Fatty acids composition of olive oil.
\r\n\tIn sum, the book presents a reflective analysis of the pedagogical hubs for a changing world, considering the most fundamental areas of the current contingencies in education.
",isbn:"978-1-83968-793-8",printIsbn:"978-1-83968-792-1",pdfIsbn:"978-1-83968-794-5",doi:null,price:0,priceEur:0,priceUsd:0,slug:null,numberOfPages:0,isOpenForSubmission:!1,hash:"b01f9136149277b7e4cbc1e52bce78ec",bookSignature:"Dr. María Jose Hernandez-Serrano",publishedDate:null,coverURL:"https://cdn.intechopen.com/books/images_new/10229.jpg",keywords:"Teacher Digital Competences, Flipped Learning, Online Resources Design, Neuroscientific Literacy (Myths), Emotions and Learning, Multisensory Stimulation, Citizen Skills, Violence Prevention, Moral Development, Universal Design for Learning, Sensitizing on Diversity, Supportive Strategies",numberOfDownloads:null,numberOfWosCitations:0,numberOfCrossrefCitations:null,numberOfDimensionsCitations:null,numberOfTotalCitations:null,isAvailableForWebshopOrdering:!0,dateEndFirstStepPublish:"September 14th 2020",dateEndSecondStepPublish:"October 12th 2020",dateEndThirdStepPublish:"December 11th 2020",dateEndFourthStepPublish:"March 1st 2021",dateEndFifthStepPublish:"April 30th 2021",remainingDaysToSecondStep:"3 months",secondStepPassed:!0,currentStepOfPublishingProcess:4,editedByType:null,kuFlag:!1,biosketch:"Dr. Phil. Maria Jose Hernandez Serrano is a tenured lecturer in the Department of Theory and History of Education at the University of Salamanca, where she currently teaches on Teacher Education. She graduated in Social Education (2000) and Psycho-Pedagogy (2003) at the University of Salamanca. Then, she obtained her European Ph.D. in Education and Training in Virtual Environments by research with the University of Manchester, UK (2009).",coeditorOneBiosketch:null,coeditorTwoBiosketch:null,coeditorThreeBiosketch:null,coeditorFourBiosketch:null,coeditorFiveBiosketch:null,editors:[{id:"187893",title:"Dr.",name:"María Jose",middleName:null,surname:"Hernandez-Serrano",slug:"maria-jose-hernandez-serrano",fullName:"María Jose Hernandez-Serrano",profilePictureURL:"https://mts.intechopen.com/storage/users/187893/images/system/187893.jpg",biography:"DPhil Maria Jose Hernandez Serrano is a tenured Lecturer in the Department of Theory and History of Education at the University of Salamanca (Spain), where she currently teaches on Teacher Education. She graduated in Social Education (2000) and Psycho-Pedagogy (2003) at the University of Salamanca. Then, she obtained her European Ph.D. on Education and Training in Virtual Environments by research with the University of Manchester, UK (2009). She obtained a Visiting Scholar Postdoctoral Grant (of the British Academy, UK) at the Oxford Internet Institute of the University of Oxford (2011) and was granted with a postdoctoral research (in 2021) at London Birbeck University.\n \nShe is author of more than 20 research papers, and more than 35 book chapters (H Index 10). She is interested in the study of the educational process and the analysis of cognitive and affective processes in the context of neuroeducation and neurotechnologies, along with the study of social contingencies affecting the educational institutions and requiring new skills for educators.\n\nHer publications are mainly of the educational process mediated by technologies and digital competences. Currently, her new research interests are: the transdisciplinary application of the brain-based research to the educational context and virtual environments, and the neuropedagogical implications of the technologies on the development of the brain in younger students. Also, she is interested in the promotion of creative and critical uses of digital technologies, the emerging uses of social media and transmedia, and the informal learning through technologies.\n\nShe is a member of several research Networks and Scientific Committees in international journals on Educational Technologies and Educommunication, and collaborates as a reviewer in several prestigious journals (see public profile in Publons).\n\nUntil March 2010 she was in charge of the Adult University of Salamanca, by coordinating teaching activities of more than a thousand adult students. She currently is, since 2014, the Secretary of the Department of Theory and History of Education. Since 2015 she collaborates with the Council Educational Program by training teachers and families in the translation of advances from educational neuroscience.",institutionString:"University of Salamanca",position:null,outsideEditionCount:0,totalCites:0,totalAuthoredChapters:"0",totalChapterViews:"0",totalEditedBooks:"0",institution:{name:"University of Salamanca",institutionURL:null,country:{name:"Spain"}}}],coeditorOne:null,coeditorTwo:null,coeditorThree:null,coeditorFour:null,coeditorFive:null,topics:[{id:"23",title:"Social Sciences",slug:"social-sciences"}],chapters:null,productType:{id:"1",title:"Edited Volume",chapterContentType:"chapter",authoredCaption:"Edited by"},personalPublishingAssistant:{id:"301331",firstName:"Mia",lastName:"Vulovic",middleName:null,title:"Mrs.",imageUrl:"https://mts.intechopen.com/storage/users/301331/images/8498_n.jpg",email:"mia.v@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, 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:"6942",title:"Global Social Work",subtitle:"Cutting Edge Issues and Critical Reflections",isOpenForSubmission:!1,hash:"222c8a66edfc7a4a6537af7565bcb3de",slug:"global-social-work-cutting-edge-issues-and-critical-reflections",bookSignature:"Bala Raju Nikku",coverURL:"https://cdn.intechopen.com/books/images_new/6942.jpg",editedByType:"Edited by",editors:[{id:"263576",title:"Dr.",name:"Bala",surname:"Nikku",slug:"bala-nikku",fullName:"Bala Nikku"}],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:"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"}},{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:"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:"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:"371",title:"Abiotic Stress in Plants",subtitle:"Mechanisms and Adaptations",isOpenForSubmission:!1,hash:"588466f487e307619849d72389178a74",slug:"abiotic-stress-in-plants-mechanisms-and-adaptations",bookSignature:"Arun Shanker and B. Venkateswarlu",coverURL:"https://cdn.intechopen.com/books/images_new/371.jpg",editedByType:"Edited by",editors:[{id:"58592",title:"Dr.",name:"Arun",surname:"Shanker",slug:"arun-shanker",fullName:"Arun Shanker"}],productType:{id:"1",chapterContentType:"chapter",authoredCaption:"Edited by"}},{type:"book",id:"878",title:"Phytochemicals",subtitle:"A Global Perspective of Their Role in Nutrition and Health",isOpenForSubmission:!1,hash:"ec77671f63975ef2d16192897deb6835",slug:"phytochemicals-a-global-perspective-of-their-role-in-nutrition-and-health",bookSignature:"Venketeshwer Rao",coverURL:"https://cdn.intechopen.com/books/images_new/878.jpg",editedByType:"Edited by",editors:[{id:"82663",title:"Dr.",name:"Venketeshwer",surname:"Rao",slug:"venketeshwer-rao",fullName:"Venketeshwer Rao"}],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"}}]},chapter:{item:{type:"chapter",id:"69338",title:"Advanced Glycation End Products: Formation, Role in Diabetic Complications, and Potential in Clinical Applications",doi:"10.5772/intechopen.89408",slug:"advanced-glycation-end-products-formation-role-in-diabetic-complications-and-potential-in-clinical-a",body:'Advanced glycation end products (AGEs) are formed through a non-enzymatic process in hyperglycemic conditions, and they impact the retinal vasculature negatively through the formation of reactive oxygen species, secretion of aberrant proteins or growth factors, alteration of the extracellular matrix, and secretion of inflammatory cytokines [1]. It is important to consider the difficulty of differentiating the effects of hyperglycemia from those of AGEs, as AGE concentration is controlled by glucose levels. Because of this, occasionally high glucose levels are measured interchangeably with high levels of AGEs. There are two primary mechanisms by which AGEs damage the retinal vasculature which will be discussed in this chapter: interactions with RAGE (AGE receptors) and damage to the extracellular matrix [2]. While these two mechanisms work differently, both pathways result in thickening of the basement membrane which impairs signaling between cells of the microvasculature hindering their structure and increasing rigidity, which leads to the hemorrhagic signs seen in patients with diabetic retinopathy (DR) [3]. Endogenous anti-stressors are important for the management of high levels of AGEs through various mechanisms, but many times are not sufficient to control the progression of DR [2]. Thus, it is important to modify the production of AGEs through exogenous mechanisms, such as nutrition, reducing smoking, or treating the condition through medication [2].
