Alberto Sols Biomedical Research Institute
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Organización del citoesqueleto de actina en las barreras celulares de la córnea y su respuesta al estrés. Búsqueda de biomarcadores de queratocono
No full textTesis Doctoral inédita leída en la Universidad Autónoma de Madrid, Facultad de Ciencias, Departamento de Biología Molecular. Fecha de Lectura: 08-03-2025Esta Tesis tiene embargado el acceso al texto completo hasta el 08-09-2025La córnea, como la primera lente del sistema ocular, desempeña un papel crucial en la salud visual. La queratoplastia, el trasplante más común, destaca la necesidad de comprender mejor el impacto del cultivo y almacenamiento en biobancos en las córneas humanas. Compuesta por un epitelio externo, un estroma y una monocapa endotelial interna, la córnea requiere la función de barrera de su endotelio para mantener la transparencia del tejido, especialmente bajo condiciones subóptimas como patologías corneales, envejecimiento y trasplantes.
La primera parte de la investigación se centra en la caracterización de la organización celular del endotelio corneal y las adaptaciones estructurales durante el almacenamiento para trasplante. Utilizando microscopía confocal de alta resolución y ratones como modelo, se observó que las células endoteliales corneales murinas, sometidas a la curvatura del tejido y la presión del humor acuoso, presentan una organización distintiva. El dominio apical es poligonal, con uniones intercelulares estrechas y adherentes conectadas a un anillo de actina filamentosa prominente. En contraste, el dominio basolateral forma contactos intercelulares sin uniones estrechas, con bajo contenido de F-actina y morfología ondulada con protrusiones que se imbrican con las células vecinas. Desde las uniones apicales poligonales, emergen haces de F-actina radial que convergen debajo del núcleo, denominadas puntos SFC (Subnuclear F-actin convergence), ricos en actomiosina, sugiriendo una zona de contracción mediada por actina.
El análisis detallado de la F-actina y las proteínas conectoras de las uniones intercelulares, ZO-1 y α-catenin, reveló que el cultivo de córneas en el medio de trasplante TissueC, induce una reorganización transitoria de las regiones SFC, aumento significativo de actomiosina y, por ende, de la contractilidad celular. La inyección de vectores lentivirales en el humor acuoso de ratones permitió expresar proteínas fluorescentes in vivo y analizar la dinámica de actomiosina y uniones intercelulares ex vivo.
Estos resultados sugieren una conexión entre las uniones intercelulares, el citoesqueleto de actomiosina y el núcleo. El análisis de componentes del complejo LINC, que conecta la pared nuclear con el citoesqueleto de actina citoplásmico, mediante la expresión de un vector lentiviral inducible del dominante negativo de nesprina-2, DN-KASH, alteró las uniones intercelulares corneales. Además, en córneas de un modelo murino de progeria, la expresión de progerina, una proteína mutante de la pared nuclear que causa envejecimiento acelerado, afectó el citoesqueleto de actina endotelial y las uniones intercelulares. En conjunto, estos resultados sugieren que alteraciones patológicas en la pared nuclear alteran la función de barrera del endotelio de córnea.
