To date, more than 50 loci associated with COVID-19 susceptibility, hospitalization, and severity have been identified using genome-wide association studies (GWAS) (Kanai et al., 2023; Pairo-Castineira et al., 2023). The COVID-19 Host Genetics Initiative (HGI) has made significant efforts (Niemi et al., 2021) to augment the power to identify disease loci by recruiting individuals from diverse populations and conducting a trans-ancestry meta-analysis. Despite this, the lack of genetic diversity and a focus on cases of European ancestries still predominate in the studies (Popejoy and Fullerton, 2016; Sirugo et al., 2019). In addition, while trans-ancestry meta-analyses are a powerful approach for discovering shared genetic risk variants with similar effects across populations (Li and Keating, 2014), they may fail to identify risk variants that have larger effects on particular underrepresented populations. Genetic disease risk has been shaped by the particular evolutionary history of populations and environmental exposures (Rosenberg et al., 2010). Their action is particularly important for infectious diseases due to the selective constraints that are imposed by host‒pathogen interactions (Karlsson et al., 2014; Kwok et al., 2021). Literature examples of this in COVID-19 severity include a DOCK2 gene variant in East Asians (Namkoong et al., 2022) and frequent loss-of-function variants in IFNAR1 and IFNAR2 genes in Polynesian and Inuit populations, respectively (Bastard et al., 2022; Duncan et al., 2022).

Including diverse populations in case‒control GWAS with unrelated participants usually requires a prior classification of individuals in genetically homogeneous groups, which are typically analyzed separately to control the population stratification effects (Peterson et al., 2019). Populations with recent admixture impose an additional challenge to GWASs due to their complex genetic diversity and linkage disequilibrium (LD) patterns, requiring the development of alternative approaches and a careful inspection of results to reduce false positives due to population structure (Rosenberg et al., 2010). In fact, there are benefits in study power from modeling the admixed ancestries either locally, at the regional scale in the chromosomes, or globally, across the genome, depending on factors such as the heterogeneity of the risk variant in frequencies or the effects among the ancestry strata (Mester et al., 2023). Despite the development of novel methods specifically tailored for the analysis of admixed populations (Atkinson et al., 2021), the lack of a standardized analysis framework and the difficulties in confidently clustering admixed individuals into particular genetic groups often lead to their exclusion from GWAS.

The Spanish Coalition to Unlock Research on Host Genetics on COVID-19 (SCOURGE) recruited COVID-19 patients between March and December 2020 from hospitals across Spain and from March 2020 to July 2021 in Latin America (https://www.scourge-covid.org). A first GWAS of COVID-19 severity among Spanish patients of European descent revealed novel disease loci and explored age- and sex-varying effects of genetic factors (Cruz et al., 2022). Here, we present the findings of a GWAS meta-analysis in admixed Latin American (AMR) populations, comprising individuals from the SCOURGE Latin American cohort and the HGI studies, which allowed us to identify two novel severe COVID-19 loci, BAZ2B and DDIAS. Further analyses modeling the admixture from three genetic ancestral components and performing a trans-ethnic meta-analysis led to the identification of an additional risk locus near CREBBP. We finally assessed a cross-ancestry polygenic risk score (PGS) model with variants associated with critical COVID-19.

We have conducted the largest GWAS meta-analysis of COVID-19 hospitalization in admixed AMR to date. While the genetic risk basis discovered for COVID-19 is largely shared among populations, trans-ancestry meta-analyses on this disease have primarily included EUR samples. This dominance of GWAS in Europeans and the subsequent bias in sample sizes can mask population-specific genetic risks (i.e., variants that are monomorphic in some populations) or be less powered to detect risk variants having higher allele frequencies in population groups other than Europeans. In this sense, after combining data from admixed AMR patients, we found two risk loci that were first discovered in a GWAS of Latin American populations. Interestingly, the sentinel variant rs77599934 in the DDIAS gene is a rare coding variant (~1% for allele G) with a large effect on COVID-19 hospitalization that is nearly monomorphic in most of the other populations. This has likely led to its exclusion from the cross-population meta-analyses conducted to date, remaining undetectable.

Fine mapping of the region harboring DDIAS did not reveal further information about which gene could be the more prone to be causal or about the functional consequences of the risk variant, but our sentinel variant was in strong LD with an eQTL that associated with reduced gene expression of DDIAS in the lung. DDIAS, known as damage-induced apoptosis suppressor gene, is itself a plausible candidate gene. It has been linked to DNA damage repair mechanisms: research has shown that depletion of DDIAS leads to an increase in ATM phosphorylation and the formation of p53-binding protein (53BP1) foci, a known biomarker of DNA double-strand breaks, suggesting a potential role in double-strand break repair (Brunette et al., 2019). Interestingly, a study found that infection by SARS-CoV-2 also triggered the phosphorylation of ATM kinase and inhibited repair mechanisms, causing the accumulation of DNA damage (Gioia et al., 2023). This gene has also been proposed as a potential biomarker for lung cancer after finding that it interacts with STAT3 in lung cancer cells, regulating IL-6 (Im et al., 2020; Im et al., 2023) and thus mediating inflammatory processes, while another study determined that its blockade inhibited lung cancer cell growth (Won et al., 2014). Another prioritized gene from this region was PRCP, an angiotensinase that shares substrate specificity with ACE2 receptor. It has been positively linked to hypertension and some studies have raised hypotheses on its role in COVID-19 progression, particularly in relation to the development of pro-thrombotic events (Angeli et al., 2023; Silva-Aguiar et al., 2020).

The risk region found in chromosome 2 harbors more than one gene. The lead variant rs13003835 is located within BAZ2B, and it increases the expression of the antisense BAZ2B gene in whole blood. BAZ2B encodes one of the regulatory subunits of the Imitation switch (ISWI) chromatin remodelers (Li et al., 2021) constituting the BRF-1/BRF-5 complexes with SMARCA1 and SMARCA5, respectively. Interestingly, it was discovered that lnc-BAZ2B promotes macrophage activation through regulation of BAZ2B expression. Its overexpression resulted in pulmonary inflammation and elevated levels of MUC5AC in mice with asthma (Xia et al., 2021). This variant was also an eQTL for LY75 (encoding lymphocyte antigen 75) in the esophageal mucosa tissue. Lymphocyte antigen 75 is involved in immune processes through antigen presentation in dendritic cells and endocytosis (Mahnke et al., 2000) and has been associated with inflammatory diseases, representing a compelling candidate for the region. Increased expression of LY75 has been detected within hours after infection by SARS-CoV-2 (Mitchell et al., 2013; Sims et al., 2013). It is worth noting that differences in AF for this variant suggest that analyses in AMR populations might be more powered to detect the association, supporting the necessity of population-specific studies.