Advanced glycation end products (AGEs) were first discovered in the early 1900s by the Maillard reaction process. Scientists discovered that when amino acids were heated in a mixture with reducing sugars, the reaction turned a yellowish brown color. Further studies indicated that reducing sugars, i.e., glucose, reacted non-enzymatically with the amino acid reagents to form Schiff bases, an early glycation product, and Amadori products, intermediate glycation products. AGE formation can utilize other reagents such as lipids, connective tissue extracellular matrix, and nucleic acids. The process of glycation is enhanced by diabetic complications and occurs in the earlier stages of the Maillard reaction; intracellular sugars, such as glycolytic pathway intermediate glucose-6-phosphate, are glycated at a faster rate than glucose. Amadori products are α-dicarbonyls (oxoaldehydes) such as 3-deoxyglucosone (3-DG) and methylglyoxal (MGO) which is formed by the non-oxidative rearrangement of Amadori adducts from fructose-3-phosphate in the polyol pathway. This pathway has also been studied as a precursor to hyperglycemia-induced damage in diabetes. Methylglyoxal and 3-deoxyglucosone are formed in the early stages of glycation processes: degradation of glucose, Schiff’s bases, and from Amadori products; these oxoaldehyde products can serve as a checkpoint in the AGE pathway since an accumulation of these products is an implication of accelerated vascular damage [4, 5].
The main mechanisms of AGE that affect cells are the adducts on proteins (including N-carboxymethyllysine, pentosidine, or hydroimidazolone) that can interact via AGE ligand-gated receptors such as RAGE on endothelium that lead to secretion of cytokines TNF-α and VEGF; AGEs can stem from exogenous and endogenous adducts due to glucose metabolism. RAGE is the most widely studied AGE receptor found on endothelial cells in vasculature and on macrophages and microglia. AGE interacts with RAGE on macrophages, leading to intracellular generation of free radicals and oxidative stress, which are then phosphorylated by MAP kinase to activate NF-κB and increase expression of NF-κB controlled genes to cause vasoconstriction, enhanced adhesion molecule expression, and induce a procoagulant state. An overexpression of RAGE leads to oxidative stress and NF-κB activation. Current studies show that cross-linked AGEs with RAGE on proteins are closely linked with diabetic retinopathy progression. In the diabetic retina, AGE and adducts are found on vascular cells, neurons, glia, and in elevated levels in Muller macroglia—these specialized retinal cells show increased dysfunction in hyperglycemic and hypoxic conditions that lead to more AGE formation. AGEs induce oxidative stress and consequent apoptosis of retinal pericytes; furthermore, AGEs induce the closure of intercellular junctions between endothelial cells [4, 5, 6, 7].
Inflammation is an important component in the progression of diabetic retinopathy (DR), and AGEs induce this process through interaction with receptors on the cell surface called RAGEs. These receptors are found on most cells, meaning that AGEs exert a wide effect on many different organs. In DR, results of AGE-RAGE interaction on inflammatory cells such as macrophages and lymphocytes, and on microvascular cells such as endothelial cells or pericytes are thought to produce a significant impact on the progression of DR [8]. Monocytes and lymphocytes secrete inflammatory cytokines through the induction of NF-κB [9], production of IL-1, IL-6, IL-8, MCP-1 and TNF-α, and upregulation of adhesion molecules such as VCAM and ICAM [9]. IL-8 and TNF-α levels are elevated in patients with nonproliferative diabetic retinopathy (NPDR), signifying the increased inflammation in the early stages of DR. These cytokines are produced by activated neuronal cells and endothelial cells, and they exert their effect by causing early neuronal cell death in the retina [9]. Inflammation negatively impacts the retinal vasculature by altering the action of vascular cells which leads to the upregulation of various proteins that contribute to the thickening of the basement membrane. MIP-1, IL-3, and IL-1 are thought to play a role in angiogenesis [9, 10], which would facilitate the progression from NPDR to proliferative diabetic retinopathy (PDR). Communication between glial cells and neurons is imperative for maintenance of the vasculature, and it has been shown that inflammation can impede the crosstalk between these cells early in the disease process [9]. Thickening of the basement membrane is one of the leading mechanisms by which crosstalk amongst cells of the retinal vasculature are impeded. This crosstalk is essential for many processes such as providing energy to retinal vascular cells and maintaining homeostasis [9]. In endothelial cells, AGE-RAGE interaction has been shown to increase proliferation via increased VEGF production induced through the MAPK pathway [10, 11]. This process contributes to angiogenesis and accelerates the progression of DR from NPDR to PDR. In pericytes, an opposite effect has been observed, as increased AGE-RAGE interaction leads to apoptosis of these cells, which is one of the first steps in the pathogenesis of DR [11]. As pericyte dropout occurs, the vasculature becomes less regulated leading to hemorrhage and leaking.
