\r\n\tThe aim of this volume is to offer an interdisciplinary perspective in which different aspects come together, for example; psychology, economics, sociology, medicine of the common subject of study: the health. All the researchers and experts of the various disciplines are invited to participate with their studies and their theoretical models to enrich the current international literature in the context of factors and aspects that can affect health.
",isbn:"978-1-83881-956-9",printIsbn:"978-1-83881-955-2",pdfIsbn:"978-1-83881-957-6",doi:null,price:0,priceEur:0,priceUsd:0,slug:null,numberOfPages:0,isOpenForSubmission:!1,hash:"0dc7792ac9e96da752948ac76275851d",bookSignature:"Dr. Rosalba Morese and Dr. Sara Palermo",publishedDate:null,coverURL:"https://cdn.intechopen.com/books/images_new/8551.jpg",keywords:"Public Health, Socioeconomic Factors, Risky Behavior, Prevention Behavior, Cognitive Processes, Model Theory, Psychological Factors, Neuroscience, Psycho Physiological Parameters, Cultural Factors, Political Factors",numberOfDownloads:null,numberOfWosCitations:0,numberOfCrossrefCitations:0,numberOfDimensionsCitations:null,numberOfTotalCitations:null,isAvailableForWebshopOrdering:!0,dateEndFirstStepPublish:"August 20th 2019",dateEndSecondStepPublish:"September 10th 2019",dateEndThirdStepPublish:"November 9th 2019",dateEndFourthStepPublish:"January 28th 2020",dateEndFifthStepPublish:"March 28th 2020",remainingDaysToSecondStep:"3 months",secondStepPassed:!0,currentStepOfPublishingProcess:4,editedByType:null,kuFlag:!1,editors:[{id:"214435",title:"Dr.",name:"Rosalba",middleName:null,surname:"Morese",slug:"rosalba-morese",fullName:"Rosalba Morese",profilePictureURL:"https://mts.intechopen.com/storage/users/214435/images/system/214435.jpg",biography:"Rosalba Morese holds a Bachelor\\'s degree in Psychology from the University of Parma and a Ph.D. in Neuroscience from the University of Turin, Italy, and has worked to develop new techniques and approaches in Cognitive Science and Social Neuroscience. \r\nShe is an expert in Experimental Neuroscience, Neuroeconomics, Psychophysiology, Cognitive and Social Neuroscience. She performs neuroimaging studies in clinical and social contexts in order to investigate neural correlates involved in neuropsychological abnormalities and during social interactions. \r\nShe is currently a research fellow at the Department of Psychology in Turin and a teaching assistant for the course on Social Psychology and Psychology of Communication at the University of Lugano, Switzerland.",institutionString:"Universita della Svizzera Italiana",position:null,outsideEditionCount:0,totalCites:0,totalAuthoredChapters:"5",totalChapterViews:"0",totalEditedBooks:"2",institution:{name:"Universita della Svizzera Italiana",institutionURL:null,country:{name:"Switzerland"}}}],coeditorOne:{id:"233998",title:"Dr.",name:"Sara",middleName:null,surname:"Palermo",slug:"sara-palermo",fullName:"Sara Palermo",profilePictureURL:"https://mts.intechopen.com/storage/users/233998/images/system/233998.jpeg",biography:"Sara Palermo is a MSc in Clinical Psychology and a PhD in Experimental Neuroscience. Moreover, she obtained the National Scientific Enabling Certificate for Associate Professorship in April 2017 (ASN-2017). She is an expert in experimental neuroscience, clinical neuropsychology and advance neuropsychological testing. Moreover, she performs multidimensional geriatric evaluation and basic neurological symptomatology detection in patients with neurodegenerative disorders. She is also engaged in Activation Likelihood Estimation meta-analysis of neuroimaging studies.\r\nShe worked as a postdoc research fellow at the Department of Neuroscience 'Rita Levi Montalcini” in Turin until July 2017. Since then she works as research fellow at the Department of Psychology in Turin. To date, she owns three research Group memberships at the University of Turin (Italy). She is a member of the 'Center for the Study of Movement Disorders” (research area: Neurology) and the 'Placebo Responses Mapping Group” (research area: Physiology) at the Department of Neuroscience, and a member of the 'Neuropsychology of cognitive impairment and central nervous system degenerative diseases Group” at the Department of Psychology (Research Area: Psychobiology and physiological psychology).\r\nThe main topics of her research are the study of awareness of illness, metacognitive-executive deficits in neuropsychiatric and neurological disorders, physical and cognitive frailty in the elderly, and placebo/nocebo phenomena. Interestingly, all of them may represent appealing perspectives from which to study how neuropsychological abnormalities can be explained in terms of brain activities and with the use of neuropsychiatric and neuropsychological batteries considering a neurocognitive approach. Given her research interests and scientific publications, she has been an ordinary member of the Italian Society of Neuropsychology (SINP), of the Italian Association of Psychogeriatrics (AIP), of the Italian Society of Neurology for Dementia (SiNdem), and – finally – of the international Society for Interdisciplinary Placebo Studies (SIPS). Importantly, she is a member of the European Innovation Partnership on Active and Healthy Aging (EIP on AHA), for which she is involved in the Action Group A3 Functional decline and frailty. \r\n\r\nSara