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Article: The FDA-approved drug nitazoxanide is a potent inhibitor of human seasonal coronaviruses acting at postentry level: effect on the viral spike glycoprotein.

Piacentini, Sara / Riccio, Anna / Santopolo, Silvia / Pauciullo, Silvia / La Frazia, Simone / Rossi, Antonio / Rossignol, Jean-Francois / Santoro, M Gabriella

Frontiers in microbiology

2023 Volume 14, Page(s) 1206951

Abstract: ... ...

Abstract	Coronaviridae
Language	English
Publishing date	2023-08-29
Publishing country	Switzerland
Document type	Journal Article
ZDB-ID	2587354-4
ISSN	1664-302X
ISSN	1664-302X
DOI	10.3389/fmicb.2023.1206951
Database	MEDical Literature Analysis and Retrieval System OnLINE

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Article ; Online: The biogenesis of SARS-CoV-2 spike glycoprotein: multiple targets for host-directed antiviral therapy.

Santopolo, Silvia / Riccio, Anna / Santoro, M Gabriella

Biochemical and biophysical research communications

2020 Volume 538, Page(s) 80–87

Abstract: Severe acute respiratory syndrome coronavirus 2 (SARS-CoV-2), the causative agent of COVID-19 (coronavirus disease-19), represents a far more serious threat to public health than SARS and MERS coronaviruses, due to its ability to spread more efficiently ... ...

Abstract	Severe acute respiratory syndrome coronavirus 2 (SARS-CoV-2), the causative agent of COVID-19 (coronavirus disease-19), represents a far more serious threat to public health than SARS and MERS coronaviruses, due to its ability to spread more efficiently than its predecessors. Currently, there is no worldwide-approved effective treatment for COVID-19, urging the scientific community to intense efforts to accelerate the discovery and development of prophylactic and therapeutic solutions against SARS-CoV-2 infection. In particular, effective antiviral drugs are urgently needed. With few exceptions, therapeutic approaches to combat viral infections have traditionally focused on targeting unique viral components or enzymes; however, it has now become evident that this strategy often fails due to the rapid emergence of drug-resistant viruses. Targeting host factors that are essential for the virus life cycle, but are dispensable for the host, has recently received increasing attention. The spike glycoprotein, a component of the viral envelope that decorates the virion surface as a distinctive crown ("corona") and is essential for SARS-CoV-2 entry into host cells, represents a key target for developing therapeutics capable of blocking virus invasion. This review highlights aspects of the SARS-CoV-2 spike biogenesis that may be amenable to host-directed antiviral targeting.
MeSH term(s)	Antiviral Agents/pharmacology ; Antiviral Agents/therapeutic use ; COVID-19/drug therapy ; COVID-19/virology ; Glycosylation ; Humans ; Molecular Targeted Therapy ; Protein Folding ; SARS-CoV-2/drug effects ; SARS-CoV-2/physiology ; Spike Glycoprotein, Coronavirus/antagonists & inhibitors ; Spike Glycoprotein, Coronavirus/biosynthesis ; Spike Glycoprotein, Coronavirus/chemistry ; Virus Internalization/drug effects
Chemical Substances	Antiviral Agents ; Spike Glycoprotein, Coronavirus ; spike protein, SARS-CoV-2
Language	English
Publishing date	2020-11-28
Publishing country	United States
Document type	Journal Article ; Research Support, Non-U.S. Gov't ; Review
ZDB-ID	205723-2
ISSN	1090-2104 ; 0006-291X ; 0006-291X
ISSN (online)	1090-2104 ; 0006-291X
ISSN	0006-291X
DOI	10.1016/j.bbrc.2020.10.080
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Article ; Online: Impairment of SARS-CoV-2 spike glycoprotein maturation and fusion activity by nitazoxanide: an effect independent of spike variants emergence.

Riccio, Anna / Santopolo, Silvia / Rossi, Antonio / Piacentini, Sara / Rossignol, Jean-Francois / Santoro, M Gabriella

Cellular and molecular life sciences : CMLS

2022 Volume 79, Issue 5, Page(s) 227

Abstract: SARS-CoV-2, the causative agent of COVID-19, has caused an unprecedented global health crisis. The SARS-CoV-2 spike, a surface-anchored trimeric class-I fusion glycoprotein essential for viral entry, represents a key target for developing vaccines and ... ...

