",isbn:"978-1-80355-367-2",printIsbn:"978-1-80355-366-5",pdfIsbn:"978-1-80355-368-9",doi:null,price:0,priceEur:0,priceUsd:0,slug:null,numberOfPages:0,isOpenForSubmission:!0,isSalesforceBook:!1,hash:"d3a491e5194cad4c59b900dd57a11842",bookSignature:" Vladimir V. Kalinin",publishedDate:null,coverURL:"https://cdn.intechopen.com/books/images_new/11782.jpg",keywords:"Variety of Traits, Historical Remarks, Modern Definitions and Descriptions, Personality Disorders, Comorbid Psychopathology, Depression, Anxiety, Obsessions, Delusion, Treatment of Personality Disorders, Phenomenology of Personality Traits, Delusional Symptoms",numberOfDownloads:null,numberOfWosCitations:0,numberOfCrossrefCitations:null,numberOfDimensionsCitations:null,numberOfTotalCitations:null,isAvailableForWebshopOrdering:!0,dateEndFirstStepPublish:"March 9th 2022",dateEndSecondStepPublish:"May 12th 2022",dateEndThirdStepPublish:"July 11th 2022",dateEndFourthStepPublish:"September 29th 2022",dateEndFifthStepPublish:"November 28th 2022",remainingDaysToSecondStep:"9 days",secondStepPassed:!0,currentStepOfPublishingProcess:3,editedByType:null,kuFlag:!1,biosketch:'A researcher with over 300 publications in psychopathology, psychopharmacology, neuropsychiatry, and epileptology, a member of the Russian Society of Psychiatry, and the Russian Society of Epileptology. Dr. Kalinin\'s biography is included in Marquis "Who’s Who in Medicine and Healthcare" (2006-2007); Who’s Who in Science and Engineering 2008-2009"; "Who’s Who in the World" (2010, 2011), and in the Cambridge International Biographical Centre.',coeditorOneBiosketch:null,coeditorTwoBiosketch:null,coeditorThreeBiosketch:null,coeditorFourBiosketch:null,coeditorFiveBiosketch:null,editors:[{id:"31572",title:null,name:"Vladimir V.",middleName:null,surname:"Kalinin",slug:"vladimir-v.-kalinin",fullName:"Vladimir V. Kalinin",profilePictureURL:"https://mts.intechopen.com/storage/users/31572/images/system/31572.png",biography:"Vladimir V. Kalinin was born in1952 into a family of physicians in Orenburg (Russian Federation). He obtained an MD from Moscow State Medical Stomatological University in 1976. In 1976-1977 he completed an internship in Psychiatry. 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\n
1. Introduction
\n
The supplementation of formula milk with vitamin D3 (cholecalciferol) prompted a rise in infants presenting with symptomatic hypercalcaemia in the United Kingdom during the 1950s [1]. While this public health initiative was proving highly successful in preventing rickets, for the small cohort of infants presenting with failure to thrive, dehydration and nephrocalcinosis, the consequences of their hypercalcaemia were at times fatal. A diagnosis of idiopathic infantile hypercalcaemia was given to many in this cohort. The apparent increased susceptibility of this minority group to vitamin D toxicity prompted research into a genetic predisposition. Fifty-nine years later, CYP24A1 mutations were identified demonstrating loss-of-function mutations encoding 1,25-hydroxyvitamin D3 24-hydroxylase, an enzyme with a key role in vitamin D metabolism [2].
\n
More recently, CYP24A1 mutations have been recognised in an adult population of patients presenting with calcium-containing renal stones. On investigation, these patients typically displayed hypercalciuria, nephrocalcinosis and occasionally chronic kidney impairment. Vitamin D supplementation was not a feature in all cases [3], demonstrating a clinically significant phenotype manifesting from normal dietary vitamin D intake. Importantly, some patients had been symptomatic for many years, undergoing extensive investigations before a diagnosis was made. A continuing focus on preventative medicine, including oral vitamin D supplementation for maintenance of bone health and during pregnancy, is likely to continue to risk triggering manifestations of vitamin D toxicity in individuals carrying biallelic mutations in CYP24A1. As diagnostic tests and successful treatments are starting to emerge, it is important to recognise clinical presentations which should prompt screening for CYP24A1 deficiency [4–6].
\n
\n
\n
2. CYP24A1 and the vitamin D pathway
\n
The crucial role of vitamin D in calcium and phosphate homeostasis means excessive levels of its active form can precipitate symptomatic hypercalcaemia. The activation of vitamin D takes place in two stages. The first stage takes place in the liver: vitamin D3 is converted to 25-hydroxyvitamin D3, a reaction catalysed by 25-hydroxylase (CYP2R1). The second stage occurs in the kidney, when 25-hydroxyvitamin D3 is hydroxylated to 1,25-dihydroxyvitamin D3, the active form. This stage is catalysed by 1α-hydroxylase, an enzyme encoded by the CYP27B1 [2].
\n
Figure 1.
Vitamin D metabolism pathway. Activation of Vitamin D: 1. Stage 1 occurs in the liver. Vitamin D3 is converted to 25-hydroxyvitamin D3 by the enzyme 25-hydroxylase. The CYP2R1 gene encodes 25-hydroxylase. 2. Stage 2 occurs in the kidney. 25-hydroxyvitamin D3 is converted to 1,25-dihydroxyvitamin D3 by the enzyme 1α-hydroxylase. The CYP27B1 gene encodes 1α-hydroxylase. 1,25-dihydroxyvitamin D3 is the physiologically most active form of vitamin D3 which binds to the vitamin D receptor. Inactivation of Vitamin D: Several hydroxylation steps occur in the catabolism of 1,25-dihydroxyvitamin D3 to calcitroic acid. The first of these steps is catalysed by the enzyme 1,25-hydroxyvitamin-D3-24-hydroxylase, which is encoded by the CYP24A1 gene.
\n
The inactivation of vitamin D metabolites relies upon two pathways which both include steps catalysed by 1,25-hydroxyvitamin-D3-24-hydroxylase; CYP24A1 encodes this mitochondrial enzyme which is part of the cytochrome P450 system [6]. The enzyme is present in vitamin D target cells, predominantly located in the intestine and kidneys (Figure 1) [5].
\n\n
\n
2.1. Phenotypes
\n
\n
2.1.1. Idiopathic infantile hypercalcaemia
\n
The first recognised phenotype of CYP24A1 mutations was in infants diagnosed with idiopathic infantile hypercalcaemia. These individuals presented with vomiting, dehydration, fevers and failure to thrive. On investigation, a typical biochemical profile of high serum calcium and suppressed parathyroid hormone levels emerged. Renal ultrasound often demonstrated nephrocalcinosis, deposition of calcium salts within the kidney. It was not initially known whether the underlying pathophysiology of idiopathic infantile hypercalcaemia (IIH) was due to excess production of vitamin D metabolites, or an inability to inactivate vitamin D. A candidate gene approach was used to investigate families with typical presentations of idiopathic infantile hypercalcaemia. This research revealed a recessive loss-of-function mutation, in which patients with CYP24A1 mutations were unable to inactivate vitamin D as they were deficient in the enzyme catalysing this pathway (1,25-hydroxyvitamin-D3-24-hydroxylase). Affected children presented either after sustained low-dose vitamin D prophylaxis or directly following bolus doses of vitamin D. One sibling in which vitamin D prophylaxis was avoided was proven to carry the same mutation but had remained clinically silent. This supported evidence directly linking exogenous vitamin D supplementation with precipitation of symptomatic hypercalcaemia [2].
\n
\n
\n
2.1.2. Adult nephrolithiasis
\n
Hypercalciuria is the most common cause of calcium-containing kidney stones. The recognition that 40–45% of patients with idiopathic hypercalciuria have at least one relative with nephrolithiasis implicates a genetic predisposition in many cases [4]. CYP24A1 mutations have now been proven in a cohort of adults presenting with nephrolithiasis, hypercalciuria, nephrocalcinosis and intermittent hypercalcaemia [4]. These patients had undergone extensive investigations before the cause of their nephrolithiasis was known, and multiple stone episodes and nephrocalcinosis may lead to progressive chronic kidney disease (CKD) [7]. This is important in highlighting the potential clinical spectrum of the phenotype, which may manifest without the trigger of vitamin D exposure. A typical biochemistry profile was found within this phenotype group, with normal/high serum calcium levels, suppressed parathyroid hormone, high levels of active vitamin D metabolites (25-hydroxyvitamin D3 and 1,25-dihydroxyvitamin D3) and low levels of inactivated vitamin D (24,25-dihydroxyvitamin D3). A recent study screening patients with known calcium nephrolithiasis for CYP24A1 mutations did not identify any biallelic variants in a cohort of 166 patients, suggesting CYP24A1 mutations are a rare cause of idiopathic nephrolithiasis [8]. However, given our increased understanding of this phenotype, it is imperative that recognition of the typical biochemical pattern (suppressed PTH, hypercalcaemia, hypercalciuria) in any patients with nephrolithiasis prompts investigation for CYP24A1 mutations [4, 6, 8]. Establishing a molecular diagnosis in this small cohort of patients can facilitate correct treatment and lifestyle modification (Table 1) [9].
\n
\n
\n
\n\n
\n
Clinical features
\n
Biochemical profile
\n
\n\n\n
\n
Idiopathic infantile hypercalcaemia:
Vomiting
Dehydration
Failure to thrive
Fever
Adult presentation:
Nephrolithiasis
Nephrocalcinosis
\n
\n
↑ 25-hydroxyvitamin D3
↑ 1,25-dihydroxyvitamin D3
↓ 24,25-dihydroxyvitamin D3
↑ or high normal serum calcium
↑ urine calcium
↓ parathyroid hormone
\n
\n
\n\n
Table 1.
Key features of CYP24A1 mutation phenotypes.
\n
\n
\n
2.1.3. Investigation
\n
In patients with CYP24A1 mutations, an elevation in total vitamin D levels is typically seen. In particular, 1,25-dihydroxyvitamin D3 levels are increased, but this assay is not routinely performed in many laboratories. Conversely, serum 24,24-dihydroxyvitamin-D3 levels are sometimes low or undetectable in patients with CYP24A1 mutations. A blood test that calculates the ratio between vitamin D metabolites could be utilised in future clinical practice as a screening tool for CYP24A1 mutations in those patients presenting with a typical biochemical profile. In the first study of this, Molin et al. used liquid chromatography–tandem mass spectrometry to calculate the ratio of active to inactive vitamin D metabolites: Molar ratio (R) of 25-hydroxyvitamin-D3: 24,25-dihydroxyvitamin D3. A large increase in the ratio of active to inactive vitamin D metabolites, usually R > 80, was demonstrated in subjects who had biallelic mutations resulting in loss of function of CYP24A1. Importantly, through use of a ratio calculation, this test can avoid misleading results in patients who might have low 24,24-dihydroxyvitamin-D3 levels due to vitamin D deficiency [6].
\n
\n
\n
\n
2.2. CYP24A1 variants
\n
Several different loss-of-function mutations have now been identified within the CYP24A1 gene. The mutations are reported to be inherited in an autosomal recessive pattern, although it is not yet clear whether partial penetrance or environmental factors may alter manifestation of a recognised phenotype. One study showed individuals with biallelic mutations presented with the clinically recognised phenotype and that heterozygous carriers were not sufficient to manifest clinical disease. However, it was hypothesised that infants with haploinsufficiency/heterozygous variants may be more sensitive to hypercalcaemia during childhood while the kidney is still developing, and this could become relevant in considering additional vitamin D supplementation which might overwhelm the 1,25-hydroxyvitamin-D3-24-hydroxylase enzyme pathway in this cohort (Table 2) [4, 6].
