Invasión trofoblástica en el embarazo normal (I): Placentación inicial

Eduardo Reyna Villasmil; Carlos Briceño Pérez; Joel Santos Bolívar

Invasión trofoblástica en el embarazo normal (I): Placentación inicial(Trophoblastic invasion in normal pregnancy (I): Initial placentation)Eduardo Reyna-Villasmil1, Carlos Briceño Pérez2, Joel Santos-Bolívar1.1 Servicio de Obstetricia y Ginecología. Hospital Central “Dr. Urquinaona”. Maracaibo. Estado Zulia. 2 Departamento de Obstetricia y Ginecología. Unidad Docente Hospital Chiquinquirá. Universidad del Zulia. Maracaibo. Estado ZuliaRecibido: 13 de Febrero de 2014. Aceptado: 09 de Mayo de 2014.Publicacion online: 11 de Junio de 2014 [ARTÍCULO DE REVISION]Resumen (español)Durante la embriogénesis y el desarrollo, el feto obtiene oxígeno y nutrientes de la madre a través de la microcirculación placentaria. La placenta es un órgano especial que se desarrolla y diferencia por sí sólo, y que organiza el crecimiento fetal y la condición materna durante la duración completa del embarazo. El conocimiento del flujo sanguíneo úteroplacentario temprano suministra un contexto fisiológico para el proceso de invasión de las arterias espirales y la remodelación asociada, evidenciándose una asociación cercana entre el incremento del flujo y los primeros pasos de la remodelación vascular.Palabras clave (español)Lecho placentario; invasión trofoblástica; remodelación arterias espiralesAbstract (english)During embryogenesis and development, the fetus obtains oxygen and nutrients from the mother through placental microcirculation. The placenta is a special organ that develops and differentiates per se, and that organizes fetal growth and maternal condition in the entire course of gestation. Knowledge of early uteroplacental blood flow has provided a physiological context for the processes of spiral artery invasion and associated remodeling, revealing a closely timed relationship between increasing flow and early steps in vascular remodeling.Keywords (english)Placental bed; trophoblast invasion; spiral artery remodeling.Introducción La placenta inicia la organogénesis en una etapa muy temprana del desarrollo embrionario, comanda el crecimiento fetal y determina su propio destino inmediatamente después del parto. La perfusión placentaria es mantenida por dos sistemas vasculares distintos: flujo sanguíneo materno y circulación fetal. Por lo tanto, la fisiopatología de la placentación está claramente asociada tanto con las condiciones maternas como con el desarrollo fetal. Una amplia variedad de moléculas angiogénicas y enzimas proteolíticas representan un papel crítico en el establecimiento de la placentación y desarrollo del sistema circulatorio placentario (1-4). Por ejemplo, el factor de crecimiento endotelial vascular (VEGF), factor de crecimiento de los fibroblastos (FGF), y el factor de crecimiento placentario (PIGF), son indispensables durante toda la gestación (5,6) y en fases más tardías del embarazo, el trofoblasto velloso y los vasos sanguíneos del lado fetal de las vellosidades terminales forman una red vascular diferenciada que aporta al feto cantidades suficientes de oxígeno y sustancias para su crecimiento exponencial (2). La circulación arterial en la placenta carece de inervación autonómica y es regulada por factores locales como la presión y el flujo (7). Si el proceso de implantación no se completa en forma exitosa, la placenta sufre de una perfusión insuficiente y secreta varias moléculas pro-inflamatorias que dañan las células endoteliales (ECs) maternas y, en consecuencia, se incrementa la resistencia vascular que posteriormente afecta los órganos maternos por la hipertensión, al igual que agrava la circulación feto-placentaria (8,9).Se considera que la respuesta inflamatoria sistémica y la disfunción de las ECs maternas representan el esquema patológico de la preeclampsia (8-10). Varias citokinas/péptidos pro-inflamatorios incluyendo la forma soluble del receptor de VEGF-1 o kinasa de tirosina soluble similar al FMS soluble (sFlt1) y la forma soluble del CD105, también conocida como forma soluble de endoglina, están elevadas en la el suero de las preeclámpticas y estos factores anti-angiogénicos juegan papeles críticos en la disfunción de las ECs (10,11). Por otra parte, aún es poco comprendido el curso patológico de la placenta y los mecanismos que llevan a la preeclampsia y la restricción del crecimiento intrauterino del feto (RCIU). Con relación a