserted-by":"publisher","first-page":"1419","DOI":"10.1038\/ng1920","volume":"38","author":"T Raedt","year":"2006","unstructured":"Raedt T, De, Stephens M, Heyns I et al (2006) Conservation of hotspots for recombination in low-copy repeats associated with the NF1 microdeletion. Nat Genet 38:1419\u20131423. https:\/\/doi.org\/10.1038\/ng1920","journal-title":"Nat Genet"},{"key":"2683_CR55","doi-asserted-by":"publisher","first-page":"435","DOI":"10.1038\/s42003-023-04819-w","volume":"6","author":"S Rajderkar","year":"2023","unstructured":"Rajderkar S, Barozzi I, Zhu Y et al (2023) Topologically associating domain boundaries are required for normal genome function. Commun Biol 6:435. https:\/\/doi.org\/10.1038\/s42003-023-04819-w","journal-title":"Commun Biol"},{"key":"2683_CR56","doi-asserted-by":"publisher","first-page":"169","DOI":"10.1002\/wdev.218","volume":"5","author":"S Remeseiro","year":"2016","unstructured":"Remeseiro S, H\u00f6rnblad A, Spitz F (2016) Gene regulation during development in the light of topologically associating domains. Wiley Interdiscip Rev Dev Biol 5:169\u2013185. https:\/\/doi.org\/10.1002\/wdev.218","journal-title":"Wiley Interdiscip Rev Dev Biol"},{"key":"2683_CR57","doi-asserted-by":"publisher","first-page":"405","DOI":"10.1038\/gim.2015.30","volume":"17","author":"S Richards","year":"2015","unstructured":"Richards S, Aziz N, Bale S et al (2015) Standards and guidelines for the interpretation of sequence variants: a joint consensus recommendation of the American College of Medical Genetics and Genomics and the Association for Molecular Pathology. Genet Med off J Am Coll Med Genet 17:405\u2013424. https:\/\/doi.org\/10.1038\/gim.2015.30","journal-title":"Genet Med off J Am Coll Med Genet"},{"key":"2683_CR58","doi-asserted-by":"publisher","first-page":"W350","DOI":"10.1093\/nar\/gky300","volume":"46","author":"CHM Rodrigues","year":"2018","unstructured":"Rodrigues CHM, Pires DEV, Ascher DB (2018) DynaMut: predicting the impact of mutations on protein conformation, flexibility and stability. Nucleic Acids Res 46:W350\u2013W355. https:\/\/doi.org\/10.1093\/nar\/gky300","journal-title":"Nucleic Acids Res"},{"key":"2683_CR59","doi-asserted-by":"publisher","first-page":"444","DOI":"10.1016\/j.ajhg.2017.01.028","volume":"100","author":"FA Santoni","year":"2017","unstructured":"Santoni FA, Stamoulis G, Garieri M et al (2017) Detection of imprinted genes by single-cell allele-specific gene expression. Am J Hum Genet 100:444\u2013453. https:\/\/doi.org\/10.1016\/j.ajhg.2017.01.028","journal-title":"Am J Hum Genet"},{"key":"2683_CR60","doi-asserted-by":"publisher","first-page":"34","DOI":"10.1186\/s13023-017-0588-2","volume":"12","author":"L Sellmer","year":"2017","unstructured":"Sellmer L, Farschtschi S, Marangoni M et al (2017) Non-optic glioma in adults and children with neurofibromatosis 1. Orphanet J Rare Dis 12:34. https:\/\/doi.org\/10.1186\/s13023-017-0588-2","journal-title":"Orphanet J Rare Dis"},{"key":"2683_CR61","doi-asserted-by":"publisher","first-page":"193","DOI":"10.1038\/bjc.2012.535","volume":"108","author":"OO Seminog","year":"2013","unstructured":"Seminog OO, Goldacre MJ (2013) Risk of benign tumours of nervous system, and of malignant neoplasms, in people with neurofibromatosis: population-based record-linkage study. Br J Cancer 108:193\u2013198. https:\/\/doi.org\/10.1038\/bjc.2012.535","journal-title":"Br J Cancer"},{"key":"2683_CR62","doi-asserted-by":"publisher","first-page":"308","DOI":"10.1093\/nar\/29.1.308","volume":"29","author":"ST