Reactive oxygen species (ROS) accumulate in DR from the conversion of glucose to fructose through the NADPH pathway. This accumulation of ROS leads to increased production of AGEs, which then exert their effects through AGE-RAGE interactions or by crosslinking extracellular matrix proteins. One of the outcomes of AGE-RAGE interactions is production of ROS as well, leading to enhanced concentrations of ROS and further progression of the disease. Aldose reductase, which is upregulated to compensate for the high levels of glucose and is essential for the conversion of glucose to fructose, activates a serine/threonine-related protein kinase PKC-δ. Protein kinase PKC-δ is known to inhibit platelet derived growth factor survival activity, an essential pathway for pericyte proliferation and survival. Considering that pericyte loss is typically the initial step in the pathogenesis of DR, this explains the role of ROS in the early stages of DR [12].
The other predominant mechanism of damage from AGEs pertains to their effect on the extracellular matrix of the retinal basement membrane. Inflammation induced by AGEs that was discussed above has a significant impact on the basement membrane, specifically from the elevated levels of inflammatory cytokines IL-1β and TNF-α which induce the production of extracellular matrix proteins. As these excess proteins accumulate in the extracellular matrix, the basement membrane begins to thicken. When AGEs attach to collagen or elastin in the extracellular matrix, it causes the collagen to be less susceptible to hydrolytic breakdown and becomes less flexible. It has also been found that glycation increases the production of collagen and other extracellular matrix proteins, along with the increase in production induced by inflammatory cytokines. This increased production and crosslinking of collagen along with the decreased elastin levels significantly increase the rigidity of the microvasculature through stiffening and thickening of the basement membrane [2, 10, 13].
The accumulation and crosslinking of extracellular matrix proteins contributes to the thickening of the basement membrane, which hinders its integrity ultimately leading to the hemorrhagic pathologies that occur as a result of diabetic retinopathy. The initial damage caused by this thickening is decreased perfusion of the retinal capillaries, leading to occlusion or degeneration of these capillaries [14]. This is one of the characteristic steps in NPDR: ischemia caused by lack of oxygen perfusion sets off the cascade of events that leads to neovascularization, the hallmark of PDR. When looking at the sequelae following the impact of AGEs on the basement membrane, it suggests that AGEs play a significant role in the progression and pathogenesis of diabetic retinopathy. A study showed rats with diabetes tested positive for AGEs (periodic acid/Schiff reagent positive material) at significantly higher levels than those under normal conditions [15]. Rats with diabetes also demonstrated a twofold increase in acellular retinal capillaries over the course of 26 weeks compared to their wild type counterparts, and diabetic rats also experienced significant capillary closure over the course of 75 weeks.
Processing of foods at high temperatures using the Maillard reaction to enhance flavoring and color subsequently leads to the formation of reactive aldehydes that leads to formation of advanced glycation end products, which are also formed naturally in body tissues. Studies depicted that canned meats, nuts, and grain-based products contained the highest levels of AGE, and coffee, butter, vegetables, and fruits as well as food prepared by steaming or boiling contained the lowest amounts of AGE [5, 16].
Research studies show that the average amount of AGE consumed on a daily basis by an individual range from 12,000 to 20,000 kilo-units (kU) of AGEs/day with diabetic subjects consuming a range of 4000–24,000 kU AGEs/day. Pyrraline is one of the most common AGE adducts and may be found in milk and bread crust, while pentosidine, another AGE adduct, is found in pretzel sticks and in its free form in coffee. Study of individual AGEs suggest that protein-bound pentosidine is not as readily absorbed as free pentosidine, therefore, increased levels free AGE in urine and plasma is correlated to AGE-rich dietary intake. Intake of elevated levels of sodium, carbohydrates, and vitamins were found to not be associated with DR risk or progression. Relationship between dietary AGE and promotion of AGE formation in the body tissues will require new research since current research has only centered on skin autofluorescence before and after intake of AGE-rich foods [5, 16].
The effects of dietary AGE were examined in several studies. AGE-poor diets depicted improved biomarkers for oxidative stress, endothelial, and inflammation in healthy subjects, and restricted AGE diets showed decreased levels of oxidative stress in diabetic patients as well as decreased insulin resistance and reduced levels of low-density-lipoprotein. Other studies have also found that dietary AGEs affect inflammatory markers including cytokine TNF-α, and AGE-poor diets have led to decreased risk for cardiovascular disease and endothelial dysfunction. Several studies have also examined the effects of dietary AGE on motor functions, finding that increases in oxidative stress and inflammation due to high levels of AGE lead to muscle stiffness and loss of elasticity [5, 16].
Hyperglycemic conditions initiate formation of AGE and promote biochemical abnormalities that involve formation of AGE. The three main AGE formation biochemical abnormalities include flux via hexosamine pathway, diacylglycerol-mediated activation of PKC-β with benfotiamine, and the stimulation of transketolase activity that induces excess triose phosphates to undergo the pentose phosphate pathway [17, 18].
The primary precursor of AGE is glucose, but other carbonyl precursors exists, though diminutively less reactive, including glyoxal, methylglyoxal, and 3-deoxyglucosone that result from glycolysis. The levels of AGE in the body tissues increase significantly in complications of disease such as diabetic retinopathy, but it is the accumulation of AGE that results in accelerated complications of diseases. Body cells have innate detoxification systems that prevent accumulation of AGE precursors such as methylglyoxal, and detoxification properties of enzymes may be essential in further research about prevention of diabetic retinopathy complications. Deterioration of kidney function leads to accumulation of AGEs, thus leading to endothelial abnormality and vascular disease [4, 5, 17].
No cure for diabetic retinopathy has been discovered yet, despite many efforts from various clinical trials. The standard pharmacological treatment currently for diabetic retinopathy is anti-VEGF injections, which aids in the stabilization and halts progression of the disease [19]. This approach has only been successful in treating about two-thirds of the population and the best second-line pharmacological therapy has not been identified [19]. These factors have spurred the search for a better alternative, especially agents which combat the AGEs and their effects directly. There are different categories of treatments against AGEs, but the most widely studied treatments include those that specifically inhibit AGEs themselves as well as lifestyle changes to reduce the production of AGEs.
