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Virus-Host Interactions in HIV Pathogenesis: Directions for Therapy

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Virus-Host Interactions in HIV Pathogenesis: Directions for Therapy Virus-Host Interactions in HIV Pathogenesis: Directions for Therapy HIV HETEROGENEITY J.A. Levy* One of the first observations made in the evaluation of various Director, Laboratory for Tumor and AIDS Virus Research, University of HIV isolates was the viral heterogeneity involved with the abil- California, San Francisco, 513 Parnassus Avenue, Suite S2180, San ity to infect and replicate in primary CD4+ T-lymphocytes and Francisco, CA 94143-1270, USA; *corresponding author, Jay.Levy@ucsf macrophages (Levy, 2007). Moreover, whereas all viruses could .edu grow in most primary CD4+ T-cells, differences were appreci- Adv Dent Res 23(1):13-18, 2011 ated when HIV isolates were added to primary macrophages or to established CD4+ T-cell lines. Some grew well in macro- phages and others in the T-cell lines. Those growing in macro- ABSTRACT phages generally did not induce cytopathology or multinucleated The challenge of controlling HIV infection involves an under- syncytial cell formation; those growing in T-cell lines were standing of the heterogeneity of the virus, its wide cellular host syncytium-inducing (SI) viruses (Cheng-Mayer et al., 1988; range, its primary routes of transmission, and the immunologic Tersmette et al., 1988). and intrinsic cellular factors that can prevent its transmission and These early observations were later explained by the recogni- replication. Identification of HIV-infected individuals who have tion that HIV, after binding to CD4, uses secondary cellular survived more than 10 years without signs of the infection and receptors for attachment to and subsequent entry into cells without therapy encourages studies examining the natural mecha- (Berger et al., 1999). These virus co-receptors are important nisms for resistance to infection and disease. Within the immune binding sites for chemokines that recruit immune cells to the system, emphasis should be given to the innate or natural response location of their secretion. In high concentration, these chemo- that appears within minutes of the infection and offers the optimal kines (e.g., MIP-1α; MIP-1β, RANTES) can compete for virus time for controlling HIV. All these parameters in HIV pathogen- attachment to the chemokine receptor (Berger et al., 1999). The esis underline the information needed to develop optimal anti- NSI viruses primarily use the CCR5 receptor and are known as HIV therapies and an effective AIDS vaccine. R5 viruses; the T-cell line viruses use the CXCR4 receptors and are known as X4 viruses (Levy, 2007). Some HIV isolates can use both receptors and are called dual/tropic or R5/X4 viruses. INTRODUCTION Generally, during acute virus infection, an R5 virus emerges as n the 25 years since the discovery of HIV, much progress has the dominant type and over time, in many cases, becomes dual/ Ibeen made in our understanding of the virus and in recogni- tropic (Singh and Collman, 2000). Later, in about half of the tion of the importance of the infected host’s response to HIV, cases, an X4 or more cytopathic strain emerges (Asjo et al., which can determine the clinical course (Levy, 2009). While the 1986; Cheng-Mayer et al., 1988). The X4 viruses are associated wide dispersion of HIV throughout the world has now affected with a more rapid progression to AIDS (Levy, 2009). people in all countries, its ability to cause the acquired immune Recognition of the co-receptors for binding sites of HIV has deficiency syndrome (AIDS) has been greatly curtailed through led to various approaches to block this interaction through spe- the development, over the past decade, of anti-retroviral drugs. cific drugs. Maraviroc prevents HIV infection via CCR5 (Deeks, They can keep the virus in check by interfering with virus entry 2006); other drugs in development target CXCR4 (Moyle et al., and replication. Thus, infected persons can have a nearly nor- 2009). The therapies directed at CCR5 were not considered to be mal life for many years. Nevertheless, over time, resistance to detrimental to the immune system, because about 1% of the these drugs occurs, and infected individuals can develop AIDS human population lack CCR5 expression (CCR5-) and have no or toxic complications from the use of these drugs. In this arti- obvious immune abnormalities (Berger et al., 1999). These indi- cle, I will review the known characteristics of HIV and vari- viduals are much less likely to be infected by HIV unless they ous aspects of its interaction with cells. The anti-viral immune encounter an X4 virus. However, these CCR5-negative people responses of infected individuals will be considered. These fea- tures have uncovered novel anti-HIV therapies and, it is hoped, will also result in an effective