Typical reservoir hosts of Leptospira.
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More than half of the publishers listed alongside IntechOpen (18 out of 30) are Social Science and Humanities publishers. IntechOpen is an exception to this as a leader in not only Open Access content but Open Access content across all scientific disciplines, including Physical Sciences, Engineering and Technology, Health Sciences, Life Science, and Social Sciences and Humanities.
\\n\\nOur breakdown of titles published demonstrates this with 47% PET, 31% HS, 18% LS, and 4% SSH books published.
\\n\\n“Even though ItechOpen has shown the potential of sci-tech books using an OA approach,” other publishers “have shown little interest in OA books.”
\\n\\nAdditionally, each book published by IntechOpen contains original content and research findings.
\\n\\nWe are honored to be among such prestigious publishers and we hope to continue to spearhead that growth in our quest to promote Open Access as a true pioneer in OA book publishing.
\\n\\n\\n\\n
\\n"}]',published:!0,mainMedia:null},components:[{type:"htmlEditorComponent",content:'
Simba Information has released its Open Access Book Publishing 2020 - 2024 report and has again identified IntechOpen as the world’s largest Open Access book publisher by title count.
\n\nSimba Information is a leading provider for market intelligence and forecasts in the media and publishing industry. The report, published every year, provides an overview and financial outlook for the global professional e-book publishing market.
\n\nIntechOpen, De Gruyter, and Frontiers are the largest OA book publishers by title count, with IntechOpen coming in at first place with 5,101 OA books published, a good 1,782 titles ahead of the nearest competitor.
\n\nSince the first Open Access Book Publishing report published in 2016, IntechOpen has held the top stop each year.
\n\n\n\nMore than half of the publishers listed alongside IntechOpen (18 out of 30) are Social Science and Humanities publishers. IntechOpen is an exception to this as a leader in not only Open Access content but Open Access content across all scientific disciplines, including Physical Sciences, Engineering and Technology, Health Sciences, Life Science, and Social Sciences and Humanities.
\n\nOur breakdown of titles published demonstrates this with 47% PET, 31% HS, 18% LS, and 4% SSH books published.
\n\n“Even though ItechOpen has shown the potential of sci-tech books using an OA approach,” other publishers “have shown little interest in OA books.”
\n\nAdditionally, each book published by IntechOpen contains original content and research findings.
\n\nWe are honored to be among such prestigious publishers and we hope to continue to spearhead that growth in our quest to promote Open Access as a true pioneer in OA book publishing.
\n\n\n\n
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In more detail, these topics are pertaining to the geological characteristics and the production response of a reservoir located offshore the Niger Delta (Nigeria), the coastal lacustrine geo-archives with the example of the Lake Bafa (Turkey), the sedimentary processes in the riparian zone of the Ruxi Tributary Channel (Three Gorges Reservoir, China), the beach morphological changes studied by means of a contour-line change model and finally, the role of the mangroves in controlling the sedimentary accretion of coastal and marine environments with the regional example of the south-eastern Asia.",isbn:"978-1-78984-765-9",printIsbn:"978-1-78984-764-2",pdfIsbn:"978-1-83880-729-0",doi:"10.5772/intechopen.77582",price:119,priceEur:129,priceUsd:155,slug:"sedimentary-processes-examples-from-asia-turkey-and-nigeria",numberOfPages:124,isOpenForSubmission:!1,hash:"e66ff69a772c2913167e9987180d7279",bookSignature:"Gemma Aiello",publishedDate:"May 27th 2020",coverURL:"https://cdn.intechopen.com/books/images_new/7768.jpg",keywords:null,numberOfDownloads:2266,numberOfWosCitations:0,numberOfCrossrefCitations:1,numberOfDimensionsCitations:3,numberOfTotalCitations:4,isAvailableForWebshopOrdering:!0,dateEndFirstStepPublish:"November 14th 2018",dateEndSecondStepPublish:"December 26th 2018",dateEndThirdStepPublish:"February 24th 2019",dateEndFourthStepPublish:"May 15th 2019",dateEndFifthStepPublish:"July 14th 2019",remainingDaysToSecondStep:"2 years",secondStepPassed:!0,currentStepOfPublishingProcess:5,editedByType:"Edited by",kuFlag:!1,biosketch:null,coeditorOneBiosketch:null,coeditorTwoBiosketch:null,coeditorThreeBiosketch:null,coeditorFourBiosketch:null,coeditorFiveBiosketch:null,editors:[{id:"100661",title:"Dr.",name:"Gemma",middleName:null,surname:"Aiello",slug:"gemma-aiello",fullName:"Gemma Aiello",profilePictureURL:"https://mts.intechopen.com/storage/users/100661/images/system/100661.jpg",biography:"Dr. Gemma Aiello was born in Aversa (CE), Italy, on 24 October\n1964. In 1989, she graduated in Geological Sciences at the University of Naples 'Federico II”. In 1993, she earned a PhD degree\nin Sedimentary Geology at the University of Naples 'Federico II”,\nDepartment of Earth Sciences, Faculty of Geological Sciences. She\ncompleted a 2-year postdoctoral fellowship at the University of\nNaples 'Federico II”, a CNR-CEE fellowship, and several contracts\nat the Research Institute 'Geomare Sud”, CNR, Naples, Italy. Since 1998, she has\nbeen a full-time researcher at the Italian CNR. Dr. Aiello has 25 years’ experience in\nthe field of sedimentary geology, marine geology, and geophysics, participating in\ndifferent research projects for the Italian National Research Council (CARG, Vector,\nCentri Regionali di Competenza). She was a contract professor of sedimentology and\nstratigraphy at the Parthenope University of Naples, Italy, and a teacher in formation\ncourses of technicians in marine science and engineering in Naples, Italy",institutionString:"Institute of Marine Sciences - National Research Council ISMAR-CNR",position:null,outsideEditionCount:0,totalCites:0,totalAuthoredChapters:"7",totalChapterViews:"0",totalEditedBooks:"4",institution:{name:"Institute for Coastal Marine Environment",institutionURL:null,country:{name:"Italy"}}}],coeditorOne:null,coeditorTwo:null,coeditorThree:null,coeditorFour:null,coeditorFive:null,topics:[{id:"667",title:"Geomorphology",slug:"sedimentology-geomorphology"}],chapters:[{id:"70756",title:"Introductory Chapter: An Introduction to Sedimentary Processes - Examples from Asia, Turkey, and Nigeria",slug:"introductory-chapter-an-introduction-to-sedimentary-processes-examples-from-asia-turkey-and-nigeria",totalDownloads:134,totalCrossrefCites:0,authors:[{id:"100661",title:"Dr.",name:"Gemma",surname:"Aiello",slug:"gemma-aiello",fullName:"Gemma Aiello"}]},{id:"67387",title:"Geologic Characteristics and Production Response of the N5.2 Reservoir, Shallow Offshore Niger Delta, Nigeria",slug:"geologic-characteristics-and-production-response-of-the-n5-2-reservoir-shallow-offshore-niger-delta-",totalDownloads:584,totalCrossrefCites:0,authors:[{id:"228567",title:"Dr.",name:"Prince Suka",surname:"Momta",slug:"prince-suka-momta",fullName:"Prince Suka Momta"}]},{id:"66556",title:"“Geo-archives of a Coastal Lacustrine Eco-system”: Lake Bafa (Mediterranean Sea)",slug:"-geo-archives-of-a-coastal-lacustrine-eco-system-lake-bafa-mediterranean-sea-",totalDownloads:258,totalCrossrefCites:0,authors:[{id:"138405",title:"Prof.",name:"M. Namik",surname:"Çagatay",slug:"m.