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17 Psychoneuroimmunology 17 Psychoneuroimmunology MICHAEL R. IRWIN AND GEORGE M. SLAVICH OVERVIEW external physical and social environment can also strongly Immune responses are mediated by the activation of influence immune system activity by affecting neural and immune response genes that encode regulatory and effec- endocrine processes that regulate immune system tor molecules, such as cytokines, antimicrobial peptides, dynamics. In addition to providing important new insights antibodies, and cytolytic molecules. Transcriptional acti- into the mechanisms underlying many mental and physi- vation of innate immune cells is triggered by two types of cal health problems, discoveries in this field have begun to signals from the body’s internal “environment”–namely, inform the development of novel interventions for improv- the presence of pathogen-associated molecular patterns ing human health. (PAMPs) and “danger signals” derived from host cell stress The goal of this chapter is to provide an overview of or death (Matzinger, 2007). A growing body of research contemporary models and recent research in psycho- suggests that a third class of macro-environmental stimuli neuroimmunology. First, we summarize the main features exists in the form of neural and endocrine signals and that of the innate and adaptive immune system. Second, we these stimuli also play a significant role in modulating describe the neural and physiologic pathways that have immune responses (Glaser & Kiecolt-Glaser, 2005). the ability to alter immune system dynamics. Third, we In addition, immune mediators such as cytokines feed review work linking a variety of different psychoneuroim- back to the brain to regulate neural and endocrine activity munological processes (e.g., life stress exposure, sleep dis- (Dantzer, O’Connor, Freund, Johnson, & Kelley, 2008). turbance, depression) with infectious and inflammatory The resulting neuroimmune circuit coordinates immune disease risk. Fourth, we examine the types of behavioral responses with other physiological processes such as interventions that have been shown to alter immune sys- fight-or-flight stress responses to maximize the overall tem processes and impact health. Finally, we conclude fitness of the organism within highly complex, contem- with ideas for future research. porary physical and social environments that bear multi- plemicrobial,physiological (e.g.,trauma,sleep loss),and social-ecological threats (e.g., predation, conspecificvio- IMMUNE SYSTEM lence, interpersonal loss, etc.) (McEwen, 2007). This neu- The immune system is primarily responsible for coordi- roimmune circuit was initially discovered in the context nating the body’s response to physical injuries and infec- of adaptive immune responses (Glaser & Kiecolt-Glaser, tions. It is thus critical for human health and well-being. 2005). Recent findings, however, suggest that this circuit Two interconnected branches, called innate immunity and originates with the innate immune system (Cole, adaptive immunity, are generally conceptualized as com- Hawkley, Arevalo, & Cacioppo, 2011b; Powell, Mays, prising the immune system. Innate immunity is the first Bailey, Hanke, & Sheridan, 2011). This review highlights and evolutionarily older branch. As the body’s first line of emerging biological themes on the reciprocal regulation defense against tissue damage and microbial infection of immune response gene expression and CNS function. (Medzhitov, 2008), innate immunity includes immune Psychoneuroimmunology is the study of how psycholo- cells such as monocytes/macrophages and dendritic cells, gical, neural, and immunologic processes interact and which circulate throughout the body. By using a fixed shape human health and behavior. Classic views of the small class of receptors to detect a wide variety of patho- immune system focused largely on how immune system gens, these cells initiate a cascade of inflammatory pro- processes are regulated by factors that are present inside cesses that signal the occurrence of injury or infection the body, such as viruses and bacteria, and chemical sig- (Medzhitov, 2008). Innate immunity is non-specific and nals released during host cell stress or death. It is now rapid, occurring over minutes or hours, and does not con- widely recognized, however, that characteristics of the fer long-lasting protection to the host. 377 Cite as: Irwin, M. R., & Slavich, G. M. (2017). Psychoneuroimmunology. In J. T. Cacioppo, L. G. Tassinary, & G. G. Berntson (Eds.), Handbook of psychophysiology, fourth edition (pp. 377-398). New York: Cambridge University Press. 