A third novel risk region was observed on chromosome 15 between the IQGAP1 and ZNF774 genes, although it did not reach genome-wide significance.

Secondary analyses revealed five TWAS-associated genes, some of which have already been linked to severe COVID-19. In a comprehensive multitissue gene expression profiling study (Gómez-Carballa et al., 2022), decreased expression of CAMP and S100A8/S100A9 genes in patients with severe COVID-19 was observed, while another study detected the upregulation of SCL25A37 among patients with severe COVID-19 (Policard et al., 2021). SMARCC1 is a subunit of the SWI/SNF chromatin remodeling complex that has been identified as proviral for SARS-CoV-2 and other coronavirus strains through a genome-wide screen (Wei et al., 2021). This complex is crucial for ACE2 expression and viral entry into the cell (Wei et al., 2023). However, it should be noted that using eQTL mostly from European populations such as those in GTEx could result in reduced power to detect associations.

To explore the genetic architecture of the trait among admixed AMR populations, we performed two cross-ancestry meta-analyses including the SCOURGE Latin American cohort GWAS findings. We found that the two novel risk variants were not associated with COVID-19 hospitalization outside the population-specific meta-analysis, highlighting the importance of complementing trans-ancestry meta-analyses with group-specific analyses. Notably, this analysis did not replicate the association at the DSTYK locus, which was associated with severe COVID-19 in Brazilian individuals with higher European admixture (Pereira et al., 2022). This lack of replication aligns with the initial hypothesis of that study suggesting that the risk haplotype was derived from European populations, as we reduced the weight of this ancestral contribution in our study by excluding those individuals.

Moreover, these cross-ancestry meta-analyses pointed to three loci that were not genome-wide significant in the HGIv7 ALL meta-analysis: a novel locus at CREBBP and two loci at ZBTB7A and CASC20 that were reported in another meta-analysis. CREBBP and ZBTB7A achieved a stronger significance when considering only the EUR, AFR, and admixed AMR GIA groups. According to a recent study, elevated levels of the ZBTB7A gene promote a quasihomeostatic state between coronaviruses and host cells, preventing cell death by regulating oxidative stress pathways (Zhu et al., 2022). This gene is involved in several signaling pathways, such as B- and T-cell differentiation (Gupta et al., 2020). On a separate note, CREBBP encodes the CREB binding protein (CBP), which is involved in transcriptional activation and is known to positively regulate the type I interferon response through virus-induced phosphorylation of IRF-3 (Yoneyama et al., 1998). In addition, the CREBP/CBP interaction has been implicated in SARS-CoV-2 infection (Yang et al., 2023) via the cAMP/PKA pathway. In fact, cells with suppressed CREBBP gene expression exhibit reduced replication of the so-called Delta and Omicron SARS-CoV-2 variants (Yang et al., 2023).

We developed a cross-population PGS model, which effectively stratified individuals based on their genetic risk and demonstrated consistency with the clinical severity classification of the patients. Only a few polygenic scores were derived from COVID-19 GWAS data. Horowitz et al., 2022 developed a score using 6 and 12 associated variants (PGS ID: PGP000302) and reported an associated OR (top 10% vs rest) of 1.38 for risk of hospitalization in European populations, whereas the OR for Latin American populations was 1.56. Since their sample size and the number of variants included in the PGS were lower, direct comparisons are not straightforward. Nevertheless, our analysis provides the first results for a PRS applied to a relatively large AMR cohort, being of value for future analyses regarding PRS transferability.

This study is subject to limitations, mostly concerning sample recruitment and composition. The SCOURGE Latino American sample size is small, and the GWAS is likely underpowered. Another limitation is the difference in case‒control recruitment across sampling regions that, yet controlled for, may reduce the ability to observe significant associations driven by different compositions of the populations. In this sense, the identified risk loci might not replicate in a cohort lacking any of the parental population sources from the three-way admixture. Likewise, we could not explicitly control for socioenvironmental factors that could have affected COVID-19 spread and hospitalization rates, although genetic principal components are known to capture nongenetic factors. Finally, we must acknowledge the lack of a replication cohort. We used all the available GWAS data for COVID-19 hospitalization in admixed AMR in this meta-analysis due to the low number of studies conducted. Therefore, we had no studies to replicate or validate the results. These concerns may be addressed in the future by including more AMR GWAS in the meta-analysis, both by involving diverse populations in study designs and supporting research from countries in Latin America.

This study provides novel insights into the genetic basis of COVID-19 severity, emphasizing the importance of considering host genetic factors by using non-European populations, especially of admixed sources. Such complementary efforts can pin down new variants and increase our knowledge on the host genetic factors of severe COVID-19.

Summary statistics from the SCOURGE Latin American GWAS and the analysis scripts are available from the public repository https://github.com/CIBERER/Scourge-COVID19 (copy archived at CIBERER, 2024).

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Author details

1. Silvia Diz-de Almeida

   1. ERN-ITHACA-European Reference Network, Soria, Spain
   2. Pediatric Neurology Unit, Department of Pediatrics, Navarra Health Service Hospital, Pamplona, Spain
   3. CIBERER, ISCIII, Madrid, Spain
   4. Centro Singular de Investigación en Medicina Molecular y Enfermedades Crónicas (CIMUS), Universidade de Santiago de Compostela, Santiago de Compostela, Spain

   Contribution

   Conceptualization, Formal analysis, Validation, Investigation, Visualization, Methodology, Writing – original draft, Writing – review and editing

   Contributed equally with

   Raquel Cruz

   Competing interests

   No competing interests declared

   "This ORCID iD identifies the author of this article:" 0000-0003-2813-8928

2. Raquel Cruz

   1. ERN-ITHACA-European Reference Network, Soria, Spain
   2. Pediatric Neurology Unit, Department of Pediatrics, Navarra Health Service Hospital, Pamplona, Spain
   3. CIBERER, ISCIII, Madrid, Spain
   4. Centro Singular de Investigación en Medicina Molecular y Enfermedades Crónicas (CIMUS), Universidade de Santiago de Compostela, Santiago de Compostela, Spain

   Contribution

   Conceptualization, Formal analysis, Validation, Investigation, Visualization, Methodology, Writing – original draft, Writing – review and editing

   Contributed equally with

   Silvia Diz-de Almeida

   Competing interests

   No competing interests declared

3. Andre D Luchessi

   Universidade Federal do Rio Grande do Norte, Departamento de Analises Clinicas e Toxicologicas, Natal, Brazil

   Contribution

   Formal analysis, Validation, Methodology, Writing – original draft, Writing – review and editing