The first direct AGE inhibitor that garners the most promise is aminoguanidine, which inhibits AGE formation on both collagen and the basement membrane [15]. As discussed above in the section about AGE’s impact on the extracellular matrix, AGEs crosslink collagen and other proteins in the basement membrane and extracellular matrix which causes it to thicken, lose its integrity and ultimately become leaky. By inhibiting the formation of these crosslinked proteins, the basement membrane and extracellular matrix can preserve their integrity and the normal communication between pericytes and endothelial cells can continue. A study demonstrated that rats treated with aminoguanidine showed significantly less AGE deposition in the basement membrane/extracellular matrix and overall healthier capillaries [15]. Treatment with aminoguanidine also reduced endothelial cell proliferation in diabetic retina, which is another pathological change associated with diabetes. The downside is that this treatment was unable to completely resolve all of the pathological processes of diabetes, namely the occurrence of retinal microaneurysms. In untreated diabetic rats, 38% demonstrated microaneurysms while those treated with aminoguanidine reduced the incidence to 20% (0% in controls). This improvement is promising, but microaneurysms lead to vessel destruction, which advances the progression of NPDR to PDR, the more detrimental stage of DR. An alternative study demonstrated an even greater decrease in microaneurysms, but their sample size was small (a single retina) and it was conducted in dogs rather than rats [20].
Another direct AGE inhibitor is pyridoxamine. This compound has been found to decrease glycation of proteins in the extracellular matrix as well as decrease the formation and production of AGEs. A study measured the success of treating diabetic retinopathy with pyridoxamine by the quantity of acellular capillaries formed over a period of time [4]. Acellular capillaries are nonperfused capillaries which result from a variety of factors onset by diabetes such as pericyte dropout, extracellular matrix, and endothelial damage [21]. After 29 weeks, it was found that diabetic rats treated with pyridoxamine showed similar amounts of acellular capillaries to controls. It also demonstrated the impact of pyridoxamine on the production of extracellular matrix proteins, which are upregulated in diabetic retinopathy. Pyridoxamine significantly reduced the production of extracellular matrix proteins like collagen type IV and laminin, close to the levels found in controls [4].
Besides the usage of direct AGE inhibitors, other drugs options are being explored. One such drug is Tanshinone IIA (Tan IIA). Tan IIA is derived from the roots of Salvia miltiorrhiza, which is a plant that is used in traditional Chinese medicine. Studies indicate that Tan IIA impacts several of the negative effects that hyperglycemic conditions have on human retinal endothelial cells. Tan IIA has an inhibitory effect on proliferation, migration, and vascularization in human endothelial cells and has some correlation to VEGF expression [22]. In terms of AGEs, a recent study explored how Tan IIA protects retinal endothelial cells from the impacts of AGEs, specifically cell dysfunction resulting from the presence of MGO. The study showed that MGO impacted cell viability negatively in a dose-dependent manner. Treatment of the cells with Tan IIA increased their viability in conditions where MGO was also present. MGO presence also resulted in mitochondrial fission in bovine retinal endothelial cells, and the presence of Tan IIA protected against this type of AGE-induced injury in the cells [23].
Exogenous AGEs are AGEs that are consumed and produced through diet and lifestyle, and they differ from the endogenous AGEs that form in hyperglycemic conditions metabolically. Because of this, diet and lifestyle changes are arguably the most important treatment in DR, as diet is a significant contributor to exogenous AGEs found in the form of foods high in protein and fat. Pertaining to lifestyle, smoking tobacco products is associated with higher levels of AGEs in serum which contributes to the progression and risk of DR [24]. Overall, a decreased calorie intake, a modified diet, and smoking cessation have been shown to increase risk and overall disease progression of DR and should be an important treatment regimen, in addition to pharmacological treatments with AGE inhibitors, in all patients with DR.
Advanced glycation end products (AGEs) are formed in increasing amounts due to hyperglycemic conditions implicated in diseases such as diabetic retinopathy. Endogenous AGEs are products from metabolic pathways that follow the Maillard reaction with the oxidation of oxoaldehydes. Exogenous AGEs may come from food sources processed at high temperatures, which increased the amount of reactive aldehydes in the food. Several studies have indicated that inhibition of AGEs holds high potential in the treatment of diabetic retinopathy. Aminoguanidine, a nonspecific inhibitor of AGE, holds the most pharmaceutical promise according to several studies conducted, but other drugs such as Tanshinone IIA are also promising. However, alterations of lifestyles may also provide highly favorable results in decreasing the amount of AGE produced and consumed by the body.
Diabetes and the complication of diabetic retinopathy are gradually on the rise and are widespread. Another disease that is nearly as widespread and also equally relevant in scientific study is Alzheimer’s disease. Recent studies indicate that there may be a link between the two and the factors that are known to impact one of those diseases, such as AGEs, also have effects on the other [25]. Of organs that may exhibit diabetic complications, the eye and its associated connections are one that are closest physically to the brain, and one long-term study on retinal health and cognitive dysfunction showed that 40% of patients with DR showed reduced cognition [26]. Future studies in relation to AGEs will not only focus on the properties of AGEs and their impact on diabetes and its complications, but also how other illnesses are impacted by them as well.
Olive oil is one of the basic components of Mediterranean diet with health protective characteristics [1]. It has been used for centuries due to its preventive and therapeutic characteristics [2, 3]. Olive oil have positive effects on preventing or improving atherosclerosis, cardiovascular diseases, serum lipoprotein levels, oxidative stress, obesity, type 2 diabetes, inflammation and cancer [4, 5, 6, 7, 8].
Health protective effect of olive oil is based on its chemical composition [4, 5, 9, 10]. Olive oil’s chemical composition may be divided into major and minor components. Major components of olive oil are related with its fatty acid content. Primary monounsaturated fatty acid (MUFA) of olive oil is oleic acid [3]. Minor components of olive oil constitute nearly 2% of the weight of olive oil and include more than 230 minor chemical compounds [3, 10]. These chemical compounds include aliphatic alcohols, triterpene alcohols, sterols, hydrocarbons, pigments, volatile compounds, and phenolic compounds [2, 3]. Olive oil has at least 30 phenolic compounds. These are phenolic acids and derivatives, phenolic alcohols, secoiridoids, lignans and flavones [10, 11, 12].
Oleic acid, which is the basic fatty acid of olive oil [8, 13], and the minor compounds of olive oil, including tocopherols, β-carotene, lutein, squalene, lipophilic and hydrophilic phenols, are considered to be responsible for its positive impacts on human health [3, 14]. Olive oil protects human health by changes in epigenetic, metabolic and physiologic mechanisms [2]. This chapter reviews the fatty acid composition and minor components of olive oil and their effects on human health.
Triglycerides constitute 98–99% of total weight of olive oil. Basic fatty acid of olive oil is oleic acid (55–83%), which is one of the MUFA. Besides, olive oil contains fatty acids such as palmitic acid, stearic acid, linoleic acid and α-linolenic acid (Table 1) [3, 10].
Fatty acids | Common name |
---|---|
Saturated | Myrictic acid |
Palmitic acid | |
Margaric acid | |
Stearic acid | |
Arachidic acid | |
Behenic acid | |
Lignosceric acid | |
Monounsaturated | Palmitoleic acid |
Heptadecenoic acid | |
Oleic acid | |
Eicosenoic acid | |
Polyunsaturated | Linoleic acid |
α-linolenic acid |
Fatty acids composition of olive oil.
