These books synthesize perspectives of renowned scientists from the world’s most prestigious institutions - from Fukushima Renewable Energy Institute in Japan to Stanford University in the United States, including Columbia University (US), University of Sidney (AU), University of Miami (USA), Cardiff University (UK), and many others.
\\n\\n
This collaboration embodied the true essence of Open Access by simplifying the approach to OA publishing for Academic editors and authors who contributed their research and allowed the new research to be made available free and open to anyone anywhere in the world.
\\n\\n
To celebrate the 50 books published, we have gathered them at one location - just one click away, so that you can easily browse the subjects of your interest, download the content directly, share it or read online.
IntechOpen and Knowledge Unlatched formed a partnership to support researchers working in engineering sciences by enabling an easier approach to publishing Open Access content. Using the Knowledge Unlatched crowdfunding model to raise the publishing costs through libraries around the world, Open Access Publishing Fee (OAPF) was not required from the authors.
\n\n
Initially, the partnership supported engineering research, but it soon grew to include physical and life sciences, attracting more researchers to the advantages of Open Access publishing.
\n\n\n\n
These books synthesize perspectives of renowned scientists from the world’s most prestigious institutions - from Fukushima Renewable Energy Institute in Japan to Stanford University in the United States, including Columbia University (US), University of Sidney (AU), University of Miami (USA), Cardiff University (UK), and many others.
\n\n
This collaboration embodied the true essence of Open Access by simplifying the approach to OA publishing for Academic editors and authors who contributed their research and allowed the new research to be made available free and open to anyone anywhere in the world.
\n\n
To celebrate the 50 books published, we have gathered them at one location - just one click away, so that you can easily browse the subjects of your interest, download the content directly, share it or read online.
\n'}],latestNews:[{slug:"intechopen-supports-asapbio-s-new-initiative-publish-your-reviews-20220729",title:"IntechOpen Supports ASAPbio’s New Initiative Publish Your Reviews"},{slug:"webinar-introduction-to-open-science-wednesday-18-may-1-pm-cest-20220518",title:"Webinar: Introduction to Open Science | Wednesday 18 May, 1 PM CEST"},{slug:"step-in-the-right-direction-intechopen-launches-a-portfolio-of-open-science-journals-20220414",title:"Step in the Right Direction: IntechOpen Launches a Portfolio of Open Science Journals"},{slug:"let-s-meet-at-london-book-fair-5-7-april-2022-olympia-london-20220321",title:"Let’s meet at London Book Fair, 5-7 April 2022, Olympia London"},{slug:"50-books-published-as-part-of-intechopen-and-knowledge-unlatched-ku-collaboration-20220316",title:"50 Books published as part of IntechOpen and Knowledge Unlatched (KU) Collaboration"},{slug:"intechopen-joins-the-united-nations-sustainable-development-goals-publishers-compact-20221702",title:"IntechOpen joins the United Nations Sustainable Development Goals Publishers Compact"},{slug:"intechopen-signs-exclusive-representation-agreement-with-lsr-libros-servicios-y-representaciones-s-a-de-c-v-20211123",title:"IntechOpen Signs Exclusive Representation Agreement with LSR Libros Servicios y Representaciones S.A. de C.V"},{slug:"intechopen-expands-partnership-with-research4life-20211110",title:"IntechOpen Expands Partnership with Research4Life"}]},book:{item:{type:"book",id:"10677",leadTitle:null,fullTitle:"Advanced Topics of Topology",title:"Advanced Topics of Topology",subtitle:null,reviewType:"peer-reviewed",abstract:"Topology is an area of mathematics that establishes relations and transformations between spaces with a certain structure depending on their position and considering the structure of the ambient space where these relations exist. This book discusses various concepts and theories of topology, including diffeomorphisms, immersions, Hausdorff spaces, cobordisms, homotopy theory, symplectic manifolds, topology of quantum field theory, algebraic varieties, dimension theory, Koszul complexes, continuum theory, and metrizability, among others.",isbn:"978-1-80355-094-7",printIsbn:"978-1-80355-093-0",pdfIsbn:"978-1-80355-095-4",doi:null,price:119,priceEur:129,priceUsd:155,slug:"advanced-topics-of-topology",numberOfPages:136,isOpenForSubmission:!1,isInWos:null,isInBkci:!1,hash:"bf964c52f9e653fac20a7fcab58070e5",bookSignature:"Francisco Bulnes",publishedDate:"July 27th 2022",coverURL:"https://cdn.intechopen.com/books/images_new/10677.jpg",numberOfDownloads:495,numberOfWosCitations:0,numberOfCrossrefCitations:1,numberOfCrossrefCitationsByBook:null,numberOfDimensionsCitations:1,numberOfDimensionsCitationsByBook:null,hasAltmetrics:0,numberOfTotalCitations:2,isAvailableForWebshopOrdering:!0,dateEndFirstStepPublish:"June 25th 2021",dateEndSecondStepPublish:"September 14th 2021",dateEndThirdStepPublish:"November 13th 2021",dateEndFourthStepPublish:"February 1st 2022",dateEndFifthStepPublish:"April 2nd 2022",currentStepOfPublishingProcess:5,indexedIn:"1,2,3,4,5,6,7",editedByType:"Edited by",kuFlag:!1,featuredMarkup:null,editors:[{id:"92918",title:"Dr.",name:"Francisco",middleName:null,surname:"Bulnes",slug:"francisco-bulnes",fullName:"Francisco Bulnes",profilePictureURL:"https://mts.intechopen.com/storage/users/92918/images/system/92918.png",biography:"Dr. Francisco Bulnes, Ph.D., is IINAMEI Director, Mathematics Research Centre, Mexico. He is a member of various international committees of science and serves as a reviewer and editor for British and American journals of mathematics and physics. He is head of the Research Department, GI-TESCHA. He has published more than 100 papers and several books in mathematics and physics. Dr. Bulnes has many theories, theorems, and math objects to his credit. He has received various honors and awards from universities as well as governmental and non-governmental organizations. He received the Doctor Honoris Causa in Education Philosophy and is a Peace Ambassador for ODAEE in Frankfurt, Germany. He is also a distinguished member of the Czech Republic Mathematics Society (JCFM). He obtained two post-doctorates in Mathematics in Cuba and Russia. His research interests include electronics, microelectronics, and spintronics.",institutionString:"Investigación Internacional Avanzada en Matemáticas e Ingeniería (IINAMEI)",position:null,outsideEditionCount:0,totalCites:0,totalAuthoredChapters:"14",totalChapterViews:"0",totalEditedBooks:"7",institution:null}],equalEditorOne:null,equalEditorTwo:null,equalEditorThree:null,coeditorOne:null,coeditorTwo:null,coeditorThree:null,coeditorFour:null,coeditorFive:null,topics:[{id:"165",title:"Geometry & Topology",slug:"geometry-and-topology"}],chapters:[{id:"80411",title:"Introductory Chapter: The Topology from Classic Studies until Its Last Frontiers",doi:"10.5772/intechopen.102527",slug:"introductory-chapter-the-topology-from-classic-studies-until-its-last-frontiers",totalDownloads:2,totalCrossrefCites:0,totalDimensionsCites:0,hasAltmetrics:0,abstract:null,signatures:"Francisco Bulnes",downloadPdfUrl:"/chapter/pdf-download/80411",previewPdfUrl:"/chapter/pdf-preview/80411",authors:[{id:"92918",title:"Dr.",name:"Francisco",surname:"Bulnes",slug:"francisco-bulnes",fullName:"Francisco Bulnes"}],corrections:null},{id:"78241",title:"More Functions Associated with Neutrosophic gsα*- Closed Sets in Neutrosophic Topological Spaces",doi:"10.5772/intechopen.99464",slug:"more-functions-associated-with-neutrosophic-gs-closed-sets-in-neutrosophic-topological-spaces",totalDownloads:108,totalCrossrefCites:1,totalDimensionsCites:1,hasAltmetrics:0,abstract:"The concept of neutrosophic continuous function was very first introduced by A.A. Salama et al. The main aim of this paper is to introduce a new concept of Neutrosophic continuous function namely Strongly Neutrosophic gsα* - continuous functions, Perfectly Neutrosophic gsα* - continuous functions and Totally Neutrosophic gsα* - continuous functions in Neutrosophic topological spaces. These concepts are derived from strongly generalized neutrosophic continuous function and perfectly generalized neutrosophic continuous function. Several interesting properties and characterizations are derived and compared with already existing neutrosophic functions.",signatures:"P. Anbarasi Rodrigo and S. Maheswari",downloadPdfUrl:"/chapter/pdf-download/78241",previewPdfUrl:"/chapter/pdf-preview/78241",authors:[{id:"426315",title:"Dr.",name:"P. Anbarasi",surname:"Rodrigo",slug:"p.-anbarasi-rodrigo",fullName:"P. Anbarasi Rodrigo"},{id:"426558",title:"Ms.",name:"S.",surname:"Maheswari",slug:"s.-maheswari",fullName:"S. Maheswari"}],corrections:null},{id:"80455",title:"4-Dimensional Canards with Brownian Motion",doi:"10.5772/intechopen.102151",slug:"4-dimensional-canards-with-brownian-motion",totalDownloads:47,totalCrossrefCites:0,totalDimensionsCites:0,hasAltmetrics:0,abstract:"Generally speaking, it is impossible to analyze slow-fast system with Brownian motion. If it becomes possible to do using a new approach, we can evaluate the rigidity of the original system. What kind of the behavior of such a system we have? Using non-standard analysis, on a“hyper finite time line” by Anderson, the Brownian motions are described by step functions. Then, the original differential equations are described by the difference equations due to using non-standard analysis. When constructing the difference equations, the corresponding measure is extended topologically. Because the interval of the difference is according to the hyper finite time line, the topological space is well defined. In this paper, we propose a two-region economic model with Brownian motions. This concrete example gives us new results.",signatures:"Shuya Kanagawa and Kiyoyuki Tchizawa",downloadPdfUrl:"/chapter/pdf-download/80455",previewPdfUrl:"/chapter/pdf-preview/80455",authors:[{id:"427472",title:"Prof.",name:"Shuya",surname:"Kanagawa",slug:"shuya-kanagawa",fullName:"Shuya Kanagawa"},{id:"427473",title:"Dr.",name:"Kiyoyuki",surname:"Tchizawa",slug:"kiyoyuki-tchizawa",fullName:"Kiyoyuki Tchizawa"}],corrections:null},{id:"79299",title:"The Topology of the Configuration Space of a Mathematical Model for Cycloalkenes",doi:"10.5772/intechopen.100723",slug:"the-topology-of-the-configuration-space-of-a-mathematical-model-for-cycloalkenes",totalDownloads:120,totalCrossrefCites:0,totalDimensionsCites:0,hasAltmetrics:0,abstract:"As a mathematical model for cycloalkenes, we consider equilateral polygons whose interior angles are the same except for those of the both ends of the specified edge. We study the configuration space of such polygons. It is known that for some case, the space is homeomorphic to a sphere. The purpose of this chapter is threefold: First, using the h-cobordism theorem, we prove that the above homeomorphism is in fact a diffeomorphism. Second, we study the best possible condition for the space to be a sphere. At present, only a sphere appears as a topological type of the space. Then our third purpose is to show the case when a closed surface of positive genus appears as a topological type.",signatures:"Yasuhiko Kamiyama",downloadPdfUrl:"/chapter/pdf-download/79299",previewPdfUrl:"/chapter/pdf-preview/79299",authors:[{id:"327075",title:"Prof.",name:"Yasuhiko",surname:"Kamiyama",slug:"yasuhiko-kamiyama",fullName:"Yasuhiko Kamiyama"}],corrections:null},{id:"82378",title:"Covers and Properties of Families of Real Functions",doi:"10.5772/intechopen.100555",slug:"covers-and-properties-of-families-of-real-functions",totalDownloads:9,totalCrossrefCites:0,totalDimensionsCites:0,hasAltmetrics:0,abstract:"We present results on the relationships of the covering property GΦΨ for Φ,Ψ∈OΛΩΓ and G∈S1SfinUfin of a topological space and the selection property GΦ0Ψ0 of the corresponding family of real functions. The result already published are presented without a proof, however with a citation of the corresponding paper. We present a general Theorem that covers almost all the result of this kind. Some results about hereditary properties are enclosed. We also present Scheepers Diagram of considered covering properties for uncountable covers.",signatures:"Lev Bukovský",downloadPdfUrl:"/chapter/pdf-download/82378",previewPdfUrl:"/chapter/pdf-preview/82378",authors:[{id:"427352",title:"Emeritus Prof.",name:"Lev",surname:"Bukovský",slug:"lev-bukovsky",fullName:"Lev Bukovský"}],corrections:null},{id:"80019",title:"Vertex Decomposability of Path Complexes and Stanley’s Conjectures",doi:"10.5772/intechopen.101083",slug:"vertex-decomposability-of-path-complexes-and-stanley-s-conjectures",totalDownloads:53,totalCrossrefCites:0,totalDimensionsCites:0,hasAltmetrics:0,abstract:"Monomials are the link between commutative algebra and combinatorics. With a simplicial complex Δ, one can associate two square-free monomial ideals: the Stanley-Reisner ideal IΔ whose generators correspond to the non-face of Δ, or the facet ideal I(Δ) that is a generalization of edge ideals of graphs and whose generators correspond to the facets of Δ. The facet ideal of a simplicial complex was first introduced by Faridi in 2002. Let G be a simple graph. The edge ideal I(G) of a graph G was first considered by R. Villarreal in 1990. He studied algebraic properties of I(G) using a combinatorial language of G. In combinatorial commutative algebra, one can attach a monomial ideal to a combinatorial object. Then, algebraic properties of this ideal are studied using combinatorial properties of combinatorial object. One of interesting problems in combinatorial commutative algebra is the Stanley’s conjectures. The Stanley’s conjectures are studied by many researchers. Let R be a Nn-graded ring and M a Zn-graded R-module. Then, Stanley conjectured that depthM≤sdepthM. He also conjectured that each Cohen-Macaulay simplicial complex is partition-able. In this chapter, we study the relation between vertex decomposability of some simplicial complexes and Stanley’s conjectures.",signatures:"Seyed Mohammad Ajdani and Francisco Bulnes",downloadPdfUrl:"/chapter/pdf-download/80019",previewPdfUrl:"/chapter/pdf-preview/80019",authors:[{id:"92918",title:"Dr.",name:"Francisco",surname:"Bulnes",slug:"francisco-bulnes",fullName:"Francisco Bulnes"},{id:"440971",title:"Dr.",name:"Seyed",surname:"Mohammad Ajdani",slug:"seyed-mohammad-ajdani",fullName:"Seyed Mohammad Ajdani"}],corrections:null},{id:"79892",title:"βI-Compactness, βI*-Hyperconnectedness and βI-Separatedness in Ideal Topological Spaces",doi:"10.5772/intechopen.101524",slug:"-em-em-sub-em-i-em-sub-compactness-em-em-sub-em-i-em-sub-hyperconnectedness-and-em-em-sub-em-i-em-su",totalDownloads:77,totalCrossrefCites:0,totalDimensionsCites:0,hasAltmetrics:0,abstract:"Let XτI be an ideal topological space. A subset A of X is said to be β-open if A⊆clintclA, and it is said to be βI-open if there is a set O∈τ with the property 1 O−A∈I and 2 A−clintclO∈I. The set A is called βI-compact if every cover of A by βI-open sets has a finite sub-cover. The set A is said to be cβI-compact, if every cover Oλ:λ∈Λ of A by β-open sets, Λ has a finite subset Λ0 such that A−∪Oλ:λ∈Λ0∈I. The set A is said to be countably βI-compact if every countable cover of A by βI-open sets has a finite sub-cover. An ideal topological space XτI is said to be βI∗-hyperconnected if X−cl∗A∈I for every non-empty βI-open subset A of X. Two subsets A and B of X is said to be βI-separated if clβIA∩B=∅=A∩clβB. Moreover, A is called a βI-connected set if it can’t be written as a union of two βI-separated subsets. An ideal topological space XτI is called βI-connected space if X is βI-connected. In this article, we give some important properties of βI-open sets, βI-compact spaces, cβI-compact spaces, βI∗-hyperconnected spaces, and βI-connected spaces.",signatures:"Glaisa T. Catalan, Michael P. Baldado Jr and Roberto N. Padua",downloadPdfUrl:"/chapter/pdf-download/79892",previewPdfUrl:"/chapter/pdf-preview/79892",authors:[{id:"425714",title:"Prof.",name:"Michael",surname:"Baldado",slug:"michael-baldado",fullName:"Michael Baldado"},{id:"438024",title:"Dr.",name:"Glaisa",surname:"Catalan",slug:"glaisa-catalan",fullName:"Glaisa Catalan"},{id:"486473",title:"Dr.",name:"Roberto N.",surname:"Padua",slug:"roberto-n.-padua",fullName:"Roberto N. Padua"}],corrections:null},{id:"79797",title:"Clairaut Submersion",doi:"10.5772/intechopen.101427",slug:"clairaut-submersion",totalDownloads:79,totalCrossrefCites:0,totalDimensionsCites:0,hasAltmetrics:0,abstract:"In this chapter, we give the detailed study about the Clairaut submersion. The fundamental notations are given. Clairaut submersion is one of the most interesting topics in differential geometry. Depending on the condition on distribution of submersion, we have different classes of submersion such as anti-invariant, semi-invariant submersions etc. We describe the geometric properties of Clairaut anti-invariant submersions and Clairaut semi-invariant submersions whose total space is a Kähler, nearly Kähler manifold. We give condition for Clairaut anti-invariant submersion to be a totally geodesic map and also study Clairaut anti-invariant submersions with totally umbilical fibers. We also give the conditions for the semi-invariant submersions to be Clairaut map and also for Clairaut semi-invariant submersion to be a totally geodesic map. We also give some illustrative example of Clairaut anti-invariant and semi-invariant submersion.",signatures:"Sanjay Kumar Singh and Punam Gupta",downloadPdfUrl:"/chapter/pdf-download/79797",previewPdfUrl:"/chapter/pdf-preview/79797",authors:[{id:"358081",title:"Dr.",name:"Sanjay",surname:"Kumar Singh",slug:"sanjay-kumar-singh",fullName:"Sanjay Kumar Singh"},{id:"426050",title:"Assistant Prof.",name:"Punam",surname:"Gupta",slug:"punam-gupta",fullName:"Punam Gupta"}],corrections:null}],productType:{id:"1",title:"Edited Volume",chapterContentType:"chapter",authoredCaption:"Edited by"},subseries:null,tags:null},relatedBooks:[{type:"book",id:"7428",title:"Advances on Tensor Analysis and their Applications",subtitle:null,isOpenForSubmission:!1,hash:"2339ac5eb978557d01451489e961b102",slug:"advances-on-tensor-analysis-and-their-applications",bookSignature:"Francisco Bulnes",coverURL:"https://cdn.intechopen.com/books/images_new/7428.jpg",editedByType:"Edited by",editors:[{id:"92918",title:"Dr.",name:"Francisco",surname:"Bulnes",slug:"francisco-bulnes",fullName:"Francisco Bulnes"}],equalEditorOne:null,equalEditorTwo:null,equalEditorThree:null,productType:{id:"1",chapterContentType:"chapter",authoredCaption:"Edited by"}},{type:"book",id:"8907",title:"Advances in Quantum Communication and Information",subtitle:null,isOpenForSubmission:!1,hash:"6b074960b5f71319aa57217e7b54216e",slug:"advances-in-quantum-communication-and-information",bookSignature:"Francisco Bulnes, Vasilios N. 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1. Introduction
Globally the area under vegetable cultivation is growing annually at the rate of 4.12% and production by 6.48%. The mean productivity is 15.49mt/ha which is quite low. In vegetables infestation of biotic stresses reflect highly on production productivity and quality. Application of chemicals leaves chemical residues in vegetables above threshold levels. Resistance is a relative attribute and refers to the ability of the plant to withstand the pest or pathogen. The susceptible plant shows severe symptoms due to which yield loss occur. A completely resistant plant shows nil reaction and a moderately resistant or field tolerant plant develops less disease development. Plants have selective resistance to some pests or pathogens and susceptible to others. They are species-specific or strain-specific. The rate of spread depends on the pest load or population, spore count and multiplication rate of pest/pathogen.
