Rifampicin dose for chemoprophylaxis [3].
\\n\\n
Released this past November, the list is based on data collected from the Web of Science and highlights some of the world’s most influential scientific minds by naming the researchers whose publications over the previous decade have included a high number of Highly Cited Papers placing them among the top 1% most-cited.
\\n\\nWe wish to congratulate all of the researchers named and especially our authors on this amazing accomplishment! We are happy and proud to share in their success!
\\n"}]',published:!0,mainMedia:null},components:[{type:"htmlEditorComponent",content:'IntechOpen is proud to announce that 179 of our authors have made the Clarivate™ Highly Cited Researchers List for 2020, ranking them among the top 1% most-cited.
\n\nThroughout the years, the list has named a total of 252 IntechOpen authors as Highly Cited. Of those researchers, 69 have been featured on the list multiple times.
\n\n\n\nReleased this past November, the list is based on data collected from the Web of Science and highlights some of the world’s most influential scientific minds by naming the researchers whose publications over the previous decade have included a high number of Highly Cited Papers placing them among the top 1% most-cited.
\n\nWe wish to congratulate all of the researchers named and especially our authors on this amazing accomplishment! We are happy and proud to share in their success!
\n'}],latestNews:[{slug:"stanford-university-identifies-top-2-scientists-over-1-000-are-intechopen-authors-and-editors-20210122",title:"Stanford University Identifies Top 2% Scientists, Over 1,000 are IntechOpen Authors and Editors"},{slug:"intechopen-authors-included-in-the-highly-cited-researchers-list-for-2020-20210121",title:"IntechOpen Authors Included in the Highly Cited Researchers List for 2020"},{slug:"intechopen-maintains-position-as-the-world-s-largest-oa-book-publisher-20201218",title:"IntechOpen Maintains Position as the World’s Largest OA Book Publisher"},{slug:"all-intechopen-books-available-on-perlego-20201215",title:"All IntechOpen Books Available on Perlego"},{slug:"oiv-awards-recognizes-intechopen-s-editors-20201127",title:"OIV Awards Recognizes IntechOpen's Editors"},{slug:"intechopen-joins-crossref-s-initiative-for-open-abstracts-i4oa-to-boost-the-discovery-of-research-20201005",title:"IntechOpen joins Crossref's Initiative for Open Abstracts (I4OA) to Boost the Discovery of Research"},{slug:"intechopen-hits-milestone-5-000-open-access-books-published-20200908",title:"IntechOpen hits milestone: 5,000 Open Access books published!"},{slug:"intechopen-books-hosted-on-the-mathworks-book-program-20200819",title:"IntechOpen Books Hosted on the MathWorks Book Program"}]},book:{item:{type:"book",id:"5747",leadTitle:null,fullTitle:"Nanowires - New Insights",title:"Nanowires",subtitle:"New Insights",reviewType:"peer-reviewed",abstract:"One-dimensional nanostructures, such as nanowires, have drawn extensive research interests in the recent years. 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It not only focuses on technological breakthroughs and roadmaps in implementing these technologies, but also presents the much-needed sharing of best practices, demonstrating the potential role of smart grid functions in improving the technical, economic, and environmental performance of modern power distribution systems. This can be achieved by allowing for massive pervasion of dispersed generating units, increasing the hosting capacity of renewable power generators, reducing active power losses and atmospheric emissions, and improving system flexibility.",isbn:"978-1-78923-892-1",printIsbn:"978-1-78923-891-4",pdfIsbn:"978-1-83968-006-9",doi:"10.5772/intechopen.77415",price:119,priceEur:129,priceUsd:155,slug:"research-trends-and-challenges-in-smart-grids",numberOfPages:170,isOpenForSubmission:!1,hash:"ca836c407ba574b88af44b497d45d42b",bookSignature:"Alfredo Vaccaro, Ahmed Faheem Zobaa, Prabhakar Karthikeyan Shanmugam and Kannaiah Sathish Kumar",publishedDate:"January 15th 2020",coverURL:"https://cdn.intechopen.com/books/images_new/7613.jpg",keywords:null,numberOfDownloads:3591,numberOfWosCitations:0,numberOfCrossrefCitations:3,numberOfDimensionsCitations:5,numberOfTotalCitations:8,isAvailableForWebshopOrdering:!0,dateEndFirstStepPublish:"September 4th 2018",dateEndSecondStepPublish:"September 25th 2018",dateEndThirdStepPublish:"November 24th 2018",dateEndFourthStepPublish:"February 12th 2019",dateEndFifthStepPublish:"April 13th 2019",remainingDaysToSecondStep:"2 years",secondStepPassed:!0,currentStepOfPublishingProcess:5,editedByType:"Edited by",kuFlag:!1,biosketch:null,coeditorOneBiosketch:null,coeditorTwoBiosketch:null,coeditorThreeBiosketch:null,coeditorFourBiosketch:null,coeditorFiveBiosketch:null,editors:[{id:"24725",title:"Dr.",name:"Alfredo",middleName:null,surname:"Vaccaro",slug:"alfredo-vaccaro",fullName:"Alfredo Vaccaro",profilePictureURL:"https://mts.intechopen.com/storage/users/24725/images/system/24725.jpeg",biography:"Alfredo Vaccaro received his MSc degree cum laude and commendation in Electronic Engineering from the University of\nSalerno and his PhD in Electrical and Computer Engineering from\nUniversity of Waterloo, Ontario, Canada. From March 2002 to\nOctober 2014 he has been Assistant Professor of Electric Power\nSystems at the Department of Engineering, Faculty of Engineering\nof the University of Sannio. From February 2011 to December\n2013 he was the Scientific Director of the Bureau of the Research Centre on Pure and\nApplied Mathematics at the Department of Engineering, University of Sannio. On\nOctober 2014 he obtained the National Scientific Qualification of Full Professor in\nElectrical Energy Engineering. Since November 2014 he has been Associate Professor\nof Electric Power Systems at the Department of Engineering of University of Sannio.\nHe is the Editor in Chief of Technology and Economics of Smart Grids and Sustainable\nEnergy, Springer Nature, Chair of the PES-IEEE Task Force on Enabling Paradigms\nfor High-Performance Computing in Wide Area Monitoring Protective and Control\nSystems, and Chair of the IEEE PES Awards and Recognition Committee.",institutionString:"University of Sannio, Benevento",position:null,outsideEditionCount:0,totalCites:0,totalAuthoredChapters:"1",totalChapterViews:"0",totalEditedBooks:"1",institution:{name:"University of Sannio",institutionURL:null,country:{name:"Italy"}}}],coeditorOne:{id:"39249",title:"Dr.",name:"Ahmed F.",middleName:null,surname:"Zobaa",slug:"ahmed-f.-zobaa",fullName:"Ahmed F. Zobaa",profilePictureURL:"https://mts.intechopen.com/storage/users/39249/images/system/39249.jpg",biography:"Ahmed Faheem Zobaa received his BSc (Hons), MSc, and PhD degrees in Electrical Power and Machines from Cairo University, Egypt, in 1992, 1997, and 2002, respectively. Also, he received his Postgraduate Certificate in Academic Practice from the University of Exeter, UK, in 2010, and his Doctorate of Science from Brunel University London, UK, in 2017. He was an instructor from 1992 to 1997, a teaching assistant from 1997 to 2002, and an assistant professor from 2002 to 2007 at Cairo University, Egypt. From 2007 to 2010, he was a senior lecturer in renewable energy at the University of Exeter, UK. Currently, he is a reader in electrical and power engineering, an MSc course director, and a full member of the Institute of Energy Futures at Brunel University London, UK. His main areas of expertise include power quality, (marine) renewable energy, smart grids, energy efficiency, and lighting applications. Dr. Zobaa is an executive editor for the International Journal of Renewable Energy Technology. He is also an executive editor-in-chief for Technology and Economics of Smart Grids and Sustainable Energy, and an editor-in-chief for the International Journal of Electrical Engineering Education. He is also an editorial board member, editor, associate editor, and editorial advisory board member for many international journals. He is a registered chartered engineer, chartered energy engineer, European engineer, and international professional engineer. He is also a registered member of the Engineering Council UK, Egypt Syndicate of Engineers, and the Egyptian Society of Engineers. He is a senior fellow of the Higher Education Academy of the UK and a fellow of the Institution of Engineering and Technology, the Energy Institute of the UK, the Chartered Institution of Building Services Engineers, the Institution of Mechanical Engineers, the Royal Society of Arts, the African Academy of Science, and the Chartered Institute of Educational Assessors. He is a senior member of the Institute of Electrical and Electronics Engineers. He is also a member of the International Solar Energy Society, the European Power Electronics and Drives Association, and the IEEE Standards Association.",institutionString:"Brunel University London",position:null,outsideEditionCount:0,totalCites:0,totalAuthoredChapters:"3",totalChapterViews:"0",totalEditedBooks:"5",institution:{name:"Brunel University London",institutionURL:null,country:{name:"United Kingdom"}}},coeditorTwo:{id:"30630",title:"Prof.",name:"Prabhakar",middleName:null,surname:"Karthikeyan Shanmugam",slug:"prabhakar-karthikeyan-shanmugam",fullName:"Prabhakar Karthikeyan Shanmugam",profilePictureURL:"https://mts.intechopen.com/storage/users/30630/images/system/30630.jpg",biography:"Prabhakar Karthikeyan Shanmugam completed his BE (EEE)\nfrom the University of Madras, Tamil Nadu (1997), his ME (Electrical Power Engineering) from the M.S. University of Baroda,\nVadodara, Gujarat (1999), and his PhD from VIT University, Tamil Nadu, India (2013) under the guidance of Prof. D. P. Kothari.