AIDS vaccine. Key Words DOI: 10.1177/0022034511398874 HIV heterogeneity, innate immunity, intracellular antiviral activity. © International & American Associations for Dental Research 13 14 Levy Adv Dent Res 23(1) 2011 VIRUSES IN DIFFERENT TISSUES Another noteworthy finding is that when viruses are isolated from different tissues of the same individual (see below), variations in 1000 their biological property can be found. For example, a virus com- x cpm/mL) 3 ing from the blood may have the ability to infect peripheral blood mononuclear cells (PBMC) and to down-modulate CD4 expres- 100 sion, whereas those from the brain can be quite infectious for macrophages and not affect CD4 expression (Cheng-Mayer et al., 1989). This observation indicates that the same virus transmitted to an individual can, through multiple replicative cycles, become ranscriptase (10 T 10 selected in various tissues as a virus related to the transmitted se virus but having different biologic properties. It may be respon- ver sible for the pathology observed in the infected tissue. Re Importantly, as cited above, when more than one virus infects 0 the same cell (perhaps at the same time, when the CD4 receptor 0 10 20 30 40 50 60 has not yet been down-modulated), interactions between the 2 Days in Culture RNA strands of each virus can take place. Recombination can then occur (Robertson et al., 1995). These recombinant viruses Fig. 1. Replication of early and late HIV-1 isolates from the same contain genetic information from 2 or 3 different viruses. Some individual. Replication of 2 strains of HIV-1 recovered from the same may have genetic regions from up to 7 different subtypes (Levy, individual over time. HIV-1SF2 (0), isolated early in the course of infec- 2009). This observation has suggested that, over time, recombi- tion, replicates slowly and to low titer in the established HUT 78 T-cell nant viruses will become more common and may take on differ- line. In contrast, HIV-1SF13 (Δ), recovered when the patient had AIDS, grows rapidly and to high titer in the same cells. These and other bio- ent biologic properties, including resistance to anti-viral drugs logic differences between these 2 strains have been used to distinguish or ability to infect many cell types. For this reason, recombina- viruses in culture as non-virulent and virulent strains. Reprinted from tion among HIV strains represents a newly emerging challenge Levy (2007) with permission. in HIV pathogenesis. have been shown to have a greater chance to develop severe dis- VIRAL-HOST CELL INTERACTIONS ease from West Nile virus (Glass et al., 2006). It is less certain what a drug against CXCR4 would do to compromise the immune Within 2 days after transmission, the HIV isolate, often a single system. Conceivably, it could lead to the emergence of HIV strain (Keele et al., 2008), enters the bloodstream and can infect strains that use other co-receptors. Currently, there are at least 9 a variety of different tissues depending on its different biologic chemokine co-receptors that can be used by HIV for attachment properties. The number of cell types reported to be susceptible to various cells of the body (Berger et al., 1999; Levy, 2007). to infection is quite large and includes cells of the brain, the GI This heterogeneity of HIV, which is evident biologically, can tract, and the kidney, and as well as oral keratinocytes and epi- also be appreciated genetically. The 2 types of HIV, HIV-1 and thelial cells. HIV-2, exist and differ by about 40% (McCutchan, 2000). In This infection of different cells occurs not only with the free addition, based on genetic variations, 3 groups of HIV-1, differ- virus, but importantly also with virus-infected cells. The latter ing by about 30% (M, N, and O) have been identified, and 8 results during the replicative cycle, when HIV integrates its groups of HIV-2 have been recognized (A-H), for which A and proviral DNA into the chromosome of the cell. These cells, pres- B are the most common HIV-2 groups in circulation (Marx, ent in blood and genital fluids, can be a major source of trans- 2005). The largest number of HIV infections has occurred with mission by interacting with cells of the immune system and group M HIV-1 strains, which can be further subdivided into 9 mucosal tissues in the body. When infected cells interact with subtypes or clades differing by at least 15% (Robertson et al., mucosal cells, HIV can be transferred readily to the individual. 2000; Peeters et al., 2003). Clade B is the most common virus Such virus-infected cells can be found at levels of 50,000 cells in the United States. It is also found in China, but has emerged in some genital fluids, and the efficiency for infected cells to as well as a recombinant between clades B and C viruses (Liu transmit the virus is often better than that of the free virus (Levy, et al., 2008) (see below).
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