List of intron retention genes and their functions.
\r\n\tThe aim of this book is to provide the reader with a comprehensive state-of-the-art in artificial neural networks, collecting many of the core concepts and cutting-edge application behind neural networks and deep learning.
",isbn:"978-1-83962-375-2",printIsbn:"978-1-83962-374-5",pdfIsbn:"978-1-83962-376-9",doi:null,price:0,priceEur:0,priceUsd:0,slug:null,numberOfPages:0,isOpenForSubmission:!1,hash:"5cc6cd7972551be6cfc4d3c87bf8fb5c",bookSignature:"Dr. Pier Luigi Mazzeo and Dr. Paolo Spagnolo",publishedDate:null,coverURL:"https://cdn.intechopen.com/books/images_new/10390.jpg",keywords:"Recurrent, Recursive Nets, Face Recognition, Crowd Analysis, Different Applications, Object Detection, Classification, Visual Tracking, Speech Recognition, Grams, Reinforcement Learning, 3-D Map",numberOfDownloads:393,numberOfWosCitations:0,numberOfCrossrefCitations:0,numberOfDimensionsCitations:0,numberOfTotalCitations:0,isAvailableForWebshopOrdering:!0,dateEndFirstStepPublish:"September 25th 2020",dateEndSecondStepPublish:"October 23rd 2020",dateEndThirdStepPublish:"December 22nd 2020",dateEndFourthStepPublish:"March 12th 2021",dateEndFifthStepPublish:"May 11th 2021",remainingDaysToSecondStep:"6 months",secondStepPassed:!0,currentStepOfPublishingProcess:5,editedByType:null,kuFlag:!1,biosketch:"Author and co-author of more than 80 works in national and international journals, conference proceedings, and book chapters, with Ph.D. in Computer Science Engineering.",coeditorOneBiosketch:"Dr. Spagnolo received the engineering degree in computer science from the University of Lecce, Italy. Since 2002 he has been with the Italian National Research Council. His work includes more than 80 publications on AI.",coeditorTwoBiosketch:null,coeditorThreeBiosketch:null,coeditorFourBiosketch:null,coeditorFiveBiosketch:null,editors:[{id:"17191",title:"Dr.",name:"Pier Luigi",middleName:null,surname:"Mazzeo",slug:"pier-luigi-mazzeo",fullName:"Pier Luigi Mazzeo",profilePictureURL:"https://mts.intechopen.com/storage/users/17191/images/system/17191.jpeg",biography:"Pier Luigi Mazzeo received the engineering degree in computer science from the University of Lecce, Lecce, Italy, in 2001. \nSince 2015 he has been with Institute of Applied Sciences and Intelligent Systems of the Italian National Research Council, Lecce, Italy. The most relevant topics, in which he is currently involved, include algorithms for video object tracking , face detection and recognition, facial expression recognition, deep neural network (CNN) and machine learning.\nHe has taken part in several national and international projects and he acts as a reviewer for several international journals and for some book publishers. 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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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Venkateswarlu",coverURL:"https://cdn.intechopen.com/books/images_new/371.jpg",editedByType:"Edited by",editors:[{id:"58592",title:"Dr.",name:"Arun",surname:"Shanker",slug:"arun-shanker",fullName:"Arun Shanker"}],productType:{id:"1",chapterContentType:"chapter",authoredCaption:"Edited by"}},{type:"book",id:"72",title:"Ionic Liquids",subtitle:"Theory, Properties, New Approaches",isOpenForSubmission:!1,hash:"d94ffa3cfa10505e3b1d676d46fcd3f5",slug:"ionic-liquids-theory-properties-new-approaches",bookSignature:"Alexander Kokorin",coverURL:"https://cdn.intechopen.com/books/images_new/72.jpg",editedByType:"Edited by",editors:[{id:"19816",title:"Prof.",name:"Alexander",surname:"Kokorin",slug:"alexander-kokorin",fullName:"Alexander Kokorin"}],productType:{id:"1",chapterContentType:"chapter",authoredCaption:"Edited by"}},{type:"book",id:"314",title:"Regenerative Medicine and Tissue Engineering",subtitle:"Cells and Biomaterials",isOpenForSubmission:!1,hash:"bb67e80e480c86bb8315458012d65686",slug:"regenerative-medicine-and-tissue-engineering-cells-and-biomaterials",bookSignature:"Daniel Eberli",coverURL:"https://cdn.intechopen.com/books/images_new/314.jpg",editedByType:"Edited by",editors:[{id:"6495",title:"Dr.",name:"Daniel",surname:"Eberli",slug:"daniel-eberli",fullName:"Daniel