Improvement of cultivated plants through tapping germplasm resources depends on introducing variability through traditional and molecular breeding techniques. Wild species provides a vast gene pool for resistance development. They have been used for decades to transfer genes of resistance or tolerance to the cultivated species. The use of wild species in breeding varieties particularly for increased vigor and resistance has been well recognized [1]. Introgression is the movement of genes or gene flow from one species into the gene pool. Inter specific hybridisation breaks the species barrier for gene transfer and makes it possible to transfer the resistant genes.
Complete exploitation of genetic variation enables the breeder to produce not only heterotic F1 hybrids but also recombinants with desirable attributes. Further, selection based on genetic nature will be highly useful to a great extent to screen out the parents and hybrids. Identification of resistance is also possible through quantifying the biochemical components present in the genotype. Further, in view of less marked host specificity, a plant breeding programme for insect resistance has to be handled separately from that of disease resistance.
In 3rd century B.C, Theophrastus observed that degree of resistance differ among varieties. It was later established in 1894, by Erikson that though pathogens are morphologically similar, they differ among each other in their ability to attack host plant. In 1911 Barrus narrated that various isolates of a pathogen differ in its ability to attack different varieties of the same plant species. This made the basis for the identification of physiological races and pathotypes. It was then called as pathogenecity i.e the infection of a host strain by a pathogen is genetically determined. In 1955, Flor formulated the of gene-for-gene hypothesis which denotes the relationship between host and pathogen. According to that disease resistance is determined by host and the genotype of the pathogen. The hosts differ in type of resistance while the pathogen differ in pathogenicity, but both are genetically controlled. The pathogen has the capacity to generate new variations in pathogenicity by reproduction methods and mutation. Therefore, the task of the breeder is to develop varieties resistant to the prevalent pathotypes of the pathogen and also for the new pathogen genotypes which will arise in future.
2. Genetic basis of resistance
According to the experimental results so far reported in vegetable crops, the genetic basis of insect resistance is monogenic. The resistance of muskmelon to melon aphid. The tolerance of muskmelon to western biotypes of Aphis gossypii in breeding line LJ 90234 was governed by a single dominant gene [2]. Inheritance studies of fruit fly resistance in pumpkin cultivar Arka Suryamukhi showed that the resistance was controlled by a dominant gene. Similar studies in water-melon also indicated that the resistance to fruit fly was governed by a single dominant gene. The work on Cucurbita pepo revealed that the resistance to squash bug, was controlled by at least 3 genes and gene action appeared to be additive in nature. In an interspecific cross between resistant Cucumis callosus and susceptible Cucumis melo it was revealed that the susceptibility to fruit fly was governed by two pairs of complimentary genes. While working with tomato for resistance to fruitworm Fery and Cuthbert [3] reported that the antibiotics factor present in Lycopersicon hirsutum appeared to be inherited recessively.
Since interest in resistant vegetable varieties started more than half a century ago work has been done on major insect pests. However, studies have shown that the Mendelian segregation has led to the identification of major genes and that the alleles for resistance were dominant over those for susceptibility in number of instances except in some where it was found to be additive or complementary gene action or recessive.
The genetics of disease resistance was first studied by Britten in 1905. Then Person and Sidhu [4] reviewed 1000 Published papers and concluded that regardless of species, resistance generally segregated in the mendelian ratios. Resistance was dominant over susceptibility. Resistance in vegetable crops have been reported to be governed by mono or oligo or polygenes and effect of genes may be additive or dominant or epistatic. The information on inheritance of various diseases of vegetables is very meager. However, some workers reported different kinds of nature of inheritance. Resistance to buck eye rot of tomato appeared to be dominant over susceptibility. Resistance to fusarium wilt of tomato was conditioned by a single dominant gene. Tomato leaf curl virus is transmitted by white fly and is most serious problem. According to Som and Chaudhary [5], resistance to TLCV was incompletely dominant and governed by polygenes.
Resistance to most of the diseases in watermelon is controlled by a single dominant gene. Walker [6] reported resistance to fusarium wilt in watermelon as recessive. Powdery mildew is a major limiting factor in the production of muskmelon in most of the parts of the world. Resistance to Erysiphae cichoracearum race- 1 and race-2 is monogenic dominant. A study on resistance to powdery mildew caused by Sphaerotheca fuligina, in two resistant varieties campo and PMR-6 indicated that they have the same locus/loci conferring resistance. Genetic studies of resistance to E. cichoracearum race-2 had indicated that resistance is partly dominant and controlled by Pm-2 [7]. Resistance to downy mildew (Pseudoperonospora cubensis) of muskmelon in PI 124111 is controlled by two independently dominant gene [8]. Whereas resistance in PI 124112 was controlled by two partially dominant genes [9].
Antonio et al. [10] studied the inheritance of resistance by antixenosis for tomato leaf miner and reported that the inheritance of antixenosis resistance of genotype BGH-1497 is ruled by a polygenes in epistatic interactions, with a phenotypic proportion of 13:3 between susceptible and resistant genotypes respectively. In another experiment Gabriele Vitelli et al. [11] reported three transgenic eggplant lines bearing a mutagenized Bacillus thuringiensis Berl. gene coding for the Cry3B toxin. The fruit production was almost twice in the highly resistant lines (3–2 and 9–8). The 6–1 transgenic line showed an intermediate level of resistance. Analysis by double antibody sandwich–enzyme linked immune sorbent assay (DAS–ELISA), performed on different tissues, revealed a lower amount of Cry3B protein in the 6–1 transgenic line.
The root knot nematode, Meloidogyne is one of the most economically damaging plant parasitic nematode and is widely distributed throughout world [12]. The genus Meloidogyne is composed of 100 species, with M. arenaria, M. incognita, M. hapla and M. javanica being considered as “major” species [13]. Natural resistance genes “R-genes” are responsible for inducing resistance against nematodes in tomato. The genes Mi-1, Mi-2, Mi-3, Mi-4, Mi-5, Mi-6, Mi-7, Mi-8, Mi-9, and Mi-HT confer resistance to the root Knot nematode [14].
3. Biochemical basis of resistance
The biochemical factors are more important than morphological and physiological factors in conferring resistance. Some biochemical constituents may act as feeding stimuli for insects. Occurrence at lower concentration or total absence of such biochemical leads to non preference, a form of insect resistance [15, 16]. The biochemical constituents like glycoalkaloid (solasodine), phenols, phenolic oxidase enzymes viz., polyphenol oxidase and peroxidase are available in plants and these biochemical constituents possess insect resistant properties [17]. It was also recorded that the maximum polyphenol oxidase activity is available in fruit (0.388 in fruit as changes in OD min-1 g-1 of sample). Several workers have reported that the biochemical constituents act as stimulants of resistance mechanism. In brinjal, Praneetha [18] and Prabhu [19] have recorded that the biochemical constituents also contribute to confer resistance to shoot and fruit borer.
Studies on the biochemical basis of resistance to Leucinodes orbonalis and their correlation with shoot and fruit borer damage in five selected brinjal genotypes were done during June to December 2005 and it was shown that less susceptible genotypes for both shoot and fruit borer had higher amount of polyphenol oxidase (PPO), phenylalanine ammonium lyase (PAL), lignin and lower reducing sugar. Significant negative correlation was established with per cent infestation of shoot and fruit borer and PPO, PAL and lignin, whereas it was positively correlated with reducing sugar. Negative correlation was observed with the biochemical constituents, PPO, PAL, lignin and reducing sugar but PPO was positively correlated with PAL and lignin content and vice-versa.
4. Biophysical basis of resistance
In a study Silva et al. [20, 21] evaluated ninety-nine F3 families derived from an interspecific cross using Solanum lycopersicum and Solanum pimpinellifolium ‘TO-937-15’ (multiple pest resistance accession with type IV glandular trichomes and acylsugar accumulation) for their resistance against the whitefly. The higher resistance levels of BTR331 were associated with a positive combination of higher type IV trichome density and higher acylsugar levels. From the breeding stand point, the genetic similarity between S. lycopersicum and S. pimpinellifolium would allow a more efficient resistance introgression by facilitating recombination and minimizing the potentially undesirable linkage drag associated with this trait.
Niranjana et al. reported that the biophysical characteristics in brinjal genotypes viz., shoot thickness at 2.5 cm below the tip, number of trichomes on lower surface of leaves, pedicel length, calyx length and diameter of fruit were correlated with the level of infestation by L. Orbonalis. Results revealed that the infestation in shoot was not significantly correlated with number of trichomes on leaves and positively correlated with shoot thickness. Fruit infestation was positively but not significantly correlated with length of pedicel and calyx whereas non-significant and negative correlation was recorded between fruit infestation and fruit characters viz., length and diameter of fruit. The shape and color of fruit had no significant influence on the level of infestation.
5. Categorization of resistance
According to the response of the plant to the pathogen.
Susceptible: In this the disease development is abundant and can not be checked by the plant.
Resistant: It is lesser disease development than the susceptible and it is a relative attribute. In this the plants will be infected and establishment take place but the progression in the host plant will be limited. As a result these plants exhibit minor symptoms than the susceptible ones.
Tolerant: Tolerance implies that the host is attacked by the pathogen but there is no less in biomass production or yield.
Based on number of genes governing the resistance trait as
Monogenic resistance: Controlled by single gene. Easy to incorporate into plants by breeding. Easy to break also.
Oligogenic resistance: Controlled by few genes
Polygenic resistance: Controlled by many genes
Resistance is classified based on biotype reaction as.
Vertical resistance: It is determined by one or few genes and is characterized by pathotype specificity. That is it is attacked by only one virulent pathotype. For all others, the host will be resistant. It is also called as specific resistance or race-specific resistance
Horizontal resistance: It is determined by polygenes. Horizontal resistance does not prevent the development of symptoms of the disease, but it slows down the rate of spread of disease in the population. It is also known by race- nonspecific, partial and field resistance.
Resistance is classified based on population/Line concept
Pureline resistance: Exhibited by lines which are phenotypically and genetically similar.
Multiline resistance: Exhibited by lines which are phenotypically similar but genotypically dissimilar
Plants once infected by a specific pathogen become resistant to further infections by the same one. This was discovered in the beginning of 20th century. This concept is involved in viral cross-protection and induced systemic resistance. Induced systemic resistance in plants is of several types of which Systemic Acquired Resistance (SAR) is the most important one. It is long lasting and effective against viral, bacterial and fungal pathogens. It ranges from a oversensitive response to necrotic lesions. SAR is due to high level of salicylic acid which is essentially needed for the development of SAR. Salicylic acid acts as a phloem translocated signal that mediates SAR. It is also due to SAR genes which is different in monocotyledonous and dicotyledonous plants. In tobacco, SAR genes cover a set of non-allelic genes that can be classified on the basis of proteins they encode such as the pathogenesis related (PR) genes. These genes play an active role in the disease resistance as their expression in transgenic plants impart significant disease resistance. SAR genes in various species differ in considerable extent.
6. Sources of resistance
Plants that may be less desirable in other ways, but carry a useful disease resistance trait. Ancient known plant varieties and wild species, cultivated varieties and land races are very important to preserve because they are the most common sources of enhanced plant disease resistance. Source of resistance are available for melon aphid, striped and spotted cucumber beetles, squash bug, squash borer, pickleworm, red pumpkin beetle, fruit fly in different cucurbits, cabbage maggot and aphid in cabbage and spinach, fruit and shoot borer in brinjal, jassids in okra, potato leaf hopper, melon fly in different beans, pea aphid and weevil. Genetic basis of insect resistance has been reported to be monogenically dominant in muskmelon aphid and in pumpkin and watermelon for fruit fly, whereas, additive gene action have been reported for resistance to both striped and spotted cucumber beetles and squash bug in squash. In interspecific crosses of muskmelon with wild melon two pairs of complimentary genes are reported to be involved for resistance to fruit fly. Maternal influence has also been indicated in inter-varietal crosses of squash for resistance to spotted cucumber beetle.
In India work on resistance of cucurbits to red pumpkin beetle and fruit fly was initiated at IARI, New Delhi as early during 1962 and at Indian Institute of Horticultural Research, Bangalore during 1969. A highly resistant source to fruit fly was obtained in pumpkin, which was utilized, in breeding a resistant variety Arka Suryamukhi. Some of the pumpkin lines were fairly resistant to red pumpkin beetle. It was observed that among the different species Citrullus colocynthis was highly damaged by fruit fly. In Lycopersicon genus L.hirsutum and L. hirsutum f. glabratum is resistant to fruitworm, and also indicated that since these were cross-compatible with L. esculentum and it was possible to transfer the resistance factor in the cultivated varieties. It was further indicated that L. hirsutum f. glabratum was also resistant to tobacco flea beetle and carmine spider mite. It was possible to incorporate resistance to more than one insect species in one genotype.
In Onion sources of resistance were identified, cause and mechanism studied and suitable varieties were developed. In Okra difference in varietal response to jassid attack was observed and fruit of the resistant line was found to have strong prickly hairs and was highly susceptible yellow vein mosaic virus. In spinach variety Manchuria was reported as resistant to aphid as early as in 1920. In Carrot resistance to fruit fly was reported and one of the amaranthus lines was observed to have high field resistance to grasshopper in Nigeria. With monogenic inheritance available for melon aphid in cantaloupe, for fruit fly in pumpkin and in watermelon it would be possible to utilize backcross method for incorporating resistant gene in commercial varieties.
7. Breeding methods
Introduction: Collections of related materials from other countries, particularly from areas where the pathogen and host species may have co-evolved, sometimes provide rich pools of resistance genes [22]. In vegetable peas, early introductions from Europe and USA were found quite successful and popular in India. These included Arkel (early maturing, dwarf type, introduction from England in 1970s) and Bonneville (main season, late maturing, tall type, introduction from USA in 1970s). These introductions were obtained at IARI, New Delhi and were released for commercial cultivation after preliminary evaluation. Early Badger a dwarf, wrinkled seeded variety introduced from USA has resistance to Fusarium wilt.
8. Utilization of wild species
The major bottleneck in the resistance breeding programme is the lack of resistant source in the cultivated germplasm. This has necessitated breeders to search resistance for genes in wild species that are taxonomically related and compatible. The use of wild forms in breeding crop plants, particularly to obtain vigor and resistance hasbeen well recognized [1]. In vegetables, several experiments involving wild species have been carried out. Selection of a genotype with high yield and resistance reduces the yield loss on one hand and increases the availability of the produce to market which is fairly free from residue on the other hand. Generally the source of genes for resistance are (i) provided by the variability within the crop species, (i) varieties from original home of insect, (ii) varieties from centres of great insect occurrence, (iii) varieties from areas of greatest morphological diversity.
In muskmelon gene for resistance is not available in the cultivated species, but Cucumis africanus was found to be fairly resistant to fruit fly. Similarly, Cucumis heptadactylis was resistant to red pumpkin beetle. An attempt to incorporate the resistance to shoot and fruit borer in the cultivated egg plant genotypes was made in interspecific hybridprogenies of the cross S. melongena x Solanum viarum and evaluating the direct segregating progenies of such interspecific crosses so as to identify recombinant inbred plants with high yield and shoot and fruit borer resistance. The data recorded from direct F9 generation derivatives of EP 65 x Solanumviarum were utilized to study thoroughly and three progenies each out of thirty in F9 generations were selected for further studies. All these selected progenies have performed very well with respect to shoot and fruit borer resistance. From this evaluation studytwo hybrid derivatives were selected and designated as HD 1 and HD 2. The progenies of the culture HD1 recorded minimum shoot (7.69%) and fruit borer infestation (6.67%). The HD2 progenies recorded the minimum shoot (9.09%) and fruit borer infestation as 6.85%. The selected progenies viz., HD 1 and HD 2 showed profuse flowering and fruiting and also cluster bearing habit. The color of the fruit was bright purple while the fruit surface was smooth, glossy along with tightly packed seeds in its flesh which again act as physical barrier for mandibles of fruit borer to chew and bore into the flesh of fruits [18].
Tomato is a self pollinated crop, which is a high demand vegetable crop. The wild species are reservoir of important genes in tomato. Solanum pimpinellifolium is the only red-fruited wild species of tomato. Because of the close phylogenetic relationship between the two species, there is little or no difficulty in initial crosses in subsequent generations of pre-breeding and breeding activities. Nineteen accessions from seven Lycopersicon species were bio assayed for their resistance to Heliothis armigera by [23]. It was found that among the various Lycopersicon spp. bioassayed, accessions of L. hirsutum f. glabratum is most potential for breeding H. armigera resistant cultivars.
