Genetic approaches for improved seed yield in cereal crops.
\\n\\n
IntechOpen Book Series will also publish a program of research-driven Thematic Edited Volumes that focus on specific areas and allow for a more in-depth overview of a particular subject.
\\n\\nIntechOpen Book Series will be launching regularly to offer our authors and editors exciting opportunities to publish their research Open Access. We will begin by relaunching some of our existing Book Series in this innovative book format, and will expand in 2022 into rapidly growing research fields that are driving and advancing society.
\\n\\nLaunching 2021
\\n\\nArtificial Intelligence, ISSN 2633-1403
\\n\\nVeterinary Medicine and Science, ISSN 2632-0517
\\n\\nBiochemistry, ISSN 2632-0983
\\n\\nBiomedical Engineering, ISSN 2631-5343
\\n\\nInfectious Diseases, ISSN 2631-6188
\\n\\nPhysiology (Coming Soon)
\\n\\nDentistry (Coming Soon)
\\n\\nWe invite you to explore our IntechOpen Book Series, find the right publishing program for you and reach your desired audience in record time.
\\n\\nNote: Edited in October 2021
\\n"}]',published:!0,mainMedia:{caption:"",originalUrl:"/media/original/132"}},components:[{type:"htmlEditorComponent",content:'With the desire to make book publishing more relevant for the digital age and offer innovative Open Access publishing options, we are thrilled to announce the launch of our new publishing format: IntechOpen Book Series.
\n\nDesigned to cover fast-moving research fields in rapidly expanding areas, our Book Series feature a Topic structure allowing us to present the most relevant sub-disciplines. Book Series are headed by Series Editors, and a team of Topic Editors supported by international Editorial Board members. Topics are always open for submissions, with an Annual Volume published each calendar year.
\n\nAfter a robust peer-review process, accepted works are published quickly, thanks to Online First, ensuring research is made available to the scientific community without delay.
\n\nOur innovative Book Series format brings you:
\n\nIntechOpen Book Series will also publish a program of research-driven Thematic Edited Volumes that focus on specific areas and allow for a more in-depth overview of a particular subject.
\n\nIntechOpen Book Series will be launching regularly to offer our authors and editors exciting opportunities to publish their research Open Access. We will begin by relaunching some of our existing Book Series in this innovative book format, and will expand in 2022 into rapidly growing research fields that are driving and advancing society.
\n\nLaunching 2021
\n\nArtificial Intelligence, ISSN 2633-1403
\n\nVeterinary Medicine and Science, ISSN 2632-0517
\n\nBiochemistry, ISSN 2632-0983
\n\nBiomedical Engineering, ISSN 2631-5343
\n\nInfectious Diseases, ISSN 2631-6188
\n\nPhysiology (Coming Soon)
\n\nDentistry (Coming Soon)
\n\nWe invite you to explore our IntechOpen Book Series, find the right publishing program for you and reach your desired audience in record time.
\n\nNote: Edited in October 2021
\n'}],latestNews:[{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"},{slug:"introducing-intechopen-book-series-a-new-publishing-format-for-oa-books-20210915",title:"Introducing IntechOpen Book Series - A New Publishing Format for OA Books"}]},book:{item:{type:"book",id:"6365",leadTitle:null,fullTitle:"Molecular Docking",title:"Molecular Docking",subtitle:null,reviewType:"peer-reviewed",abstract:"Molecular docking has always been and will be on the forefront of developments in the eminent field of drug design and medicinal chemistry. At the early days, drug discovery was based on blackboard drawings and expert intuition. However, as times move on, the amount of available information and overall knowledge base that needs to be analyzed cannot be processed manually. This, coupled by the rapid growth in computational infrastructure and processing power, has allowed for the efficient use of molecular docking tools and algorithms to be considered in the greater field of drug discovery. In the postgenomic era, molecular docking has become the key player for the screening of hundreds of thousands of compounds against a repertoire of pharmacological targets.",isbn:"978-1-78923-355-1",printIsbn:"978-1-78923-354-4",pdfIsbn:"978-1-83881-432-8",doi:"10.5772/intechopen.69830",price:119,priceEur:129,priceUsd:155,slug:"molecular-docking",numberOfPages:188,isOpenForSubmission:!1,isInWos:null,isInBkci:!1,hash:"ca23ec77de0bb7a434608335e1d6a963",bookSignature:"Dimitrios P. Vlachakis",publishedDate:"July 11th 2018",coverURL:"https://cdn.intechopen.com/books/images_new/6365.jpg",numberOfDownloads:14789,numberOfWosCitations:26,numberOfCrossrefCitations:23,numberOfCrossrefCitationsByBook:1,numberOfDimensionsCitations:37,numberOfDimensionsCitationsByBook:1,hasAltmetrics:0,numberOfTotalCitations:86,isAvailableForWebshopOrdering:!0,dateEndFirstStepPublish:"May 25th 2017",dateEndSecondStepPublish:"June 15th 2017",dateEndThirdStepPublish:"September 11th 2017",dateEndFourthStepPublish:"December 10th 2017",dateEndFifthStepPublish:"February 8th 2018",currentStepOfPublishingProcess:5,indexedIn:"1,2,3,4,5,6",editedByType:"Edited by",kuFlag:!1,featuredMarkup:null,editors:[{id:"179110",title:"Dr.",name:"Dimitrios",middleName:"P.",surname:"Vlachakis",slug:"dimitrios-vlachakis",fullName:"Dimitrios Vlachakis",profilePictureURL:"https://mts.intechopen.com/storage/users/179110/images/system/179110.jpeg",biography:"Dr. Dimitrios Vlachakis is an Assistant Professor at the Genetics Laboratory at the Biotechnology Department of the Agricultural University of Athens, Greece. He leads the Genetics and Computational Biology Group and his main scientific interests revolve around the investigation of genetic polymorfisms, genetic variability in viral strains and the in silico drug design of novel antiviral and anticancer agents. To date, Dr. Vlachakis has published more than 90 original research articles in international peer-reviewed journals with impact factor, 100+ articles in international conference proceedings, 5 monograph ISBN books, 2 scientific patents and has been on the receiving end of numerous grants and awards. Since 2012 Dr. Vlachakis has been serving as the Editor in Chief and Associate Editor of relevant international journals.",institutionString:"Agricultural University of Athens",position:null,outsideEditionCount:0,totalCites:0,totalAuthoredChapters:"4",totalChapterViews:"0",totalEditedBooks:"3",institution:{name:"Agricultural University of Athens",institutionURL:null,country:{name:"Greece"}}}],equalEditorOne:null,equalEditorTwo:null,equalEditorThree:null,coeditorOne:null,coeditorTwo:null,coeditorThree:null,coeditorFour:null,coeditorFive:null,topics:[{id:"380",title:"Molecular Biology",slug:"biochemistry-genetics-and-molecular-biology-biochemistry-molecular-biology"}],chapters:[{id:"61751",title:"Introductory Chapter: Molecular Docking - Overview, Background, Application and What the Future Holds",doi:"10.5772/intechopen.78266",slug:"introductory-chapter-molecular-docking-overview-background-application-and-what-the-future-holds",totalDownloads:1572,totalCrossrefCites:1,totalDimensionsCites:2,hasAltmetrics:0,abstract:null,signatures:"Dimitrios Vlachakis",downloadPdfUrl:"/chapter/pdf-download/61751",previewPdfUrl:"/chapter/pdf-preview/61751",authors:[{id:"179110",title:"Dr.",name:"Dimitrios",surname:"Vlachakis",slug:"dimitrios-vlachakis",fullName:"Dimitrios Vlachakis"}],corrections:null},{id:"61097",title:"Molecular Docking Studies of Enzyme Inhibitors and Cytotoxic Chemical Entities",doi:"10.5772/intechopen.76891",slug:"molecular-docking-studies-of-enzyme-inhibitors-and-cytotoxic-chemical-entities",totalDownloads:1576,totalCrossrefCites:1,totalDimensionsCites:2,hasAltmetrics:0,abstract:"Docking is a powerful approach to perform virtual screening on large library of compounds, rank the conformations using a scoring function, and propose structural hypotheses of how the ligands inhibit the target, which is invaluable in lead optimization. Using experimentally proven active compounds, detailed docking studies were performed to determine the mechanism of molecular interaction and its binding mode in the active site of the modeled yeast α-glucosidase and human intestinal maltase-glucoamylase. All active ligands were found to have greater binding affinity with the yeast α-glucosidase as compared to that of human homologs, intestinal, and pancreatic