la disfunción placentaria, los análisis están basados principalmente en los estudios Doppler y sobre los estudios patológicos de placentas a término (12), por lo que la dinámica de las señales vasoactivas en la placenta durante la aparición de la hipertensión continúa siendo un enigma. El objetivo de la presente revisión una visión general de la implantación normal y de los diferentes mecanismos implicados en la implantación, placentación inicial y regulación de la invasión trofoblástica.Estructura de la red vascular placentariaLa placenta normal a término está dividida en tres capas: 1) Membrana basal en la superficie materna y las vellosidades de anclaje, en las extensiones más dístales de las vellosidades primarias, que interactúan directamente con el endometrio materno. 2) Unidad vellosa terminal donde ocurre en forma activa el intercambio de gas y nutrientes. 3) Lámina coriónica en la superficie fetal y las vellosidades que se mezclan en un tejido conectivo denso que contienen vasos fetales. El amnios y el corión son membranas que cubren la lámina coriónica y el cordón umbilical recoge la sangre de las arterias y venas coriónicas sobre la lámina coriónica generalmente en vasos propios en su parte central (5).La estructura fundamental de la placenta se establece durante la primera mitad del embarazo (5). En la placenta humana, la unidad vellosa terminal (vellosidades terciarias que se desarrollan de las secundarias) está compuesta principalmente de capilares del lado fetal alineados con ECs, colágeno mesenquimal y rodeado por sinciciotrofoblasto. En estadios tempranos, el citotrofoblasto se alinea por debajo del sinciciotrofoblasto y a medida que progresa el embarazo, las láminas del citotrofoblasto se vuelven indetectables y los capilares fetales se colocan en cercana proximidad a la circulación intervellosa materna, con el objetivo de hacer más eficiente el intercambio de nutrientes y gases. Los espacios con sangre materna están directamente alineados por el sinciciotrofoblasto diferenciado terminal y no por las ECs en la interfase hemocorial (13).El peso fetal se incrementa casi dos veces durante la etapa final del embarazo. Sin embargo, el peso de la placenta no se incrementa significativamente. Por el contrario, las redes vasculares en las vellosidades terminales se diferencian aún más se incrementa la capacidad funcional tanto en los capilares del lado fetal como los sinusoides sanguíneos maternos (2, 14). En el RCIU, la diferenciación de las vellosidades terminales generalmente está alterada, y se hace evidente la falta de crecimiento de las vellosidades dístales. El flujo sanguíneo del cuerpo fetal en condiciones de hipoxia se acumula en los órganos más críticos como el cerebro y corazón. Esta redistribución reduce el flujo e incrementa la resistencia vascular (15).Zona de unión del miometrio137Tanto el endometrio como la zona de unión del miometrio (ZUM) se derivan de los conductos paramesonéfricos embrionarios, mientras que el miometrio exterior es de origen diferente (16). El hecho que la porción interna del miometrio humano constituye una entidad separada dentro de la musculatura uterina tiene más de un siglo de antigüedad, y ha sido descrita como “arquimiometrio” (17). A pesar de esta descripción temprana, fue con la disponibilidad de la ecografía de alta resolución y la resonancia magnética que finalmente se confirmó que el miometrio interno es una estructura distinta, no solo por su origen embrionario sino también desde el punto de vista de sus funciones especializadas (18). En las imágenes de resonancia magnética, la zona de unión representa una banda de baja densidad, ubicada entre el miometrio exterior y el endometrio cíclico. También se ha descrito como “una banda de unión de baja densidad” (19).Figura 1. Arteria uteroplacentaria con marcada distensión y reemplazo del tejido muscular y elástico de la pared por fibrinoide y trofoblasto invasor.Existe una considerable variación en el grosor y la apariencia de la ZUM, no solo entre individuos sino según el estado hormonal (menarquía, menopausia, fase del ciclo menstrual y uso de anticonceptivos orales), lo que hace difícil definir una zona anatómica uterina “normal” en la resonancia magnética (20). Inicialmente se pensó que la ZUM podría corresponder al “estrato basal” del endometrio (21), debido a que es un área de perfusión aumentada; sin embargo, esto también fue una explicación inadecuada debido a que la ZUM permanecía