Positive impact of olive oil over human health is related with its oleic acid content [9]. MUFA in olive oil is considered to reduce hypercholesterolemia, hypertension, atherosclerosis and cardiovascular mortality [8, 13]. Besides, olive oil consumption increases MUFA intake rather than saturated fatty acids (SFA), which, in turn, protects from cardiovascular diseases [15, 16, 17].
In 1985, it was noticed that the replacement of SFA with MUFA intake in diets reduce low-density lipoprotein (LDL) cholesterol level [18]. It was found that the replacement of carbohydrates in diet with MUFA reduced triglycerides (TG), very low-density lipoprotein (VLDL) cholesterol, blood pressure, C-reactive protein (CRP) and increased high-density lipoprotein (HDL) cholesterol and apolipoprotein A1 (Apo-A1) levels [19, 20, 21]. In an another study with the replacement of SFA with olive oil, which has the same amount of energy in diet, LDL cholesterol levels decreased whereas HDL cholesterol levels did not change [22]. Replacement of partially hydrogenated vegetable oil with MUFA had positive impact on cardiovascular risk parameters, such as LDL cholesterol, TG, Apo-A1 and Apo-B levels [23]. It was indicated that following a MUFA rich diet increased HDL cholesterol levels and decreased TG levels [24]. In a meta-analysis it was found that replacement of SFA with MUFA or polyunsaturated fatty acids (PUFA) significantly reduced total and LDL cholesterol levels [25]. In an another meta-analysis it was determined that consuming diets high in MUFA can improve metabolic risk factors among patients with type 2 diabetes [26]. In a meta-analysis of 12 randomized controlled studies high MUFA diets (>12%) were compared to those with low MUFA diets (≤12%). Significant differences between high- and low-MUFA diets were determined with respect to fat mass, systolic blood pressure and diastolic blood pressure. Therefore, high MUFA diets are important dietary regimens for obesity and cardiovascular disease [27].
Such positive impacts of olive oil on human health are related with high oleic acid, proper amount of linoleic and α-linolenic acid, and limited amount of SFA that it contains. High amount of oleic acid and proper amount of linoleic acid helps cells to protect their integrity and slows down the process of aging. Besides, low levels of α-linolenic acid demonstrate anti-inflammatory and vasodilatory effects [2].
Olive oil is rich in terms of minor compounds with antioxidant characteristics, such as polyphenols, carotenoids, squalene and tocopherols. Tyrosol, hydroxytyrosol, flavonoids (apigenin, luteolin), oleuropein and oleocanthal are among the phenolic compounds of olive oil with antioxidant characteristics (Table 2) [2, 6].
Minor components |
---|
|
Minor components of olive oil.
The amount of squalene, which is the basic hydrocarbon in olive oil ranges between 0.8 and 13 g/kg. Squalene has antioxidant activity and helps to lower serum cholesterol levels. Regular squalene intake by using olive oil in diets protects human health from cancer and cardiovascular diseases [28]. The most important carotenoids in olive oil are luteolin and β-carotene. These compounds have antioxidant characteristics that maintain the neutralization of reactive oxygen species (ROS) [2].
Triterpenes have also antioxidant and anti-inflammatory effects [11]. Regular consumption of olive oil maintains antioxidant intake. These compounds reduce free radicals and prevent damages to the cellular membrane, mitochondria, and DNA, with beneficial effects on aging and cancer risk. Especially, phenolic compounds in olive oil have strong antioxidant effect [2, 6].
Polyphenols are compounds with diverse characteristics, which protects human health from chronic diseases [6]. They are described as phenolic compounds, which consists of one or more hydroxyl groups and aromatic rings [10]. Phenolic compounds in olive oil are classified as phenolic acids, flavonoids, secoiridoids, lignans, and phenolic alcohols (Table 3).
Phenolic compounds | Common compound name |
---|---|
Flavonoids | |
|
|
|
|
Lignans |
|
Phenolic acids and derivatives | |
|
|
|
|
|
|
Phenolic alcohols |
|
Secoiridoids |
|
Phenolic compounds in olive oil.
Phenolic compounds in olive oil may have lipophilic and hydrophilic characteristics. Tocopherols are among the lipophilic phenols. Alpha-tocopherols are the most common type of tocopherols in olive oil. Tocopherols especially prevent lipid oxidation in cellular membrane. Due to this reason, they are considered as the most important antioxidant agents in structures that contain lipid [14]. Tocopherols may be found in other oil types. However, hydrophilic phenols (phenolic acids and alcohols, secoiridoids, flavonoids and lignans) do not exist in oil types other than olive oil [3, 10]. Oleuropein, hydroxytyrosol and tyrosol are among the important phenolic compounds of olive oil [2, 6].
On average, olive oil consists of 500 mg/kg phenol [29] and the amount of phenol in different types of olive oil may range between 40 and 1000 mg/kg [10]. A study conducted in Spain found that daily polyphenol intake from olive and olive oil was 90.4 mg, which constitutes 11% of total daily polyphenol intake [30].
Phenolic compounds display a board spectrum of health promoting characteristics, including lipid-improving, anti-oxidant, anti-inflammatory, anti-atherogenic, anti-thrombotic, anti-mutagenic, anti-microbial effects [6, 7, 8]. Beneficial effects of olive oil’s phenolic compounds on human health include improvements in oxidative stress, lipid metabolism, platelet and endothelial functions, and inflammation [6, 8, 14]. Oleuropein and hydroxytyrosol are principal minor phenolic components of olive oil and these antioxidant components prevents diabetes, cardiovascular diseases, neurodegenerative diseases, cancer, inflammation and oxidative stress through its nutrigenomic and immunomodulatory effects [4, 5]. Also oleacein has antioxidant, anti-inflammatory, anti-proliferative and antimicrobial properties and it may play a special role in decreasing the progression of atherosclerosis [31].
A study conducted on 200 healthy male participants, administered 25 ml/day of three olive oils that had low (2.7 mg/kg), medium (164 mg/kg) and high (366 mg/kg) phenolic content. The study found that higher phenolic content was associated with decrease in oxidative stress markers and improvements in lipid profile [32].
Phenolic compounds in olive oil have strong antioxidant characteristics and radical scavenging activities [29]. Regular olive oil consumption maintains phenolic compound intake and protects human health [14]. Taking these into consideration, we may suggest that biological characteristics of olive oil decrease the prevalence of chronic diseases [6, 8, 14].
Olive oil rich diet protects human health from cardiovascular diseases, hypertension, inflammation, oxidative stress, obesity, type-2 diabetes, and cancer [4, 5, 6, 7, 8].