   Competing interests

   No competing interests declared

4. José M Lorenzo-Salazar

   Genomics Division, Instituto Tecnológico y de Energías Renovables, Santa Cruz de Tenerife, Spain

   Contribution

   Formal analysis, Methodology, Writing – original draft, Writing – review and editing

   Competing interests

   No competing interests declared

5. Miguel López de Heredia

   CIBERER, ISCIII, Madrid, Spain

   Contribution

   Data curation, Project administration, Writing – review and editing

   Competing interests

   No competing interests declared

6. Inés Quintela

   Fundación Pública Galega de Medicina Xenómica, Sistema Galego de Saúde (SERGAS), Santiago de Compostela, Spain

   Contribution

   Data curation, Project administration, Writing – review and editing

   Competing interests

   No competing interests declared

7. Rafaela González-Montelongo

   Genomics Division, Instituto Tecnológico y de Energías Renovables, Santa Cruz de Tenerife, Spain

   Contribution

   Data curation, Writing – review and editing

   Competing interests

   No competing interests declared

8. Vivian Nogueira Silbiger

   Universidade Federal do Rio Grande do Norte, Departamento de Analises Clinicas e Toxicologicas, Natal, Brazil

   Contribution

   Data curation, Writing – review and editing

   Competing interests

   No competing interests declared

   "This ORCID iD identifies the author of this article:" 0000-0002-9252-0278

9. Marta Sevilla Porras

   1. CIBERER, ISCIII, Madrid, Spain
   2. Instituto de Genética Médica y Molecular (INGEMM), Hospital Universitario La Paz IDIPAZ, Madrid, Spain

   Contribution

   Data curation, Writing – review and editing

   Competing interests

   No competing interests declared

10. Jair Antonio Tenorio Castaño

    1. ERN-ITHACA-European Reference Network, Soria, Spain
    2. CIBERER, ISCIII, Madrid, Spain
    3. Instituto de Genética Médica y Molecular (INGEMM), Hospital Universitario La Paz IDIPAZ, Madrid, Spain

    Contribution

    Data curation, Writing – review and editing

    Competing interests

    No competing interests declared

11. Julian Nevado

    1. ERN-ITHACA-European Reference Network, Soria, Spain
    2. CIBERER, ISCIII, Madrid, Spain
    3. Instituto de Genética Médica y Molecular (INGEMM), Hospital Universitario La Paz IDIPAZ, Madrid, Spain

    Contribution

    Data curation, Writing – review and editing

    Competing interests

    No competing interests declared

12. Jose María Aguado

    1. Unit of Infectious Diseases, Hospital Universitario 12 de Octubre, Instituto de Investigación Sanitaria Hospital 12 de Octubre (imas12), Madrid, Spain
    2. Spanish Network for Research in Infectious Diseases (REIPI RD16/0016/0002), Instituto de Salud Carlos III, Madrid, Spain
    3. CIBERINFEC, ISCIII, Madrid, Spain

    Contribution

    Data curation, Writing – review and editing

    Competing interests

    No competing interests declared

13. Carlos Aguilar

    Hospital General Santa Bárbara de Soria, Soria, Spain

    Contribution

    Data curation, Writing – review and editing

    Competing interests

    No competing interests declared

14. Sergio Aguilera-Albesa

    1. Pediatric Neurology Unit, Department of Pediatrics, Navarra Health Service Hospital, Pamplona, Spain
    2. Navarra Health Service, NavarraBioMed Research Group, Pamplona, Spain

    Contribution

    Data curation, Writing – review and editing

    Competing interests

    No competing interests declared

15. Virginia Almadana

    Hospital Universitario Virgen Macarena, Neumología, Sevilla, Spain

    Contribution

    Data curation, Writing – review and editing

    Competing interests

    No competing interests declared

16. Berta Almoguera

    1. CIBERER, ISCIII, Madrid, Spain
    2. Department of Genetics & Genomics, Instituto de Investigación Sanitaria-Fundación Jiménez Díaz University Hospital - Universidad Autónoma de Madrid (IIS-FJD, UAM), Madrid, Spain

    Contribution

    Data curation, Writing – review and editing

    Competing interests

    No competing interests declared

17. Nuria Alvarez

    Spanish National Cancer Research Centre, Human Genotyping-CEGEN Unit, Madrid, Spain

    Contribution

    Data curation, Writing – review and editing

    Competing interests

    No competing interests declared

18. Álvaro Andreu-Bernabeu

    1. Department of Child and Adolescent Psychiatry, Institute of Psychiatry and Mental Health, Hospital General Universitario Gregorio Marañón (IiSGM), Madrid, Spain
    2. School of Medicine, Universidad Complutense, Madrid, Spain

    Contribution

    Data curation, Writing – review and editing

    Competing interests

    No competing interests declared

19. Eunate Arana-Arri

    1. Biocruces Bizkai HRI, Bizkaia, Spain
    2. Cruces University Hospital, Osakidetza, Bizkaia, Spain

    Contribution

    Data curation, Writing – review and editing

    Competing interests

    No competing interests declared

20. Celso Arango

    1. Department of Child and Adolescent Psychiatry, Institute of Psychiatry and Mental Health, Hospital General Universitario Gregorio Marañón (IiSGM), Madrid, Spain
    2. School of Medicine, Universidad Complutense, Madrid, Spain
    3. Centre for Biomedical Network Research on Mental Health (CIBERSAM), Instituto de Salud Carlos III, Madrid, Spain

    Contribution

    Data curation, Writing – review and editing

    Competing interests

    No competing interests declared

21. María J Arranz

    Fundació Docència I Recerca Mutua Terrassa, Barcelona, Spain

    Contribution

    Data curation, Writing – review and editing

    Competing interests

    No competing interests declared

22. Maria-Jesus Artiga

    Spanish National Cancer Research Centre, CNIO Biobank, Madrid, Spain

    Contribution

    Data curation, Writing – review and editing

    Competing interests

    No competing interests declared

23. Raúl C Baptista-Rosas

    1. Hospital General de Occidente, Zapopan Jalisco, Mexico
    2. Centro Universitario de Tonalá, Universidad de Guadalajara, Tonalá Jalisco, Mexico
    3. Centro de Investigación Multidisciplinario en Salud, Universidad de Guadalajara, Tonalá Jalisco, Mexico

    Contribution

    Data curation, Writing – review and editing

    Competing interests

    No competing interests declared

    "This ORCID iD identifies the author of this article:" 0000-0002-0273-4740

24. María Barreda- Sánchez

    1. Universidad Católica San Antonio de Murcia (UCAM), Murcia, Spain
    2. Instituto Murciano de Investigación Biosanitaria (IMIB-Arrixaca), Murcia, Spain