Abstract	SARS-CoV-2, the causative agent of COVID-19, has caused an unprecedented global health crisis. The SARS-CoV-2 spike, a surface-anchored trimeric class-I fusion glycoprotein essential for viral entry, represents a key target for developing vaccines and therapeutics capable of blocking virus invasion. The emergence of SARS-CoV-2 spike variants that facilitate virus spread and may affect vaccine efficacy highlights the need to identify novel antiviral strategies for COVID-19 therapy. Here, we demonstrate that nitazoxanide, an antiprotozoal agent with recognized broad-spectrum antiviral activity, interferes with SARS-CoV-2 spike maturation, hampering its terminal glycosylation at an endoglycosidase H-sensitive stage. Engineering multiple SARS-CoV-2 variant-pseudoviruses and utilizing quantitative cell-cell fusion assays, we show that nitazoxanide-induced spike modifications hinder progeny virion infectivity as well as spike-driven pulmonary cell-cell fusion, a critical feature of COVID-19 pathology. Nitazoxanide, being equally effective against the ancestral SARS-CoV-2 Wuhan-spike and different emerging variants, including the Delta variant of concern, may represent a useful tool in the fight against COVID-19 infections.
MeSH term(s)	Antiviral Agents/pharmacology ; COVID-19/drug therapy ; Humans ; Nitro Compounds/pharmacology ; SARS-CoV-2/drug effects ; Spike Glycoprotein, Coronavirus/antagonists & inhibitors ; Thiazoles/pharmacology
Chemical Substances	Antiviral Agents ; Nitro Compounds ; Spike Glycoprotein, Coronavirus ; Thiazoles ; spike protein, SARS-CoV-2 ; nitazoxanide (SOA12P041N)
Language	English
Publishing date	2022-04-07
Publishing country	Switzerland
Document type	Journal Article
ZDB-ID	1358415-7
ISSN	1420-9071 ; 1420-682X
ISSN (online)	1420-9071
ISSN	1420-682X
DOI	10.1007/s00018-022-04246-w
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Article ; Online: MicroRNA analysis of Natural Killer cell-derived exosomes: the microRNA let-7b-5p is enriched in exosomes and participates in their anti-tumor effects against pancreatic cancer cells.

Di Pace, Anna Laura / Pelosi, Andrea / Fiore, Piera Filomena / Tumino, Nicola / Besi, Francesca / Quatrini, Linda / Santopolo, Silvia / Vacca, Paola / Moretta, Lorenzo

Oncoimmunology

2023 Volume 12, Issue 1, Page(s) 2221081

Abstract: Natural Killer (NK) cells are important components of the immune system in the defense against tumor growth and metastasis. They release exosomes containing proteins and nucleic acids, including microRNAs (miRNAs). NK-derived exosomes play a role in the ... ...

Abstract	Natural Killer (NK) cells are important components of the immune system in the defense against tumor growth and metastasis. They release exosomes containing proteins and nucleic acids, including microRNAs (miRNAs). NK-derived exosomes play a role in the anti-tumor NK cell function since they are able to recognize and kill cancer cells. However, the involvement of exosomal miRNAs in the function of NK exosomes is poorly understood. In this study, we explored the miRNA content of NK exosomes by microarray as compared to their cellular counterparts. The expression of selected miRNAs and lytic potential of NK exosomes against childhood B acute lymphoblastic leukemia cells after co-cultures with pancreatic cancer cells were also evaluated. We identified a small subset of miRNAs, including miR-16-5p, miR-342-3p, miR-24-3p, miR-92a-3p and let-7b-5p that is highly expressed in NK exosomes. Moreover, we provide evidence that NK exosomes efficiently increase let-7b-5p expression in pancreatic cancer cells and induce inhibition of cell proliferation by targeting the cell cycle regulator CDK6. Let-7b-5p transfer by NK exosomes could represent a novel mechanism by which NK cells counteract tumor growth. However, both cytolytic activity and miRNA content of NK exosomes were reduced upon co-culture with pancreatic cancer cells. Alteration in the miRNA cargo of NK exosomes, together with their reduced cytotoxic activity, could represent another strategy exerted by cancer to evade the immune response. Our study provides new information on the molecular mechanisms used by NK exosomes to exert anti-tumor-activity and offers new clues to integrate cancer treatments with NK exosomes.
MeSH term(s)	Humans ; Child ; Exosomes/genetics ; MicroRNAs/genetics ; Pancreatic Neoplasms/genetics ; Pancreatic Neoplasms/therapy ; Killer Cells, Natural ; Pancreatic Neoplasms
Chemical Substances	MicroRNAs
Language	English
Publishing date	2023-06-07
Publishing country	United States
Document type	Journal Article ; Research Support, Non-U.S. Gov't
ZDB-ID	2645309-5
ISSN	2162-402X ; 2162-402X
ISSN (online)	2162-402X
ISSN	2162-402X
DOI	10.1080/2162402X.2023.2221081
Database	MEDical Literature Analysis and Retrieval System OnLINE

