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International Journal of Bioprinting                                 Coronavirus-infected bioprinted intestine




            also provide protection against intestinal infection and   cell types.  Moreover, under certain culture conditions,
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            potential fecal shedding. This valuable knowledge can   intestinal organoids are capable of virtually indefinite
            guide vaccine development and strategies to improve   propagation. 16,17  Additionally, these organoids recapitulate
            overall vaccine efficacy.                          the in vivo tissue architecture, displaying multiple cell type
                                                               heterogeneity and interactions in an in vitro setting (Figure
            3. Utilization of intestinal organoid in           2A).  When fully developed, human intestinal organoids
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            studying human coronavirus infection               can form crypt and villi domains that house intestinal stem
                                                               cells, progenitors, and differentiated enterocytes.  The
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            3.1. Intestinal organoids                          high-throughput scalability  and physiological relevance
            Intestinal organoids, which are 3D structures derived from   of intestinal organoids make them valuable tools for
            human intestinal stem cells, provide a physiologically-  studying disease phenotypes and screening potential
            relevant representation of the intestinal epithelium.   drug candidates. 14,19,20
            These organoids were among the first successfully
            established, and their development has paved the way for   Given their advantages, intestinal organoids have been
            the establishment of protocols for other organoid types   extensively utilized to model pathogen–host interactions
            and yielded remarkable findings in the field of intestinal   in the human intestine with various types of pathogens. 21,22
                   14
            research.   One  of  the  key  advantages  of  intestinal   For example, Heo et al. established an infection model
            organoids is their composition, which includes multi-  of  the  protozoan  parasite  Cryptosporidium  using  human
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            potent tissue-specific stem cells and differentiated daughter   small intestinal organoids.  Bartfeld et al. successfully








































            Figure 2. Employing intestinal organoids to investigate HCoV infection in the intestinal context. (A) Well-developed intestinal organoids mirror the
            complexity of in vivo intestinal structures, encompassing distinct cell types and architectural features such as crypt and villi domains (created with
            Biorender.com). (B) The versatility of organoid systems facilitates the exploration of pathogen–host dynamics using techniques like microinjection, apical-
            out organoid assays, and organoid dissociation approaches (created with Biorender.com). Studies employing intestinal organoids have (C) elucidated the
            role of TMPRSS2 as a co-factor in SARS-CoV-2 infection (reprinted under the terms of the Creative Commons CC-BY license),  (D) probed the impact
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            of interferons on restraining SARS-CoV-2 infection (reprinted under the terms of the Creative Commons CC-BY-NC-ND license),  and (E) unveiled
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            inflammatory responses triggered by SARS-CoV-2 infection (licensed under a Creative Commons Attribution 4.0 International (CC BY 4.0) license and
            reprinted with permission granted by American Association for the Advancement of Science). 30
            Volume 10 Issue 2 (2024)                       168                                doi: 10.36922/ijb.1704
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