Cardiovascular diseases arise from malfunctioning of heart and blood vessels and include problems, such as coronary heart disease, atherosclerosis, cerebrovascular diseases, peripheral artery disease, congenital heart disease, deep venous thrombosis, pulmonary embolism, myocardial infarction and stroke. Eating habits, and especially total fat and fatty acid intake are among the reasons of cardiovascular diseases [33]. ‘Prevención con Dieta Mediterránea’ (PREDIMED) study that conducted in Spain and randomly assigned participants, who were at high cardiovascular risk, to one of three diets: a Mediterranean diet supplemented with extra-virgin olive oil (EVOO), a Mediterranean diet supplemented with mixed nuts, and a control diet that was advised to reduce dietary fat. Compared to the control group, the Mediterranean diet supplemented with EVOO or mixed nuts was associated with 30% lower risk of major cardiovascular events for a period of 5 years [34]. In an another study, it was found that olive oil consumption, specifically EVOO, is associated with reduced risks of cardiovascular disease and mortality in individuals at high cardiovascular risk. For each 10 g/day increase in extra-virgin olive oil consumption, cardiovascular disease and mortality risk decreased by 10 and 7%, respectively [35].
A study that followed stroke incidence in three French cities for an average of 5.25 years found that the participants with high olive oil consumption had a 41% lower risk of stroke compared to those who never used olive oil. The same study found that higher plasma oleic acid was associated with lower stroke incidence and that the participants with higher plasma oleic acid levels had a 73% reduction of stroke risk compared to the participants with lower plasma oleic acid levels [36]. Another study, which followed Italian women for an average of 7.85 years, found that cardiovascular disease risk was lower for women in the highest quartile of olive oil intake (35.0 ± 0.1 g/day) compared to those in the lowest quartile of olive oil intake (16.8 ± 0.1 g/day) [37].
Another study conducted in five Spanish regions compared non-consumers of olive oil with participants that consumed ≥29.4 g/2000 kcal/day and found that the highest quartile of olive oil consumption was associated with a 26% reduction in risk of overall mortality and a 44% reduction in cardiovascular disease mortality. The study also found that for each increase in olive oil intake of 10 g/2000 kcal/day, there was a 7% decreased risk of overall mortality and a 13% decreased risk of cardiovascular disease mortality [38].
EVOO that is frequently consumed in Mediterranean countries, tends to produce a less prothrombotic environment, promoting antioxidant and anti-inflammatory effects with a greater endothelial protective capacity, which in turn, prevents cardiovascular diseases [22]. Besides, another study in which, healthy participants were administered virgin olive oil (VOO) for 3 weeks found that VOO supplementation altered the expression of 10 genes related to atherosclerosis development and progression [39].
Phenolic compounds of olive oil protects blood lipids from oxidative damage [8]. LDL oxidation is among the main risk factors that contribute to the development of atherosclerosis and cardiovascular diseases. Oxidation of LDL results in the formation of plaque within the arterial wall. Oxidized LDL levels should be taken into consideration as an indicator of oxidative damage and subclinical atherosclerosis. Oxidation of lipids and LDL apolipoproteins is taken by scavenger receptors on monocytes, smooth muscle cells and macrophages in an uncontrolled process, which, in turn, leads to formation of foam cells as an early feature of atherosclerosis [40]. Phenolic compounds delay atherosclerosis by reducing the expression of oxidized LDL and cellular adhesion molecules [2, 14]. Consumption of VOO, which is rich in terms of phenolic compounds significantly decreased LDL cholesterol levels in 1 week [41].
Due to vessel damage, endothelial adhesion molecule expression, platelet activity and aggregation is stimulated. Circulating macrophages and other molecules are adhered to the endothelium, which scavenge LDL and TG, becoming foam cells [42]. Phenolic compounds in olive oil prevent endothelial adhesion molecule expression and platelet aggregation. Hydroxytyrosol, oleuropein, aglycon and luteolin are crucial to prevent platelet aggregation [8]. Oleocanthal-rich extra virgin olive oil may influence platelet aggregation responses in healthy male adults [43]. Oleacein by inhibiting neutral endopeptidase activity, adhesion molecules expression and elastase release might play a role in the protective effects of olive oil against endothelial injuries [44]. Oleacein enhances anti-inflammatory activity of human macrophages by increasing CD163 expression. It could play a potential role in the prevention of inflammatory disease related to atherosclerosis [45]. Additionally, oleacein possess ability to diminish the destabilization of carotid plaque and it could be useful in the reduction of ischemic stroke risk [46]. For that reasons it is possible to emphasize that olive oil rich diet protects human health from cardiovascular diseases.
Positive effects of EVOO consumption on blood pressure have been noticed. EVOO intake contributes to the decrease in diastolic and systolic blood pressure in hypertensive individuals. When compared with vegetable oil that is rich in terms of PUFA, EVOO consumption has a positive effect on blood pressure [47]. A study conducted in five European countries found that consumption of 25 ml/day olive oil significantly decreased systolic blood pressure [48].
In a meta-analysis conducted in the recent years compared dietary regimes with a high amount of MUFA (>12%) with those ≤12% for a period of more than 6 months and found that diets with high amount of MUFA decreased systolic blood pressure, and diastolic blood pressure [27].
Antioxidant effect of EVOO is related with the fact that EVOO consumption reduces the generation of ROS. This effect prevents endothelial dysfunction, which is responsible for hypertension [49]. Related with this, the comparison of EVOO with olive oil or corn oil reveals that inflammatory markers (TXB2 and LTB4) decreased and serum antioxidant capacity increased only in the group of participants, who were administered EVOO [50].
Increase in oxidative stress causes an increase in ROS. Superoxide anion (O2−), hydroxyl radical (OH), and hydrogen peroxide (H2O2), which are also known as ROS, are the oxidizing agents that highly aggressive against the principal biological components of the organism, including lipids, proteins, mitochondria, and DNA. When lipids in the cell membrane are subjected to this action, oxidation that alters membrane permeability takes place. This, in turn, results with early aging of cells. When enzymes, mitochondria and proteins are subjected to this phenomenon, they can lead to metabolic disorders and inflammation in blood vessels, heart, kidney or joints. On the other hand, if DNA undergoes oxidation, the risk of cancer increases. Minor components of olive oil protects mitochondria and DNA from oxidation and decreases the generation of free radicals [2, 11]. Both oleuropein and oleacein are important antioxidant compounds of olive oil and they are stronger scavengers of O−2 and H2O2 [51].