\nHe also completed his Postdoctoral Fellowship from the Central\nPower Research Institute, Bengaluru, Karnataka, India. He is\npresently with VIT University as an associate professor. He is a Senior Member-IEEE\nwith 31 published articles in peer-reviewed journals and 77 works in national and\ninternational conferences. His area of interest includes deregulation and restructured\npower systems under smart grid environments, electric vehicles, and issues related to\ndistribution system studies.",institutionString:"Vellore Institute of Technology",position:null,outsideEditionCount:0,totalCites:0,totalAuthoredChapters:"2",totalChapterViews:"0",totalEditedBooks:"0",institution:{name:"Vellore Institute of Technology University",institutionURL:null,country:{name:"India"}}},coeditorThree:{id:"291294",title:"Dr.",name:"Kannaiah Sathish",middleName:null,surname:"Kumar",slug:"kannaiah-sathish-kumar",fullName:"Kannaiah Sathish Kumar",profilePictureURL:"https://mts.intechopen.com/storage/users/291294/images/system/291294.jpg",biography:"Dr. Kannaiah Sathish Kumar, PhD from VIT University, Vellore, India, has 14 years of total teaching experience and 9 years\nof research experience working in various research teams to\ndevelop new applications of evolutionary computing algorithms\nfor solving various power system problems such as unit commitment, economic dispatch, emission reduction, smart grids,\npower system reconfiguration, and restoration. His current area\nof interest is studying interconnection problems in linkages of HVDC and HVAC\n(765 kV) transmission lines with existing 230 kV high-voltage lines, nano-additives\nfor high-voltage XLPE cables, and optimization of smart grids. He is a member of\nIEEE, IEEE-PES, and SSI. He has published 51 research papers in different journals\nand conferences of international repute and authored a book on power quality. At\npresent he is working for the School of Electrical Engineering as an associate professor. He is a reviewer for various SCI journals such as Electrical Power and Energy\nSystems, ISA Transactions, AAI, etc.",institutionString:"Vellore Institute of Technology",position:null,outsideEditionCount:0,totalCites:0,totalAuthoredChapters:"0",totalChapterViews:"0",totalEditedBooks:"0",institution:null},coeditorFour:null,coeditorFive:null,topics:[{id:"770",title:"Renewable Energy",slug:"engineering-energy-engineering-renewable-energy"}],chapters:[{id:"67471",title:"Introductory Chapter: Open Problems and Enabling Methodologies for Smart Grids",slug:"introductory-chapter-open-problems-and-enabling-methodologies-for-smart-grids",totalDownloads:230,totalCrossrefCites:0,authors:[{id:"24725",title:"Dr.",name:"Alfredo",surname:"Vaccaro",slug:"alfredo-vaccaro",fullName:"Alfredo Vaccaro"},{id:"39249",title:"Dr.",name:"Ahmed F.",surname:"Zobaa",slug:"ahmed-f.-zobaa",fullName:"Ahmed F. Zobaa"},{id:"301321",title:"Dr.",name:"Antonio",surname:"Pepiciello",slug:"antonio-pepiciello",fullName:"Antonio Pepiciello"}]},{id:"65607",title:"Connected Autonomous Electric Vehicles as Enablers for Low-Carbon Future",slug:"connected-autonomous-electric-vehicles-as-enablers-for-low-carbon-future",totalDownloads:524,totalCrossrefCites:2,authors:[{id:"27900",title:"Dr.",name:"Hussein T.",surname:"Mouftah",slug:"hussein-t.-mouftah",fullName:"Hussein T. Mouftah"},{id:"277362",title:"Ph.D.",name:"Binod",surname:"Vaidya",slug:"binod-vaidya",fullName:"Binod Vaidya"}]},{id:"65420",title:"A Distributed Optimization Method for Optimal Energy Management in Smart Grid",slug:"a-distributed-optimization-method-for-optimal-energy-management-in-smart-grid",totalDownloads:834,totalCrossrefCites:1,authors:[{id:"28587",title:"Dr.",name:"Tatsuo",surname:"Narikiyo",slug:"tatsuo-narikiyo",fullName:"Tatsuo Narikiyo"},{id:"31886",title:"Prof.",name:"Michihiro",surname:"Kawanishi",slug:"michihiro-kawanishi",fullName:"Michihiro Kawanishi"},{id:"275140",title:"Dr.",name:"Dinh Hoa",surname:"Nguyen",slug:"dinh-hoa-nguyen",fullName:"Dinh Hoa Nguyen"},{id:"276982",title:"Dr.",name:"Tran Huynh",surname:"Ngoc",slug:"tran-huynh-ngoc",fullName:"Tran Huynh Ngoc"}]},{id:"66100",title:"Voltage Regulation in Smart Grids",slug:"voltage-regulation-in-smart-grids",totalDownloads:445,totalCrossrefCites:0,authors:[{id:"275562",title:"Dr.",name:"Maher",surname:"Azzouz",slug:"maher-azzouz",fullName:"Maher Azzouz"}]},{id:"67026",title:"A Reliable Communication Model Based on IEEE802.15.4 for WSANs in Smart Grids",slug:"a-reliable-communication-model-based-on-ieee802-15-4-for-wsans-in-smart-grids",totalDownloads:253,totalCrossrefCites:0,authors:[{id:"274565",title:"M.A.",name:"Jafar",surname:"Rasouli",slug:"jafar-rasouli",fullName:"Jafar Rasouli"},{id:"295377",title:"Mr.",name:"Ahmad",surname:"Motamedi",slug:"ahmad-motamedi",fullName:"Ahmad Motamedi"},{id:"305377",title:"Dr.",name:"Mohamad",surname:"Baseri",slug:"mohamad-baseri",fullName:"Mohamad Baseri"},{id:"306082",title:"Dr.",name:"Mahshad",surname:"Parsa",slug:"mahshad-parsa",fullName:"Mahshad Parsa"}]},{id:"69786",title:"Microgrid",slug:"microgrid",totalDownloads:431,totalCrossrefCites:0,authors:[{id:"300179",title:"Dr.",name:"Nithiyananthan",surname:"Kannan",slug:"nithiyananthan-kannan",fullName:"Nithiyananthan Kannan"}]},{id:"65727",title:"Solid-State Transformer for Energy Efficiency Enhancement",slug:"solid-state-transformer-for-energy-efficiency-enhancement",totalDownloads:632,totalCrossrefCites:0,authors:[{id:"279261",title:"M.Sc.",name:"Fernando",surname:"Vaca-Urbano",slug:"fernando-vaca-urbano",fullName:"Fernando Vaca-Urbano"},{id:"279479",title:"MSc.",name:"Manuel S.",surname:"Alvarez-Alvarado",slug:"manuel-s.-alvarez-alvarado",fullName:"Manuel S. 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From chapter submission and review, to approval and revision, copyediting 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:"3633",title:"Solar Energy",subtitle:null,isOpenForSubmission:!1,hash:null,slug:"solar-energy",bookSignature:"Radu D Rugescu",coverURL:"https://cdn.intechopen.com/books/images_new/3633.jpg",editedByType:"Edited by",editors:[{id:"8615",title:"Prof.",name:"Radu",surname:"Rugescu",slug:"radu-rugescu",fullName:"Radu Rugescu"}],productType:{id:"1",chapterContentType:"chapter",authoredCaption:"Edited by"}},{type:"book",id:"1288",title:"Solar Cells",subtitle:"Dye-Sensitized Devices",isOpenForSubmission:!1,hash:"05a255471069664ecf5fbf8778b92076",slug:"solar-cells-dye-sensitized-devices",bookSignature:"Leonid A. 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Zobaa"}],productType:{id:"1",chapterContentType:"chapter",authoredCaption:"Edited by"}}]},chapter:{item:{type:"chapter",id:"50834",title:"Molecular Mechanisms Involved in the Acquisition of Resistance to Treatment of Colon Cancer Cells",doi:"10.5772/63524",slug:"molecular-mechanisms-involved-in-the-acquisition-of-resistance-to-treatment-of-colon-cancer-cells",body:'\nThe inherent or developed resistance of many cancer cells to chemotherapy, targeted types of therapy, and irradiation is the primary cause for treatment failure in clinical oncology. Through basic research efforts aimed at gaining a better understanding of the mechanisms responsible for these effects, it is hoped that more effective strategies to manage this disease will be developed.