visible en úteros recién extraídos (22). El único análisis morfométrico e inmunohistoquímico fue realizado a finales del siglo pasado en el que se encontró que los miocitos tenían una mayor relación núcleo-citoplasma, comparado con los músculos lisos de la zona miometrial externa, junto con una intensa coloración con el marcador endotelial vascular CD31, reflejando una mayor vascularidad o un mayor nivel de “activación” del endotelio (23). Esta arquitectura celular sugiere que el contenido general de agua es relativamente más bajo, lo cual altera la impedancia acústica y explica su apariencia en la ecografía. La ZUM muestra un patrón cíclico de expresión de receptores de estrógenos y progesterona, similar al del endometrio, sufriendo cambios cíclicos en el grosor, los cuales son similares, pero menos pronunciados que los observados en el endometrio (24). Los avances en la ecografía han demostrado que las contracciones miometriales en el útero no gestante se originan exclusivamente de la ZUM y son dependientes de la fase del ciclo menstrual. Una baja frecuencia de contractilidad de la ZUM antes de la transferencia de embriones se asocia con una mayor tasa de implantación y de embarazos (25). Todo lo anterior indica que mientras la ZUM es morfológicamente similar al miometrio exterior, pero es funcionalmente más parecida al endometrio, especialmente en función de los cambios dinámicos en respuesta a las variaciones de las hormonas ováricas.Flujo sanguíneo úteroplacentarioRemodelación de las arterias espirales.138Después que se identificaron los cambios fisiológicos de las arterias espirales, se propuso que estos eran el resultado de la acción destructiva del trofoblasto sobre la vasculatura muscular y la membrana elástica (figura 1). Sin embargo, posteriormente se observó que los cambios asociados con la invasión del trofoblasto eran precedidos por edema de la pared y cambios en las células musculares lisas con pérdida del núcleo y miofibrillas, cuerpos densos y acumulación de glicógeno (26).Investigaciones con muestras de tejido obtenidas de histerectomías entre 8-18 semanas de embarazo llevaron a dos hallazgos importantes (27,28). Primero, los cambios vasculares que incluyen desorganización de la pared muscular no pueden contribuir exclusivamente a la actividad del trofoblasto, ya que se ha descrito que el músculo liso vascular se desorganiza antes de la llegada del trofoblasto endovascular; sin embargo, esta desorganización es acentuada por la presencia del trofoblasto intersticial. El segundo hallazgo fue la aparente presencia de invasión endovascular en la ZUM, lo cual fue considerado la segunda “oleada” de la invasión trofoblástica, la cual ocurre después de un periodo de 4 semanas del trofoblasto dentro de la decidua. Aunque el “concepto de dos oleadas” no es aceptado en forma universal (29), suministra un modelo valioso para considerar los posibles mecanismos responsables para la alteración de la placentación profunda. La representación de los periodos de la invasión intersticial y endovascular del trofoblasto a la decidua y a la zona de unión del miometrio se muestra en la figura 2.Figura 2. Representación de los periodos de la invasión intersticial y endovascular del trofoblasto a la decidua y a la zona de unión del miometrioUn aspecto clave es la contribución relativa del trofoblasto y la decidua en la remodelación vascular de las arterias espirales. Craven y col. (30) compararon las características histológicas de las arterias espirales en la fase secretora del ciclo menstrual usando muestras de biopsias endometriales y arterias deciduales de pacientes sometidas a terminación electiva del embarazo y concluyeron que las fases iniciales de los cambios fisiológicos de las arterias espirales ocurren sin la presencia de invasión trofoblástica. Sin embargo, Herr y col. (31) describieron que la “necrosis fibrinoide” de la pared no ocurre en ausencia de trofoblasto. Kam y col. (32) compararon los vasos sanguíneos de los sitios de implantación en embarazos tempranos con muestras en las cuales el trofoblasto está ausente. Los resultados confirmaron que la verdadera transformación fisiológica de las arterias espirales ocurre solo en presencia del trofoblasto. Una investigación que examinó las muestra de decidua basal (8-12 semanas de gestación) utilizando inmunohistoquímica y se observó que las células natural killer uterinas (uNK o también llamadas células natural killer deciduales) y los