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Article ; Online: The proteostasis guardian HSF1 directs the transcription of its paralog and interactor HSF2 during proteasome dysfunction.

Santopolo, Silvia / Riccio, Anna / Rossi, Antonio / Santoro, M Gabriella

Cellular and molecular life sciences : CMLS

2020 Volume 78, Issue 3, Page(s) 1113–1129

Abstract: Protein homeostasis is essential for life in eukaryotes. Organisms respond to proteotoxic stress by activating heat shock transcription factors (HSFs), which play important roles in cytoprotection, longevity and development. Of six human HSFs, HSF1 acts ... ...

Abstract	Protein homeostasis is essential for life in eukaryotes. Organisms respond to proteotoxic stress by activating heat shock transcription factors (HSFs), which play important roles in cytoprotection, longevity and development. Of six human HSFs, HSF1 acts as a proteostasis guardian regulating stress-induced transcriptional responses, whereas HSF2 has a critical role in development, in particular of brain and reproductive organs. Unlike HSF1, that is a stable protein constitutively expressed, HSF2 is a labile protein and its expression varies in different tissues; however, the mechanisms regulating HSF2 expression remain poorly understood. Herein we demonstrate that the proteasome inhibitor anticancer drug bortezomib (Velcade), at clinically relevant concentrations, triggers de novo HSF2 mRNA transcription in different types of cancers via HSF1 activation. Similar results were obtained with next-generation proteasome inhibitors ixazomib and carfilzomib, indicating that induction of HSF2 expression is a general response to proteasome dysfunction. HSF2-promoter analysis, electrophoretic mobility shift assays, and chromatin immunoprecipitation studies unexpectedly revealed that HSF1 is recruited to a heat shock element located at 1.397 bp upstream from the transcription start site in the HSF2-promoter. More importantly, we found that HSF1 is critical for HSF2 gene transcription during proteasome dysfunction, representing an interesting example of transcription factor involved in controlling the expression of members of the same family. Moreover, bortezomib-induced HSF2 was found to localize in the nucleus, interact with HSF1, and participate in bortezomib-mediated control of cancer cell migration. The results shed light on HSF2-expression regulation, revealing a novel level of HSF1/HSF2 interplay that may lead to advances in pharmacological modulation of these fundamental transcription factors.
MeSH term(s)	Boron Compounds/chemistry ; Boron Compounds/metabolism ; Bortezomib/chemistry ; Bortezomib/metabolism ; Bortezomib/pharmacology ; Cell Line, Tumor ; Cell Movement/drug effects ; Cell Nucleus/metabolism ; Electrophoretic Mobility Shift Assay ; Glycine/analogs & derivatives ; Glycine/chemistry ; Glycine/metabolism ; Heat Shock Transcription Factors/antagonists & inhibitors ; Heat Shock Transcription Factors/genetics ; Heat Shock Transcription Factors/metabolism ; Heat-Shock Proteins/antagonists & inhibitors ; Heat-Shock Proteins/genetics ; Heat-Shock Proteins/metabolism ; Humans ; Promoter Regions, Genetic ; Proteasome Endopeptidase Complex/chemistry ; Proteasome Endopeptidase Complex/metabolism ; Proteasome Inhibitors/chemistry ; Proteasome Inhibitors/metabolism ; Proteasome Inhibitors/pharmacology ; RNA Interference ; RNA, Messenger/metabolism ; RNA, Small Interfering/metabolism ; Transcription Factors/antagonists & inhibitors ; Transcription Factors/genetics ; Transcription Factors/metabolism ; Transcription Initiation Site ; Transcription, Genetic
Chemical Substances	Boron Compounds ; HSF1 protein, human ; Heat Shock Transcription Factors ; Heat-Shock Proteins ; Proteasome Inhibitors ; RNA, Messenger ; RNA, Small Interfering ; Transcription Factors ; HSF2 protein, human (142297-91-4) ; Bortezomib (69G8BD63PP) ; ixazomib (71050168A2) ; Proteasome Endopeptidase Complex (EC 3.4.25.1) ; Glycine (TE7660XO1C)
Language	English
Publishing date	2020-06-30
Publishing country	Switzerland
Document type	Journal Article
ZDB-ID	1358415-7
ISSN	1420-9071 ; 1420-682X
ISSN (online)	1420-9071
ISSN	1420-682X
DOI	10.1007/s00018-020-03568-x
Database	MEDical Literature Analysis and Retrieval System OnLINE