\nCYP24A1 mutations lead to calcium stone formation, and conventional treatments for calcium stones are recommended. These would include maintaining a high fluid intake and avoiding excess dietary sodium. Specific measures would include avoiding dietary vitamin D supplements (in foods and drinks) and avoidance of excessive sunlight exposure [7]. Ketoconazole was first demonstrated as an effective treatment for reducing the effects of vitamin D toxicity in patients with CYP24A1 mutations. As a non-specific P450 enzyme inhibitor ketoconazole inhibits the enzyme catalysing production of 1,25-dihydroxyvitamin D3, (1α-hydroxylase), thereby decreasing levels of active vitamin D3. However, CYP24A1-deficient individuals require lifelong treatment as they will always lack the enzyme to inactivate vitamin D, and the side-effect profile of ketoconazole, which includes hepatotoxicity, hypogonadism and adrenal insufficiency, makes it unsuitable for this purpose [4]. More recently, low-dose fluconazole, also acting as a P450 enzyme inhibitor, has been shown to reduce serum calcium levels and urinary calcium excretion in a patient with CYP24A1 mutation. It is likely that this drug, alongside lifestyle modifications such as avoiding excess sun exposure and following a low calcium and oxalate diet, will become the main treatment offered to patients diagnosed with CYP24A1 mutations (Figure 2) [7, 18, 19].
\n
Figure 2.
Chemical structures of ketoconazole, an imidazole antifungal agent, and fluconazole, a triazole antifungal agent. Azole agents are cytochrome inhibitors primarily used as antifungal agents. They are heterocyclic ring compounds and are generally classified as either imidazoles (e.g. ketoconazole) or triazoles (e.g. fluconazole), containing two or three nitrogen atoms, respectively, in the azole ring. They exhibit their antifungal action through inhibition of lanosterol 14-α demethylase, a cytochrome P450 enzyme important for the synthesis of a fungal plasma membrane constituent.
\n
\n
\n
2.4. Evidence for genetic heterogeneity of idiopathic infantile hypercalcaemia
\n
Since the discovery of CYP24A1 mutations underlying idiopathic infantile hypercalcaemia (IIH) in 2011, a cohort of IIH patients has been identified without CYP24A1 mutations. In 2015, a new loss-of-function mutation in SLC34A1, which encodes the renal sodium–phosphate cotransporter 2A (NaPi-IIa), was recognised in this group [20]. These patients presented with a classical IIH phenotype, with symptoms of hypercalcaemia. Importantly, however, their symptoms did not resolve with removal of vitamin D supplementation. Instead, their hypercalcaemia corrected rapidly after commencing phosphate replacement, highlighting the different mechanism driving the hypercalcaemia. In patients with SLC34A1 mutations, renal phosphate wasting leads of inappropriately high levels of 1,25-dihydroxyvitamin-D3, which in turn causes hypercalcaemia. It is crucial to distinguish between patients carrying mutations in CYP24A1 versus SLC43A1, as different intervention is required to successfully treat their hypercalcaemia [20]. As SLC34A1 mutations have also been identified as a cause of nephrolithiasis, there is overlap between SLC34A1 and CYP24A1 mutation phenotypes in both paediatric and adult presentations [21].
\n
\n
\n
\n
3. Conclusions
\n
Overall, CYP24A1 mutations are rare and account for a small proportion of symptomatic hypercalcaemia or nephrolithiasis cases. However, a greater awareness of their phenotypes will increase clinical suspicion in patients presenting with a typical biochemical profile. Testing for mutations in CYP24A1 can establish a definitive diagnosis, avoiding protracted further investigations and allowing treatment to commence. Alongside dietary and lifestyle advice, aimed at minimising vitamin D intake, fluconazole is proving a promising lifelong treatment to prevent effects of vitamin D toxicity.
\n
\n
Acknowledgments
\n
JAS is supported by the Northern Counties Kidney Research Fund.
\n
\n',keywords:"CYP24A1, vitamin D, hypercalcaemia, idiopathic infantile hypercalcaemia, nephrolithiasis",chapterPDFUrl:"https://cdn.intechopen.com/pdfs/51750.pdf",chapterXML:"https://mts.intechopen.com/source/xml/51750.xml",downloadPdfUrl:"/chapter/pdf-download/51750",previewPdfUrl:"/chapter/pdf-preview/51750",totalDownloads:1520,totalViews:642,totalCrossrefCites:4,totalDimensionsCites:4,totalAltmetricsMentions:12,impactScore:3,impactScorePercentile:87,impactScoreQuartile:4,hasAltmetrics:1,dateSubmitted:"November 7th 2015",dateReviewed:"June 6th 2016",datePrePublished:null,datePublished:"April 12th 2017",dateFinished:"July 21st 2016",readingETA:"0",abstract:"The CYP24A1 gene encodes 1,25-hydroxyvitamin-D3-24-hydroxylase, a key enzyme responsible for the catabolism of active vitamin D (1,25-dihydroxyvitamin D3). Loss-of-function mutations in CYP24A1 lead to increased levels of active vitamin D metabolites. Clinically, two distinct phenotypes have been recognised from this: infants with CYP24A1 mutations present with infantile idiopathic hypercalcaemia, often precipitated by prophylactic vitamin D supplementation. A separate phenotype of nephrolithiasis, hypercalciuria and nephrocalcinosis often presents in adulthood. CYP24A1 mutations should be suspected when a classical biochemical profile of high active vitamin D metabolites, high or normal serum calcium, high urine calcium and low parathyroid hormone is detected. Successful treatment with fluconazole, a P450 enzyme inhibitor, has been shown to be effective in individuals with CYP24A1 mutations. Although CYP24A1 mutations are rare, early recognition can prompt definitive diagnosis and ensure treatment is commenced.",reviewType:"peer-reviewed",bibtexUrl:"/chapter/bibtex/51750",risUrl:"/chapter/ris/51750",book:{id:"5269",slug:"a-critical-evaluation-of-vitamin-d-basic-overview"},signatures:"Fay Joanne Hill and John A. Sayer",authors:[{id:"181499",title:"Prof.",name:"John",middleName:"Andrew",surname:"Sayer",fullName:"John Sayer",slug:"john-sayer",email:"john.sayer@ncl.ac.uk",position:null,profilePictureURL:"//cdnintech.com/web/frontend/www/assets/author.svg",institution:null}],sections:[{id:"sec_1",title:"1. Introduction",level:"1"},{id:"sec_2",title:"2. CYP24A1 and the vitamin D pathway",level:"1"},{id:"sec_2_2",title:"2.1. Phenotypes",level:"2"},{id:"sec_2_3",title:"2.1.1. Idiopathic infantile hypercalcaemia",level:"3"},{id:"sec_3_3",title:"Table 1.",level:"3"},{id:"sec_4_3",title:"2.1.3. Investigation",level:"3"},{id:"sec_6_2",title:"2.2. CYP24A1 variants",level:"2"},{id:"sec_7_2",title:"2.3. Treatment",level:"2"},{id:"sec_8_2",title:"2.4. Evidence for genetic heterogeneity of idiopathic infantile hypercalcaemia",level:"2"},{id:"sec_10",title:"3. Conclusions",level:"1"},{id:"sec_11",title:"Acknowledgments",level:"1"}],chapterReferences:[{id:"B1",body:'\nBritish Paediatric Association. Hypercalcemia in infants and vitamin D. British Medical Journal 1956; 2: 149.\n'},{id:"B2",body:'\nSchlingmann KP, Kaufmann M, Weber S, Irwin A, Goos C, John U et al. Mutations in CYP24A1 and idiopathic infantile hypercalcemia. The New England Journal of Medicine 2011; 365(5): 410–421.\n'},{id:"B3",body:'\nJobst-Schwan T, Pannes A, Schlingmann KP, Eckardt K-U, Beck BB, Wiesener MS. Discordant clinical course of vitamin-D-hydroxylase (CYP24A1) associated hypercalcemia in two adult brothers with nephrocalcinosis. Kidney and Blood Pressure Research 2015; 40: 443–451.\n'},{id:"B4",body:'\nDinour D, Beckerman P, Ganon L, Tordjman K, Eisenstein Z, Holtzman EJ. Loss-of-function mutations of CYP24A1, the vitamin D 24-hydroxlylase gene, cause long-standing hypercalciuric nephrolithiasis and nephrocalcinosis. The Journal of Urology 2013; 190: 552–557.\n'},{id:"B5",body:'\nNesterova G, Malicdan MC, Yasuda K, Sakaki T, Vilboux T, Ciccone C. 1,25-(OH)2D-24 hydroxylase (CYP24A1) deficiency as a cause of nephrolithiasis. Clinical Journal of the American Society of Nephrology 2013; 8: 649–657.\n'},{id:"B6",body:'\nMolin A, Baudoin R, Kaufmann M, Souberbielle JC, Ryckewaert A, Vantyghem MC. CYP24A1 mutations in a cohort of hypercalcemic patients: evidence for a recessive trait. Journal of Clinical Endocrinology and Metabolism 2015; 100(10): E1343–E1352.\n'},{id:"B7",body:'\nSayers J, Hynes AM, Srivastava S, Dowen F, Quinton R, Datta HK, Sayer JA. Successful treatment of hypercalcaemia associated with a CYP24A1 mutation with fluconazole. Clinical Kidney Journal 2015; 8(4): 453–455.\n'},{id:"B8",body:'\nSayers J, Hynes AM, Rice SJ, Hogg P, Sayer JA. Searching for CYP24A1 mutations in cohorts of patients with calcium nephrolithiasis. OA Nephrology 2013; 1(1): 1–6.\n'},{id:"B9",body:'\nSayer JA. Re: Loss-of-function mutations of CYP24A1, the vitamin D 24-hydroxlylase gene, cause long-standing hypercalciuric nephrolithiasis and nephrocalcinosis. The Journal of Urology 2015; 68: 164–165.\n'},{id:"B10",body:'\nDauber A, Nguyen TT, Sochett E, Cole DEC, Horst R, Abrams SA et al. Genetic defect in CYP24A1, the vitamin D 24-hydroxylase gene, in a patient with severe infantile hypercalcaemia. The Journal of Clinical Endocrinology and Metabolism 2012; 97(2): E268–E274.\n'},{id:"B11",body:'\nTebben PJ, Milliner DS, Horst RL, Harris PC, Singh RJ, Wu Y et al. Hypercalcaemia, hypercalciuria, and elevated calcitriol concentrations with autosomal dominant transmission due to CYP24A1 mutations: effects of ketoconazole therapy. The Journal of Clinical Endocrinology and Metabolism 2012; 97(3): E423–E427.\n'},{id:"B12",body:'\nFencl F, Bláhová K, Schlingmann KP, Konrad M, Seeman T. Severe hypercalcemic crisis in an infant with idiopathic infantile hypercalcaemia caused by mutation in CYP24A1 gene. European Journal of Pediatrics 2013; 172: 45–49.\n'},{id:"B13",body:'\nSkalova S, Cerna L, Bayer M, Kutilek S, Konrad M, Schlingmann KP. Intravenous pamidronate in the treatment of severe idiopathic infantile hypercalcaemia. Iranian Journal of Kidney Diseases 2013; 7(2): 160–164.\n'},{id:"B14",body:'\nMeusburger E, Mündlein A, Zitt E, Obermayer-Pietsch B, Kotzot D, Lhotta K. Medullary nephrocalcinosis in an adult patient with idiopathic infantile hypercalcaemia and a novel CYP24A1 mutation. Clinical Kidney Journal 2013; 6: 211–215.\n'},{id:"B15",body:'\nJacobs TP, Kaufman M, Jones G, Kumar R, Schlingmann KP, Shapses S. A lifetime of hypercalcaemia and hypercalciuria, finally explained. The Journal of Clinical Endocrinology and Metabolism 2014; 99(3): 708–712.\n'},{id:"B16",body:'\nTray KA, Laut J, Saidi A. Idiopathic infantile hypercalcaemia, presenting in adulthood- no longer idiopathic nor infantile: two case reports and review. Connecticut Medicine 2015; 79(10): 593–597\n'},{id:"B17",body:'\nDinour D, Davidovits M, Aviner S, Ganon L, Michael L, Modan-Moses D et al. Maternal and infantile hypercalcaemia caused by vitamin-D-hydroxylase mutations and vitamin D intake. Pediatric Nephrology 2015; 30: 145–152.\n'},{id:"B18",body:'\nDusso AS, Gomez-Alonso C, Cannata-Andia JB. The hypercalcaemia of CYP24A1 inactivation: new ways to improve diagnosis and treatment. Clinical Kidney Journal 2015; 8(4): 456–458.\n'},{id:"B19",body:'\nFigueres M-L, Linglart A, Bienaime F, Allain-Launay E, Roussey-Kessler G, Ryckewaert A. Kidney function and influence of sunlight exposure in patients with impaired 24-hydroxylation of vitamin D due to CYP24A1 mutations. American Journal of Kidney Disease 2015; 65(1): 122–126.\n'},{id:"B20",body:'\nSchlingmann KP, Ruminska J, Kaufmann M, Dursun I, Patti M, Kranz B et al. Autosomal-recessive mutations in SLC34A1 encoding sodium-phosphate cotransporter 2a cause idiopathic infantile hypercalcemia. Journal of American Society of Nephrology 2015; 27: 604–614.\n'},{id:"B21",body:'\nOddsson A, Sulem P, Helgason H, Edvardsson VO, Thorlefisson G, Sveinbjörnsson G et al. Common and rare variants associated with kidney stones and biochemical traits. Nature Communications 2015; 6(7975): 1–9\n'}],footnotes:[],contributors:[{corresp:null,contributorFullName:"Fay Joanne Hill",address:null,affiliation:'
Renal Services, Newcastle upon Tyne NHS Foundation Trust Hospitals, Newcastle upon Tyne, United Kingdom
'},{corresp:"yes",contributorFullName:"John A. Sayer",address:"john.sayer@ncl.ac.uk",affiliation:'
Renal Services, Newcastle upon Tyne NHS Foundation Trust Hospitals, Newcastle upon Tyne, United Kingdom
Institute of Genetic Medicine, Newcastle University, Newcastle upon Tyne, United Kingdom
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1. Introduction
Defensins are a family of antimicrobial peptides that are part of the innate and adaptive immune system. They provide protection against too broad spectrum of pathogens, viruses, bacteria and fungi [1]. The defensins are small cationic peptides of 3–4 kb, and its structure is composed of beta sheets, six cysteines joined by disulfide bonds. In humans, it is classified in two groups: α- and β-defensins [2, 3]. The β-defensin-2 (HBD2) is found in the second group and it is a peptide of 41 amino acids; it is expressed mainly in the epithelial cells, mucous and skin [1, 4, 5]. It was isolated in 1997 from skin lesions in patients with psoriasis; later, it was detected in the epithelium of almost entire human body [6, 7, 8, 9, 10, 11, 12, 13, 14]. In recent years, HBD2 has been considered as a multifunctional peptide with antimicrobial activity and with immunomodulatory functions [15, 16, 17]. On the other hand, the HBD2 is expressed throughout the respiratory epithelium from the mouth to the lungs; it is believed that this defensin has a very important role in the defense against respiratory infections [2]. The alteration of the expression of the HBD2 in the respiratory epithelium has been associated with the pathogenesis of several respiratory diseases such as asthma, pulmonary fibrosis, pneumonia, tuberculosis, rhinitis, etc. [2, 9, 17, 18, 19, 20].
