PMC:7275137 / 2784-8488
Annnotations
{"target":"https://pubannotation.org/docs/sourcedb/PMC/sourceid/7275137","sourcedb":"PMC","sourceid":"7275137","source_url":"https://www.ncbi.nlm.nih.gov/pmc/7275137","text":"1 Introduction\nThe main objective of all viruses is to hijack the intracellular machinery of the host cell for genome replication and assembly. To achieve this, the virus first has to bypass the cellular membrane barrier. The structure of some viruses enables direct fusion at the plasma membrane after recognition of the binding site [1]. However, most enveloped viruses depend on the endocytic pathway prior to fusion with the host cell [2]. Once endocytosis has been initiated, survival and replication of the virus depend on successful delivery of the viral nucleocapsid into the correct intracellular department, as different viral families target distinct compartments as primary replication sites [3]. Moreover, in order to survive, the virus has to escape from the endosomal compartment silently, without triggering the cellular defence mechanisms [4]. As a result, viral evolution involves highly complex escape strategies relying on different cues of endosomal maturation and trafficking [5]. Establishment of an acidic endosomal lumen is the key requirement for normal functioning of the endocytic pathway [6], so pH sensing has evolved as one of the fundamental strategies used by viruses to monitor endosomal maturation [7]. Two important sensing strategies are used by most viruses for induction of membrane fusion mechanisms: direct pH sensing mediated by a conformational change of the envelope glycoproteins, and endosomal protease-mediated proteolytic cleavage [7]. From a pharmacological perspective, this makes endosomal pH an attractive drug target. Neutralization of endosomal pH obstructs endosomal trafficking, impedes sorting mechanisms, and inhibits activity of the endosomal proteases [6]. As a consequence, pharmacological modulation of endolysosomal pH could prevent viral fusion and impair the formation of viral replication complexes. Moreover, as pH modulation is important for regulation of exocytosis, vesicular pH modulation could also affect viral dissemination [8]. Endosomal trafficking is necessary for normal cellular functioning, so long-term disruption of endosomal function could have considerable pathophysiological consequences [9]. Moreover, the infectivity of some viruses that use alternative strategies to enter the cell could be increased by pH-mediated inhibition of the endosomal pathway [10]. For this reason, most antiviral drug development strategies have focused on more specific fusion mechanisms rather than general inhibition of endocytosis. However, short-term inhibition of endocytosis could be a viable strategy for the management of viral infections with no alternative treatment options, such as coronavirus disease 2019 (COVID-19). Recent evidence suggests that chloroquine (CQ) and hydroxychloroquine (HCQ) may be effective therapeutic strategies for COVID-19 [11], [12], [13], [14]. Although the exact mechanism of action responsible for the antiviral activity of CQ and HCQ remains unclear, the elevation of lysosomal pH seems to play a key role [14]. This is in concordance with previous findings, as proteolytic processing of glycoprotein S by lysosomal proteases seems to be necessary for the fusion of coronavirus with the host cell [2]. Moreover, treatment of cells with glycoprotein S – the main protein responsible for viral fusion of severe acute respiratory syndrome coronavirus (SARS-CoV) and severe acute respiratory syndrome coronavirus-2 (SARS-CoV-2) – resulted in translocation of the main viral receptor, angiotensin-converting enzyme 2 (ACE-2), to endosomal vesicles [15]. Furthermore, the same effect was demonstrated for spike-bearing pseudoviruses, emphasizing the importance of glycoprotein S for viral fusion [15]. It is important to mention that although many groups have demonstrated that SARS-CoV enters the cells by receptor-mediated, pH-sensitive endocytosis [15], some groups have reported that viral fusion might also occur by pH-independent direct fusion with the plasma membrane [16]. Although the exact mechanism responsible for the entrance of SARS-CoV-2 into the host cell remains to be elucidated, the authors believe that data from clinical trials on CQ and HCQ support the importance of pH-dependent endocytosis, and that other widely available pharmacological agents with lysosomotropic properties may also demonstrate clinical effectiveness in the treatment of COVID-19. This review discusses commonly used pharmacological agents in the context of their lysosomotropic effects and pH-modulation-mediated interaction with cytoplasmic vesicle trafficking (Fig. 1 .). Although almost 5 million people had been diagnosed with COVID-19 at the time of writing (20th May 2020), organized clinical data that would facilitate the evaluation of potential antiviral effects of the drugs discussed below on SARS-CoV-2 are unavailable at present. Colleagues in possession of such data are urged to evaluate the possible benefits of these drugs, as they might offer new hope in terms of prevention and/or treatment of this devastating disease.\nFig. 1 Possible interaction of lysosomotropic agents and lysosome targeting drugs with the replication cycle of severe acute respiratory syndrome corovavirus-2 (SARS-CoV-2) in a cell. Lysosome targeting drugs and lysosomotropic drugs, such as chloroquine, have the ability to interfere with endosomal trafficking, affecting both endosomal maturation and exocytotic pathways. Taking into consideration the dependence of viral replication on the normal functioning of endocytic trafficking, commonly used pharmacological agents that demonstrate lysosomotropic properties should be examined as a possible therapeutic option for the treatment of coronavirus disease 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