-namik-cagatay",fullName:"M. Namik Çagatay"},{id:"290307",title:"Dr.",name:"Özlem",surname:"Bulkan",slug:"ozlem-bulkan",fullName:"Özlem Bulkan"},{id:"290457",title:"MSc.",name:"Bilgehan",surname:"Toksoy",slug:"bilgehan-toksoy",fullName:"Bilgehan Toksoy"}]},{id:"66126",title:"Fingerprinting Sources of the Sediments Deposited in the Riparian Zone of the Ruxi Tributary Channel of the Three Gorges Reservoir (China)",slug:"fingerprinting-sources-of-the-sediments-deposited-in-the-riparian-zone-of-the-ruxi-tributary-channel",totalDownloads:236,totalCrossrefCites:0,authors:[{id:"290401",title:"Dr.",name:"Zhonglin",surname:"Shi",slug:"zhonglin-shi",fullName:"Zhonglin Shi"},{id:"292470",title:"Dr.",name:"Dongchun",surname:"Yan",slug:"dongchun-yan",fullName:"Dongchun Yan"},{id:"292825",title:"Prof.",name:"Anbang",surname:"Wen",slug:"anbang-wen",fullName:"Anbang Wen"},{id:"292827",title:"Dr.",name:"Yongyan",surname:"Wang",slug:"yongyan-wang",fullName:"Yongyan Wang"}]},{id:"66184",title:"A Long-Term Prediction of Beach Changes around River Delta using Contour-Line-Change Model",slug:"a-long-term-prediction-of-beach-changes-around-river-delta-using-contour-line-change-model",totalDownloads:312,totalCrossrefCites:0,authors:[{id:"13491",title:"Dr.",name:"Takaaki",surname:"Uda",slug:"takaaki-uda",fullName:"Takaaki Uda"},{id:"122917",title:"Dr.",name:"Masumi",surname:"Serizawa",slug:"masumi-serizawa",fullName:"Masumi Serizawa"},{id:"287386",title:"Ms.",name:"Shiho",surname:"Miyahara",slug:"shiho-miyahara",fullName:"Shiho Miyahara"},{id:"288256",title:"Mr.",name:"Toshiro",surname:"San-Nami",slug:"toshiro-san-nami",fullName:"Toshiro San-Nami"}]},{id:"67292",title:"The Role of Mangroves in Coastal and Estuarine Sedimentary Accretion in Southeast Asia",slug:"the-role-of-mangroves-in-coastal-and-estuarine-sedimentary-accretion-in-southeast-asia",totalDownloads:743,totalCrossrefCites:1,authors:[{id:"216896",title:"Dr.",name:"Punarbasu",surname:"Chaudhuri",slug:"punarbasu-chaudhuri",fullName:"Punarbasu Chaudhuri"},{id:"290438",title:"Ms.",name:"Raktima",surname:"Ghosh",slug:"raktima-ghosh",fullName:"Raktima Ghosh"},{id:"290966",title:"Dr.",name:"Subhamita",surname:"Chaudhuri",slug:"subhamita-chaudhuri",fullName:"Subhamita Chaudhuri"}]}],productType:{id:"1",title:"Edited Volume",chapterContentType:"chapter",authoredCaption:"Edited by"},personalPublishingAssistant:{id:"287827",firstName:"Gordan",lastName:"Tot",middleName:null,title:"Mr.",imageUrl:"https://mts.intechopen.com/storage/users/287827/images/8493_n.png",email:"gordan@intechopen.com",biography:"As an Author Service Manager my responsibilities include monitoring and facilitating all publishing activities for authors and editors. From chapter submission and review, to approval and revision, copyediting and design, until final publication, I work closely with authors and editors to ensure a simple and easy publishing process. I maintain constant and effective communication with authors, editors and reviewers, which allows for a level of personal support that enables contributors to fully commit and concentrate on the chapters they are writing, editing, or reviewing. I assist authors in the preparation of their full chapter submissions and track important deadlines and ensure they are met. I help to coordinate internal processes such as linguistic review, and monitor the technical aspects of the process. As an ASM I am also involved in the acquisition of editors. 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Leptospirosis is a worldwide but more neglected zoonotic disease that usually affects humans and animals around the world [1, 2, 3] with case records 350,000 annually [4]. Death occurs from 10 to 50% in severe infection cases [5]. Pathogenic Leptospira are bacteria that cause the disease through penetration into the mucous membrane or damaged skin of the host [6]and then transfer to the proximal tubule of the kidneys [7]. Infected animals remain asymptomatic and secrete infectious organisms in urine throughout their lifetime [8]. Leptospira are excreted in the urine for weeks, months, or longer [9]. It pollutes moist soil, farm land, and rivers. Infection occurs when a person or animal is in intimate contact with a contaminated climate or infected urine of host [6]. The famous Leptospira carriers are wild, domestic, and some other mammalian species [10]. Occurrence of leptospirosis in humans is also associated with high temperature, high humidity, and basic hygienic conditions [11, 12]. Human leptospirosis is biphasic, with the general symptoms of pyrogens associated with acute or leptospiremic phase, continuing for about 1 week, and subsequent of recovery or immunization which is defined by antibody production [13]. In the past few decades, leptospirosis has become an infectious disease of urban environments especially in industrialized and developing countries. Rural areas are also affected with high mortality due to delay in diagnosis and lack of infrastructure with adequate clinical suspicion and other unknown causes like inherent pathogenicity of Leptospira strains [8]. Leptospirosis is very common in tropical and subtropical areas where people are very close to animals. Warm and humid environment favors in distribution and survival of pathogen [14]. Sporadic cases are reported throughout the year with incidence ranges from 0.1 to 10 per 100,000 people; and during the epidemic, it can reach over 50 per 10,000 people. Most cases are reported in Sri Lanka, India, Indonesia, Maldives, and Thailand. In the past, Sri Lanka (2008), Jakarta (2003), and Mumbai (2005) have been reported as epidemic areas of Southeast Asia [10]. The molecular classification of Leptospira in different species is described on DNA correlation base [3, 15]. The genome of Leptospira consists of two round chromosomes whose entire sequence has been established recently [8, 16]. The genome is larger than the genomes of other spirochetes, which shows the viability of Leptospira in different harsh environments [17]. Symptoms usually appear suddenly after an incubation period of 2–20 days, with a duration of 14 days at least. About 10% of patients diagnosed with leptospirosis having signs like Weil’s disease, which manifests itself as jaundice, kidney failure, and hemorrhagic in pulmonary arteries [18]. Leptospirosis is usually related to headache, fever with muscular pain in both adults and children. Drowsiness, vomiting, abdominal pain, diarrhea, cough, photophobia, arthralgia, and constipation may also occur [19].
Leptospirosis is a prominent communicable disease caused by spirochete bacteria. The bacterial species belong to genus Leptospira that have ability to cause a disease in a variety of wild and domestic animal bases [3]. Spirochetes bacteria are motile having hook form or question-mark shape and range in size from 6 to 20 μm in length and 0.1 μm in thickness [4]. Family Leptospiraceae includes genus Leptospira, which is further divided into two strains, that is, pathogenic and saprophytic [4, 15]. Pathogenic Leptospira have 21 species, 25 serogroups, and 250 serovars [3, 5]. Leptospira spp. are obligate aerobes having sluggish growth. Ideal growth temperature for Leptospira is 28–30°C [16]. There are different other characteristics of Leptospira like size, number of genes and pseudogenes, etc. (Figure 1). Serovars “pomona and grippotyphosa” are expectedly found to be the most prevalent candidates [17]. Leptospirosis in cattle is caused by Leptospira borgpetersenii and Leptospira interrogans serovar Hardjo, strains (well adapted to cattle) Hardjo bovis and Hardjo prajitno [18]. In Brazil as well as in Latin America, L. borgpetersenii serovar Hardjo strain Hardjo bovis were isolated from naturally infected cattle. Before this study, only serological studies had shown reactive animals with the serovar Hardjo in various countries [20, 21]. In Brazil, Chile, England, and Columbia, serovar Hardjo was found most prevalent among cattle [22]. Between 1988 and 2007 in France, serovar Serjoe (34%) was most common in cattle [23]. There are different reservoir hosts of Leptospira (Table 1).
Main characteristics of pathogenic and trophic genome Leptospira spp. [20].
Worldwide, the prevalence of animal leptospirosis is reported between 2 and 46% depending upon animal species [12]. More than 15 serogroups of Leptospira is observed and isolated from cattle, for example, icterohaemorrhagiae, canicola, pomona and grippotyphosa, etc. (Rocha). Seroprevalence of different serovars is different in all countries or regions. L. bratislava and L. grippotyphosa are more in Spain in those cattle which do not have good reproductive health [21]. Latin American countries, like Venezuela, have high prevalence of leptospirosis (80.51%) along with predominance of Sejroe serovar. Lesser incidence (2.6%) of disease has been observed in Peru during desiccated season when there are less chances for bacterial survival and transmission [24]. Similar results have been observed in Colombia and Mexico with the prevalence of 16.4–60.9% and 28.4–52%, respectively [25, 26]. Particularly, in countries like India, the bovine leptospirosis is highly prevalent, that is, up to 87% [27], 89.9% in Poland, and 88.2% in Mexico [28]. In contrary, certain states present lesser incidence, for example, 31.3% in Brazil [29], 27.4% in Australia [30], 30.3% in Tanzania [31], 20.3% in Sri Lanka [32], and 19.1% in Iran [33] (Table 2). These variations could be due to altered topographical localities, husbandry and farm management applications, infection immunity among diverse rears, and intensities of normal resistance [29]. In urban areas, leptospirosis is broadly prevalent, as stated by Platts-Mills [34].
