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Metabolismo De Isoflavonas Y Formación De Equol Por Bacterias Del Tracto Gastrointestinal Humano

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Metabolismo De Isoflavonas Y Formación De Equol Por Bacterias Del Tracto Gastrointestinal Humano Programa de Doctorado en Ingeniería Química, Ambiental y Bioalimentaria Metabolismo de isoflavonas y formación de equol por bacterias del tracto gastrointestinal humano. Lucía Vázquez Iglesias Tesis Doctoral Oviedo, 2020 Programa de Doctorado en Ingeniería Química, Ambiental y Bioalimentaria Metabolismo de isoflavonas y formación de equol por bacterias del tracto gastrointestinal humano Lucía Vázquez Iglesias Tesis Doctoral Oviedo, 2020 Este trabajo ha sido realizado en el Instituto de Productos Lácteos de Asturias (IPLA-CSIC) AUTORIZACIÓN PARA LA PRESENTACIÓN DE TESIS DOCTORAL Año Académico: 2019/2020 1.- Datos personales del autor de la Tesis Apellidos: Nombre: Vázquez Iglesias Lucía DNI/Pasaporte/NIE: Teléfono: Correo electrónico: 71668563B 609562520 [email protected] 2.- Datos académicos Programa de Doctorado cursado: Programa en Ingeniería Química, Ambiental y Bioalimentaria Órgano responsable: ) Universidad de Oviedo 8 1 Departamento/Instituto en el que presenta la Tesis Doctoral: 0 2 Departamento de Ingeniería Química y Tecnología del Medio Ambiente . g e Título definitivo de la Tesis R ( Español/Otro Idioma: Inglés: 9 Metabolismo de isoflavonas y formación de 0 0 equol por bacterias del tracto production by bacteria from the human - A gastrointestinal humano. O Rama de conocimiento: V - Ingeniería y Arquitectura T A M 3.- Autorización del Director/es y Tutor de la tesis - R D/Dª: Baltasar Mayo Pérez DNI/Pasaporte/NIE: 10820266-P O Departamento/Instituto: F Departamento de Microbiología y Bioquímica de Productos Lácteos Instituto de Productos Lácteos de Asturias ( IPLA-CSIC) D/Dª: Ana Belén Flórez García DNI/Pasaporte/NIE: 09432953-D Departamento/Instituto/Institución: Departamento de Microbiología y Bioquímica de Productos Lácteos Instituto de Productos Lácteos de Asturias ( IPLA-CSIC) Autorización del Tutor de la tesis D/Dª: Clara González de los Reyes-Gavilán DNI/Pasaporte/NIE: 10594470-A Departamento/Instituto: Departamento de Microbiología y Bioquímica de Productos Lácteos Instituto de Productos Lácteos de Asturias ( IPLA-CSIC) Autoriza la presentación de la tesis doctoral en cumplimiento de lo establecido en el Art. 32 del Reglamento de los Estudios de Doctorado, aprobado por el Consejo de Gobierno, en su sesión del día 20 de julio de 2018 (BOPA del 9 de agosto de 2018) Villaviciosa, 27 Agosto del 2020 Director/es de la Tesis Tutor de la Tesis Fdo.: Baltasar Mayo/ Ana Belén Flórez Fdo.: Clara G. de los Reyes-Gavilán SR. PRESIDENTE DE LA COMISIÓN ACADÉMICA DEL PROGRAMA DE DOCTORADO EN ________________________________________________________________________ ÍNDICE INDEX ÍNDICE/INDEX Lista de abreviaturas i Lista de figuras y tablas iii Resumen/ Summary iv INTRODUCCIÓN /INTRODUCTION 1. ISOFLAVONAS Y EQUOL 1.1 Isoflavonas 1.1.1 Polifenoles y fitoestrógenos 1 1.1.2 Isoflavonas 2 1.1.3 Isoflavonas y soja 3 1.1.4 Metabolismo de isoflavonas 4 1.2 Equol 1.2.1 Características generales 5 1.2.2 Metabolismo de daidzeína y formación de equol 5 1.3 Isoflavonas, equol