\nWnt signaling has been recognized as one of the most important contributors to malignant transformation in many types of solid tumors. Canonical Wnt signaling is altered in most cases of colorectal cancer (CRC). Indeed, a great amount of experimental evidence has shown that mutations in the adenomatous polyposis coli (APC) gene, a key negative modulator of canonical Wnt signaling, trigger the molecular pathogenesis of this type of cancer [1, 2]. The Wnt pathway has also been demonstrated to play an important role in the regulation of adult stem cell systems, and canonical Wnt signaling regulates the self-renewal and maintenance of both stem and cancer stem cells (CSCs). Cross talk has been reported between canonical Wnt signaling and hypoxia-inducible factor (HIF) signaling in tumor progression and metastasis [3]. However, the molecular mechanisms involved in this cross talk remain poorly understood.
\nDiminished oxygen availability (hypoxia) with the hypoxia-inducible factors (HIFs) mediating adaptation to it causes autophagy establishment, particularly in RAS-driven and BRAF-driven cancers, such as in most colorectal carcinomas. However, the relevance of the relation between HIFs and autophagy in drug resistance is not well understood. We have examined the effects of stable knockdown of HIF-1α or HIF-2α expression on malignant phenotype maintenance and in the canonical Wnt activation (β-catenin dependent). Our results indicated that although both HIF-1α and HIF-2α are essential for stemness and malignancy maintenance, these two proteins exert different effects and play opposing roles in canonical Wnt signaling [3]. We have also examined the effects of HIFs silencing on autophagy and drug resistance displayed by cancer cells. In agreement with other groups, we have observed that cancer cells exhibit high basal levels of autophagy and co-expression of HIF-1α and HIF-2α, compared with control nonmalignant cells and that the combination of mTOR inhibition with the autophagy inhibitor hydroxychloroquine (HCQ) dramatically induced cell death via apoptosis [4].
\nIn addition, it has been demonstrated recently that challenging cancer cells with stresses that they would typically encounter during tumor progression or as a part of a therapeutic regiment to treat or manage the disease, including chemotherapeutic agents, gamma irradiation, and hypoxic and serum-limiting conditions, increase the rate of microvesicles (MVs, referred also as oncosomes) formation and shedding by cells [5, 6]. These MVs are secreted from numerous types of cells and function in intercellular communication by transporting intracellular contents, such as protein, mRNA, and microRNAs (miRNAs) [7]. Oncosomes secreted by cancer cells may play an important role in cancer progression by promoting angiogenesis, neutrophil infiltration, and the education of bone marrow-derived cells [8]. Indeed, recent findings suggest that MVs can promote cell survival and contribute to drug resistance.
\nIn this review we focus on the molecular mechanisms whereby cancer cells develop resistance to antineoplastic drugs, particularly focused in the cell signaling pathways involved. A better understanding of the mechanisms that drive such resistance would enable the development of approaches to overcome it.
\nTargeted blockade of aberrantly activated signaling pathways is an attractive therapeutic strategy for solid tumors, but drug resistance is common. Several selective inhibitors, particularly against kinases and receptor tyrosine kinases (RTKs), have shown promising initial efficacy [9]; however, with few exceptions, the duration of response is limited; drug resistance rapidly emerges. This underscores the difficulty of successfully treating an adept, heterogeneous disease with a single targeted therapy and highlights the fact that monotherapy is often not a tractable long-term therapeutic approach.
\nResistance mechanisms can include alterations in the drug target itself, the pathway in which the target signals, or a parallel pathway that can alleviate the pressure on the cell due to blockade of the target [10]. Alternatively, drug resistance may be mediated by epigenetic reprogramming [11], by epithelial-to-mesenchymal transition (EMT) [12], or by emergence of a less differentiated, progenitor cell type [13]. Inducers of EMT, such as growth factors, transforming growth factor beta (TGFβ), and Wnt ligands, induces the expression of a gene program that leads to the suppression of the expression of the cell adhesion protein E-cadherin via the expression of transcriptional repressors such as Snail, Slug, Zeb1, Zeb2, and Twist. Besides these genes, other typical markers of EMT are N-cadherin, vimentin, and fibronectin-1, which are usually expressed in mesenchymal cells [3].
\nThe emergence of acquired resistance to targeted therapy against cancer is very common and the most frequent cause of treatment failure in cancer patients. This resistance can be mediated by signaling pathway reactivation [14] or by genetic or epigenetic events occurring within cancer cells [10]. Ultimately, these events lead to the activation of growth and survival signaling pathways within cancer cells that enable them to survive the stressful conditions. However, for most drugs, the identities of potential resistance pathways are unknown. The signaling pathways more frequently associated with cancer resistance to treatment are the following:
\nThe phosphoinositide 3-kinase (PI3K) signaling pathway, which lies downstream of various growth factor receptor tyrosine kinases, including the EGFR, is often aberrantly activated in human cancers. It plays critical roles in the regulation of cell growth, proliferation, differentiation, motility, survival, and intracellular trafficking. When deregulated, it is a major driver of tissue hyperplasia, oncogenesis [15], and is implicated in many aspects of tumorigenesis, including inappropriate cellular proliferation, angiogenesis, metastasis, and resistance to cell death.
\nThere are three classes of PI3K enzymes: the Class I PI3Ks play a central role in the transmission of regulatory signals through the metabolism-signaling supernetwork [15]. Class I PI3Ks are expressed as heterodimers consisting of four p110 catalytic subunits and one of two p85 regulatory subunits and are activated primarily by tyrosine kinase signaling pathways and heterotrimeric G protein-coupled signaling pathways. Upon activation, Class I PI3Ks preferentially phosphorylate PI-4,5-P2 to yield the second messenger PI-3,4,5-P3 [15]. The activation of PI3K activation is dampened by the lipid phosphatase PTEN, which dephosphorylates PI-3,4,5-P3 to regenerate PI-4,5-P2, thereby disengaging the proximal signaling proteins from the network. The local accumulation of PI-3,4,5-P3 at the inner leaflet of the plasma membrane attracts proteins containing pleckstrin homology (PH) domains, which bind PI-3,4,5-P3 and PI-3,4-P2 to act as proximal signal transducers in the PI3K pathway [15] (Figure 1).
\nThe PI3K and AMPK pathways are shown. Different members of the PI3K pathway increase apoptosis resistance. AMPK also regulates cellular metabolism by phosphorylating directly the tumor suppressor tuberous sclerosis 1 and 2 complex (TSC1/TSC2) to activate mTOR via Rheb GTPase.
The best-studied member of this set of PH domain-containing proteins is the serine-threonine kinase AKT, which is juxtaposed with its upstream activating protein kinase, PDK1. AKT is activated through phosphorylation of its threonine 308 residue by PDK1 and of serine 473 by mTOR complex 2 to mediate many PI3K responses, including growth, metabolism, survival, and glucose homeostasis [16]. AKT promotes apoptosis resistance by phosphorylating and thereby inhibiting proapoptotic substrates and profoundly influences cellular metabolism through its direct effects on metabolic enzymes and, more indirectly, through the stimulation of mTOR complex 1 (mTORC1) activity (Figure 1).
\nThe PI3K-AKT pathway is central to the integration of growth factor-derived signals and nutrient availability with anabolic metabolism, growth, and cell cycle progression in multicellular organisms. In diseases such as cancer, this pathway is reprogrammed to fuel stress resistance and uncontrolled growth and couples growth factors and other hormonal stimuli to the metabolic and autophagy networks.