Palermo is Panel Editor for 'EC Psychology and Psychiatry'. She was recently appointed as Specialty Chief Editor for 'Frontiers in Psychology - Neuropsychology'.",institutionString:"University of Turin",position:null,outsideEditionCount:0,totalCites:0,totalAuthoredChapters:"3",totalChapterViews:"0",totalEditedBooks:"1",institution:{name:"University of Turin",institutionURL:null,country:{name:"Italy"}}},coeditorTwo:null,coeditorThree:null,coeditorFour:null,coeditorFive:null,topics:[{id:"16",title:"Medicine",slug:"medicine"}],chapters:null,productType:{id:"1",title:"Edited Volume",chapterContentType:"chapter",authoredCaption:"Edited by"},personalPublishingAssistant:{id:"280415",firstName:"Josip",lastName:"Knapic",middleName:null,title:"Mr.",imageUrl:"https://mts.intechopen.com/storage/users/280415/images/8050_n.jpg",email:"josip@intechopen.com",biography:"As an Author Service Manager my responsibilities include monitoring and facilitating all publishing activities for authors and editors. From chapter submission and review, to approval and revision, copy-editing and design, until final publication, I work closely with authors and editors to ensure a simple and easy publishing process. I maintain constant and effective communication with authors, editors and reviewers, which allows for a level of personal support that enables contributors to fully commit and concentrate on the chapters they are writing, editing, or reviewing. I assist authors in the preparation of their full chapter submissions and track important deadlines and ensure they are met. I help to coordinate internal processes such as linguistic review, and monitor the technical aspects of the process. As an ASM I am also involved in the acquisition of editors. Whether that be identifying an exceptional author and proposing an editorship collaboration, or contacting researchers who would like the opportunity to work with IntechOpen, I establish and help manage author and editor acquisition and contact."}},relatedBooks:[{type:"book",id:"8262",title:"The New Forms of Social Exclusion",subtitle:null,isOpenForSubmission:!1,hash:"29bf235aa7659d3651183fe9ea49dc0d",slug:"the-new-forms-of-social-exclusion",bookSignature:"Rosalba Morese and Sara Palermo",coverURL:"https://cdn.intechopen.com/books/images_new/8262.jpg",editedByType:"Edited by",editors:[{id:"214435",title:"Dr.",name:"Rosalba",surname:"Morese",slug:"rosalba-morese",fullName:"Rosalba Morese"}],productType:{id:"1",chapterContentType:"chapter",authoredCaption:"Edited by"}},{type:"book",id:"5810",title:"Socialization",subtitle:"A Multidimensional Perspective",isOpenForSubmission:!1,hash:"bfac2e9c0ec2963193e9d15d617c6a01",slug:"socialization-a-multidimensional-perspective",bookSignature:"Rosalba Morese, Sara Palermo and Juri Nervo",coverURL:"https://cdn.intechopen.com/books/images_new/5810.jpg",editedByType:"Edited by",editors:[{id:"214435",title:"Dr.",name:"Rosalba",surname:"Morese",slug:"rosalba-morese",fullName:"Rosalba Morese"}],productType:{id:"1",chapterContentType:"chapter",authoredCaption:"Edited by"}},{type:"book",id:"6550",title:"Cohort Studies in Health Sciences",subtitle:null,isOpenForSubmission:!1,hash:"01df5aba4fff1a84b37a2fdafa809660",slug:"cohort-studies-in-health-sciences",bookSignature:"R. Mauricio Barría",coverURL:"https://cdn.intechopen.com/books/images_new/6550.jpg",editedByType:"Edited by",editors:[{id:"88861",title:"Dr.",name:"René Mauricio",surname:"Barría",slug:"rene-mauricio-barria",fullName:"René Mauricio Barría"}],productType:{id:"1",chapterContentType:"chapter",authoredCaption:"Edited by"}},{type:"book",id:"3161",title:"Frontiers in Guided Wave Optics and Optoelectronics",subtitle:null,isOpenForSubmission:!1,hash:"deb44e9c99f82bbce1083abea743146c",slug:"frontiers-in-guided-wave-optics-and-optoelectronics",bookSignature:"Bishnu Pal",coverURL:"https://cdn.intechopen.com/books/images_new/3161.jpg",editedByType:"Edited by",editors:[{id:"4782",title:"Prof.",name:"Bishnu",surname:"Pal",slug:"bishnu-pal",fullName:"Bishnu Pal"}],productType:{id:"1",chapterContentType:"chapter",authoredCaption:"Edited by"}},{type:"book",id:"72",title:"Ionic Liquids",subtitle:"Theory, Properties, New Approaches",isOpenForSubmission:!1,hash:"d94ffa3cfa10505e3b1d676d46fcd3f5",slug:"ionic-liquids-theory-properties-new-approaches",bookSignature:"Alexander Kokorin",coverURL:"https://cdn.intechopen.com/books/images_new/72.jpg",editedByType:"Edited by",editors:[{id:"19816",title:"Prof.",name:"Alexander",surname:"Kokorin",slug:"alexander-kokorin",fullName:"Alexander Kokorin"}],productType:{id:"1",chapterContentType:"chapter",authoredCaption:"Edited by"}},{type:"book",id:"1591",title:"Infrared Spectroscopy",subtitle:"Materials Science, Engineering and Technology",isOpenForSubmission:!1,hash:"99b4b7b71a8caeb693ed762b40b017f4",slug:"infrared-spectroscopy-materials-science-engineering-and-technology",bookSignature:"Theophile Theophanides",coverURL:"https://cdn.intechopen.com/books/images_new/1591.jpg",editedByType:"Edited by",editors:[{id:"37194",title:"Dr.",name:"Theophanides",surname:"Theophile",slug:"theophanides-theophile",fullName:"Theophanides Theophile"}],productType:{id:"1",chapterContentType:"chapter",authoredCaption:"Edited by"}},{type:"book",id:"1373",title:"Ionic Liquids",subtitle:"Applications and Perspectives",isOpenForSubmission:!1,hash:"5e9ae5ae9167cde4b344e499a792c41c",slug:"ionic-liquids-applications-and-perspectives",bookSignature:"Alexander