A diferencia del endotelio, el epitelio de córnea regenera fácilmente y es accesible para análisis moleculares. En colaboración con Cornea Project S.L en este doctorado industrial, en la segunda parte de esta investigación se identificaron biomarcadores específicos en el epitelio de córnea humana para el diagnóstico temprano y pronóstico del queratocono. Cambios de expresión en seis genes humanos permitieron generar un clasificador para la enfermedad en sus fases iniciales basado en el análisis de tres genes, indicando un nuevo método potencial para el diagnóstico temprano o pronóstico de esta patologí
Nuevas técnicas de análisis para la optimización del uso de la cesárea
No full textTesis Doctoral inédita leída en la Universidad Autónoma de Madrid, Facultad de Ciencias, Departamento de Biología. Fecha de Lectura: 08-03-2024Esta Tesis tiene embargado el acceso al texto completo hasta el 08-09-202
Un análisis transcultural de los conceptos "justicia, inocencia, culpa y vergüenza" en inglés, español y ruso basado en corpus
No full textTesis Doctoral inédita leída en la Universidad Autónoma de Madrid, Facultad de Medicina, Departamento de Filología Inglesa. Fecha de Lectura: 15-03-2024Esta Tesis tiene embargado el acceso al texto completo hasta el 15-09-202
Silicon (Si) mitigates the negative effects of iron deficiency in common bean (Phaseolus vulgaris L.) by improving photosystem activities and nutritional status
No full textSilicon (Si) is the second most abundant element in the Earth's crust after oxygen. Its beneficial impact on crop development and yield, particularly under stressful conditions such as iron (Fe) deficiency, has been well documented. Fe deficiency is a critical constraint that limits crop production globally. The objective of this study was to investigate the effects of silicon (Na2SiO3) on common bean (Phaseolus vulgaris L. ‘Coco Rose’ variety) under iron-deficient conditions. The common bean plants were subjected to six treatments, which included three sufficient iron treatments (50 μM Fe) each paired with three varying silicon concentrations (0, 0.25, and 0.5 mM Si), and three iron-deficient treatments (0.1 μM Fe) each associated with the same silicon concentrations (0, 0.25, and 0.5 mM Si). The results indicate that iron deficiency had a negative impact on almost all the measured parameters. However, under silicon treatments, especially with 0.5 mM Si, the depressive effects of iron deficiency were significantly mitigated. The addition of 0.5 mM Si alleviated leaf chlorosis and improved biomass production, nutritional status, photosynthetic pigment content, photosynthetic gas exchange, and photosystem (PSI and PSII) activities. Interestingly, a greater beneficial effect of silicon was observed on PSII compared to PSI. This was accompanied by a significant augmentation in leaf iron concentration by 42%. Therefore, by enhancing the photosystem activities and nutritional status, among other mechanisms, silicon is capable of mitigating the adverse effects of iron-deficient conditions, making it a successful and effective solution to cope with this nutritional stressThis work was funded by the Tunisian Ministry of Higher Education and Scientific Research under grant LR15CBBC02. A portion of the work was also supported by the PHC Maghreb 2019 network (Partenariat Hubert CURIEN Maghreb 2019 no. 19MAG41) and the Young Researchers Encouragement Program (Programme d’Encouragement des Jeunes Chercheurs - 21PEJCD1P22 for Tarek SLATNI and 21PEJCD4P1 for Walid ZORRIG
Bio-recovery of CuS nanoparticles from the treatment of acid mine drainage with potential photocatalytic and antibacterial applications
No full textIn the present work, CuS nanoparticles were biorecovered from a real acid mine drainage (AMD) and its photocatalytic and antibacterial activities were studied. CuS were formed by delivering biogenic H2S produced by a continuous sulfidogenic bioreactor to an off-line vessel containing the AMD. The main physico-chemical properties of CuS nanoparticles were analyzed by UV–vis spectroscopy, TEM, FE-SEM, XRD and XPS. Moreover, its photocatalytic activity on the photodegradation of organic dyes in water and its antibacterial activity against several bacterial strains were studied and compared with CuS nanoparticles synthetized from a CuSO4 aqueous solution based on the same synthesis method. CuS nanoparticles from the real AMD showed similar physico-chemical properties and photocatalytic and antibacterial activities in comparison to CuS nanoparticles formed with the copper solutions. These results open the way to recover valorous CuS nanoparticles from AMD with potential industrial applications using a metal bioremediation process based on sulfidogenic bioreactorsThe authors of this work gratefully thank the financial support of FONDECYT Regular 1230412. IN is funded by FONDEF ID21I10111 and FONDECYT 1221604. Financial support from grant PID2020- 116712RB-C21 funded by MCIN/AEI/10.13039/501100011033 is acknowledged by A.A
Aprendiendo de la biología viral: componentes celulares involucrados en la respuesta inmunitaria frente a EBV y SARS-CoV-2