The acute respiratory infections caused by viruses are the main cause of morbidity and mortality in the world, and HBD2 has got antiviral activity against some respiratory viruses (influenza, respiratory syncytial virus, rhinovirus) [21, 22, 23]. The mechanisms of viral inactivation vary and include not only direct binding of the virus to the peptide but also indirect methods of inactivation via intracellular modulation of the viral replication, modulation of signaling pathways necessary for antiviral effects, and recruitment of immune cells that contribute to antiviral activity [2, 22]. This revision chapter focuses on the structural and general characteristics of HBD2, multifunctional activities and expression in respiratory diseases. We present some studies concerning the effect of respiratory viruses and their relationship with HBD2, mechanisms of action and their relevance as therapeutic agents.
2. General characteristics and classification
Defensins are a family of antimicrobial peptides that form part of the innate and adaptive immune system and constitute the first line of host defense against microorganisms. It has shown the broad antimicrobial activity spectrum against bacteria, fungi and viruses [1, 24].
The defensins are small cationic peptides of 28–42 amino acids, characterized by a β-sheet structure linked by tree disulfide bonds, which are formed by six cysteine residues [2, 25]. Based on the distribution of their cysteines and disulfide bonds, defensins are classified into two groups in humans: α- and β-defensins [1]. α-defensins have 29–35 amino acids and the positions of the cysteines are C1-C6, C2-C4 and C3-C5, while the β-defensins are composed of 38–42 amino acids with the positions of the cysteines of C1-C5, C2-C4 and C3-C6 [26].
The α-defensins are expressed mainly in neutrophils and called human neutrophil peptides with four types (HNP1–4). There are other α-defensins (HD5 and HD6), known as enteric defensins, that are expressed in Paneth cells in the small intestine [27]. The expression of HBD6 has also been confirmed in the female genitourinary system [1, 28].
β-defensins are mainly expressed in epithelial cells throughout the body, including mucous membranes and skin [29, 30, 31, 32]. Four types (HBD1–4) have been identified in humans, but several analyses indicate that there may be approximately 31 types, of which HBD5 and 6 are expressed in epididymis with their importance in defense against infections. The other defensins are known only for their antimicrobial activity [33, 34]. The β-defensin-1 is expressed constitutively, while the other three (HBD2–4) are expressed by the effect of proinflammatory cytokines or during the infectious process [1, 28].
3. Molecular structure of β-defensin-2
The HBD2 is a small cationic peptide with a positive net charge (+6). It has 41 amino acid residues; the complete gene is approximately 4 kb. It consists of six cysteines in positions 1–5, 2–4 and 3–6, joined by three disulfide bonds. Its secondary structure consists of an N-terminal region linked to an alpha helix, three β-strands arranged in an antiparallel sheet and a C-terminal region [35, 36, 37] (Figure 1). Its structure has an amphipathic nature, with hydrophilic and hydrophobic amino acids on the surface of protein; it is stabilized by the disulfide bonds, which protect it from degradation by proteases [38, 39]. The alpha helix is also stabilized by the disulfide bonds (I and V) and by the first beta sheet that has a domain (Gly-X-CysIV), which is responsible for the native structure and the correct folding of the peptide [40, 41]. The N-terminal region binds to the membrane of microorganisms. The specific conformation of N-terminal region of β-defensins may be important for the biological properties of these proteins. [37].
Figure 1.
Secondary structure of HBD-2 that consists of an N-terminal region linked to an alpha helix, three β-strands arranged in an antiparallel sheet and a C-terminal region [41].
The C-terminal region consists mainly of cationic amino acids (lysine and arginine) that are distributed asymmetrically with their positive charges and are important in antimicrobial activity [3, 38, 40].
4. Genetic structure of β-defensin-2
The gene codify for HBD2 is located on chromosome 8p23. The gene is approximately 4 kb and composed of a 5‘and 3’ non-translatable region, two exons separated by an intron. The first exon has 81 bp and codes for the signal peptide and the second exon has 238 bp and codes for a short anion segment called propeptide and the mature peptide [3, 25, 42]. The 5′ region of HBD2 has specific sites that bind to the transcription factor kB (NF-kB). In addition, there are sites that bind to other transcription factors such as C/EBP, and NF-IL-6, which are important for their expression [42, 43, 44].
The HBD2 have a polymorphic nature, and the number of copies of the gene varies in each individual [45]. It has been reported that there are 2–12 copies per diploid genome, and the number of copies is related to the level of expression. It has been suggested that these variations may have consequences on the function of immune system. Some infectious and inflammatory diseases are related to the number of copies of the HBD2 gene [44, 46] (Figure 2).
Figure 2.
Genetic structure of HBD2. It encompasses untranslated 5 ‘and 3’ regions and two exons separated by an intron. The first exon codes for the signal peptide and the second exon codes for the propeptide and the mature peptide [3].
5. Expression and regulation of β-defensin-2
HBD2 is expressed mainly in all the epithelia of human body (respiratory, digestive, urogenital, conjunctive epithelium), mucous, peripheral blood and skin. In the respiratory system, HBD2 is expressed in mucous from the mouth to the epithelium of the lungs and is induced by bacteria, fungi and virus infections and by proinflammatory stimuli such as interleukin 1α (IL-1α), interleukin 1β (IL-1β), tumor necrosis factor alpha (TNF-α), interferon gamma (IFN-γ) and interleukin IL-6 (IL-6) [29, 47, 48, 49, 50, 51]. Recently, a study showed that the cytokines IL-17 and IL-22 are produced by Th17 cells, which are reportedly known to regulate the expression of HBD2 on mucosal surfaces [32]. HBD2 is produced as a functionally inactive peptide (prepro-defensin), and to achieve its biological activity, it must go through a post-translational modification process and form a mature peptide. Prepro-defensin is composed of a highly hydrophobic pre-peptide signal, a propeptide and the mature peptide. The prepropeptide is cut by proteases in the Golgi apparatus, and once removed, the mature peptide with antimicrobial activity is secreted on the surface of the epithelial cells [3, 52].
The regulation of the expression of the HBD2 in the respiratory epithelium involves multiple signaling pathways; most of these have been studied with bacterial infections. The bacterial proteins induce the expression of HBD2 in the respiratory epithelium through transcription factors (NF-kB) [53], the myeloid ELF-1-like factor (MEF) [54], the nuclear factor interleukin 6 (NF-IL6) [55] or the activated protein 1–3 (AP1–3) [56]. Signaling pathways involve mitogen-activated protein kinase (MAPKs) [49, 57, 58], phosphatidylinositol-3-kinase (PI3K) and protein kinase C (PKC) [59].
Another form to induce HBD2 expression is through cellular receptors, the respiratory epithelium expresses various receptors on its surface. Toll-like receptors (TLRs) 1–6 are expressed on the epithelial surface, and in the intracellular vesicles, the TLR3 and TLR7–9 are expressed in the endosomes or endoplasmic reticulum. These receptors recognize pathogens and have a cytoplasmic domain that is homologous to the IL-1 receptor and is responsible for initiating intracellular signaling pathways. This signaling cascade includes the activation of NF-kB. This transcription factor promotes the gene expression that contributes to the cytokines, chemokines, adhesion molecules, co-stimulatory molecules release as well as the expression of HBD2 [60].
In a study with lung epithelial cells infected with L. pneumophila, it was observed that the infection induces the release of HBD2, and its expression is mediated by the receptors TLR2 and TLR5 and activation of MAPKs (p38, JNK) and transcription factors NF-kB and AP-1 [50, 57, 58, 61].
Another way to regulate the expression of HBD2 is through the interaction with receptors; its chemotactic property was initially discovered in different types of cells such as monocytes, T cells and immature dendritic cells. The activity is carried through the CCR6 receptor binding [32].
Other receptors involved in the regulation of the HBD2 expression are the receptors for vitamin D and protease-coupled receptors (PARs) [62, 63, 64, 65].
Nowadays, HBD2 is considered a multifunctional molecule; the ability to bind to several ligands suggests that there may be more receptors and signaling pathways still to be discovered and that may have important biological activities in the immune system.
6. Mechanisms of action of β-defensin-2
Several mechanisms of action have been proposed for defensins; however, the main mechanism of action of the HBD2 is to eliminate microorganisms directly through the interaction with the microorganism membrane. The first step is given by the electrostatic attraction between the cationic defensin with positive charge and the microorganism’s membrane components with negative charges [37].
Components of the bacteria membrane have been identified as targets for the HBD2. Lipopolysaccharides are targets for Gram-negative bacteria, teichoic acid for Gram-positive bacteria and phospholipids for both bacteria. In Gram-negative bacteria, the peptides are inserted into the membrane by hydrophobic interactions. It is thought that it possibly involves a folding of the peptide within the structure of the membrane [38, 66, 67]. After the electrostatic interaction of the peptide with the membrane and the displacement of the lipids, the defensin is added to the surface of membrane. There are several modes of action that have been proposed to describe how defensins are oriented to form pores and how the structure of the membrane is altered, becomes permeable, such as cell lysis and finally results in the death of the microorganism (Figure 3) [3, 37, 68].