Kokorin",coverURL:"https://cdn.intechopen.com/books/images_new/1373.jpg",editedByType:"Edited by",editors:[{id:"19816",title:"Prof.",name:"Alexander",surname:"Kokorin",slug:"alexander-kokorin",fullName:"Alexander Kokorin"}],productType:{id:"1",chapterContentType:"chapter",authoredCaption:"Edited by"}},{type:"book",id:"4816",title:"Face Recognition",subtitle:null,isOpenForSubmission:!1,hash:"146063b5359146b7718ea86bad47c8eb",slug:"face_recognition",bookSignature:"Kresimir Delac and Mislav Grgic",coverURL:"https://cdn.intechopen.com/books/images_new/4816.jpg",editedByType:"Edited by",editors:[{id:"528",title:"Dr.",name:"Kresimir",surname:"Delac",slug:"kresimir-delac",fullName:"Kresimir Delac"}],productType:{id:"1",chapterContentType:"chapter",authoredCaption:"Edited by"}},{type:"book",id:"57",title:"Physics and Applications of Graphene",subtitle:"Experiments",isOpenForSubmission:!1,hash:"0e6622a71cf4f02f45bfdd5691e1189a",slug:"physics-and-applications-of-graphene-experiments",bookSignature:"Sergey Mikhailov",coverURL:"https://cdn.intechopen.com/books/images_new/57.jpg",editedByType:"Edited by",editors:[{id:"16042",title:"Dr.",name:"Sergey",surname:"Mikhailov",slug:"sergey-mikhailov",fullName:"Sergey Mikhailov"}],productType:{id:"1",chapterContentType:"chapter",authoredCaption:"Edited by"}},{type:"book",id:"3092",title:"Anopheles mosquitoes",subtitle:"New insights into malaria vectors",isOpenForSubmission:!1,hash:"c9e622485316d5e296288bf24d2b0d64",slug:"anopheles-mosquitoes-new-insights-into-malaria-vectors",bookSignature:"Sylvie Manguin",coverURL:"https://cdn.intechopen.com/books/images_new/3092.jpg",editedByType:"Edited by",editors:[{id:"50017",title:"Prof.",name:"Sylvie",surname:"Manguin",slug:"sylvie-manguin",fullName:"Sylvie Manguin"}],productType:{id:"1",chapterContentType:"chapter",authoredCaption:"Edited by"}}]},chapter:{item:{type:"chapter",id:"62945",title:"Dendritic Cell Subsets, Maturation and Function",doi:"10.5772/intechopen.79926",slug:"dendritic-cell-subsets-maturation-and-function",body:'\nDendritic cells (DCs) are rare, heterogeneous bone marrow (BM)-derived professional APCs that are disseminated ubiquitously in blood, lymphoid, and peripheral tissues, particularly at the gates of antigen entry. They originate from hematopoietic stem cells throughout specialized progenitor subsets and are essential in innate and adaptive immune capacity and in managing the balance between immunity and tolerance [1]. Under normal conditions, DCs are present throughout the body at low numbers representing ≈1–2% of white blood cells [2]. In the steady state, DCs reside in immature or semi-mature states in the periphery where they regularly take up and process self-Ags and maintain self-tolerance [3]. Immuno-stimulatory DCs have undergone maturation after recognition of exogenous and endogenous danger signals by Toll-like receptors (TLRs). These signals include pathogen-associated molecular patterns in the form of microbial products, such as products of damaged or dying cells [4].
\nDCs are matured by CD40 ligation and by pro-inflammatory cytokines that can produce DC maturation ex vivo, detached of CD40 ligation. Maturation is correlated with up-regulation of cell surface MHC gene products, co-stimulatory molecules (CD40, CD80, and CD86 and CD83), and relevant chemokine receptors that improve the ability of DCs to migrate to secondary lymphoid tissue, where they present Ag to Ag-specific T cells and induce T-cell activation and generation. Consequently, activated T cells drive DCs toward terminal maturation [5].
\nDCs produce from Hematopoietic stem cells (HSCs) in the BM and are originated from both myeloid and lymphoid progenitors, as illustrated in Figure 1. Both subsets, conventional DC (cDC) and plasmacytoid DC subsets (pDC), are derived from a common CD34+ progenitor [6]. The hematopoietic growth factor fms-like tyrosine kinase 3 ligand (Flt3L) represents a fundamental function in steady-state DC expansion; this is evidenced by the preponderance of DC precursors being Flt3+ (CD135+) and culture with Flt3L appearing in cDC and pDC subsets. GM-CSF is also crucial in DC hematopoiesis, as it provides DCs from monocytes and immature progenitors in the deficiency of intact Flt3L signaling and provides DCs under inflammatory conditions [1].
\nDendritic cell hematopoiesis.
DCs are divided into two principal cell populations, conventional DCs (cDCs) and plasmacytoid DCs (pDCs). In the steady state, cDCs present typical DC characteristics (e.g., cytoplasmic dendrites) and function (e.g., Ag uptake, processing, and exhibition). cDCs can be divided into migratory DCs, such as skin epidermal Langerhans cells (LCs), dermal DCs, which present Ag in lymph nodes following its uptake in peripheral tissue and resident DCs, which take up and process Ag within a lymphoid organ, such as splenic or thymic DCs [1]. Thymic DCs remove self-Ag-specific thymocytes and stimulate the expansion of immunoregulatory T cells (Treg). Thymic conventional DCs (cDC) readily received MHC class I and II from thymic epithelial cells (TEC), but plasmacytoid DCs (pDC) were less effective. Intercellular MHC shift was donor cell-specific; thymic DC readily gained MHC from TEC plus thymic or splenic DC, whereas thymic or splenic B cells were smaller donors [7].