macrófagos participan en la remodelación a través de la inducción de apoptosis o degradación de la matriz extracelular y en fases tempranas, las células del músculo liso vascular muestran una dramática alteración y desorganización antes de la presencia de trofoblasto endovascular (28).Placentación profunda. Dos factores determinan el flujo de sangre materno hacia la placenta. El primero es el tamaño del lecho placentario, el cual está determinado por el número de arterias espirales que se comunican con el espacio intervelloso. En una investigación en la cual se reconstruyó la membrana basal de las placentas de embarazadas normales, se encontró una distribución irregular de las aperturas arteriales en el espacio intervelloso y encontraron que estaban reunidas en grupos de 2-3, en cercana proximidad al septo placentario (33). En un estudio posterior se encontró que cada apertura correspondía a una densidad de 1 arteria por cada 2 cm2 de la membrana basal. Los cortes seriados de muestras de histerectomías demostraron que las arterias radiales se dividían aproximadamente a 0,5 centímetros más allá del endometrio en 2-3 arterias con cambios o transformación fisiológica. 139El segundo factor es el que la profundidad de la transformación fisiológica de las arterias espirales es mayor en el centro del lecho placentario que en la periferia (28,34). Esto es consistente con las observaciones que el grado de invasión trofoblástica es menor en la periferia que en el centro del lecho placentario, demostrando que el trofoblasto intersticial está ausente o es escaso en la periferia, y la transformación fisiológica del segmento miometrial de las arterias espirales es parcial o está ausente, aún en el embarazo normal. Sin embargo, tal fenómeno involucra aproximadamente el 10% de las arterias espirales del lecho placentario (33). Los estudios de las biopsias del lecho placentario confirman que la mayoría de las arterias espirales muestran transformación completa en la ZUM. Estos hallazgos son consistentes con las observaciones reportadas por ultrasonido. Los estudios con ecografía Doppler realizados durante el segundo trimestre del embarazo han demostrado una más baja impedancia del flujo sanguíneo del área central del lecho placentario que de la periferia (35).Fase temprana del embarazo: transformación epitelial endotelial140Es bien conocido que durante la implantación el trofoblasto extravelloso invade el endometrio y también el tercio interno del miometrio (36). Para establecer un flujo sanguíneo suficiente desde la circulación materna, el trofoblasto sufre una especialización similar a ECs y expresa algunos marcadores de ECs como por ejemplo CD31, VE-caderina, VCAM-1 e integrina αvβ3 (37). Además de la integrina αvβ3, este regula la expresión de las integrinas α5β1 y α1β1 y disminuye la expresión de la integrina α6β4 (38). Este proceso es llamado “transformación epitelial-endotelial” o “pseudovasculogénesis” (39). Un proceso similar de remodelación vascular ha sido reportado en varios tipos de neoplasias, como el melanoma agresivo, carcinomas prostático, ovárico y mamario, las células tumorales producen sinusoides sanguíneas (6). Las células tumorales que forman esta red vascular alternativa expresan ciertos marcadores de ECs y moléculas relacionadas con la vasculogénesis embrionaria como caderina-VE, CD34 y CD105 (40). La invasión cancerígena es destructiva y desorganizada, mientras que la invasión trofoblástica es finamente controlada por microambientes proinflamatorios locales bajo condiciones fisiológicas. Las células deciduales e inmunes gobiernan las interacciones no solo al aceptar células “alogénicas”, sino al restringir la invasión excesivaPapel de las células inmunes locales en la determinación de la respuesta hormonalExiste evidencia que la polarización del endometrio y el miometrio es efectuado a través de la interacción entre las hormonas ováricas y factores liberados localmente (41). Las citokinas y los factores de crecimiento liberados por las células inmunes (incluyendo células T, células uNK, neutrófilos polimorfonucleares, macrófagos y monocitos) juegan un papel primordial en el establecimiento de microambientes específicos de las láminas endometrial superficial y basal (42,43). Se considera que el interferón gamma secretado por los agregados linfoideos en la lámina endometrial basal contribuye a la baja actividad apoptoica y proliferativa en estas láminas y esto puede llevar a un aumento en