    Contribution

    Data curation, Writing – review and editing

    Competing interests

    No competing interests declared

25. Moncef Belhassen-Garcia

    Hospital Universitario de Salamanca-IBSAL, Servicio de Medicina Interna-Unidad de Enfermedades Infecciosas, Salamanca, Spain

    Contribution

    Data curation, Writing – review and editing

    Competing interests

    No competing interests declared

26. Joao F Bezerra

    Escola Tecnica de Saúde, Laboratorio de Vigilancia Molecular Aplicada, Brasilia, Brazil

    Contribution

    Data curation, Writing – review and editing

    Competing interests

    No competing interests declared

27. Marcos AC Bezerra

    Federal University of Pernambuco, Genetics Postgraduate Program, Recife, Brazil

    Contribution

    Data curation, Writing – review and editing

    Competing interests

    No competing interests declared

28. Lucía Boix-Palop

    Hospital Universitario Mutua Terrassa, Barcelona, Spain

    Contribution

    Data curation, Writing – review and editing

    Competing interests

    No competing interests declared

29. María Brion

    1. Instituto de Investigación Sanitaria de Santiago (IDIS), Xenética Cardiovascular, Santiago de Compostela, Spain
    2. CIBERCV, ISCIII, Madrid, Spain

    Contribution

    Data curation, Writing – review and editing

    Competing interests

    No competing interests declared

30. Ramón Brugada

    1. CIBERCV, ISCIII, Madrid, Spain
    2. Cardiovascular Genetics Center, Institut d’Investigació Biomèdica Girona (IDIBGI), Girona, Spain
    3. Medical Science Department, School of Medicine, University of Girona, Girona, Spain
    4. Hospital Josep Trueta, Cardiology Service, Girona, Spain

    Contribution

    Data curation, Writing – review and editing

    Competing interests

    No competing interests declared

31. Matilde Bustos

    Institute of Biomedicine of Seville (IBiS), Consejo Superior de Investigaciones Científicas (CSIC)- University of Seville- Virgen del Rocio University Hospital, Seville, Spain

    Contribution

    Data curation, Writing – review and editing

    Competing interests

    No competing interests declared

32. Enrique J Calderón

    1. Institute of Biomedicine of Seville (IBiS), Consejo Superior de Investigaciones Científicas (CSIC)- University of Seville- Virgen del Rocio University Hospital, Seville, Spain
    2. Departamento de Medicina, Hospital Universitario Virgen del Rocío, Universidad de Sevilla, Seville, Spain
    3. CIBERESP, ISCIII, Madrid, Spain

    Contribution

    Data curation, Writing – review and editing

    Competing interests

    No competing interests declared

33. Cristina Carbonell

    1. Hospital Universitario de Salamanca-IBSAL, Servicio de Medicina Interna, Salamanca, Spain
    2. Universidad de Salamanca, Salamanca, Spain

    Contribution

    Data curation, Writing – review and editing

    Competing interests

    No competing interests declared

34. Luis Castano

    1. CIBERER, ISCIII, Madrid, Spain
    2. Biocruces Bizkai HRI, Bizkaia, Spain
    3. Osakidetza, Cruces University Hospital, Bizkaia, Spain
    4. Centre for Biomedical Network Research on Diabetes and Metabolic Associated Diseases (CIBERDEM), Instituto de Salud Carlos III, Madrid, Spain
    5. University of Pais Vasco, UPV/EHU, Bizkaia, Spain

    Contribution

    Data curation, Writing – review and editing

    Competing interests

    No competing interests declared

35. Jose E Castelao

    Oncology and Genetics Unit, Instituto de Investigacion Sanitaria Galicia Sur, Xerencia de Xestion Integrada de Vigo-Servizo Galego de Saúde, Vigo, Spain

    Contribution

    Data curation, Writing – review and editing

    Competing interests

    No competing interests declared

36. Rosa Conde-Vicente

    Hospital Universitario Río Hortega, Valladolid, Spain

    Contribution

    Data curation, Writing – review and editing

    Competing interests

    No competing interests declared

37. M Lourdes Cordero-Lorenzana

    Servicio de Medicina intensiva, Complejo Hospitalario Universitario de A Coruña (CHUAC), Sistema Galego de Saúde (SERGAS), A Coruña, Spain

    Contribution

    Data curation, Writing – review and editing

    Competing interests

    No competing interests declared

38. Jose L Cortes-Sanchez

    1. Tecnológico de Monterrey, Monterrey, Mexico
    2. Department of Microgravity and Translational Regenerative Medicine, Otto von Guericke University, Magdeburg, Germany

    Contribution

    Data curation, Writing – review and editing

    Competing interests

    No competing interests declared

39. Marta Corton

    1. CIBERER, ISCIII, Madrid, Spain
    2. Department of Genetics & Genomics, Instituto de Investigación Sanitaria-Fundación Jiménez Díaz University Hospital - Universidad Autónoma de Madrid (IIS-FJD, UAM), Madrid, Spain

    Contribution

    Data curation, Writing – review and editing

    Competing interests

    No competing interests declared

40. M Teresa Darnaude

    Hospital Universitario Mostoles, Unidad de Genética, Madrid, Spain

    Contribution

    Data curation, Writing – review and editing

    Competing interests

    No competing interests declared

41. Alba De Martino-Rodríguez

    1. Instituto Aragonés de Ciencias de la Salud (IACS), Zaragoza, Spain
    2. Instituto Investigación Sanitaria Aragón (IIS-Aragon), Zaragoza, Spain

    Contribution

    Data curation

    Competing interests

    No competing interests declared

    "This ORCID iD identifies the author of this article:" 0000-0002-2436-9852

42. Victor del Campo-Pérez

    Preventive Medicine Department, Instituto de Investigacion Sanitaria Galicia Sur, Xerencia de Xestion Integrada de Vigo-Servizo Galego de Saúde, Vigo, Spain

    Contribution

    Data curation, Writing – review and editing

    Competing interests

    No competing interests declared

43. Aranzazu Diaz de Bustamante

    Hospital Universitario Mostoles, Unidad de Genética, Madrid, Spain

    Contribution

    Data curation, Writing – review and editing

    Competing interests

    No competing interests declared

44. Elena Domínguez-Garrido

    Unidad Diagnóstico Molecular, Fundación Rioja Salud, La Rioja, Spain

    Contribution

    Data curation, Writing – review and editing

    Competing interests

    No competing interests declared

45. Rocío Eirós

    Hospital Universitario de Salamanca-IBSAL, Servicio de Cardiología, Salamanca, Spain