\nPlasmacytoid DCs (pDCs) are a subset of precursor DCs which possess an immature phenotype in the steady-state and plasma cell morphology (e.g., lack dendrites). On activation, pDCs strictly match cDCs in form and function. Monocyte-derived DCs or inflammatory DCs are similar to cDCs in form and function and related to in vitro GM-CSF-generated DCs [3].
\nUnder steady-state conditions, human pDCs display lower levels of MHC and costimulatory molecules compared with conventional myeloid DCs (mDCs). pDCs are less efficient in Ag processing and loading ability to excite T cells than mDCs. After their activation via TLR, pDCs produce high levels of type 1 interferon (IFN) and incite CD4+ and CD8+ T cells. This is in opposition to activated mDCs, which secrete IL-12 and enhance T-helper type-1 (Th1) cell differentiation and CD8+ cytotoxic T lymphocyte (CTL) responses [8]. Plasmacytoid DCs (pDCs) are a subset of precursor DCs which have an immature phenotype in the steady-state and plasma cell morphology (e.g., lack dendrites). On activation, pDCs closely resemble cDCs in form and function. Monocyte-derived DCs or inflammatory DCs are similar to cDCs in form and function and correlate with in vitro GM-CSF-generated DCs [3].
\nUnder steady-state conditions, human pDCs display lower levels of MHC and costimulatory molecules compared with conventional myeloid DCs (mDCs). pDCs are less efficient in Ag processing and loading ability to stimulate T cells than mDCs. After their activation via TLR, pDCs secrete high levels of type 1 interferon (IFN) and stimulate CD4+ and CD8+ T cells. This is in contradiction to activated mDCs, which produce IL-12 and increase T-helper type-1 (Th1) cell differentiation and CD8+ cytotoxic T lymphocyte (CTL) responses [8].
\npDCs have intrinsic tolerogenic features; in the steady position, human thymic pDCs provoke Treg, whereas liver and airway pDCs control oral and mucosal tolerance, respectively. pDCs have also been involved in the management of disease activity in experimental models of autoimmunity and revealed to exert disease-suppressing capacity [9].
\nIt may be important after transplantation regarding donor engraftment (tolerance), which has clinical features that overlap with autoimmune disease. Epidermal LCs may be immunostimulatory or tolerogenic, depending on their state of maturity, inciting immunogen, and the cytokine environment [10].
\nDCs are characterized by high versatility, flexibility and multiple functional activities combined with their dual capacity to induce self-tolerance or trigger immune responses. The principal function of DCs is to scare the immune system toward heterogeneous and dangerous invasions and to defend self-tissues from destruction to keep self-tolerance [11]. The coordination of these supposedly multiple functions may open up new roads for stimulating or controlling immune responses and to promote defensive or therapeutic remedies for controlling inflammatory and autoimmune diseases or cancer, as well as designing unusual varieties of vaccines based on DCs biology [12].
\nA basic biological role of DCs relies on the constant sampling of their tissue environment, reacting to stress, risk signals and transducing the gathered molecular information to other cell classes of the immune system [13]. DCs are implemented with characteristics sets of pattern-recognition receptors, such as TLRs (Toll-like receptors), NLRs (NOD-like receptors), and RLRs (RIG-I-like receptors), which are specialized to recognize exogenous pathogen-associated molecular patterns (PAMPs) and endogenous danger signals, damage-associated molecular patterns (DAMPs) [14].
\nThe response of DCs to MAMPs and DAMPs is achieved by the activation of pausing DCs by microbial components, noxious or toxic abuses. Activation of DCs sequences in the expression of costimulatory molecules, the generation of cytokines, chemokines and additional soluble mediators. Both are resting and stimulated DCs can switch their tissue position and transfer through peripheral and lymphoid tissues. Activation of DCs by MAMPs and DAMPS appears in the prompt, chemokine-mediated translocation of DCs to peripheral lymph nodes where they have the possibility to communicate naive T-lymphocytes to induct adaptive immune responses [15]. This process assures the transformation of molecular message obtained in the periphery toward other cell varieties of both innate and adaptive immunity such as neutrophils, granulocytes, NKs, killer T cells, T- and B-lymphocytes [16].
\nThe response of DCs can be divided into the perception phase followed by phases of signal transduction pathways supported by adaptors and interfered by post-translational changes such as phosphorylation and ubiquitination reactions leading to the activation of transcription factors, and gene transcription followed by the secretions of soluble factors [17].
\nIn this cascade, few receptor complexes ligated by their specific ligands allow substantial signal amplification. It has also been shown that the generation of fully active and stable DCs requires the parallel activation of multiple signaling pathways [18]. Signs through a particular receptor may produce partial stimulation only, which may be regressed by signals which promote the differentiation of regulative DCs. Signals produced by Toll-like receptors (TLRs), cytokines, chemokines, eicosanoids, free oxygen radicals, and several inflammatory mediators provide a signaling matrix and determine the phenotype and functional activities of DCs [19].
\nFive types of PRRs have been recognized: (i) transmembrane TLRs, which are combined to cell surface or endosomal membranes of different cell types, (ii) membrane C-type lectin receptors (CLRs) identified by the appearance of a carbohydrate-binding domain, (iii) three further classes of intracellular sensors, which are confined to the cytosol of multiple cell types and include NOD-like receptors (NLRs), RIG-like receptors (RLRs), and the latterly expressed AIM2-like receptors (ALRs), all with nucleotide recognition capacities [20].
\nUpon binding of their specific ligands, TLRs activate the NF-κB/AP-1 and the interferon-regulatory factor 7/3 (IRF-7/3) pathways to coordinate innate and initiate adaptive immunity [21].