maltase, by an average value of ~−1.3 and ~−0.8 kcal/mol, respectively. Thirty quinoline derivatives have been synthesized and evaluated against β-glucuronidase inhibitory potential. Twenty-four analogs, which showed outstanding β-glucuronidase activity, have IC50 values ranging between 2.11 ± 0.05 and 46.14 ± 0.95 μM than standard D-saccharic acid 1,4-lactone (IC50 = 48.4 ± 1.25 μM). Structure activity relationship and the interaction of the active compounds and enzyme active site with the help of docking studies were established. In addition, Small series of morpholine hydrazones synthesized to form morpholine hydrazones scaffold. The in vitro anti-cancer potential of all these compounds were checked against human cancer cell lines such as HepG2 (Human hepatocellular liver carcinoma) and MCF-7 (Human breast adenocarcinoma). Molecular docking studies were also performed to understand the binding interaction.",signatures:"Sadia Sultan, Gurmeet Kaur Surindar Singh, Kamran Ashraf and\nMuhammad Ashraf",downloadPdfUrl:"/chapter/pdf-download/61097",previewPdfUrl:"/chapter/pdf-preview/61097",authors:[{id:"176737",title:"Dr.",name:"Sadia",surname:"Sultan",slug:"sadia-sultan",fullName:"Sadia Sultan"},{id:"222831",title:"Dr.",name:"Kamran",surname:"Ashraf",slug:"kamran-ashraf",fullName:"Kamran Ashraf"},{id:"256881",title:"Dr.",name:"Gurmeet Kaur Surindar",surname:"Singh",slug:"gurmeet-kaur-surindar-singh",fullName:"Gurmeet Kaur Surindar Singh"},{id:"398577",title:"Dr.",name:"Muhammad",surname:"Ashraf",slug:"muhammad-ashraf",fullName:"Muhammad Ashraf"}],corrections:null},{id:"59277",title:"Molecular Docking for Detoxifying Enzyme Studies",doi:"10.5772/intechopen.73920",slug:"molecular-docking-for-detoxifying-enzyme-studies",totalDownloads:1436,totalCrossrefCites:0,totalDimensionsCites:1,hasAltmetrics:0,abstract:"In this chapter, we pointed some relevant results obtained by protein-ligand docking simulations in the context of insecticide and herbicide resistance performed by glutathione S-transferases (GSTs), a detoxifying superfamily enzyme. We present here some in silico evidences of GST binding against chemical insecticides in the malaria and dengue vectors (Anopheles gambiae and Aedes aegypti mosquitoes) and against chemical herbicides used on rice (Oryza sativa) culture. Our findings suggest that some members from epsilon class (GSTE2, GSTE5) can metabolize some insecticide compounds and that a tau class member (GSTU4) can metabolize some herbicides. The results reinforce the importance of docking studies for enzyme activity comprehension. These information can allow in the future the implementation of new strategies for mosquito control and herbicide management on rice culture through biotechnological improvements designed to specific GST targets. Induced mutations on catalytic binding sites of GSTU4 could improve rice herbicide resistance and minimize produce damage, while rational compounds can be designed to inhibit GSTE members to decline insecticide resistance on mosquito control. In both cases, biotechnological tools could be developed focusing on GSTs that would reduce environmental impact by the use of insecticide and herbicide.",signatures:"Rafael Trindade Maia and Vinícius Costa Amador",downloadPdfUrl:"/chapter/pdf-download/59277",previewPdfUrl:"/chapter/pdf-preview/59277",authors:[{id:"212393",title:"Prof.",name:"Rafael",surname:"Trindade Maia",slug:"rafael-trindade-maia",fullName:"Rafael Trindade Maia"},{id:"214489",title:"BSc.",name:"Vinícius",surname:"Amador",slug:"vinicius-amador",fullName:"Vinícius Amador"}],corrections:null},{id:"61865",title:"A Click Chemistry Approach to Tetrazoles: Recent Advances",doi:"10.5772/intechopen.75720",slug:"a-click-chemistry-approach-to-tetrazoles-recent-advances",totalDownloads:2633,totalCrossrefCites:1,totalDimensionsCites:3,hasAltmetrics:0,abstract:"Introduction to tetrazole and click chemistry approaches was briefed in a concise way in order to help the readers have a basic understanding. Tetrazole and its derivatives play very important role in medicinal and pharmaceutical applications. The synthesis of tetrazole derivatives can be approached in ecofriendly approaches such as the use of water as solvent, moderate conditions, nontoxic, easy extractions, easy setup, low cost, etc. with good to excellent yields.",signatures:"Ravi Varala and Bollikolla Hari Babu",downloadPdfUrl:"/chapter/pdf-download/61865",previewPdfUrl:"/chapter/pdf-preview/61865",authors:[{id:"212519",title:"Dr.",name:"Varala",surname:"Ravi",slug:"varala-ravi",fullName:"Varala Ravi"},{id:"221476",title:"Dr.",name:"Bollikolla",surname:"Hari Babu",slug:"bollikolla-hari-babu",fullName:"Bollikolla Hari Babu"}],corrections:null},{id:"58925",title:"Docking Studies on Novel Analogues of 8-Chloro-Quinolones against Staphylococcus aureus",doi:"10.5772/intechopen.72995",slug:"docking-studies-on-novel-analogues-of-8-chloro-quinolones-against-staphylococcus-aureus",totalDownloads:1929,totalCrossrefCites:2,totalDimensionsCites:3,hasAltmetrics:0,abstract:"Molecular docking studies have been carried out for a better understanding of the drug-receptor interactions. All the synthesized compounds have been subjected to molecular docking against targets that have been chosen based on the specific mechanism of action of the quinolones used in the antibacterial activity screening. A study of the characteristics and molecular properties of the small molecule known as ligand has been realized. In the first stage of the study, the 2D and 3D structures have been generated. The most stable conformer for each structure was obtained by geometry optimization and energy minimization. A series of topological, conformational characteristics and QSAR properties, important to assess the flexibility and the ability of the studied conformer to bind to the protein receptor, were determined and analyzed. These properties were discussed in order to assess the flexibility and the binding ability of studied conformers to bind to the receptor protein. The docking studies have been carried out. The score and hydrogen bonds formed with the amino acids from group interaction atoms are used to predict the binding modes, the binding affinities and the orientation of the docked quinolones in the active site of the protein receptor.",signatures:"Lucia Pintilie and Amalia Stefaniu",downloadPdfUrl:"/chapter/pdf-download/58925",previewPdfUrl:"/chapter/pdf-preview/58925",authors:[{id:"94504",title:"Dr.",name:"Lucia",surname:"Pintilie",slug:"lucia-pintilie",fullName:"Lucia Pintilie"},{id:"213696",title:"Dr.",name:"Amalia",surname:"Stefaniu",slug:"amalia-stefaniu",fullName:"Amalia Stefaniu"}],corrections:null},{id:"58703",title:"Molecular Docking in Halogen Bonding",doi:"10.5772/intechopen.72994",slug:"molecular-docking-in-halogen-bonding",totalDownloads:1258,totalCrossrefCites:1,totalDimensionsCites:2,hasAltmetrics:0,abstract:"Molecular modeling applies several computational chemistry tools as molecular docking; this latter has been useful in medicinal chemistry for prediction of interactions between small ligands and biological targets measuring angles, enthalpy and other physical-chemical properties involved in the supramolecular entities. In this chapter, we present molecular docking advances with a perspective to the improvement of parameterization including halogen bonding interactions (XB) and the modification of scoring functions based on halogen sigma-hole polarization. At the same time, we have included the current computational methods to study halogen bonding that increased the accuracy of predicted entities. Finally, we present examples of the main force fields including electronic distribution and modifications for halogen atoms.",signatures:"Abel Suárez-Castro, Mario Valle-Sánchez, Carlos Jesús Cortés-García\nand Luis Chacón-García",downloadPdfUrl:"/chapter/pdf-download/58703",previewPdfUrl:"/chapter/pdf-preview/58703",authors:[{id:"214359",title:"Dr.",name:"Luis",surname:"Chacon",slug:"luis-chacon",fullName:"Luis Chacon"},{id:"214403",title:"M.Sc.",name:"Abel",surname:"Suárez",slug:"abel-suarez",fullName:"Abel Suárez"},{id:"214405",title:"Dr.",name:"Carlos",surname:"Cortés",slug:"carlos-cortes",fullName:"Carlos Cortés"},{id:"215264",title:"MSc.",name:"Mario",surname:"Valle",slug:"mario-valle",fullName:"Mario Valle"}],corrections:null},{id:"59207",title:"A Combined Molecular Docking and Electronic Structure Study for a Breast Cancer Drug