la expresión local de sustancias dependientes del interferón gamma, como los antígenos del complejo de histocompatibilidad mayor clase II y la proteína 70 de choque de calor (42). También regula los genes involucrados en la proliferación y contractilidad del músculo liso como la sintetasa inducible de óxido nítrico y endotelina 1. Por lo tanto, parece posible que el interferón gamma producido en la ZUM está dirigido a los miocitos de esta área. Comparado con el endometrio exterior, un gran número de mastocitos están presentes y producen una amplia variedad de moléculas reguladoras, como histamina, serotonina, heparina, factor de necrosis tumoral alfa, interleucina 1, proteasas específicas y triptasa, que son capaces de participar en varias funciones biológicas incluyendo la remodelación de la matriz extracelular, angiogénesis, quimiotaxis y diferenciación del músculo liso (44).La distribución espacio-temporal de las células inmunes uterinas está controlada por las hormonas ováricas. Los agregados linfocitarios en la membrana basal son pequeños durante la fase proliferativa temprana pero aumentan en forma significativa durante la segunda mitad del ciclo (45). Durante la fase proliferativa, la lámina endometrial superficial contienen solo unas pocas células uNK, macrófagos y células T dispersa en el estroma y glándulas. Después de la ovulación, el número de células uNK (pero no las células T y los macrófagos) aumentan dramáticamente hasta unos días antes de la menstruación (43). En el embarazo temprano, 30% de las células del estroma decidual son leucocitos, de los cuales aproximadamente 70% son células NK y 30% son macrófagos (46). Aunque la función de las células uNK no está completamente descrita, se piensa que representan un papel importante en la reacción decidual, remodelación de las arterias espirales en la decidua y regulación inmune de la invasión trofoblástica (47,48).Aspectos moleculares de la implantación141Tabla 1. Características generales de los pacientes con anemia hemolitica hereditaria estudiados.FactorNombrePapel sugeridoCitosinas yFactoresde crecimientoInterleucina 1Interleucina 2Factor inhibidor de la leucemiaFactor de crecimiento similar a la insulina I y IIFactor I estimulante de coloniasFactor transformante y de desarrolloFactor de crecimiento de hepatocitosFactor de crecimiento de fibroblastosFactor de crecimiento epidérmico unido al heparanFactor I inhibidor de la hipoxiaFactor de crecimiento vascular endotelialFacilitar la comunicación entre el blastocisto y el útero, promoción de la proliferación y diferenciación endometrial y regulación de la angiogénesis endometrial y permeabilidad vascular.Hormonas esteroideasEstradiol-17Proliferación y diferenciación de las células epiteliales uterinasProgesteronaProliferación y diferenciación de las células del estroma endometrialCatecolestrogénosUn metabolito del estrógeno que activa el blastocistos en preparación para la implantaciónFactoresinmunológicosInterleucina 10Inmunosupresiónindolamina 2,3-dioxigenasaUna enzima que degrada el triptófano el cual es esencial para la acción de los macrófagosCrry (regulador complementario)InmunosupresiónCambios en elepitelio luminalPinopodosMuc-1GlicodelinaIntegrina αvβ3CalcitoninaPueden ser importantes para el reconocimiento y adhesión del blastocistoFactores detranscripciónMash-2Hand-1Inhibidores de las proteínas de unión del ADNFactor 4 de transcripción de octamerosReceptores de estrógenoReceptor activado de proliferación de peroxisomasDefinición de los mecanismos moleculares por los cuales los factores reguladores ejercen su efecto a nivel celularOtros factoresHoxa 10 y Hoxa-11Genes que se expresan en las células del estroma endometrial al momento de la implantaciónCiclooxigenasa 2Regulación la producción de prostaglandinasTensión de oxigenoFacilitar la invasión vascular trofoblástica al afectar la expresión de las integrinasLa implantación exitosa es el resultado final de un dialogo molecular complejo, entre un útero modificado por el efecto hormonal y un blastocisto activo y maduro. Aunque los mecanismos celulares y moleculares responsables de la implantación no son bien conocidos, está claro que son necesarias múltiples señales para sincronizar la maduración del blastoscito y la receptividad uterina, incluyendo hormonas esteroideas y peptídicas, factores de crecimiento, citoquinas y factores inmunológicos. Un resumen de estos factores en estudios en animales y en humanos, in vitro e