\nRLRs are crucial viral sensors in the cytoplasm and contain retinoic acid-inducible gene-I (RIG-I), melanoma differentiation-associated gene-5 (MDA5), and laboratory of genetics and physiology 2 (LGP2), sequentially. RIG-I and MDA5 have been recognized as receptors toward double-stranded RNA [22].
\nNucleotide-binding oligomerization domain (NOD)-like receptors mediate primarily antibacterial immunity through the activation of NF-κB or inflammasomes, whereas RIG-I-like helicases have a fundamental role in the induction of antiviral immune responses [23] (Figure 2).
\nTLR and RLR signaling.
The collaboration of PRRs and the resulting secretion of type I interferons and inflammatory cytokines can be extremely potent toward pathogens. Following infections, innate defense mechanisms are stimulated immediately and support the expansion of adaptive immune responses. DCs perform a crucial role in the orchestration of humoral and cellular immunity and the initiation and sustaining of long-term immunological memory [24]. Interaction of microbes with the innate immune system involves the induction of multiple PRR pathways triggered simultaneously by various PAMPs of the whole pathogen [25].
\nThe possible interaction of two or more signaling pathways in biochemical systems can either be potentiating or hampering. For example, in moDCs and monocyte-derived Langerhans cells (moLCs), co-ligation of TLR3/TLR7 and TLR3/Dectin-1 lead to increased Th1/Th17 responses, in contrast to TLR3 and Langerin ligation, which had an opposite effect [26]. Similarly, another group found that RLR/TLR co-activation caused decreased Th1/Th17 responses upon bacterial infection. This cross-interference of RLR and TLR signaling might have significant implications in the design of future vaccination strategies, and the possible spectrum may be expanded to other non-immune cell types as well [27].
\nIn vaccine construction, a primary purpose is to produce efficient, specific T-cell responses. This is accomplished by targeting antigen to cell surface molecules on DCs that efficiently direct the antigen into endocytic chambers for packing onto MHC molecules and stimulation of T-cell responses. Toll-like receptors (TLRs) expressed on DCs employed as intentions for antigen presentation for cancer and different disorders [28].
\nDepending on phenotypic and functional requirements, DCs may develop immunogenic or tolerogenic responses. Although several Toll-like receptors, such as TLR3, TLR4, TLR5, TLR7, and TLR8, provoke immune activation, others can quiet immune responses by tolerance initiation in DCs. Under certain conditions, TLR2 activation can lead to IL-10 production or Treg cell activation via repression of TLR7/TLR9 signaling and prevention of IFN-α and -β secretion from pDCs [29].
\nDespite the immunogenic capability of DCs in mounting immune responses, which has been assigned to the only target in the immune system, they have also been ascribed several roles in tolerance installation and silencing of immune responses. DCs express a fundamental role in the induction of several subsets of T cells, such as Th1, Th2, Th17 and regulatory T cells (Tregs). In the steady state, DCs play a critical role in the induction of tolerance against self-antigens. Complete ablation of DCs breaks self-tolerance of CD4+ T cells and results in fatal autoimmunity [30] (Figure 3).
\nDendritic cells in the choice between immunity and tolerance.
Although the general state of knowledge considers cDCs as inducers of immunity, while pDCs serve as the inducer of tolerance [31], their functions in the immune response to a diverse range of antigens are more complex.
\npDCs are believed to be the critical effector cells in the early antiviral innate immune response by providing large quantities of type I interferons upon viral infection. pDCs increase immune responses by cross-talking with cDCs by the secretion of IFN-α, through performing a crucial role in active stimulation of adaptive immunity as well. In the interest to IFN-α secretion, it has been described that pDCs also express CD40L, which stimulates cDCs to secrete IL-12 [32] (Figure 4).
\nCooperative action of different DC subsets to tackle both innate and adaptive immunity.
An association between the appearance and deficiency of multiple surface markers has been employed to identify DC subsets. These include the presence of significant expression of class II MHC antigens and the insufficiency of several progenitors’ markers such as CD3 (T cell marker), CD14 (monocyte marker), CD19 (B cell marker), CD56 (natural killer cell marker) and CD66b (granulocyte marker). DCs further express a modification of adhesion molecules including CD11a (LFA-1), CD11c, CD50 (ICAM-2), CD54 (ICAM-1), CD58 (LFA-3), and CD102 (ICAM-3). DCs also represent costimulatory molecules including CD80 (B7.1), and CD86 (B7.2), which are upregulated through DC activation. CD86 designates to be a marker of primary DC maturation, while CD80 only increases in mature DC. Two additional markers of mature DC in humans are CD83 and CMRF-44. CD83 also is exposed by stimulated B cells, and CMRF-44 will also be exposed by macrophages and monocytes [33].
\nThe identification of DCs by surface phenotyping may be accomplished by merely demonstrating a high level of MHC class II or a costimulatory molecule such as CD80 and the absence of lineage markers [34].
\nThe conventional or myeloid DCs (cDCs) are characterized by a high exhibit of the phenotype LIN-CD11c and low HLA-DR + CD123, while plasmacytoid DCs (pDCs), derived from a lymphoid precursor, manifest low expression of the phenotype LIN-CD11c and high HLA-DR + CD123 [35]. The maturation state of DCs can categorize DCs. Immature DCs are located mainly in peripheral tissues, where they capture antigens, initiate their maturation and migrate to lymphoid organs, where they become mature to present antigen and stimulate naive T lymphocytes [36].
\nBuckley et al. [37] revealed that macrophages and DCs are positioned in the same splenic anatomical sections and yield monocyte-macrophage markers, proposing that both cell classes are relevant and probably originated from a familiar precursor. Vandenabeele et al. [38] illustrated in human thymus main classes of DCs, showing the low of the phenotype CD11b-CD11 + CD45RO, great CD83, CD86, HLA-DR and fewer DCs with high CD11b + CD11c, CD45RO population. They also recorded the appearance of pDCs with great CD123 in the thymic cortex. The role of DCs is tightly correlated to their anatomical location. In secondary lymphoid tissues, mature DCs present antigens, caught in the periphery, to naive T cells and produce immunity, while in the thymus DCs present self-antigens, produce negative determination of autoreactive T cells and improve the positive selection of regulatory T cells [39].