\nWnt signaling is a key pathway in embryonic development and adult homeostasis and aberrant activation of this pathway plays an important role in the development of many human cancer types [1, 2]. Indeed, aberrant Wnt signaling is a hallmark of the majority of colorectal cancers and it is implicated in maintenance of tumor-initiating cells, drug resistance, tumor progression, and metastasis.
\nCanonical Wnt signals are transduced through Frizzled family receptors and LRP5/LRP6 co-receptor to regulate the phosphorylation and degradation of the transcription co-activator β-catenin (Figure 2). Noncanonical Wnt signals are transduced independently of β-catenin through Frizzled family receptors and ROR2/RYK co-receptors to the Rho family guanosine triphosphatases, c-jun-NH2-terminal kinase, or the Ca2+-dependent signaling cascades.
\nThe canonical Wnt pathway shown controls β-catenin intracellular levels and localization. When it is activated, β-catenin is stabilized and translocated to the nucleus to bind TCF transcription factor and activate Wnt-responsive genes. Noncanonical Wnt pathway transduces signals in a β-catenin-independent manner and its activation has been associated with aggressive malignant phenotype.
In the absence of Wnt ligands, β-catenin is assembled into the so-called destruction complex assembled by adenomatous polyposis coli (APC) tumor suppressor, axin, glycogen synthase kinase-3β (GSK-3β), and casein kinase 1 (CK1). This complex promotes phosphorylation of β-catenin that targets it for ubiquitination and subsequent proteolysis via the proteasome [2, 3]. Upon Wnt stimulation β-catenin breakdown is inhibited, thereby causing its accumulation and entry to the nucleus, to activate Wnt target genes [2]. In noncanonical Wnt pathways, Wnt signals are transduced independently of β-catenin. The best-studied noncanonical Wnt pathways are planar cell polarity (PCP) pathway and Ca2+ pathway, which play central roles in developmental morphogenesis, cell polarity, and cell migration. In Wnt/Ca2+ pathway, Wnt-Frizzled (Fzd) binding activates phospholipase C (PLC) via G proteins, producing an increase of intracellular Ca2+ concentration and the activation of downstream effectors including protein kinase C, as it can be seen in Figure 2 [1].
\nA great effort in developing selective drugs to target components of the Wnt pathway, particularly the β-catenin-dependent pathway, with anticancer activity, is underway but only a few of them have reached phase I clinical trials. In this respect, in models of KRAS-mutant colorectal cancer, it has been found by RNA-Seq data analysis for differential expression of canonical Wnt target genes that resistant cells display increased Wnt-β-catenin transcriptional activity [10], but there is also evidence, using this technique, that activation of noncanonical Wnt signaling exists in cancer-resistant cells and in different subsets of circulating tumor cells (CTCs) obtained from patients [17].
\nAs mentioned before, oncogenic drivers often elicit a strong tumor dependence on the pathway that the driver controls, leading to the so-called pathway addiction. One such oncogenic driver that elicits a pathway addiction is mutant KRAS which is directly implicated in the simplified linear RAS-RAF-MEK ERK (extracellular signal-regulated kinase) signaling axis, as well as the PI3K-mTOR axis [10]. Mutations within RAS-Raf-Mex pathway have frequently detected in colorectal cancer, being BRAFV660E the most frequently found (Figure 3).
\nThe Ras-Raf-MEK and NF-kB signaling pathways promote resistance. Mutations and amplification of Ras-Raf-MEK are frequently found in colorectal cancer allowing the development of resistance to treatment via autophagy activation. NF-kB upregulates Beclin 1 expression allowing positive regulation of autophagy. On the other hand, HIF-1α also modulates Beclin 1. IKK can promote the autophagy in an NF-kB-dependent manner.
Little et al. [18] and Corcoran et al. [19] examined the mechanisms whereby colorectal cancers develop resistance. Both studies focused on the effects produced by inhibition of MEK (mitogen-activated or extracellular signal-regulated protein kinase kinase), which is a downstream effector of the oncogene BRAF or KRAS [9]. Both groups found that resistance arose through amplification of the driving oncogene (BRAF or KRAS) rather than through amplification or mutation of the targeted kinase itself (MEK).
\nThere is also evidence that BRAF oncogene induces the expression of key autophagic markers, like microtubule-associated protein 1 light chain 3 (LC3) and Beclin 1 (BECN1) in colorectal tumor cells. Goulielmaki et al. [20] provided strong evidence that pretreatment with the autophagy inhibitor 3-methyl adenine (3-MA) followed by its combination with BRAFV600E targeting drug PLX4720 can synergistically sensitize resistant colorectal tumors.
\nAMPK is the central metabolic sensor activated by elevated AMP/ATP ratios. LKB1 is a human tumor suppressor kinase that is a crucial upstream molecule for the activation of AMPK and hence links cell metabolism to growth control and cell polarity [21]. During nutrient and energy depletion, ATP becomes depleted while the AMP/ATP ratio rises, activating the energy-sensing kinase, LKB1, which consequently activates AMPK. The AMPK regulates mTORC1 by direct phosphorylation of the tumor suppressor tuberous sclerosis complex 2 (TSC2) to activate mTORC1 via Rheb GTPase (Figure 1).
\nAberrant activation of the nuclear factor kappa B (NF-kB) family of dimeric transcription factors has been linked to most cellular processes in tumor evolution including inflammation, transformation, proliferation, invasion, metastasis, and chemoresistance [22]. The most common constitutively active form reported in human malignancies is the p50/RelA dimer, but other forms, such as p50/p50, p52/p52, p52/RelA, p50/c-Rel, c-Rel/c-Rel, p52/RelB, and p50/RelB, have also been identified [22]. In normal inactivated state, this transcription factor is sequestered in the cytoplasm by its inhibitor IkB (Figure 3). In order to be activated, IkB must be phosphorylated by the IkB kinase (IKK) complex and degraded via proteasome, causing the liberation and translocation of NF-kB into the nucleus where it modulates gene expression.
\nExperimental evidence demonstrates that NF-kB can positively regulate autophagy (Figure 3). For instance, RelA upregulates Beclin 1 expression through direct binding to the NF-kB-binding site on the Beclin 1 gene promoter to induce autophagy [23]. The multilevel control of autophagy by the IKK/IkB/NF-kB axis is also highlighted by the finding that IKK may also promote the autophagic pathway in a manner independent of NF-kB. All these findings support the idea that NF-kB activation promotes autophagy.
\nMetabolism is the process by which cells convert relatively simple extracellular nutrients into energy and building blocks necessary for their growth and survival. In cancer cells, metabolism is dramatically altered compared with normal cells.
\nMetabolic stress in tumors arises from multiple factors, including cell growth and proliferation in the setting of fluctuating supplies of oxygen and nutrients. Lack of adequate blood supply leads to several stress types in tumors: hypoxia, nutrient deprivation, and the accumulation of metabolic waste products [15]. These stress types are relieved in part by the over-expression of hypoxia-inducible factors (HIFs) to release proangiogenic factors for the construction of a tumor-associated vasculature. Persistent metabolic stress in tumors provokes the rewiring of the metabolic network to accommodate the stressful tumor microenvironment [17].
\nMetabolic reprogramming is a hallmark of cancer cells and is used by them for growth and survival. Their metabolism is highly dependent on glycolysis instead of mitochondrial oxidative phosphorylation, regardless of oxygen availability, a process termed as Warburg effect [15]. Glycolysis alone, although relatively inefficient means to produce ATP, provides a mechanism for rapid energy generation and a source of carbon for macromolecular synthesis. In addition, the shift to glycolytic metabolism allows rapidly proliferating cells to support both energy production and biosynthesis [15]. In addition, the glycolytic pathway generates metabolites that can be efficiently diverted into pathways that support nucleotide and amino acid biosynthesis in cycling cells [15].
\nOne interesting facet, from a chemotherapeutic point of view, is that, in a large number of cases, cells undergoing a Warburg effect exhibit a marked dependence upon glutamine, to the extent that these cells are referred to as being “glutamine addicted” [24]. Glutamine addiction arises from the need for extracellular glutamine to be consumed for anaplerotic input in the citric acid cycle, which accounts for the majority of the bioenergetic needs of normal (nontransformed) cells. Cancer cells rely heavily on glutamine as a source of carbon and nitrogen for the synthesis of ATP, proteins, lipids, nucleic acids, and the antioxidant glutathione. Expression of the Wnt target MYC proto-oncogene increases glutamine uptake by stimulating expression of the glutamine transporters. At the same time, MYC increases the levels of glutaminase 1 (GLS1), the enzyme that converts glutamine to glutamate [15, 24].