Reservoir host | Serovar | Reference |
---|---|---|
Pigs | Pomona, Tarassovi | [8] |
Cattle | Hardjo, Pomona | |
Horse | Bratislava | |
Dog | Canicola | |
Sheep | Hardjo | |
Rats | Icterohaemorrhagiae, Copenhageni | |
Bats | Cynopteri, Wolffi |
Typical reservoir hosts of Leptospira.
Country | Year | Diagnostic methods | Specie | Prevalence% | Reference |
---|---|---|---|---|---|
India | 1983 2011 | — MAT | Cattle — | 68 87 | [35] [36] |
Malaysia | 1987 | MAT | Cattle Buffalo | 40.5 31 | [37] |
Sri Lanka | 2011 2014 | MAT Nested PCR | Cattle Cattle | 20.31 12.2 | [38] [6] |
Iran | 2011 | MAT | Cattle | 19.10 | [39] |
Pakistan | 2018 | Indirect ELISA | Cattle Buffalo | 25.52 20.72 | [40] |
Bangladesh | 2015 | ELISA | Cattle | 47.27 | [41] |
Prevalence of bovine leptospirosis in different countries of Asia.
Leptospirosis is termed as “storm of abortion” and is farm economy jeopardizing malaise [42]. Leptospira spread in direct and indirect ways, while the latter is a more pronounced method of transfer. Direct transmission involves through infected urine, post abortion uterine discharge, sexual contact, and infected placentae. Indirect involves contact with environment. Bacteria get entry to skin through abraded skin that follows hematogenous spread in the body. Bacteria result vasculitis that in turn results into either direct cytotoxic injury and immunological reactions or massive migration of fluid from intravascular to interstitial compartment. The latter results in renal dysfunction and vascular injury to internal organs. Pathogenic Leptospira could not be phagocytosed by macrophages and neutrophils, but if there are specific antibodies present, it can be phagocytosed [22]. It has been suggested that the animals are susceptible to severe or acute leptospirosis caused by increase in production of anti-inflammatory cytokines and chemokines [23]. Although pathogenic Leptospira are complementary to bactericidal activity, it has long been known that Leptospira has antimicrobial activity [43]. In most studies, leptospiral proteins that bind to one or more components are usually identified in recombinant form. Adhesin LenA (LfhA, Lsa 24) and Len B also bind to complementary regulatory protein factor H [44, 45]. Complement resistance to pathogenic Leptospira can also bind to the complement module C4BP, which catalyzes the cleavage of C4b [24]. It leads to decrease in surface deposition of subsequent components of the complement, where decaying species are not available. In subsequent studies, this activity was attributed to the new leptospiral proteins LcpA and Lsa30 [46]. Interestingly, ligand proteins that interact with many host ECM hosts and other proteins also interact with the complement regulators H, FHL-1, FHR1, and C4BP [25]. Surprisingly, the Leptospira elongation factor Tu shows a superficial effect and interaction with factor H, as well as binding to many purified host proteins, which leads to its diversity, the so-called “moonlighting protein” [25]. Most of the above studies provide indirect evidence of the role of Leptospira protein in the prevention of complement, but recently, it has been shown that the pathogenicity and non-urogenital fecundity of the three complement pathways, including factor B [26]. C2 and C4b are identified in the culture supernatant of the Leptospira cleavage component. In fact, inactivation of the above two proteins, Lig B and Len B, does not have a significant effect on pathogenicity [27, 28, 47]. Similarly, functional redundancy must also be considered: LenA and LenB. All proteins have structural and functional similarities to the endostatin of mammals [44].
The emergence of the mutagenesis system revealed a small number of Leptospira genes that encode the components necessary for the manifestation of pathogenicity. The first leptospiral protein, identified as virulence, is Loa 22, and the outer membrane protein containing the C-terminal OmpA domain appears to mediate the connection between the outer membrane and the peptidoglycan layer (mutant hama loa 22). The presence of a homolog of Loa 22 in L. biflexa has an indirect effect on pathogenicity. Since lipopolysaccharide (LPS) is a pathogenic factor for all Gram-negative bacteria, it is not surprising that this hypothesis cannot be obtained until a certain mutant is identified in the Leptospira, studies eliminate motility by inactivating flagellated structures or genes involved in biosynthesis. Inactivation of fliY-encoding choleretic switching protein reduces toxicity in guinea pigs [29]. Mutations in the gene encoding the sensory protein LB139 reduce motility and down-regulation of a number of chemotactic genes and weaken the mutant for hamsters [30]. The lack of similar genes in vegetative bacilli strongly suggests the survival of mammalian species as well as in nutrition intake. Indeed, studies have shown toxicity related activities of Leptospira sphingomyelinase such as pore formation and cytotoxicity [31]. However, the key role of leptospiral sphingomyelinase in the pathogenesis is not genetically established. Recently, it has been shown that Leptospira-LruA [48] plays an important role in autoimmune response, which is associated with mammalian apolipoprotein A1 [49].
Transmission of Leptospira occurs often with direct contact with infected urine, placenta, or milk. Transmission through venereal or transplacental route is also possible, whereas the most common route is infected urine. If there are leptospiral infected animals present in a dairy farm, the environment is also contaminated. Dairy feeder calves are probably the largest carriers of Leptospira in commercial feed yards. Dairy calves have the habit of sucking the scrotum of other calves in the pen, so this would be direct contamination of infected urine from carriers by suckling habit. Leptospira survives in the moist, damp, and moderately warm environment and can be easily killed by freezing, dehydration, and direct sunlight (Figure 2).
Mechanism of Leptospira.
Infection occurred by pathogenic leptospires is divided into two stages, first stage is acute stage or septicaemia (because septicaemia is in this stage), which lasts from 7 to 10 days with headache and myalgia. The second stage is immune stage which is after first week of infection and lasts 4–30 days [34]. During first stage, leptospires are present in blood and can say bacterial count is high in the blood, while when second stage starts, then the level of antibodies IgM and IgG start to increase and this increase in antibodies titer is correlated to elimination of leptospires from blood. Leptospira antigens and DNA sometimes may not be detected from the blood; this may be due to late sampling, or sampling in acute stage where proper level of leptospiremia is not developed and due to antibiotic administration, leptospires are eliminated from the blood. False negative results will be there, if we detect antibodies prior to sero-conversion during acute stage. Coagulated blood has serum and clot while non-coagulated blood has plasma, RBCs, WBCs and platelets. That can be collected according to the tests. If you are going to do gene amplification than EDTA, plasma gives the best results [50]. Leptospires can be detected in urine and cerebrospinal fluid samples. Many kits are available in the market for rapid detection of leptospires from blood, urine, and CSF sample; these kits basically detect nucleic acid of leptospires, but for these tests, purification of nucleic acid is required [51].
Different tools are being developed for the study of virulence factors, pathogenicity, and basic cell biology of organisms [52]. These are essential for proper treatment and reduction of the severity of the disease. During acute infection, nonspecific symptoms of leptospires mimic the febrile condition, which are essential for proper treatment and reduce the severity of the disease. Therefore, the diagnosis of leptospirosis is highly dependent on the particular laboratory tests [13]. Serology is the dominant one in diagnosis, while the micro-aggregation test (MAT) is the standard serological reference method. MAT is a sensitive test due to the antigenic heterogeneity of Leptospira, which require a large number of serovars as antigens. Furthermore, it is useless at early stage of the disease when antibodies are not present or present in less quantity [18]. Detection of disease in early stage helps the epidemiological investigators. However, antigen detection at this stage is more expensive and complex [13]. Current diagnostic tools for Leptospira detections other than MAT are rapid antibody-based tests, direct examination of blood, the rapid nucleic-acid diagnosis, [53], dark field microscopy (DFM), IgM ELISA, and polymerase chain reaction (PCR) [13].
This method is cheap, but for direct examination, dark field microscope is required [54]. Theoretically, leptospires may be diagnosed by direct examination of blood during first week after onset of symptoms. Leptospires are 6–20 μm long and their diameter is 0.15 μm. Because of their size, dark field microscopy is required; 10−2–10−6 leptospires/mL of blood may be observed during the acute stage of leptospirosis [55].
The use of PCR is increasing in recent years and it has replaced the serological methods in endemic areas, because it is more sensitive and has capacity to give early diagnosis. Real-time PCR is faster than regular PCR [56]. The threshold level in the urine or blood is 10–100 leptospires/mL (Figures 3 and 4) [50, 57].
Specificity of different diagnostic tests during acute phase of leptospirosis [56, 58].