\nDCs can be generated via culturing CD34+ cells in the presence of several cytokines. One procedure which has been developed includes depleting the CD34+ cells of differentiated ancestors and next culture the cells in the presence of GM-CSF and IL-4 ± TNF-α. CD34+ cells can be collected from bone marrow or cord blood. Further procedure is to generate DC-like cells by culturing CD14+ monocyte-enriched peripheral blood mononuclear cells [40]. In the presence of GM-CSF and IL-4, these cultures lead to large numbers of DC like cells. These monocyte-derived DCs require additional conditioning in vitro with either TNF-α or lipopolysaccharides added to culture media to enable adequately function as a DC accomplished of preparing antigen-specific T cell responses [41].
\nBecause of the established role of DCs in maintaining the balance between immunity and tolerance, tolerogenic (tol)DCs might be novel therapeutic targets to prevent undesirable (auto-)immune responses. The idea behind tolDC therapy is that it is a highly targeted, antigen-specific treatment that only affects the auto-reactive inflammatory response [42]. A tolerogenic state in DCs can be induced using several pharmacological agents, such as cyclosporine A, rapamycin, dexamethasone, vitamin A, vitamin D or other cytokines and growth factors [43].
\nIsolation and culture of leukocytes (buffy coats) obtained from heparinized human peripheral blood provide a valuable model for studies on DCs biology and may help uncover new means to manipulate DCs differentiation and function in therapeutic settings [44]. The buffy coat layer from human peripheral blood was cultured in the presence of GM-CSF and IL-4 to generate dendritic cell populations which were allowed to differentiate into mature DCs by TNF-α within 9 days [45] (Figure 5).
\nMorphological changes during generation and differentiation of DCs (40×), (a) adherent monocytes on day 0, (b) transforming monocytes on day 3, (c) generated DCs on day 7 and (d) mature DCs on day 9. The culture of buffy coat layer from human peripheral blood leads to generation of dendritic cell populations that in response to granulocyte-macrophage colony-stimulating factor (GM-CSF) plus interleukin-4 (IL-4) in 7 days and differentiate into mature DCs in response to maturation stimulus tumor necrosis factor-α (TNF-α). Morphological changes were examined under inverted microscope Carl Zeiss® using ZEN 2012® software, Germany.
The in vitro effect of dexamethasone (DEX) on generation and differentiation of DCs through microscopic and phenotypic analysis was studied. The addition of DEX to the culture on day 0 prevented the differentiation of DCs to be tolerogenic. On the other hand, addition of DEX to the culture on day 7 or 8, either preceded or followed by addition of TNF-α, resulted in significant increase of CD83 expressing DCs; the greatest percent of tolerogenic DCs was obtained in the culture media to which DEX (1 μM) was added on day 8 and TNF-α (10 ng mLG1) was added on day 7. Although the addition of TNF-α to the culture 1 day prior to addition of DEX enhanced the differentiation of DCs (high percent of CD83 expressing DCs), TNF-α did not affect the morphological changes of DCs which became mature even in the absence of TNF-α. Opposite studies were reported that TNF-α is a maturation factor essential for the appearance of the morphological characteristics of DCs [46].
\nCD83 is an important marker for activated/mature DCs. It was recorded that both stimulated DCs and B cells secrete soluble form of CD83 and so low concentration of soluble CD83 are present in normal human sera [47]. The CD83 seems to possess regulatory roles for immune response. The soluble form of CD83 can repress immune responses, while being strongly up-regulated during DCs maturation and activation [48].
\nFujimoto and Tedder (2006) revealed that CD83 has immunosuppressive roles such as the inhibition of surface molecules, such as MHC-II, reducing the dendritic cell-mediated T cell stimulation. The allogeneic stimulatory capacity of the DCs and immunosuppressant mechanisms of CD83 were illustrated significantly inhibiting anti-donor antibody responses [49]. The study of Ge et al. [50] reported that CD83 is capable of down-modulating expression of various DC [50]. The elevated CD83 expression suggests the possibility of DEX-generated cells to initiate a Th2-biased response where CD83 is able to inhibit DCs mediated T cells stimulation [51]. Furthermore, dexamethasone treated DCs possessed the capacity to convert CD4+ T cells into IL-10-secreting Treg potently suppressing the proliferation of responder T cells [52].
\nThe CD83 is a surface marker that distinguishes immature and mature human dendritic cell populations. The CD83 is type 1 glycoprotein belonging to the immunoglobulin superfamily and has been known to be one of the best markers. There is an outstanding deal of attention in how DCs might be developed as a manner of immunotherapy. DCs are being examined as adjuvants for vaccines or as a principal therapy to aggravate immunity against cancer. That DCs may show valuable in cancer has been most often studied in animal models. DCs burdened with tumor lysates, tumor antigen-derived peptides, MHC class I modified peptides, or whole protein have all been shown to yield anti-cancer immune responses and actions, including in some cases the ability to begin broad relapse of existing tumor [53] (Figure 6).
\nGeneration of immunogenic cancer cells fused to activated dendritic cells.
In conclusion, there is a pronounced hope to study these strategies and use tumor-antigen bearing DCs as a vaccine in humans. Human clinical investigations are continuing in numerous institutions to use DCs to initiate immunity to antigens against breast cancer, lung cancer, melanoma, prostate and renal cell cancers [54]. The study of immune-mediated mechanisms could be of value in avoiding and managing main immune disorders [45].