Figure 3.
Mechanisms of action of HBD2. The defensin with its positive charge is attached by electrostatic attraction to the membrane of the pathogen forming pores [3].
The high content of negatively charged amino acids in the membranes of bacteria is the main factor that makes them more susceptible to being targeted by defensins. The membrane of the eukaryotic cells predominates lipids with neutral charge without net charge, they have a high level of cholesterol, and bacteria do not contain cholesterol in their membranes. Cholesterol causes the membrane to condense and prevents the peptide from penetrating; this also has an asymmetric distribution of phospholipids contributing to the resistance against defensing. These mechanisms explain why these peptides are not toxic in eukaryotic cells [69, 70].
7. Multifunctional activities of β-defensin-2
HBD2 was isolated and characterized by its antimicrobial activity. Currently, several studies have described different biological activities of HBD2, and it is considered as a multifunctional protein. Some of the account on its biological activities is given below.
7.1 Antimicrobial activities
HBD2 has a broad spectrum of activity against a wide variety of bacteria, fungi and viruses. The mechanism of action of this defensin begins with the interaction of the negative charges of the pathogen membrane, the formation of pores and finally the lysis of the microorganism. The variability of the composition of the membranes of the different pathogens explains in part the different antimicrobial effects [2, 22].
7.2 Innate and adaptive immunity
HBD2 is important in innate immunity and constitutes the first line of host defense against infections by microorganisms. Its role in adaptive immunity is attributed mainly to its chemotactic activity in immature dendritic cells and memory T lymphocytes through the CCR6 receptor [26].
Several mechanisms have been proposed for HBD2 that contributes to the adaptive immunity: (a) Increase in the recruitment of immature dendritic cells. Immature dendritic cells are recruited from circulating blood or tissue near the site of inflammation by chemoattractants that interact with their corresponding receptors (CCR1, CCR5, CCR6). (b) Formation of defensin-antigen complexes. HBD2 forms defensin-antigen complexes facilitating the presentation to dendritic cells. (c) Maturation of dendritic cells. HBD2 induced the maturation of dendritic cells for direct production of IL-2 or indirect production of TNF and IL-1 by monocytes and macrophages. (d) Recruitment of memory T cells. It facilitates the recruitment of memory T cells that are the effector cells of adaptive immunity [15, 26].
HBD2 contributes to unite the innate and adaptive immune response, and this property has been applied for the development of vaccine adjuvants, since it promotes adaptive immunity when it is administered together with antigens [15, 26].
7.3 Inflammation
Inflammation is a protective reaction by the host to eliminate injurious stimuli (microorganism, damage cell or irritants). Some viral infections cause severe inflammation, and the tissue can be damaged and then it must be repaired. The mechanisms include the production of anti-inflammatory cytokines, lipid mediators, glucocorticoids, immune cell apoptosis, etc. [3].
HBD2 plays a critical role in regulating inflammation processes in the respiratory system and modulates the production of inflammatory cytokines and chemokines. An increase in local expression has been observed; but in severe cases, the HBD2 can be detected systemically [71].
HBD-2 can promote histamine release and prostaglandin D2 production in mast cells, suggesting a role in allergic reactions [1, 26, 72].
7.4 Anti-inflammatory activity
HBD2 and the complement system are two important innate immune mechanisms against a broad range of microorganisms. The complement is composed of more than 30 proteins found in the human serum, and it is activated by three different pathways (classical, alternative and lectin) [73, 74]. It has been described that HBD2 binds to C1q (first component of the complement system) and inhibits the classical complement pathway. HBD2 have a dual protective role not only as an antimicrobial agent but also to provide protection against uncontrolled activation of complement system [73, 75, 76].
7.5 Immunomodulatory properties
Recently, the study of the HBD2 has focused on the modulation of the immune response. It has been suggested that multiple mechanisms of action may be involved from direct binding to the membrane of the microorganism to the union with different types of cellular receptors that can induce transduction signals, gene transcription and various signaling pathways [77]. This mechanism paves the way for the development of new therapies for infectious respiratory diseases.
7.5.1 Moxifloxacin/HBD2
In a study with epithelial lung cells (A549) stimulated with LPS, it was demonstrated that the association of moxifloxacin/HBD2 has an anti-inflammatory effect. Moxifloxacin is a fluoroquinolone against Gram-positive and Gram-negative bacteria, which may have affected the immune system. The treatment induced a reduction of proinflammatory cytokines (IL-1 and IL-6). These data support the hypothesis of its immunomodulatory capacity of HBD2 to neutralize the components of bacteria that induce the activation of cytokines [78].
7.5.2 Vitamin D (VitD)
VitD plays an important role for the calcium homeostasis in the bones. Currently, the interest has focused on the modulation of the immune response in fighting viral respiratory infections. Some studies show that patients with deficiencies in VitD are at higher risk of suffering respiratory infections in the upper respiratory tract [79]. One study showed the association of polymorphisms in the VitD receptor and severe bronchiolitis in patients infected by the respiratory syncytial virus [80].
The immunomodulatory effect has been of great importance in some viral infections; several types of cells including epithelial cells treated with VitD induce the expression of the receptor and the production of antimicrobial peptides such as the HBD2. In the patients infected with HIV, the high levels of VitD and its receptor increase the amount of IL-10 and HBD2, which are associated with a natural resistance to HIV infection [62].
The VitD receptor is expressed in several types of cells (monocytes, B cells, T cells and NK), is an endogenous immunomodulator and induces the transcription of the HBD2. This work shows that the expression of HBD2 is important to render tolerance to viral infections [62, 64].
7.5.3 PARs (receptors coupled to proteases)
It is a family of receptors coupled to the G-protein, which is formed by seven transmembrane domains with an amino-terminal extracellular domain and a C-terminal domain. PARs are activated by proteolytic cleavage in the N-terminal domain by serine proteases. N-terminal serves as a ligand to carry out intracellular signaling. They are expressed in epithelial, endothelial and immune cells such as leukocytes, mast cells, eosinophils, neutrophils and mastoid cells. Four types of receptors have been described (PAR-1–4). PAR-1, 3 and 4 are activated mainly by thrombin and are involved in the aggregation of platelets. PAR-2 is activated by trypsin, tryptase from mastoid cells, protease 3 from neutrophils and tissue factor, factor VIIa and Xa [81]. Recently, PARs have been implicated in the regulation of the expression of antimicrobial peptides found in epithelial cells such as defensins [65, 82]. The researchers identified the expression of the HBD2 in human gingiva by P. gingivalis infection. Further, proteases of the bacteria induced the expression of the defensin through PAR2. The authors suggest that this signaling pathway can lead to the development of preventive therapies in mucosal infections [65].
7.5.4 Triptolide
This is an immunosuppressive and anti-inflammatory agent that was extracted from an herb of Chinese origin (Tripterygium wilfordii). It decreases the expression of the NF-kB and genes related to inflammatory processes. This review chapter shows that this agent suppresses the expression of HBD2 induced by IL-1β in A549 cells and the suppression is associated with the inhibition of NF-kB [83].
7.5.5 Neutrophilic elastase
It is a serine protease that is expressed in neutrophils and stored in their granules. It has been reported that neutrophilic elastase in the bronchial epithelium has a direct effect against bacteria; in addition, it can regulate the increased expression of the HBD2 [84].
7.5.6 Dexamethasone
It is a synthetic glucocorticoid that is used in the treatment of respiratory, allergic or autoimmune diseases. Its effect is to decrease the expression of proinflammatory cytokine genes through NF-kB. The clinical use of glucocorticoids can increase the susceptibility to infections by decreasing the expression of antimicrobial peptides such as the HBD2. In this study, they investigated the molecular mechanism by which dexamethasone modulated HBD2 expression in response to IL-1b in A549 cells and, the role of MAPKs, MKP-1, AKT, and NF-kB transcription factor. They demonstrated that dexamethasone suppresses the expression of HBD2 for this signaling pathway [83].
7.5.7 Isoleucine
Isoleucine is an essential amino acid that can induce the expression of the HBD2 in the epithelium. Its expression involves the activation of NF-kB/rel family of trans-activating factors. The authors suggest that isoleucine or analogues may have clinical utility as immunostimulants that could bolster the defense of the respiratory epithelium and mucosae [85].
7.5.8 Hyaluronic acid
When the skin epithelium suffers damage, the hyaluronic acid, which is found in the extracellular matrix, is fragmented and activates keratinocytes which in turn stimulate the HBD2 production. The induction is mediated by toll receptors (TLR2 and TLR4) as well as other signaling pathways such as c-Fos and protein kinase C; thus, the epithelium is protected from infections [86].
7.5.9 Sirtuin1 (SIRT1)
It is a nicotinamide adenine dinucleotide-dependent histone deacetylase, which regulates several processes of the innate and adaptive immune system. The anti-inflammatory properties of SIRT1 are reportedly known to show [87] that the infection with S. pneumoniae in alveolar epithelial cells (A549) induces HBD2 production involving the SIRT1. Furthermore, HBD2 production induced by S pneumoniae is mediated by the MAPK activation (p38), and the expression of IL-8 is regulated by the phosphorylation of ERK.
7.6 Wound repair
This process has been described in the epithelium of skin and can be achieved by various means, which includes the modulation of cytokines production, cell proliferation and migration and in some cases angiogenesis [88, 89]. It has been demonstrated that HBD2 is expressed in normal skin [90], and its expression increases when the skin is damaged or during the chronic infection [91].
Patients with diabetes mellitus suffer from skin ulcers, and the expression of HBD2 does not increase when compared to normal skin. The scarce expression of HBD2 is seen during the chronic disease. The authors supposed that high glucose levels inhibit the expression of HBD2 in human keratinocytes [92].
HBD2 is reportedly seen to elicit intracellular Ca+2 mobilization and increased keratinocyte migration and proliferation [93]. Besides, this peptide induced phosphorylation of EGFR, signal transducer and activator of transcription STAT1 and STAT3. These are intracellular signaling molecules involved in keratinocyte migration and proliferation.
7.7 Angiogenesis
It is a process by which endothelial cells proliferate and migrate towards the angiogenic stimulus to form new blood vessels. This process is part of the repair of damaged tissues and severe inflammatory processes. The present paper demonstrates that HBD2 stimulates the migration, proliferation and formation of capillary tubes of endothelial cells [94].
7.8 Cytokines and chemokines
Peripheral blood mononuclear cells when stimulated by the HBD2 induce a strong cytokine response. The cytokines that were detected in the highest concentration were interleukin 6 (IL-6), interleukin 8 (IL-8) and interleukin 10 (IL10). It was also found that monocyte chemotactic protein 1 (MCP-1) induces a strong response and also induces RANTES, IL-1β, ENA-78 and GRO, so that the induction patterns of cytokines/chemokines can be crucial in the development and amplification of the immune response against pathogenic microorganisms [95].
7.9 Hypertension
It has been proven that HBD2 can regulate blood pressure and provide a new mechanism for the treatment of hypertension [96].
8. Respiratory diseases associated with the expression of β-defensin-2
The respiratory epithelium is the largest surface of the human body in contact with external medium, and it exposed to a large number of pathogens. The respiratory epithelium counts upon many defense effectors and one of them is the production of defensins. They act as a first line of host defense against invading microorganism. The cells of respiratory epithelium produce four types of defensins (HBD1–4). HBD1 is expressed constitutively and HBD2–4 are expressed during infectious or inflammatory processes. HBD2 is the most expressed in all respiratory epithelium from the oral cavity, pharynx, larynx, trachea, serous cells and submucosal glands to the lung [1, 29, 97]. Several authors have reported expression of the HBD2 in these tissues and also detected the protein in healthy and diseased people in different secretions of the body such as bronchoalveolar fluid, saliva, blood, plasma, milk, sputum, nasal secretions, etc. [32, 98, 99, 100, 101, 102].