Sensitivity of different diagnostic tests during acute phase of leptospirosis [58].
In recent years, many isothermal amplification techniques are developed like isothermal technique [59]. This technique can be used as alternative to the PCR. There is no need for constant maintenance of temperature at 60–65°C and no thermal recycler is required; so, these things make it best for developing countries. For this, an effective and specific amplification is performed by DNA polymerase and six primers in 1 hour under isothermal conditions. Now the amplified DNA can be easily detected by eye observation of fluorescence without using gel electrophoresis [60]. Loop-mediated isothermal amplification (LAMP) methods are recently developed for the quick diagnosis of pathogenic leptospires, and lipL41 and rrs are the genes targeted by LAMP. The specificity of these methods is weak because these can detect the threshold between 2 and 100 leptospires/reactive mixture [61].
This microscopic agglutination test is developed in Pasteur Institute. Dark field microscopy is required to see agglutination of live leptospires cultures with patient’s serum. This is the gold standard test for leptospirosis. It determines the anti-Leptospira immunoglobulin titers in human and animal serum at the serogroup level, so it is used for clinical and epidemiological investigations [62]. MAT is performed on micro titration plates, dilutions of serum which is collected from the patient is made and then equal volume of leptospiral culture is added to form agglutinations of distinctive patterns that consist of highly dense packs of partly intact leptospires. The test is read by DFM.
Normal ELISA is commonly used to diagnose leptospirosis. Enzyme immunoassay of leptospirosis can be performed using a commercially available kit or antigen obtained internally. Which is commonly used to detect IgM, and sometimes to detect IgG antibodies against leptospiral antigens. The presence of IgM antibodies indicates current or recent leptospirosis. The commercially available Leptospira IgM ELISA is used for the serological detection of acute leptospirosis infection in a patient’s serum sample. This ELISA is based on the principle that any Leptospira IgM antibody present in the patient’s serum binds to the Leptospira antigen that adheres to the microporous surface of the microwell. Residual serum was removed from these wells by washing with 1% buffer (included in the kit). Peroxidase-conjugated anti-human IgM is presented after adding to the wells, and the plate is reincubated so that the bound antigen-antibody complex binds to the conjugate. The wells are washed again and a colorless substrate system, tetramethylbenzidine hydroperoxide, is added. The substrate is hydrolyzed, and the chromogen is blue. When the reaction is stopped with phosphoric acid, TMB turns yellow. The development of color indicates the presence of visual acuity IgM antibody against Leptospira in serum samples [63].
Cows with acute leptospirosis are characterized by anemia, jaundice, hemoglobinuria, and lower lobe hemorrhage. An ulcer and bleeding may be present on the mucous membrane of the peritoneum. Pulmonary edema and emphysema are also common in cattle. Histologically, there is a progressive and diffuse interstitial nephritis and liver necrosis in the centre of the lobules. Sometimes the vascular lesions of the meninges are transferred to chronic infections. Leptospira can be seen in the silvery spots of a part of tortuous tubules proximal to the kidneys. In acute infection, there is minimal inflammation, and in the middle of the leaflet, there are only tubes filled with hemoglobin and visible liver necrosis. At a later stage, progressive interstitial nephritis is characterized by a small white cortical lesion, which initially slightly increases or decreases with increasing age of the lesion. The fruit of a broken cow is usually automated by the fact that there is no damage or bacteria. Even fresh fruit, positive identification of leptospirosis in lesions is not easy. Although the use of fluorescent antibody technology facilitates the identification of organisms, false positive results are common unless experienced diagnosticians interpret the test. Although dark field microscopy can be attempted, it is not suitable for tissue collected at dissection. Although PCR technology is important, in some cases, several primer sequences may be required.
Samples for confirmation of diagnosis are kidney, liver and placenta. Histology of kidney, liver, brain, heart, lungs and placenta can be performed. While for serological analysis heart blood serum or pericardial fluid from foetus can also be obtained.
The zoonotic potential of this organism should keep in mind during handling of carcasses and submitting specimens.
Treatment is based on severity of illness being presented by animal which in most of the cases is mild and self-limiting requiring no care. Other considerations, while treatment is considered, include differential diagnosis, cost, and availability of drugs. Treatment obtained based on in-vitro studies presented doxycycline, ampicillin, azithromycin or amoxicillin [64]. The double-blind randomized trials conducted on 29 patients produced promising results by reducing symptoms of malaise in 2 days preventing leptospiremia. The treatment, however, was not conclusive prevention from progression to severity [65]. Doxycycline or azithromycin is the drug of choice in endemic areas while contraindicated in pregnancy [64]. Sever cases are responsive to penicillin G sodium in studies conducted before 90s. The emerging resistance has narrowed spectrum of antibiotic use against infections [66]. Open randomized trial conducted with experiment involving 256 patients proved nonsignificant difference among penicillin G, cefotaxime, and doxycycline antibiotics [67]. Some of meta-analysis studies have reported nonsignificant difference between penicillin G and placebo on mortality [68]. Mortality is reported to increase up to 70% with pulmonary involvement which is due to immune-mediated inflammatory response. The therapeutic indicated for this complication is steroidal drugs. Early steroid administration was found responsive but methodologically flawed in various studies. Desmopressin was evaluated in various randomized studies as adjunct therapy with nonsignificant mortality benefits [69]. Therapy is considered beneficial with doxycycline or azithromycin along with steroid administration in mild and severe cases. Variations in studies are reported with nonsignificant benefits to mortality reduction.
Leptospirosis is a zoonosis with worldwide distribution. It is more prevalent in the developing countries. Hemorrhagic manifestations constitute the common clinical feature in leptospirosis [70]. In cattle, acute hemolytic syndrome of leptospirosis has been reported characterized by fever, icterus, anemia, and hemoglobinuria [71]. Without effective treatment, hemolytic syndrome in cattle may result in death. A high mortality rate of severe disease was determined to be associated with certain serotypes of Leptospira [72]. The disease, for instance, causes a decrease in erythrocyte and platelet counts, leading to anemia and hemorrhagic diathesis, respectively. Elevated bilirubin levels result from hemolysis and hepatorenal failure, indicating the characteristic nature of clinical signs [73]. Blood transfusion was reported to be quite effective in cases of life-threatening anemia in cattle. Previous reports suggest that timely transfusion of whole fresh blood be administrated to overcome severe hemolytic leptospirosis. Indeed, transfusion providing the vital components such as erythrocytes, platelets, and plasma contributes to repair the present collapses, that is, anemia, hemorrhagic diathesis, septicemia, and hepatorenal failure, in affected cattle [71]. A PCV value of 15% or less developing acutely may require transfusion, while chronic anemia can be tolerated in cattle without any transfusion [74].
The optimal control regime for leptospirosis is to prevent clinical disease and exfoliation in the urine in animals exposed to different serotypes of Leptospira. The most common method of controlling leptospirosis in cattle is vaccination and selective treatment. In addition, proper quarantine procedures should be implemented to prevent the introduction of hajo in the herd by buying infected animals. However, leptospirosis and wild animals gave rise as a carrier of the prevalence of serotypes of hardjo infections in cattle, mainly to prevent the overall impact of leptospirosis in most dairy products and Rieben beef failure. This is impossible. Thus, vaccination depends on an increase in the resistance of an animal to leptospiral serotype infection in this area. In all cases, the effort (buildings, under the control of rodents around swamps and creeks, for example, surround) must be made in order to limit direct and indirect contact between the cattle and Leptospirosetragern. In addition, proper quarantine procedures should be implemented to prevent the introduction of hajo in the herd by buying infected animals. However, leptospirosis and wild animals gave rise as a carrier of the prevalence of serotypes of hardjo infections in cattle, mainly to prevent the overall impact of leptospirosis in most dairy products and Rieben beef failure. This is impossible. Thus, vaccination depends on an increase in the resistance of an animal to leptospiral serotype infection in this area. The leptoral vaccine currently available for cattle in the United States is a 5-fold bacterial whole cell vaccine, including Pomona, Canicola, Icterohaemorrhagiae, Grippotyphosa, and Hardjo serotypes. These antigens can also be used in various combinations of other viral and bacterial vaccines. In the United States, a series of experimental studies and field data are available from the United States. Typical leptospirosis vaccines are Hajo kidney serotype infection, urinary tract infection or fetus (ha-ha type). This does not exclude the fact that the state indicates that the country is isolated from the United States. Many of the available Hardjo vaccines were approved many years ago in rigorous efficacy studies that mimic the natural route of exposure, and the last method to determine whether the Hardjo serotypes are infected with cattle is Hardjo stocks which did not use the serotype. However, recently two Hardjo vaccine serotypes have been widely studied using appropriate strains and methods. Compared to many other Hardjo serotype vaccines, these two products have shown excellent protection against infection and Hardjo hemoglobin isolation [75].