Design",doi:"10.5772/intechopen.72895",slug:"a-combined-molecular-docking-and-electronic-structure-study-for-a-breast-cancer-drug-design",totalDownloads:1291,totalCrossrefCites:2,totalDimensionsCites:2,hasAltmetrics:0,abstract:"The molecular docking of tamoxifen’s metabolites, 4-hydroxy-tamoxifen, N-desmethyl-tamoxifen, and 4-hydroxy-N-desmethyl-tamoxifen, in estrogen and progesterone hormone receptors was studied in aqueous solution. The metabolites 4-hydroxy-tamoxifen, N-desmethyl-tamoxifen, and 4-hydroxy-N-desmethyl-tamoxifen exhibit a binding energy in the estrogen receptor cavity of −10.69 kcal/mol, −10.9 kcal/mol, and −11.35 kcal/mol, respectively, and −1.45 kcal/mol, −9.29 kcal/mol, and −0.38 kcal/mol in the progesterone receptor. This indicates a spontaneous interaction between the metabolites and the active sites in the hormone receptors. Docking has an adequate accuracy for both receptors, and from this calculation the active site residues were defined for the different metabolites and the estrogen and progesterone receptors. Also, the chemical reactivity of the amino acids of the active sites of each metabolite was determined. These reactivity properties were obtained within the framework of density functional theory, using the functional M06 with the basis set 6-31G (d). The results indicate that in the estrogen receptor, the highest charge transfer of the three analyzed metabolites is in the union of the metabolite and the Leu346-Thr347 residue. The progesterone receptor shows minor tendency to react with higher hardness values than the estrogen receptor. The hydrogen bonds are three for the estrogen receptor in two different metabolites, while in progesterone only one is formed with the N-desmethyl-tamoxifen metabolite.",signatures:"Linda-Lucila Landeros-Martinez, Daniel Glossman-Mitnik, Erasmo\nOrrantia-Borunda and Norma Flores-Holguin",downloadPdfUrl:"/chapter/pdf-download/59207",previewPdfUrl:"/chapter/pdf-preview/59207",authors:[{id:"34191",title:"Prof.",name:"Erasmo",surname:"Orrantia-Borunda",slug:"erasmo-orrantia-borunda",fullName:"Erasmo Orrantia-Borunda"},{id:"154505",title:"Dr.",name:"Norma",surname:"Flores-Holguín",slug:"norma-flores-holguin",fullName:"Norma Flores-Holguín"},{id:"198499",title:"Dr.",name:"Daniel",surname:"Glossman-Mitnik",slug:"daniel-glossman-mitnik",fullName:"Daniel Glossman-Mitnik"},{id:"214378",title:"MSc.",name:"Linda-Lucila",surname:"Landeros-Martinez",slug:"linda-lucila-landeros-martinez",fullName:"Linda-Lucila Landeros-Martinez"}],corrections:null},{id:"59054",title:"Has Molecular Docking Ever Brought us a Medicine?",doi:"10.5772/intechopen.72898",slug:"has-molecular-docking-ever-brought-us-a-medicine-",totalDownloads:3097,totalCrossrefCites:15,totalDimensionsCites:22,hasAltmetrics:1,abstract:"Molecular docking has been developed and improving for many years, but its ability to bring a medicine to the drug market effectively is still generally questioned. In this chapter, we introduce several successful cases including drugs for treatment of HIV, cancers, and other prevalent diseases. The technical details such as docking software, protein data bank (PDB) structures, and other computational methods employed are also collected and displayed. In most of the cases, the structures of drugs or drug candidates and the interacting residues on the target proteins are also presented. In addition, a few successful examples of drug repurposing using molecular docking are mentioned in this chapter. It should provide us with confidence that the docking will be extensively employed in the industry and basic research. Moreover, we should actively apply molecular docking and related technology to create new therapies for diseases.",signatures:"Mark Andrew Phillips, Marisa A. Stewart, Darby L. Woodling and\nZhong-Ru Xie",downloadPdfUrl:"/chapter/pdf-download/59054",previewPdfUrl:"/chapter/pdf-preview/59054",authors:[{id:"214567",title:"Prof.",name:"Zhong-Ru",surname:"Xie",slug:"zhong-ru-xie",fullName:"Zhong-Ru Xie"},{id:"223007",title:"Ms.",name:"Marisa A.",surname:"Stewart",slug:"marisa-a.-stewart",fullName:"Marisa A. Stewart"},{id:"223009",title:"Mr.",name:"Darby L.",surname:"Woodling",slug:"darby-l.-woodling",fullName:"Darby L. Woodling"},{id:"223013",title:"Mr.",name:"Mark Andrew",surname:"Phillips",slug:"mark-andrew-phillips",fullName:"Mark Andrew Phillips"}],corrections:null}],productType:{id:"1",title:"Edited Volume",chapterContentType:"chapter",authoredCaption:"Edited by"},subseries:null,tags:null},relatedBooks:[{type:"book",id:"8322",title:"Gene Expression Profiling in Cancer",subtitle:null,isOpenForSubmission:!1,hash:"d1f92f63e67400b51488fe66c51fc342",slug:"gene-expression-profiling-in-cancer",bookSignature:"Dimitrios Vlachakis",coverURL:"https://cdn.intechopen.com/books/images_new/8322.jpg",editedByType:"Edited by",editors:[{id:"179110",title:"Dr.",name:"Dimitrios",surname:"Vlachakis",slug:"dimitrios-vlachakis",fullName:"Dimitrios Vlachakis"}],equalEditorOne:null,equalEditorTwo:null,equalEditorThree:null,productType:{id:"1",chapterContentType:"chapter",authoredCaption:"Edited by"}},{type:"book",id:"8646",title:"Systems Biology",subtitle:null,isOpenForSubmission:!1,hash:"e7735d2fe6193b7b256c7098be4adcf4",slug:"systems-biology",bookSignature:"Dimitrios Vlachakis",coverURL:"https://cdn.intechopen.com/books/images_new/8646.jpg",editedByType:"Edited by",editors:[{id:"179110",title:"Dr.",name:"Dimitrios",surname:"Vlachakis",slug:"dimitrios-vlachakis",fullName:"Dimitrios Vlachakis"}],equalEditorOne:null,equalEditorTwo:null,equalEditorThree:null,productType:{id:"1",chapterContentType:"chapter",authoredCaption:"Edited by"}},{type:"book",id:"3203",title:"Oxidative Stress and Chronic Degenerative Diseases",subtitle:"A Role for Antioxidants",isOpenForSubmission:!1,hash:"7014dbaa632114f7220802475ccd0402",slug:"oxidative-stress-and-chronic-degenerative-diseases-a-role-for-antioxidants",bookSignature:"José A. 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He obtained his Master in 1995 and his Doctorate in 1997 both from UNIFESP. He is a specialist in laparoscopy (Cleveland Clinic Foundation, 2000) and has a PhD in reproductive medicine (2008). Due to the relevance of his research he obtained the title of Associate Professor of the Department of Obstetrics of UNIFESP (2006). He has experience in the field of Medicine, with emphasis in Gynecology and Obstetrics, acting mainly in the following subjects: ectopic pregnancy, methotrexate, ultrasonography, beta-hCG, polymorphisms, serological markers and multiple pregnancy. He has published a predictive score for medical treatment in ectopic pregnancy and reference charts for twins. He has published 68 articles in international and national journals, is a reviewer in several national and international journals, and a speaker at a number of national and international congresses. 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Jovandaric and Svetlana J. Milenkovic",coverURL:"https://cdn.intechopen.com/books/images_new/9160.jpg",editedByType:"Edited by",editors:[{id:"268043",title:"Dr.",name:"Miljana Z.",surname:"Jovandaric",slug:"miljana-z.-jovandaric",fullName:"Miljana Z. Jovandaric"}],productType:{id:"1",chapterContentType:"chapter",authoredCaption:"Edited by"}}]},chapter:{item:{type:"chapter",id:"77893",title:"Molecular and Transcriptional Regulation of Seed Development in Cereals: Present Status and Future Prospects",doi:"10.5772/intechopen.99318",slug:"molecular-and-transcriptional-regulation-of-seed-development-in-cereals-present-status-and-future-pr",body:'Cereal seeds are the major source of starch and proteins in staple foods, animal feed, and raw materials for food and fiber-based industries all over the world [1]. Considerable efforts have been made to elucidate the molecular mechanism regulating important agronomic traits in order to improve the cereal seed production. Several agronomic traits, including grain number per spike, spike length, thousand seed weight, seed size and many others, have contributed to grain yield improvement in many cereals plants, with the development of embryo, endosperm and integuments being the most important [2]. As a result, better understanding of the genetic and molecular processes governing seed development is crucial. Here in this book chapter, we provide a comprehensive review on the ontogeny of seed development, followed by genetics, molecular and transcriptional regulation of seed development for improved crop yield.