Eberli"}],productType:{id:"1",chapterContentType:"chapter",authoredCaption:"Edited by"}},{type:"book",id:"57",title:"Physics and Applications of Graphene",subtitle:"Experiments",isOpenForSubmission:!1,hash:"0e6622a71cf4f02f45bfdd5691e1189a",slug:"physics-and-applications-of-graphene-experiments",bookSignature:"Sergey Mikhailov",coverURL:"https://cdn.intechopen.com/books/images_new/57.jpg",editedByType:"Edited by",editors:[{id:"16042",title:"Dr.",name:"Sergey",surname:"Mikhailov",slug:"sergey-mikhailov",fullName:"Sergey Mikhailov"}],productType:{id:"1",chapterContentType:"chapter",authoredCaption:"Edited by"}},{type:"book",id:"1373",title:"Ionic Liquids",subtitle:"Applications and Perspectives",isOpenForSubmission:!1,hash:"5e9ae5ae9167cde4b344e499a792c41c",slug:"ionic-liquids-applications-and-perspectives",bookSignature:"Alexander Kokorin",coverURL:"https://cdn.intechopen.com/books/images_new/1373.jpg",editedByType:"Edited by",editors:[{id:"19816",title:"Prof.",name:"Alexander",surname:"Kokorin",slug:"alexander-kokorin",fullName:"Alexander Kokorin"}],productType:{id:"1",chapterContentType:"chapter",authoredCaption:"Edited by"}},{type:"book",id:"2270",title:"Fourier Transform",subtitle:"Materials Analysis",isOpenForSubmission:!1,hash:"5e094b066da527193e878e160b4772af",slug:"fourier-transform-materials-analysis",bookSignature:"Salih Mohammed Salih",coverURL:"https://cdn.intechopen.com/books/images_new/2270.jpg",editedByType:"Edited by",editors:[{id:"111691",title:"Dr.Ing.",name:"Salih",surname:"Salih",slug:"salih-salih",fullName:"Salih Salih"}],productType:{id:"1",chapterContentType:"chapter",authoredCaption:"Edited by"}}]},chapter:{item:{type:"chapter",id:"55619",title:"Ovary Differentiation and Activity in Teleostei Fish",doi:"10.5772/intechopen.69022",slug:"ovary-differentiation-and-activity-in-teleostei-fish",body:'Teleostei fish represents about 50% of the vertebrates [1]. The bony fishes, within the Teleostei, are divided into Ostariophysi, Protacanthopterygii, and Neoteleostei.
Ostariophysi is the second largest superorder of fish, and it is considered the most basal among the Teleostei, representing about three quarters of the world’s freshwater fish [1]. This diverse group contains important fish in the area of feeding, sport fishing, aquarium, and research, such as the common carp and zebrafish (Cypriniformes), the characids and tetras (Characiformes), the catfishes (Siluriformes), and the electric eels (Gymnotiformes) [1, 2].
The Neoteleostei is another large clade of bony fish that includes most derived species among Teleostei, and it is also very important in several areas. Among the Neoteleostei, the most relevant group is the Perciformes, which presents the greatest diversity among all orders of fish, being the largest order among vertebrates [1]. The most popular Perciformes are the cichlids, such as tilapia.
Regardless of their position on the phylogenetic scale, in most Teleostei species, the reproduction is cyclic and seasonal, determining a series of morphophysiological modifications in their gonads [3]. Teleostei present several reproductive strategies [3, 4]. Among these, there are mechanisms of release of gametes in the aquatic environment for external fertilization, development of specialized organs for internal fertilization, posture of fertilized eggs after internal fertilization, and even internal gestation of the embryos [5].
In Teleostei, the sexual determination and gonadal differentiation are controlled by genetic, physiological, and behavioral factors [6]. The genetic sex of the embryo is determined at the time of the fertilization by the combination of the chromosomes from the male and female gametes, and sexual determination is defined as the sum of the genes responsible for the formation of the gonads and their characteristics [7, 8]. In this aspect, the genetic control is one of the main determinants of the gender, even though environmental factors, such as temperature, photoperiod, or salinity, also have a great influence on the process, determining the physiological gender of the fish [9].