Several studies have suggested that there is a relationship between HBD2 and the pathogenesis of several respiratory diseases. The increase or decrease of its expression can result in the protection or amplification of the disease.
8.1 Lung cancer
This type of disease has a high mortality rate and the treatments are very expensive. It is difficult to detect early stages of the disease or if the tumor is benign or malignant, especially in the preliminary condition. The higher concentrations of HBD2 in the serum of patients did not show any relation with the histopathological classification [103].
8.2 Pneumonia
It is a disease of the lower respiratory system that mainly affects young children and older adults and has a high mortality rate and a great social and economic impact due to long periods of hospitalization and resistance to antibiotics. The alteration of the immune function of the mucosa of the respiratory system in these age groups constitutes a risk factor for contracting pneumonia. Bacterial infections are frequent, such infections lead to a serious deterioration of lung function, and are usually associated with persistent colonization by bacteria such as Pseudomonas aeruginosa, Haemophilus influenzae, Streptococcus pneumonia and Legionella pneumophila. Patients with acute pneumonia caused by bacterial infections have shown that the HBD2 is expressed in lung tissue and the concentration of the protein increases in blood and alveolar fluid [98, 101]. When the HBD2 levels in plasma are below 12.5 mg/ml, the patients with pneumonia have a high possibility of needing mechanical ventilation and development of new complications or death [104].
In an in vitro model, L. pneumophila induces the release of hBD-2 in A549 cells in a manner dependent on TLR2 and TLR5. The activation of p38 MAPK and JNK, as well as NF-B and AP-1, is involved in the production of hBD-2 induced by L. pneumophila. Therefore, regulation of the release of hBD-2 by L. pneumophila in A549 cells appears to be determined by multiple signaling molecules and may contribute to host defense in Legionnaires disease. [57].
8.3 Cystic fibrosis
Cystic fibrosis is a chronic lung disease with high morbidity and mortality rates. It is caused by a recessive mutation in the transmembrane conductance regulator gene (CFTR), which is located on chromosome 7 on the apical surface of epithelial cells [19, 24, 105]. This mutation causes abnormal transport of chlorine ions, which induce an increase in the salinity of the alveolar fluid. The high salt concentration abrogates the antimicrobial activity of HBD2; this might explain the recurrent bacterial infections in the lungs of these patients [32, 105].
Bacterial infections in patients with cystic fibrosis are very common; in the acute phase, the patients are infected mainly with Haemophilus influenzae and Staphylococcus aureus. While in the chronic phase, the infection is always caused by Pseudomonas aeruginosa, a Gram-negative bacterium, opportunistic and resistant to antibiotics [32].
The decrease in the expression and degradation of HBD2 plays an important role in the pathogenesis of pulmonary infection for P. aeruginosa in patients with cystic fibrosis [105]. They explain that in the chronic phase of infection, the phenotype of the bacteria changes and it loses the flagella. The flagella are composed of flagellin, which is a virulence factor that promotes mobility and adhesion. The flagellum is the ligand that triggers the signaling pathway for the expression of HBD2. When losing the flagella, the bacterium does not recognize the receptor (TLR5) of the epithelium and transcription of the gene is not executed through NF-kB. The bacterium causes damage to the epithelium of the lung producing a severe inflammatory process, with production of IL-8. IL-8 causes the accumulation of neutrophils; later, they enter apoptosis and phagocytosed by macrophages. Many macrophages accumulate in the lung and secrete cathepsin, a cysteine protease that remodels the extracellular matrix found in large quantities in bronchoalveolar lavage. This enzyme degrades the disulfide bonds of the defensins by inactivating them, thus losing their antimicrobial activity [9].
The patients with cystic fibrosis and polyps showed high levels of HBD2 and TLR2 expression with an increase in IL-8 [106]. In nasal epithelium of patients with CF, HBD2 is not upregulated in response to inflammatory stimuli. The higher levels of inflammatory markers were seen but without any correlation with the expression of HBD2. They explain that the epithelium of patients with cystic fibrosis is chronically exposed to inflammatory stimuli and lost the capacity to upregulate defensin synthesis. The inflammatory stimuli may not be the sole inducers of defense expression [107].
8.4 Chronic obstructive pulmonary disease (COPD)
COPD is an inflammatory disease of the respiratory tract that is characterized by recurrent infections, severe inflammation and is associated with a reduction of airflow and a decrease in lung function [108]. The main environmental risk factor is smoking; some authors suggest that tobacco smoke inhibits the activation of the host’s innate immune system. The in vitro studies with respiratory epithelium exposed to tobacco smoke are reported to inhibit the expression of HBD2 when infected with bacterium [109]. COPD patients who smoke have lowered basal hBD-2 expression in the epithelial cells of the central airways, which correlates with the amount of cigarette pack years. The decreased HBD2 expression is associated with current or former smoking, which makes the population more susceptible towards infections by microorganisms [108, 110].
Moreover, genetic factors can contribute to the progression of the disease. The variation in the number of copies of HBD2 gene in epithelial cells is associated with the pathogenesis of the disease [110]. Moreover, genetic factors can contribute to the progression of the disease. The variation in the number of copies of HBD2 gene in epithelial cells is associated with the pathogenesis of COPD. In this study showed a significantly higher proportion of the patients with severe COPD had high diploid β-defensin copy numbers (five or more) compared with the control sample.
In another study, the expression of HBD2 in peripheral lung tissue in patients with COPD is elevated and associated with the habit of smoking and with the high levels of IL-8 as well as severity of the disease [111].
8.5 Allergic rhinitis (AR)
AR is an inflammatory disorder that occurs in the nasal mucosa and triggered by the exposure to allergens (mites, pets, insects, pollen, latex items, tobacco particles, ozone, nitrogen oxide, sulfur dioxide, aspirin, etc.) producing inflammation mediated by IgE. Clinically, it is characterized by symptoms such as rhinorrhea, sneezing, nasal congestion and itching. These symptoms worsen a person’s productivity and quality of life and cause sleep disturbances, fatigue or depression. Although it occurs more frequently in young children, adults are affected as well [18].
Studies with tonsil tissue (lymphocytes) and pharyngeal epithelial cells from patients with allergic rhinitis found a reduction in the expression of HBD2. When cells are exposed to Th2 cytokines (IL-4, IL-5 and IL-13) and histamine in vitro, they also cause a decrease in the expression of HBD2. Therefore, patients with allergic rhinitis are more susceptible to respiratory infections and severe exacerbations [112, 113, 114].
8.6 Rhinosinusitis
Rhinosinusitis is characterized by the presence of at least two respiratory symptoms, nasal obstruction and nasal discharge, wherein the presence or absence of nasal polyps is seen. In one study, the sinonasal epithelial cells of patients with rhinosinusitis with nasal polyps showed a decrease in the expression of β-defensin-2 in response to the presence of IL-4 and IL-13 [114, 115, 116].
8.7 Otitis media
It is a very common disease mainly affecting the young children under 3 years of age having episode of otitis media. The pathogenesis of the disease is multifactorial, and among the most important factors are viral infections (respiratory syncytial virus, rhinovirus, influenza A, adenovirus) and bacterial infections (Streptococcus pneumoniae, nontypeable Haemophilus influenzae (NTHI), and Moraxella catarrhalis). The acute illness resolves quickly but the chronic or recurrent disease can cause hearing loss [39, 117].
The epithelial cells of the middle ear express HBD2, and two signaling pathways have been described. HBD2 expression is induced by pro-inflammatory stimuli such as interleukin 1 alpha (IL-1a), tumor necrosis factor alpha (TNF-a), and lipopolysaccharide (LPS). Their transcriptional activation is mediated through an Src-dependent Raf-MEK1/2-ERK signaling pathway [119]. The other known route is when the bacterium is recognized by the TLR2 of epithelial cells through the MyD88-IRAKI-TRAF6-MKK3/6-p38 MAP kinase signal transduction pathway [117, 118].
8.8 Asthma
It is a common chronic obstructive disease characterized by obstruction, hyperreactivity and inflammation. It affects all age groups throughout the world and has high morbidity and mortality rates [119]. Epidemiological studies indicate that the pathogenesis of asthma is multifactorial. The viral respiratory infections are the main cause of exacerbations or asthma attacks; influenza viruses, parainfluenza, rhinovirus and respiratory syncytial are those that have been identified more frequently [120, 121].
The mechanisms that explain how viruses cause or exacerbate asthma are diverse and some are not yet fully characterized. Recent studies indicate that HBD2 may play an important role in the pathogenesis of asthma. Some mechanisms have been suggested to explain the exacerbations of asthma such as (a) patients with asthma develop a TH2 type response; it induced the production of cytokines (IL-4, IL-5, IL-13) that inhibit the production of HBD2, causing susceptibility to infections [122]. (b) HBD2 induces the activation and degranulation of mastoid cells that release histamine and prostaglandins, substances that increase asthma exacerbations [26].
8.9 Recurrent respiratory papillomatosis
It is a disease in the respiratory tract caused by infection with human papilloma virus. The papilloma virus belongs to the Papovaviridae family. It is a circular double-stranded DNA virus and has no envelope, and its genome is 7200–8000 bp. There are more than 100 serotypes, which due to their oncogenic capacity are divided into high risk and low risk [123].
The disease characterized by abnormal proliferation of epithelial keratinocytes forming a papilloma. It occurs more frequently in the larynx but might spread to the trachea or even lungs. It is common in children 2–4 years of age and in children over 12 years of age and in young adults. The first symptom is progressive dysphonia, with symptoms of variable respiratory obstruction, dyspnea and stridor. Some patients have spontaneous remission and others may have a very rapid growth that may require multiple surgical procedures or cause obstruction resulting in death. Papilloma viruses have a high morbidity rate because of its recurrence, and there are no satisfactory treatments [124, 125].
The serotypes of papilloma that have been found with higher frequency are the serotypes of low risk 6 and 11; nevertheless, a 2% can be infected with other subtypes like the 16 and 18 of high risk, but these are associated with transformation of cells causing carcinoma [123].
The expression of HBD2 in samples of papillomavirus induced lesions in patients with recurrent respiratory papillomatosis, and its association with IL-8 [126] was studied. The lesions showed high levels of HBD2 expression, and the inflammatory process was not significant. Despite the higher expression of HBD2 in the deep-seated tissues, the persistence of the virus leads to the disease progression. They suggest that HBD2 can serve as a signaling molecule to induce the adaptive immune response against viral infection with acceptable inflammatory events.
8.10 Diffuse panbronchiolitis (DP)
DP is a disease with chronic inflammation in the bronchioles of respiratory system. The immune cells are activated, and neutrophils are found in large quantities and activate T lymphocytes. There are often bacterial infections (P. aeruginosa and H influenzae). The pathogenesis of this disease is not clear; but recently, HBD2 has been implicated. In a study with patients with DP, high levels of HBD2 in bronchoalveolar fluid and plasma were found, suggesting that it could play an important role in the defense against infections. The levels of HBD2 in BAL fluid may be a useful marker of airway inflammation in patients with DPB [127].
8.11 Tuberculosis
Tuberculosis is one of the most frequent infectious diseases that cause more deaths; its incidence is still a global health problem. Mycobacterium tuberculosis is the causative agent of tuberculosis, and it can cause a progressive disease or a latent infection. It has been reported that a third part of the population is latently infected and 10% have active disease. There are many difficulties due to highly resistant strains and fewer therapeutic agents [128].
However, HBD2 is suggested to play an important role in the control of the disease by direct inactivation of the bacteria or as an immunostimulant in vaccines. This bacterium mainly infects macrophages and lung epithelial cells. The in vitro studies have shown that M. tuberculosis induces the expression of HBD2 in human lung epithelial cells (A549) and is associated with the destruction of bacteria [129, 130].
The transfection of monocytes derived from macrophages with the HBD2 gene increases the ability to control the growth of M tuberculosis when compared with non-transfected cells [131]. Children infected with M. tuberculosis present high concentrations of HBD2 in bronchoalveolar lavage suggesting its involvement in the pathogenesis of disease.