Biologically, seed is a mature fertilized ovule that consists primarily three parts: the embryo, endosperm, and seed coat (integuments) [3, 4]. The development of seed begins with double fertilization, in which one of the male gamete fertilizes with haploid egg cell to form an embryo and the other male gamete fertilizes the megagametophyte’s diploid central cell to form the triploid nuclear endosperm [5]. The event of seed development, which described below can be divided into three phases: a morphogenesis and cell divisions for endosperm development (0–7 Day post anthesis, DPA), embryo development (7–15 DPA), and maturation (14 to 28 DPA), which includes embryo growth at the expense of endosperm, seed desiccation and storage materials accumulation [6].
The nuclear type of endosperm development is the most common in monocot plants, particularly cereals, where initial endosperm nucleus divides repeatedly without cell wall formation, resulting in a characteristic coenocyte-stage endosperm [7, 8]. The morphogenetic event of the early stages of endosperm development was observed in wheat [9] and rice [10, 11]. The first division of the triploid endosperm nucleus, in which the daughter nuclei are separated in the central cell, without cell wall formation in subsequent mitotic divisions, results in a 256 to 512 multinucleate cell (the endosperm coenocyte) [8, 12]. The nuclei enter a 2-day mitotic hiatus, lead to the formation of interzonal phragmoplast, occurs 3 days after pollination. While much information about the regulation of phragmoplast formation and expansion remain unknown, recent evidence suggests that the mitogen-activated protein kinase cascade plays a key role in this process [13]. The development of cellularization in the coenocytic endosperm then begins with the formation of radial microtubule on all nuclear surfaces. Soon after, the microtubules from the adjacent nuclei meet, creating interzones where callose-based wall material is deposited. Further, radial microtubules that encase each nucleus undergo reorganization, anchoring the nuclei to the central cell wall while extending toward the central vacuole in a canopy of microtubules. In cereals, the endosperms become fully cellular during 6 to 8 days after pollination if this process is repeated four to five times [14, 15].
The fully developed cereal endosperm consists of four main cell types: the aleurone layer, transfer cells, starchy endosperm, and cells of the embryo-surrounding region [16]. The former two cells, i.e. Aleurone layer, transfer cells remain alive at the end of cereal seed development, while later two including starchy endosperm, and cells of the embryo-surrounding have undergone programmed cell death (PCD) with characteristic DNA laddering and organelle degradation [17].
The cereal endosperm has attracted attention from researchers because of its economic importance, and much insight has accumulated about the genes underlying the accumulation of storage products such as proteins and starch. Additionally, the endosperm protects the embryo from atmospheric oxygen that eventually leads to the formation of hydroperoxides and cell death [18] and critical cross-talk between abscisic acid (ABA) and gibberellin (GA) regulating seed development, size, dormancy or storage breakdown during germination are also the results of endosperm—embryo interactions [19, 20]. Considerably less is known about the genes that regulate the developmental biology of these cell types, which is the topic of this section. Cell fate specification in cereal endosperm is believed to occur by positional signaling at an early developmental stage [12]. For simplicity, each cell type is described separately below, although cell fate specification occurs simultaneously with the cellularization process described above. How this integration occurs is unknown, but elucidation of the molecular controls for each of the four cell types should lay the foundation for understanding the genetic specification of the entire endosperm body plan.
Starchy endosperms, which accumulate starch and storage proteins, encoded by transcripts that are expressed differentially in these cells, make up the largest body of cell in the endosperm [21]. There are two types of starchy endosperm present in the cereal crop. The first, and most important, is the inner cells of cell files that remain after endosperm cellularization is complete. The second source of starchy endosperm cells is the inner daughter cells of aleurone cells that divide periclinally. These cells redifferentiate to become starchy endosperm cells and likely are the source of the so-called subaleurone cells found adjacent to the aleurone layer in the starchy endosperm in all cereals. Several collections of mutants such as
The aleurone layer covers the perimeter of the endosperm with the exception of the transfer cell region. Wheat have one layer of aleurone cells, while rice has one to several layers, functions in seed germination by mobilizing starch and storage protein reserves in the starchy endosperm through the production of hydrolases (α-amylase), glucanases, and proteinases after hormone (gibberellic acid) stimulation from the embryo [26]. In the mature grain of cereals, the aleurone layer consists of an estimated 250,000 aleurone cells derived by an estimated 17 rounds of anticlinal divisions. Toward the end of seed maturation, a specialized developmental program confers desiccation tolerance to the aleurone cells, allowing them to survive the maturation process.
Transfer cells develop in the basal endosperm over the main vascular tissue of the maternal plant, where they facilitate solute (mainly of amino acids, sucrose, and monosaccharides), transfer across the plasmalemma between the symplastic (maternal plant) and apoplastic (endosperm) compartments [30]. However, sucrose is not delivered in this form to transfer cell; instead, it is converted into monosaccharide glucose and fructose through the major activity of cell-wall invertase, offering a mechanism for controlling cell division and even cell differentiation in developing kernels [31].
In cereals, the
Several groups of transcripts, for instance, OsPR602 and OsPR9a in rice and Endosperm 1 (
The embryo-surrounding region (ESR) lines the cavity of the endosperm in which the embryo develops and has been studied most extensively in maize. The exact role of the ESR is unknown, but possible functions include a role in embryo nutrition, the establishment of a physical barrier between the embryo and the endosperm during seed development, and providing a zone for communication between the embryo and the endosperm. The ESR development is under the control of CLAVATA3, a peptide hormone with the conserved domain composed on 12 to 14 amino acids, regulates embryo and endosperm development, cotyledon establishment, and pollen wall formation in Arabidopsis [37], while root and stem development in wheat plants [38].
The seed coat (also known as testa) is made up of two structures covering the nucellus [39], while the single integuments ovules can be found in members of certain families. The seed coat provides a mechanical shield protecting the embryo and the endosperm from the environment, but it also regulates phloem unloading of assimilates in growing seeds [40], fluid and gas exchanges with the environment, and seed dormancy and germination [41]. Generally, seed coat development and maturation precede that of filial tissues. In cereals, after an initial phase of cell division during the first two days after flowering (DAF), pericarp differentiation involves cell elongation along the longitudinal axis between 3 and 10 DAF coupled to PCD, and it coincides with the cellularization of the endosperm [42]. PCD in the pericarp may contribute to redistribution of nutrients, relaxation of physical constraints of the maternal tissue to allow inner growth of the filial tissue, and the re-activation, together with PCD in the nucellus and the nucellar projections, of post-phloem transport functions to allow passage of solutes [42]. Crosstalk among embryo, endosperm, and seed coat appears to be complex, but gene networks that coordinate development of these three seed compartments are being elucidated [41, 43].
Seed yield is a quantitative trait that is influenced by the genetics and environment. It is usually determined by plant height, number of primary and secondary branches, plant density, date of flowering, number of panicle per plant, number of seed per panicle, seed size including seed length and seed width, and finally seed weight [44, 45]. The last two traits, i.e. seed number and weight, were found to be trade-off [46], but recent evidence from studies in wheat suggests that increasing one yield component without reducing the other is possible [47]. The grain number has maintained higher phenotypic plasticity throughout domestication events when compared with grain weight, which enables crop to effectively respond to resource availability during early reproductive stages [46]. The critical periods for determination of grain number and weight are also generally considered separated by the developmental stage of anthesis (flowering), although Ugarte et al. [48] found that grain weight was affected by pre-anthesis environmental conditions in other cereals including wheat. The genotype × environment interaction for grain yield is likely strong in winter wheat [49] and rice [50].
To explore candidate genes underlying yield related traits, GWAS were conducted to identify underlying loci for each phenotype. Association mapping has been used to successfully discover significant marker–trait associations in cereal crops including rice [51, 52, 53, 54] and wheat [55, 56, 57, 58]. A large number of well-characterized QTLs such as GW2, GIF1, qSW5, GS3 and qGL7 in rice [59, 60, 61, 62, 63] and more than 40 QTL including TaGW2 [64, 65, 66] associated with kernel morphological traits such as kernel length, kernel width, kernel thickness, kernel length/width ratio, kernel length/thickness ratio, kernel width/thickness ratio, flag leaf width, length and area have been recently identified and mapped in wheat [67, 68, 69, 70]. A variety of QTLs regulating seed size have been identified in other crop species, but they have yet to be functionally characterized [47, 71]. The additional genetic approaches on key agronomic traits for improved yield are presented in Table 1.