No full textTesis Doctoral inédita leída en la Universidad Autónoma de Madrid, Facultad de Ciencias, Departamento de Biología Molecular. Fecha de Lectura: 12-04-2024Esta tesis tiene embargado el acceso al texto completo hasta el 12-10-2025Immunity against the Epstein-Barr virus has been a subject of research since its
discovery in the 1960s, although it has never been studied with a sufficient level of
complexity to describe the essential cellular populations in the memory cellular immune
response to this virus. In this work, we were able to establish a protocol for PBMC infection,
detecting lymphocytes that respond to EBV through proliferation or expression of activation
markers and cytokines. This system allowed us to sequence, for the first time at a single-cell
level, the transcriptome of all cells capable of responding to the virus in healthy individuals
and in certain immunodeficient patients with diseases associated with EBV infection. In this
way, we established that cytotoxic CD8 T cells, expressing CD27 and EOMES, seem to be crucial
in the proper elimination of EBV-infected cells, as these cells are either absent or appear to
undergo certain functional modifications in our samples from individuals with Inborn Errors of
Immunity. CD4 T cells comprised a variety of subpopulations, among which the presence of
Th1 cells (CXCR3+CCR2+) and regulatory cells (IL2RA+FOXP3+) is noteworthy, but these exhibited
a much lower capacity for clonal expansion than CD8 T cells. NK cells largely aligned with what
has been described in the literature, predominantly fitting a dim phenotype with expression of
KLRK1, KLRC1, and FCGR3A, while TRDV2 cells were able to express CD27 and TNF abundantly,
highlighting their helper role in anti-tumoral immunity. We also detected TRDV1 cells, with an
activated and cytotoxic phenotype, as well as MAIT cells, with a basal transcriptome similar to
TRDV2. In most of these populations, we confirmed by flow cytometry, in general, an increase
in molecules such as IFN-γ or LAMP1 after incubating PBMCs with EBV. This led us to
hypothesize that TRDV1, rare in blood but highly functional against the virus, could be a great
candidate for a potential allogeneic therapy in patients with diseases arising from EBV
infection. Thus, we established a protocol for expanding cytotoxic TRDV1 cells and confirmed
their ability to kill and degranulate against LCLs, primarily through innate interactions. All these
results represent a level of characterization of the immune response to EBV never before
undertaken, and pave the way for further research that may ultimately lead to the
development of methods to improve the lives of individuals with EBV-associated problems.
On the other hand, despite all the research conducted, many questions still remain regarding
humoral and cellular immunity against SARS-CoV-2. One key question is whether the presence
of memory T or B cells with affinity for seasonal coronavirus antigens can recognize and be
functional against SARS-CoV-2 antigens. To delve into this, we studied sera and peripheral
blood samples from convalescent COVID-19 patients and prepandemic donors. First and
foremost, we confirmed that antibodies against the SARS-CoV-2 nucleoprotein were only
detectable in serum samples from COVID-19 patients. However, we detected cross-reactivity
in T lymphocytes, which were capable of proliferating and producing IFN-γ in response to
SARS-CoV-2 antigens in approximately 30% of the donor samples collected before the
pandemic. Interestingly, these T cell responses exhibited very weak activation of TFH cells
when stimulated with the nucleoprotein of HCoV-OC43 or SARS-CoV-2. In conclusion, our
observations confirm the presence of T cell cross-reactivity with SARS-CoV-2 antigens in
individuals not exposed to SARS-CoV-2. Nonetheless, they suggest that the T cell response to
HCoV-OC43 may be deficient in certain aspects, such as TFH cell expansion, which could
compromise the generation of cross-reactive TFH cells and antibodies. A deeper understanding
of these aspects of the cellular immune response would help to advance in our knowledge of
immunity against SARS-CoV-
El aprendizaje de historia en contextos bilingües (inglés/español) y no bilingües en la Comunidad de Madrid: oportunidades y desafíos
No full textTesis Doctoral inédita leída en la Universidad Autónoma de Madrid, Facultad de Filosofía y Letras, Departamento de Filología Inglesa. Fecha de Lectura: 03-04-2024Esta tesis tiene embargado el acceso al texto completo hasta el 03-10-2025Bilingual programmes in Spain make the educational scenario where the teaching of school subjects such as history and the teaching of a foreign language converge. This research investigates if the learning of the history subject in bilingual and non-bilingual public secondary schools is different according to the language of instruction (Spanish L1 and English L2). The study is aimed at comparing the outcomes in groups of students from the Madrid Bilingual Schools Network: the Bilingual Section (BS) and the Bilingual Programme (BP), and schools from the Non-Bilingual mainstream. The educational approach that best characterizes the models of bilingual education depicted here is Content and Language Integrated Learning (CLIL), where content and language are addressed in integration, serving as a converging point for several discourses, processes and practices (Nikula et al., 2016).