Wilt, little leaf and phomopsis blight are the serious diseases of brinjal. Solanum incanum is resistant to Fusarium wilt. In humid tropical areas brinjal is highly infected with bacterial wilt. Wild species of Solanum viz., S. torvum, S. xanthocarpum, S. nigrum, and S. sisymbriifolium are resistant [24]. The wild species Solanum viarum showed no infection and was immune, whereas the species S. incanum and S. sisymbrifolium were resistant to little leaf disease.
Pinheiro et al. [25] conducted two assays, to evaluate the resistance to root knot nematode, M. incognita race 1 in Citrullus lanatus cv. Citroides, Lagenaria vulgaris, Sicana odorifera, Cucurbita facifolia, Cucurbita moschata, Cucurbita moschata x Cucurbita maxima, Luffa sp., Cucumis melo and Cucumis metuliferus accessions. The results revealed that three accessions of Cucumis metuliferus (‘Kino’) were resistant to M. incognita race 1 in the first experiment. In the second experiment conducted to evaluate the reaction to nematode M. incognita race 1, M. javanica and M. enterolobii all the seedlings in pots were inoculated with 2nd stage (J2) juveniles and 5000 eggs of each Meloidogyne species. The observations on egg mass index (IMO), gall index (IG), number of eggs per gram of root (NEGR) and reproduction factor (RF) was observed on 53 and 84 days after inoculation, respectively. The melon Cucumis metuliferus was resistant to root-knot nematode.
9. Screening varieties
Varietal difference with regard to resistance for fruit and shoot borer was observed in brinjal. Choudhary et al. [26] screened eight varieties against L. Orbonalis. The order of susceptibility ofbrinjal varieties was recorded as Pant Samrat< Pant Rituraj<Manjarigota<Pusa Purple Long < Pant Brinjal-5 < Kavach< MHB-80 < BR-112 during 2014–2015 and 2015–2016, respectively. In another study, Amit et al. [27] screened twenty five brinjal varieties against brinjal shoot and fruit borer and red spider mite and IBH-3, IBL-116, Rajindra brinjal, KS-356, JB-24, JBH-8, IBH-02 andCHBR-1 were found tolerant. The research carried out at RARS, Jamalpur, Bangladesh showed that, the brinjal varieties Jumki-1 and Jumki-2 were highly resistant (HR), Islampuri-3, BL-34 and Muktakeshi were fairly resistant (FR), Singnath long and Singnath-4 were tolerant to brinjal shoot and fruit borer [28].
Similarly, in okra Rehman et al. [29] screened four varieties (SabzPari, SadaBahar, PusaSawani, Arka Anamika) and those varieties showed some degree of resistance against sucking insect pests. Okra variety Sada Bahar was less infested with jassid (1.30/leaf) and whitefly (5.36/leaf) compared to other tested varieties and resulted in maximum yield (1529.62 kg/ha). Number of fruits pods per plant was found non significantly different on all the tested okra varieties.
In another experiment, Jackson and Bohac [30] evaluated sweet potato accessions by using bio assay techniques using the adults of banded cucumber beetle and spotted cucumber beetle. A single beetle was placed on a piece of sweetpotato peel that was embedded periderm-side up in plaster in a petridish. Feeding and longevity of insects on sweet potato genotypes were evaluated. Durability of feeding with respect to banded cucumber beetles on sweet potato peels ranged from 12 d for the most-resistant genotype to 123 d for a susceptible control cultivar (SC1149–19). The feeding longevity of spotted cucumber beetles was slightly shorter than banded cucumber beetles. For the highly resistant genotypes, both the species exhibited a significant delay in feeding initiation, and most beetles died before they had fed the sweet potato. Thus it was evident that both antibiosis and non preference (antixenosis) are important mechanisms of resistance in sweetpotato genotypes.
Seventy seven eggplant genotypes were tested for resistance to root-knot nematode by classical testing. As a result it was determined that P29 and P52 genotypes were resistant.Ditylenchus destructor and Ditylenchus dipsaci are economically important plant-parasitic nematodes, affecting potato production mostly in temperate climates. Mwaura and Vidal [31] screened 25 potato varieties for resistance to and tolerance for D. destructor and D. dipsaci infections. Reproduction factor (RF) and relative susceptibility (RS) were used to evaluate resistance, Based on Reproduction factor, sixteen varieties were assessed as susceptible (S) and five were identified as resistant (R) to D. destructor. The varieties Innovator, Aveka and Spunta were identified as resistant to Ditylenchus dipsaci. The highly susceptible one for D. destructor and D. dipsaci in both experiments was Desiree and was used as the standard susceptible control variety for the calculation of RS. An 1–9 scale was used to assess and classify the potato varieties based on level of resistance to D. destructor and D. dipsaci, where 9 indicated the highest level of resistance. Among the varieties screened six had significantly lower relative susceptibility (RS) to D. dipsaci than the standard susceptible control. Few varieties were also observed to be tolerant to both the nematodes. The suitable indices for resistance and tolerance determination were relative susceptibility (RS) and external potato tuber damage.
Nayak and Pandey [32], screened one hundred fifty brinjal varieties/cultivars against root-knot nematode, only twenty varieties have shown resistant reaction with least gall index (1.1 to 2.0) viz., Gachhabaigan, Azadkranti, Kantabaigen, Athagara Local, Kamaghara local, Solanum indicum, PBR 129–5, ARU-1, BB1–3, BB 45-C, BB-49, KS-224, Utkal madhuri, BR-112, LB-13, LB-25, LB-28, LB-30, LB-44, LB-55.
Akhter and Khan [33], screened thirty brinjal varieties for their resistance/susceptibility to root-knot nematode (Meloidogyne incognita race-1) infestation. Out of 30 brinjal varieties, eighteen varieties viz., Black Beauty, Brinjal 1 hybrid, Brinjal No.38, Chamak, Govinda, Green round, Nagina, Nav Kiran, Neel Kamal, Nishant, P.K-123, Prabha Kiran, Prasad, Sukhda, Surya Kiran, i9Utkal, VNR-51 and VNR-60 were highly susceptible, seven varieties (Brinjal Advance, Brinjal BSS1013, Green long, Harshit, Prapti, Shamli and Ujjwal) were susceptible, two varieties (Mahy 112 and Mahy Ruby) were tolerant, two varieties (Hybrid green and JK Kajal) were moderately resistant and only one variety Mahy 80 was resistant against Meloidogyne incognita race-1. Mahy 80 variety was ported to be resistant against root-knot nematode, M. incognita race-1 for the first time.
10. Selection
Selection is an important method for breeding of varieties resistant to biotic stresses. It is an important means of isolating or identifying sources of disease resistance. Normally the sources of resistance are available in natural populations, wild species, introductions and spontaneous mutants. In the earlier periods selection was accomplished by sequestering the resistant survivors of natural epiphytotics. Now a days advanced artificial epiphytotics are being created and selection of resistant types are being done rather than escaped suscepts [34]. Selection of resistant plants from a commercial variety is the cheapest and quickest method of developing a resistant variety. IIHR Bangalore developed tomato varieties through pureline selection viz., Arka Alok and Arka Ahuti which showing resistant against bacterial wilt.
The production of garden pea is seriously limited by major diseases namely, wilt, powdery mildew and rust. Fusarium species cause root rot (F. solani f.sp. pisi and F. avenaceum) or wilt (F. oxysporum f.sp. pisi). Coyne et al. [35] developed three breeding lines (W6 26,740, W6 26,743, W6 26,745) having high level of resistance to Fusarium root rot caused by F. solani f.sp. pisi with acceptable agronomic traits. In melons the genotypes A19, A32, A30, JAB-11, JAB-20,JAB-3, JAB-7, C384, C67, JAB-9, JAB-18 were identified for powdery mildew resistance.
Pinheiro et al. [36] studied thirty seven pepper genotypes, Capsicum chinense, C. annuum and C. frutescens, were characterized for resistance to three root-knot nematode species (Meloidogyne javanica, M. incognita race 1 and M. enterolobii). Three experiments were carried out, in 2013, 2014 and 2016, in a greenhouse. Among the genotypes of Capsicum frutescens evaluated all were resistant or immune to M. javanica and M. incognita race 1. In C. chinense six accessions were susceptible. In the second experiment all genotypes of C. chinense and C. annuum, evaluated were resistant to M. incognita. In the third experiment, with C. annuum genotypes, most were susceptible to M. incognita race 1 while CNPH 30118 and CNPH 6144 were resistant to M. enterolobii, the most aggressive species. A greater degree of resistance was observed in few accessions of C. chinense and C. frutescens.
11. Hybridization
The common method used for resistance breeding is hybridization. In this the resistance is transferred by two means. In the first, by backcross method the resistance is transferred from a wild species or a variety with undesirable horticultural attributes to a susceptible but otherwise a desirable variety. In pedigree method the resistance is combined with some desirable characters of one variety and superior characteristics of another variety.
Heterosis a complex biological phenomenon manifested in the superiority of hybrids over parental forms due to the rate of development of one or more complex traits. Heterosis values should be negative for getting tolerant hybrids through heterosis breeding. As far as pest infestation is concerned a hybrid with least incidence might be due to complementation of genes. With the genetic knowledge available on insect resistance in vegetable crops it would be necessary to employ hybridization to transfer single, multiple or additive genes into commercial varieties in the advanced filial generations. The major achievement is expected through inter specific hybridization. In order that fruit quality, yield and resistance to insect pest may preferrably be incorporated in one genotype, it may possibly involve number of parents to achieve desired results.
Inter-varietal and inter-specific crosses, followed by selection, have accounted for the development of resistant hybrids. The production of inter-specific hybrids is useful for the transfer of desirable genes from wild to cultivated species. The Source, mechanism, biochemical and genetical basis of resistance in squash, muskmelon, cucumber and watermelon were studied by several authors.
By utilizing interspecific breeding technique and recombination, borer-free brinjal can be developed to protect the high yield and satisfy the preference of consumers. The parental lines EP 65 and Pusa Uttam had recorded higher yield together with lower fruit and shoot borer incidence. Both of them were found to be good combiners for the above traits. So these parents can be involved in multiple crossing programmes to transfer the resistant genes and to isolate desirable hybrids with high yield and low fruit and shoot borer incidence.
Introgression breeding has been extensively used in the genetic improvement of potato and tomato. In potato twelve traits have been introgressed from wild species viz., S.demissum, S. spegazzinii, S. stoloniferum, S. chacoense, S. acaule, S. vernei and in tomato from the wild species S. peruvianum, S. cheesmanii, S. pennellii and S. chilense. In watermelon, the F 1 (between resistant and susceptible) showed pronounced resistance to fruit fly.
Kishaba et al. [37] studied resistance by using several melon aphid-susceptible (MAS) recurrent parents from an initial cross of ‘PMR 45’ with P1 414,723. Fifteen advanced melon breeding lines with different levels of melon aphid resistance (MAR), their recurrent parents and P1 414,723 were compared in a naturally infested field test for susceptibility to feeding damage by CB. None of the MAR entries were more susceptible than their recurrent parents for fruit damage by CB. P1 414,723 was found to have a low level of resistance to seedling damage, and a high level of resistance to fruit damage from feeding by Cucumber beetle.
Hybridization was undertaken in brinjal with Solanum viarum to combine the resistance trait with high yield. The derivatives of the inter specific cross of Solanum melangena and Solanum viarum EP 65 (accession of Solanum melangena) x Solanum viarum were assessed till F9 generation. As a result two recombinant progenies viz., 7 and 9 were chosen and carried to the next generation as they had recorded high marketable yield and least infestation of shoot and fruit borer. Molecular confirmation with RAPD primers was done which depicted the introgression of the genes from donor parent Solanum viarum to brinjal and thus hybridity was confirmed.
Backcross method: This method is widely used for incorporation of resistant gene from a wild species or any variety with undesirable traits to a susceptible variety which is good in other agronomic attributes. The parents from which the resistant gene is to transferred is called as donor parent or non-recurrent parent. The susceptible parent in which the resistance gene is transmitted is known as the recurrent parent. The generation up to which the selection process has to be effected in a backcross program would differ according to the allelic relationship of the resistance genes i.e., whether it is resistance or dominant to the allele. Normally the backcross method help in the recovery of recurrent parent phenotype along with the transfer of resistant genes.
[38] investigated hybrid progenies of late blight resistant potato somatic hybrids developed through hybridization with common potato varieties, and also linked ISSR markers with resistant parent/progenies. Potato somatic hybrids (Solanum tuberosum di haploid ‘C-13’ + S. pinnatisectum) were back-crossed with potato varieties (S. tuberosum) and true potato seed (TPS) were produced. TPS-raised seedlings were advanced to back-cross progenies clones (BC1-C1, BC1-C2, BC1- C3 and BC1-C4) during the five years based on tuber traits in field trials and field resistance to late blight. The BC1-C2 progenies were profiled by ISSR markers and alleles linked to late blight resistant somatic hybrid parent P8 and their progenies (P8 × Kufri Jyoti) were identified. Eight promising advanced hybrids for late blight resistance were identified. This is the first ever report in India towards widening the genetic base of potato by exploitation of interspecific somatic hybrids. The methodology followed was as follows (Figure 1).
Figure 1.
A schematic presentation of development of interspecific potato somatic hybrids (S. tuberosum + S. pinnatisectum).
Powdery mildew is a serious disease of pea and for which new good resistant donors are available, a typical backcross breeding approach as applicable to a character governed by a single recessive gene was outlined by Gritton [39].
Pedigree method: It is quite suited when the resistance is governed by horizontal or polygenic. In breeding for disease resistance, artificial disease epidemics are generally produced to help in selection for disease resistance. Pedigree breeding is mainly used when the resistance is governed by major genes and have higher heritability. If it is used for low heritability traits then the selection process will be time consuming as it takes several generations usually F5 or F6 to attain homozygosity [40, 41], implemented a quantitative trait locus (QTL) mapping approach to study the inheritance of anthracnose resistance in an F2 population derived from the pedigree of C. annuum × C. chinense.
The major crops in which break through research was done and the resistant varieties developed through introgression breeding methods is tabulated below.
Literature on mutation breeding of vegetable crops for resistance to insect attack is meager. In IIHR, Bangalore varietal differences, have been indicated for resistance to jassids in the M1and M 2 populations of okra (dry seed irradiated with 55 to 60 KR gamma rays), to aphid in the M population of muskmelon (30–40 KR gamma rays); and to aphid in the MQ population of watermelon (dry seed irradiated with 50 KR gamma rays). This was indicative that there was a great potential in this approach for resistance genes. It is particularly applicable in case of muskmelon to fruit fly, brinjal fruit and shoot borer, tomato fruit borer, melon fruit fly and others where useful source of resistance has not been obtained. Further in cases where one or two sources have been located in nature, it would be desirable to obtain more sources of resistance through mutation breeding. It is evident that very little experience has been gained in the use of induced mutations in resistance breeding against insect pests but the prospects of this approach are great.
Selection of spontaneous and induced mutant plants with resistance through the use of mutagenesis. In germplasm repositories sometimes natural spontaneous mutations may occur. During that occurrence some resistant types will be developed naturally. If such instances does not occur then mutations can be induced by artificial methods. The short comings in this method is that mostly duplication of naturally occurring genes will be the outcome and induced mutants will have mono factorial behavior and the inheritance is short-lived [4]. The genetic stability has to be tested by repeated screening of mutants for the confirmation of resistance.
Potato is a major vegetable crop and is infected by many diseases which induces losses in yield and quality. Artificial mutations through irradiations were tried for developing resistant varieties against stem canker and black scurf of potato caused by Rhizoctonia solani. The mutants developed were resistant to these diseases. Aslı Kara and Şerife Evrim Arici [42] investigated the effect of gamma radiation on the susceptibility of the potato plant to Rhizoctonia solani at 22, 33, 54, 57 and 109 Gy. The best results were found with a dose of 22 Gy. The application of gamma irradition in this study may offer a new approach for potato breeders for developing plants resistant to R. solani.
13. Biotechnological approaches
The Bt-brinjal has been developed by inserting cry1Ac gene from a soil bacterium called Bacillus thuringiensis through Agrobacterium tumefaciens mediated method. Bt brinjal contains three genes, cry1Ac gene, which encodes an insecticidal protein Cry1Ac, is derived from a common soil bacterium and is driven by CaMV 35S promoter (cauliflower mosaic virus 35S). It also has nptII gene (neomycin phosphor transferase-II) which contains an antibiotic resistance marker and another marker gene “aad “for amino glycoside adenyl transferase. The cry1AC protein formed in Bt brinjal is analogous in structure and function to that found in nature. The resistance against fruit and shoot borer of brinjal is provided by cry1Ac genes and it minimizes damages and facilitate for the reduction in pesticide sprays and thus it is eco-friendly. Bacillus thuringiensis and B. t var. Kenyae (B.t.k) microbial formulations have been found to be highly specific to target insect pests, and do not have deleterious effects on non-target organisms such as beneficial insects, birds, fish, and mammals including human beings. The confirmation by ELISA revealed the presence of the Cry1Ac protein. The quantitative estimates established significant levels of Cry1Ac protein (2.46–4.33 ng ml−1) in the leaf extract of the transformed plants. The expression of this insecticidal protein in high levels resulted in significant amount of mortality of larvae and also stunted the growth of any surviving larva on transformed plant tissue.
Incongruity occurs in inter-specific crosses as a result of a lack of genetic information in one partner to complete pre- and post-pollination processes in the other. After fertilization the growth of the embryo is restricted due to some post-fertilization barriers. For the developmental process an equilibrium has to be established between embryo and endosperm for sharing the nutrients. When this equilibrium in the development of the zygote is disturbed, first division is delayed and abortion of the young embryo or disintegration of endosperm happens. This abortion may occur in any stages of development of the young seed. Based on the stage of embryo abortion in vitro methods have been developed to overcome post-fertilization barriers in a number of plant species. Embryo rescue techniques are the oldest and most successful in vitro procedures.
In many instances, progeny from wild crosses of inter-varietal and inter-specific is difficult to produce owing to several barriers like pre zygotic and post-zygotic barriers. The development of young zygote may be arrested by hybrid breakdown, hybrid sterility and hybrid non viability.Post-zygotic barriers such as endosperm abortion and, at later stages, embryo degeneration are of common occurrence, leading to low fertility but these have been overcome through the use of embryo rescue. Embryo culture is one of the earliest forms of in vitro culture applied to practical problems that has proven of greatest value to breeders. Among the very important strategies hybrid embryo rescue, and related applications like ovule/ovary/placental cultures through sequential embryo cultureis especially useful in vegetable crops and culture of embryos has also been demonstrated in tomato, brinjal, capsicum, hot pepper, onion, potato, tomato including cucurbits for rescuing useful hybrids.