Leptospirosis is a major zoonotic disease resulting in high mortality in humans and animals. The disease is diagnosed clinically by fever, headache, vomiting, abdominal pain, and arthralgia. Leptospirosis is caused by more than 250 serovars, while pomona and grippotyphosa being the most prevalent serovars among them. However, among cattle, serovar Hardjo is the most important in causation of disease. Among the Asian countries, the highest prevalence of leptospirosis was found in India. Leptospirosis is mainly transmitted by direct contact with infected urine, and bacteria are mainly entered through ruptured skin. In house IgM ELISA is highly specific technique for Leptospira diagnosis. However, among the serological test, ELISA is more sensitive test for Leptospira diagnosis. Most effective treatment for Leptospira is doxycycline or azithromycin; however, former is not recommended in pregnancy. However, in severe cases, blood transfusion is also a best choice to save the life of animal. At last, the most effective way to control the disease is vaccination at early age of life following booster doses to avoid from more severe economic losses.
As part of secondary lymphoid organs, mucosa-associated lymphoid tissue (MALT) is an aggregate of unencapsulated lymphoid tissue that is located diffusely in the mucosa of the aerodigestive tract and consists of the tonsils, vermiform appendix, and Peyer’s patch [1]. As part of MALT, the tonsils serve as a protection ring including nasopharynx-associated lymphoid tissue (NALT), which is known as Waldeyer’s tonsillar ring, around the entrance of the upper aerodigestive tract to start the initial immunological barrier to infections [1, 2]. This annular-shaped lymphoid ring contains four types of tonsils in a fixed position [3].
Pharyngeal (adenoid) tonsil
Eustachian tube tonsils (Gerlach’s tonsils)
Lingual tonsils and lymphoid aggregations close to the epiglottis
The palatine (faucial) tonsils
Due to the close proximity of the palatine tonsil with the surrounding spaces including parapharyngeal, retropharyngeal, masticator, and parotid spaces, the tumors and inflammation of the tonsil commonly spread into these spaces and result in secondary lesions [4, 5]. A detailed knowledge of the surgical anatomic landmarks in the tonsillar region and the spaces around it is required for preoperative planning and to prevent iatrogenic complications.
The tonsils are lymphoepithelial organs acting as a guardian at the entrance of the upper aerodigestive tract. Lymphoid system, as a component of the immune system, consists of lymph vessels, nodes, and organs that regulate the immune response directly or indirectly. The lymph vessels play a key role in the drainage of interstitial fluid from the tissues to the blood and fat absorption, whereas the lymphoid organs mediate the proliferation and maturation of the cells of the immune system, which protect the body against ingested or inhaled foreign pathogens. The cell groups of the immune system, which is known as the ability to distinguish self from nonself, produce two reactions that are called the innate (natural) and adaptive (acquired) immunity [1, 2]. Initially, lymphocytes and accessory cells are developed and matured to the stage of antigen recognition in primary lymphoid organs including the thymus and bone marrow, and then they are activated and differentiated to effector cells of the immune response by antigen presentation in secondary lymphoid organs. The lymph nodes, the spleen, and mucosa-associated lymphoid tissue are secondary lymphoid organs which allow lymphocytes to become functional to produce a defense mechanism against microorganisms such as viruses, parasites, and bacteria. The structures of MALT have 70% of all the cells of the immune system, and the percentages of the lymphocytes in each of them are variable [1].
As part of the upper aerodigestive tract, the pharynx is located between the skull base and the inferior border of cricoid cartilage and consists of three portions; the nasopharynx (upper nasal), oropharynx (middle oral), and laryngopharynx (lower laryngeal). It is a musculomembranous tube covered with three external (circular, the superior, middle, and inferior constrictors) and three internal (longitudinal, voluntary) muscles, which play a key role in swallowing, respiration, and phonation [6]. The tonsils are located posterior to the nasal and oral portions of the pharynx to form a circumferential ring, known as the Waldeyer’s tonsillar ring, which was first described by German anatomist Heinrich Wilhelm Gottfried von Waldeyer-Hartz [3]. The unpaired nasopharyngeal and lingual tonsils and the paired palatine and tubal tonsils form Waldeyer’s lymphoid ring at the opening of the upper aerodigestive tract and are responsible for both innate and adaptive immunological responses which have a crucial role in the defense mechanism of the pharynx (Figure 1) [7].
The localization of the tonsils in Waldeyer’s tonsillar ring.
As a part of the upper respiratory system, the nasopharynx is bounded by the choanae anteriorly, the upper surface of the soft palate inferiorly, and the oropharyngeal isthmus (OPI) posterolaterally. The nasopharyngeal and tubal tonsils are located in the posterolateral wall of the nasopharynx [6]. The palatoglossal arch (PGa) and palatopharyngeal arch (PPa) join with each other to form the OPI which allows the communication with the oropharynx. The boundaries of the OPI are formed by the soft palate (velum) anteriorly and the lateral and posterior pharyngeal walls posterolaterally. During swallowing and speaking, the levator veli palatini (LVPm), palatopharyngeus (PPm), superior pharyngeal constrictor (SPCm), salpingopharyngeus (SPm), and the uvula, which are called a velopharyngeal sphincter, play a role in the closure of the OPI. During velopharyngeal function, a transverse mucosal ridge called Passavant’s ridge (palatopharyngeal sphincter) runs along the posterior wall of the OPI between the most lateral part (transverse fibers) of the PPm and the most superior part of the SPCm [7, 8].
Pharyngeal tonsil is the superior-most of the Waldeyer’s ring and located above the soft palate in the posterosuperior roof of the nasopharynx as a single median unencapsulated mass with 12–15 shallow, crypt-like invaginations. The pharyngeal bursa, a blind mucosal sac, may be seen in the posterior median wall of the nasopharynx above the SPCm. A median longitudinal groove extends from this sac inferiorly [6]. Anterosuperiorly, the pharyngeal tonsil is usually lined by pseudostratified ciliated columnar epithelium (respiratory epithelium), whereas posteroinferiorly the areas adjacent to the oropharynx is covered by stratified epithelium. These mucosal folds containing numerous lymphoid nodules commonly enlarge and become adenoid which results in respiratory difficulties and nasal obstruction during childhood and often start to involute after 7 years of age or even atrophied in the adult. Chronic inflammation of the pharyngeal tonsil results in hyperplasia and hypertrophy of the lymphoid tissue known as adenoid [7].
The arterial supply of it comes from ascending pharyngeal artery, pharyngeal branch of the maxillary artery, artery of the pterygoid canal, basisphenoid artery, ascending palatine, and tonsillar branch of the facial artery. It has a lymphatic drainage into upper deep cervical within the parapharyngeal space (PPS) and retropharyngeal lymph nodes [7].
Eustachian tube (ET) tonsils, small aggregates of lymphoid tissue, form the upper lateral aspect of the ring and are located bilaterally around the pharyngeal ostium of the ET (torus tubarius) which is below and in front of the pharyngeal recess (fossa of Rosenmüller) in the posterolateral wall of the nasopharynx [6]. Because of their close relationship to the torus tubarius, they are called tubal or Gerlach’s (German anatomist) tonsils. This triangular pharyngeal ostium has three prominences: anterior, posterior, and inferior. The anterior fold continues as a plica salpingopalatina descends into the soft palate. The posterior prominence is conspicuous and formed by the projecting cartilage of the auditory tube, called the torus tubarius, and also lies as plica salpingopharyngeus which is composed of the SPm. The torus tubarius can be used for ET catheterization. On the lower aspect of the ostium, the LVPm insertion forms a slightly rounded prominence [6, 7].
Tubal tonsils are covered by pseudostratified ciliated columnar epithelium with no crypts. They receive arterial supply via the ascending pharyngeal artery. Their lymphatic drainage is the same as the pharyngeal tonsil’s [6, 7].
The oropharynx extends from the OPI at the level of the soft palate to hyoid bone (C3 vertebra level). Anteriorly, the oropharynx communicates with the oral cavity via isthmus faucium which is limited by the PGa bilaterally, the uvula superiorly, and the posterior one third of the tongue that is in line with the sulcus terminalis inferiorly. According to the oncologic description, the oropharynx consists of four parts: the soft palate, the pharyngeal wall, the base of the tongue, and the palatine tonsillar fossa. So, a thorough understanding of the anatomy of oropharyngeal parts and adjacent structures is paramount in differential diagnosis and surgical interventions. It contains the palatine tonsils laterally and lingual tonsil in the retrolingual region anteriorly [6, 9].