The present research encompasses the institutional component in the organization of bilingual education in schools as well as the pedagogical component. It studies how students learn historical content, how they incorporate the communicative, cultural and conceptual factors in their written discourse (Coyle, 2007), how they are assessed, and the methodologies and practices developed by teachers in their history lessons. The data consists of 900 tests administered to 450 students from Y1 and Y3ESO (grades 7 and 9), conducted in six public secondary schools in the Madrid region (three with a bilingual programme, three non-bilingual); 450 student questionnaires; and 22 questionnaires to history teachers (11 in bilingual programmes, 11 in non-bilingual). The data consist of the test results, participating students’ history writing, and the students’ and teachers’ responses to questionnaires. The data was collected through tests that were analyzed using rubrics, and questionnaires created ad hoc in an empirical design.
The researcher proposes the H-CLIL model as a suitable tool to facilitate the measurement of history learning in bilingual and non-bilingual contexts, support teachers in choosing the specific text genres and concepts for their lessons and promote assessment through rubrics. The researcher explored the presence of Dalton-Puffer’s (2013) CDFs (Cognitive Discourse Functions) in the teaching and learning process in the classroom. The CDF construct provides teachers with a tangible means to perceive how the curricular content and language can be integrated in the history classroom (Lorenzo, 2017). The articulation of the content in the historical discourse that secondary students are able to produce across the school levels was observed through the identification of the role that the language plays in the classrooms (Llinares et al., 2012). Quantitative and qualitative data were reported to support the effectiveness of the H-CLIL tool of assessment for the application in both bilingual and non-bilingual groups.
Findings show that there was no significant difference between students from bilingual and non-bilingual schools in the content they provided and the history knowledge they demonstrated. There were more differences in terms of the expression of the content. Within the limited scope of this study, it may be stated that students proved a progression in their learning of history with the ability to retrieve content learnt in previous academic years and apply that knowledge to their current learning. While students from the non-bilingual groups presented slightly higher scores in the tests -0.13 points in Y1 and 0.88 points in Y3- but insufficient to deduce statistical significance, students from the Bilingual Section outperformed in the richness of historical lexicon, a higher degree of nominalizations, and the level of abstraction and complexity in their answers to the questions and essay. An evolution was found in all the groups of students between the levels Y1ESO and Y3ESO. Y3 ESO students presented longer paragraphs, more elaborated sentences with time and consequence markers, and an evolution was also observed from mere narrative in Y1 to argumentative writing in Y3 in the historical discours
Lapides loquentes: La Urbs Iulia Gaditana a través de sus inscripciones
No full textTesis Doctoral inédita leída en la Universidad Autónoma de Madrid, Facultad de Filosofía y Letras, Departamento de Filología Clásica. Fecha de Lectura: 18-04-2024Esta tesis tiene embargado el acceso al texto completo hasta el 18-10-2025La presente tesis doctoral versa sobre las aportaciones de la epigrafía latina al estudio de la ciudad romana de Gades. A través del análisis pormenorizado de un conjunto de 489 epígrafes, seleccionados de entre los más de 1000 conocidos para la civitas gaditana, hemos elaborado un retrato político, económico y social basado en los testimonios realizados por y para sus habitantes desde finales de la República y el Principado hasta la Tardoantigüedad. Dicha investigación se recoge en la primera parte de esta tesis, mientras que la segunda está compuesta por el catálogo de inscripciones, donde se incluyen los textos y los comentarios individualizados de la selección de epígrafes antes mencionada. La elección de las inscripciones está basada en la relevancia de la información aportada por las mismas, especialmente aquella relacionada con los habitantes de la ciudad, por lo que se han excluido todos los fragmentos ininteligibles o simplemente formulares.
Gracias a esta investigación hemos podido profundizar en cómo las cualidades de Gades, en calidad de una de las ciudades portuarias más importantes del Mediterráneo, influyeron en su sociedad y en el desarrollo de esta. Encontramos gran variedad de gentilicios reflejados en sus inscripciones —fundamentalmente funerarias— la mayoría de los cuales, sin embargo, se registran en menos de diez ocasiones. Esto indicaría una tendencia constante a la movilidad poblacional, con individuos que, atraídos por las posibilidades económicas de la ciudad, permanecen en ella tan solo una o dos generaciones. En esta misma línea constatamos que, si bien la mayor parte de los personajes mencionados en los epígrafes gaditanos son de origen ingenuo, hay una cantidad notable de cognomina griegos y orientales y de nomina unica, asociados tradicionalmente a liberti y servi respectivamente (ver Cap. 9.3). Aunque no podemos asegurar a ciencia cierta el origen servil de estos individuos, en caso de que así fuera, parece darse una tendencia a ocultar dicho origen, pues las menciones explícitas a relaciones de servidumbre o patronato son llamativamente limitadas.