    Contribution

    Data curation, Writing – review and editing

    Competing interests

    No competing interests declared

46. María Carmen Fariñas

    1. IDIVAL, Cantabria, Spain
    2. Hospital U M Valdecilla, Cantabria, Spain
    3. Universidad de Cantabria, Cantabria, Spain

    Contribution

    Data curation, Writing – review and editing

    Competing interests

    No competing interests declared

47. María J Fernandez-Nestosa

    Universidad Nacional de Asunción, Facultad de Politécnica, Paraguay, United States

    Contribution

    Data curation, Writing – review and editing

    Competing interests

    No competing interests declared

    "This ORCID iD identifies the author of this article:" 0000-0001-7764-7791

48. Uxía Fernández-Robelo

    Urgencias Hospitalarias, Complejo Hospitalario Universitario de A Coruña (CHUAC), Sistema Galego de Saúde (SERGAS), A Coruña, Spain

    Contribution

    Data curation, Writing – review and editing

    Competing interests

    No competing interests declared

49. Amanda Fernández-Rodríguez

    1. CIBERINFEC, ISCIII, Madrid, Spain
    2. Unidad de Infección Viral e Inmunidad, Centro Nacional de Microbiología (CNM), Instituto de Salud Carlos III (ISCIII), Madrid, Spain

    Contribution

    Data curation, Writing – review and editing

    Competing interests

    No competing interests declared

50. Tania Fernández-Villa

    1. CIBERESP, ISCIII, Madrid, Spain
    2. Grupo de Investigación en Interacciones Gen-Ambiente y Salud (GIIGAS) - Instituto de Biomedicina (IBIOMED), Universidad de León, León, Spain

    Contribution

    Data curation, Writing – review and editing

    Competing interests

    No competing interests declared

51. Manuela Gago-Dominguez

    1. Fundación Pública Galega de Medicina Xenómica, Sistema Galego de Saúde (SERGAS), Santiago de Compostela, Spain
    2. IDIS, Seongnam, Republic of Korea

    Contribution

    Data curation, Writing – review and editing

    Competing interests

    No competing interests declared

52. Belén Gil-Fournier

    Hospital Universitario de Getafe, Servicio de Genética, Madrid, Spain

    Contribution

    Data curation, Writing – review and editing

    Competing interests

    No competing interests declared

53. Javier Gómez-Arrue

    1. Instituto Aragonés de Ciencias de la Salud (IACS), Zaragoza, Spain
    2. Instituto Investigación Sanitaria Aragón (IIS-Aragon), Zaragoza, Spain

    Contribution

    Data curation, Writing – review and editing

    Competing interests

    No competing interests declared

54. Beatriz González Álvarez

    1. Instituto Aragonés de Ciencias de la Salud (IACS), Zaragoza, Spain
    2. Instituto Investigación Sanitaria Aragón (IIS-Aragon), Zaragoza, Spain

    Contribution

    Data curation, Writing – review and editing

    Competing interests

    No competing interests declared

55. Fernan Gonzalez Bernaldo de Quirós

    Ministerio de Salud Ciudad de Buenos Aires, Buenos Aires, Argentina

    Contribution

    Data curation, Writing – review and editing

    Competing interests

    No competing interests declared

56. Anna González-Neira

    Spanish National Cancer Research Centre, Human Genotyping-CEGEN Unit, Madrid, Spain

    Contribution

    Data curation, Writing – review and editing

    Competing interests

    No competing interests declared

57. Javier González-Peñas

    1. Department of Child and Adolescent Psychiatry, Institute of Psychiatry and Mental Health, Hospital General Universitario Gregorio Marañón (IiSGM), Madrid, Spain
    2. School of Medicine, Universidad Complutense, Madrid, Spain
    3. Centre for Biomedical Network Research on Mental Health (CIBERSAM), Instituto de Salud Carlos III, Madrid, Spain

    Contribution

    Data curation, Writing – review and editing

    Competing interests

    No competing interests declared

58. Juan F Gutiérrez-Bautista

    Hospital Universitario Virgen de las Nieves, Servicio de Análisis Clínicos e Inmunología, Granada, Spain

    Contribution

    Data curation, Writing – review and editing

    Competing interests

    No competing interests declared

59. María José Herrero

    1. IIS La Fe, Plataforma de Farmacogenética, Valencia, Spain
    2. Universidad de Valencia, Departamento de Farmacología, Valencia, Spain

    Contribution

    Data curation, Writing – review and editing

    Competing interests

    No competing interests declared

60. Antonio Herrero-Gonzalez

    Data Analysis Department, Instituto de Investigación Sanitaria-Fundación Jiménez Díaz University Hospital - Universidad Autónoma de Madrid (IIS-FJD, UAM), Madrid, Spain

    Contribution

    Data curation, Writing – review and editing

    Competing interests

    No competing interests declared

61. María A Jimenez-Sousa

    1. CIBERINFEC, ISCIII, Madrid, Spain
    2. Unidad de Infección Viral e Inmunidad, Centro Nacional de Microbiología (CNM), Instituto de Salud Carlos III (ISCIII), Madrid, Spain

    Contribution

    Data curation, Writing – review and editing

    Competing interests

    No competing interests declared

62. María Claudia Lattig

    1. Universidad de los Andes, Facultad de Ciencias, Bogotá, Colombia
    2. SIGEN Alianza Universidad de los Andes - Fundación Santa Fe de Bogotá, Bogotá, Colombia

    Contribution

    Data curation, Writing – review and editing

    Competing interests

    No competing interests declared

    "This ORCID iD identifies the author of this article:" 0000-0003-2113-9266

63. Anabel Liger Borja

    Hospital General de Segovia, Medicina Intensiva, Segovia, Spain

    Contribution

    Data curation, Writing – review and editing

    Competing interests

    No competing interests declared

64. Rosario Lopez-Rodriguez

    1. CIBERER, ISCIII, Madrid, Spain
    2. Department of Genetics & Genomics, Instituto de Investigación Sanitaria-Fundación Jiménez Díaz University Hospital - Universidad Autónoma de Madrid (IIS-FJD, UAM), Madrid, Spain
    3. Facultad de Farmacia, Universidad San Pablo-CEU, CEU Universities, Urbanización Montepríncipe, Boadilla del Monte, Spain

    Contribution

    Data curation, Writing – review and editing

    Competing interests

    No competing interests declared

65. Esther Mancebo

    1. Hospital Universitario 12 de Octubre, Department of Immunology, Madrid, Spain
    2. Instituto de Investigación Sanitaria Hospital 12 de Octubre (imas12), Transplant Immunology and Immunodeficiencies Group, Madrid, Spain