\nDuring oncogenesis, the survival signaling pathways such as PI3K, intermediate metabolism, and autophagy constitute the metabolism-signaling supernetwork reprogrammed in ways that support aberrant cell growth, proliferation, and stress resistance. Indeed, emerging evidence suggests that autophagy is an important source of amino acids that supports the stressed cell’s biosynthetic and bioenergetic needs.
\nDiminished oxygen availability (hypoxia) is a hallmark of the tumor microenvironment. A major regulator of cellular adaptation to hypoxia is the hypoxia-inducible factor (HIF) family of transcription factors, which play key roles in many crucial aspects of cancer biology including angiogenesis, stem cell maintenance, metabolic reprogramming, resistance to apoptosis, autocrine growth factor signaling, the epithelial-mesenchymal transition (EMT) program, genetic instability, invasion, metastasis, and radiation resistance [25] (Figure 4). HIFs also cause autophagy establishment, particularly in RAS-driven and BRAF-driven cancers, such as most colorectal carcinomas (Figure 4). However, the relevance of the relation between HIFs and autophagy in drug resistance is not well understood.
\nHIF target genes. The cellular processes regulated by HIF-1α and HIF-2α genes are indicated and also the target genes for each one.
HIFs are heterodimeric transcription factors consisting of an O2-sensitive subunit HIF-α and a stable subunit HIF-β (or ARNT) that are expressed constitutively at the transcriptional and translational levels (Figure 5). In mammals, three HIF-α isoforms have been identified but HIF-1α and HIF-2α are the two best-studied members of the HIF-α family. Under normoxic conditions, the cellular stability and activity of HIF-α subunits are highly dependent on oxygen supply. Prolyl hydroxylases hydroxylate key proline residues on HIFs, which allows them to interact with the von Hippel-Lindau (pVHL) tumor suppressor, which is a component of an E3 ubiquitin ligase complex that targets HIF-α for proteasomal degradation (Figure 5). Hypoxic conditions stabilize HIF-α by inhibiting its hydroxylation and proteasomal degradation, making HIF-α capable to translocate to the nucleus and dimerize with ARNT activating the transcription of hypoxia-associated genes [25, 26]. Importantly, high levels of HIFs expression have also been detected in tumor cells in the absence of hypoxia (Figure 5) because the sustained oncogenic signaling mediated by growth factors in cancer cells can induce HIF-α expression through O2-independent mechanisms, including increased transcription and/or translation of HIF-α mRNA [27, 28]. In this respect, we have reported that colon carcinoma cells co-express HIF-1α and HIF-2α under normoxic conditions, in contrast to nonmalignant colon cells, which do not express these factors under these conditions [3].
\nHIF activation. HIFs are heterodimeric transcription factors subjected to O2-sensitive or O2-independent mode of activation, as indicated in the figure. Both types of regulation converge in HIF protein stabilization, which can enter to the nucleus to bind HIF-1β at HIF-responsive elements (HRE) to activate target genes.
Hypoxia can lead to therapeutic resistance through diverse mechanisms [29, 30]: (1) direct effects due to the requirement of some drugs and radiation of lack of oxygen in order to be maximally cytotoxic, (2) indirect effects by altering cellular metabolism to decrease drug cytotoxicity, and (3) enhanced genetic instability which in turn leads to more rapid induction of drug resistance in tumor cells [29, 30]. HIF-1α and to a lesser extent HIF-2α have been associated with radio- and chemotherapy failure for decades [30] and interference with HIF function holds great promise to improve future anticancer therapy.
\nOne of the possible mechanisms of resistance to anticancer therapy in hypoxic tumor is the switching of cellular metabolism from oxidative phosphorylation to aerobic glycolysis (Warburg effect), a hallmark of cancer cells. As a result, cancer cells eventually develop a system that uses cytoplasmic glycolysis to generate ATP instead of mitochondrial oxidative respiration, even in the presence of oxygen. Several reports have indicated that activation of the HIF-1α signaling pathway under glucose deprivation induces resistance to cell death by apoptosis in human colon cancer cells and that targeting the HIF-1α signaling pathway may provide an effective way to treat resistant cancers to conventional therapy [31].
\nHIF-1α upregulation has been shown to induce expression of drug efflux transporters, to alter the activity of DNA repair mechanisms, and to shift the balance between pro- and antiapoptotic factors toward cell survival [32].
\nDrug efflux is an important mechanism to chemoresistance in many solid tumors, including colon cancer. The multidrug resistance 1 (MDR1) gene, encoding the membrane-resident P-glycoprotein (P-gp) that belongs to a family of ATP-binding cassette (ABC) transporters, has been found to be a HIF-1α target gene [33]. The multidrug resistance-associated protein 1 (MRP1) is another ABC transporter encoded by the ABCC1/MRP1 gene that confers cellular resistance to a broad range of structurally and functionally chemotherapeutic agents. Recently, it has been demonstrated that MRP1 is a downstream target gene of HIF-1α in human colon cancer LoVo cells. Genetic inhibition of HIF-1α by siRNA and dominant-negative HIF-1α reduced the expression of MRP1, which provides a potential novel mechanism for HIF-1α-mediated drug resistance [34].
\nDefective apoptosis and/or changes in cell cycle regulation represent pivotal causes for drug resistance [35]. It has been proposed that increased cell survival, due to a shift favoring antiapoptotic pathways, is a primary mechanism of hypoxia-induced drug resistance [36]. In the vast majority of transformed cells, HIF-1α functions as a suppressor of apoptosis and functional interference with HIF-1α results in enhanced cell death upon treatment with chemotherapeutic agents in tumors of different origins [26]. Through the inhibition of proapoptotic and induction of antiapoptotic genes, HIF-1α can inhibit apoptosis and promote tumor cell survival in chemotherapy-treated cancer cells.
\nIn colorectal cancer, the combination therapy of rapamycin, inhibitor of mTOR which regulates autophagy, and irinotecan, which is able to inhibit the accumulation of HIF-1α, has been reported to induce massive death of colon cancer cells under hypoxic, but not normoxic conditions in vitro, and a great reduction of tumor volume in vivo [37].
\nEnhanced autophagy has been associated with the elevated level of HIF-1α in several cancer types. In this regard, it has been observed that hypoxia-mediated failure of cytotoxic treatment in vitro can be conferred via HIF-1α-dependent induction of autophagy [14], but the relevance of the intriguing relation between HIF-1α and autophagy for drug resistance is still not known.
\nAutophagy is a term derived from the Greek words “auto” (self) and “phagy” (to eat) and refers to a multistep lysosomal degradation process in which a cell degrades damaged organelles and longlived proteins to maintain cellular homeostasis, particularly during exposure to stressful conditions. Three forms of autophagy have been identified based upon the mode of delivery to the lysosome, namely, macroautophagy, microautophagy, and chaperone-mediated autophagy. Macroautophagy (autophagy), the best characterized, is a major regulated catabolic process that involves the delivery of cytoplasmic cargo sequestered inside double-membrane vesicles to the lysosome [38]. The other two forms, microautophagy and chaperone-mediated autophagy, involve a direct membrane invagination to engulf damaged proteins and the translocation of soluble cytosolic proteins by chaperone-dependent selection across the lysosomal membrane, respectively [39]. Thirty-six genes (ATGs for AuTophaGy), have been identified that are required for the autophagy process to occur. Dysregulation of autophagy, which alters the rate of protein degradation and the metabolic state of the cells, has severe consequences and is associated with several pathophysiological conditions, such as cancer.
\nThe best-characterized regulator of autophagy is mTOR, which integrates growth factor and nutrient signals to influence protein synthesis, growth, autophagy, and ribosomal biogenesis. M-TOR occurs in two multiprotein complexes, mTORC1, containing the specific binding proteins raptor and PRAS40, and mTORC2 containing rictor and other binding partners [38]. When nutrients are available, mTORC1 phosphorylates Unc-51-like kinase (ULK1) and ATG13 to block autophagy initiation. When nutrients are scarce, mTORC1 dissociates from the ULK1 complex, initiating the autophagy process [15, 40]. The initiation of autophagy occurs with the assembly of the double-membrane phagophore, the precursor of the autophagosome. This requires the formation of a protein complex constituted by Class III phosphatidylinositol 3-kinase (PtdIns3K) and the proteins VPS34, p150, ATG14, and BECLIN 1. In tumors, autophagy is stimulated by metabolic stress (e.g., nutrient/growth factor deprivation, hypoxia, and acidosis), cellular damage, or inhibition of pro-survival signals caused by anticancer therapies [41]. Through autophagy, cancer cells utilize a highly plastic and dynamic mechanism to either repress initial steps of carcinogenesis or support the survival and growth of established tumors [42, 43].