Increase in ROS is associated with atherosclerosis, cancer and neurodegenerative diseases. Oxidative stress increases the levels of lipid peroxide and oxidized glutathione (GSSG), and decreases the levels of glutathione (GSH) and glutathione peroxidase (GSH-Px). Consumption of olive oil that is rich in terms of phenols improves the balance of GSH and GSSG, increases GSH-Px levels and decreases lipid peroxide levels. Consequently, cellular oxidative damage may be decreased by consumption of olive oil with high phenol content [32, 52]. In a controlled before and after supplementation trial with 45 healthy adults found that regular consumption of 50 ml/day EVOO rich in phenolic compounds during 30 days increases plasma antioxidant capacity and antioxidant enzyme activity (catalase and superoxide dismutase). Also it was observed increase in superoxide dismutase and decrease in catalase gene expression [53].
Cell aging, atherosclerosis, diabetes, rheumatoid arthritis, pulmonary emphysema, cataract, Alzheimer and Parkinson diseases, dementia, and development of breast, prostate, colon and skin cancers are related with continuous oxidative damage of cells. Oxidative damage may be partially prevented when the activation of free radicals is inhibited by nutrients, such as minor compounds of EVOO [2, 11]. In order to protect aging that is caused by the damage of free radicals, EVOO intake should start during the early childhood period [54].
EVOO may prevent inflammation when chronic inflammation is associated with pathological cases, such as obesity. Anti-inflammatory effects of EVOO is depends on its fatty acid content and antioxidant compounds [54]. In a randomized controlled trial in healthy adults found that VOO consumption decreased plasma oxidative and inflammatory status and the gene expression related with both inflammation and oxidative stress [55].
Increase in the concentration of inflammatory markers is associated with a higher risk of cardiovascular disease. Plasma thromboxane B2 (TXB2) and leukotriene B4 (LTB4) are known as the pro-inflammatory agents. TXB2 increases platelet aggregation in blood whereas LTB4 leads to cellular damage [56]. Bogani et al. [50] determined that consumption of EVOO that is rich in phenolic components decreases the concentration of inflammatory markers such as TXB2 and LTB4 and increases the serum antioxidant capacity whereas no such decrease was found for other oil types such as olive oil and corn oil.
Interleukin-6 (IL-6) and CRP, which are among the inflammatory markers increase in case of cardiovascular diseases. Olive oil with high phenolic compounds has anti-inflammatory effect and decreases CRP and IL-6 levels in circulation [57]. Oleocanthal, which is among the phenolic compounds of olive oil, prevents cyclooxygenase-1 (COX-1) and cyclooxygenase-2 (COX-2) activities, which have roles in the inflammation process. Inhibition of cyclooxygenase (COX) enzymes results in the reduction of arachidonate to the eicosanoids, prostaglandins and thromboxane [58]. It have been proven to possess that oleacein have antioxidant and anti-inflammatory activities. Oleacein enhances anti-inflammatory activity of human macrophages by increasing CD163 expression [45]. For that reasons, anti-inflammatory effects of olive oil enables protection from diseases that are related with inflammation [8].
Additionally, EVOO reduces the expression of genes involved in the inflammatory response, including intercellular adhesion molecule-1 (ICAM-1), vascular cells adhesion molecule-1 (VCAM-1) and monocyte chemotactic protein-1 (MCP-1), and interferes with the activation of major transcription factor that controls the inflammatory endothelial activation, namely nuclear factor kB (NF-kB). It was known that minor compounds of EVOO, such as phenols, carotenoids and tocopherols, prevent the activation of NF-kB at cellular level [2].
The anti-inflammatory effects that arise from the consumption of olive oil phenolic compounds have been shown to provide protection against inflammatory diseases. Thus, due to the reduction of the risk of inflammation, it can be said that the Mediterranean populations have low rate of cardiovascular mortality and certain types of cancer [8].
Positive effects of olive oil consumption over weight control are widely known. Olive oil increases postprandial thermogenesis. Besides, it may contribute to an increase in fat oxidation. Furthermore, oleic acid may increase satiety, thus reducing food intake. The effect of fatty acids over weight gain may be related with neurotransmitters, intestinal peptides or thermogenesis [59]. The presence of fatty acids in the small intestine lumen induces a number of changes in the gastrointestinal function and inhibits appetite and energy intake. Gastrointestinal hormones, including cholecystokinin, glucagon-like peptide-1 and peptide YY are crucial to regulate appetite and control nutrition intake [60]. One of the studies found that oleic acid caused a slower gastric emptying, promoted the release of cholecystokinin and peptide YY, and a lower subsequent energy intake for both normal weight and obese participants [61].
Comparison of EVOO with cream, which has the same amount of energy but higher SFA levels indicated that olive oil significantly promoted postprandial fat oxidation and stimulated diet-induced thermogenesis [62]. Due to this reason, SFA may be replaced by MUFA for overweight and obese persons in order to lower body weight and fat weight [63].
Oleoylethanolamide (OEA), which is a by-product of oleic acid, acts as a hormone and may lead to satiety and decrease meal frequency. Nutritional intake of oleic acid stimulates the activation of OEA mobilization in the proximal small intestine, which, in turn, leads to satiety. Oleic acid in diets is precursor of OEA synthesis in erythrocytes. OEA decreases during fasting and increases after meals [64]. OEA production in small intestine serves as a molecular sensor linking fat intake to satiety [65]. In addition, OEA resembles to endocannabinoid anandamide in structural terms. Anandamide has an appetizing effect and activates cannabinoid receptors (CB1). Nevertheless, independent of the cannabinoid receptors, OEA has anorectic effect and increases satiety between meals [65].
OEA regulates satiety and appetite, thus contributes to a decrease in body weight. This effect is related with the engagement of peroxisome proliferator-activated receptors-α (PPAR-α). Following food intake, activation of PPAR-α receptor in intestine plays a role in the stimulation of vagus nerves, creating satiety in paraventricular nucleus (PVH) of hypothalamus [64]. OEA decreases meal frequency, increases lipolysis, and modulates the inflammatory response together with PPAR-α by reducing the activity of nuclear factor NF-kB activity, and increasing the catabolism of LTB4 in macrophages [2]. For that reasons, it has been suggested that the increase in OEA production may prevent overnutrition [2].
Additionally, proinflammatory cytokines, such as leptin, tumor necrosis factor alpha (TNF-α), MCP-1, and IL-6 are released from adipose tissue and this phenomenon triggers chronic inflammation [66]. EVOO prevents inflammation in cases such as obesity, which are associated with chronic inflammation [54]. EVOO consumption by obese individuals may reduce inflammatory responses [22]. Anti-inflammatory effect of EVOO may be related with its fatty acid content and antioxidant compounds [54].