Ovule and ovary culture are more suitable than embryo culture for small seeded species or very young embryos. When abortion occurs in a very young stage and maternal tissue has no negative influence on the development of seeds, ovary culture can be applied. Ovary culture has been applied in many Brassica species. The ovule culture is applied in Lycopersicon. Depending on the genotypic combination of the inter-specific crossing, the percentage of seedlings obtained from ovule culture varied from 0.5–22.5 per cent, whereas in the in vivo situation on the plant no seeds could be harvested.
Embryo rescue technique was attempted in an interspecific hybridisation of tomato variety MT-3 and Kashi Amrit with wild relative S. peruvianum (WIR-3957). The optimum time for rescuing the embryos was standardized as twenty five days after pollination. The most effective media for germination of the immature putative hybrid embryos was Murashige and Skoog’s medium supplemented with 1 mg/L GA3, 0.1 mg/L NAA and 0.5 mg/L BAP. The confirmation of hybridity of this inter-specific crosses was done using RAPD markers. Verba et al. [43] made inter-specific reciprocal crossings between Solanum melongena, S. aethiopicum.
Confirmation of resistance: The confirmation of resistance in the developed varieties/hybrids are usually undertaken by artificial screening studies.
Artificial Screening: This is artificial epidemics created by inoculation of pathogen onto the plant population. The most common methods used to inoculate Colletotrichum on chili plants either involve using the fruit puncture method or the spraying method in the laboratory or in the field [44]. The injection of very small amount of conidia suspension into the fruit pericarp is known as microinjection or fruit puncture method. The resistance can be known by the lesions shown at the injection or inoculation area [20, 21, 45]. Secondly, spraying method in which the conidia suspension is sprayed on plants at flowering and fruiting. But this method is not safe and risky. So the puncture of detached fruits in the laboratory has paid an outstanding development of anthracnose resistance evaluation in chili.
Different artificial screening methods are employed based on the mode of spread of the diseases as given below
Soil borne diseases- Sick plots are created for testing resistance to such diseases.
Air borne diseases- Spraying a suspension of spores.
Seed borne diseases-dry spores are dusted on seeds or seeds may be soaked in a suspension of pathogen spores
14. Conclusion
In vegetables the biotic stresses are pests, diseases and nematodes. The damages induced by these factors affect the production, productivity and quality. Breeding of resistant varieties offer the cheapest means of pest/disease/nematode management. Resistant varieties obviate the use of chemicals, thus reduce environmental pollution. Effectiveness of resistant varieties depend upon the stability of their performance in various environmental conditions and changing climatic scenario. However like any other concept, resistant breeding also has its own limitations. That is breakdown of resistance. However, horizontal resistance being durable but difficulty relates to an accurate & reliable assessment of the level of resistance. A variety resistant to a stress may be susceptible for other. So future planning should require far greater effort for introgression of genes from different resistant sources and develop multiple resistant varieties against several biotic stresses than that required for single.
\n',keywords:"vegetables, biotic stress, breeding methods, varieties",chapterPDFUrl:"https://cdn.intechopen.com/pdfs/74317.pdf",chapterXML:"https://mts.intechopen.com/source/xml/74317.xml",downloadPdfUrl:"/chapter/pdf-download/74317",previewPdfUrl:"/chapter/pdf-preview/74317",totalDownloads:400,totalViews:0,totalCrossrefCites:0,totalDimensionsCites:0,totalAltmetricsMentions:0,introChapter:null,impactScore:0,impactScorePercentile:41,impactScoreQuartile:2,hasAltmetrics:0,dateSubmitted:"July 21st 2020",dateReviewed:"November 10th 2020",datePrePublished:null,datePublished:"May 5th 2021",dateFinished:"December 4th 2020",readingETA:"0",abstract:"In vegetables the factors for biotic stress are pests, diseases and nematodes. The damages induced by these factors reflect highly on production, productivity and quality. Although application of pesticides/fungicides and nematicides has managed these stresses, excessive use of unsafe chemicals results in environmental pollution and leave residues in vegetables which are above threshold levels and also promote the development of new races/biotypes of pests and pathogens. Therefore vegetable improvement works concentrate on high yielding varieties with multiple resistance to these biotic stresses. For such studies, the knowledge on the genetic basis of resistance and plant-pest/pathogen interactions is necessary which will in turn improve the efficiency of the breeding programmes by introducing resistant genes and result in high-yielding genetically resistant cultivars. For the development of resistant varieties and pre-breed lines, information on sources of resistance is prerequisite and serve as a backbone in the breeding programme. Further, gene action responsible for the inheritance of characters helps in the choice of suitable breeding methods for the improvement of the crop. Work has been done by using the various breeding methods and resistant varieties have been bred and they offer the cheapest means of pest/disease/nematode control. Resistant varieties obviate the use of chemicals, thus reduce environmental pollution and facilitate safe food for human consumption.",reviewType:"peer-reviewed",bibtexUrl:"/chapter/bibtex/74317",risUrl:"/chapter/ris/74317",book:{id:"10365",slug:"plant-breeding-current-and-future-views"},signatures:"Ramakrishna Swarnapriya",authors:[{id:"332629",title:"Prof.",name:"Swarna",middleName:null,surname:"Priya",fullName:"Swarna Priya",slug:"swarna-priya",email:"swarnapriya@tnau.ac.in",position:null,profilePictureURL:"//cdnintech.com/web/frontend/www/assets/author.svg",institution:{name:"Tamil Nadu Agricultural University",institutionURL:null,country:{name:"India"}}}],sections:[{id:"sec_1",title:"1. Introduction",level:"1"},{id:"sec_2",title:"2. Genetic basis of resistance",level:"1"},{id:"sec_3",title:"3. Biochemical basis of resistance",level:"1"},{id:"sec_4",title:"4. Biophysical basis of resistance",level:"1"},{id:"sec_5",title:"5. Categorization of resistance",level:"1"},{id:"sec_6",title:"6. Sources of resistance",level:"1"},{id:"sec_7",title:"7. Breeding methods",level:"1"},{id:"sec_8",title:"8. Utilization of wild species",level:"1"},{id:"sec_9",title:"9. Screening varieties",level:"1"},{id:"sec_10",title:"10. Selection",level:"1"},{id:"sec_11",title:"11. Hybridization",level:"1"},{id:"sec_12",title:"12. Mutation breeding",level:"1"},{id:"sec_13",title:"13. Biotechnological approaches",level:"1"},{id:"sec_14",title:"14. Conclusion",level:"1"}],chapterReferences:[{id:"B1",body:'Sarvayya V. 1936. The first generation of an interspecific cross in Solanums between Solanum melongena and S.xanthocarpum. Madras Agric. J., 24(4): 139-142'},{id:"B2",body:'Bonn GW, Kishaba AIT, Principe JA, Toba HH. Tolerance to melon aphid in Cucumismejlo L. J. Amer. Soc. Hort. Sci. 1973;98(l):37-40'},{id:"B3",body:'Fery RL, Cuthbert. Antibiosis in Lycopersicon to the tomato fruit worm (Heliothiszea). J. Am. Soc. 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Resistance to Bemisia tabaci biotype B of Solanum pimpinellifolium is associated with higher densities of type IV glandular trichomes and acylsugar accumulation. Entomologia Experimentalis Et Applicata. 218-230'},{id:"B21",body:'Silva SAM, Rodrigues R, Goncalves LSA, Sudre CP, Bento CS, Carmo MGF, et al. Resistance in Capsicum spp. to anthracnose affected by different stages of fruit development during pre- and postharvest. Trop. Plant Pathology. 2014b;39:335-341'},{id:"B22",body:'Leppik EE. Gene centers of plants as sources of disease resistance. Annual Review of Phytopathology. 1970;8:323-344'},{id:"B23",body:'Kashyap, R. K., M. K. Banerjee, Kalloo and A. N. Verma. 1990. Survival and development of fruit borer, Heliothis Armigera (Lepidoptera: Noctuidae) on Lycopersicon Spp. (Hübner), Intl J. of Tropical Insect Science, 11 (6) : 877-881'},{id:"B24",body:'Sugha SK, Pathania NK, Kumar S. Evaluation of Brinjal germplasm against Phomopsis disease. 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Bangladesh Journal of Agricultural Research. 2009;34:705-712'},{id:"B29",body:'Rehman H, Ayyaz M, Nadeem M, Begum HA. Screening of okra varieties resistance against insect pests under agro climatic conditions of Deraismail khan, Pakistan. Russian Agricultural Sciences. 2017;43(2):149-152'},{id:"B30",body:'Jackson DM, Bohac JR. Resistance of sweetpotato genotypes to adult Diabroticabeetles. Journal of Economic Entomology. 2007;100(2):566-572'},{id:"B31",body:'Mwaura, B. Niere & S. Vidal 2014. Resistance and tolerance of potato varieties to potato rot nematode (Ditylenchus destructor) and stem nematode (Ditylenchus dipsaci). Annals of Applied Biology ISSN 0003-4746. doi:10.1111/aab.12180'},{id:"B32",body:'Nayak DK, Pandey R. Screening and evaluation of brinjal varieties/cultivars against root-knot nematode, Meloidogyne incognita. International Journal of Advanced Research. 2015;3(10):476-479'},{id:"B33",body:'Akhter G, Khan TA. Response of brinjal (Solanum melongena L.) varieties for resistance against root-knot nematode, Meloidogyne incognita race-1. The Journal of Phytopharmacology. 2018;7(3):222-224'},{id:"B34",body:'Hayes, H. K., Immer, F. R., Smith, D. C. 1955. Methods of plant breeding. 2nd ed. McGrawHill: New York 5 5 1 pp'},{id:"B35",body:'Coyne CJ, Porter LD, Inglis DA, et al. Registration of W6 26740, W6 26743 and W6 26745 green pea germplasm resistant to fusarium root rot. J Plant Registration. 2008;2:137-139p'},{id:"B36",body:'Pinheiro, JB; Silva, GO; Macêdo, AG; Biscaia, D; Ragassi, CF; Ribeiro, CSC; Carvalho, SIC; Reifschneider, FJB. 2020. New resistance sources to root-knot nematode in Capsicum pepper. Horticultura Brasileira 38: 33-40. DOI - doi:10.1590/S0102-053620200105'},{id:"B37",body:'Kishaba AN, Castle SJ, Coudriet DL, McCreight JD, Bohn GW. Resistance to western spotted and striped cucumber beetle in melon. Cucirbitacea. pp. 1998;101-105'},{id:"B38",body:'Jagesh Kumar Tiwari, SK Luthra, Sapna Devi, Vinod Kumar, Nilofer Ali, Rasna Zinta and SK Chakrabarti (2018). Development of advanced back-cross Progenies of potato somatic hybrids and linked ISSR markers for late blight resistance with diverse genetic base- first ever produced in indian potato breeding. Potato J. 45 (1): January – June'},{id:"B39",body:'Gritton ET. Pea breeding. In: Basset MJ, editor. Breeding Vegetable Crops. Westport, Connecticut: AVI Publishing Company, Inc.; 1986. pp. 283-319'},{id:"B40",body:'Collard BC, Mackill DJ. Marker-assisted selection: An approach for precision plant breeding in the twenty-first century. Philos. Trans. R. Soc. Lond. B Biol. Sci. 2008;363:557-572'},{id:"B41",body:'Voorrips RE, Finkers R, Sanjaya L, Groenwold R. QTL mapping of anthracnose (Colletotrichum spp.) resistance in a cross between Capsicum annum and C. chinense. Theor. Appl. Genet. 2004;109:1275-1282'},{id:"B42",body:'Aslı Kara and Şerife Evrim Arici (2019). Determination of Gamma Rays Efficiency against Rhizoctonia solani in Potatoes. Open Chem.,17: 254-259'},{id:"B43",body:'Verba VM, Mamedov MI, Pyshnaya ON, Suprunova TN, Shmykova NA. Isolation of eggplant interspecific hybrids by the method of embryo culture. Agricultural Biology. 2010;5:66-71'},{id:"B44",body:'Susheela K. Evaluation of screening method for anthracnose disease in chili. Pest Manag. Hortic. Ecosyst. 2012;18:188-193'},{id:"B45",body:'Mahmodi F, Kadir J, Puteh A. Genetic diversity and pathogenic variability of Colletotrichum truncatum causing anthracnose of pepper in Malaysia. Journal of Phytopathology. 2014;162:456-465'}],footnotes:[],contributors:[{corresp:"yes",contributorFullName:"Ramakrishna Swarnapriya",address:"swarnapriya@tnau.ac.in",affiliation:'
Department of Vegetable Science, Horticultural College and Research Institute, Tamil Nadu Agricultural University, Coimbatore, India
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1. Introduction
Ulcerative colitis (UC) is a chronic inflammatory disorder of the colon characterized by remissions and relapses. The disease almost always begins in the rectum and extends proximally to involve the colon. In some patients, the disease is confined to the rectum (ulcerative proctitis) and in others, the inflammation extends to a variable distance along the colon in a continuous manner. The entire colon is involved in some patients (pancolitis). The clinical presentation and the disease course vary widely amongst patients. Majority of the patients present with an acute episode that clinically mimics any of the acute colitis. The patients on surveillance could show minimal clinical symptoms and minimal histological changes during the periods of remission. Some patients have continuous low activity while some may have an initial episode of active disease followed by long periods of quiescence where the accuracy of the initial diagnosis becomes questionable [1]. Some patients present with the fulminant disease either as the first attack or in an acute exacerbation and this may lead to resection of the colon as an emergency measure.
The pathologist plays a major role in the diagnosis and the management of this chronic relapsing disease. The differing clinical presentations, the chronic relapsing and remitting nature of the disease resulting in recurrent mucosal damage and healing, and the iatrogenic interventions both medical and surgical lead to a variable pathological picture (both macroscopic and microscopic) making the pathologists task a difficult one. Therefore, the diagnosis of UC should always be a combined effort of the clinician, endoscopist, and pathologist. The pathologist should be provided with information regarding clinical symptoms, their duration, the clinical course of the disease, the treatment received by the patient, and the endoscopic appearance of the bowel.
The pathologists are called upon to play various roles during the management of UC. These include the initial diagnosis of the disease and its differentiation from the other forms of colitis and in the classification of inflammatory bowel disease (IBD) in differentiating it from Crohn disease (CD). The morphological differential diagnosis of UC is wide and depends on whether the biopsies are performed during an acute relapse, in remission, or while on treatment.
Assessment of disease activity in UC is another responsibility of the pathologist. There are various clinical and endoscopic activity indices; however, histology is considered to be the gold standard in assessing the disease activity in UC [2]. There are various activity indices used by pathologists. The selection of the activity index that is used in practice is largely determined by the preference of the reporting pathologist and the responsible clinicians [3].
Pathologists are also required to identify dysplasia associated with UC. Dysplasia is an indicator of poor prognosis in UC patients, with a high risk of evolution towards invasive colorectal adenocarcinoma in the absence of treatment. Diagnosis and classification of dysplasia in UC is a challenge to the pathologist and it is a crucial step in the surveillance and management of the patients.
Basic knowledge of the histopathology of this disease is important for clinicians managing UC to interpret and act on the information provided in the pathology report. Here, the morphological features of UC are reviewed, with an emphasis on typical features as well as atypical features that could cause diagnostic pitfalls. The challenges the pathologist faces when examining the diagnostic material at different stages of the disease are highlighted. Further, the use of histological indices for the evaluation of disease activity, identification, and grading of dysplasia associated with UC are also discussed.
2. Macroscopic appearance of ulcerative colitis
UC is characterized by diffuse, continuous inflammation without skip lesions, restricted to the rectal and colonic mucosa. The bowel is characteristically filled with blood-stained dark fluid mixed with mucus. At the onset of the disease, the mucosa shows diffuse granularity, oedema, and erythema justifying the term ‘red velvety’ appearance. With the progression of the disease, the mucosa becomes friable with the appearance of punctate ulcers followed by irregular broad-based ulcers of various sizes. Ulceration may undermine the mucosa creating mucosal bridges. Ulcers distributed along the long axis of the colon extending down to muscularis propria are seen in UC but not the serpentine ulcers that are characteristic of CD. These mucosal changes involve the rectum and variable lengths of the proximal colon in continuity. The distal colon is more severely diseased than the proximal colon. The margin between the inflamed and normal mucosa is distinct and abrupt.
Against a backdrop of mucosal ulceration, there are islands of non-ulcerated residual and regenerating mucosa, bulging into the lumen creating inflammatory pseudopolyps which are usually small, multiple, and bizarre in shape. Sometimes these pseudopolyps could be large and may mimic malignancy (Figure 1) [4].
Figure 1.
Macroscopic appearance of the colon in UC showing involvement of the entire colon with ulceration and pseudopolyp formation.
The mucosa is atrophic and smooth with the absence of mucosal folds in long-standing UC. In the quiescent stage of UC, the mucosa may appear normal or exhibit diffuse granularity and inflammatory pseudopolyps.
The extent of UC is classified according to the Montreal classification as follows [5];
a. Ulcerative proctitis
Only the rectum is affected
b. Left-sided or distal UC
Colonic involvement distal to the splenic flexure
c. Extensive UC or pancolitis
Involvement of the colon extending proximal to the splenic flexure
3. Microscopy appearance of ulcerative colitis
For the optimum assessment of the microscopy, the pathologist should be supplied with adequate and good-quality colonoscopic biopsies. The endoscopist should obtain samples from the ileum, at least four colonic sites, and the rectum, with a minimum of two biopsies from each site [6].
Biopsies from each colonic segment should be submitted in separate containers for each colonic segment, which should be accurately labeled. This is of paramount importance as the pathologist has no means of identifying the separate colonic segments/sites based on histology. This in turn will hinder the pathologists’ attempts at mapping the pattern and distribution of disease activity throughout the colon.
Samples should be fixed immediately by immersion in buffered formalin solution before transport and should be accompanied by clinical information, including endoscopic findings, duration of disease, and current treatment. It is important to sample both endoscopically normal as well as abnormal mucosa as there could be histological activity identified in even endoscopically normal mucosa [7, 8]. Proper orientation of the biopsy during tissue embedding is important as tangentially sectioned biopsies hinder the assessment of the crypt architecture. Serial sectioning is also vital as certain focal features like granulomas may appear at different levels. However, the ideal number of sections is not defined and varies with the laboratory. At least two tissue levels, and preferably three, are advisable [2]. Some laboratories produce step-sections on two slides, but this may incur extra costs. Routine staining with hematoxylin and eosin is appropriate. Special stains and immunohistochemistry are not routinely necessary for the diagnosis or classification of IBD [2].