Lingual tonsils are the inferior-most of the ring and composed of numerous lymphoid nodules in the posterior third of the tongue. The stratified squamous nonkeratinized epithelium covers this lymphoid tissue aggregates forming large, irregular protrusions. Also, they have less branching shallow crypts which are covered by the reticulated epithelium and mucous salivary glands which are drained through several ducts into these crypts which appear after birth [6, 9].
Vascular supply to the lingual tonsils is provided with the dorsal lingual branches of the lingual artery and vein. Efferent lymphatic vessels of the lingual tonsil passing through the pharyngeal wall drain into the deep cervical lymph nodes [6, 7, 9].
The palatine tonsils are two large, conspicuous almond-shaped mass of the lymphoid tissue forming the lower lateral aspect of the ring and localized in a triangular tonsillar fossa along the anterolateral border of the oropharynx on each side. The dimensions of the tonsils are about 10–15 mm in width and 20–25 mm in length in adults, but increase in children. The surface landmark of the tonsil corresponds to the lower part of masseter muscle in front of the angle of mandible [3, 6, 9]. The palatoglossal (anterior pillar) and palatopharyngeal (posterior pillar) mucosal folds diverge from the soft palate to form the boundaries of the tonsillar fossa, which lodges the palatine tonsils. These mucosal arches consist of the palatoglossal muscle (PGm) anteriorly and the PPm posteriorly. The palatine tonsil has two poles, upper and lower; two borders, anterior and posterior; two surfaces, medial and lateral; three mucosal folds, plica semilunaris, plica triangularis, and plica retrotonsillaris; and two depressions, supratonsillar and anterior tonsillar fossa [3, 6, 7].
Superiorly, the tonsil is free and expands into the soft palate where both arches join.
Inferiorly, the suspensory ligament, a band of fibrous tissue, connects the lower pole with the posterior one third of the tongue. Most of carcinomas develop in the tonsillolingual sulcus which separates the tonsil from tongue anteroinferiorly [3, 6].
The tonsillar fossa or sinus is a triangular space between the anterior pillar in front, the posterior pillar behind, and the dorsal surface of the posterior one third of the tongue inferiorly (Figure 2). Because the tonsils are positioned in it, its borders also limit the tonsil [7].
The mucosal folds and arches of the palatine tonsil.
The anterior boundary is formed by the PGa which is composed of the PGm. A cylindrical muscle extends from the palatine aponeurosis to the posterolateral surface of the tongue and becomes continuous with the intrinsic transverse muscles [6, 7]. It acts as an antagonist of the LVPm and constricts the OPI during swallowing. All of the muscles of the tongue derive from the occipital myotomes except the PGm which is derivation of the fourth branchial arch. According to the variations of the origin of the PGm, the tongue elevator’s function increases or decreases. During lateral pharyngoplasty, the relaxation of the SPCm and PGm is provided by the myotomy of these muscles [3, 10].
The posterior boundary is formed by the PPa including the PPm which originates from the palatine aponeurosis and the median part of soft palate by two heads and consists of muscle bundles medial and lateral to the LVPm. The lateral fibers of the PPm are composed of the longitudinal and transverse parts. The transverse part inserts into the pharyngeal raphe to join with the contralateral side, whereas the longitudinal part joins with the medial fibers at the posterior border of the soft palate and afterward are merged by the SPm [7, 8, 11]. This muscle bundle is observed to course downward along the inner surface of pharyngeal wall and inserts into the palatal tonsil to form the posterior pillar. Also, some of its fibers insert into the posterior border of the thyroid cartilage with the stylopharyngeus muscle (StPm) and into the SPCm [11].
During velopharyngeal closure, the PPm performs various functions such as a sphincter with the SPCm, a puller of the pharyngeal wall medially in collaboration with the SPCm and StPm, and an elevator with the StPm because of the fibers of the PPm running in various directions [8, 11].
In the 14th–15th week of gestation, the primitive tonsil and tonsillar fossa develop indirectly from the endoderm part of the second pharyngeal arch. At first, the tonsil has two lobes and a plica intratonsillaris (intratonsillar cleft) between them. This plica later usually disappears, but it may infrequently transform into crypta magna [6, 12]. Because the tonsil does not completely fill this fossa, two small depressions exist at the upper and anteroinferior parts of the tonsillar fossa. They are separated from the tonsil by mucosal folds, known as the plica semilunaris and triangularis, which are remnants of the primitive tonsillar fossa (Figure 2) [6, 7].
Superiorly the plica semilunaris originates from the upper aspect of the PGa and extends backward toward the PPa along the upper pole of the tonsil. It encloses a small depression that is known as supratonsillar fossa which separates the tonsil from the uvula [6, 7].
Anteroinferiorly the plica triangularis, an inconstant mucosal fold, arises from the PGa and covers the anteroinferior part of the tonsil. It encloses a smaller fossa that is known as anterior tonsillar fossa, which is then obscured by its walls and forms the imbedded portion of the tonsil [6, 7].
Also, the plica infratonsillaris or retrotonsillaris may extend to the PPa at the posteroinferior part of the tonsil [7]. At first there is no lymphoid tissue in these fossae, but in childhood, they are usually transformed into lymphoid tissues, which are an exclusive hiding place for a constant lithified secretion and foreign bodies, causing an inflammation or quinsy [6, 7].
Medial surface is the free mucosal part of the tonsil that faces the oropharynx and contains bulging lymphoid projects. It is lined by stratified squamous nonkeratinized epithelium which contains polygonal superficial cells with microridges and numerous tubule-like long invaginations or orifices leading into tonsillar crypts. There are about 10–30 branching (primary and secondary) and anastomosing crypts, small pores, ranging in size between 5 and 25 μm. They increase the surface area of the tonsil up to 300 cm2 except the anterior part for interactions between antigens and the nodular lymphoid tissue. Secondary crypts are branching part of the primary crypts and continue deeply into the lymphoid tissue and forms the lymphoid fronds. The largest and deepest crypt is called crypta magna or intratonsillar cleft which is localized near the upper part of the tonsil [6, 7, 9].
The transitional type nonkeratinized stratified epithelium, reticulated lymphoepithelium, with a discontinuous basement membrane covers the crypts with fenestrated capillaries and represents pores that are filled with large oval microvillus cells (M cells or dendritic cells) or lymphocytes (T and B cells). Dendritic cells play a role in the uptake and transport of antigens to extrafollicular T cell and B cell follicles [9, 12].
At about 5th month of gestation, there are no germinal centers, and the lymphocytes develop from the connective tissue cells or are relocated in the blood and lymph vessels [13]. After birth, the exogenous antigens cause immune response which is represented by the transformation of effector B cell into extrafollicular plasma cell in 2 weeks, and secondary follicles containing active germinal centers develop and rapidly expand not invade the surrounding tissue in the first decade of life [7, 12]. The tonsillar lymphoid follicles consist of the lymphoid (germinal centers) and non-lymphoid cells (reticular cells and dendritic cells/macrophages). The germinal center is composed of a central area of proliferating B cells which is surrounded by resting B and T cells. Between these follicles, high endothelial venules allow the entrance of T and B cells from the blood and the release of mature lymphocytes into blood [6, 9, 13]. So, the tonsils have efferent lymphatic vessels to connect to lymph nodes, but no afferent vessels unlike a lymph node. The lymphoid fronds are separated from the tonsillar bed by a capsule, which is firmly coherent to the lymphoid tissue by multiple septa or trabeculae that dissect the tonsillar parenchyma. The trabeculae consist of elastin fibers and reticular fibers that are composed of type III collagen and provide cytoskeletal support. So, each tonsil is in a fixed position, in contrast to other MALTs, which are distributed throughout the body, and to disconnect the tonsil from its capsule is impossible. Also, the nerves, lymphatic and blood vessels, pass through the trabeculae [6, 9, 12].
The tonsils are most immunologically active at 4–10 years of age, whereas the adenoids are at 4–6 years. Age-dependent involution of the tonsil which refers to the regression of the germinal centers and the proliferation of fibrous tissue including the capsule and trabeculae occurs by adolescence. Also, fat deposition in tonsils starts and increases after 25 years of age [12, 13].
Lateral surface is a base of the tonsil that is covered by well-defined fibrous capsule at the lateral wall of the tonsillar fossa, which is composed of five layers from within outward (Figure 3):
Tonsillar capsule, a thin false or surgical sheet, covers the tonsillar fossa as an appendage of the pharyngobasilar fascia. The upper part of this condensed capsule is even and loosely fixed, whereas the lower part is irregular and intermingled with the pharyngeal muscle fibers and is firmly attached anteroinferiorly by the suspensory ligament to the posterior one third of the tongue. The tonsillar artery enters near this ligament. So, the surgical removal of the upper part of the capsule up to the lower third is very easy, but the lower part requires cautious resection [6, 7, 14].