Cereal Crop | Traits | Gene/QTL/Markers | References |
---|---|---|---|
Wheat | 1000-grain weight | qTgw.nwipb-4DS; qTgw.nwipb-6AL | [72] |
wsnp_Ex_c32624_41252144, BS00021705_51 | [73] | ||
Grain yield, TKW, spike weight, spike length | rs36032, rs4772, rs736, rs50187, rs59282 | [74] | |
Heading and flowering dates | RAC875_c41145_189, Excalibur_c60164_137, RAC875_c50422_299, Ppd-D1, Vrn-B1, Vrn-D1 | [75] | |
Grain weight and grain number | TaGW2-6A, Rht-B1, Vrn-D1a | [56] | |
Rice | Yield associated loci | qSN8 and qSPB1 | [76] |
Heading date | Ghd8/OsHAP3H | [77, 78] | |
Panicle trait | DENSE AND ERECT PANICLE 1 (DEP1) | [79, 80] | |
Grain length and yield | OsLG3 | [81] | |
Heading date and yield potential | Hd1, Ghd7, and DTH7 | [82] | |
Grain yield and quality traits | qPH1/OsGA20ox2, qDF3/OsMADS50, PL, QDg1, qGW-5b, grb7–2, qGL3/GS3, Amy6/Wx gene and OsNAS3 | [83] |
Genetic approaches for improved seed yield in cereal crops.
Overexpression, targeted mutagenesis and mutation breeding are examples of recent biotechnological strategies that have been used to manage seed development for increased yield. The activity of ADP-glucose pyrophosphorylase (AGPase), starch synthase (SS) includes granule bound starch synthase (GBSS) and soluble starch synthase (SSS), starch branching enzyme (SBE), debranching enzyme (DBE), and amylase catalyzes the synthesis and accumulation of endosperm storage components, primarily starch, in cereal crops [84, 85, 86, 87]. AGPase catalyzes the first committed step of starch biosynthesis, namely the conversion of Glc-1-P and ATP to ADP-glucose and pyrophosphate (PPi). Through a new −1,4-linkage, the glucose moiety from ADP-glucose is transferred to the non-reducing end of the -glucan receptor of existing chains of amylose and amylopectin [86]. In addition few transporters and transcription factors also play an important role in the regulation of the biosynthesis of starch [88, 89]. Modification of these enzymes has the drastic effect on different aspects of starch such as composition, and finally grain yield and summarized in Table 2.
Gene | Crop | Mechanism | Function/phenotypes | References |
---|---|---|---|---|
AGPase | Wheat/Rice | Over expression +Chemical mutagens | Enhanced ADP-glucose pyrophosphorylase activity in endosperm and seed yield | [90, 91, 92, 93] |
GBSS | Wheat | Combining null alleles | Low amylose and lower yield | [94] |
SSI,SSII/SSIII | Wheat/Rice | RNAi silencing | Reduced SSI enzyme activity with novel starch structure | [95] |
SSSIIIa | Rice | Chemical mutagen | High amylose | [96] |
BEIIa | Wheat/Durum wheat | RNAi silencing TILLING | High amylose and resistant starch | [97, 98, 99] |
ISA | Rice | RNAi silencing | Alters the physicochemical properties of starch | [100] |
AMY | Wheat | Overexpression | Increased the soluble carbohydrate (mainly sucrose) in dry seed | [101] |
Rice | Overexpression | Regulates the expression of starch biosynthetic genes in rice endosperm | [102] |
Molecular approaches for improved seed yield in cereal crops.
In the context of seed development, genotype-specific and stage-dependent temporal shifts in gene expression profile have been reported in the aleurone, embryo and endosperm, and other cell-type of maturing seeds, potentially leading to seed phenotypic differences [103, 104]. Transcriptomic studies in several plant systems has led to the identification of transcriptional programs and regulatory networks underlying molecular functions associated with cellular activities in endosperm [105, 106], starch metabolism [107], seed storage substances and high molecular weight glutenin genes [108, 109, 110], grain quality (glycemic index) [111], post-transcriptional regulations occurs at the end of seed development [17] and programming of seed developmental and maturation processes, and elucidation of the underlying functional transitions (Table 3) [103].
Cereal Crop | Traits | Transcription factor/gene | References |
---|---|---|---|
Wheat | Grain number per spike | [112] | |
Endosperm specific transcription factor | bHLH (seven tissue-specific bHLH TF clusters were identified according to their expression patterns in endosperm, aleurone, seedlings, heading-stage spikes, flag leaves, shoots and roots). | [113] | |
Starch biosynthesis | bZIP (TabZIP 151, TabZIP121, TabZIP69.1, howing moderate negative to moderate positive correlation with GBSSI and SBEIIb, respectively | [89] | |
Embryo and endosperm specific transcriptome | Identification of genes underlying macromolecules biosynthesis (starch, protein, lipid, protein translation) | [17] | |
ABA mediated transcriptional mechanisms controlling seed maturation | [103] | ||
Identification of key genes for processing quality | [105] | ||
Rice | Seed germination, grain size and yield | [114] | |
Fatty acid metabolism | [115] | ||
Panicle branching | miR156 targeting OsSPL13, OsSPL14 and OsSPL16 | [116] | |
[117] | |||
Seed setting | [118] | ||
Metabolism of sugars, fatty acids, amino acids, and phytosterols | Mutation on | [119] | |
Transcriptome analysis of colored rice | Flavonoid biosynthetic pathway | [120] | |
Accumulation of seed storage substance | NF-YC12 | [121] | |
Regulation of grain size | OsPIL15, targeting purine permease gene OsPUP7 | [122] | |
Early seed development | [123] | ||
Plant architecture, longer panicles, more grain number and yield | [124] | ||
Leaf angle, grain size and seed quality | OsmiR1848 regulating OsCYP51C expression and mediates BR biosynthesis | [125] |
Transcriptional approaches for improved seed yield in cereal crops.
In rice, Nie et al. [15], identified 12 classes of endosperm-specific genes, including transcription factor, stress/defense, seed storage protein (SSP), carbohydrate and energy metabolism, seed maturation, protein metabolism, lipid metabolism, transport, cell wall related, hormone related, signal transduction, and one unclassified category. In addition, several cis-regulator elements were found in the promoter region of endosperm-specific expressed genes including, AACA box, ACGT box, GCN4 motif (TGA (G/C) TCA), the prolamin box (P box: AAAG), SKN-1
Based on the cis-element, the corresponding transcription factor were also determined. For example, the MYB protein specifically binds to the AACA box, and the GNC4 motif is bounded by transcription factors of the Opaque2-like basic leucine zipper (bZIP) activators (rice RISBZ1), ABRE motif by bZIP transcription factors, the P box by plant-specific DNA binding with one finger (DOF) zinc-finger transcription factors (rice RPBF), and FUSCA3 (FUS3) recognizes the RY repeats [29, 127, 128]. In addition, synergy between RPBF and RISBZ1 has been implicated in mediating the regulatory networks essential for seed development by binding to the GCN4 motif to trans-activate the expression of seed storage proteins in rice [29, 129]. Recently, Grimberg et al. [130] identified an oat endosperm homolog of WRINKLED1 transcription factor (
Polyamines such as putresceine, Spermidine (Spd), and Spermine (Spm) have been implicated in regulation of spikelets postanthesis development [131]. Exogenous Spd and Spm are applied to rice panicles to improve grain filling and grain weight in inferior spikelets [132]. Furthermore, the concentrations of Spd and Spm are related to rice grain size. The
During plant reproductive growth, cell-to-cell communication via receptor-like kinases (RLKs) regulates a wide range of biological processes. FLORALORGANNUMBER1 (FON1), a potential ortholog of CLAVATA1 (CLV1), interacts with the putative ligand FON2/FON4, a CLV3-related protein, to maintain the inflorescence meristem [134]. The orthologous
Seed development is a multi-step process that includes the production of an embryo and endosperm. The synthesis and accumulation of storage product in the seed is controlled by genetics, molecular and transcriptional regulation, which is critical for maximum yield. For instance, seed yield improvement can be achieved directly under genetic control by selecting and applying markers, QTL linked to agronomic and physiological traits, and improved grain yield potential. Intensive use of molecular tools such as Genetic engineering, Gene silencing and Genome editing together with increase access of system biology tools would provide researchers to gain a better understanding of the pathways and genes that control seed size and number, resulting greater yield as shown in Figure 1. It is envisaged that a more detailed investigation is urgently required for understanding of metabolic control of seed development, storage, product partitioning, epigenetic controls, phytohormone regulation and their interplay would appear to be sufficient to solve global food security challenges faced by the world in future.
Summary of Molecular Approaches for Regulation of Seed Development through Plant Breeding & Genetics, Genetic Engineering & Genome Editing and at Transcriptional Levels.