3.1 Main histological characteristics of ulcerative colitis
The knowledge of the normal histology of the gastrointestinal mucosa is essential for the optimal interpretation of biopsy specimens in IBD. Four main histological characteristics are assessed in the diagnostic process of IBD.
3.1.1 Crypt architectural abnormalities
Normal colonic crypts are straight, parallel, and extend from the surface, up to the muscularis mucosae. The crypt architecture is assessed based on the crypt size, crypt branching, crypt shortening, and variability in inter-crypt spacing. The crypt architectural distortion observed in UC is characterized by irregularly arranged, dilated, branched, fused, and shortened crypts. The crypt size and spacing could vary. This is a manifestation of ongoing inflammation and regeneration [9]. However, the crypts in the anorectal junction and ileocaecal junction can show minor abnormalities resembling architectural distortion due to regional variations. Another point to remember is that the crypt architecture should not be assessed close to lymphoid aggregates as these could result in distortion.
3.1.2 Cellularity in the lamina propria
Lymphocytes and plasma cells are always found in the colorectal lamina propria (LP). Normally, these chronic inflammatory cells are most dense in the upper third of the mucosa and their density decreases towards the base, resulting in the ‘plasma cell gradient’. The absence of the plasma cell gradient is accepted as normal in the caecum and ascending colon. The cellularity in the LP varies depending on the anatomical site of the colon. In general, the caecum and the right colon are most cellular with a progressive decrease in the cellularity from the right to the left side. Dense lymphoid tissue may be found in the normal large bowel, particularly in the rectum. The abundance of eosinophil granulocytes varies a lot between normal individuals and is found to have a seasonal and geographic variation and is more in the right colon and the ileum than the left colon [2, 10].
Abnormal cellularity of the LP refers both to increased and altered distribution of cell types that are normally present. Basal plasmacytosis or plasma cells extending below crypt endings in more than two foci in a biopsy is considered to denote an increase in chronic inflammation (Figure 2).
Figure 2.
Main histological features of UC. (A) Basal plasmacytosis and mucin depletion. (B) Villiform surface, variation in size and shape and branching of crypts. (C) Surface ulceration, diffuse heavy inflammatory cell infiltrate in the lamina propria and crypt abscess formation ((A–C): H&E ×200).
3.1.3 Neutrophil granulocytes in the lamina propria
Normal colonic mucosa does not contain neutrophils except a few scattered neutrophils, that may occur as a result of bowel preparation [11]. Neutrophils are the hallmark of activity in IBD/UC. These are found in the LP or they can infiltrate the crypt surface epithelium (termed cryptitis) or enter the lumen of crypts forming crypt abscesses (Figure 2).
3.1.4 Epithelial abnormalities
These include mucin depletion, surface epithelial damage, and metaplastic changes. Mucin depletion can be defined as a decreased number of goblet cells or decreased amount of intracellular mucin. Focal epithelial cell loss, flattening, erosions, and ulcers denote epithelial damage and reflect the activity of the disease. Metaplastic changes are seen in the form of Paneth cell or pyloric gland metaplasia. None of these findings is disease-specific and might be observed in UC, CD, and other types of colitis.
Working definitions of some of the microscopic features of IBD/UC are given in Table 1 [11].
Microscopic abnormality
Definition
Additional remarks
Crypt distortion
Branching, loss of parallelism, irregularity, tortuosity, dilatation, and variation in shape and size of the crypts
Should not be assessed adjacent to crypt abscesses/lymphoid aggregates/follicles Anal transition zone/columnar cuff unsuitable for assessment
Crypt branching
Two or more branched crypts in a well-oriented biopsy
Branching between mucosal hillocks is normal
Crypt atrophy
Crypt shortening, with an increased gap between crypt base and muscularis mucosae Additional evidence is the wider spacing of crypts; >1 crypt diameter between crypts (normal: 6 crypts/mm in a biopsy with muscularis mucosa)
Caution adjacent to lymphoid aggregates/follicles Anal transition zone/columnar cuff is unsuitable for assessment Mucosal oedema may mimic atrophy
Villiform/irregular mucosal surface
Undulating or broadly villiform surface Wide crypt mouths
Basal plasmacytosis
Plasma cells at the base of mucosa. May separate crypts from muscularis mucosae but not always sub cryptal Loss of plasma cell gradient
Basal plasma cells are normal in the caecum and the ascending colon
Basal lymphoid aggregates
Nodular collections of lymphocytes with or without germinal centres. It may be seen between muscularis mucosae and crypts
One or two transmucosal lymphoid nodules can be seen in normal mucosa; can extend across muscularis mucosae. Pathological aggregate may be difficult to distinguish from normal
Cryptitis
Neutrophils in the crypt epithelium
Crypt abscess
Neutrophils in crypt lumen
Often located near the crypt base
Granuloma
A discrete collection of at least five epithelioid macrophages
Consider crypt rupture as a cause. Serial sections may help
Mucin depletion
Unequivocal reduction of goblet cell mucin in the crypt epithelium
Mucin in normal mucosa may be reduced near lymphoid follicles. Depletion can reflect bowel preparation
Ulceration/erosion
Loss of epithelium replaced by ‘immature’ granulation tissue or the presence of a fibrinopurulent exudate An ulcer extends more deeply than the muscularis mucosae while an erosion does not
It May be difficult to differentiate ulcer and erosion The epithelium can denude artifactually during biopsy procedures or processing
Paneth cell metaplasia
Pyramidal crypt epithelial cells with supranuclear eosinophilic granular cytoplasm
Normal in the caecum and right colon, probably as far as the splenic flexure
Diffuse chronic inflammation
An overall increase in chronic inflammatory cell density throughout the biopsy/biopsies
The caecum and ascending colon may have a higher density of chronic inflammatory cells
Patchy chronic inflammation
Areas of increased chronic inflammatory cell density in a background of variable cellularity
Focal chronic inflammation
Well circumscribed foci of increased chronic inflammatory cell density in a normocellular background
Differentiate from lymphoid aggregates
Focal active inflammation
Focal crypt infiltration by neutrophils in the absence of significant inflammation in the adjacent LP
Uncommon in UC but could be seen in early or treated UC
Table 1.
Working definitions of histological features seen in UC/IBD.
3.2 Typical histological features of ulcerative colitis
The histologic findings in UC vary depending on the clinical phase of the disease and the grade of inflammatory activity. The histological features that define chronicity are crypt architectural distortion, crypt atrophy, diffuse mixed lamina propria inflammation, basal plasmacytosis, basally located lymphoid aggregates, and Paneth cell metaplasia [9]. Inflammatory activity is defined by the presence of neutrophils. Neutrophilic cryptitis, crypt abscesses, hemorrhage, erosions, ulceration, and necrosis are features of active inflammation.
It is convenient to divide the histologic appearances into those seen inactive disease, resolving disease, and disease in remission.
3.2.1 Active ulcerative colitis
The characteristic features of acute UC include architectural distortion which is more in the distal colon than the proximal bowel, diffuse chronic inflammatory cell infiltrates extending up to the muscularis mucosae, and neutrophil infiltration. The neutrophils seem to migrate directly from capillaries into the crypt epithelium (cryptitis) and often form crypt abscesses. Ulcers covered with granulation tissue and regenerative epithelium could be seen. The surface epithelium may take an undulating or low villiform appearance. The inflammation may extend into the superficial submucosa but the muscularis propria and serosa remain free of inflammation, except in fulminant colitis.
Neutrophils are predominant within the lumina of the crypts in UC and comparatively small numbers are seen migrating between the epithelial cells. Crypt abscesses are the precursors of mucosal ulceration and inflammatory polyp formation. In severe active UC, crypt abscesses burst into the loose submucosal tissues and there is a tendency to spread beneath the mucosal membrane, which sloughs off leaving an ulcer. The remaining mucosa is relatively raised from the ulcerated area and forms ‘inflammatory pseudopolyps’.
The inflammatory damage to the crypts produces a variety of degenerative and regenerative changes in the crypt epithelium. There is loss of mucin from goblet cells, often with enlargement and hyperchromatism of nuclei of the absorptive cells. Such changes must not be mistaken for dysplasia. In the presence of attenuated or restituting superficial epithelium, the changes in the crypts are almost certainly reactive [1].
There is a heavy diffuse infiltrate of inflammatory cells in the LP. These include neutrophils, lymphocytes, plasma cells, eosinophils, and mast cells. The presence of deep plasma cells is characteristic of longstanding UC. Many eosinophils may be seen in the LP in some of the biopsies but the role of eosinophils in UC is uncertain and has been subject to many studies [10]. Lymphoid aggregates, that may show germinal centres, are common in UC. These are situated basally between crypt bases and the muscularis mucosae (Figure 3). Epithelioid granulomas, which are a hallmark of CD, are not identified in UC, where only foreign body granulomas evolved as a response to mucin from ruptured crypts (termed cryptolytic granulomas) are observed.
Figure 3.
Histological appearances of active UC. (A) Depletion of mucin in the epithelium, cryptitis, heavy infiltrate of lymphocytes, plasma cells, and eosinophils in the LP (H&E ×400). (B) Marked crypt distortion and crypt abscess formation. (C) Surface ulceration. (D) A lymphoid aggregate with the germinal centre formation in the LP ((B–D): H&E ×200).
3.2.2 Resolving ulcerative colitis
With the resolution of disease, the numbers of inflammatory cells of all types show a variable density, therefore, the LP could be either hyper or hypocellular. Further, the distribution of inflammatory cells becomes uneven. The goblet cell population returns to normal. The crypt architecture will show variable evidence of distortion, depending on the severity of the attack. The crypts may appear short and branched, the changes invariably being more marked distally. The resolution may occur at different rates in different parts of the colon, and this may give rise to the false impression of segmental disease (Figure 4).
Figure 4.
Resolving UC; a mild crypt architectural distortion, normal mucin content in the glands, hypocellular LP, no activity (H&E ×100).
3.2.3 Quiescent ulcerative colitis
Varying degrees of crypt atrophy and distortion are the hallmarks of the quiescent disease. Active inflammation is absent. The muscularis mucosa is thickened and a characteristic finding in UC, especially in rectal biopsies, is the double muscularis mucosae [1]. Paneth cell metaplasia in the left colon and pyloric-type metaplasia in any location of the colon are also features of chronic disease [1].
Although most UC patients have some residual changes of previous damage, such as crypt distortion, atrophy, and Paneth cell metaplasia, it has become increasingly recognized that a group of UC patients may show complete resolution with no evidence of previous disease [12, 13]. This must always raise the question of whether the original diagnosis is UC or infective colitis. In such situations, a careful review of the previous clinical records including the initial biopsies and ancillary investigations may be of help [1].
4. Unusual morphological patterns of ulcerative colitis
There are some exceptions to the classic morphological patterns described above, that may lead to diagnostic confusion. A summary of the unusual morphologic patterns of disease in UC is noted in Table 2. Pathologists need to recognize these patterns to avoid falling into a diagnostic trap.
1. Treatment effect
2. Appendiceal ‘skip’ lesion and ‘caecal patch’
3. Ileal involvement
4. Rectal sparing
5. Paediatric UC-initial presentation
6. Fulminant colitis
Table 2.
Unusual patterns of disease in UC.
4.1 Morphological features of treated ulcerative colitis
Endoscopically or histologically discontinuous disease may be observed in the setting of medically treated UC. This occurs as a result of uneven healing [6]. The same process may also lead to absolute or relative rectal sparing in 30–40% of patients [9]. As patchiness of the disease and rectal sparing mimicking CD is commonly seen in treated UC, evaluation of disease distribution to subtype IBD should not be attempted in this setting. This emphasizes the importance of pre-treatment histology and the value of communication between the clinician and the pathologist.
4.2 Appendiceal ‘skip’ lesions and the ‘caecal patch’ in ulcerative colitis
Appendiceal involvement is demonstrated in about 75% of the total colectomies performed for UC. This may be continuous with extensive colitis or may represent a ‘skip lesion’ of UC with involvement of the more distal colon only [9]. Such a skip lesion may raise the suspicion of CD and be erroneously considered as a contraindication for pouch surgery. The mucosal inflammation in the appendix may extend to the appendiceal orifice and contiguous large bowel as a periappendiceal patch.
Similar to the appendiceal skip lesion, there could be isolated involvement of the caecum and/or ascending colon, which is discontinuous with the left-sided colitis. It is shown that patchy right-sided inflammation in patients with left-sided colitis has little clinical significance but should be recognized by pathologists to prevent a false diagnosis of CD in this setting [14].
It is also interesting that in appendices removed for possible acute appendicitis, inflammation confined to the mucosa with associated crypt distortion should raise the possibility of an appendiceal involvement in UC.
4.3 Involvement of the ileum
Ileitis is found in about 10% of patients with UC, the extent of involvement varying from 50–250 mm [1]. The mucosal changes are similar to those seen in the colon and are always in continuity with the disease in the large bowel. The involvement of the ileum is associated with an open dilated and incompetent ileocaecal valve. Although the term ‘backwash ileitis’ is in common use for this condition, it is not necessarily accurate as evidence for such a mechanism is not yet proven [1]. This condition should not be confused with CD of the terminal ileum which typically shows longer lengths of involvement and is normally associated with chronic active inflammation, and other features of CD. Unfortunately, strict histopathologic criteria for backwash ileitis have not been defined [15].
4.4 Rectal sparing
According to traditional belief, UC is a diffuse continuous disease that begins in the rectum and extends proximally, without skip areas. The term ‘absolute rectal sparing’ refers to the rectum with a normal endoscopic appearance and normal histology. Another term sometimes used is ‘relative rectal sparing’, in which the rectum has inflammation that is less severe than the more proximal colon [16]. Rectal sparing and patchiness of inflammation are seen in medically treated UC, especially with therapeutic steroid enemas. This too emphasizes the importance of the provision of clinical details to the pathologist. The pathologists should also be vigilant not to interpret these findings as definite evidence of CD.
4.5 Ulcerative colitis in pediatric populations
Pediatric-onset of IBD can show fewer characteristic symptoms and histological findings than adult-onset IBD. In general, diagnostic biopsies from children with UC often show less severe inflammation, fewer architectural abnormalities, and less epithelial injury despite extensive disease [2, 6, 17, 18].
The available evidence strongly suggests that UC in children is typically a pancolitis with variable degrees of inflammation on histology [16]. In a subset of pediatric patients, relative rectal sparing and patchy inflammation both endoscopically and histologically may occur at the pre-treatment stage of UC. These features are also observed in treated pediatric patients presumably similar to adults. In a minority, absolute rectal sparing occurs [16]. It is prudent not to preclude the diagnosis of UC in children when these atypical features are present. The precise reason why pre-treatment stage pediatric patients have a higher prevalence of rectal sparing compared to adults is unclear. Younger age (<10 years) at presentation and shorter duration between the symptoms and endoscopy are proposed explanations [19].
4.6 Fulminant ulcerative colitis
Fulminant colitis is defined as severe, acute inflammation of the colon with associated systemic toxicity. Most cases (89%) of fulminant colitis represent IBD, with the remainder relating to ischemia or infection, amongst other aetiologies [20].
In fulminant UC, the inflammatory cell infiltrate extends beyond the mucosa with thinning of the wall. There is separation and oedema of the muscle layer known as myocytolysis. There is diffuse haemorrhagic necrosis of the mucosa, deep fissuring ulcers, and transmural polymorphous inflammation. Unlike in classical UC where the serosa is shiny and intact, there is a purulent or seropurulent exudate seen on the peritoneal surface in many cases of fulminant colitis. The bowel wall is also markedly thinned out and dilated and this usually occurs in the region of the transverse colon. Macroscopic features, such as dilation, skip lesions, rectal sparing, linear ulcers, terminal ileal disease, pseudopolyps, and creeping fat, are poor discriminators of UC and CD, in the setting of fulminant colitis.
5. Histological differential diagnosis of ulcerative colitis and diagnostic pitfalls
5.1 Infective colitis
Infective colitis may clinically mimic acute UC. However, most cases of infectious colitis demonstrate a histological pattern of acute colitis, which may be diffuse, patchy, or focal, without evidence of architectural distortion. Less commonly, chronic infectious colitis may produce a histological pattern of chronic active colitis resembling IBD. Most of these cases have no specific diagnostic features on histological examination and in such cases, knowledge of the clinical history and correlation with serologic studies or stool cultures are required for diagnosis.
Some features help in differentiating acute self-limiting colitis from UC in the acute stage. In acute self-limiting colitis, the neutrophils are plentiful in the LP and are more superficially arranged. In UC, the neutrophils are predominant within the lumina of the crypts and comparatively small numbers are seen migrating between the epithelial cells [1]. Chronic changes, such as crypt distortion in UC, take about 4–6 weeks to develop and this could, therefore, cause a diagnostic difficulty in the early stages of the disease.
In amoebic colitis, presenting as chronic active colitis, there could be trophozoites of Entamoeba histolytica in biopsy material and identifying the trophozoites becomes crucial because, if immunosuppressive therapy is started for presumed IBD in these patients, it can result in perforation due to fulminant amoebic colitis. Superimposed infection with many organisms can occur with established UC and cytomegalovirus (CMV), campylobacter and Clostridium difficile are some of the important secondary infections to be considered when UC presents with an acute exacerbation [21].
5.2 Chronic ischaemic colitis
Chronic ischemia may produce a pattern of chronic active colitis and can present a difficult differential diagnosis. Chronic or recurrent ischemia may cause significant crypt distortion, Paneth cell metaplasia, and chronic active inflammation, all features which mimic UC. However, atrophic and regenerative changes in the epithelium, hyalinization of the LP, and the presence of microthrombi in the adjacent mucosa should suggest ischemia [21]. Overall, in ischemia, the chronic active inflammation is mild relative to the degree of epithelial injury [21]. Further confounding this differential diagnosis, UC has been reported to cause a hypercoagulable state, particularly in genetically predisposed patients. In this setting, a superimposed arterial and venous thrombosis may occur, leading to severe steroid-refractory colitis [21].