Loose areolar tissue refers to the peritonsillar space between the tonsillar capsule and the pharyngobasilar fascia and contains the paratonsillar veins. A collection of pus in this space result in peritonsillar abscess or quinsy. It allows free movement of the pharyngeal muscles in the bed and makes easy to dissect the tonsil with capsule during tonsillectomy [6, 7].
Pharyngobasilar fascia or pharyngeal aponeurosis originates from the pharyngeal tubercle and covers the first layer of the SPCm and is limited with the inferior fibers of the muscle. Efferent lymphatic vessels from the tonsil pierce through the buccopharyngeal fascia [7, 15].
The lateral wall of tonsillar fossa or tonsillar bed is mostly made up of the SPCm and pharyngobasilar fascia superiorly, the StPm posteriorly, and the stylohyoid ligament, middle pharyngeal constrictor (MPCm), the glossopharyngeal nerve (GPn), and styloglossus (StGm) muscles anteroinferiorly [7, 14, 15].
The SPCm narrows the superior part of the pharynx and is composed of four portions depending on their origins;
The pterygopharyngeal portion originates from the posterior margin of the medial pterygoid plate and pterygoid hamulus.
The buccopharyngeal portion arises from the pterygomandibular raphe.
The mylopharyngeal portion originates from the posterior end of the mylohyoid line of the mandible.
The glossopharyngeal portion arises from the side of the tongue.
All of the muscle fibers are inserted into the median pharyngeal raphe posteriorly [7, 11]. Frequently, there is a space of 1–3 cm between the SPCm and MPCm. The GPn between the stylohyoid ligament and StGm curve forward and medially and pass through this space at the level of the lower pole of the palatine tonsil. The StGm and stylohyoid ligament originate from the anterior margin of the styloid process near its apex. The StGm inserts into the inferolateral surface of the tongue and interdigitates with intrinsic longitudinal lingual muscle, whereas the stylohyoid ligament lies between the StPm and StGm and attaches to the hyoid bone medially [7, 14, 16]. The StGm functions to elevate and retract the base of the tongue. Inferolaterally the lingual artery crosses the StGm and gives the dorsal lingual branches medial to the attachment of the StGm to the base of tongue [16].
At the junction of pharyngeal constrictor muscles beneath the tonsil, the GPn gives tonsillar branch and afterward, extends into the base of tongue between the StGm and the stylohyoid ligament posteroinferiorly. The StPm originates from the posterior margin of the styloid process and courses downward along the posterolateral part of the stylohyoid ligament. Between the SPCm and MPCm, it passes and inserts to the PPm, MPCm, and pharyngeal mucosa [14, 17].
The buccopharyngeal fascia covers the lateral aspect of the SPCm medially and the medial pterygoid muscle anterolaterally. It forms anteromedial wall of the PPS and contains the pharyngeal plexus of nerves and vessels. The PPS like an inverted pyramid is situated between the lateral pharyngeal wall and the pterygoid musculature (Figure 3) [4, 15].
The layers of the lateral pharyngeal wall at the level of tonsillar fossa, the parapharyngeal space compartments, and the structures between external and internal carotid arteries: SPm, salpingopharyngeus muscle; SPCm, superior pharyngeal constrictor muscle; PPm, palatopharyngeus muscle; PGm, palatoglossal muscle; StPm, stylopharyngeus muscle; StHm, stylohyoid muscle; StGm, styloglossus muscle; MPm, medial pterygoid muscle; GPn, glossopharyngeal nerve; PPS, parapharyngeal space; ICA, internal carotid artery; ECA, external carotid artery; X, vagus nerve; XI, accessory nerve; XII, hypoglossal nerve.
The base of the parapharyngeal pyramid is located at the skull base and its apex at the greater cornu of the hyoid bone. The PPS is bounded by the following structures:
The buccopharyngeal fascia which covers the SPCm, the LVPm, and tensor veli palatini muscles medially,
The fascia overlying the masticator space, the medial pterygoid muscle, the sphenomandibular ligament, the ramus of the mandible, and the deep lobe of the parotid gland anterolaterally,
The styloid process, the StGm and StPm posterolaterally,
The pterygomandibular raphe between the medial pterygoid plate and the mylohyoid line of the mandible and interpterygoid fascia anteriorly,
The prevertebral fascia and muscles posteriorly.
Inferiorly, the direct communication of the PPS with the submandibular space may be seen at the apex [4, 7, 10, 15].
Prasad et al. reported that the PPS is composed of three compartments as follows: the upper part of the PPS is located between the skull base and the axial plane passing through the inferior border of the lateral pterygoid muscle, the lower border of the middle part is formed by the axial plane passing through the mandibular insertion of medial pterygoid muscle, and the lower part is limited with the hyoid bone. The middle part of the PPS is situated at the level of the tonsillar fossa. Also, the upper and middle parts are divided into prestyloid and poststyloid compartments in relation to the styloid diaphragm. Thus, the PPS consists of five parapharyngeal subspaces [18].
The styloid diaphragm is a thick gray fascia which is composed of the posterior belly of the digastric muscle, the styloid musculature (StPm, StGm and stylohyoid muscle-StHm), and the stylohyoid and stylomandibular ligaments. It divides the lower PPS into the prestyloid and poststyloid compartments by extending from the styloid process to the parotid fascia (Figure 3). The prestyloid space is localized between the medial pterygoid muscle and SPCm [7, 15, 18].
In the prestyloid part of the upper PPS, minor salivary glands, the posterior division of the mandibular nerve, the internal maxillary artery, fat pad, and tensor veli palatini muscle are located. In the poststyloid part of the upper PPS, the carotid sheath which consists of the internal carotid artery (ICA), internal jugular vein (IJV), vagus nerve, and also just in this superior section the ascending pharyngeal artery, cervical sympathetic chain, and the lower cranial nerves, IX, XI, and XII, are situated [7, 18].
In the prestyloid part of the middle PPS, the fat pad, a deep lobe of the parotid gland, from superior to inferior numerous tonsillar branches of the descending palatine, the ascending pharyngeal, and the ascending palatine arteries between the StGm and StPm are located. In the poststyloid part of the middle PPS, the curves of the internal maxillary, facial, and lingual arteries, cervical sympathetic chain, and the carotid sheath which consists of the ICA, IJV, and the lower cranial nerves (CNIX–CNXII) are situated [5, 7, 18].
Different surgical procedures can be used in treatment of the upper airway obstruction due to tonsillar or adeno-tonsillar hypertrophy and peritonsillar abscess. Classic tonsillectomy consists of full removal of the tonsil with its capsule by dissecting the peritonsillar space with or without adenoidectomy. In the post-acute tonsillitis, a peritonsillar abscess may spread into the PPS through the buccopharyngeal fascia. Due to the close proximity of the PPS with the surrounding spaces including pharyngeal mucosal, retropharyngeal, masticator, and parotid spaces, the lesions in these spaces commonly spread into the PPS and result in secondary lesions [4, 5].
Sun et al. reported that the localization of the tumors in the PPS can be identified by some anatomical landmarks during surgical approaches. Because of the tumors in the upper PPS are mostly benign and located in the prestyloid space, the endoscopic transnasal transpterygoid approaches to this region require detailed anatomic knowledge of the surgical anatomic landmarks in this space. They demonstrated that the surgical anatomic landmarks in the prestyloid part of the upper PPS are as follows: the pterygoid process with medial and lateral plates, the tensor veli palatini, the SPCm, the lateral and medial pterygoid muscles, and the fat pad. In the prestyloid part of the lower PPS, the PGa, the SPCm, the pterygomandibular raphe, fat tissue, and the styloid diaphragm could be used as surgical anatomical landmarks during endoscopic transoral approach (Figure 3) [19].
Approximately 80% of primary oropharyngeal tumors originate from the tonsillar fossa and their incidence in younger patients increases. The tumors in the tonsillar fossa and the PPS can be removed by endoscope-assisted lateral oropharyngectomy approaches, transoral robotic surgery, or laser microsurgery. The lateral pharyngeal wall is composed of three deep fascia layers from inward to outward: the capsule of the tonsil, the pharyngobasilar fascia, and the buccopharyngeal fascia [5, 15, 19]. Depending on these fascia layers, De Virgilio et al. reported their lateral oropharyngectomy classification based on three types of surgical procedures and four possible extensions (superior, soft palate; posterior, pharyngeal wall; inferior, base of the tongue; anterior, retromolar trigone) [5].