The mandibular central and lateral incisors have a single conical root. The root dimensions of both incisors vary corresponding to the crown. They are narrow in mesiodistal dimension and wide in labiolingual dimension and taper uniformly on both proximal sides from the CEJ to the apex. The apical end may curve slightly to the distal. Longitudinal root depressions can be seen in both incisors from the mesial and distal views. Multiple comparisons revealed that, among all permanent teeth, mandibular central incisor has the shortest root. Furthermore, in contrary to maxillary incisors, the root of mandibular lateral incisor is longer than that of mandibular central incisor [1]. It has been reported that the average lengths of mandibular central incisor and lateral incisor roots are 12.6 mm (7.7–17.9) and 13.5 (9.4–18.1), respectively [2]. Kim et al. [3] measured the mandibular incisor root lengths using CBCT in Korean population and found that no significant differences in crown and root lengths were noted between the CBCT-based and direct measurements. The R/C ratios were higher for the mandibular lateral incisors (1.4 ± 0.1) than mandibular central incisors (1.3 ± 0.1) [4]. Therefore, crown lengthening may not be possible in the case of traumatic fracture or iatrogenic orthodontic extrusion due to the short root length in these teeth. Variations in root length between males and females have been reported. According to Zorba et al. [5], it was observed that root length was greater in males than in females. Haghanifar et al. [6] found similar results when comparing crown and root lengths between males and females. He found that females had longer crowns while males had longer roots.
Many authors reported that the external crown and root morphology of mandibular central and lateral incisors are similar [2, 7, 8]. Mandibular incisors usually have a single root, which is wider buccolingually than mesiodistally and tapers toward the apex. The lateral incisor root is larger than that of the central incisor in mesiodistal and labiolingual directions [8, 9]. Variation in number of roots has not been reported in literature. However, Loushine et al. [10] have found two rooted mandibular lateral incisors. However, the shape may vary from conical to round in different populations. Sexual variation in the number and shape of roots has not been reported [9]. Mandibular incisor roots are commonly reported to be straight and in rare occasions curved in the apical region. Curvature can be in the mesial, distal, labial, or lingual direction [9].
Orban stated that the shape of the root canal “to a large degree, conforms to the shape of the root. A few canals are round and tapering, but many are elliptical, broad and thin” [11].
The internal anatomy of permanent mandibular incisors does not usually reproduce the simplicity of external anatomy. Its internal anatomy is complicated by the presence of lingual canals, lateral canals, isthmus, and apical deltas [12]. The pulp cavity is the central cavity within a tooth and is entirely enclosed by dentin except at apical foramen. It is divided into coronal portion (pulp chamber) and radicular portion. The pulp chamber is wide and ovoid labiolingually and it tapers incisally. The size of the pulp chamber is not constant throughout life. It decreases in size with aging as a result of secondary dentin deposition [13]. The pulp horn is well developed in this tooth. The root canal systems of these single-rooted teeth often have a single root with a single root canal. However, studies have shown that the root canal anatomy of these teeth is not simple. It may not be single and straight as it appears on the periapical radiograph. Indeed, these teeth have a high prevalence of bifurcation, second canals, lateral canals, and apical deltas which would complicate surgical and nonsurgical endodontic treatment. Mandibular incisor’s anatomy presents a challenge when an endodontic access is made, because of its small size and high prevalence of two canals. The main reason for failure in endodontic treatment of mandibular incisors is the inability to detect the presence of a second canal which can then not be prepared and filled during treatment [14]. In literature, the incidence of mandibular incisor teeth with more than one canal has been reported to range from 11 to 68% [15, 16, 17, 18, 19]. The differences between these morphology studies may be related to variations of examination methods, classification systems, sample sizes, and ethnic background of tooth sources. Many researchers have studied the prevalence of a second canal in mandibular permanent incisors on different populations and showed that the root canal morphology varies with race, sex, and age [20, 21, 22, 23, 24].
Routine clinical radiographs may mislead clinicians to be under an impression that all root canals are round in shape. A high prevalence of oval root canals in human teeth was reported [25, 26].
The pulp canal of the permanent mandibular central incisor is wider buccolingually than mesiodistally [9]. These dimensions are not constant along the root from the orifice till the apex. Oval canals and long oval canals are the most common canal shape seen in the coronal and middle third [27]. As we approach the apex, the canal shape becomes more rounded [28]. This canal shape morphology corresponds to the shape of the root.
The root canal morphology of mandibular central and lateral incisors is very similar. Although they have only one root and a high prevalence of Type 1 root canal morphology, surgical and nonsurgical root canal treatment may fail in these teeth if there is a lack of awareness in their internal anatomy which is complicated by the presence of the lingual canal, bifurcation, lateral anatomy, and isthmus [17, 29]. The morphological characteristics of the root canal system were studied using a number of techniques [18, 27, 30]. The prevalence of a second canal in mandibular permanent incisors is different between populations. Vertucci [18] reported that the incidence of the presence of a second canal was 25.7% among American population, whereas the incidence in Chinese population for the mandibular central and lateral incisors was 5.71 and 27.36%, respectively [31], 30% in Saudi population [32], 26.2% in north Jordanian population [33], and 36.25% in North-East Indian population [34]. In Iranian population, the incidence of mandibular central and lateral incisors having two canals was 27.3 and 29.4%, respectively [35]. The highest incidence (63%) of a second canal in mandibular incisors has been reported in a study in Turkish population [19].
Rankine-Wilson and Henry [36] filled the root canals of mandibular anterior teeth with radio-opaque material, sectioned them in a horizontal plane, and exposed radiographs. They reported two canals in 40.5% of mandibular incisors. Later, Vertucci [18] studied the root canal morphology of 300 extracted mandibular anterior teeth using the clearing technique. In 30% of mandibular central incisors and in 25% of mandibular lateral incisors, there was a second canal. On the other hand, higher prevalence of a second canal in Chinese population was reported in lateral mandibular incisors 25.5% compared with 10.9% in central mandibular incisors [37].
Many researches have shown that root canal systems also vary according to gender. In Turkish population, Sert and Bayirli [19] reported the incidence of second canal in central incisors in females (70%) was higher than in males (65%). Also in Turkish population, Arslan et al. [38] found the frequency of mandibular incisors with a second root canal in males (63%) was higher than in females (35%). The differences among both studies may be due to the fact that Sert and Bayirli examined the root canal morphology in vitro, whereas Arslan et al. studied the root canal anatomy in vivo. In Chinese population, Zhengyan et al. [30] found a significant difference between sex. The result of his study showed that 9.4% of the mandibular lateral incisors in males had a second canal, whereas this value was 11.9% in females. Among Iranian population, Haji et al. [39] reported that there was no significant difference between males and females in the incidence of a second canal in mandibular incisors.
It has become clear that teeth have complicated root canal systems rather than simplified canals [40]. Most investigators have shown that the root canal systems for most, if not all, permanent teeth are complex and canals may branch, divide, and rejoin. In addition to the complexity of root canal anatomy, root canal morphology varies from tooth to tooth. Concerning root canal treatment, these variations in root canal morphology of permanent teeth may result in missing root canals, nonsurgical endodontic treatment failure, and a need for surgical procedures. Weine et al. [41] classified root canal systems into four basic types, but Vertucci [18] subsequently classified them into eight configurations. The Vertucci classification may give consideration to the complex reality of canal systems in a way that the Weine et al. system did not.
Weine [42] described each of the canal types as below:
Vertucci [24] classified canal configurations into eight types as described below:
Although mandibular incisors are usually single-rooted teeth, their root canal system cannot be predicted not only between different populations but also between the same population, with respect to the Vertucci’s configuration. Studies reported that Vertucci’s Type I configuration has the highest prevalence among the other Vertucci configurations [43, 44, 45]. When a second canal is present, Vertucci’s Type III configuration is the most common for central and lateral incisors. Scarlatescu [46] found Type III has higher incidence than Type II, of 25 and 6.3% respectively in a Romanian population. de Almeida [47] reported that Vertucci’s Type I and III configurations represented 92% of the sample. Leoni investigated the root canal anatomy of mandibular central (
Lateral canals are accessory canals located in the coronal or middle third of the root, extending horizontally from the main canal to the external surface of the root. Their formation is due to the entrapment of periodontal vessels in Hertwig’s epithelial root sheath or when blood vessels running from the dental sac through the dental papilla persist during calcification [50]. Lateral canals communicate with the periodontal ligament space and this increases risk of spread of periodontal disease into the pulp canal. According to their location, Vertucci classified lateral canals into coronal, middle, apical, or furcation. He observed lower occurrence of canal ramifications in the middle 11.4% and coronal 6.3% thirds compared to the apical 73.5% third [18, 24]. Recent micro-CT studies on root canal morphology of mandibular anterior teeth reported that lateral canals are rare [48, 51]. Miyashita et al. [17] reported that out of mandibular incisors with lateral branches, single lateral branch had the highest prevalence (82.2%) and multiple branches were extremely narrow. Al-Qudah and Awawdeh [33] found that there was an increasing prevalence of lateral canals toward the apical third of the root with approximately 64% occurring in the apical part of the roots. On the other hand, other studies reported that lateral canals were frequently found in the middle of the canal [34, 46]. Clinically, lateral canals are not usually visible in preoperative radiographs, but its presence can be suspected when there is a localized thickening of the periodontal ligament or a lesion on the lateral surface of the root [50]. It is also important to note that lateral canals cannot be instrumented. Its contents can only be neutralized by the action of effective irrigation with appropriate tissue dissolvent properties and antimicrobial activity solution or with the addition of use of intracanal medications.