5.3 Diverticular disease-associated colitis (DAC)
Chronic active colitis resembling UC may be seen in the setting of diverticulosis. In addition, diverticulosis is a relatively common disease of the elderly and, thus, both diverticulosis and IBD (either CD or UC) may coexist in the same patient. Unlike chronic active colitis of UC, DAC is confined to segments involved by diverticular disease, most commonly the sigmoid colon, and, by definition, spares the rectum. However, UC and diverticular colitis may in some cases represent overlapping entities, as a small subset of diverticular colitis patients has progressed to typical rectosigmoid UC and DAC may respond to medical therapy utilized for IBD [21].
5.4 Drug-induced colitis
Some forms of medication-induced colitis may demonstrate chronic active colitis, which may enter the differential diagnosis of UC. Nonsteroidal anti-inflammatory drugs (NSAIDs) may result in the reactivation of UC. However, in some patients, this group of drugs has also been implicated in initiating UC [1].
The most useful histological feature in distinguishing NSAID-related colitis from UC is an increase in apoptotic bodies in the crypt epithelium and lymphocytes and mononuclear cells of the superficial LP. The morphological changes that occur with NSAIDs include a generalized increase in chronic inflammatory cells in the LP, a prominent eosinophil infiltrate, increased intra-epithelial T lymphocytes and thickening of the subepithelial collagen plate may resemble eosinophilic colitis, lymphocytic colitis, and collagenous colitis, respectively [1].
5.5 Diversion colitis
Diversion colitis develops in segments of the bowel that have been excluded from the fecal stream, such as in a Hartmann’s pouch. This chronic inflammatory condition usually develops within a few months to several years following surgical diversion, and typically regresses completely within 3–6 months of re-establishment of the fecal stream [22]. The disease may mimic IBD on biopsy samples and may show crypt atrophy, distortion, and lymphoid hyperplasia, involving the mucosa and/or submucosa. Symptomatic patients can show superimposed cryptitis, crypt abscesses, and superficial aphthous-type erosions or frank ulceration.
This warrants the importance of obtaining a biopsy of the segment at the time of the surgery to get a baseline analysis and review of the clinical, radiological, and endoscopic information prior to diagnosis.
5.6 Crohn disease (CD)
In most instances, UC and CD may be readily distinguished from each other pathologically, particularly when each exhibits classic histological features assisted by clinical data and other ancillary investigations. There are several circumstances in which the ‘classic’ morphological features that help to distinguish UC from CD are altered or absent. When these atypical morphological features are present, they may mimic CD (Tables 3 and 4). Most of these atypical features have already been discussed in Section 3.
Pathological feature
Ulcerative colitis
Crohn’s disease
Disease distribution
Diffuse and continuous
Segmental
Rectal involvement
Almost always (adults)
Occasionally
Disease severity
Increased distally
Patchy and variable
Ileal involvement
Occasional (‘backwash’)
Often
Inflammation of the colonic wall
Superficial (mucosal)
Transmural
Transmural lymphoid aggregates
Rare, underneath ulcers
Any location
Fissures
Rare superficial in fulminant colitis
Deep, any location
Sinuses and fistulas
Absent
Present
Granulomas
Related to ruptured crypts ‘cryptolytic granuloma’
Not crypt-related and are epithelioid cell granulomas
Table 3.
Classic morphological features helpful in differentiating UC from CD [13].
1.
Discontinuous or patchy disease (‘caecal patch’)
2.
Absolute or relative rectal sparing
3.
Inflammatory changes in the ileum (‘backwash’ ileitis)
4.
Treatment-related change
5.
Granuloma formation
6.
Transmural inflammation
Table 4.
Unusual morphological patterns of UC that may mimic CD.
5.6.1 Granulomas in UC
Approximately 30–40% of CD cases contain either mucosal or mural, non-necrotic granulomas [22]. When present, it is a helpful feature to confidently diagnose CD, especially in mucosal biopsies. Granulomas in CD are composed of loose collections of CD 68 immunostain positive epithelioid histiocytes. When there is rupture of a crypt or extravasated mucin in UC, there could be the formation of a granuloma which is termed a ‘cryptolytic granuloma’ that could be difficult to distinguish from granulomas of CD (Figure 5). Examination of multiple tissue levels that will demonstrate the relationship between granulomas and the crypt epithelium is a helpful measure in this situation. Cryptolytic granulomas often contain an admixture of neutrophils and lymphocytes, in addition to foamy macrophages and multinucleated foreign body-type giant cells and these are not usually seen in CD-related granulomas [22].
Figure 5.
(A) Cryptolytic granuloma following a rupture of a crypt in UC. Extravasated mucin, multinucleated giant cells, and inflammatory cells are seen. (B) Epithelioid cell granuloma formed by histiocytic cells in CD ((A and B) H&E ×400).
5.6.2 Transmural inflammation in UC
In CD, transmural lymphoid aggregates are seen randomly in the wall of the bowel. In fulminant UC when superficial fissuring ulcers that extend into the deep submucosa or superficial muscularis propria are present and in toxic megacolon when myocyte necrosis and serosal inflammation are prominent, there could be mural mononuclear inflammation. However, in contrast to CD, these do not form typical discrete lymphoid aggregates and are usually seen underlying the areas of severe ulceration [22]. Thus, lymphoid aggregates in areas under intact mucosa are not a feature of UC and, in fact, favor a diagnosis of CD.
5.6.3 CD with UC-like features
Typical features of CD such as granulomas, fissuring ulcers, and transmural lymphoid aggregates are seen less commonly in the colon compared to the small intestine [15]. Therefore, some cases of colonic CD may mimic UC by demonstrating only superficial mucosal involvement without inflammatory changes in the submucosa or muscularis propria, diffuse and continuous disease, and even pancolitis. Nearly 20% of CD patients develop colitis without the involvement of the upper GI tract [22]. In these cases, careful evaluation of colonic and ileal biopsies for granulomas, identifying focal or patchy inflammation and activity within the LP, identifying transmural lymphoid aggregates in resections, correlation with a detailed clinical history and imaging will be of help in differentiating UC from CD.
6. Indeterminate colitis (IC) and inflammatory bowel disease unclassified (IBDU)
In up to 5% of IBD cases, an exact classification of IBD into UC or CD proves difficult due to either the overlapping histological features of the two diseases or to the fact that UC and CD represent two ends of the spectrum of a single disease [1, 6].
Several different terms have been used to refer to this condition, including ‘indeterminate colitis’ (IC), ‘inflammatory bowel disease unclassified’ (IBDU), ‘chronic inflammatory bowel disease unclassified’, and ‘chronic idiopathic inflammatory bowel disease not otherwise specified’. The European Crohn’s and Colitis Organization (ECCO) and the European Society of Pathology (ESP) jointly addressed the ambiguous usage of this terminology, in their consensus report in 2013 [6]. Accordingly, the term IBDU could be used for patients with chronic colitis who have IBD based on the clinical history, but endoscopy and histology of the biopsies show no definitive features of either UC or CD [6]. This term is reserved for biopsy examination as the post-operative examination of resections of such IBDU cases usually provides definitive evidence of UC or CD.
The entity of IBDU is more common in the pediatric population. The possible reasons for this being more colitis than ileitis occurs in CD in early cases and the presence of rectal sparing in UC in the pediatric population [6]. Upper GI biopsies are particularly helpful in these cases.
The pathological diagnosis of IC is made only on resected specimens with the presence of overlapping features or the absence of a clear diagnostic pattern to distinguish CD from UC.
Usually, macroscopically IC shows diffuse disease with involvement of transverse and right colon and less severe inflammation in the distal colon. There is extensive ulceration. Microscopy confirms extensive ulceration with a sharp transition to normal adjacent mucosa and multiple V-shaped ulcers lacking surrounding inflammation. The overlapping histological features of IC are given in Table 5.
Severe mucosal and wall involvement
Non-aggregated transmural inflammation
Fissures reaching the muscularis propria
Discontinuous pattern
Diffuse mucosal disease with normal ileum
Deep mural lymphoid aggregation
Non-necrotizing granulomas in lymph nodes
Anal fistula
Table 5.
Overlapping histological features in indeterminate colitis (IC).
It makes no difference whether the large bowel resection is called ‘UC’ or ‘IC’, but CD needs to be excluded conclusively since an ileal pouch-anal anastomosis (IPAA)/‘pouch’ procedure is generally contradicted in CD.
7. Measuring the disease activity in ulcerative colitis
The complete assessment of disease activity in UC involves symptomatic evaluation, physical examination, measurement of laboratory indices, endoscopic visualization, and the histological assessment of the mucosal inflammation [3]. However, measuring disease activity using all these different parameters is cumbersome and time-consuming in practice and will delay therapy. In routine clinical practice, the disease activity and subsequent medical treatment are usually assessed largely by the clinical symptomatology. Histological assessment of the degree of inflammation is the gold standard for evaluating the true disease activity but its conventional use is limited owing to its inconvenience, invasiveness, and cost [3].
Traditionally, clinical and endoscopic remission were the two main therapeutic targets for UC. However, up to 40% of patients in clinical and endoscopic remission show persistent histological activity [2]. Furthermore, histological activity predicts the worst outcome and histological inflammation represents a significant risk factor for the subsequent development of UC-related colorectal neoplasia. Therefore, histological remission is now increasingly regarded as an important therapeutic target for UC [2].
There are about 30 histological activity indices in IBD, that have been introduced over the last few decades [23]. These systems use different stepwise grading scales for the assessment of inflammation which is used as the basis of grading the disease activity. These scales have four to seven steps and quantitatively assess the following features—architectural changes in the mucosa, chronic inflammatory cell infiltrate, amount and location of neutrophils within the mucosa, crypt abscess formation, erosion, and ulcers.
Of these scores, the ‘Geboes score’ developed in 2000 has been the most widely used and can serve as an independent risk factor for disease progression in UC [24, 25]. The more recent ‘Nancy histological index’ (NHI) and ‘Robarts histopathological index’ (RHI) both from 2016 have proven feasible, easy to use, and are the most extensively validated [26, 27]. There is currently no general agreement on which index should be used. The 2020 ECCO position paper concluded that the NHI can be recommended for daily clinical practice and for clinical trials both the NHI and RHI are feasible [2].
RHI requires assessment of four features which include ulceration/erosion, neutrophils in the epithelium, neutrophils in the LP, and the chronic inflammatory cell infiltrate. Each of the features is subdivided on a scale of 0–3 to calculate the ultimate disease activity score. This may reduce its clinical usefulness and probably is more useful for clinical trials and in research. In NHI, three main histological characteristics which include ulceration, the acute inflammatory cell infiltrate and the chronic inflammatory cell infiltrate are assessed. The NHI is defined by a 5-level classification ranging from grade-0 (absence of significant histological disease activity) to grade-4 (severely active disease) (Table 6) [27].
Histological criteria and defining features
Disease activity
Score
Ulceration Loss of colonic crypts replaced with immature granulation tissue or presence of a fibrinopurulent exudate
Severe
4
Acute inflammatory cells infiltrate Presence of neutrophils in LP and/or epithelial cells
Moderate to severe Presence of multiple clusters of neutrophils in LP and/or in the epithelium that is easily apparent.
3
Mild Few or rare neutrophils in LP or in the epithelium that are difficult to see
2
Absence Assess the chronic inflammatory cell infiltrate
Chronic inflammatory infiltrate Presence of lymphocytes and/or plasma cells and/or eosinophils in LP
Moderate to severe Presence of an increase in chronic inflammatory cell number that is easily apparent
1
Mild No or mild increase in chronic inflammatory cell number
0
Table 6.
Nancy histological index (NHI).
In the NHI, chronic inflammation includes lymphocytes, plasma cells, and eosinophils and it is assessed without quantification. Furthermore, if neutrophilic inflammation is present, regardless of extent, the degree of chronic inflammation is not assessed.
It has been shown that both NHI and RHI have a similar degree of inter and intra-observer agreement and share equivalent feasibility in terms of time taken for scoring the biopsies [28].
Despite the development and validation of novel histologic scoring systems, there are no agreed definitions for histologic healing and remission. Histologic healing is the ultimate goal of the treatment and could be defined as complete normalization of the mucosa [7]. What constitutes complete normalization needs to be precisely defined. Rare architecturally distorted crypts should not be overinterpreted as evidence of persistent architectural abnormalities. A rare, branched crypt can be seen even in a normal colon. Furthermore, the crypts in a normal rectum often do not extend to the muscularis mucosae [7].
The best definition of histologic remission in UC is also unclear. Traditionally, this has been regarded as persistent architectural abnormalities without neutrophilic (active) inflammation, with varying degrees of lymphoplasmacytic inflammation. The presence of mucosal eosinophils is allowed [7]. Ideally, remission includes clinical, endoscopic, and histological resolution, which is called complete remission.
8. Dysplasia and malignancy in ulcerative colitis
The risk of colorectal carcinoma (CRC) is increased in patients with long-term UC compared to the general population. Carcinogenesis in UC is inflammation-driven and has a different pathway than usual colorectal carcinogenesis. Epithelial cells acquire early mutations of TP53 and KRAS genes and no mutations of APC genes, while in non-inflammatory carcinogenesis of the colon, APC mutation is the earliest event [7].
Epithelial dysplasia is the precursor lesion of UC-associated CRC. The features associated with increased risk of dysplasia/CRC in UC include the duration of the disease, the anatomical extent of the disease, early age of onset, concomitant sclerosing cholangitis, family history of CRC, and endoscopic/histological activity of the disease [29]. The prognosis of CRC in IBD may be worse than CRC in the general population and shows higher mortality [9].
8.1 Colorectal dysplasia in ulcerative colitis
The presence of dysplasia in endoscopic biopsies is the most reliable marker of cancer risk. There are no specific clinical features related to dysplasia in UC. Most cases of dysplasia occur in the left/distal colon, and this mirrors the higher incidence of UC-associated colorectal carcinoma (CRC) in the rectosigmoid region. The endoscopic appearance of dysplasia is categorized according to the SCENIC classification and includes visible and invisible lesions. The visible lesions are subdivided into polypoidal (either pedunculated or sessile) and non-polypoidal (superficial, flat, depressed [29].
Dysplasia of the colorectum is defined as an unequivocal epithelial alteration that remains confined within the basement membrane within which it originated. The microscopic features of dysplasia in UC are based on a combination of cytoarchitectural features of the crypt epithelium that remains confined to the mucosa and are identical to those used in the general assessment of dysplasia elsewhere [9]. Dysplasia is classified according to either the Riddell or the Vienna system [30, 31]. In the Riddell system, there are four categories for dysplasia, which are negative, indefinite, low grade, and high grade. The Vienna system has five categories with the addition of invasive carcinoma.
The most common histological subtypes of dysplasia include intestinal (adenomatous) and serrated types [29]. Regardless of these subtypes, dysplasia is divided into low grade and high grade according to the cytoarchitectural features. In low-grade dysplasia (LGD) the crypts may be tubular and/or villous or serrated and they show either no or only mild crypt budding or crowding. The dysplastic cells show enlarged, hyperchromatic nuclei with a high nuclear/cytoplasmic ratio, nuclear stratification limited to the basal half of the cytoplasm, and clumped chromatin or multiple small nucleoli. In serrated dysplasia, the dysplastic cells may show hypereosinophilic, mucin depleted cytoplasm, or a microvesicular epithelium that is similar to the sporadic sessile serrated adenomas. These atypical nuclear features usually involve both the crypt and surface epithelium.
High-grade dysplasia (HGD) exhibits enlarged nuclei with marked nuclear hyperchromasia, pleomorphism, stratification involving the full thickness of the cytoplasm, increased mitoses, and loss of nuclear polarity. It shows complex glandular architecture with crowding, cribriforming, complex branching, and budding [29].
The category indefinite for dysplasia refers to ambiguous epithelial alterations that cannot with certainty be classified either as negative or positive for dysplasia. Some of the settings in which indefinite for dysplasia is considered are shown in Table 7.
1.
Atypical cytological features in an inflammatory background where differentiating regenerative change from LGD/HGD could be difficult
2.
Marked cytological atypia in the crypt bases where surface maturation cannot be assessed due to ulceration or poor orientation
3.
Various artifactual cytological changes occur as a result of poor histological techniques—processing, cutting, staining
4.
Only a very small focus (only a few crypts) shows dysplasia.
Table 7.
Some settings where indefinite for dysplasia are considered.
8.2 Some issues in histopathological reporting in UC-associated dysplasia
Examination of multiple biopsies is necessary to rule out the possibility of dysplasia confidently, as dysplasia is also identified in endoscopically normal mucosa.
The active and resolving phase of UC may cause diagnostic difficulties as the damaged or the regenerative epithelium may harbor mucin depleted cells and atypical nuclear changes, such as enlargement, hyperchromasia, stratification, and brisk mitoses. Careful observation for surface maturation, which is a feature in reactive conditions helps to solve this problem. Another good practice is to perform colonoscopic surveillance during a period of remission of UC.
Inter-observer agreement amongst pathologists for dysplasia associated with UC is suboptimal. Poor reproducibility is seen mainly in indefinite dysplasia and LGD groups. (6). Reporting the biopsies for dysplasia ideally by two histopathologists will help to overcome this problem. A review of the biopsies with the diagnosis of dysplasia by a more experienced gastrointestinal pathologist, before any surgical intervention is undertaken, will optimize the management of these patients.
Adenomas can arise in both affected and non-affected mucosa in UC patients. They are treated like any other adenomas by complete local excision. The stalk or the tissue around the base of the lesion needs to be examined carefully to confirm that these are adenomas or part of a more widespread area of dysplasia associated with UC.
Colonic biopsies of patients treated with immunosuppressive agents, such as cyclosporin for severe UC, are known to show ‘pseudo dysplastic’ changes in the epithelium [32]. Perhaps the most helpful feature is that the ‘pseudo-dysplasia’ induced by cyclosporin is strikingly diffuse, with many, and sometimes all, crypts showing similar changes, a pattern not usually seen in UC associated dysplasia. Therefore, the clinician needs to alert the pathologist to the fact that the patient has been on cyclosporin and that the pathologist in turn should be cautious when diagnosis dysplasia in this situation (Figure 6).
Figure 6.
Dysplasia in UC. (A) Flat low-grade dysplasia with villous configuration. There is no significant pleomorphism or loss of polarity (H&E ×100). (B) High-grade dysplasia with crowding glandular proliferation (H&E ×100). Reprinted by ([21], pp. 178-192). Published by Oxford University press and digestive science publishing Co Limited.