Type 1 contains the removal of the palatine tonsil deep to the pharyngobasilar fascia with the resection of all or part of the anterior pillar excluding the SPCm. The aim of this procedure is mostly diagnostic, but it can be used in surgical treatment of noninvasive hyperplasia, dysplasia, or carcinoma in situ of the tonsil.
Type 2 is resection of the palatine tonsil, the PGm, the PPm, and the SPCm deep to the buccopharyngeal fascia. It can be therapeutic for invasive malignant tumors not grossly infiltrating the SPCm.
Type 3 includes the resection of the buccopharyngeal fascia with extension to the pterygoid muscle and PPS adipose corpus in addition to Type 2 contents. According to the extension of the tumor, the resection of the PPS tissue up to the exposure of the ICA could be included, and also a flap coverage for the ICA is required [5].
Similarly, Mirapeix et al. identified an applicable dissection method based on the anatomic stratification and evident anatomic landmarks [4]. They performed the dissections layer by layer from within outward and described this technique by dividing the lateral oropharyngeal wall into three layers:
The first layer, medial to styloid muscles, includes important surgical landmarks such as the SPCm, PGm, PPm, and StGm, the pharyngobasilar fascia, and a vascular network, which is composed of the branches of the descending and ascending palatine arteries and the ascending pharyngeal artery. The vascular supply of the tonsillar fossa can be identified by the PGm and PPm, and also the lingual branch of the GPn mostly crosses at the midpoint between PGm and PPm or along the posteroinferior edge of the StGm.
The second layer is observed after resection of the constrictor muscles and located in the PPS medial to the styloid diaphragm. The surgical landmarks are composed of the styloid musculature, the buccopharyngeal fascia, the stylohyoid ligament, the pharyngeal venous plexus, and the GPn. The insertion point of the StGm refers to junction of the tongue with anterior pillar, and the lingual branch of the GPn can be identified along the posteroinferior border of the StGm. The pharyngeal venous plexus is located in a space between the StGm and SPCm. The facial artery and the hypoglossal nerve cross the StHm which extends parallel to the stylohyoid ligament. The GPn travels downward along the posterolateral aspect of the StPm.
The third layer lateral to styloid diaphragm refers to the poststyloid part of the PPS. Surgical landmarks in this layer consist of the styloid musculature, the posterior belly of the digastric muscle, the ICA, the hypoglossal nerve, and lingual and facial arteries. Especially, the StGm is an essential landmark to identify the localization of the ICA posterolaterally, the lingual nerve anteriorly, and the submandibular gland inferolaterally. The hypoglossal nerve crosses laterally to medially over the ascending pharyngeal originating from the superolateral border of the external carotid artery (ECA) in the poststyloid part of the lower PPS [4].
During transoral robotic surgery (TORS), the dissection of the SPCm from the pterygomandibular raphe refers to a window into the prestyloid compartment of the PPS. The tendon of the medial pterygoid muscle leads to identification of the buccopharyngeal fascia and indicates a safe plane in the prestyloid compartment of the PPS [7, 19]. Also, the plane that is constituted by the styloid musculature and the stylohyoid ligament is an essential surgical landmark for ICA identification. Wang et al. demonstrated that the styloid process, styloid diaphragm, pharyngeal venous plexus, GPn, and pharyngeal branch of the vagus are located between the ECA and the ICA and subdivide the PPS into prestyloid and poststyloid spaces (Figure 3). The curves of the branches of the ECA (lingual, facial, ascending pharyngeal, internal maxillary arteries) are located in the prestyloid space, and also the ascending pharyngeal artery crosses the StGm at the distal third near the tonsillar fossa surgical field [20]. In addition, the lingual artery and hypoglossal nerve are located lateral to the StGm, and the lingual artery passes between greater cornu of hyoid and the StGm where it has high risk of hemorrhage during the resection of the base of the tongue [21]. The fact that in the PPS the facial artery is located inferolateral to the StGm is of great importance, because a dissection lateral to the StGm or resection of tonsillar malignancy may result in significant hemorrhage. In PPS after branching from the facial artery, the tonsillar and ascending palatine arteries course between the StGm and the StPm and then pierce the SPCm to supply the tonsil [22]. So, the fact that the StGm is in close relationship with the branches of the ECA should be kept in mind when the transoral dissection space at the level of the tonsillar fossa is dissected in the superolateral direction, and the dissection deep into the plane of this muscle must be performed rigorously and accurately [16, 20].
In the PPS lateral to the styloid diaphragm, the ICA lies about 10–20 mm behind the palatine tonsil at the level of the epiglottis apex, whereas its distance to the ET is approximately 23.5 mm. So, it is closer to the lateral pharyngeal wall in the poststyloid part of the lower PPS than in the upper PPS, and the risk of arterial trauma during tonsillectomy increases with a decrease in the distance to the pharyngeal wall. Also, the level of the common carotid artery bifurcation higher than the epiglottis apex is more susceptible to common carotid artery trauma during surgery [23]. During radical tonsillectomy, because the lingual nerve lies lateral to the SPCm, it may be injured at the anterior border of the medial pterygoid muscle [15].
The GPn extends from the jugular foramen to the base of the tongue in the lateral wall of the pharynx. Because of the close relationship of the GPn with the StPm, it is divided into three parts: upper (jugular foramen, upper border of the StPm), middle (upper-lower borders of the StPm), and lower (lower borders of the StPm, the base of the tongue) [17].
The upper part travels between the ICA and IJV behind the styloid process and gives the carotid body and carotid sinus branches in the poststyloid part of the upper PPS.
The middle part extends downward along the inferolateral border of the StPm and gives off branches to the StPm and pharyngeal wall in the poststyloid part of the lower PPS. Particularly, this part passes obliquely anterior to the distal segment of the ICA and may result in vascular injury.
The lower part passes through a space or slit between the SPCm and MPCm to enter the pharynx. Between the StGm and StPm, it lies along the inferior border of the palatine tonsil or beneath the capsule and gives the tonsillar branch. Generally, it gives terminal branches at the junction of the PPa with the base of tongue, known as glossotonsillar sulcus, which is anatomic landmark for the terminal part of the GPn deep to the SPCm [17].
During surgical interventions including transoral tonsillectomy, tumor resection, and the SPCm block, the integrity of this nerve may be damaged and result in dysphagia and taste disturbance. In recurrent tonsillitis, the adherence of the capsule with surrounding structures makes it difficult to remove the hypertrophic tonsillar capsule from the tonsillar bed, or the dissection of the capsule which is firmly adherent with the lingual branch of this nerve causes disturbance of the nerve functions [3, 7, 17]. During transoral surgery, early description of the StPm allows to specify the GPn which crosses over the ICA and serves as a surgical landmark to protect it in the PPS. Also, the surgeon should keep in mind the association of the GPn with a venous plexus in the glossotonsillar sulcus to prevent iatrogenic bleeding during surgical dissection.
The tonsil and tonsillar fossa with boundaries are supplied by the branches of the ECA including lingual, facial, ascending pharyngeal, and internal maxillary arteries (Figure 4).
The vascular supply of the tonsil: PPm, palatopharyngeus muscle; PGm, palatoglossal muscle; SPCm, superior pharyngeal constrictor muscle.
The upper part is supplied by descending palatine artery branch of the internal maxillary artery and the middle and inferior branches of the ascending pharyngeal artery.
The middle part is supplied by tonsillar branch of the facial artery.
The lower part is supplied by an ascending palatine artery branch of the facial artery and dorsal lingual branch of the lingual artery [20, 23].
The veins of the tonsil and tonsillar fossa drain into the paratonsillar vein and then into the pharyngeal venous plexus. This plexus drains through the facial vein into the IJV (Figure 4) [6, 7].
Benign or malign lesions in the tonsil and tonsillar fossa may penetrate the lateral wall of the pharynx, or the PPS may be distorted evidently by the tumors. Due to the anatomical complexity with vital neurovascular structures in the PPS, transoral robotic approach to this region makes it necessary to identify the surgical anatomic landmarks which are required to perform effective surgical intervention quickly and accurately. The detailed and precise anatomic knowledge of the tonsillar region and the PPS allows surgeon to carry out wide resections in a confined space. In transoral approaches, the classification of the dissection method based on the anatomic stratification or the surgical procedures which is oriented to cardinal points is essential for preoperative planning and to prevent the iatrogenic complications.
The author reports no conflict of interest concerning the materials used in this paper. And the author has no personal financial or institutional interest in this article.
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