378 IRWIN AND SLAVICH The second branch of the immune system, called adaptive (Slavich, 2015), there has recently been intense interest in immunity, becomes active when innate immune system understanding the mechanisms that underlie inflamma- defenses are insufficient (Barton, 2008). Adaptive immunity tion and the role inflammation plays in disease. involves the proliferation of microbial-specific white blood Inflammatory cytokines are very important in this context cells (i.e., lymphocytes) that attempt to neutralize or elim- because they are released from immune cells, such as inate microbes based on an immunological memory of hav- monocytes/macrophages, dendritic cells, and neutrophils, ing responded to a specific pathogen in the past (Gruys, and coordinate cell-to-cell communication; they also alter Toussaint, Niewold, & Koopmans, 2005). Compared with neurochemical and neuroendocrine processes with wide- innate immunity, the adaptive immune response takes days ranging effects on physiology and behavior (Curfs, Meis, & to fully develop (Barton, 2008), and by virtue of the devel- Hoogkamp-Korstanje, 1997). Similar to neurotransmit- opment and maintenance of immunological memory, con- ters and hormones, inflammatory cytokines mediate fers lasting protection to the organism. physiological responses, rely on receptor–ligand interac- tions, and have self (autocrine), local (paracrine), and dis- tal (endocrine) effects. Cytokines either increase or Innate Immune System decrease inflammatory activity, and these cytokines are Highly conserved features of microbes or pathogen- called pro-inflammatory and anti-inflammatory, respec- associated molecular patterns (i.e., pathogen-associated tively. A description of cytokines commonly studied in molecular patterns, or PAMPs) are recognized by receptors psychoneuroimmunology is presented in Table 17.1. of innate immune cells. This “hard-wired” recognition strat- Among the pro-inflammatory cytokines, (IL)-1, IL-6, egy, which uses a relatively small number of immune cell and TNF- coordinate several cell functions that stimulate types to detect and generate a response to a wide range of and enhance inflammation. These pro-inflammatory cyto- microbial diversity, is termed pattern recognition.Hence, kines also have effects on adaptive immunity; for example, innate immune receptors that use this strategy are called they can promote the differentiation of lymphocytes called pattern recognition receptors, and activation of these recep- cytotoxic T cells that kill pathogens. Moreover, inflamma- tors triggers an acute-phase response, which leads to an tory cytokines induce increased vascular permeability and increase in inflammatory activity that can occur both cellular adhesion, which enable the migration of immune locally and throughout the body (Medzhitov, 2008). cells from the circulation into the tissues where they can Toll-like receptors (TLRs), one class of pattern recogni- eliminate pathogens (Dhabhar, Malarkey, Neri, & tion receptors, are found on macrophages, neutrophils, McEwen, 2012). By activating the expression of the and dendritic cells (Medzhitov, 2008), and these TLRs endothelial adhesion molecule intercellular adhesion recognize conserved components of microbes including molecule-1, which then binds integrin (e.g., LFA-1) on bacteria, viruses, and fungi. Within the “family” of TLRs, the surface of immune cells, IL-1 promotes adhesion to there is some specificity of ligand recognition. For exam- endothelial cells and eventual extravasation (Smith, ple, TLR4 is one type of TLR, and this receptor binds Marlin, Rothlein, Toman, & Anderson, 1989). Similarly, lipopolysaccharide (LPS), an endotoxin that is the major TNF- stimulates the production of the adhesion molecule component of the outer membrane of Gram-negative bac- E-selectin on the endothelium, which binds to adhesion teria. Hence LPS is a prototypical PAMP (Raetz & molecules on neutrophils (Hubbard & Rothlein, 2000). Whitfield, 2002). Response to TLR4 activation can be char- A family of small cytokines, called chemokines, are acti- acterized by examining activation of a conserved signaling vated by TNF- , IL-6, and IL-1 and help redistribute cells cascade, which includes key intra-cellular transcription to sites of injury or infection by acting as chemoattractants factors such as nuclear factor-κB (NF-κB) and activator that recruit immune cells to the site of inflammatory activ- protein 1 (AP-1) (Karin, 2006). Activation of NF-κB, for ity (Murphy, 2001). example, leads to the transcription of pro-inflammatory Intra-cellular processes at the genomic level regulate the immune response genes such as tumor necrosis factor inflammatory response, and the specific pathway involved (TNF)- and interleukin (IL)-1 and the translation and depends on the type of danger signal present (Amit et al., production of pro-inflammatory cytokines that help coor- 2009). During exposure to extracellular pathogens such as dinate the inflammatory response (Karin,
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