8.3 The demise of the term ‘DALM’ in ulcerative colitis
A diagnosis of dysplasia is made on biopsy material taken from a polyp or a mass evident on endoscopy was historically termed ‘dysplasia associated lesion or mass’ (DALM) and was considered as an indication for colectomy to rule out the possibility of invasive malignancy. In 2015 SCENIC international consensus statement on the surveillance and management of dysplasia in IBD, abandoned the term DALM and replaced it with endoscopically visible and non-visible lesions [33]. With advancements in endoscopic polypectomy and endoscopic mucosal resections (EMR), the concept of DALM is now outdated because most lesions that are noninvasive can be removed using these techniques [1].
8.4 Carcinoma in ulcerative colitis
Carcinomas arising in UC are mostly similar to their counterpart in non-colitis patients except for the background colitis. However, there are some features that are more frequent in UC-associated carcinomas. The tumors could be multiple and often are flat lesions with ill-defined edges, therefore, these tumors are easily felt than seen. Histologically there is a higher incidence of high-grade tumors and mucinous subtypes [1].
9. Conclusions
The pathologist plays a vital role in the diagnosis and follows up of patients with UC. The histological features in the biopsies vary widely depending on the stage of this chronic relapsing and remitting disease, making the differential diagnosis lengthy and challenging. The final diagnosis should be ideally concluded at a clinicopathological meeting. Understanding the typical and atypical histological features of UC is vital in the task of differentiating UC from other types of colitis, mainly CD. Histological disease activity and identifying histological remission are increasingly considered important therapeutic targets. Identifying dysplasia associated with UC and its grading is a crucial step in the surveillance and management of this chronic disease.
Acknowledgments
The authors gratefully thank Dr. D.T.T. Jayasinghe and Dr. R.D.K. Medonza, Postgraduate trainee in Histopathology, Faculty of Medicine, University of Kelaniya, for their assistance in photomicrography.
\n',keywords:"ulcerative colitis, histological features, diagnostic pitfalls, histological disease activity, dysplasia in ulcerative colitis",chapterPDFUrl:"https://cdn.intechopen.com/pdfs/80116.pdf",chapterXML:"https://mts.intechopen.com/source/xml/80116.xml",downloadPdfUrl:"/chapter/pdf-download/80116",previewPdfUrl:"/chapter/pdf-preview/80116",totalDownloads:81,totalViews:0,totalCrossrefCites:0,dateSubmitted:"November 27th 2021",dateReviewed:"December 17th 2021",datePrePublished:"January 19th 2022",datePublished:null,dateFinished:"January 19th 2022",readingETA:"0",abstract:"Pathologists have an essential and wide role in the management of patients with ulcerative colitis (UC) which is a chronic inflammatory disorder of the bowel with remissions and relapses. The initial diagnosis of UC itself is challenging as the histological features vary widely with the clinical phase of the disease. Differentiating UC from other types of acute and chronic colitis, especially Crohn’s disease is crucial in the management. Understanding the characteristic morphological features of UC as well as unusual morphological features of the disease are important in this task. The histological disease activity has now been identified as important in therapeutic decisions. There are several histological activity indices in UC and currently, the Nancy histological index has been recommended to be used in daily clinical practice. Identifying dysplasia associated with UC and its grading is a challenging task for the pathologist and it is a crucial step in the surveillance and management of this chronic disease.",reviewType:"peer-reviewed",bibtexUrl:"/chapter/bibtex/80116",risUrl:"/chapter/ris/80116",signatures:"Gayana Mahendra and Janaki Hewavisenthi",book:{id:"11268",type:"book",title:"Ulcerative Colitis",subtitle:null,fullTitle:"Ulcerative Colitis",slug:null,publishedDate:null,bookSignature:"Dr. Partha Pal",coverURL:"https://cdn.intechopen.com/books/images_new/11268.jpg",licenceType:"CC BY 3.0",editedByType:null,isbn:"978-1-80355-295-8",printIsbn:"978-1-80355-294-1",pdfIsbn:"978-1-80355-296-5",isAvailableForWebshopOrdering:!0,editors:[{id:"345620",title:"Dr.",name:"Partha",middleName:null,surname:"Pal",slug:"partha-pal",fullName:"Partha Pal"}],productType:{id:"1",title:"Edited Volume",chapterContentType:"chapter",authoredCaption:"Edited by"}},authors:null,sections:[{id:"sec_1",title:"1. Introduction",level:"1"},{id:"sec_2",title:"2. Macroscopic appearance of ulcerative colitis",level:"1"},{id:"sec_3",title:"3. Microscopy appearance of ulcerative colitis",level:"1"},{id:"sec_3_2",title:"3.1 Main histological characteristics of ulcerative colitis",level:"2"},{id:"sec_3_3",title:"3.1.1 Crypt architectural abnormalities",level:"3"},{id:"sec_4_3",title:"3.1.2 Cellularity in the lamina propria",level:"3"},{id:"sec_5_3",title:"3.1.3 Neutrophil granulocytes in the lamina propria",level:"3"},{id:"sec_6_3",title:"Table 1.",level:"3"},{id:"sec_8_2",title:"3.2 Typical histological features of ulcerative colitis",level:"2"},{id:"sec_8_3",title:"3.2.1 Active ulcerative colitis",level:"3"},{id:"sec_9_3",title:"3.2.2 Resolving ulcerative colitis",level:"3"},{id:"sec_10_3",title:"3.2.3 Quiescent ulcerative colitis",level:"3"},{id:"sec_13",title:"4. Unusual morphological patterns of ulcerative colitis",level:"1"},{id:"sec_13_2",title:"4.1 Morphological features of treated ulcerative colitis",level:"2"},{id:"sec_14_2",title:"4.2 Appendiceal ‘skip’ lesions and the ‘caecal patch’ in ulcerative colitis",level:"2"},{id:"sec_15_2",title:"4.3 Involvement of the ileum",level:"2"},{id:"sec_16_2",title:"4.4 Rectal sparing",level:"2"},{id:"sec_17_2",title:"4.5 Ulcerative colitis in pediatric populations",level:"2"},{id:"sec_18_2",title:"4.6 Fulminant ulcerative colitis",level:"2"},{id:"sec_20",title:"5. Histological differential diagnosis of ulcerative colitis and diagnostic pitfalls",level:"1"},{id:"sec_20_2",title:"5.1 Infective colitis",level:"2"},{id:"sec_21_2",title:"5.2 Chronic ischaemic colitis",level:"2"},{id:"sec_22_2",title:"5.3 Diverticular disease-associated colitis (DAC)",level:"2"},{id:"sec_23_2",title:"5.4 Drug-induced colitis",level:"2"},{id:"sec_24_2",title:"5.5 Diversion colitis",level:"2"},{id:"sec_25_2",title:"5.6 Crohn disease (CD)",level:"2"},{id:"sec_25_3",title:"5.6.1 Granulomas in UC",level:"3"},{id:"sec_26_3",title:"5.6.2 Transmural inflammation in UC",level:"3"},{id:"sec_27_3",title:"5.6.3 CD with UC-like features",level:"3"},{id:"sec_30",title:"6. Indeterminate colitis (IC) and inflammatory bowel disease unclassified (IBDU)",level:"1"},{id:"sec_31",title:"7. Measuring the disease activity in ulcerative colitis",level:"1"},{id:"sec_32",title:"8. Dysplasia and malignancy in ulcerative colitis",level:"1"},{id:"sec_32_2",title:"8.1 Colorectal dysplasia in ulcerative colitis",level:"2"},{id:"sec_33_2",title:"8.2 Some issues in histopathological reporting in UC-associated dysplasia",level:"2"},{id:"sec_34_2",title:"8.3 The demise of the term ‘DALM’ in ulcerative colitis",level:"2"},{id:"sec_35_2",title:"8.4 Carcinoma in ulcerative colitis",level:"2"},{id:"sec_37",title:"9. Conclusions",level:"1"},{id:"sec_38",title:"Acknowledgments",level:"1"}],chapterReferences:[{id:"B1",body:'Jain D, Warren BF, Riddell RH. Inflammatory disorders of the large intestine. In: Shepherd NA, Warren BF, Williams GT, Greenson JK, Lauwers GY, Novelli MR, editors. Morson and Dawson′s Gastrointestinal Pathology. 5th ed. Chichester, West Sussex, UK: Wiley-Blackwell; 2013. pp. 552-635'},{id:"B2",body:'Magro F, Doherty G, Peyrin-Biroulet L, Svrcek M, Borralho P, Walsh A, et al. 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DOI: 10.1111/j.1365-2559.2005.02248.x'},{id:"B23",body:'Mosli MH, Parker CE, Nelson SA, Baker KA, Macdonald JK, Zou GY, et al. Histologic scoring indices for evaluation of disease activity in ulcerative colitis. Cochrane Database of Systematic Reviews. 2017;2017:1-34. DOI: 10.1002/14651858.CD011256.pub2'},{id:"B24",body:'Geboes K, Riddell R, Öst A, Jensfelt B, Persson T, Löfberg R. A reproducible grading scale for histological assessment of inflammation in ulcerative colitis. Gut. 2000;47:404-409. DOI: 10.1136/gut.47.3.404'},{id:"B25",body:'Magro F, Alves C, Lopes J, Lopes S, Tavares de Sousa H, Cotter J, et al. Histologic features of colon biopsies (Geboes score) associated with progression of ulcerative colitis for the first 36 months after biopsy. Clinical Gastroenterology and Hepatology. 2021;19:2567-2576. DOI: 10.1016/j.cgh.2020.09.017'},{id:"B26",body:'Marchal-Bressenot A, Salleron J, Boulagnon-Rombi C, Bastien C, Cahn V, Cadiot G, et al. Development and validation of the Nancy histological index for UC. Gut. 2017;66:43-49. DOI: 10.1136/gutjnl-2015-310187'},{id:"B27",body:'Mosli MH, Feagan BG, Zou G, Sandborn WJ, D’Haens G, Khanna R, et al. Development and validation of a histological index for UC. Gut. 2017;66:50-58. DOI: 10.1136/gutjnl-2015-310393'},{id:"B28",body:'MedMadonsa RDK, Mahendra BAGG, Silva H, Jayathunga DNU, Fernando J, Ediriweera DS, et al. Manchester Pathology 2021. 13th Joint Meeting of the British Division of the International Academy of Pathology and the Pathological Society of Great Britain &; Ireland, 6-8 July 2021. The Journal of Pathology. 2021;255:S1-S54. DOI: 10.1002/path.5793'},{id:"B29",body:'Odze RD, Harpaz N. Inflammtory bowel disease associated colorectal dysplasia. In: WHO Classification of Tumours Editorial Board, editor. WHO Classification of Tumours: Digestive System Tumours. 5th ed. Vol. 1. Lyon, France: IARC Publications; 2019. pp. 174-176'},{id:"B30",body:'Riddell RH, Goldman H, Ransohoff DF, Appelman HD, Fenoglio CM, Haggitt RC, et al. Dysplasia in inflammatory bowel disease: Standardized classification with provisional clinical applications. Human Pathology. 1983;14:931-969. DOI: 10.1016/S0046-8177(83)80175-0'},{id:"B31",body:'Schlemper RJ, Riddell RH, Kato Y, Borchard F, Cooper HS, Dawsey SM, et al. The vienna classification of gastrointestinal epithelial neoplasia. Gut. 2000;47:251-255. DOI: 10.1136/gut.47.2.251'},{id:"B32",body:'Lichtiger S, Present DH, Kornbluth A, Gelernt I, Bauer J, Galler G, et al. Cyclosporine in severe ulcerative colitis refractory to steroid therapy. New England Journal of Medicine. 1994;330:1841-1845. DOI: 10.1056/nejm199406303302601'},{id:"B33",body:'Laine L, Kaltenbach T, Barkun A, McQuaid KR, Subramanian V, Soetikno R. SCENIC international consensus statement on surveillance and management of dysplasia in inflammatory bowel disease. Gastroenterology. 2015;148:639-651. DOI: 10.1053/j.gastro.2015.01.031'}],footnotes:[],contributors:[{corresp:"yes",contributorFullName:"Gayana Mahendra",address:"baggmahendra@yahoo.co.uk",affiliation:'
Faculty of Medicine, Department of Pathology, University of Kelaniya, Sri Lanka
Faculty of Medicine, Department of Pathology, University of Kelaniya, Sri Lanka
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Several approaches for modelling horizontal well performance have been studied and reported in the literature. Analytical approach is the easiest with large inaccuracy in the prediction of the horizontal well performance because of inability to apply it in reservoir-wellbore coupling equation. Numerical approach is more reliable for field application than analytical approach. However, it involves iterative nature that requires longer computational times. Semi-analytical approach is simpler and sufficiently exact for field applications if the governing fundamental flow equation is accurately modelled. This study presents a new semi-analytical model for predicting horizontal and multilateral well performance, which includes friction, acceleration and accumulation induced pressure drop along horizontal well length into the governing fundamental flow equations. The outcomes of the proposed model have been validated by field data gotten from gauge rate of 5660stb/d at steady-state condition. 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Various textile production methods used for the formation of textile preforms are explained. Composite fabrication methods are introduced. Engineering properties of textile composites are reviewed with regard to specific application areas. 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\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\t
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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\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\t
\r\n
\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\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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Saxena",profilePictureURL:"https://s3.us-east-1.amazonaws.com/intech-files/0030O00002bRET3QAO/Profile_Picture_2022-05-10T10:10:26.jpeg",institutionString:"King George's Medical University",institution:{name:"King George's Medical University",institutionURL:null,country:{name:"India"}}}],equalEditorOne:null,equalEditorTwo:null,equalEditorThree:null}]},subseriesFiltersForPublishedBooks:[{group:"subseries",caption:"Bacterial Infectious Diseases",value:3,count:2},{group:"subseries",caption:"Parasitic Infectious Diseases",value:5,count:4},{group:"subseries",caption:"Viral Infectious Diseases",value:6,count:7}],publicationYearFilters:[{group:"publicationYear",caption:"2022",value:2022,count:2},{group:"publicationYear",caption:"2021",value:2021,count:4},{group:"publicationYear",caption:"2020",value:2020,count:3},{group:"publicationYear",caption:"2019",value:2019,count:3},{group:"publicationYear",caption:"2018",value:2018,count:1}],authors:{paginationCount:229,paginationItems:[{id:"318170",title:"Dr.",name:"Aneesa",middleName:null,surname:"Moolla",slug:"aneesa-moolla",fullName:"Aneesa Moolla",position:null,profilePictureURL:"https://mts.intechopen.com/storage/users/318170/images/system/318170.png",biography:"Dr. Aneesa Moolla has extensive experience in the diverse fields of health care having previously worked in dental private practice, at the Red Cross Flying Doctors association, and in healthcare corporate settings. She is now a lecturer at the University of Witwatersrand, South Africa, and a principal researcher at the Health Economics and Epidemiology Research Office (HE2RO), South Africa. Dr. Moolla holds a Ph.D. in Psychology with her research being focused on mental health and resilience. In her professional work capacity, her research has further expanded into the fields of early childhood development, mental health, the HIV and TB care cascades, as well as COVID. She is also a UNESCO-trained International Bioethics Facilitator.",institutionString:"University of the Witwatersrand",institution:{name:"University of the Witwatersrand",country:{name:"South Africa"}}},{id:"419588",title:"Ph.D.",name:"Sergio",middleName:"Alexandre",surname:"Gehrke",slug:"sergio-gehrke",fullName:"Sergio Gehrke",position:null,profilePictureURL:"https://s3.us-east-1.amazonaws.com/intech-files/0033Y000038WgMKQA0/Profile_Picture_2022-06-02T11:44:20.jpg",biography:"Dr. Sergio Alexandre Gehrke is a doctorate holder in two fields. The first is a Ph.D. in Cellular and Molecular Biology from the Pontificia Catholic University, Porto Alegre, Brazil, in 2010 and the other is an International Ph.D. in Bioengineering from the Universidad Miguel Hernandez, Elche/Alicante, Spain, obtained in 2020. In 2018, he completed a postdoctoral fellowship in Materials Engineering in the NUCLEMAT of the Pontificia Catholic University, Porto Alegre, Brazil. He is currently the Director of the Postgraduate Program in Implantology of the Bioface/UCAM/PgO (Montevideo, Uruguay), Director of the Cathedra of Biotechnology of the Catholic University of Murcia (Murcia, Spain), an Extraordinary Full Professor of the Catholic University of Murcia (Murcia, Spain) as well as the Director of the private center of research Biotecnos – Technology and Science (Montevideo, Uruguay). Applied biomaterials, cellular and molecular biology, and dental implants are among his research interests. He has published several original papers in renowned journals. In addition, he is also a Collaborating Professor in several Postgraduate programs at different universities all over the world.",institutionString:null,institution:{name:"Universidad Católica San Antonio de Murcia",country:{name:"Spain"}}},{id:"342152",title:"Dr.",name:"Santo",middleName:null,surname:"Grace Umesh",slug:"santo-grace-umesh",fullName:"Santo Grace Umesh",position:null,profilePictureURL:"https://mts.intechopen.com/storage/users/342152/images/16311_n.jpg",biography:null,institutionString:null,institution:{name:"SRM Dental College",country:{name:"India"}}},{id:"333647",title:"Dr.",name:"Shreya",middleName:null,surname:"Kishore",slug:"shreya-kishore",fullName:"Shreya Kishore",position:null,profilePictureURL:"https://mts.intechopen.com/storage/users/333647/images/14701_n.jpg",biography:"Dr. Shreya Kishore completed her Bachelor in Dental Surgery in Chettinad Dental College and Research Institute, Chennai, and her Master of Dental Surgery (Orthodontics) in Saveetha Dental College, Chennai. She is also Invisalign certified. She’s working as a Senior Lecturer in the Department of Orthodontics, SRM Dental College since November 2019. She is actively involved in teaching orthodontics to the undergraduates and the postgraduates. Her clinical research topics include new orthodontic brackets, fixed appliances and TADs. She’s published 4 articles in well renowned indexed journals and has a published patency of her own. Her private practice is currently limited to orthodontics and works as a consultant in various clinics.",institutionString:null,institution:{name:"SRM Dental College",country:{name:"India"}}},{id:"323731",title:"Prof.",name:"Deepak M.",middleName:"Macchindra",surname:"Vikhe",slug:"deepak-m.-vikhe",fullName:"Deepak M. Vikhe",position:null,profilePictureURL:"https://mts.intechopen.com/storage/users/323731/images/13613_n.jpg",biography:"Dr Deepak M.Vikhe .