    Contribution

    Data curation, Writing – review and editing

    Competing interests

    No competing interests declared

66. Caridad Martín-López

    Hospital General de Segovia, Medicina Intensiva, Segovia, Spain

    Contribution

    Data curation, Writing – review and editing

    Competing interests

    No competing interests declared

67. Vicente Martín

    1. CIBERESP, ISCIII, Madrid, Spain
    2. Grupo de Investigación en Interacciones Gen-Ambiente y Salud (GIIGAS) - Instituto de Biomedicina (IBIOMED), Universidad de León, León, Spain

    Contribution

    Data curation, Writing – review and editing

    Competing interests

    No competing interests declared

68. Oscar Martinez-Nieto

    1. SIGEN Alianza Universidad de los Andes - Fundación Santa Fe de Bogotá, Bogotá, Colombia
    2. Fundación Santa Fe de Bogota, Departamento Patologia y Laboratorios, Bogotá, Colombia

    Contribution

    Data curation, Writing – review and editing

    Competing interests

    No competing interests declared

69. Iciar Martinez-Lopez

    1. Unidad de Genética y Genómica Islas Baleares, Islas Baleares, Spain
    2. Hospital Universitario Son Espases, Unidad de Diagnóstico Molecular y Genética Clínica, Islas Baleares, Spain

    Contribution

    Data curation, Writing – review and editing

    Competing interests

    No competing interests declared

70. Michel F Martinez-Resendez

    Tecnológico de Monterrey, Monterrey, Mexico

    Contribution

    Data curation, Writing – review and editing

    Competing interests

    No competing interests declared

71. Angel Martinez-Perez

    Genomics of Complex Diseases Unit, Research Institute of Hospital de la Santa Creu i Sant Pau, IIB Sant Pau, Barcelona, Spain

    Contribution

    Data curation, Writing – review and editing

    Competing interests

    No competing interests declared

    "This ORCID iD identifies the author of this article:" 0000-0002-5934-1454

72. Juliana F Mazzeu

    1. Universidade de Brasília, Faculdade de Medicina, Brasília, Brazil
    2. Programa de Pós-Graduação em Ciências Médicas (UnB), Brasília, Brazil
    3. Programa de Pós-Graduação em Ciencias da Saude (UnB), Brazila, Brazil

    Contribution

    Data curation, Writing – review and editing

    Competing interests

    No competing interests declared

73. Eleuterio Merayo Macías

    Hospital El Bierzo, Unidad Cuidados Intensivos, León, Spain

    Contribution

    Data curation, Writing – review and editing

    Competing interests

    No competing interests declared

74. Pablo Minguez

    1. CIBERER, ISCIII, Madrid, Spain
    2. Department of Genetics & Genomics, Instituto de Investigación Sanitaria-Fundación Jiménez Díaz University Hospital - Universidad Autónoma de Madrid (IIS-FJD, UAM), Madrid, Spain

    Contribution

    Data curation, Writing – review and editing

    Competing interests

    No competing interests declared

75. Victor Moreno Cuerda

    1. Hospital Universitario Mostoles, Medicina Interna, Madrid, Spai, Spain
    2. Universidad Francisco de Vitoria, Madrid, Spain

    Contribution

    Data curation, Writing – review and editing

    Competing interests

    No competing interests declared

76. Silviene F Oliveira

    1. Programa de Pós-Graduação em Ciencias da Saude (UnB), Brazila, Brazil
    2. Departamento de Genética e Morfologia, Instituto de Ciências Biológicas, Universidade de Brasília, Brasília, Brazil
    3. Programa de Pós-Graduação em Biologia Animal (UnB), Brasília, Brazil
    4. Programa de Pós-Graduação Profissional em Ensino de Biologia (UnB), Brasília, Brazil

    Contribution

    Data curation, Writing – review and editing

    Competing interests

    No competing interests declared

    "This ORCID iD identifies the author of this article:" 0000-0002-7741-0257

77. Eva Ortega-Paino

    Spanish National Cancer Research Centre, CNIO Biobank, Madrid, Spain

    Contribution

    Data curation, Writing – review and editing

    Competing interests

    No competing interests declared

78. Mara Parellada

    1. Department of Child and Adolescent Psychiatry, Institute of Psychiatry and Mental Health, Hospital General Universitario Gregorio Marañón (IiSGM), Madrid, Spain
    2. School of Medicine, Universidad Complutense, Madrid, Spain
    3. Centre for Biomedical Network Research on Mental Health (CIBERSAM), Instituto de Salud Carlos III, Madrid, Spain

    Contribution

    Data curation, Writing – review and editing

    Competing interests

    No competing interests declared

79. Estela Paz-Artal

    1. Hospital Universitario 12 de Octubre, Department of Immunology, Madrid, Spain
    2. Instituto de Investigación Sanitaria Hospital 12 de Octubre (imas12), Transplant Immunology and Immunodeficiencies Group, Madrid, Spain
    3. Universidad Complutense de Madrid, Department of Immunology, Ophthalmology and ENT, Madrid, Spain

    Contribution

    Data curation, Writing – review and editing

    Competing interests

    No competing interests declared

80. Ney PC Santos

    Universidade Federal do Pará, Núcleo de Pesquisas em Oncologia, Belém, Brazil

    Contribution

    Data curation, Writing – review and editing

    Competing interests

    No competing interests declared

81. Patricia Pérez-Matute

    Infectious Diseases, Microbiota and Metabolism Unit, CSIC Associated Unit, Center for Biomedical Research of La Rioja (CIBIR), Logroño, Spain

    Contribution

    Data curation

    Competing interests

    No competing interests declared

82. Patricia Perez

    Inditex, A Coruña, Corunna, Spain

    Contribution

    Data curation, Writing – review and editing

    Competing interests

    No competing interests declared

83. M Elena Pérez-Tomás

    Instituto Murciano de Investigación Biosanitaria (IMIB-Arrixaca), Murcia, Spain

    Contribution

    Data curation, Writing – review and editing

    Competing interests

    No competing interests declared

    "This ORCID iD identifies the author of this article:" 0000-0002-5429-3414

84. Teresa Perucho

    GENYCA, Madrid, Spain

    Contribution

    Data curation, Writing – review and editing

    Competing interests

    No competing interests declared

85. Mellina Pinsach-Abuin

    1. CIBERCV, ISCIII, Madrid, Spain
    2. Cardiovascular Genetics Center, Institut d’Investigació Biomèdica Girona (IDIBGI), Girona, Spain

    Contribution

    Data curation, Writing – review and editing

    Competing interests

    No competing interests declared

86. Guillermo Pita

    Spanish National Cancer Research Centre, Human Genotyping-CEGEN Unit, Madrid, Spain