Sherry","year":"2001","unstructured":"Sherry ST (2001) dbSNP: the NCBI database of genetic variation. Nucleic Acids Res 29:308\u2013311. https:\/\/doi.org\/10.1093\/nar\/29.1.308","journal-title":"Nucleic Acids Res"},{"key":"2683_CR63","doi-asserted-by":"publisher","unstructured":"Shirokova OM, Pchelin PV, Mukhina IV (2020) MERCs. The Novel Assistant to Neurotransmission? Front Neurosci 14:. https:\/\/doi.org\/10.3389\/fnins.2020.589319","DOI":"10.3389\/fnins.2020.589319"},{"key":"2683_CR64","doi-asserted-by":"publisher","first-page":"453","DOI":"10.1038\/s41576-018-0007-0","volume":"19","author":"M Spielmann","year":"2018","unstructured":"Spielmann M, Lupi\u00e1\u00f1ez DG, Mundlos S (2018) Structural variation in the 3D genome. Nat Rev Genet 19:453\u2013467. https:\/\/doi.org\/10.1038\/s41576-018-0007-0","journal-title":"Nat Rev Genet"},{"key":"2683_CR65","doi-asserted-by":"publisher","first-page":"511","DOI":"10.1007\/s00439-018-1904-1","volume":"137","author":"A Summerer","year":"2018","unstructured":"Summerer A, Mautner V-F, Upadhyaya M et al (2018) Extreme clustering of type-1 NF1 deletion breakpoints co-locating with G-quadruplex forming sequences. Hum Genet 137:511\u2013520. https:\/\/doi.org\/10.1007\/s00439-018-1904-1","journal-title":"Hum Genet"},{"key":"2683_CR66","doi-asserted-by":"publisher","first-page":"D941","DOI":"10.1093\/nar\/gky1015","volume":"47","author":"JG Tate","year":"2019","unstructured":"Tate JG, Bamford S, Jubb HC et al (2019) COSMIC: the catalogue of somatic mutations in Cancer. Nucleic Acids Res 47:D941\u2013D947. https:\/\/doi.org\/10.1093\/nar\/gky1015","journal-title":"Nucleic Acids Res"},{"key":"2683_CR68","doi-asserted-by":"publisher","unstructured":"Tritto V, Eoli M, Paterra R et al (2022) Characterization of 22q12 microdeletions causing position effect in rare NF2 patients with Complex Phenotypes. Int J Mol Sci 23. https:\/\/doi.org\/10.3390\/ijms231710017","DOI":"10.3390\/ijms231710017"},{"key":"2683_CR67","doi-asserted-by":"publisher","unstructured":"Tritto V, Capitanio D, Gelfi C, Riva P (2023a) Changes of RAS Pathway Phosphorylation in Lymphoblastoid Cell lines from Noonan Syndrome patients carrying hypomorphic variants in two NS genes. Int J Mol Sci 24. https:\/\/doi.org\/10.3390\/ijms24044035","DOI":"10.3390\/ijms24044035"},{"key":"2683_CR69","doi-asserted-by":"publisher","first-page":"181","DOI":"10.1007\/s10048-023-00718-8","volume":"24","author":"V Tritto","year":"2023","unstructured":"Tritto V, Grilli F, Milani D, Riva P (2023b) Deregulated expression of polycomb repressive complex 2 target genes in a NF1 patient with microdeletion generating the RNF135-SUZ12 chimeric gene. Neurogenetics 24:181\u2013188. https:\/\/doi.org\/10.1007\/s10048-023-00718-8","journal-title":"Neurogenetics"},{"key":"2683_CR70","doi-asserted-by":"publisher","first-page":"1978","DOI":"10.1200\/JCO.2015.65.3576","volume":"34","author":"E Uusitalo","year":"2016","unstructured":"Uusitalo E, Rantanen M, Kallionp\u00e4\u00e4 RA et al (2016) Distinctive Cancer associations in patients with neurofibromatosis type 1. J Clin Oncol 34:1978\u20131986. https:\/\/doi.org\/10.1200\/JCO.2015.65.3576","journal-title":"J Clin Oncol"},{"key":"2683_CR71","doi-asserted-by":"publisher","first-page":"35","DOI":"10.1136\/jmg.2003.014761","volume":"41","author":"M Venturin","year":"2004","unstructured":"Venturin M (2004) Mental retardation and cardiovascular malformations in NF1 microdeleted patients point to candidate genes in 17q11.2. J Med Genet 