in-vivo, se presenta en la tabla 1 y 2 (49,50). Las alteraciones en uno o más de estos factores pueden llevar a la pérdida del embarazo (aborto espontáneo) o defectos de la implantación que se traduce en complicaciones clínicas como la preeclampsia o el parto pretérmino.Tabla 2. Factores ovulares asociados a la implantación y el mantenimiento del embarazo temprano.FactorNombrePapel sugeridoCitosinas yfactoresde crecimientoInterleucina 1Interleucina 6Factor inhibidor de la leucemiaFactor transformante y de crecimientoFactor II similar a la insulinaFactor I estimulante de coloniasFacilitar la comunicación entre el blastocisto y el útero, puede promover la diferenciación e invasión trofoblasticaProteinasas einhibidoresMetaloproteinasas de la matriz 9/ inhibidor tisular de la metaloproteinasa 3Regulación la invasión del trofoblastoActivador del plasminógeno tipo urocinasa/inhibidor 4 del activador del plasminógenoFacilitar la invasión vascular del trofoblastoCaptesina B y LRegulación la invasión del trofoblastoHormonasGonadotropina coriónica humanaMantenimiento de la liberación de progesterona del cuerpo lúteoFactoresinmunológicosHLA-GPuede prevenir el rechazo inmune al aloinjerto fetalExpresión demoléculas de adhesiónIntegrina α6β4 E-caderina Integrina α1v3β3 VE-caderinaPromoción de la diferenciación e invasión trofoblásticaOtros factoresProstaglandina E2 Factor activador de plaquetas Ciclooxigenasa 2 Tensión de oxigenoPromoción de la apoptosis endometrial Estimulación de la producción de prostaglandinas uterinas Regulación la producción de prostaglandinas Facilitar la invasión vascular trofoblástica al afectar la expresión de las integrinasPapel de las prostaglandinas en la implantación y placentación142Las concentraciones de prostaglandinas endógenas en la decidua humana son menores en el embarazo temprano que en el endometrio en cualquier fase del ciclo menstrual (49,50), debido principalmente a la disminución de la síntesis de prostaglandinas y al mantenimiento del metabolismo (50). La administración de prostaglandinas exógenas (intravenosa, intraamniótica o vaginal) en cualquier momento de la gestación temprana induce la pérdida del embarazo. Más aún, la incapacidad de suprimir las prostaglandinas deciduales en el momento de la implantación se ha asociado con pérdida temprana del embarazo. Todos estos datos en conjunto sugieren que el embarazo es mantenido por un mecanismo que suprime la síntesis de prostaglandinas uterinas. Debido a que la producción de prostaglandinas endometriales está disminuida en el embarazo ectópico (49), parece posible que estén involucrados factores sistémicos más que locales. La candidata más probable para esta regulación es la progesterona. Los antagonistas de los receptores de progesterona (como el RU-486) inducen el aborto si se administra antes de las 7 semanas de embarazo (51). En forma similar, la remoción quirúrgica del cuerpo lúteo, la fuente de progesterona durante el primer trimestre, produce pérdida espontánea del embarazo (52). Estos datos sugieren que la producción adecuada de progesterona es crítica para el mantenimiento del embarazo hasta que la placenta comienza a funcionar alrededor de las 7-9 semanas. El mecanismo de acción de la progesterona no es bien comprendido, pero parece que es parcialmente independiente a la interacción con los receptores de progesterona o glucocorticoides, pero es posible que actúe disminuyendo la producción de prostaglandinas endometriales, directamente inhibiendo la liberación de ácido araquidónico de las células endometriales (53,54) e indirectamente aumentando la inhibición de la síntesis de prostaglandinas (50). La progesterona también puede actuar alterando la actividad de los receptores de prostaglandinas. En los ratones con ausencia del gen HOXA-10, por ejemplo, tienen un fenotipo reproductivo caracterizado por anomalías de la decidualización e implantación que ha sido relacionada con modificaciones en la expresión de dos subtipos de receptores de prostaglandinas reguladas por la progesterona, EP3 y EP4, en el estroma decidual (55). Se ha identificado la presencia en la decidua de dos inhibidores endógenos: componente secretor y proteína A surfactante de la producción de prostaglandinas que pueden ser importantes en la implantación (50). El paso limitante por la tasa de la síntesis de prostaglandinas de la serie 2 es la hidrólisis de ácido araquidónico no esterificado (libre) de los fosfolípidos de la membrana, la