Llamativa es también la escasez o ausencia de los otros grandes tipos de inscripciones urbanas, como las votivas, las edilicias o las evergéticas. En el primer caso, destaca el hecho de que no se han recogido hasta el momento epígrafes dedicados a las dos principales divinidades gaditanas (véase Cap. 7), Hercules Gaditanus y Venus Marina, cuyo influjo en la ciudad debe ser rastreado epigráficamente a partir de teóforos y de ciertos formularios relacionados con la hierodoulia. Gracias a las pocas inscripciones relacionadas con el gobierno de la ciudad se constatan los privilegios que recibió la civitas de mano de Augusto —ya municipium civium Romanorum desde época cesariana— , y que conllevaron un cambio de magistraturas, la adscripción a la tribu Galeria y la adopción del epíteto Augusta en la nomenclatura urbana (ver Caps. 5.3 y 5.4). A partir de este pequeño conjunto,
además, hemos podido identificar algunas familias que, contrariamente a la dinámica general de la ciudad, no solo perduran en el tiempo, sino que se mantienen en el poder durante generaciones.
Tampoco es posible identificar reminiscencias del pasado púnico de Gades, cuyas inscripciones latinas muestran una sociedad totalmente romana desde finales del siglo I a. C., momento en que se datan sus epígrafes más antiguos. No obstante, la presencia de algunos cognomina de raíz semítica y el empleo frecuente de antropónimos latinos típicos del norte de África en la onomástica gaditana, muestran un continuo y estrecho contacto entre la ciudad y las provincias norteafricanas, tal vez fomentado por ese pasado común.
El estudio individualizado y minucioso de cada uno de los epígrafes seleccionados nos ha permitido, además, corregir lecturas tradicionalmente aceptadas, así como proponer nuevas interpretaciones de algunas inscripciones basándonos en el estudio comparativo de los propios textos gaditanos. Incorporamos además por primera vez al conjunto epigráfico gaditano piezas atribuidas historiográficamente a otras ciudades de la Bética (Corduba, Asido Caesarina), presentando de este modo una investigación acerca de la sociedad de la Urbs Iulia Gaditana lo más amplia y detallada posibl
Motor de aprendizaje máquina para la optimización de correcciones de errores de refracción
No full textTesis Doctoral inédita leída en la Universidad Autónoma de Madrid, Escuela Politécnica Superior, Departamento de Tecnología Electrónica y de las Comunicaciones. Fecha de Lectura: 22-05-2024Esta tesis tiene embargado el acceso al texto completo hasta el 22-11-2025This thesis has been focused on improving access to eyecare through the design and development of portable medical devices that use low cost components, and the implementation of advanced algorithms that allows the detection and diagnosis of different eye diseases.
Uncorrected refractive errors (UREs) are a global issue impacting over 900 million individuals. This issue is particularly pronounced in low resource settings, where access to vision care is limited due to a shortage of eyecare professionals (ECP) capable of providing accurate eyeglass prescriptions. Although autorefraction-based prescriptions are a practical approach to addressing the global problem of UREs, there is always a measurable difference between the autorefractor measurement the clinical gold standard, the subjective refraction (SR). To try to reduce the variability between objective and SR measurements we have assessed the performance of machine learning (ML) ensemble models for predicting patient SR using demographic factors, wavefront aberrometry data, and measurement quality related metrics taken with the QuickSee aberrometer. The ML model significantly outperformed the autorefractor, reducing in a ±0.63 D, ±0.14 D, and ±0.08 D the 95% limits of agreement of the error distribution for M, J0, and J45, respectively. These results suggest that machine learning effectively enhances accuracy when predicting a subjective refraction of a patient from objective measurements acquired with a low-cost portable device.