\nIn multicellular organisms, autophagy also clears ubiquitinated or malfunctioning aggregated proteins. This selective degradative process is mediated through the recognition of ubiquitin-tagged cargos by the autophagy receptor p62/sequestosome 1 [44]. This protein directs them to the autophagosome through concomitant binding to the LC3 (microtubule associated protein 1 light chain 3) molecules localized at the inner and outer autophagosome membranes. Autophagy is responsible for the degradation of p62, and therefore, when autophagy is inhibited, p62 accumulates in mammalian cells.
\nAutophagy facilitates cancer cell resistance to chemotherapy treatment, and the inhibition of autophagy may potentiate the resensitization of therapeutic-resistant cancer cells to anticancer drugs. Chemotherapy treatment conferred resistance by triggering key autophagy signaling molecules in malignant cells. For example, in response to irinotecan, pro-survival autophagy was induced by activating MAPK14/p38 signaling, which lead to drug resistance [45]. It is likely that protective autophagy in 5-Fluouracil (5-FU) resistance occurs through c-Jun N-terminal Kinase (JNK) activation [46]. Given the propensity of PI3K-mTOR inhibition to robustly induce autophagy, increased autophagic flux is a suspect contributor to the modest efficacy of PI3K inhibitors. Several recent studies have provided evidence that combined inhibition of autophagy and PI3K inhibitors or a combination therapy with a mTORC1 inhibitor (temsirolimus) and autophagy inhibitor [hydroxychloroquine (HCQ)] can sensitize cells to chemotherapeutic agents [15, 47]. Several other trials that combine HCQ with radiation therapy or chemotherapeutic agents have also been used on the basis of the accumulating preclinical evidence supporting the notion that increased autophagy promotes therapy-induced resistance in tumor cells.
\nExamining the role of autophagy in RAS- and BRAF-induced transformation in colon cancer cell lines, Goulielmaki et al. [20] found that the MEK/ERK pathway can increase the protein levels of LC3, unlike the AKT/MTOR pathway, which has been shown to abolish the autophagic process. They showed that using specific autophagy inhibitors not only cancer cell proliferation rate can be reduced, but the otherwise resistant mutant BRAF colon cell lines to targeted BRAF agents, like PLX4720 (Vemurafenib), can be sensitized to apoptosis in a synergistic manner. This study proposed a promising rational combinatorial treatment using BRAF and autophagy inhibitors that will potentially provide efficient therapeutic protocols for these otherwise untreatable tumors.
\nThe intestinal epithelium is an example of self-renewing tissue on expense of stem cells that reside at the crypt bottom. Tumor growth is sustained by a subpopulation of highly malignant cancer stem cells (CSCs) or tumor-initiating cells, which are characterized by a life-long capacity to self-renew, are multipotent, and can reversibly enter quiescent or even dormant states and resist cytotoxic drugs [48]. Successful treatment is thus dependent on the selective elimination of these highly resistant subpopulations, instead of only the main tumor mass. Over the past decade, several cell surface markers have been identified in these cell populations. CD133, Lgr5, and CD44 are the most frequently proposed stem cell markers in colorectal cancer, but their distribution differs between patients and tumor cell lines [48, 49].
\nCD133 (also called prominin-1) is a pentaspan transmembrane glycoprotein identified as cell surface stem cell marker that has been associated with tumorigenicity and progression of colon cancer [50], but its precise role and functions are unknown. CD44 is a transmembrane glycoprotein involved in cell-cell and cell-matrix adhesion through its affinity for hyaluronic acid. CD44 is encoded by a single gene, including 20 exons. The standard form, expressed in normal adult stem cells (referred to as CD44), consists of exons 1–5 and 15–20. Importantly, it has been demonstrated that cancer cells express different exon variants, such as CD44v6, produced by alternative splicing mRNA processing [48]. Thus, universal targeting of CD44 might be deleterious for patients and they can be avoided targeting different isoforms of CD44.
\nSeveral studies have implicated the potential contribution of a subpopulation of stem-like progenitor cells in resistance to both chemotherapeutic and targeted therapies [51]. Various groups have characterized the drug-resistant aspects of such stem-like subpopulations, including a quiescent state refractory to agents targeting rapidly dividing cells, enhanced DNA damage repair mechanisms, and decreased apoptotic machinery [52]. Recent studies have implicated a potential mechanistic link between EGFR activation and the acquisition of stem-like properties including the increase in known stem cell markers and enhanced spheroid formation [53]; however, the role of EGFR in promoting stem cell properties in CRC has not been fully characterized.
\nThe radioresistance of cancer stems cells has been supported by several research groups in glioma and head and neck, breast, pancreatic, and colorectal cancer [48, 54]. Radiation itself has also been shown to increase the expression of AKT and CD133 and reduce the expression of CD44 in colorectal cancer cells [55]. Sahlberg et al. [48] found that cells with a CD44high/CD133high expression demonstrated a higher radioresistance compared to CD44low/CD133low cells and that different AKT isoforms have varying effects on the expression of cancer stem cell markers, which is an important consideration when targeting AKT in a clinical setting.
\nThere is experimental evidence that hypoxia is associated with the maintenance and formation of CSCs, promoting their phenotype and tumorigenesis [56]. It has also been shown that hypoxia, by means of HIF-1α activation, is capable of maintaining CSCs phenotype in colon cancer cells [57]. However, it has been recognized that the resident microenvironment of cancer cells, also known as a “niche,” plays an important role in the genetic instability, metastasis, and therapeutic resistance of CSCs [25]. Mao et al. [58] demonstrated that most CD133+ colon CSCs are located in a hypoxic niche, where oxaliplatin, rather than 5-FU, inhibits proliferation of these colon CSC cells. Recently several drugs have discovered to be selective against CSC. Examples of them are microbe-derived and plant-derived biomolecules; small inhibitors that target key signaling pathways of CSCs such as metformin, tranilast, and thioridazine; and also antibodies directed against CSC-specific cell surface molecules, such as the CD44, CD47, EpCAM, CD123, GD2, Lgr5, IGF-IR, Dll4, and FZD receptors [59].
\nIt has been demonstrated recently that challenging cancer cells with stresses that they would typically encounter during tumor progression, or as a part of a therapeutic regiment to treat or manage the disease, increase the rate of microvesicles (MVs referred also as oncosomes or exosomes), formation, and shedding by cells [5–7]. Interestingly, the noncanonical Wnt signaling pathways PCP and Wnt/Ca2+ appear to be involved in the regulation of MV biogenesis and budding in human cancer cells, since one of their downstream effectors, the Rho subfamily GTPase, that induce cell spreading and migration, has been demonstrated to promote the rearrangements of the actin cytoskeleton to stimulate MV budding [60, 61].
\nMVs generally range in size from 0.1 to 2 μm in diameter. In addition to containing conventional paracrine signaling molecules, such as growth factors and pro-inflammatory cytokines, MVs also contain membrane-associated, cytosolic, and nuclear molecules not normally released by normal cells such as metabolic enzymes, metalloproteases, molecular chaperones, and miRNAs and RNA transcripts [6, 7]. The uptake of MVs by cells has the potential to protect them from a variety of apoptotic challenges by up-regulating the expression and/or activation of proteins that work to counter the actions of cell death machinery. That is why recent findings suggesting that MVs can promote cell survival and contribute to drug resistance are possibly of significant value.
\nIn addition, because cancer cell-derived MVs often contain oncogenic proteins that reflect their cell of origin, and their abundance and protein concentration correlate with the tumor grade/aggressiveness, they have been converted in the focus for searching cancer biomarkers and/or for monitoring tumor progression.
\nMammalian cells coordinate cell metabolism and growth with environmentally induced stress. In cancer cells, survival signaling cascades, metabolism, and autophagy are corrupted and work in a highly integrated network to cope with stressful conditions such as hypoxia, limited nutrients, and drugs, in order to maintain viability and proliferative activity.
\nUnderstanding the signaling networks that contribute to cancer cell survival and how the changes in those networks allow cells to adapt in the presence of drugs that target key components implicated in cancer cell survival and proliferation is the challenge. Thus, we are hopeful that a better understanding of the signaling pathways involved and the development of strategies to inhibit autophagy and/or hypoxia represent a new approach to enhance the efficacy of cancer therapy to overcome therapeutic resistance in cancer cells.