    Contribution

    Data curation, Writing – review and editing

    Competing interests

    No competing interests declared

87. Ericka N Pompa-Mera

    1. Instituto Mexicano del Seguro Social (IMSS), Centro Médico Nacional Siglo XXI, Unidad de Investigación Médica en Enfermedades Infecciosas y Parasitarias, Mexico City, Mexico
    2. Instituto Mexicano del Seguro Social (IMSS), Centro Médico Nacional La Raza, Hospital de Infectología, Mexico City, Mexico

    Contribution

    Data curation, Writing – review and editing

    Competing interests

    No competing interests declared

88. Gloria L Porras-Hurtado

    Clinica Comfamiliar Risaralda, Pereira, Colombia

    Contribution

    Data curation, Writing – review and editing

    Competing interests

    No competing interests declared

89. Aurora Pujol

    1. CIBERER, ISCIII, Madrid, Spain
    2. Bellvitge Biomedical Research Institute (IDIBELL), Neurometabolic Diseases Laboratory, L’Hospitalet de Llobregat, Barcelona, Spain
    3. Catalan Institution of Research and Advanced Studies (ICREA), Barcelona, Spain

    Contribution

    Data curation, Writing – review and editing

    Competing interests

    No competing interests declared

    "This ORCID iD identifies the author of this article:" 0000-0002-9606-0600

90. Soraya Ramiro León

    Hospital Universitario de Getafe, Servicio de Genética, Madrid, Spain

    Contribution

    Data curation, Writing – review and editing

    Competing interests

    No competing interests declared

91. Salvador Resino

    1. CIBERINFEC, ISCIII, Madrid, Spain
    2. Unidad de Infección Viral e Inmunidad, Centro Nacional de Microbiología (CNM), Instituto de Salud Carlos III (ISCIII), Madrid, Spain

    Contribution

    Data curation, Writing – review and editing

    Competing interests

    No competing interests declared

92. Marianne R Fernandes

    1. Universidade Federal do Pará, Núcleo de Pesquisas em Oncologia, Belém, Brazil
    2. Hospital Ophir Loyola, Departamento de Ensino e Pesquisa, Belém, Brazil

    Contribution

    Data curation, Writing – review and editing

    Competing interests

    No competing interests declared

93. Emilio Rodríguez-Ruiz

    1. IDIS, Seongnam, Republic of Korea
    2. Unidad de Cuidados Intensivos, Hospital Clínico Universitario de Santiago (CHUS), Sistema Galego de Saúde (SERGAS), Santiago de Compostela, Spain

    Contribution

    Data curation, Writing – review and editing

    Competing interests

    No competing interests declared

94. Fernando Rodriguez-Artalejo

    1. CIBERESP, ISCIII, Madrid, Spain
    2. Department of Preventive Medicine and Public Health, School of Medicine, Universidad Autónoma de Madrid, Madrid, Spain
    3. IdiPaz (Instituto de Investigación Sanitaria Hospital Universitario La Paz), Madrid, Spain
    4. IMDEA-Food Institute, CEI UAM+CSIC, Madrid, Spain

    Contribution

    Data curation, Writing – review and editing

    Competing interests

    No competing interests declared

95. José A Rodriguez-Garcia

    Complejo Asistencial Universitario de León, León, Spain

    Contribution

    Data curation, Writing – review and editing

    Competing interests

    No competing interests declared

96. Francisco Ruiz-Cabello

    1. IDIS, Seongnam, Republic of Korea
    2. Instituto de Investigación Biosanitaria de Granada (ibs GRANADA), Granada, Spain
    3. Universidad de Granada, Departamento Bioquímica, Biología Molecular e Inmunología III, Granada, Spain

    Contribution

    Data curation, Writing – review and editing

    Competing interests

    No competing interests declared

97. Javier Ruiz-Hornillos

    1. Hospital Infanta Elena, Allergy Unit, Valdemoro, Madrid, Spain
    2. Instituto de Investigación Sanitaria-Fundación Jiménez Díaz University Hospital - Universidad Autónoma de Madrid (IIS-FJD, UAM), Madrid, Spain
    3. Faculty of Medicine, Universidad Francisco de Vitoria, Madrid, Spain

    Contribution

    Data curation, Writing – review and editing

    Competing interests

    No competing interests declared

98. Pablo Ryan

    1. CIBERINFEC, ISCIII, Madrid, Spain
    2. Hospital Universitario Infanta Leonor, Madrid, Spain
    3. Complutense University of Madrid, Madrid, Spain
    4. Gregorio Marañón Health Research Institute (IiSGM), Madrid, Spain

    Contribution

    Data curation, Writing – review and editing

    Competing interests

    No competing interests declared

    "This ORCID iD identifies the author of this article:" 0000-0002-4212-7419

99. José Manuel Soria

    Genomics of Complex Diseases Unit, Research Institute of Hospital de la Santa Creu i Sant Pau, IIB Sant Pau, Barcelona, Spain

    Contribution

    Data curation, Writing – review and editing

    Competing interests

    No competing interests declared

100. Juan Carlos Souto

     Haemostasis and Thrombosis Unit, Hospital de la Santa Creu i Sant Pau, IIB Sant Pau, Barcelona, Spain

     Contribution

     Data curation, Writing – review and editing

     Competing interests

     No competing interests declared

101. Eduardo Tamayo

     1. Hospital Clinico Universitario de Valladolid, Servicio de Anestesiologia y Reanimación, Valladolid, Spain
     2. Universidad de Valladolid, Departamento de Cirugía, Valladolid, Spain

     Contribution

     Data curation, Writing – review and editing

     Competing interests

     No competing interests declared

102. Alvaro Tamayo-Velasco

     Hospital Clinico Universitario de Valladolid, Servicio de Hematologia y Hemoterapia, Valladolid, Spain

     Contribution

     Data curation, Writing – review and editing

     Competing interests

     No competing interests declared

103. Juan Carlos Taracido-Fernandez

     Data Analysis Department, Instituto de Investigación Sanitaria-Fundación Jiménez Díaz University Hospital - Universidad Autónoma de Madrid (IIS-FJD, UAM), Madrid, Spain

     Contribution

     Data curation, Writing – review and editing

     Competing interests

     No competing interests declared

104. Alejandro Teper

     Hospital de Niños Ricardo Gutierrez, Buenos Aires, Argentina

     Contribution

     Data curation, Writing – review and editing

     Competing interests

     No competing interests declared

105. Lilian Torres-Tobar

     Fundación Universitaria de Ciencias de la Salud, Bogotá, Colombia

     Contribution

     Data curation, Writing – review and editing

     Competing interests

     No competing interests declared

106. Miguel Urioste

     Spanish National Cancer Research Centre, Familial Cancer Clinical Unit, Madrid, Spain