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Zs.A 195: Show issues

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ab Jg. 2022: Lesesaal (EG)

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Article ; Online: Human coronaviruses activate and hijack the proteostasis guardian HSF1 to enhance viral replication

Pauciullo, Silvia / Riccio, Anna / Rossi, Antonio / Santopolo, Silvia / Piacentini, Sara / Santoro, Maria Gabriella

bioRxiv

Abstract: Organisms respond to proteotoxic stress by activating a cellular defense mechanism, known as the heat shock response (HSR), that triggers the expression of cytoprotective heat shock proteins (HSP) to counteract the damaging effects of proteostasis ... ...

Abstract	Organisms respond to proteotoxic stress by activating a cellular defense mechanism, known as the heat shock response (HSR), that triggers the expression of cytoprotective heat shock proteins (HSP) to counteract the damaging effects of proteostasis disruption. The HSR is regulated by a family of transcription factors (heat shock factors, HSFs); among six human HSFs, HSF1 acts as a proteostasis guardian regulating acute and severe stress-driven transcriptional responses. Seasonal coronaviruses HCoV-229E, HCoV-NL63, HCoV-OC43 and HCoV-HKU1 (sHCoV) are globally circulating in the human population. Although sHCoV generally cause only mild upper respiratory diseases in immunocompetent hosts, severe complications may occur in specific populations. There is no effective treatment for sHCoV infections, also due to the limited knowledge on sHCoV biology. We now show that both Alpha- and Beta- sHCoV are potent inducers of HSF1, selectively promoting HSF1 phosphorylation at serine-326 residue and nuclear translocation, and triggering a powerful HSF1-driven transcriptional response in infected cells at late stages of infection. Despite the coronavirus-mediated shut-down of the host cell translational machinery, high levels of selected canonical and non-canonical HSF1-target genes products, including HSP70, HSPA6 and the zinc-finger AN1-type domain-2a gene/AIRAP, were found in HCoV-infected cells. Interestingly, silencing experiments demonstrate that HSR activation does not merely reflect a cellular defense response to viral infection, but that sHCoV activate and hijack the HSF1-pathway for their own gain. Notably, nuclear HSF1 pools depletion via Direct-Targeted HSF1 inhibitor (DTHIB) treatment was highly effective in hindering sHCoV replication in lung cells. Altogether the results open new scenarios for the search of innovative antiviral strategies in the treatment of coronavirus infections.
Keywords	covid19
Language	English
Publishing date	2022-12-22
Publisher	Cold Spring Harbor Laboratory
Document type	Article ; Online
DOI	10.1101/2022.12.22.519205
Database	COVID19

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Article: Impairment of SARS-CoV-2 spike glycoprotein maturation and fusion activity by nitazoxanide: an effect independent of spike variants emergence

Riccio, Anna / Santopolo, Silvia / Rossi, Antonio / Piacentini, Sara / Rossignol, Jean-Francois / Santoro, M. Gabriella