Results are favorable in animal model studies in mice, mice were vaccinated with DNA vaccines (gene encoding β-defensin-2 and antigens of M tuberculoisis) and months later were challenged with M tuberculoisis strains. The level of protection was evaluated by survival, and tissue damage. DNA vaccines showed protection with significant higher survival and less tissue damage in the mice [132]. When a person is infected with M. tuberculosis, the microenvironment is reduced in oxygen; this triggers the expression of the vitamin D receptor and HBD2 inhibits the growth bacteria. The lack of local oxygen benefits the macrophage for its elimination.
Other signaling pathways that have been discovered are in monocytes through the CD40L and IFN receptors that converge in the activation of vitamin D. HBD2 is expressed and acts against the bacterium M. tuberculosis [134].
8.12 Sepsis
The infectious diseases can cause severe sepsis and the patient’s immune system to suffer drastic changes. This study investigated the concentration and the expression of HBD2 in peripheral whole blood cells from patients with severe sepsis. They detected that the peripheral blood cells expressed a decrease in the expression of HBD2. The patient serum showed higher concentrations of HBD2. The patients with severe sepsis have severe inflammatory processes and the proinflammatory cytokines are at high concentrations (IL-1 and TNF); these cytokines also induce the expression of HBD2. It is suggested that these cytokines may be involved in the overproduction of HBD2 in the blood of patients with sepsis. The decreased HBD2 induced in peripheral blood cells was not associated with decreased plasma levels, suggesting that peripheral blood cells do not represent the exclusive source of released protein [135].
9. Antiviral activity of β-defensin-2 in respiratory infections
The HBD2 was initially studied as an antibacterial peptide, and its antiviral activity has been recently demonstrated. The notion was that it only acted in enveloped viruses, but other studies have proven that they also act against naked viruses and any type of genome DNA, RNA and retroviruses. The mechanisms of antiviral inactivation have been classified as direct and indirect. The direct ones occur when HBD2 binds to the viral membrane by electrostatic attraction, causing pore formation and lysis of microorganism. The indirect mechanism occurs when it inactivates an intracellular pathway of the virus replication cycle or when there is recruitment of immune cells that contribute to their antiviral activity [2].
Acute respiratory infections caused by viruses are very common, causing high rates of mobility and mortality. There are few studies that relate to HBD2 and viral infection, and some of them are presented in the following paragraphs.
9.1 Influenza virus
The influenza virus belongs to the Orthomyxoviridae family and renders acute respiratory infection affecting mainly children and adults. It is an RNA virus, enveloped and has the ability to mutate rapidly, causing epidemics and pandemics [136].
Influenza virus induces the production of HBD2 in the epithelial cells of the respiratory tract in vivo and in vitro [137]. In the in vitro study with infected MDCK cells with influenza virus, recombinant murine β-defensin-2 prevents infection by blocking the entry of the virus. The lungs of murine model infected with influenza virus showed an increased expression of β-defensin and therefore confer protection against infection [21, 138].
9.2 Respiratory syncytial virus (VSR)
This virus belongs to the family Paramyxoviridae and affects principally infants, young children and older adults causing bronchiolitis and pneumonia. VSR has a high rate of morbidity and mortality having RNA enveloped virus of negative sense. There are no vaccines available and only fewer antivirals available which have been seen ineffective [22, 139].
The human lung cells (A549) infected with RSV expressed HBD2 [140]. The expression of the peptide depends on the activation of NF-kB and the action of TNF produced by the virus. The elimination of the virus is due to damage to the membrane.
9.3 Adenovirus
The adenoviruses belong to the Adenoviridae family; 51 serotypes divided into six species have been recognized (HAd A-F). Species B, C and E produce respiratory infections. Adenoviruses are double-stranded linear DNA viruses, lack envelope and replicate in the nucleus, and their genome has a size of 36 kb. These viruses spread rapidly in closed environments such as military camps, orphanages, boarding schools and prisons. The acute respiratory infections are transmitted mainly by aerosols and by direct inoculation through fingers. Although this disease is benign and with little severity, the infection might be severe in immunosuppressed patients afflicted with HIV and with kidney transplants. The HBD2 inactivates the infection for adenovirus in vitro; the mechanisms of action are not yet known due to lack of in vivo studies [140].
9.4 Rhinovirus
The rhinovirus is the main cause of the common cold belonging to Picornaviridae family. They are small, single-chained, naked RNA viruses. Respiratory infections caused by rhinoviruses are associated with asthma exacerbations in children and adults. Rhinovirus infection in the A549 cells (human lung cells) and bronchial epithelial cells induces the expression of HBD2. The virus replication appears essential for the expression of peptide [141].
10. Conclusions
Initially, β-defensin-2 was considered an antimicrobial peptide. The advance in the study of these molecules has made it possible to know that β-defensin-2 is a peptide with multiple functions. However, its role in the pathology of respiratory diseases is unclear. During the infectious processes of the respiratory epithelium, HBD2 is expressed by the effect of the infection, and it has been described that it may be closely associated with the severity of the disease. The study of HBD2 during viral respiratory infections is unclear, so it is necessary to continue investigating the role of this molecule in the immune response activated by viral infections.
\n',keywords:"β-defensin-2, pathogenesis, respiratory diseases, viral infections",chapterPDFUrl:"https://cdn.intechopen.com/pdfs/63967.pdf",chapterXML:"https://mts.intechopen.com/source/xml/63967.xml",downloadPdfUrl:"/chapter/pdf-download/63967",previewPdfUrl:"/chapter/pdf-preview/63967",totalDownloads:1143,totalViews:147,totalCrossrefCites:1,dateSubmitted:"April 30th 2018",dateReviewed:"July 31st 2018",datePrePublished:"November 5th 2018",datePublished:"April 17th 2019",dateFinished:"October 8th 2018",readingETA:"0",abstract:"Human β-defensin-2 is a small cationic peptide that is part of the innate and adaptive immunity. It is expressed mainly in the epithelium and has a broad spectrum of antimicrobial activity against bacteria, fungi and viruses. In addition to its antimicrobial activity, it has other biological functions. The alteration of the expression of β-defensin-2 in the respiratory epithelium has been associated with the pathogenesis of several respiratory diseases such as asthma, pulmonary fibrosis, pneumonia, tuberculosis, rhinitis, etc. The acute respiratory infections caused by viruses are the main cause of morbidity and mortality in the world; there are few studies and it is necessary to study this peptide to understand its role in the viral pathogenesis. In addition, it also becomes relevant in its potential to take advantage of its properties in the development of alternative therapies that allow the prevention or treatment of viral respiratory infections.",reviewType:"peer-reviewed",bibtexUrl:"/chapter/bibtex/63967",risUrl:"/chapter/ris/63967",signatures:"Dora Patricia Rosete Olvera and Carlos Cabello Gutiérrez",book:{id:"6963",type:"book",title:"Immune Response Activation and Immunomodulation",subtitle:null,fullTitle:"Immune Response Activation and Immunomodulation",slug:"immune-response-activation-and-immunomodulation",publishedDate:"April 17th 2019",bookSignature:"Rajeev K. Tyagi and Prakash S. Bisen",coverURL:"https://cdn.intechopen.com/books/images_new/6963.jpg",licenceType:"CC BY 3.0",editedByType:"Edited by",isbn:"978-1-78985-152-6",printIsbn:"978-1-78985-151-9",pdfIsbn:"978-1-83962-135-2",isAvailableForWebshopOrdering:!0,editors:[{id:"201069",title:"Dr.",name:"Rajeev",middleName:"K.",surname:"Tyagi",slug:"rajeev-tyagi",fullName:"Rajeev Tyagi"}],productType:{id:"1",title:"Edited Volume",chapterContentType:"chapter",authoredCaption:"Edited by"}},authors:[{id:"95406",title:"Prof.",name:"Dora Patricia",middleName:null,surname:"Rosete",fullName:"Dora Patricia Rosete",slug:"dora-patricia-rosete",email:"Dorosete67@yahoo.com.mx",position:null,profilePictureURL:"//cdnintech.com/web/frontend/www/assets/author.svg",institution:{name:"Instituto Nacional de Enfermedades Respiratorias",institutionURL:null,country:{name:"Mexico"}}},{id:"259774",title:"Dr.",name:"Carlos",middleName:null,surname:"Cabello-Gutierrez",fullName:"Carlos Cabello-Gutierrez",slug:"carlos-cabello-gutierrez",email:"carloscginer@gmail.com",position:null,profilePictureURL:"//cdnintech.com/web/frontend/www/assets/author.svg",institution:{name:"Instituto Nacional de Enfermedades Respiratorias",institutionURL:null,country:{name:"Mexico"}}}],sections:[{id:"sec_1",title:"1. Introduction",level:"1"},{id:"sec_2",title:"2. General characteristics and classification",level:"1"},{id:"sec_3",title:"3. Molecular structure of β-defensin-2",level:"1"},{id:"sec_4",title:"4. Genetic structure of β-defensin-2",level:"1"},{id:"sec_5",title:"5. Expression and regulation of β-defensin-2",level:"1"},{id:"sec_6",title:"6. Mechanisms of action of β-defensin-2",level:"1"},{id:"sec_7",title:"7. Multifunctional activities of β-defensin-2",level:"1"},{id:"sec_7_2",title:"7.1 Antimicrobial activities",level:"2"},{id:"sec_8_2",title:"7.2 Innate and adaptive immunity",level:"2"},{id:"sec_9_2",title:"7.3 Inflammation",level:"2"},{id:"sec_10_2",title:"7.4 Anti-inflammatory activity",level:"2"},{id:"sec_11_2",title:"7.5 Immunomodulatory properties",level:"2"},{id:"sec_11_3",title:"7.5.1 Moxifloxacin/HBD2",level:"3"},{id:"sec_12_3",title:"7.5.2 Vitamin D (VitD)",level:"3"},{id:"sec_13_3",title:"7.5.3 PARs (receptors coupled to proteases)",level:"3"},{id:"sec_14_3",title:"7.5.4 Triptolide",level:"3"},{id:"sec_15_3",title:"7.5.5 Neutrophilic elastase",level:"3"},{id:"sec_16_3",title:"7.5.6 Dexamethasone",level:"3"},{id:"sec_17_3",title:"7.5.7 Isoleucine",level:"3"},{id:"sec_18_3",title:"7.5.8 Hyaluronic acid",level:"3"},{id:"sec_19_3",title:"7.5.9 Sirtuin1 (SIRT1)",level:"3"},{id:"sec_21_2",title:"7.6 Wound repair",level:"2"},{id:"sec_22_2",title:"7.7 Angiogenesis",level:"2"},{id:"sec_23_2",title:"7.8 Cytokines and chemokines",level:"2"},{id:"sec_24_2",title:"7.9 Hypertension",level:"2"},{id:"sec_26",title:"8. Respiratory diseases associated with the expression of β-defensin-2",level:"1"},{id:"sec_26_2",title:"8.1 Lung cancer",level:"2"},{id:"sec_27_2",title:"8.2 Pneumonia",level:"2"},{id:"sec_28_2",title:"8.3 Cystic fibrosis",level:"2"},{id:"sec_29_2",title:"8.4 Chronic obstructive pulmonary disease (COPD)",level:"2"},{id:"sec_30_2",title:"8.5 Allergic rhinitis (AR)",level:"2"},{id:"sec_31_2",title:"8.6 Rhinosinusitis",level:"2"},{id:"sec_32_2",title:"8.7 Otitis media",level:"2"},{id:"sec_33_2",title:"8.8 Asthma",level:"2"},{id:"sec_34_2",title:"8.9 Recurrent respiratory papillomatosis",level:"2"},{id:"sec_35_2",title:"8.10 Diffuse panbronchiolitis (DP)",level:"2"},{id:"sec_36_2",title:"8.11 Tuberculosis",level:"2"},{id:"sec_37_2",title:"8.12 Sepsis",level:"2"},{id:"sec_39",title:"9. Antiviral activity of β-defensin-2 in respiratory infections",level:"1"},{id:"sec_39_2",title:"9.1 Influenza virus",level:"2"},{id:"sec_40_2",title:"9.2 Respiratory syncytial virus (VSR)",level:"2"},{id:"sec_41_2",title:"9.3 Adenovirus",level:"2"},{id:"sec_42_2",title:"9.4 Rhinovirus",level:"2"},{id:"sec_44",title:"10. Conclusions",level:"1"}],chapterReferences:[{id:"B1",body:'Gallo RL, Murakami M, Ohtake T, Zaiou M. Biology and clinical relevance of naturally occurring antimicrobial peptides. The Journal of Allergy and Clinical Immunology. 2002;110:823-831. DOI: 10.1067/mai.2002.129801'},{id:"B2",body:'Diamond G, Beckloff N, Ryan LK. Host defense peptides in the oral cavity and the lung: Similarities and differences. Journal of Dental Research. 2008;87:915-927. DOI: 10.1177/154405910808701011'},{id:"B3",body:'Hazlett L, Wu M. Defensins in innate immunity. Cell and Tissue Research. 2011;343:175-188. DOI: 10.1007/s00441-010-1022-4'},{id:"B4",body:'Schrόder JM, Harder J. 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DOI: 10.1016/S0928-8244 (03) 00106-8'}],footnotes:[],contributors:[{corresp:null,contributorFullName:"Dora Patricia Rosete Olvera",address:null,affiliation:'
