PMC:7463108 / 67443-72854 JSONTXT

{"target":"https://pubannotation.org/docs/sourcedb/PMC/sourceid/7463108","sourcedb":"PMC","sourceid":"7463108","source_url":"https://www.ncbi.nlm.nih.gov/pmc/7463108","text":"Neural Progenitors as an HIV Reservoir and Target for Opioids\nEven though neural progenitors (Krathwohl and Kaiser 2004; Lawrence et al. 2004; Rothenaigner et al. 2007; Schwartz et al. 2007; Balinang et al. 2017), neuroblast cell lines (Ensoli et al. 1994; Rothenaigner et al. 2007), and/or immature astroglia (Atwood et al. 1993; Tornatore et al. 1994; Barat et al. 2018) can harbor HIV infection (reviewed by Hauser and Knapp 2014; Putatunda et al. 2019), the degree to which they are a source of active infection or serve as a latent viral reservoir (Blankson et al. 2002; Bruner et al. 2019) by retaining intact proviral DNA within incipient macroglial progeny is uncertain. In fact, spurious reports of HIV-infected adult neurons (Torres-Munoz et al. 2001; Canto-Nogues et al. 2005) may result from the retention of proviral genes that integrated into pluripotent neural progenitors or neuroblasts at earlier stages during maturation. Importantly, prolonged exposure to opioids can increase the production of HIV in human neural progenitor cells (hNPCs). Exposure of R5-tropic HIVBaL-infected hNPCs to morphine continuously for 21 d increased viral production compared to HIVBaL infection alone in vitro (Balinang et al. 2017).\nBesides being able to infect hNPCs, HIV may also affect their maturation and the fate of neural stem cells. That HIV or gp120 can inhibit adult neurogenesis (Okamoto et al. 2007; Lee et al. 2013; Putatunda et al. 2018) has been the topic of past reviews (Schwartz and Major 2006; Venkatesan et al. 2007; Peng et al. 2008, 2011; Ferrell and Giunta 2014; Hauser and Knapp 2014; Putatunda et al. 2019). How HIV inhibits the self-renewal, tripotential differentiation, and survival of neural progenitors/stem cells or the genesis of adult neurons in the subgranular zone (SGZ) of the dentate gyrus is uncertain. HIV and gp120 [via actions at the same chemokine receptor(s) (Tran and Miller 2005; Li and Ransohoff 2008)] are proposed to modulate the adult neurogenesis via Notch (Fan et al. 2016), by obstructing a cell cycle checkpoint via the activation MAPK-activated protein kinase 2 and Cdc25B/C (Okamoto et al. 2007), or through signaling by platelet-derived growth factor BB (Chao et al. 2014) or BDNF (Lee et al. 2013). The extent that HIV regulates the genesis of neural progenitors within the subventricular zone of the developing CNS through similar mechanisms as in the adult SGZ of the dentate gyrus is uncertain—even though HIV disrupts the generation of neurons and glia during maturation or in adults. For example, MAPK/ERK1/2 enhances p53- and p21-dependent downregulation of cyclin D1 hindering progression through the G1 phase of the cell cycle in hNPCs (Mishra et al. 2010; Malik et al. 2014). Importantly, opioids too can affect the genesis of neurons and glia during maturation or in the adult directly via convergent pathways (Hauser and Knapp 2018; Kibaly et al. 2018) suggesting yet another site of opioid and HIV interactions in dysregulating the creation and fate of new neurons and glia.\nFew studies have examined the interplay between opioids, neural progenitors and HIV/HIV proteins. Sustained exposure (4 d) to morphine (500 nM) and Tat1–72 (100 nM) decreased the viability of MOR-expressing striatal glial precursors, and to a lesser extent astrocytes, and this coincided with caspase-3 activation (Khurdayan et al. 2004). By contrast, comparably administered morphine or Tat alone was sufficient to decrease the viability of immature glia/glial progenitors in spinal cord cultures, while Tat and morphine failed to interact (Buch et al. 2007). Collectively, these findings were the first to indicate that opioid and/or Tat could enhance programmed cell death in subpopulations of glial precursors in a developmentally regulated and region-dependent manner (Khurdayan et al. 2004; Buch et al. 2007). In human glial progenitors, co-administering morphine (500 nM) increased the antiproliferative effects of Tat (12–48 h) or conditioned medium from HIV-1SF162-infected MDMs (12 h), while paradoxically reversing the antiproliferative effects from HIV-1IIIB conditioned medium (12 h) (Hahn et al. 2012). In these studies, Tat or HIV exposure reduced the proliferation of Sox2+ and Olig2+ undifferentiated glial and oligodendroglial progenitors, respectively, while progenitor viability was unchanged (Hahn et al. 2012). In human neural progenitor cells (hNPCs), sustained infection with R5-tropic HIVBaL increased the proliferation and premature differentiation of hNPCs into both neurons and astrocytes, and both measures were significantly enhanced by morphine co-exposure (Balinang et al. 2017). Importantly, immunoneutralizing antibodies (Hahn et al. 2012) or the selective antagonist, maraviroc (Balinang et al. 2017), were able to significantly attenuate the consequences of R5-tropic HIV infection on hNPC differentiation and fate confirming a direct role of CCR5 in these processes. Lastly, decreases in the proliferation of hNPCs seen with morphine and Tat are, in part, regulated by ERK1/2-dependent increases in p53 and p21 expression (Malik et al. 2014) and can be modulated by PDGF BB suggesting a possible therapeutic target (Malik et al. 2011). Thus, morphine can exaggerate R5-tropic HIV-induced alterations in the maturation and fate of human and rodent NPCs, thereby further disrupting the balance of neural cell types and CNS 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