\n\n\t\n\tDr Deepak M.Vikhe , completed his Masters & PhD in Prosthodontics from Rural Dental College, Loni securing third rank in the Pravara Institute of Medical Sciences Deemed University. He was awarded Dr.G.C.DAS Memorial Award for Research on Implants at 39th IPS conference Dubai (U A E).He has two patents under his name. He has received Dr.Saraswati medal award for best research for implant study in 2017.He has received Fully funded scholarship to Spain ,university of Santiago de Compostela. He has completed fellowship in Implantlogy from Noble Biocare. \nHe has attended various conferences and CDE programmes and has national publications to his credit. His field of interest is in Implant supported prosthesis. Presently he is working as a associate professor in the Dept of Prosthodontics, Rural Dental College, Loni and maintains a successful private practice specialising in Implantology at Rahata.\n\nEmail: drdeepak_mvikhe@yahoo.com..................",institutionString:null,institution:{name:"Pravara Institute of Medical Sciences",country:{name:"India"}}},{id:"204110",title:"Dr.",name:"Ahmed A.",middleName:null,surname:"Madfa",slug:"ahmed-a.-madfa",fullName:"Ahmed A. Madfa",position:null,profilePictureURL:"https://mts.intechopen.com/storage/users/204110/images/system/204110.jpg",biography:"Dr. Madfa is currently Associate Professor of Endodontics at Thamar University and a visiting lecturer at Sana'a University and University of Sciences and Technology. He has more than 6 years of experience in teaching. His research interests include root canal morphology, functionally graded concept, dental biomaterials, epidemiology and dental education, biomimetic restoration, finite element analysis and endodontic regeneration. Dr. Madfa has numerous international publications, full articles, two patents, a book and a book chapter. Furthermore, he won 14 international scientific awards. Furthermore, he is involved in many academic activities ranging from editorial board member, reviewer for many international journals and postgraduate students' supervisor. Besides, I deliver many courses and training workshops at various scientific events. Dr. Madfa also regularly attends international conferences and holds administrative positions (Deputy Dean of the Faculty for Students’ & Academic Affairs and Deputy Head of Research Unit).",institutionString:"Thamar University",institution:null},{id:"210472",title:"Dr.",name:"Nermin",middleName:"Mohammed Ahmed",surname:"Yussif",slug:"nermin-yussif",fullName:"Nermin Yussif",position:null,profilePictureURL:"https://mts.intechopen.com/storage/users/210472/images/system/210472.jpg",biography:"Dr. Nermin Mohammed Ahmed Yussif is working at the Faculty of dentistry, University for October university for modern sciences and arts (MSA). Her areas of expertise include: periodontology, dental laserology, oral implantology, periodontal plastic surgeries, oral mesotherapy, nutrition, dental pharmacology. She is an editor and reviewer in numerous international journals.",institutionString:"MSA University",institution:null},{id:"204606",title:"Dr.",name:"Serdar",middleName:null,surname:"Gözler",slug:"serdar-gozler",fullName:"Serdar Gözler",position:null,profilePictureURL:"https://mts.intechopen.com/storage/users/204606/images/system/204606.jpeg",biography:"Dr. Serdar Gözler has completed his undergraduate studies at the Marmara University Faculty of Dentistry in 1978, followed by an assistantship in the Prosthesis Department of Dicle University Faculty of Dentistry. Starting his PhD work on non-resilient overdentures with Assoc. Prof. Hüsnü Yavuzyılmaz, he continued his studies with Prof. Dr. Gürbüz Öztürk of Istanbul University Faculty of Dentistry Department of Prosthodontics, this time on Gnatology. He attended training programs on occlusion, neurology, neurophysiology, EMG, radiology and biostatistics. In 1982, he presented his PhD thesis \\Gerber and Lauritzen Occlusion Analysis Techniques: Diagnosis Values,\\ at Istanbul University School of Dentistry, Department of Prosthodontics. As he was also working with Prof. Senih Çalıkkocaoğlu on The Physiology of Chewing at the same time, Gözler has written a chapter in Çalıkkocaoğlu\\'s book \\Complete Prostheses\\ entitled \\The Place of Neuromuscular Mechanism in Prosthetic Dentistry.\\ The book was published five times since by the Istanbul University Publications. Having presented in various conferences about occlusion analysis until 1998, Dr. Gözler has also decided to use the T-Scan II occlusion analysis method. Having been personally trained by Dr. Robert Kerstein on this method, Dr. Gözler has been lecturing on the T-Scan Occlusion Analysis Method in conferences both in Turkey and abroad. Dr. Gözler has various articles and presentations on Digital Occlusion Analysis methods. He is now Head of the TMD Clinic at Prosthodontic Department of Faculty of Dentistry , Istanbul Aydın University , Turkey.",institutionString:"Istanbul Aydin University",institution:{name:"Istanbul Aydın University",country:{name:"Turkey"}}},{id:"256417",title:"Associate Prof.",name:"Sanaz",middleName:null,surname:"Sadry",slug:"sanaz-sadry",fullName:"Sanaz Sadry",position:null,profilePictureURL:"https://mts.intechopen.com/storage/users/256417/images/8106_n.jpg",biography:null,institutionString:null,institution:{name:"Istanbul Aydın University",country:{name:"Turkey"}}},{id:"240870",title:"Ph.D.",name:"Alaa Eddin Omar",middleName:null,surname:"Al Ostwani",slug:"alaa-eddin-omar-al-ostwani",fullName:"Alaa Eddin Omar Al Ostwani",position:null,profilePictureURL:"https://mts.intechopen.com/storage/users/240870/images/system/240870.jpeg",biography:"Dr. Al Ostwani Alaa Eddin Omar received his Master in dentistry from Damascus University in 2010, and his Ph.D. in Pediatric Dentistry from Damascus University in 2014. Dr. Al Ostwani is an assistant professor and faculty member at IUST University since 2014. \nDuring his academic experience, he has received several awards including the scientific research award from the Union of Arab Universities, the Syrian gold medal and the international gold medal for invention and creativity. Dr. Al Ostwani is a Member of the International Association of Dental Traumatology and the Syrian Society for Research and Preventive Dentistry since 2017. He is also a Member of the Reviewer Board of International Journal of Dental Medicine (IJDM), and the Indian Journal of Conservative and Endodontics since 2016.",institutionString:"International University for Science and Technology.",institution:{name:"Islamic University of Science and Technology",country:{name:"India"}}},{id:"42847",title:"Dr.",name:"Belma",middleName:null,surname:"Işik Aslan",slug:"belma-isik-aslan",fullName:"Belma Işik Aslan",position:null,profilePictureURL:"https://mts.intechopen.com/storage/users/42847/images/system/42847.jpg",biography:"Dr. Belma IşIk Aslan was born in 1976 in Ankara-TURKEY. After graduating from TED Ankara College in 1994, she attended to Gazi University, Faculty of Dentistry in Ankara. She completed her PhD in orthodontic education at Gazi University between 1999-2005. Dr. Işık Aslan stayed at the Providence Hospital Craniofacial Institude and Reconstructive Surgery in Michigan, USA for three months as an observer. She worked as a specialist doctor at Gazi University, Dentistry Faculty, Department of Orthodontics between 2005-2014. She was appointed as associate professor in January, 2014 and as professor in 2021. Dr. Işık Aslan still works as an instructor at the same faculty. She has published a total of 35 articles, 10 book chapters, 39 conference proceedings both internationally and nationally. Also she was the academic editor of the international book 'Current Advances in Orthodontics'. She is a member of the Turkish Orthodontic Society and Turkish Cleft Lip and Palate Society. She is married and has 2 children. Her knowledge of English is at an advanced level.",institutionString:"Gazi University Dentistry Faculty Department of Orthodontics",institution:null},{id:"202198",title:"Dr.",name:"Buket",middleName:null,surname:"Aybar",slug:"buket-aybar",fullName:"Buket Aybar",position:null,profilePictureURL:"https://mts.intechopen.com/storage/users/202198/images/6955_n.jpg",biography:"Buket Aybar, DDS, PhD, was born in 1971. She graduated from Istanbul University, Faculty of Dentistry, in 1992 and completed her PhD degree on Oral and Maxillofacial Surgery in Istanbul University in 1997.\r\nDr. Aybar is currently a full-time professor in Istanbul University, Faculty of Dentistry Department of Oral and Maxillofacial Surgery. She has teaching responsibilities in graduate and postgraduate programs. Her clinical practice includes mainly dentoalveolar surgery.\r\nHer topics of interest are biomaterials science and cell culture studies. She has many articles in international and national scientific journals and chapters in books; she also has participated in several scientific projects supported by Istanbul University Research fund.",institutionString:null,institution:{name:"Marmara University",country:{name:"Turkey"}}},{id:"178412",title:"Associate Prof.",name:"Guhan",middleName:null,surname:"Dergin",slug:"guhan-dergin",fullName:"Guhan Dergin",position:null,profilePictureURL:"https://mts.intechopen.com/storage/users/178412/images/6954_n.jpg",biography:"Assoc. Prof. Dr. Gühan Dergin was born in 1973 in Izmit. He graduated from Marmara University Faculty of Dentistry in 1999. He completed his specialty of OMFS surgery in Marmara University Faculty of Dentistry and obtained his PhD degree in 2006. In 2005, he was invited as a visiting doctor in the Oral and Maxillofacial Surgery Department of the University of North Carolina, USA, where he went on a scholarship. Dr. Dergin still continues his academic career as an associate professor in Marmara University Faculty of Dentistry. He has many articles in international and national scientific journals and chapters in books.",institutionString:null,institution:{name:"Marmara University",country:{name:"Turkey"}}},{id:"178414",title:"Prof.",name:"Yusuf",middleName:null,surname:"Emes",slug:"yusuf-emes",fullName:"Yusuf Emes",position:null,profilePictureURL:"https://mts.intechopen.com/storage/users/178414/images/6953_n.jpg",biography:"Born in Istanbul in 1974, Dr. Emes graduated from Istanbul University Faculty of Dentistry in 1997 and completed his PhD degree in Istanbul University faculty of Dentistry Department of Oral and Maxillofacial Surgery in 2005. He has papers published in international and national scientific journals, including research articles on implantology, oroantral fistulas, odontogenic cysts, and temporomandibular disorders. Dr. Emes is currently working as a full-time academic staff in Istanbul University faculty of Dentistry Department of Oral and Maxillofacial Surgery.",institutionString:null,institution:{name:"Istanbul University",country:{name:"Turkey"}}},{id:"192229",title:"Ph.D.",name:"Ana Luiza",middleName:null,surname:"De Carvalho Felippini",slug:"ana-luiza-de-carvalho-felippini",fullName:"Ana Luiza De Carvalho Felippini",position:null,profilePictureURL:"https://mts.intechopen.com/storage/users/192229/images/system/192229.jpg",biography:null,institutionString:"University of São Paulo",institution:{name:"University of Sao Paulo",country:{name:"Brazil"}}},{id:"256851",title:"Prof.",name:"Ayşe",middleName:null,surname:"Gülşen",slug:"ayse-gulsen",fullName:"Ayşe Gülşen",position:null,profilePictureURL:"https://mts.intechopen.com/storage/users/256851/images/9696_n.jpg",biography:"Dr. Ayşe Gülşen graduated in 1990 from Faculty of Dentistry, University of Ankara and did a postgraduate program at University of Gazi. \nShe worked as an observer and research assistant in Craniofacial Surgery Departments in New York, Providence Hospital in Michigan and Chang Gung Memorial Hospital in Taiwan. \nShe works as Craniofacial Orthodontist in Department of Aesthetic, Plastic and Reconstructive Surgery, Faculty of Medicine, University of Gazi, Ankara Turkey since 2004.",institutionString:"Orthodontist, Assoc Prof in the Department of Aesthetic, Plastic and Reconstructive Surgery, Faculty of Medicine, University of Gazi",institution:null},{id:"255366",title:"Prof.",name:"Tosun",middleName:null,surname:"Tosun",slug:"tosun-tosun",fullName:"Tosun Tosun",position:null,profilePictureURL:"https://mts.intechopen.com/storage/users/255366/images/7347_n.jpg",biography:"Graduated at the Faculty of Dentistry, University of Istanbul, Turkey in 1989;\nVisitor Assistant at the University of Padua, Italy and Branemark Osseointegration Center of Treviso, Italy between 1993-94;\nPhD thesis on oral implantology in University of Istanbul and was awarded the academic title “Dr.med.dent.”, 1997;\nHe was awarded the academic title “Doç.Dr.” (Associated Professor) in 2003;\nProficiency in Botulinum Toxin Applications, Reading-UK in 2009;\nMastership, RWTH Certificate in Laser Therapy in Dentistry, AALZ-Aachen University, Germany 2009-11;\nMaster of Science (MSc) in Laser Dentistry, University of Genoa, Italy 2013-14.\n\nDr.Tosun worked as Research Assistant in the Department of Oral Implantology, Faculty of Dentistry, University of Istanbul between 1990-2002. \nHe worked part-time as Consultant surgeon in Harvard Medical International Hospitals and John Hopkins Medicine, Istanbul between years 2007-09.\u2028He was contract Professor in the Department of Surgical and Diagnostic Sciences (DI.S.C.), Medical School, University of Genova, Italy between years 2011-16. \nSince 2015 he is visiting Professor at Medical School, University of Plovdiv, Bulgaria. \nCurrently he is Associated Prof.Dr. at the Dental School, Oral Surgery Dept., Istanbul Aydin University and since 2003 he works in his own private clinic in Istanbul, Turkey.\u2028\nDr.Tosun is reviewer in journal ‘Laser in Medical Sciences’, reviewer in journal ‘Folia Medica\\', a Fellow of the International Team for Implantology, Clinical Lecturer of DGZI German Association of Oral Implantology, Expert Lecturer of Laser&Health Academy, Country Representative of World Federation for Laser Dentistry, member of European Federation of Periodontology, member of Academy of Laser Dentistry. Dr.Tosun presents papers in international and national congresses and has scientific publications in international and national journals. He speaks english, spanish, italian and french.",institutionString:null,institution:{name:"Istanbul Aydın University",country:{name:"Turkey"}}},{id:"260116",title:"Dr.",name:"Mehmet",middleName:null,surname:"Yaltirik",slug:"mehmet-yaltirik",fullName:"Mehmet Yaltirik",position:null,profilePictureURL:"https://mts.intechopen.com/storage/users/260116/images/7413_n.jpg",biography:"Birth Date 25.09.1965\r\nBirth Place Adana- Turkey\r\nSex Male\r\nMarrial Status Bachelor\r\nDriving License Acquired\r\nMother Tongue Turkish\r\n\r\nAddress:\r\nWork:University of Istanbul,Faculty of Dentistry, Department of Oral Surgery and Oral Medicine 34093 Capa,Istanbul- TURKIYE",institutionString:null,institution:{name:"Istanbul University",country:{name:"Turkey"}}},{id:"171887",title:"Prof.",name:"Zühre",middleName:null,surname:"Akarslan",slug:"zuhre-akarslan",fullName:"Zühre Akarslan",position:null,profilePictureURL:"https://mts.intechopen.com/storage/users/171887/images/system/171887.jpg",biography:"Zühre Akarslan was born in 1977 in Cyprus. She graduated from Gazi University Faculty of Dentistry, Ankara, Turkey in 2000. \r\nLater she received her Ph.D. degree from the Oral Diagnosis and Radiology Department; which was recently renamed as Oral and Dentomaxillofacial Radiology, from the same university. \r\nShe is working as a full-time Associate Professor and is a lecturer and an academic researcher. \r\nHer expertise areas are dental caries, cancer, dental fear and anxiety, gag reflex in dentistry, oral medicine, and dentomaxillofacial radiology.",institutionString:"Gazi University",institution:{name:"Gazi University",country:{name:"Turkey"}}},{id:"272237",title:"Dr.",name:"Pinar",middleName:"Kiymet",surname:"Karataban",slug:"pinar-karataban",fullName:"Pinar Karataban",position:null,profilePictureURL:"https://mts.intechopen.com/storage/users/272237/images/8911_n.png",biography:"Assist.Prof.Dr.Pınar Kıymet Karataban, DDS PhD \n\nDr.Pınar Kıymet Karataban was born in Istanbul in 1975. After her graduation from Marmara University Faculty of Dentistry in 1998 she started her PhD in Paediatric Dentistry focused on children with special needs; mainly children with Cerebral Palsy. She finished her pHD thesis entitled \\'Investigation of occlusion via cast analysis and evaluation of dental caries prevalance, periodontal status and muscle dysfunctions in children with cerebral palsy” in 2008. She got her Assist. Proffessor degree in Istanbul Aydın University Paediatric Dentistry Department in 2015-2018. ın 2019 she started her new career in Bahcesehir University, Istanbul as Head of Department of Pediatric Dentistry. In 2020 she was accepted to BAU International University, Batumi as Professor of Pediatric Dentistry. She’s a lecturer in the same university meanwhile working part-time in private practice in Ege Dental Studio (https://www.egedisklinigi.com/) a multidisciplinary dental clinic in Istanbul. Her main interests are paleodontology, ancient and contemporary dentistry, oral microbiology, cerebral palsy and special care dentistry. She has national and international publications, scientific reports and is a member of IAPO (International Association for Paleodontology), IADH (International Association of Disability and Oral Health) and EAPD (European Association of Pediatric Dentistry).",institutionString:null,institution:null},{id:"172009",title:"Dr.",name:"Fatma Deniz",middleName:null,surname:"Uzuner",slug:"fatma-deniz-uzuner",fullName:"Fatma Deniz Uzuner",position:null,profilePictureURL:"https://mts.intechopen.com/storage/users/172009/images/7122_n.jpg",biography:"Dr. Deniz Uzuner was born in 1969 in Kocaeli-TURKEY. After graduating from TED Ankara College in 1986, she attended the Hacettepe University, Faculty of Dentistry in Ankara. \nIn 1993 she attended the Gazi University, Faculty of Dentistry, Department of Orthodontics for her PhD education. After finishing the PhD education, she worked as orthodontist in Ankara Dental Hospital under the Turkish Government, Ministry of Health and in a special Orthodontic Clinic till 2011. Between 2011 and 2016, Dr. Deniz Uzuner worked as a specialist in the Department of Orthodontics, Faculty of Dentistry, Gazi University in Ankara/Turkey. In 2016, she was appointed associate professor. Dr. Deniz Uzuner has authored 23 Journal Papers, 3 Book Chapters and has had 39 oral/poster presentations. She is a member of the Turkish Orthodontic Society. 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