Department of Virology and Mycology, National Institute of Respiratory Diseases, Ismael Cosío Villegas, Ciudad de México, Mexico
Department of Virology and Mycology, National Institute of Respiratory Diseases, Ismael Cosío Villegas, Ciudad de México, Mexico
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Tsvetkov"}],equalEditorOne:null,equalEditorTwo:null,equalEditorThree:null,productType:{id:"1",chapterContentType:"chapter",authoredCaption:"Edited by"}}],booksByTopicTotal:16,seriesByTopicCollection:[],seriesByTopicTotal:0,mostCitedChapters:[{id:"66882",doi:"10.5772/intechopen.85919",title:"World’s Demand for Food and Water: The Consequences of Climate Change",slug:"world-s-demand-for-food-and-water-the-consequences-of-climate-change",totalDownloads:2126,totalCrossrefCites:10,totalDimensionsCites:35,abstract:"This study focused on analysis of global food demand and supply situation by 2030 and 2050, water demand-availability, impact of climate change on world water resource, food security and desalination challenges and development opportunities. The population of the world will be 8.6 billion in 2030 and 9.8 billion in 2050; Africa will be the major contributor. World cereal equivalent (CE) food demand is projected to be around 10,094 million tons in 2030 and 14,886 million tons in 2050, while its production is projected to be 10,120 million tons in 2030 and 15,970 million tons in 2050 having a marginal surplus. India and China are capturing large share of global food demand. The developing country will demand more animal origin foods due to income growth in the future. The growth rate of world demand for cereals will decline till 2050. Global water demand is projected to increase by 55% between 2000 and 2050 from 3500 to 5425 km3. Evidence showed that climate change will have adverse impact on world water resources and food production with high degree of regional variability and scarcity. A number of options are suggested for development of global water resource and food production.",book:{id:"7645",slug:"desalination-challenges-and-opportunities",title:"Desalination",fullTitle:"Desalination - Challenges and Opportunities"},signatures:"Sheikh Mohammad Fakhrul Islam and Zahurul Karim",authors:[{id:"288119",title:"Prof.",name:"S.M. Fakhrul",middleName:null,surname:"Islam",slug:"s.m.-fakhrul-islam",fullName:"S.M. Fakhrul Islam"},{id:"288121",title:"Prof.",name:"Zahurul",middleName:null,surname:"Karim",slug:"zahurul-karim",fullName:"Zahurul Karim"}]},{id:"60850",doi:"10.5772/intechopen.76624",title:"Wastewater Treatment Using Membrane Technology",slug:"wastewater-treatment-using-membrane-technology",totalDownloads:2961,totalCrossrefCites:14,totalDimensionsCites:29,abstract:"Water contamination by heavy metals, cyanides and dyes is increasing globally and needs to be addressed as this will lead to water scarcity as well as water quality. Different techniques have been used to clean and renew water for human consumption and agricultural purposes but they each have limitations. Among those techniques, membrane technology is promising to solve the issues. Nanotechnology present a great potential in wastewater treatment to improve treatment efficiency of wastewater treatment plants. In addition, nanotechnology supplement water supply through safe use of modern water sources. This chapter reviews recent development in membrane technology for wastewater treatment. Different types of membrane technologies, their properties, mechanisms advantages, limitations and promising solutions have been discussed.",book:{id:"6539",slug:"wastewater-and-water-quality",title:"Wastewater and Water Quality",fullTitle:"Wastewater and Water Quality"},signatures:"Azile Nqombolo, Anele Mpupa, Richard M. Moutloali and Philiswa\nN. Nomngongo",authors:[{id:"191669",title:"Dr.",name:"Philiswa",middleName:null,surname:"Nomngongo",slug:"philiswa-nomngongo",fullName:"Philiswa Nomngongo"}]},{id:"9113",doi:"10.5772/7588",title:"Photocatalytic Processes on the Oxidation of Organic Compounds in Water",slug:"photocatalytic-processes-on-the-oxidation-of-organic-compounds-in-water",totalDownloads:5773,totalCrossrefCites:3,totalDimensionsCites:24,abstract:null,book:{id:"3704",slug:"new-trends-in-technologies",title:"New Trends in Technologies",fullTitle:"New Trends in Technologies"},signatures:"C. J. Philippopoulos and M. D Nikolaki",authors:null},{id:"70242",doi:"10.5772/intechopen.90256",title:"Advancements in the Fenton Process for Wastewater Treatment",slug:"advancements-in-the-fenton-process-for-wastewater-treatment",totalDownloads:1873,totalCrossrefCites:9,totalDimensionsCites:21,abstract:"Fenton is considered to be one of the most effective advanced treatment processes in the removal of many hazardous organic pollutants from refractory/toxic wastewater. It has many advantages, but drawbacks are significant such as a strong acid environment, the cost of reagents consumption, and the large production of ferric sludge, which limits Fenton’s further application. The development of Fenton applications is mainly achieved by improving oxidation efficiency and reducing sludge production. This chapter presents a review on fundamentals and applications of conventional Fenton, leading advanced technologies in the Fenton process, and reuse methods of iron containing sludge to synthetic and real wastewaters are discussed. Finally, future trends and some guidelines for Fenton processes are given.",book:{id:"9415",slug:"advanced-oxidation-processes-applications-trends-and-prospects",title:"Advanced Oxidation Processes",fullTitle:"Advanced Oxidation Processes - Applications, Trends, and Prospects"},signatures:"Min Xu, Changyong Wu and Yuexi Zhou",authors:[{id:"307479",title:"Dr.",name:"Changyong",middleName:null,surname:"Wu",slug:"changyong-wu",fullName:"Changyong Wu"},{id:"307546",title:"Prof.",name:"Yuexi",middleName:null,surname:"Zhou",slug:"yuexi-zhou",fullName:"Yuexi Zhou"},{id:"311139",title:"Dr.",name:"Min",middleName:null,surname:"Xu",slug:"min-xu",fullName:"Min Xu"}]},{id:"67689",doi:"10.5772/intechopen.86952",title:"Membrane Distillation: Basics, Advances, and Applications",slug:"membrane-distillation-basics-advances-and-applications",totalDownloads:1444,totalCrossrefCites:9,totalDimensionsCites:20,abstract:"Membrane technology as an emerging separation process has become competitive with other separation techniques in recent decades. Among pressure-driven and isothermal membrane processes, membrane distillation (MD) as a thermally driven process has come out to put an end to hardships of such processes like distillation. MD process can be used in a wide variety of applications such as desalination and wastewater treatment. Generally, MD is a process which water is a main component of the feed solution and only water vapor can pass through a hydrophobic membrane pores. With four main configurations different from each other by their condensation procedure, the performance of MD process is limited due to the lack of appropriate module, membrane, and energy consumption rate. In recent years, many experiments have been carried out to find well-suited membrane type and module. Also, applying solar or waste heat as heat source and the capability of coupling with other processes like forward osmosis and osmotic distillation distinguish MD process from other membrane processes. This chapter addresses membrane characteristics, MD applications, transport mechanisms, and process challenges.",book:{id:"8915",slug:"advances-in-membrane-technologies",title:"Advances in Membrane Technologies",fullTitle:"Advances in Membrane Technologies"},signatures:"Mohammad Reza Shirzad Kebria and Ahmad Rahimpour",authors:[{id:"289042",title:"Associate Prof.",name:"Ahmad",middleName:null,surname:"Rahimpour",slug:"ahmad-rahimpour",fullName:"Ahmad Rahimpour"},{id:"289043",title:"Mr.",name:"Mohammad Reza",middleName:null,surname:"Shirzad Kebria",slug:"mohammad-reza-shirzad-kebria",fullName:"Mohammad Reza Shirzad Kebria"}]}],mostDownloadedChaptersLast30Days:[{id:"70242",title:"Advancements in the Fenton Process for Wastewater Treatment",slug:"advancements-in-the-fenton-process-for-wastewater-treatment",totalDownloads:1873,totalCrossrefCites:9,totalDimensionsCites:21,abstract:"Fenton is considered to be one of the most effective advanced treatment processes in the removal of many hazardous organic pollutants from refractory/toxic wastewater. It has many advantages, but drawbacks are significant such as a strong acid environment, the cost of reagents consumption, and the large production of ferric sludge, which limits Fenton’s further application. The development of Fenton applications is mainly achieved by improving oxidation efficiency and reducing sludge production. This chapter presents a review on fundamentals and applications of conventional Fenton, leading advanced technologies in the Fenton process, and reuse methods of iron containing sludge to synthetic and real wastewaters are discussed. Finally, future trends and some guidelines for Fenton processes are given.",book:{id:"9415",slug:"advanced-oxidation-processes-applications-trends-and-prospects",title:"Advanced Oxidation Processes",fullTitle:"Advanced Oxidation Processes - Applications, Trends, and Prospects"},signatures:"Min Xu, Changyong Wu and Yuexi Zhou",authors:[{id:"307479",title:"Dr.",name:"Changyong",middleName:null,surname:"Wu",slug:"changyong-wu",fullName:"Changyong Wu"},{id:"307546",title:"Prof.",name:"Yuexi",middleName:null,surname:"Zhou",slug:"yuexi-zhou",fullName:"Yuexi Zhou"},{id:"311139",title:"Dr.",name:"Min",middleName:null,surname:"Xu",slug:"min-xu",fullName:"Min Xu"}]},{id:"71660",title:"Applications of Chemical Kinetics in Heterogeneous Catalysis",slug:"applications-of-chemical-kinetics-in-heterogeneous-catalysis",totalDownloads:1104,totalCrossrefCites:3,totalDimensionsCites:3,abstract:"Chemical kinetics is a key subdiscipline of physical chemistry that studies the reaction rate in every elemental step and corresponding catalytic mechanism. It mainly concludes molecular reaction dynamics, catalytic dynamics, elemental reaction dynamics, macrodynamics, and microdynamics. Such a research field has wide applications in heterogeneous catalysis. Based on the Arrhenius plot fitted by the catalytic conversions below 15% without the mass transfer effect and heat transfer effect, the apparent activation energy echoing with the intrinsically catalytic sites and the pre-exponential factor echoing with the relative number of active sites can be, respectively, derived from the slope and intercept of the Arrhenius plots, which can be used to compare the intrinsically catalytic activity of different catalysts and the relative amount of active sites. Reaction orders of both reactants and products are derived from the reaction rate equation and also fitted by the catalytic conversions below 15% without the mass transfer effect and heat transfer effect. According