Most Teleostei are dioecious or gonochoristic, that is, they present individuals with separated sexes. These fishes may present two types of gonadal development, classified as undifferentiated or differentiated gonochoristics. In undifferentiated gonochoristics, the undifferentiated gonad begins its development resembling an ovary. Subsequently, part of the individuals becomes male, while another part remains female. This natural condition is known as juvenile hermaphroditism. In the differentiated gonochoristics, the gonad differentiates directly in a testis or in an ovary [6, 10].
However, at the beginning of embryogenesis, the gender of the fish is not morphologically defined, since it does not have gonads differentiated in testes or ovaries, and there is no other developed characteristic which is associated to the reproductive system. There are only embryological precursors that will give rise to the ovaries and testes: the primordial germ cells (PGCs) and the somatic cells. At this stage of development, these cells are totipotent, and they can give rise to male or female gonads [6]. At some point during gonadal development, through hormonal chemical signaling, the gonad differentiates into the ovary or testis. Once this occurs and the gonadal tissue completes its differentiation, the fish becomes physiologically male or female [11].
The gonadal differentiation in the Teleostei includes changes in both somatic and germ cells [9], as mitotic divisions of oogonia or spermatogonia from the primordial germ cells, to structural changes, including mitotic proliferation of the somatic cells [12]. As a result, there is the formation of the ovarian cavity and spermatic ducts and lobules that will give origin to the ovaries and testes, respectively [9, 13].
In males, it is known that primordial germ cells establish specific positions, depending on the pattern of testicular organization [14]. This pattern, found in adult males, differs between basal (Ostariophysi) and derived taxa (Neoteleostei) [15]. However, the same does not happen to adult females. In this aspect, this chapter will describe the gonadal morphogenesis, with special attention to the formation of the ovarian cavity and establishment of the germinal epithelium, in basal taxa (
These three representatives were chosen because they are small ornamental species, quite resistant and known in ornamental aquarium. In addition, they can be reproduced in aquarium, presenting a fast period of differentiation. Although gonadal differentiation is quick in these representatives, the data showed here can be extrapolated to the species of their groups, since there is no difference in the events along the differentiation. Thus, despite the time of differentiation being species specific, the events and morphological changes are the same [16].
In Teleostei, as in other vertebrates, the primordial germ cells (PGCs) differentiate from yolk sac cells, originating from extragonadal regions, and migrate to the genital ridge during embryonic development [18, 19]. The genital ridge is formed by a thickening of the intermediate mesoderm, which protrudes ventrally into the coelomic cavity of the embryo being delimited by a mesothelium [6, 18, 19].
The primordial germ cells (PGCs) migrate to the genital ridge and are disposed between the somatic cells. Both cell populations proliferate by mitosis constituting the gonadal primordium [14, 16, 19, 20], as observed in
The gonadal primordium in
The gonadal primordia of the species here taken as representatives of the basal (
These characteristics of the gonadal primordium are also found in the adult form of the species that present an odd gonad, such as Poeciliids, a viviparous species, considered derived taxa [14, 23, 24]. In these, the formation of bilateral gonadal primordia is common. However, both gonadal primordia merge during the development of the gonadal tissue, forming a single organ in the adult individual [3, 14, 23].
Histologically, the gonadal primordia are formed by primordial germ cells (PGCs) and somatic cells. The somatic cells show varied forms, being predominantly squamous, with basophilic nucleus and scarce cytoplasm. The PGCs are large oval cells with voluminous nucleus and show quite evident nucleolus. Their cytoplasm is scarce and rich in “nüage,” presenting positive response to metanil yellow, indicating the presence of proteins in its constitution (Figure 1C).
The primordial germ cells (PGCs) are distributed along the gonadal primordium, which is filiform, long, and thin, composed by only one or two layers of PGCs. Mitotic activity of PGCs may be occasionally visualized.
At this stage, in any of the species, there are no morphological differences which suggest the evolution toward a male or female gonad. The number of germ and somatic cells gradually increases throughout the gonadal development. As a result, there is an increase in the size of the gonadal tissue, which becomes an undifferentiated gonad.