cual es mediada por las enzimas de la familia de la fosfolipasa, principalmente por la fosfolipasa A2 (PLA2). En 1985, se identificó un inhibidor endógeno de la PLA2 en el líquido amniótico durante el tercer trimestre del embarazo, un compuesto llamado “gravidina”, pero posteriormente se ha que es un homologo fetal al componente secretor del adulto de las inmunoglobulinas poliméricas (56). El componente secretor es una glicoproteína de 58 kD que se une a la inmunoglobulina (Ig) A y M sobre la superficie luminal de las células glandulares epiteliales, transportados a través de las glándulas y liberados en las secreciones exocrinas unidos en forma covalente a la inmunoglobulina. En su forma no unida (libre) dentro de las células glandulares endometriales el componente secretor, al igual que la gravidina, es capaz de inhibir la actividad de la PLA2 (56). La producción del componente secretor y prostaglandinas ha sido localizada principalmente en las células epiteliales glandulares en la decidua y se ha demostrado que la progesterona estimula la expresión de componente secretor en estas células produciendo una rápida y marcada disminución en la liberación de ácido araquidónico y la producción de prostaglandinas (50). Debido a que la producción de progesterona por el cuerpo lúteo es mantenida por la liberación de la gonadotropina coriónica por el sinciciotrofoblasto, este puede ser el mecanismo por el cual el blastocisto mantiene el embarazo en fases tempranas.Por su parte, el surfactante es un complejo de lipoproteína compartimentalizado producido por los neumocitos tipo II en el pulmón y que actúa disminuyendo la tensión superficial evitando el colapso alveolar y contiene varias glicoproteínas asociadas. Las proteínas B y C del surfactante son críticas para la función pulmonar; las proteínas A y D, por otro lado, son lectinas tipo C (colectinas) y son importantes reguladores del sistema inmune innnato. La proteína A ha sido identificada también en tejidos extra-pulmonares, incluyendo el tracto reproductivo femenino (57). Se ha demostrado que las altas dosis de proteína A (100 μg/mL) inhiben selectivamente la producción de prostaglandinas por las células estromales deciduales in vitro sin afectar la producción de otros mediadores inflamatorios y factores angiogénicos. Además, parece ejercer sus efectos por la unión directa a la proteína citoplasmática peroxiredonina 6, la cual tienen una actividad endógena sobre la PLA2 (50,58). El papel de la proteína A en la implantación no ha sido completamente delineado. Microambiente proinflamatorio en el sitio de implantaciónLos microambientes de la unión úteroplacentario son cruciales para el proceso de inflamación y desarrollo fetal durante el embarazo. En respuesta a la aparición de la gonadotropina coriónica (hCG), el endometrio sufre cambios deciduales (59) y las células mononucleares se acumulan en el endometrio decidualizado. Las uNK son células inmunes CD45+/CD69+/CD56/CD16- que representan la mayoría de los leucocitos en el sitio de implantación (47,60,61).En el embarazo en humanos, el trofoblasto expresa antígenos leucocitarios humanos (HLA) del tipo C, E y G, un repertorio único de ligandos para los receptores de las células uNK (47,61,62). Las HLA-A y HLA-B, moléculas clase I del complejo mayor de histocompatibilidad (MHC) con los polimorfismos que inician el rechazo de los aloinjertos, no se expresa en el trofoblasto velloso y tampoco se expresan las moléculas clase II del MHC (47). Es también de hacer notar que en el sitio feto-placentario, el sinciciotrofoblasto no expresa antígenos MHC en la superficie celular (61).143Existe más de una vía de control de la función de las células uNK. Tienen receptores de superficie tanto estimuladores como inhibidores incluyendo los de la familia NKG2 (CD94), 2B4 (CD244) y NKp46 (CD335) (60,63). El HLA-E del trofoblasto interactúa con los receptores NKG2A de las células uNK (62), lo cual explica en parte el mecanismo de supresión de la citotoxicidad de las células uNK contra el trofoblasto invasor. Es necesaria la activación cruzada entre el trofoblasto invasor y las células uNK. Como se mencionó anteriormente, las moléculas clase I HLA-A y HLA-B del MHC no se expresan en el trofoblasto extravelloso, pero otra molécula clase I clásica, HLA-C, está presente en el primer trimestre (63).ReferencesKharfi A, Giguère Y, Sapin V, Massé J, Dastugue B, Forest JC. 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