y salud 1.3.1 Mecanismos de acción 7 1.3.2 Efectos beneficiosos del equol en la salud 8 1.3.2.1 Menopausia 10 1.3.2.2 Sistema cardiovascular 10 1.3.2.3 Salud ósea 11 1.3.2.4 Cánceres hormonodependientes 12 1.3.2.5 Sistema nervioso central 12 1.3.2.6 Otros efectos beneficiosos 13 2. MICROBIOTA INTESTINAL HUMANA 2.1. Composición y enterotipos 13 2.2. Funciones de la microbiota 15 2.2.1. Función metabólica 16 2.2.2. Función inmunológica 17 2.2.3. Función protectora 17 2.3. Factores que influyen en la microbiota 18 2.3.1. Microbiota y dieta 18 2.3.2. Alimentación funcional, probióticos y prebióticos 19 2.4. Métodos de estudio de la microbiota intestinal 20 2.4.1. Modelos intestinales 21 3. MICROBIOTA Y EQUOL 3.1. Fenotipo productor de equol 23 3.1.1. Influencia de la dieta en el fenotipo productor de equol 24 3.2. Microorganismos productores de equol 25 3.3. Caracterización molecular de la formación de equol 28 3.4. Producción biotecnológica de equol 29 3.5. Equol y alimentos funcionales 30 OBJETIVOS/OBJECTIVES 31 TRABAJO EXPERIMENTAL /EXPERIMENTAL WORK CAPÍTULO 1 37 Desarrollo de métodos para identificar y cuantificar poblaciones intestinales involucradas en el metabolismo de las isoflavonas y la producción de equol. CAPÍTULO 2 70 Estudio de las relaciones e interacciones entre isoflavonas, equol y poblaciones bacterianas intestinales. CAPÍTULO 3 100 Caracterización de la producción de equol en muestras fecales y bacterias productoras con el fin de maximizar su síntesis endógena y biotecnológica. DISCUSIÓN/ DISCUSSION 148 CONCLUSIONES/ CONCLUSIONS 162 BIBLIOGRAFÍA/BIBLIOGRAPHY 166 ANEXOS/ANNEXES . Anexo I.- Revisión bibliográfica sobre el equol y sus efectos en la salud humana 186 . Anexo II.- Informe sobre la calidad de los artículos 206 LISTA DE ABREVIATURAS ADN: Ácido desoxirribonucleico FI: Factor de impacto AF: Alimentos funcionales FISH: “Fluorescence in situ hybridization” (Hibridación fluorescente in situ) AGCC: Ácidos grasos de cadena corta FOS: Fructooligosacáridos AOS: Arabinooligosacáridos g: Gramo ARN: Ácido ribonucleico GABA: “Gamma-aminobutyric acid” (Ácido ARNr: Ácido ribonucleico ribosomal gamma-aminobutírico) ATTC: “American Type Culture Collection” GOS: Galactooligosacáridos (Colección Americana de Cultivos Tipo) IgA: Inmunoglobulina A BAL: Bacterias ácido-lácticas ISAPP: “International Scientific Association for CCR: Cáncer colorrectal Probiotics and Prebiotics” (Asociación CIM: Concentración inhibitoria mínima Científica Internacional de Probióticos y cLDL: Colesterol unido a lipoproteínas de baja Prebióticos) densidad Kpb: Kilo pares de bases Ct: “Cycle threshold ” (ciclo umbral en qPCR) L: Litro DGGE: “Denaturing gradient gel electro- mL: Mililitro phoresis” (Electroforesis en geles de gradiente mg: Miligramos desnaturalizantes) MRSA: “Methicillin-resistant Staphylococcus DHD: Dihidrodaidzeína aureus” (S.aureus resistente a la meticilina) DSMZ: “Deutsche Sammlung von Mikroor- NADP(H): Nicotinamida adenina dinucleótido ganismen und Zellkulturen” (Colección fosfato Alemana de Microorganismos y Cultivos Celulares) NGS: “Next-generation sequencing” (Tecnologías de secuenciación masiva) ECV: Enfermedades cardiovasculares nmol: Nanomol EEUU: Estados Unidos de América O-DMA: O-desmetilangolensina EFSA: “European Food Safety