to the acquired reaction orders, the reaction mechanism can be proposed and even defined in some simple reactions. Therefore, investigations of chemical kinetics are of extreme importance and meaning in heterogeneous catalysis.",book:{id:"9415",slug:"advanced-oxidation-processes-applications-trends-and-prospects",title:"Advanced Oxidation Processes",fullTitle:"Advanced Oxidation Processes - Applications, Trends, and Prospects"},signatures:"Zhenhua Zhang, Li-Ping Fan and Yue-Juan Wang",authors:[{id:"312555",title:"Prof.",name:"Zhenhua",middleName:null,surname:"Zhang",slug:"zhenhua-zhang",fullName:"Zhenhua Zhang"},{id:"316868",title:"Ms.",name:"Li-Ping",middleName:null,surname:"Fan",slug:"li-ping-fan",fullName:"Li-Ping Fan"},{id:"316869",title:"Prof.",name:"Yue-Juan",middleName:null,surname:"Wang",slug:"yue-juan-wang",fullName:"Yue-Juan Wang"}]},{id:"77416",title:"Application of Water Quality Index for the Assessment of Water from Different Sources in Nigeria",slug:"application-of-water-quality-index-for-the-assessment-of-water-from-different-sources-in-nigeria",totalDownloads:516,totalCrossrefCites:0,totalDimensionsCites:1,abstract:"Water quality index (WQI) provides a single number that expresses the overall water quality, at a certain location and time, based on several water quality parameters. The objective of WQI is to turn complex water quality data into information that is understandable and usable by the public. A number of indices have been developed to summarize water quality data in an easily expressible and easily understood format. The WQI is basically a mathematical means of calculating a single value from multiple test results. This chapter discusses, in detail, the application of a water quality index for the assessment of water quality to different several water sources in Nigeria.",book:{id:"9921",slug:"promising-techniques-for-wastewater-treatment-and-water-quality-assessment",title:"Promising Techniques for Wastewater Treatment and Water Quality Assessment",fullTitle:"Promising Techniques for Wastewater Treatment and Water Quality Assessment"},signatures:"Ruth Olubukola Ajoke Adelagun, Emmanuel Edet Etim and Oko Emmanuel Godwin",authors:[{id:"256167",title:"Dr.",name:"Emmanuel",middleName:null,surname:"Edet Etim",slug:"emmanuel-edet-etim",fullName:"Emmanuel Edet Etim"},{id:"345734",title:"Mr.",name:"Oko",middleName:null,surname:"Emmanuel Godwin",slug:"oko-emmanuel-godwin",fullName:"Oko Emmanuel Godwin"},{id:"345735",title:"Dr.",name:"Ruth",middleName:null,surname:"Olubukola Ajoke Adelagun",slug:"ruth-olubukola-ajoke-adelagun",fullName:"Ruth Olubukola Ajoke Adelagun"}]},{id:"71348",title:"Water Treatment and Desalination",slug:"water-treatment-and-desalination",totalDownloads:1049,totalCrossrefCites:2,totalDimensionsCites:3,abstract:"Water covers a large area of the earth that reaches about three quarters of the surface of this planet, but we cannot say that all of this water is fresh or drinkable; according to many statistics, the percentage of fresh water reaches about 1% of the total water on earth. But with the great need for fresh water, whether for drinking or other purposes such as agriculture, the search for water treatment methods has become much larger. One of the most important of these methods that have been developed is desalination of seawater using desalination plants; therefore, we will address here the most important methods used in desalination and water treatment.",book:{id:"7645",slug:"desalination-challenges-and-opportunities",title:"Desalination",fullTitle:"Desalination - Challenges and Opportunities"},signatures:"Mona M. Amin Abdel-Fatah and Ghada Ahmed Al Bazedi",authors:[{id:"286268",title:"Associate Prof.",name:"Mona",middleName:null,surname:"Abdel-Fatah",slug:"mona-abdel-fatah",fullName:"Mona Abdel-Fatah"},{id:"295973",title:"Dr.",name:"Ghada",middleName:null,surname:"Al-Basedi",slug:"ghada-al-basedi",fullName:"Ghada Al-Basedi"}]},{id:"69228",title:"Advances in Passive Cooling Design: An Integrated Design Approach",slug:"advances-in-passive-cooling-design-an-integrated-design-approach",totalDownloads:2073,totalCrossrefCites:1,totalDimensionsCites:4,abstract:"Incorporating passive cooling devices within building design requires analysis of device variables and actions to improve cooling performance, maximize efficiency, and integrate with building elements. Improving devices performance requires understanding the relation of devices to design stages, building elements, and working mechanism, and actions performed by devices to enhance cooling process and effectiveness. Therefore, designers could integrate passive devices as intrinsic design elements. The current research introduces SARS as an innovative classification of passive devices based on cooling actions that are performed by a device like storing, avoidance, removal or slowing (SARS). All actions, devices, and variables were discussed and analyzed to help integrate them within design stages: analysis, designing, and performance. Understanding actions will help maximize the performance of the devices, combine two or more devices together, and integrate the devices’ deign in design process. Combining more devices together to perform more than one function will move passive design to a new level to become as whole building design approach and to be a core design element.",book:{id:"8496",slug:"zero-and-net-zero-energy",title:"Zero and Net Zero Energy",fullTitle:"Zero and Net Zero Energy"},signatures:"Ahmed A.Y. Freewan",authors:[{id:"284866",title:"Dr.",name:"Ahmed A.Y.",middleName:null,surname:"Freewan",slug:"ahmed-a.y.-freewan",fullName:"Ahmed A.Y. Freewan"}]}],onlineFirstChaptersFilter:{topicId:"287",limit:6,offset:0},onlineFirstChaptersCollection:[],onlineFirstChaptersTotal:0},preDownload:{success:null,errors:{}},subscriptionForm:{success:null,errors:{}},aboutIntechopen:{},privacyPolicy:{},peerReviewing:{},howOpenAccessPublishingWithIntechopenWorks:{},sponsorshipBooks:{sponsorshipBooks:[],offset:8,limit:8,total:0},allSeries:{pteSeriesList:[{id:"14",title:"Artificial Intelligence",numberOfPublishedBooks:9,numberOfPublishedChapters:87,numberOfOpenTopics:6,numberOfUpcomingTopics:0,issn:"2633-1403",doi:"10.5772/intechopen.79920",isOpenForSubmission:!0},{id:"7",title:"Biomedical Engineering",numberOfPublishedBooks:12,numberOfPublishedChapters:98,numberOfOpenTopics:3,numberOfUpcomingTopics:0,issn:"2631-5343",doi:"10.5772/intechopen.71985",isOpenForSubmission:!0}],lsSeriesList:[{id:"11",title:"Biochemistry",numberOfPublishedBooks:27,numberOfPublishedChapters:287,numberOfOpenTopics:4,numberOfUpcomingTopics:0,issn:"2632-0983",doi:"10.5772/intechopen.72877",isOpenForSubmission:!0},{id:"25",title:"Environmental Sciences",numberOfPublishedBooks:1,numberOfPublishedChapters:9,numberOfOpenTopics:4,numberOfUpcomingTopics:0,issn:"2754-6713",doi:"10.5772/intechopen.100362",isOpenForSubmission:!0},{id:"10",title:"Physiology",numberOfPublishedBooks:11,numberOfPublishedChapters:139,numberOfOpenTopics:4,numberOfUpcomingTopics:0,issn:"2631-8261",doi:"10.5772/intechopen.72796",isOpenForSubmission:!0}],hsSeriesList:[{id:"3",title:"Dentistry",numberOfPublishedBooks:8,numberOfPublishedChapters:129,numberOfOpenTopics:0,numberOfUpcomingTopics:2,issn:"2631-6218",doi:"10.5772/intechopen.71199",isOpenForSubmission:!1},{id:"6",title:"Infectious Diseases",numberOfPublishedBooks:13,numberOfPublishedChapters:107,numberOfOpenTopics:3,numberOfUpcomingTopics:1,issn:"2631-6188",doi:"10.5772/intechopen.71852",isOpenForSubmission:!0},{id:"13",title:"Veterinary Medicine and Science",numberOfPublishedBooks:10,numberOfPublishedChapters:103,numberOfOpenTopics:3,numberOfUpcomingTopics:0,issn:"2632-0517",doi:"10.5772/intechopen.73681",isOpenForSubmission:!0}],sshSeriesList:[{id:"22",title:"Business, Management and Economics",numberOfPublishedBooks:1,numberOfPublishedChapters:12,numberOfOpenTopics:2,numberOfUpcomingTopics:1,issn:null,doi:"10.5772/intechopen.100359",isOpenForSubmission:!0},{id:"23",title:"Education and Human Development",numberOfPublishedBooks:0,numberOfPublishedChapters:0,numberOfOpenTopics:2,numberOfUpcomingTopics:0,issn:null,doi:"10.5772/intechopen.100360",isOpenForSubmission:!1},{id:"24",title:"Sustainable Development",numberOfPublishedBooks:0,numberOfPublishedChapters:10,numberOfOpenTopics:4,numberOfUpcomingTopics:1,issn:null,doi:"10.5772/intechopen.100361",isOpenForSubmission:!0}],testimonialsList:[{id:"6",text:"It is great to work with the IntechOpen to produce a worthwhile collection of research that also becomes a great educational resource and guide for future research endeavors.",author:{id:"259298",name:"Edward",surname:"Narayan",institutionString:null,profilePictureURL:"https://mts.intechopen.com/storage/users/259298/images/system/259298.jpeg",slug:"edward-narayan",institution:{id:"3",name:"University of Queensland",country:{id:null,name:"Australia"}}}},{id:"13",text:"The collaboration with and support of the technical staff of IntechOpen is fantastic. The whole process of submitting an article and editing of the submitted article goes extremely smooth and fast, the number of reads and downloads of chapters is high, and the contributions are also frequently cited.",author:{id:"55578",name:"Antonio",surname:"Jurado-Navas",institutionString:null,profilePictureURL:"https://s3.us-east-1.amazonaws.com/intech-files/0030O00002bRisIQAS/Profile_Picture_1626166543950",slug:"antonio-jurado-navas",institution:{id:"720",name:"University of Malaga",country:{id:null,name:"Spain"}}}}]},series:{item:{id:"24",title:"Sustainable Development",doi:"10.5772/intechopen.100361",issn:null,scope:"
\r\n\tTransforming our World: the 2030 Agenda for Sustainable Development endorsed by United Nations and 193 Member States, came into effect on Jan 1, 2016, to guide decision making and actions to the year 2030 and beyond. Central to this Agenda are 17 Goals, 169 associated targets and over 230 indicators that are reviewed annually. The vision envisaged in the implementation of the SDGs is centered on the five Ps: People, Planet, Prosperity, Peace and Partnership. This call for renewed focused efforts ensure we have a safe and healthy planet for current and future generations.
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\r\n\tThis Series focuses on covering research and applied research involving the five Ps through the following topics:
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\r\n\t1. Sustainable Economy and Fair Society that relates to SDG 1 on No Poverty, SDG 2 on Zero Hunger, SDG 8 on Decent Work and Economic Growth, SDG 10 on Reduced Inequalities, SDG 12 on Responsible Consumption and Production, and SDG 17 Partnership for the Goals
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\r\n\t2. Health and Wellbeing focusing on SDG 3 on Good Health and Wellbeing and SDG 6 on Clean Water and Sanitation
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\r\n\t3. Inclusivity and Social Equality involving SDG 4 on Quality Education, SDG 5 on Gender Equality, and SDG 16 on Peace, Justice and Strong Institutions
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\r\n\t4. Climate Change and Environmental Sustainability comprising SDG 13 on Climate Action, SDG 14 on Life Below Water, and SDG 15 on Life on Land
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\r\n\t5. Urban Planning and Environmental Management embracing SDG 7 on Affordable Clean Energy, SDG 9 on Industry, Innovation and Infrastructure, and SDG 11 on Sustainable Cities and Communities.
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\r\n
\r\n\tThe series also seeks to support the use of cross cutting SDGs, as many of the goals listed above, targets and indicators are all interconnected to impact our lives and the decisions we make on a daily basis, making them impossible to tie to a single topic.
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