Cellular and molecular life sciences. 2022 May, v. 79, no. 5

2022

Abstract	SARS-CoV-2, the causative agent of COVID-19, has caused an unprecedented global health crisis. The SARS-CoV-2 spike, a surface-anchored trimeric class-I fusion glycoprotein essential for viral entry, represents a key target for developing vaccines and therapeutics capable of blocking virus invasion. The emergence of SARS-CoV-2 spike variants that facilitate virus spread and may affect vaccine efficacy highlights the need to identify novel antiviral strategies for COVID-19 therapy. Here, we demonstrate that nitazoxanide, an antiprotozoal agent with recognized broad-spectrum antiviral activity, interferes with SARS-CoV-2 spike maturation, hampering its terminal glycosylation at an endoglycosidase H-sensitive stage. Engineering multiple SARS-CoV-2 variant-pseudoviruses and utilizing quantitative cell–cell fusion assays, we show that nitazoxanide-induced spike modifications hinder progeny virion infectivity as well as spike-driven pulmonary cell–cell fusion, a critical feature of COVID-19 pathology. Nitazoxanide, being equally effective against the ancestral SARS-CoV-2 Wuhan-spike and different emerging variants, including the Delta variant of concern, may represent a useful tool in the fight against COVID-19 infections.
Keywords	COVID-19 infection ; Severe acute respiratory syndrome coronavirus 2 ; antiviral properties ; etiological agents ; glycoproteins ; glycosylation ; pathogenicity ; progeny ; therapeutics ; vaccines ; virion ; viruses
Language	English
Dates of publication	2022-05
Size	p. 227.
Publishing place	Springer International Publishing
Document type	Article
ZDB-ID	1358415-7
ISSN	1420-9071 ; 1420-682X
ISSN (online)	1420-9071
ISSN	1420-682X
DOI	10.1007/s00018-022-04246-w
Database	NAL-Catalogue (AGRICOLA)

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Article ; Online: Nitazoxanide is a potent inhibitor of human seasonal coronaviruses acting at postentry level: effect on viral spike glycoprotein

Piacentini, Sara / Riccio, Anna / Santopolo, Silvia / Pauciullo, Silvia / Rossi, Antonio / Rossignol, Jean-Francois / Santoro, M. Gabriella

bioRxiv

Abstract: Coronaviridae is recognized as one of the most rapidly evolving virus family as a consequence of the high genomic nucleotide substitution rates and recombination. The family comprises a large number of enveloped, positive-sense single-stranded RNA ... ...

Abstract	Coronaviridae is recognized as one of the most rapidly evolving virus family as a consequence of the high genomic nucleotide substitution rates and recombination. The family comprises a large number of enveloped, positive-sense single-stranded RNA viruses, causing an array of diseases of varying severity in animals and humans. To date, seven human coronaviruses (HCoV) have been identified, namely HCoV-229E, HCoV-NL63, HCoV-OC43 and HCoV-HKU1, which are globally circulating in the human population (seasonal HCoV, sHCoV), and the highly pathogenic SARS-CoV, MERS-CoV and SARS-CoV-2. Seasonal HCoV are estimated to contribute to 15-30% of common cold cases in humans; although diseases are generally self-limiting, sHCoV can sometimes cause severe lower respiratory infections, as well as enteric and neurological diseases. No specific treatment is presently available for sHCoV infections. Herein we show that the anti-infective drug nitazoxanide has a potent antiviral activity against three human endemic coronaviruses, the Alpha-coronaviruses HCoV-229E and HCoV-NL63, and the Beta-coronavirus HCoV-OC43 in cell culture with IC50 ranging between 0.05 and 0.15 μg/ml, and high selectivity indexes. We found that nitazoxanide does not affect HCoV adsorption, entry or uncoating, but acts at postentry level and interferes with the spike glycoprotein maturation, hampering its terminal glycosylation at an endoglycosidase H-sensitive stage. Altogether the results indicate that nitazoxanide, due to its broad-spectrum anti-coronavirus activity, may represent a readily available useful tool in the treatment of seasonal coronavirus infections.
Keywords	covid19
Language	English
Publishing date	2022-07-13
Publisher	Cold Spring Harbor Laboratory
Document type	Article ; Online
DOI	10.1101/2022.07.13.499346
Database	COVID19

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Article ; Online: Impairment of SARS-CoV-2 spike glycoprotein maturation and fusion activity by the broad-spectrum anti-infective drug nitazoxanide

Riccio, Anna / Santopolo, Silvia / Rossi, Antonio / Piacentini, Sara / Rossignol, Jean-Francois / Santoro, Maria Gabriella

bioRxiv

Abstract: The emergence of the highly-pathogenic severe acute respiratory syndrome coronavirus-2 (SARS-CoV-2), the causative agent of COVID-19 (coronavirus disease-2019), has caused an unprecedented global health crisis, as well as societal and economic disruption. ...