41:35\u201341. https:\/\/doi.org\/10.1136\/jmg.2003.014761","journal-title":"J Med Genet"},{"key":"2683_CR72","doi-asserted-by":"publisher","first-page":"508","DOI":"10.1111\/j.1529-8817.2005.00203.x","volume":"69","author":"M Venturin","year":"2005","unstructured":"Venturin M, Bentivegna A, Moroni R et al (2005) Evidence by Expression Analysis of Candidate Genes for Congenital Heart Defects in the NF1 microdeletion interval. Ann Hum Genet 69:508\u2013516. https:\/\/doi.org\/10.1111\/j.1529-8817.2005.00203.x","journal-title":"Ann Hum Genet"},{"key":"2683_CR73","doi-asserted-by":"publisher","first-page":"436","DOI":"10.1136\/jmedgenet-2013-102240","volume":"51","author":"M Venturin","year":"2014","unstructured":"Venturin M, Carra S, Gaudenzi G et al (2014) ADAP2 in heart development: a candidate gene for the occurrence of cardiovascular malformations in NF1 microdeletion syndrome. J Med Genet 51:436\u2013443. https:\/\/doi.org\/10.1136\/jmedgenet-2013-102240","journal-title":"J Med Genet"},{"key":"2683_CR74","doi-asserted-by":"publisher","DOI":"10.1371\/journal.pone.0049229","author":"R Visser","year":"2012","unstructured":"Visser R, Landman EBM, Goeman J et al (2012) Sotos Syndrome is Associated with Deregulation of the MAPK\/ERK-Signaling pathway. PLoS ONE. https:\/\/doi.org\/10.1371\/journal.pone.0049229","journal-title":"PLoS ONE"},{"key":"2683_CR75","doi-asserted-by":"publisher","first-page":"R80","DOI":"10.1186\/gb-2014-15-6-r80","volume":"15","author":"J Vogt","year":"2014","unstructured":"Vogt J, Bengesser K, Claes KBM et al (2014) SVA retrotransposon insertion-associated deletion represents a novel mutational mechanism underlying large genomic copy number changes with non-recurrent breakpoints. Genome Biol 15:R80. https:\/\/doi.org\/10.1186\/gb-2014-15-6-r80","journal-title":"Genome Biol"},{"key":"2683_CR76","doi-asserted-by":"publisher","first-page":"e164","DOI":"10.1093\/nar\/gkq603","volume":"38","author":"K Wang","year":"2010","unstructured":"Wang K, Li M, Hakonarson H (2010) ANNOVAR: functional annotation of genetic variants from high-throughput sequencing data. Nucleic Acids Res 38:e164. https:\/\/doi.org\/10.1093\/nar\/gkq603","journal-title":"Nucleic Acids Res"},{"key":"2683_CR77","doi-asserted-by":"publisher","first-page":"158","DOI":"10.1016\/j.devcel.2011.11.012","volume":"22","author":"R Wimmer","year":"2012","unstructured":"Wimmer R, Cseh B, Maier B et al (2012) Angiogenic sprouting requires the fine tuning of endothelial cell cohesion by the Raf-1\/Rok-\u03b1 complex. Dev Cell 22:158\u2013171. https:\/\/doi.org\/10.1016\/j.devcel.2011.11.012","journal-title":"Dev Cell"},{"key":"2683_CR78","doi-asserted-by":"publisher","first-page":"1589","DOI":"10.1093\/bioinformatics\/btw031","volume":"32","author":"DK Witvliet","year":"2016","unstructured":"Witvliet DK, Strokach A, Giraldo-Forero AF et al (2016) ELASPIC web-server: proteome-wide structure-based prediction of mutation effects on protein stability and binding affinity. Bioinformatics 32:1589\u20131591. https:\/\/doi.org\/10.1093\/bioinformatics\/btw031","journal-title":"Bioinformatics"},{"key":"2683_CR79","doi-asserted-by":"publisher","DOI":"10.1038\/s41431-018-0196-1","author":"WL Wooderchak-Donahue","year":"2018","unstructured":"Wooderchak-Donahue WL, Johnson P, McDonald J et al (2018) Expanding the clinical and molecular findings in RASA1 capillary malformation-arteriovenous malformation. Eur J Hum Genet. https:\/\/doi.org\/10.1038\/s41431-018-0196-1","journal-title":"Eur J Hum