\nOrthodontic treatment in the present day does not just require to meet the demands of creating the functional harmony in occlusion and improving the aesthetic outlook of but is should also be completed in the most efficient duration that is accepted by the patient and the orthodontist. We live in a fast-paced world where the treatment duration has clearly made the field of orthodontic treatment to revolve around it. Accelerated orthodontic tooth movement is not something that has recently emerged; it has been studied and tried out for many years. In an attempt of producing faster tooth movement during orthodontic treatment, there are numerous methods of accelerating tooth movements that have been introduced over the years which range from surgical means to the use of laser therapy. Now let us look at each method explained in this chapter.
\nWe can categorise the methods of accelerated tooth movement into the following categories:
Pharmacological methods
Surgical methods
Physical methods
Orthodontic forces cause a fluid movement in the periodontal ligament space and distortion of the matrix and cells. There is release of molecules which initiate bone remodelling for tooth movement [1]. There are a number of researches on pharmacological agents that act as biomodulators for increased orthodontic tooth movement. These are examples of such biomodulators:
Prostaglandin E2 and Prostaglandin E1
Misoprostol
1,25-Dihydroxycholecalciferol
Parathyroid hormones
Intravenous immunoglobulins
Prostaglandin E2 (PGE2) is an arachidonic acid metabolite is an often-tested substance to increase orthodontic tooth movement [2]. Animal studies have shown PGE2 to increase tooth movement and facilitate bone resorption [3, 4, 5, 6]. Camacho and Velásquez Cujar conducted a study that showed that it required repeated injections due to its short half-life [7]. Particular synthases that are required for the synthesis of PGE2 could be targeted to control the production of the prostaglandins [8].
\nAnother prostaglandin that has been reported to speed up orthodontic tooth movement is Prostaglandin E1 (PGE1). Prostaglandin E1 (PGE1) has also been seen to be induced by mechanical stress and cause bone remodelling, Patil and his co-workers had shown that even minimal amounts of PGE1 injection had significant increase in tooth movement [9]. Due to the hyperalgesia that accompanies with the local injection of PGE1, an analogue of it which is misoprostol was tried out. It was seen that it was effective in increasing orthodontic tooth movement (Figure 1) [10].
\nInjection of a biomodulator in the periodontium.
The parathyroid hormone (PTH) acts directly on osteoblasts and on osteoclasts indirectly by binding to the PTH type 1 receptor on osteoblasts. This causes the expression of insulin like growth factor 1. There is promotion of osteoblast survival, osteoblastogenesis and receptor activator for nuclear factor κ B ligand (RANKL) which induces osteoclast activation [2]. PTH facilitates bone remodelling in intermittent treatment by enhancing activities of osteoblasts and osteoclasts [11].
\nCalcitriol or 1,25-dihydroxycholecalciferol which is the most active metabolite of vitamin D acts in a similar fashion to PTH by facilitating osteoblastic proliferation and function [12]. Calcitriol facilitates alveolar bone remodelling which leads to increase in tooth movement while force application [6, 13].
\nRecently intravenous immunoglobulin (IVIg) preparations which are used in immunodeficient patients as replacement therapy. These preparations have been shown to induce COX-2 mediated PGE2 and cytokine production [14, 15]. Future potential of these preparations could be used to modulate orthodontic movement via PGE2 synthesis.
\nBichmalyr in 1931, put forward a surgical technique with orthodontic appliances for rapid correction of severe maxillary protrusion. First, wedges of bone were removed to reduce the volume for which the roots of the maxillary anterior teeth would require for retraction. Kӧle further looked into this technique in 1959 by including special movements like crossbite correction and space closure. He believed that he was able to move bony blocks using the crowns of teeth as handles as the blocks were connected by only less-dense medullary bone [16]. Currently there are few surgical methods being practiced, they are:
Periodontally accelerated osteogenic orthodontics
Piezocision
Micro-osteoperforations
In 2001, Wilcko et al. had introduced a method which combines corticotomy surgery and alveolar bone grafting which is referred to as accelerated osteogenic orthodontics or recently termed as periodontally accelerated osteogenic orthodontics (PAOO) [16]. This procedure which enables rapid tooth movement is due to a healing event that was described by Frost [17] and termed as regional acceleratory phenomenon (RAP).
\nRAP is the acceleration of the normal regional healing process from the original injury. It usually occurs after osteotomy, bone-grafting procedure, arthrodesis and fractures and there might be involvement and activation of precursor cells required for healing at the injury site. RAP can increase both soft and hard tissue healing processes by two- to tenfold [17]. It usually starts in the first few days of injury, peaks at the first or second month and may last for 3–4 months [16].
\nOrthodontic treatment can be started 1 week before or within 2 weeks after the surgery. Surgery begins with flap reflection and decortication with low-speed round burs. Bone graft is then laid over these areas of corticotomies. The flaps are then closed and sutured [18]. Several studies have been done related to corticotomies, an example is one by Uzuner and her co-workers where they showed that canine retraction assisted by corticotomy had reduced duration of retraction by 20% ratio [19]. PAOO has shown to have reduced treatment time, produce lower cortical bone resistance leading to reduced root resorption, enhancement of post-orthodontic stability, increased bone support since there is supplementation of the bone graft. However, PAOO still has risks since it is an invasive procedure and is expensive [20, 21, 22, 23, 24].