Apical deltas are defined as an intricate system of spaces within the root canal that allows free passage of blood vessels and nerves from the periapical compartment to the pulp tissue [52, 53]. The apical delta is different from the accessory canal in which the main pulp canal is still distinguishable. The prevalence of apical deltas in human permanent teeth varies among populations, and the type and locations of tooth and methods of study. High prevalence of apical deltas is found in maxillary second premolars, mandibular lateral incisors, and mandibular second premolars [22]. Among American population, Vertucci [18] reported that the incidence of apical deltas was 5, 6, and 8% in the mandibular central incisors, lateral incisors, and canines, respectively. However, Çalişkan et al. [22] reported that the prevalence of apical deltas in those teeth was 9.8, 23.5 and 7.8% in a Turkish population. Apical deltas have been reported to be of great importance in endodontics because they are difficult to be instrumented during chemical-mechanical preparation. Furthermore, their long vertical extension may cause failure of the apical surgery if not involved during apical resection [54]. Gao et al. [55] reported that the median vertical distance of the apical delta was 1.87 mm with 13% of them more than 3 mm. Therefore, resection of the apical 3 mm of a root may include the whole apical delta and residual microorganisms from 87% of roots with apical delta.
A thin communication can occur between two or more canals in the same root or between vascular elements in tissues [56]. Green [23] described this corridor as a “ribbon shaped passage.” He found this corridor in 22% of mandibular incisors. An isthmus is formed when an individual root projection is unable to close itself off. Any root that contains two root canals has the potential to contain an isthmus [57]. It may contain tissue remnants and necrotic debris, which participate in microorganisms’ growth resulting in root canal treatment failure [58]. Therefore, knowledge of the root canal anatomy is essential for complete cleaning of the root canal and successful endodontic treatment [11]. Isthmus classification was described by Hsu and Kim et al. [59]. They classified isthmus into five types: Type I—two canals with no notable communication; Type II—a hair-thin connection between the two main canals; Type III—differs from Type II because of the presence of three canals instead of two; Type IV—an isthmus with extended canals into the connection; and Type V—there is a true connection or wide corridor of tissue between two main canals. Mauger reported that isthmus was present in 20% of the teeth at the 1-mm level, 30% at 2 mm, and 55% at 3 mm [27]. Estrela et al. [60] demonstrated high prevalence of both partial and complete isthmii in mandibular lateral incisors (47.6%) compared with mandibular central incisors (33.3%). On the other hand, Arslan et al. [38] found a low incidence (3.7%) of intracanal communication among Turkish population. A similar study done by Haghanifar [61] found the prevalence of complete isthmus in the mandibular anterior teeth ranged from 3 to 5%.
As a result of large width of the root canal buccolingually than mesiodistally, mandibular incisors have oval and flattened canals [25]. The overall prevalence of long oval root canals in the apical region in mandibular incisors is >50% [25]. When using rotary files, these oval-shaped canals are a challenge for proper shaping of the canal. This is because rotary instrumentation cannot touch all the canal walls, leaving behind untouched area. To improve mechanical apical debridement, the use of instruments up to an ISO size 100 is required to avoid leaving untouched area on the buccal and/or lingual walls of the canal [62]. However, using files with large taper or tip may cause lateral or apical perforation of the root as the root has a narrower diameter in the mesiodistal direction. Therefore, it stresses the use of good chemical disinfection protocol on these teeth. Canals are considered as oval, long oval, and flattened when the ratio between the maximum and the minimum cross-sectional diameter is <2:1, 2–4:1, and >4:1, respectively. Apical foramina are the main apical opening of the root canal. It is the main exit of the root canal onto the external root surface. Variation in the number and position of apical foramina is especially seen in mandibular incisors with two canals. The apical foramen coincides with the anatomical apex in 17–52.2% of the cases [19, 22, 33, 57, 63].
A number of studies (17.33%) reported that the position and the number of the apical foramen vary according to the race. Al-Qudah and Awawdeh [33] reported that more than half of the roots (52.2%) had centrally located foramina and 47.8% had laterally located foramina. Apical deltas were observed in only eight teeth (1.8%), and among mandibular incisors with two canals, single foramen was more prevalent than two apical foramina. Miyashita et al. [17] reported that only 3% of the mandibular incisors containing two canals had two foramina. He also found that 67.9% of mandibular incisors with curved root had eccentrically located foramina toward the labial direction and none of the canals were curved lingually.
According to Walker [63], the distance between the apical foramen and the most apical end of the root ranges between 0.2 and 2.0 mm. The diameter of the apical foramen of mandibular incisors has been reported to be as 262.5 μm.
Anomaly (Gk, anomalos; irregular) is a deviation from what is regarded as normal [64]. These abnormalities may occur, in terms of size or shape, to either crown or root. WHO listed the following dental anomalies: concrescence, fusion, gemination, dens evaginatus, dens in dente, dens invaginatus, enamel pearls, macrodontia, microdontia, peg-shaped teeth, taurodontism, and tuberculum paramolare [65]. Anomalies of permanent mandibular incisors regarding the crown and root shape are extremely rare. However, few case reports have registered anomalies associated with mandibular incisors. As an example, dens invaginatus, a deep surface invagination of the crown or root, which is lined by enamel and resulting from the invagination of the enamel organ into the dental papilla during odontogenesis, can be seen in these teeth [66]. Dens invaginatus has been classified into three categories according to the depth of invagination and communication with the periapical tissue or periodontal ligament [67].
The prevalence of this anomaly has been found to range from 0.25 to 5.1% of the population [66]. More commonly, dens invaginatus occurs in the maxillary permanent lateral incisors. Also, it may occur in maxillary central incisors, premolars, canines, and molars. It usually occurs unilaterally, but bilateral cases have also been reported [68]. Occurrence of dens invaginatus in mandibular teeth is very rare. When it occurs in mandibular incisors, the central incisor has a higher incidence compared with lateral incisor [69, 70].
Talon cusp is also a rare developmental anomaly defined as an additional cusp that projects predominantly from the labial or lingual surface of primary or permanent anterior teeth [71]. Mellor and Ripa [72] named this anomaly “talon cusp” as it resembles the shape of an eagle’s talon. Talon cusp was classified by Hattab [73] as follows:
Radiographically, the talon cusp may appear typically as a V-shaped radiopacity, starting from the cervical third of the crown. Most of the talon cusps occur in the maxillary lateral incisors (55%), followed by maxillary central incisors (32%) and maxillary canines (9%) [71]. Although it is rarely seen in mandibular teeth [74], Gündüz and Celenk [43] studied the site distribution of talon cusp among Turkish population and found only 3% of talon cusp was seen in the mandibular right central incisors.
Another rare developmental anomaly that has been reported to occur in mandibular central incisor is “Gemination” [75]. It is a rare anomaly that arises when the tooth bud of a single tooth attempts to divide. The structure most often presents as two crowns, either totally or partially separated, with a single root and one root canal [76]. In the anterior region, gemination can cause poor esthetic appearance due to irregular morphology. In addition, these teeth are more susceptible to periodontal disease and caries, if deep groove is present [77, 78].
Fusion is another developmental anomaly which can occur in these teeth. Contrary to gemination, fusion is defined as the union of two or more separately developing tooth germs during odontogenesis, when the crown is not yet mineralized at the dentinal level, yielding a single large tooth [79]. Depending on the stage of development at the time of union, the pulp might be merged or separated [80]. Fusion is more frequently seen in primary dentition, but it may occur in both dentitions. If it occurs in permanent dentition, the vast majority of permanent teeth fusion cases are seen in maxillary teeth. Although, the incidence of fusion of mandibular incisors is rare, mandibular central incisors have been reported to fuse with a supernumerary tooth [81] and bilaterally with the adjacent lateral incisor [82].