In addition to all these characteristics, the consumption of olive oil together with legumes, vegetable dishes and salads is believed to have positive effects on digestive system, glycemic responses and weight control [59]. A 3 years follow-up of a Mediterranean diet supplemented by VOO and nuts found that Mediterranean diet, especially rich in VOO is associated with higher levels of plasma antioxidant capacity and the reduction in body weight after 3 years of intervention in a high cardiovascular risk population [67]. All these reflect the fact that olive oil consumption together with a Mediterranean diet may prevent weight gain [8, 13].
Olive oil with its MUFA content improves glucose metabolism [8, 13]. When high-MUFA diets were compared with high-PUFA diets, there was a significant reduction in fasting plasma glucose level [26]. In addition, it was determined that decreases of SFA levels and increases of oleic acid levels in cellular membranes do not change insulin secretion but improves insulin sensitivity [2].
Prevention of diabetes might be attributable to the antioxidant property of EVOO, thus, oxidative stress seems to be implicated in β-cells dysfunction and eventually diabetes [68]. After 6 hours of consumption of 50 ml VOO, significant changes occurred in gene expression related with insulin sensitivity [69]. On the other hand, four cohort studies, including 15,784 T2D cases and 29 trials, were included in a recent meta-analysis, which indicated that the highest olive oil intake showed a 16% reduced risk of type 2 diabetes compared with the lowest intake. Additionally, in case of patients with type 2 diabetes, olive oil supplementation resulted in a significant reduction of HbA1c and fasting plasma glucose compared to the control group [70]. Additionally, change from polyunsaturated to monounsaturated diet in type 2 diabetes reduced insulin resistance [71]. For these reasons, olive oil could be beneficial for the prevention and management of type 2 diabetes [70].
Obesity causes an increase in inflammation [54]. Additionally, free fatty acids and glycerol release from adipocytes. Increase in the levels of proinflammatory cytokines, ROS and free fatty acids, result with risk of insulin resistance, which, in turn, may lead to type-2 diabetes [66]. Olive oil consumption contributes to weight loss, which is one of the strongest risk factors for type 2 diabetes [72]. All these reveal that olive oil, which is the fundamental ingredient of Mediterranean diet, may have a preventive role for type 2 diabetes [4, 5, 6, 7, 8].
Epidemiologic studies found that cancer incidence was lower in countries such as Greece, Italy and Spain, which are characterized by high EVOO consumption [54]. Meta-analysis of 19 case-controlled studies reveals that cancer risk for the highest olive oil consuming group was 60% lower than the lowest olive oil consuming group. Besides, the study demonstrated that the increase in olive oil consumption was associated with protection from breast and gastrointestinal cancer [73].
High corn oil diet allows the development of malignant adenocarcinomas in rats whereas high EVOO consumption does not have such an effect. Excessive intake of oleuropein, which is one of the phenols in olive oil, has no toxic effects. Additionally, it has antimicrobial, antioxidant, hypotensive, hypoglycemic and antiangiogenic properties. Due to this reason, it is believed that oleuropein has anti-tumor activities. In case of rats with one or more large tumors (>2 cm diameter), addition of 1% oleuropein to drinking water showed strong antineoplastic effects and lead to the disappearance of tumor within 9-to-12 weeks due to antiangiogenic mechanisms that have direct inhibitor effects over cells. The study found that oleuropein in olive oil is an important compound with antineoplastic activity [74].
EVOO has antineoplastic effect on breast cancer in females. EVOO consumption reduces breast cancer risk in postmenopausal women [72]. In the presence of high levels of fatty acid synthase enzyme, oleic acid inhibits the oncogenic effect of epidermal growth factor receptor (HER2) gene by reducing the transcription activity of this gene. Furthermore, EVOO had a strong tumoricidal action on HER2 as a result of phenols, mainly oleuropein aglycon, which is related to inhibition of HER2 gene [75].
Oxidative damage of DNA starts carcinogenesis. Consumption of olive oil that is rich of phenols may inhibit oxidative DNA damage [76]. Cell proliferation and prevention of cell death are among the other factors that lead to tumor formation and development. Hydroxytyrosol in olive oil may prevent cell proliferation. Besides, oleuropein and hydroxytyrosol induce the death of breast cancer cells [77]. Due to these reasons, phenolic compounds in olive oil maintains the integrity of cells and prevents the development of tumors [8, 77].
Compounds of EVOO may show antitumoral effects as a result of metabolic and pathophysiological mechanisms, and may prevent the transformation of human cells into malignant cells and form metastases [2].
Olive oil has protective effects against inflammation and oxidative stress. Besides, it protects human body from various diseases, including, cardiovascular diseases, hypertension, obesity, type 2 diabetes and cancer. The basic fatty acid of olive oil, namely oleic acid, and minor compounds of olive oil, primarily phenolic compounds with their antioxidant activities, are responsible for the positive effects of olive oil over human health. Olive oil polyphenols have antioxidant, anti-inflammatory, antimicrobial, antiviral, anti-atherogenic, anti-thrombotic, anti-mutagenic and hypoglycemic characteristics. Hydroxytyrosol, tyrosol, and oleuropein are the phenolic compounds that are mainly responsible for antioxidant activity of olive oil. Through antioxidant and anti-inflammatory mechanisms, olive oil leads to epigenetic, metabolic and physiologic changes, which protects human health. Due to this reason, regular consumption of olive oil may be effective to decrease the risk of chronic diseases.
MUFA | monounsaturated fatty acids |
SFA | saturated fatty acids |
LDL | low-density lipoprotein |
TG | triglycerides |
VLDL | very low-density lipoprotein |
CRP | C-reactive protein |
HDL | high-density lipoprotein |
Apo-A1 | apolipoprotein A1 |
PUFA | polyunsaturated fatty acids |
VOO | virgin olive oil |
EVOO | extra-virgin olive oil |
PREDIMED | Prevención con Dieta Mediterránea |
ROS | reactive oxygen species |
O2− | superoxide anion |
OH | hydroxyl radical |
H2O2 | hydrogen peroxide |
GSSG | oxidized glutathione |
GSH | glutathione |
GSH-Px | glutathione peroxidase |
TXB2 | thromboxane B2 |
LTB4 | leukotriene B4 |
IL-6 | interleukin-6 |
COX-1 | cyclooxygenase-1 |
COX-2 | cyclooxygenase-2 |
COX | cyclooxygenase |
ICAM-1 | intercellular adhesion molecule-1 |
VCAM-1 | vascular cells adhesion molecule-1 |
MCP-1 | monocyte chemotactic protein-1 |
NF-kB | nuclear factor kB |
OEA | oleoylethanolamide |
CB1 | cannabinoid receptors |
PPAR-α | proliferator-activated receptors-α |
TNF-α | tumor necrosis factor alpha |
HER2 | epidermal growth factor receptor |
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