Abstract	The emergence of the highly-pathogenic severe acute respiratory syndrome coronavirus-2 (SARS-CoV-2), the causative agent of COVID-19 (coronavirus disease-2019), has caused an unprecedented global health crisis, as well as societal and economic disruption. The SARS-CoV-2 spike (S), a surface-anchored trimeric class-I fusion glycoprotein essential for entry into host cells, represents a key target for developing vaccines and therapeutics capable of blocking virus invasion. The emergence of several SARS-CoV-2 spike variants that facilitate virus spread and may affect the efficacy of recently developed vaccines, creates great concern and highlights the importance of identifying antiviral drugs to reduce SARS-CoV-2-related morbidity and mortality. Nitazoxanide, a thiazolide originally developed as an antiprotozoal agent with recognized broad-spectrum antiviral activity in-vitro and in clinical studies, was recently shown to be effective against several coronaviruses, including SARS-CoV-2. Using biochemical and pseudovirus entry assays, we now demonstrate that nitazoxanide interferes with the SARS-CoV-2 spike biogenesis, hampering its maturation at an endoglycosidase H-sensitive stage, and hindering its fusion activity in human cells. Besides membrane fusion during virus entry, SARS-CoV-2 S-proteins in infected cells can also trigger receptor-dependent formation of syncytia, observed in-vitro and in COVID-19 patients tissues, facilitating viral dissemination between cells and possibly promoting immune evasion. Utilizing two different quantitative cell-cell fusion assays, we show that nitazoxanide is effective in inhibiting syncytia formation mediated by different SARS-CoV-2 spike variants in human lung, liver and intestinal cells. The results suggest that nitazoxanide may represent a useful tool in the fight against COVID-19 infections, inhibiting SARS-CoV-2 replication and preventing spike-mediated syncytia formation.
Keywords	covid19
Language	English
Publishing date	2021-04-12
Publisher	Cold Spring Harbor Laboratory
Document type	Article ; Online
DOI	10.1101/2021.04.12.439201
Database	COVID19

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Article ; Online: The roles of different forms of IL-15 in human melanoma progression.

Frontiers in immunology

2023 Volume 14, Page(s) 1183668

Abstract: Background: Melanoma is a lethal skin cancer, and the risk of developing it is increased by exposure to ultraviolet (UV) radiation. The production of cytokines such as interleukin-15 (IL-15), induced by the exposure of skin cells to UV rays, could also ... ...

Abstract	Background: Melanoma is a lethal skin cancer, and the risk of developing it is increased by exposure to ultraviolet (UV) radiation. The production of cytokines such as interleukin-15 (IL-15), induced by the exposure of skin cells to UV rays, could also promote melanoma development. The aim of this study is to investigate the possible role of Interleukin-15/Interleukin-15 Receptor α (IL-15/IL-15Rα) complexes in melanoma development. Methods: The expression of IL-15/IL-15Rα complexes by melanoma cells was evaluated both Results: Analysis of a melanoma tissue microarray shows a significant increase in the number of IL-15 Conclusions: Membrane-bound and secreted IL-15/IL-15Rα complexes are continuously present during progression in melanoma. It is notable that, although IL-15/IL-15Rα initially promoted the production of cytotoxic T and NK cells, at stage IV promotion of the development of anergic and dysfunctional cytotoxic NK cells was observed. In a subgroup of melanoma metastatic patients, the continuous secretion of high amounts of the soluble complex could represent a novel NK cell immune escape mechanism.
MeSH term(s)	Humans ; Antineoplastic Agents ; Cell Line, Tumor ; Interleukin-15/metabolism ; Interleukin-15 Receptor alpha Subunit/genetics ; Interleukin-15 Receptor alpha Subunit/metabolism ; Killer Cells, Natural ; Melanoma/metabolism
Chemical Substances	Antineoplastic Agents ; Interleukin-15 ; Interleukin-15 Receptor alpha Subunit ; IL15 protein, human
Language	English
Publishing date	2023-06-02
Publishing country	Switzerland
Document type	Journal Article ; Research Support, Non-U.S. Gov't
ZDB-ID	2606827-8
ISSN	1664-3224 ; 1664-3224
ISSN (online)	1664-3224
ISSN	1664-3224
DOI	10.3389/fimmu.2023.1183668
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