A novel AI-based approach for the detection of manifest keratoconus from wavefront aberrometry images has been evaluated. The model consists of two components: a first convolutional neural network (CNN) trained to classify individual wavefront maps as either keratoconic positive or keratoconic negative, and a second component that leverages the dynamic nature of the QuickSee device. This second component provides the final classification label for a measurement based on the most frequent prediction among the individual predictions contained in the video. The algorithm demonstrated remarkable performance, achieving a 100% accuracy rate across training, validation, and test subgroups. This represents an advance in keratoconus detection based solely on total wavefront aberrometry data.
We have designed a new portable wavefront autorefractor, the QuickSee Free, that has been evaluated using an alternative study protocol that considers the variability of the SR. The proposed protocol compares the differences between the device and the SR performed by two independent ECPs to the differences between the SR measurements performed by the two ECPs, thus providing a measure of the device’s accuracy against the inherent variability of the SR procedure. The average difference between the device and each of the two SRs were found to be smaller than those found between the two SRs for the M, and comparable for the astigmatic components. Overall, the device had a high level of agreement with both SRs but was found to be more comparable to SR2 than to SR1. These results indicate the potential of the device to be used as a reliable tool for clinical and screening purposes.
In conclusion, this research aims to address the global problem of UREs and presents innovative solutions. These solutions include the design of the QuickSee Free device and various ML algorithms, all of which work towards improving access to eyecare, increasing the accuracy of eyeglass prescriptions, and detecting keratoconu
Relevancia de TAP y del proteasoma en la presentación antigénica durante una infección por citomegalovirus
No full textTesis Doctoral inédita leída en la Universidad Autónoma de Madrid, Facultad de Ciencias, Departamento de Biología Molecular. Fecha de Lectura: 10-05-2024Esta tesis tiene embargado el acceso al texto completo hasta el 10-11-2025CD8+ cytotoxic T lymphocytes (CTLs) recognize peptides presented by major histocompatibility complex (MCH) class I molecules at the surface of altered cells and selectively detect and kill them. These peptides are mainly generated and presented via the classical antigen processing and presentation pathway. This pathway typically involves protein processing by the proteasome in the cytosol and the transport of processed peptides into the endoplasmic reticulum (ER) by the transporter associated with antigen processing (TAP) to then be loaded onto MHC class I molecules and be finally presented at the cell surface. However, several reports have shown that certain peptides can still be presented when one or both elements are absent or inhibited. Studying these alternative antigen presentation pathways is highly relevant to the development of new vaccination strategies, as certain viruses and cancer cells have developed mechanisms that disrupt this pathway and thus, perturb the immune control. The aim of the present work is to study antigen presentation in the absence of TAP and when proteasome function is impaired during infection with murine cytomegalovirus (MCMV).
First, we studied the relevance of TAP in the control of MCMV infection in mice. We observed that TAP-deficient mice were able to efficiently control MCMV infection, although with a slight delay. Moreover, MCMV latency establishment and generation of MCMV neutralizing antibodies were not affected by TAP absence. Lastly, we characterized splenic NK cells derived from TAP-deficient mice during MCMV infection.
Secondly, we studied TAP-independent antigen presentation to CD8+ T lymphocytes during MCMV infection in vitro and in vivo. We demonstrated that TAP-deficient cells were capable of presenting MCMV peptides in vitro. Even though TAP-independent presentation was very limited, we were able to identify an epitope from the m164 ORF of MCMV that could be presented in the absence of TAP. Unfortunately, we could not demonstrate the presentation of the m164 epitope in vivo, as we were unable to induce CD8+ T lymphocyte responses against this epitope in TAP-deficient mice by several different approaches.
Finally, we investigated MCMV antigen presentation to CD8+ T lymphocytes when proteasome function was inhibited. We found out that around half of the TAP-dependent MCMV antigen presentation could still take place under the inhibition of the proteasome. Furthermore, proteasome inhibition affected the presentation of MCMV epitopes differently. While presentation of some epitopes was completely abrogated in the absence of proteasome function, presentation of others was even increased when the proteasome was inhibited.
Remarkably, we demonstrated that TAP-independent presentation of the m164 epitope required proteasomal activity.
Thus, we demonstrated in vitro that MCMV antigen presentation can still take place either when TAP is absent or when the proteasome is inhibited, and we identified a TAP-independent MCMV epitope that required proteasomal processing to be presented. However, we were unable to demonstrate the relevance of this TAP-independent antigen presentation in the control of MCMV infection in viv