\nSince the corticotomy procedure is still invasive, Dibart et al. introduced a new minimally invasive method called piezocision. Piezocision involves microincisions which are confined to the buccal side that allows the use of piezoelectric knife and selective tunnelling which enables hard and soft tissue grafting [25]. Piezocision is usually done a week after orthodontic appliance placement. The procedure involves vertical incisions made buccally and interproximally. The mid portion of the incision between the roots enables the piezoelectric knife to be inserted. A piezotome is then inserted in the gingival openings that were made and piezoelectrical corticotomy of 3 mm is made. Hard or soft tissue grafts can then be added via a tunnelling procedure (Figure 2) [26].
\nPiezocision.
Piezocision can be used as an adjunct to treat a number of malocclusions and aid in rapid orthodontic treatment in adults. Since it is much more minimally invasive than corticotomy, it is having high degree of patient acceptance, short surgical time and has less postoperative discomfort [25, 26]. Dibart and coworkers in 2013 showed that there was an increase in the rate of tooth movement in their animal study and preliminary human studies are being conducted to correlate with the animal studies [26, 27].
\nTo further reduce the amount of invasive nature of surgical intervention, a method called micro-osteoperforation (MOP). It is procedure in which small pinhole-sized perforations are created within the alveolar bone surrounding the dentition. This initiates cytokine release to call in osteoclasts to increase bone resorption. Thus, acceleration of tooth movement occurs during orthodontic treatment. The site of perforation is within the attached gingiva and close to the target teeth on the mesial and distal aspect of the roots of the teeth which will be moved. The most favourable place for placement of the perforation is the buccal cortical plate but lingual plate can also be approached with a contra-angled appliance. Two to four perforations are ideal amounts with depths of 3–7 mm into the bone [28].
\nIn 2013, Alikhani et al. showed that MOP increased expression of cytokines for osteoclast differentiation, increased canine retraction, reduced orthodontic treatment by 62% with mild discomfort in patients [29]. In an animal study, Alikhani and co-workers found that the expression of inflammatory markers and bone resorption was significant. Their human clinical trial found distalisation was twice as much with MOP than the forces alone [30].
\nDespite all the attempts in making surgical methods being minimally invasive, they still remain as an invasive procedure. This had led to discoveries in other tools that can accelerate tooth movement during orthodontic treatment. The two most common physical methods used in the present day are:
Vibratory stimulus
Low level laser therapy
Low-intensity pulsed ultrasound
Bone has the ability to respond to the mechanical stimuli that is applied to it as a mechanism to withstand functional activity. Rubin et al. showed the rate of remodelling in mechanically loaded long bones have been increased following vibrations or low level mechanical oscillatory signals [31]. In 2008, Nishimura et al. did an animal study which gave an insight on how resonance vibration could be able to accelerate tooth movement through the expression of RANKL in the periodontal ligament [32].
\nA novel device that was introduced by OrthoAccel Technologies is the AcceleDent device. The device has an activator and a mouthpiece. The patient bites on the mouthpiece component when in use. The activator which is extraorally positioned generates and transmits vibrations to the teeth. It can provide 0.2 N of vibration at 30 Hz for 20 minutes. It was fabricated to work in tandem with existing bracket systems and not replace them. The device produces cyclic forces to move teeth within the alveolus via accelerated bone remodelling [33]. Pavlin and co-workers in 2015 showed low-level cyclic loading with AcceleDent increased the rate of orthodontic movement (Figure 3) [34].
\nAcceleDent device.
Another treatment modality to speed up orthodontic tooth movement is by the use of low-level laser therapy (LLLT). Laser irradiation on tissues has a biostimulating effect with not more than 1°C rise in local temperature. Biostimulation potency of laser irradiation utilised by treatment are called low-level laser therapy [35]. Other than accelerating tooth movement, LLLT can enhance stability of orthodontic mini-implants [36], reduce post-adjustment pain [37], and induce bone growth in midpalatal suture area following rapid maxillary expansion [38].
\nStudies done by Fujita et al. and Yamaguchi et al. showed that LLLT enhances osteoclastogenesis on the compressed side of teeth being moved. There was stimulation of RANKL and macrophage colony-stimulating factor [39, 40]. Coordination of bone remodelling had been facilitated by RANKL and osteoprotegerin following orthodontic force with LLLT. LLLT stimulates bone formation on the tension side [41]. Kim et al. observed osteopontin localisation in the periodontal tissue in their study subjects, indicating LLLT may stimulate osteogenesis as well in orthodontic treatment [42]. Although much findings show LLLT stimulates osteoblast and osteoclast function, further studies are still required to optimise the effect of LLLT on tooth movement (Figure 4) [43].
\nLow-level laser therapy.
Apart from physical agents, low-intensity pulsed ultrasound (LIPUS) has also been suggested. It uses mechanical energy which passes through the tissues as acoustic pressure waves [44]. This leads to biochemical changes at molecular and cellular levels. It can increase the healing of both soft tissue and hard tissue [45]. LIPUS is usually used at frequency pulses of 1.5 MHz with 200 μs pulse width, which is repeated at 1KHz a for 20 minutes a day with an intensity of 30 mW/cm2 [46].
\nRecent studies on LIPUS using animal models by Xue et al. showed that there is induction of alveolar bone remodelling. The remodelling occurred due to an increase in the gene expression of HGF/Runx2/BMP-2 signalling pathway with LIPUS. This led to an increase in the velocity of tooth movement during orthodontic treatment [47]. El-Bialy et al. observed that LIPUS may reduce the root resorption that was orthodontically-induced by deposition of dentin and cementum to create a preventive layer from root resorption [48].
\nOver the years, the methods of reducing treatment time has risen along with its’ demand. The options that are available on the orthodontist’s plate are numerous ranging from surgical means to photostimulation. Much studies will still need to be done for newer methods to emerge and obtaining a clearer understanding on the methods that already exist. At present, the clinician should use all the knowledge obtained for deciding which treatment option is best for the patient to meet the healthcare needs of the patient and achieving an optimum treatment outcome.
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