It should be emphasized that special attention is required during root canal treatment owing to the abnormal morphology of the crown and the complexity of the root canal system in fused teeth.
Mandibular incisors are prone to endodontic treatment as a result of several reasons. Due to their location in the jaw, they are prone to traumas that result in tooth fracture which may necessitate root canal therapy. Moreover, their proximity to the opening of the sublingual and submandibular ducts increases the incidence of dental caries as a result of lingual deposition of calculus. Therefore, an accurate knowledge of the external and internal anatomy of these teeth is an essential prerequisite to carry out root canal treatment. They often have two canals that are buccolingually located and the lingual canal usually is missed. Therefore, the dentist should extend the access preparation in lingual direction to locate the lingual canal which is usually below the cingulum. In case of two canals, Type II canal is the most prevalent configuration where the buccal canal is the most straight and easiest to be located. Consequently, it is recommended to instrument and fill these canals till the apex whereas the lingual canal merges with the labial canal. Presence of an isthmus may complicate the root canal disinfection as it may contain tissue remnants and necrotic debris, hence irrigation and activation are very essential to overcome these anatomical difficulties.
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From Old Problems to New Challenges"},signatures:"Andreas Schmeling, Pedro Manuel Garamendi, Jose Luis Prieto and María Irene Landa",authors:[{id:"34264",title:"Prof.",name:"Pedro Manuel",middleName:null,surname:"Garamendi Gonzalez",slug:"pedro-manuel-garamendi-gonzalez",fullName:"Pedro Manuel Garamendi Gonzalez"}]},{id:"19161",doi:"10.5772/19234",title:"Diagnostic of Drowning in Forensic Medicine",slug:"diagnostic-of-drowning-in-forensic-medicine",totalDownloads:8196,totalCrossrefCites:9,totalDimensionsCites:18,abstract:null,book:{id:"243",slug:"forensic-medicine-from-old-problems-to-new-challenges",title:"Forensic Medicine",fullTitle:"Forensic Medicine - From Old Problems to New Challenges"},signatures:"Audrey Farrugia and Bertrand Ludes",authors:[{id:"34146",title:"Dr.",name:"Audrey",middleName:null,surname:"Farrugia",slug:"audrey-farrugia",fullName:"Audrey Farrugia"},{id:"49284",title:"Dr.",name:"Bertrand",middleName:null,surname:"Ludes",slug:"bertrand-ludes",fullName:"Bertrand Ludes"}]},{id:"19172",doi:"10.5772/22792",title:"Advanced Medical Imaging and Reverse Engineering Technologies in Craniometric Study",slug:"advanced-medical-imaging-and-reverse-engineering-technologies-in-craniometric-study",totalDownloads:4535,totalCrossrefCites:2,totalDimensionsCites:6,abstract:null,book:{id:"243",slug:"forensic-medicine-from-old-problems-to-new-challenges",title:"Forensic Medicine",fullTitle:"Forensic Medicine - 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These include situation of death, anatomical and histological analysis, toxicology and microbiological study. A low part of autopsies remain without a conclusive cause of death, but all these cases are usually seen in young population, apparently healthy who died suddenly and unexpectedly. In these situations a cardiac arrhythmia is suspected as cause of death and genetic testing is recommended despite not regularly performed. Sudden death is a natural and unexpected decease that occurs in apparently healthy people, or whose disease was not severe enough to expect a fatal outcome. It can be due to several pathologies, usually of cardiac cause and called sudden cardiac death. In infants and young people, both long QT syndrome and catecholaminergic polymorphic ventricular tachycardia are main causes in negative autopsies. These genetic diseases lead to ventricular fibrillation, syncope and sudden cardiac death in a normal heart. Unfortunately, sudden cardiac death could be the first manifestation of the diseases, being early identification and prevention a crucial point in current medical practice. This chapter focuses on sudden death and negative autopsy in young population, mainly due to cardiac arrhythmias.",book:{id:"6262",slug:"post-mortem-examination-and-autopsy-current-issues-from-death-to-laboratory-analysis",title:"Post Mortem Examination and Autopsy",fullTitle:"Post Mortem Examination and Autopsy - Current Issues From Death to Laboratory Analysis"},signatures:"Georgia Sarquella-Brugada, Sergi Cesar, Anna Fernandez-Falgueras,\nMaria Dolores Zambrano, Anna Iglesias, Josep Brugada, Ramon\nBrugada and Oscar Campuzano",authors:[{id:"54165",title:"Prof.",name:"Ramon",middleName:null,surname:"Brugada",slug:"ramon-brugada",fullName:"Ramon Brugada"},{id:"54168",title:"Dr.",name:"Oscar",middleName:null,surname:"Campuzano",slug:"oscar-campuzano",fullName:"Oscar Campuzano"},{id:"218478",title:"Dr.",name:"Georgia",middleName:null,surname:"Sarquella-Brugada",slug:"georgia-sarquella-brugada",fullName:"Georgia Sarquella-Brugada"},{id:"218479",title:"Dr.",name:"Sergi",middleName:null,surname:"Cesar",slug:"sergi-cesar",fullName:"Sergi Cesar"},{id:"218480",title:"MSc.",name:"Anna",middleName:null,surname:"Fernandez-Falgueras",slug:"anna-fernandez-falgueras",fullName:"Anna Fernandez-Falgueras"},{id:"218482",title:"Dr.",name:"Maria Dolores",middleName:null,surname:"Zambrano",slug:"maria-dolores-zambrano",fullName:"Maria Dolores Zambrano"},{id:"218483",title:"MSc.",name:"Anna",middleName:null,surname:"Iglesias",slug:"anna-iglesias",fullName:"Anna Iglesias"},{id:"218484",title:"Prof.",name:"Josep",middleName:null,surname:"Brugada",slug:"josep-brugada",fullName:"Josep Brugada"}]},{id:"57778",title:"Defining Dental Age for Chronological Age Determination",slug:"defining-dental-age-for-chronological-age-determination",totalDownloads:2574,totalCrossrefCites:1,totalDimensionsCites:3,abstract:"Dental age assessment is one of the most reliable methods of chronological age estimation used for criminal, forensic and anthropologic purposes. Visual, radiographic, chemical and histological techniques can be used for dental age estimation. Visual method is based on the sequence of eruption of the teeth and morphological changes that are caused due to function such as attrition, changes in color that are indicators of aging. Radiographs of the dentition can be used to determine the stage of dental development of the teeth from initial mineralization of a tooth, crown formation to root apex maturation. Histological methods require the preparation of the tissues for detailed microscopic examination. The chemical analysis of dental hard tissues determines alterations in ion levels with age, whereas the histological and chemical methods are invasive methods requiring extraction/sectioning of the tooth. In this chapter, the different techniques and considered studies were overviewed in conjunction with their advantages and disadvantages. It needs to be taken into consideration that rather than restricting on one age estimation technique, using the other available techniques additionally and performing repetitive measurements may be beneficial for accurate age estimation.",book:{id:"6262",slug:"post-mortem-examination-and-autopsy-current-issues-from-death-to-laboratory-analysis",title:"Post Mortem Examination and Autopsy",fullTitle:"Post Mortem Examination and Autopsy - Current Issues From Death to Laboratory Analysis"},signatures:"Fatma Deniz Uzuner, Emine Kaygısız and Nilüfer Darendeliler",authors:[{id:"172009",title:"Dr.",name:"Fatma Deniz",middleName:null,surname:"Uzuner",slug:"fatma-deniz-uzuner",fullName:"Fatma Deniz Uzuner"},{id:"200985",title:"Dr.",name:"Emine",middleName:null,surname:"Kaygisiz",slug:"emine-kaygisiz",fullName:"Emine Kaygisiz"},{id:"222232",title:"Prof.",name:"Nilufer",middleName:null,surname:"Darendeliler",slug:"nilufer-darendeliler",fullName:"Nilufer Darendeliler"}]},{id:"77222",title:"Forensic Analysis and Interpretation of Tool Marks",slug:"forensic-analysis-and-interpretation-of-tool-marks",totalDownloads:492,totalCrossrefCites:0,totalDimensionsCites:0,abstract:"The forensic analysis and interpretation of tool marks raise for consideration key methods and advances in the field of tool marks in forensic science. This chapter shows how tool mark analysis can be utilized in the course of criminal investigations. The focus of the chapter is on bringing together as much scientific knowledge in the area as possible in an accessible manner. It covers all aspects of tool mark evidence from the crime scene to the courtroom. This chapter provides information about tool marks in an effort to assist tool mark examiners as well as people practicing forensic science, crime scene examiners, crime investigating officers and members of the legal profession. 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