\nMycobacterium leprae (M. leprae) is an acid fast bacilli that is the causative agent of leprosy disease which mainly effects the skin and peripheral nerves. In olden times leprosy was common in temperate climates (e.g. Europe), today it is mainly confined to tropical and subtropical regions. Mode of transmission in leprosy is mainly through inhalation of droplets containing the bacteria. But skin contact is also claimed by many leprologists. The disabilities and deformities associated with leprosy due to neuropathy leads to long-term consequences, including. This in turn is associated with stigma.
The immunity of the host plays an important role in disease progress and control. Thus, fortunately 95% of patients exposed to M. leprae will not develop this disease. The variation in incubation period ranges from 2 to 20 years, or even longer.
Leprosy has been successfully eliminated as a public health problem in 2000 globally and at the national level in 113 countries out of 122 by 2005 [1]. Elimination of leprosy is defined by World Health Organization as a point prevalence below 1 per 10,000 population [2]. However, the number of new patients diagnosed with leprosy is still significant, at more than 200,000 in 2016 globally. The new case detection rate of the disease (NCDR) is only slowly declining (Figure 1) [3].
Trend in case detection and case detection rate, by WHO region, 2006–2016 [3].
The long incubation period, silent symptoms, long duration MDT and unavailability of effective vaccine makes this disease difficult to identify, treat and eradicate. To add to the misery the stigma associated with the disease is another challenge. In such circumstances, prevention and control of disease gains utmost importance.
In 2017, 192,713 patients were on treatment globally which makes the prevalence rate of 0.25 per 10,000 population [4]. Total of 210,671 new cases were reported in same year from 150 countries making NCDR of 2.77 per 100,000 population. Figure 2 below shows the trends over the past decade (2008–2017) in new case detection of leprosy cases globally in the reporting countries of World Health Organization (WHO) [4].
Country-wise trends of detection of new leprosy cases from 2008 to 2017 [4].
The three main goals of control of leprosy are
To detect the pathology early and treat the patient completely.
To prevent the transmission to the others.
To prevent the disabilities and other complications.
Thus the following modalities are adopted to control leprosy:
Medical measures
Social support
Program management
Evaluation
The control of leprosy starts with the estimation of size and magnitude of the problem. Most common epidemiological survey method of collection of data is “Quick random sample survey.” Information about the prevalence of leprosy, age and sex-wise distribution, various forms of leprosy and the health facilities available should be gathered. Roughly the total prevalence of leprosy in an area would be about 4 times that of the cases found among school children [5, 6]. These estimates are essential to plan, implement and to evaluate the results of the control program.
The objective is to detect all the cases as early as possible and to register them. Active case finding is important as the disease is symptomless in the early stages. Cases can be detected by the Contact surveys, Group surveys and Mass surveys. Contact surveys consists of examination of all household contacts with a lepromatous case, particularly children, in areas with prevalence less than 1 per 1000. Contact surveillance of households is recommended for a minimum period of 10 years after case is declared bacteriologically negative, and for 5 years in households with a non-lepromatous case from the time of diagnosis of the index case. Group surveys are done in areas where prevalence of leprosy is more than 1 in 1000 population. This consists of screening certain groups such as school children, slum dwellers, military recruits, industrial workers, etc. through “Skin camps.” Lastly, mass surveys consists of examination of each and every individual by house-to-house visits in hyperendemic areas (prevalence – 10 or more per 1000 population). These are generally carried out by repeated annual examinations of school children which yield better results at relatively low cost [5, 6]. The data of each case is entered in the standardized proforma developed by WHO.
Since an effective vaccine is unavailable for leprosy the secondary prevention (early treatment) becomes more important. Until 1981, Dapsone (Diamino Diphenyl Sulphone—DDS) was used to treat leprosy which resulted in the development of resistance and relapse, making leprosy control difficult.
Multidrug Therapy: In 1982, WHO recommended Multidrug Therapy (MDT) for all leprosy patients. Introduction of MDT has opened a new avenue in the control of leprosy in the world. Aim of MDT is to convert the infectious case into noninfectious as soon as possible, so as to reduce the reservoir of infection in the community.
The main objectives of MDT are:
To ensure early detection of the cases.
To interrupt the transmission of infection.
To prevent drug resistance, relapse and reaction.
The advantages of MDT over dapsone monotherapy are:
Shorter duration of treatment,
Better patient compliance,
High cure rate,
Cost-effectiveness and
Ease in health delivery system.
There are two types of MDT regimens used depending on the symptoms and signs shown by the patients - Paucibacillary (PB) and Multibacillary (MB). Recommended Regimens are discussed below [3, 5, 6, 7]:
i. Multibacillary leprosy:
MDT is recommended for following groups of patients:
All smear positive cases.
Skin lesions more than five in number.
More than one nerve trunk thickening.
All cases of relapse/reactivation and all cases who have been treated with Dapsone monotherapy earlier.
The drugs used in Multibacillary MDT and dosages are:
Rifampicin: 600 mg once monthly, supervised.
Dapsone: 100 mg daily, self administered.
Clofazimine: 300 mg once monthly, supervised and 50 mg daily, self administered.
Duration of treatment for Multibacillary leprosy is 12 months, can be extended to 18 months and continued where possible up to smear negativity. Sometimes LL/BL patients with high bacilli may need 2–3 years or more of MDT for achieving bacteriological negativity.
ii. Paucibacillary leprosy:
The drugs and dose schedule is:
Rifampicin 600 mg once a month for 6 months supervised.
Dapsone 100 mg daily for 6 months self administered.
Paucibacillary leprosy is treated for 6 months.
MDT is not contraindicated in patients with HIV infection.
Each MDT blister pack contains tablets for 4 weeks treatment. For easy identification color coding of the blister pack is done, that is, with different colors for multibacillary and paucibacillary cases both in adults and children.
The treatment in both PB and MB cases varies depending on the age of the patient. The patients between 10 to 14 years are treated as paediatric cases, while >14 years are considered adult. The standard treatment regimen for MB leprosy in adults is given for 12 months. The drugs in each blister pack are (Figure 3):
Two capsules of Rifampicin of 300 mg (600 mg once a month) to be taken as single dose under supervision.
Clofazimine 3 capsules of 100 mg each to be consumed once a month as single dose under supervision and 50 mg daily for next 28 days.
Dapsone 100 mg as single dose and then daily once for 1 month.
MDT for adult MB type of leprosy [2, 7].
The standard adult treatment regimen for PB leprosy is (Figure 4):
Rifampicin: 600 mg once a month.
Dapsone: 100 mg daily.
Duration: 6 months (6 blister packs of 28 days each).
MDT for adult PB type of leprosy [2, 7].
Treatment regimen for MB leprosy in children (ages 10–14 years) is (Figure 5):
Rifampicin: 450 mg once a month.
Clofazimine: 150 mg once a month, and 50 mg every other day.
Dapsone: 50 mg daily.
Duration: 12 months (12 blister packs of 28 days each).
MDT for pediatric MB type of leprosy [2, 7].
Treatment regimen for PB leprosy in children (ages 10–14 years) is (Figure 6):
Rifampicin: 450 mg once a month.
Dapsone: 50 mg daily.
Duration: 6 months (6 blister packs of 28 days each).
MDT for pediatric PB type of leprosy [2, 7].
MDT is provided free-of-charge globally through an agreement between a pharmaceutical company and WHO. WHO manages distribution of MDT to countries in coordination with national leprosy programs.
Clinical surveillance of the patients after completion of treatment is an important part of MDT to ensure complete cure. For paucibacillary cases follow up for at least once a year for 2 years after completion of treatment and for multibacillary cases at least once a year for 5 years [3, 4, 5].
Early diagnosis of cases, aggressive treatment and proactive measures to avoid complications and disabilities is the backbone for the success of any comprehensive program. In addition to accurate reporting and control measures, effective preventions will be needed to achieve elimination. Search for an effective vaccine either to be used alone or in combination with a drug has been going for a long time.
Presently BCG (Bacillus Calmette-Guerin) is the only vaccine that has shown some protection against M. leprae bacillus. A single dose of BCG gives 50 percent or higher protection against the disease. It is the most widely used vaccine in the world, yet the degree of protection it confers is not yet confirmed. The meta-analysis of many experimental studies concludes that the vaccine gives approximately 26% protection against leprosy. But the protection level decreases with time. To overcome this problem more than one dose of vaccine is advised.