Genet"},{"key":"2683_CR81","doi-asserted-by":"publisher","first-page":"151","DOI":"10.1042\/0264-6021:3510151","volume":"351","author":"W YIP-SCHNEIDER MT, MIAO","year":"2000","unstructured":"YIP-SCHNEIDER MT, MIAO W, LIN A et al (2000) Regulation of the Raf-1 kinase domain by phosphorylation and 14-3-3 association. Biochem J 351:151. https:\/\/doi.org\/10.1042\/0264-6021:3510151","journal-title":"Biochem J"},{"key":"2683_CR82","doi-asserted-by":"publisher","first-page":"NP13","DOI":"10.1177\/0883073813492384","volume":"29","author":"YA Zarate","year":"2014","unstructured":"Zarate YA, Lichty AW, Champion KJ et al (2014) Unique cerebrovascular anomalies in Noonan Syndrome with RAF1 mutation. J Child Neurol 29:NP13\u2013NP17. https:\/\/doi.org\/10.1177\/0883073813492384","journal-title":"J Child Neurol"},{"key":"2683_CR83","doi-asserted-by":"publisher","first-page":"11291","DOI":"10.1038\/srep11291","volume":"5","author":"J Zhang","year":"2015","unstructured":"Zhang J, Tong H, Fu X et al (2015) Molecular characterization of NF1 and neurofibromatosis type 1 genotype-phenotype correlations in a Chinese Population. Sci Rep 5:11291. https:\/\/doi.org\/10.1038\/srep11291","journal-title":"Sci Rep"}],"container-title":["Human Genetics"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/link.springer.com\/content\/pdf\/10.1007\/s00439-024-02683-0.pdf","content-type":"application\/pdf","content-version":"vor","intended-application":"text-mining"},{"URL":"https:\/\/link.springer.com\/article\/10.1007\/s00439-024-02683-0\/fulltext.html","content-type":"text\/html","content-version":"vor","intended-application":"text-mining"},{"URL":"https:\/\/link.springer.com\/content\/pdf\/10.1007\/s00439-024-02683-0.pdf","content-type":"application\/pdf","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2024,6,20]],"date-time":"2024-06-20T02:02:41Z","timestamp":1718848961000},"score":1,"resource":{"primary":{"URL":"https:\/\/link.springer.com\/10.1007\/s00439-024-02683-0"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2024,6]]},"references-count":82,"journal-issue":{"issue":"6","published-print":{"date-parts":[[2024,6]]}},"alternative-id":["2683"],"URL":"https:\/\/doi.org\/10.1007\/s00439-024-02683-0","relation":{},"ISSN":["0340-6717","1432-1203"],"issn-type":[{"value":"0340-6717","type":"print"},{"value":"1432-1203","type":"electronic"}],"subject":[],"published":{"date-parts":[[2024,6]]},"assertion":[{"value":"15 March 2024","order":1,"name":"received","label":"Received","group":{"name":"ArticleHistory","label":"Article History"}},{"value":"3 June 2024","order":2,"name":"accepted","label":"Accepted","group":{"name":"ArticleHistory","label":"Article History"}},{"value":"14 June 2024","order":3,"name":"first_online","label":"First Online","group":{"name":"ArticleHistory","label":"Article History"}},{"order":1,"name":"Ethics","group":{"name":"EthicsHeading","label":"Declarations"}},{"value":"The study was conducted according to the guidelines of the Declaration of Helsinki and approved by the Fondazione IRCCS Istituto Neurologico Carlo Besta Ethical Committee and Scientific Board (N\u00b050\u2009\u2212\u200919\/3\/2018).","order":2,"name":"Ethics","group":{"name":"EthicsHeading","label":"Ethical approval"}},{"value":"Written informed consent was obtained from all subjects involved in the study.","order":3,"name":"Ethics","group":{"name":"EthicsHeading","label":"Consent to participate"}},{"value":"The authors declare no competing interests.","order":4,"name":"Ethics","group":{"name":"EthicsHeading","label":"Competing interests"}}]}}