     Contribution

     Data curation, Writing – review and editing

     Competing interests

     No competing interests declared

107. Juan Valencia-Ramos

     University Hospital of Burgos, Burgos, Spain

     Contribution

     Data curation, Writing – review and editing

     Competing interests

     No competing interests declared

108. Zuleima Yáñez

     Universidad Simón Bolívar, Facultad de Ciencias de la Salud, Barranquilla, Colombia

     Contribution

     Data curation, Writing – review and editing

     Competing interests

     No competing interests declared

109. Ruth Zarate

     Centro para el Desarrollo de la Investigación Científica, Asunción, Paraguay

     Contribution

     Data curation, Writing – review and editing

     Competing interests

     No competing interests declared

110. Itziar de Rojas

     1. Centre for Biomedical Network Research on Neurodegenerative Diseases (CIBERNED), Instituto de Salud Carlos III, Madrid, Spain
     2. Research Center and Memory clinic, ACE Alzheimer Center Barcelona, Universitat Internacional de Catalunya, Barcelona, Spain

     Contribution

     Data curation, Writing – review and editing

     Competing interests

     No competing interests declared

111. Agustín Ruiz

     1. Centre for Biomedical Network Research on Neurodegenerative Diseases (CIBERNED), Instituto de Salud Carlos III, Madrid, Spain
     2. Research Center and Memory clinic, ACE Alzheimer Center Barcelona, Universitat Internacional de Catalunya, Barcelona, Spain

     Contribution

     Data curation, Writing – review and editing

     Competing interests

     No competing interests declared

112. Pascual Sánchez

     CIEN Foundation/Queen Sofia Foundation Alzheimer Center, Madrid, Spain

     Contribution

     Data curation, Writing – review and editing

     Competing interests

     No competing interests declared

113. Luis Miguel Real

     Hospital Universitario de Valme, Unidad Clínica de Enfermedades Infecciosas y Microbiología, Sevilla, Spain

     Contribution

     Data curation, Writing – review and editing

     Competing interests

     No competing interests declared

     "This ORCID iD identifies the author of this article:" 0000-0003-4932-7429

114. SCOURGE Cohort Group

     Contribution

     Data curation, Methodology, Resources

     Competing interests

     No competing interests declared

115. Encarna Guillen-Navarro

     1. Instituto Murciano de Investigación Biosanitaria (IMIB-Arrixaca), Murcia, Spain
     2. Sección Genética Médica - Servicio de Pediatría, Hospital Clínico Universitario Virgen de la Arrixaca, Servicio Murciano de Salud, Murcia, Spain
     3. Departamento Cirugía, Pediatría, Obstetricia y Ginecología, Facultad de Medicina, Universidad de Murcia (UMU), Murcia, Spain
     4. Grupo Clínico Vinculado, Centre for Biomedical Network Research on Rare Diseases (CIBERER), Instituto de Salud Carlos III, Madrid, Spain

     Contribution

     Data curation, Writing – review and editing

     Competing interests

     No competing interests declared

116. Carmen Ayuso

     1. CIBERER, ISCIII, Madrid, Spain
     2. Department of Genetics & Genomics, Instituto de Investigación Sanitaria-Fundación Jiménez Díaz University Hospital - Universidad Autónoma de Madrid (IIS-FJD, UAM), Madrid, Spain

     Contribution

     Data curation, Writing – review and editing

     Competing interests

     No competing interests declared

117. Esteban Parra

     Department of Anthropology, University of Toronto at Mississauga, Mississauga, Canada

     Contribution

     Investigation, Methodology, Writing – review and editing

     Competing interests

     No competing interests declared

     "This ORCID iD identifies the author of this article:" 0000-0002-2057-8577

118. José A Riancho

     1. CIBERER, ISCIII, Madrid, Spain
     2. IDIVAL, Cantabria, Spain
     3. Hospital U M Valdecilla, Cantabria, Spain
     4. Universidad de Cantabria, Cantabria, Spain

     Contribution

     Conceptualization, Supervision, Funding acquisition, Investigation, Methodology, Project administration, Writing – review and editing

     Competing interests

     No competing interests declared

     "This ORCID iD identifies the author of this article:" 0000-0003-0691-8755

119. Augusto Rojas-Martinez

     Tecnologico de Monterrey, Escuela de Medicina y Ciencias de la Salud, Monterrey, Mexico

     Contribution

     Conceptualization, Supervision, Funding acquisition, Investigation, Project administration, Writing – review and editing

     Competing interests

     No competing interests declared

     "This ORCID iD identifies the author of this article:" 0000-0003-3765-6778

120. Carlos Flores

     1. Genomics Division, Instituto Tecnológico y de Energías Renovables, Santa Cruz de Tenerife, Spain
     2. Research Unit, Hospital Universitario Nuestra Señora de Candelaria, Instituto de Investigación Sanitaria de Canarias, Santa Cruz de Tenerife, Spain
     3. Department of Clinical Sciences, University Fernando Pessoa Canarias, Las Palmas de Gran Canaria, Spain
     4. Centre for Biomedical Network Research on Respiratory Diseases (CIBERES), Instituto de Salud Carlos III, Madrid, Spain

     Contribution

     Conceptualization, Supervision, Funding acquisition, Investigation, Methodology, Writing – original draft, Project administration, Writing – review and editing

     For correspondence

     cflores@ull.edu.es

     Competing interests

     No competing interests declared

     "This ORCID iD identifies the author of this article:" 0000-0001-5352-069X

121. Pablo Lapunzina

     1. ERN-ITHACA-European Reference Network, Soria, Spain
     2. CIBERER, ISCIII, Madrid, Spain
     3. Instituto de Genética Médica y Molecular (INGEMM), Hospital Universitario La Paz IDIPAZ, Madrid, Spain

     Contribution

     Conceptualization, Supervision, Funding acquisition, Investigation, Project administration, Writing – review and editing

     Competing interests

     No competing interests declared

122. Ángel Carracedo

     1. CIBERER, ISCIII, Madrid, Spain
     2. Centro Singular de Investigación en Medicina Molecular y Enfermedades Crónicas (CIMUS), Universidade de Santiago de Compostela, Santiago de Compostela, Spain
     3. Fundación Pública Galega de Medicina Xenómica, Sistema Galego de Saúde (SERGAS), Santiago de Compostela, Spain
     4. IDIS, Seongnam, Republic of Korea

     Contribution

     Conceptualization, Supervision, Funding acquisition, Investigation, Methodology, Writing – original draft, Project administration, Writing – review and editing

     For correspondence

     angel.carracedo@usc.es

     Competing interests

     No competing interests declared

     "This ORCID iD identifies the author of this article:" 0000-0003-1085-8986