Other variants of vaccination are also suggested.
Adding killed M. leprae to BCG: Various modifications have been suggested, such as the addition of killed M. leprae to BCG. This method almost doubles the vaccine efficacy in some populations as concluded by few studies. But the same cannot be said for patients below 15 years.
Vaccination with M. indicus pranii (Mycobacterium W): This strain discovered in India. Testing of the MIP vaccine took place in 2005 and showed that it was effective for seven to 8 years, after which a booster dose would be needed to maintain the immunity. Recently the vaccine was approved by the Drug Controller General of India to be rolled out in a project involving five districts in the states of Bihar and Gujarat, where there are high rates of leprosy. Leprosy patients and their close contacts will benefit from this project, making India the first country in the world to have a large-scale leprosy vaccination initiative [8].
Another milestone in prevention of leprosy is the discovery of the vaccine candidate, called LepVax. Scientists at Infectious Disease Research Institute (IDRI), along with national and international collaborators including the National Hansen’s Disease Program and the National Institute of Allergy and Infectious Diseases (NIAID), part of the National Institutes of Health, with financial support from American Leprosy Missions, have developed this leprosy vaccine. Based on the preclinical studies, the LepVax, has progressed to Phase I clinical testing in the United States, the first stage of safety testing in human volunteers. The clinical trial is focused not only on safety but also evaluates the immune response of the individual to the vaccine.
Indian cancer research center (ICRC) bacilli: Another variant belonging to the M. avium intracellulare group, the ICRC bacilli are thought to induce lepromin conversion in lepromatous leprosy patients and in lepromin-negative leprosy-free individuals. Its efficacy was reported to be 65.5 percent [8].
M. vaccae: The studies with this soil-dwelling mycobacterial species combined with BCG showed to provide greater protection against leprosy, but a Vietnamese trial contradicted the results [8].
M. Habana: This bacilli has been reported to induce lepromin conversion when used as a live vaccine in monkeys, and protected mice against the development of leprosy [8].
Chemoprophylaxis alone provides two-year protective window while effective immunization will provide a much broader protective window. Thus many studies and research is going on to provide both chemoprophylaxis and immunization for immediate and short-term protection and longer-term protection respectively. This strategy could have better impact and distinct appeal in controlling and preventing leprosy. Such trials could also provide a gateway for the assessment and implementation of new emerging vaccines (Figure 7).
Locations of leprosy vaccine testing.
Chemoprohylaxis using effective antibiotics focuses on providing protection to people at risk such as close contacts – family members, neighbors, co-workers, health care providers for lepers etc. Due to the stigma of disease the leprosy cases are found in clusters in all endemic regions, rather than being evenly dispersed over the whole area. Thus these high risk people can be identified and prophylaxis provided along with secondary prevention strategies. The process includes focused surveillance, contact tracing, early diagnosis and treatment. This helps in reducing the incidence and breaking the chain of transmission.
Chemoprophylaxis, as recommended by WHO Guideline Development Group (GDG), is done using single dose rifampicin (SDR) for contacts of leprosy patients both in adults and children of 2 years of age and above. Before starting the drug leprosy and TB disease are to be excluded. There should be no contraindications also for the use of rifampicin.
Other important considerations for the implementation of this chemoprophylaxis by programs are:
Adequate management of contacts.
Consent of the index case to disclose his/her disease.
An RCT found that SDR reduces risk of leprosy over 5–6 years in leprosy contacts. For every 1000 contacts treated with SDR, there were four leprosy cases prevented after 1–2 years and three cases prevented after 5–6 years.
Recommended dosage schedules for SDR are given in Table 1.
Age/weight | Rifampicin single dose |
---|---|
Adults (≥15 years) | 600 mg |
10–14 years | 450 mg |
Children 6–9 years (weight ≥ 20 kg) | 300 mg |
Children <20 kg (≥2 years) | 10–15 mg/kg |
Rifampicin dose for chemoprophylaxis [3].
The limitations of this approach are:
The protection is approximately for only 2 years.
High bacillary load cannot be eliminated using single dose.
Specific screening test needed to distinguish between contacts with high and low bacillary load.
Among communicable diseases, leprosy remains a leading cause of peripheral neuropathy and disability in the world, despite extensive efforts to reduce the disease burden. It is an important aspect of leprosy control. It means the medical, surgical, social, educational, and vocational restoration as far as possible of treated patients to normal activity so that they resume their place in the home, in society and industry [5, 6, 7]. Early treatment helps in disability limitation.
Rehabilitation: WHO has defined rehabilitation as “the combined and coordinated use of medical, social, educational and vocational measures for training and retraining the individual to the highest possible level of functional ability.”
Preventive rehabilitation consists of prevention of development of disabilities in a leprosy patient by early diagnosis and prompt treatment. But once the patient becomes handicapped and suffers from the damage caused, should be trained and retrained to the maximum functional ability so that the patient becomes useful to self, to the family and to community at large by various measures such as medical (physical), surgical, psychological, vocational and social rehabilitation (Flow chart 20.10).
Health education is given to the patient, to the family and to the community at large about leprosy. The education should be directed to ensure general public and patients help them develop their own actions and efforts to change the perception about the disease and seeking professional help whenever required. Early recognition of symptoms, prompt diagnosis, health seeking behavior, personal care, treatment adherence and rehabilitation are important aspects of health education. The key messages included are about the cause of disease and the complete cure available to encourage people for early diagnosis and treatment. It also aims at helping people to change their attitude and behavior by removing the misunderstandings and misconceptions. Mass Health education also helps to eradicate social stigma, social ostracism and social prejudice associated with leprosy which is the biggest hindrance for the eradication of disease.
The complications of the disease cause disfigurement and disabilities which in turn gives way to the stigma and strong discrimination of these patients. This results not only in physical and social isolation also financial dependency, ultimately forcing the leprosy patients to beg on streets for their survival. To address this issue WHO introduced the strategy of community-based rehabilitation (CBR). This intended to enhance the quality of life for lepers with disabilities through community initiatives. Community participation and using local resources to support the rehabilitation of people with disabilities within their own communities is the foundation of this concept [9, 10].
“Enhanced Global Strategy for Further Reducing the Disease Burden due to Leprosy for 2011–2015” was launched in 2009 by the World Health Organization. The target of the program was to reduce Grade 2 Disability rate (G2DR) in leprosy patients by at least 35% by the end of 2015 (G2DR is the number of new cases with grade 2 disability per 100,000 population). Since the elimination of leprosy in 2005, the prevalence is very less and thus G2DR has been proposed as an indicator. The advantage of G2DR as indicator is that, it is less susceptible to operational factors such as detection delay and is a more robust marker for mapping cases of leprosy in any country. This will also help the program implementers to focus on interventions that reduce visible deformities by enhancing early detection and treatment of leprosy patients and ultimately reduce the number of new leprosy cases in the population. However by the end of 2015, only Thailand was able to achieve this target [11].
In 2016, WHO launched the “Global Leprosy Strategy 2016–2020: Accelerating towards a leprosy-free world” [9].
The program aims to reinvigorate efforts to control leprosy and avert disabilities, especially among children still affected by the disease in endemic countries.
The strategy is built around three major pillars:
Strengthen government ownership and partnerships;
Stop leprosy and its complications; and
Stop discrimination and promote inclusion.
The strategy of this program is:
To sustain expertise and increase the number of skilled leprosy staff;
To improve the participation of affected persons in leprosy services;
To reduce visible deformities and stigma associated with the disease;
To call for renewed political commitment and enhanced coordination among partners;
To highlight the importance of research and improved data collection and analysis.
The key interventions needed to achieve these targets include:
Early case detection especially in children before visible disabilities occur thus reduce transmission;
In highly endemic areas or communities detection of disease among higher risk groups through campaigns;
Improving health care coverage and access for marginalized populations such as poor patients, patients in the difficult to reach areas and the areas of conflicts.
Customization of the strategic interventions in endemic countries is permitted to suit the national plans to meet the new targets. E.g. Screening all close contacts of persons affected by leprosy; initiating a shorter and uniform treatment regimen; and incorporating specific interventions against stigmatization and discrimination.
Its ultimate goal of this program is to further reduce the global and local leprosy burden, that is, (a) zero disabilities in children with leprosy-affected, (b) G2DR less than one per million population and (c) repeal of laws that discriminate leprosy patients of their rights.
Author declares no conflict of interest.
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