Authority” (Autoridad Europea de Seguridad OMS: Organización Mundial de la Salud Alimentaria) ORFs: “Open reading frame” (Pauta abierta de EII: Enfermedad inflamatoria intestinal lectura) EPS: Exopolisacáridos PCR: “Polymerase chain reaction” (Reacción en cadena de la polimerasa) FAD: Flavín adenín dinucleótido Q: Cuartil FAO: “Food and Agriculture Organization of the United Nations” (Organización de las qPCR: “Quantitative polymerase chain Naciones Unidas para la Alimentación y la reaction” (PCR cuantitativa o PCR en tiempo Agricultura) real) FDA: “Food and Drug Administration” R-DHD: R-dihidrodaidzeína (Agencia Americana de Medicamentos y REs: Receptores estrogénicos Alimentación) REα: Receptor estrogénico subtipo alfa Fg: Femtogramo i REβ: Receptor estrogénico subtipo beta Tm: “Melting temperature” (Temperatura de fusión) RT-PCR: “Reverse transcription polymerase chain reaction” (PCR con transcriptasa TNF α: “Tumor necrosis factor alpha” (Factor inversa) de necrosis tumoral alfa) RT-qPCR: “Quantitative reverse TNO: “Nederlandse Organisatie voor Toegepast transcription PCR” (PCR cuantitativa con Natuurwetenschappelijk Onderzoek” transcriptasa inversa) (Organización Holandesa para la Investigación Científica Aplicada) SCI: “ Science citation index” Ufc: Unidades formadoras de colonia S-DHD: S-dihidrodaizeína UHPLC: “Ultra-high-pressure liquid S-THD: S-tetrahidrodaizeína chromatography” (Cromatografía líquida de SHBG: “Sex hormone binding globulin” ultra-alta resolución) (Globulina fijadora de hormonas sexuales) UV: Ultravioleta TGI: Tracto gastrointestinal WHI: “Women’s Health Initiative” THD: Tetrahidrodaidzeína YIT: “Yeoju Institute of technology” (Instituto THS: Terapia hormonal sustitutiva de Tecnología de Yeoju) TIM-2: “The TNO in vitro model of the colon” μ: Tasa específica de crecimiento (Modelo in vitro de colon desarrollado por el TNO) μg: Microgramo ii LISTA DE FIGURAS Y TABLAS Figura/Tabla Título Página Figura 1 Clasificación de los polifenoles según su estructura química: ácidos 1 fenólicos, lignanos, estilbenos y flavonoides, dentro de los cuales se incluyen las isoflavonas. Adaptado de Choi y Shin (2016). Figura 2 Estructura química básica de las isoflavonas glicosiladas y agliconas. Se 3 indican los dos anillos fenólicos y el anillo cromano con las letras A, B y C, respectivamente, así como la numeración de los carbonos y la posición de los radicales cuyos sustituyentes se indican en la tabla adyacente. Figura 3 Metabolismo de la daidzeína y ruta de formación de equol. Se indican las 6 enzimas de la microbiota intestinal humana involucradas en el metabolismo. Adaptado de Mayo et al. (2019). Figura 4 Analogía estructural entre el 17 β-estradiol y el equol. 8 Figura 5 Efectos beneficiosos para la salud humana asociados al consumo de 9 equol. Figura 6 Esquema de la distribución de la composición microbiana a lo largo del 14 tracto gastrointestinal humano en el que se representan los géneros más abundantes, el pH de la sección y la concentración bacteriana aproximada por gramos de contenido fecal. Adaptado de Jandhyala et al. (2015). Figura 7 Esquema del modelo intestinal in vitro TIM-2. El sistema está integrado 22 por: (a) compartimentos con movimientos peristálticos que albergan la materia fecal
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