The undifferentiated gonads of
The undifferentiated gonads in
In these species, as in most Teleostei, the undifferentiated gonads are pairs, long, and thin, occupying two-thirds of the coelomatic cavity from the urogenital papillae. They are formed by primordial germ cells (PGCs) dispersed among somatic cells.
In parasagittal sections, the undifferentiated gonads are thicker in comparison with the gonadal primordium, mainly due to the greater amount of somatic cells, which present irregular and squamous forms. The primordial germ cells (PGCs) remain in small numbers and are initially isolated between somatic cells, scattered throughout the gonad (Figure 2).
Since the undifferentiated gonads are formed only by primordial germ cells and somatic cells, they are very similar in any group of fishes, from the basal to the most derived taxa, including primitive fish such as sturgeon [25] or species with indirect gonochoristic development as Cypriniformes
Morphological changes in the gonadal tissue, such as the formation of the ovarian cavity and the entrance into meiosis of the germ cells in females or the formation of the testicular ducts and lobules in males, are the main parameters used for the sexual distinction of gonochoristic gonads. Although these characteristics gather the consensus among different authors [6, 13], who use them as parameters for gonadal differentiation, the distinction between the female and male gonads may be detected prior to the entrance of the primordial germ cells into meiosis or before the formation of gonadal structures by the somatic cells. This detection of presumed female or male gonads is possible when considering the organization of germ and somatic cells in the gonadal tissue [14]. In other words, the gonadal differentiation in many species of Teleostei is closely related to the organization of the cellular types, which constitute the early gonadal tissue [14, 16].
A peculiarity observed here in
However, this pattern of organization of the Neoteleostei differs from the one found in the representatives of the Ostariophysi here utilized (
In these, the supposedly female animals present primordial germ cells distributed in the central region of the early gonadal tissue, which has a smaller number of somatic cells concentrated mainly in the peripheral region of the gonad. At this stage, among most basal fish, it is possible to differentiate female from male gonads. In female gonads, the oogonia proliferate, form continuous cords of cells, and enter into meiosis, originating the first oocytes [16], while in male gonads, spermatogonia are organized in acinar structure or cell clusters, after forming continuous cords [14].
Thus, before the appearance of structures such as the ovarian cavity formation, the female gonadal differentiation in both Ostariophysi and Neoteleostei is initially marked by the appearance of meiotic figures in gonadal tissue [12, 13, 16].
The gonads of
In
In
In
As it can be observed in these species, as well as in most Teleostei, the events involved in ovarian differentiation are quite similar, distinguishing between the species only in the chronology and the way in which the processes concurs to achieve the same final result—the formation of a cavity organ delimited by a germinal epithelium. Thus, the first stages of ovarian differentiation, characterized by the entrance of the germ cells into meiosis and the beginning of the folliculogenesis process, did not present significant differences between the species.
The gonadal tissue of the analyzed species is thin, elongated, and formed by primordial germ cells (PGCs), now differentiated into oogonia, and somatic cells. The oogonia, immersed within the gonadal tissue, may be associated with somatic cells or remain isolated (Figure 3A). Isolated oogonia proliferate by mitosis giving rise to new oogonia (Figure 3B). When associated with somatic cells, they form a cyst of oogonium (Figure 3C and D), which originates the initial prophase oocytes, upon entering into meiosis, analogous to what occurs in the germinal epithelium of the ovigerous lamellae in sexually adult females [28–30]. The development of germ cells within each cyst is synchronous, due to the presence of cytoplasmic bridges between oogonia (Figure 3E and F) and prophase oocytes [16, 22, 31]. Thus, the cytokinesis is incomplete.
Transmission electron microscopy of
Since each oogonium gives rise to a cyst and the cellular divisions begin, different cysts are formed next to each other, giving rise to cell clusters, delimited gradually by a sole basement membrane in formation (Figure 3C and D). Thus, throughout the gonad, it is possible to observe individual isolated oogonia between somatic cells and cysts delimited by somatic cells derived from the epithelium, containing oogonia and/or early prophase oocytes (Figure 3G).
The oogonia are oval cells that present scarce cytoplasm with granulations corresponding to “nüages.” Their nuclei are large and spherical with one or more evident nucleoli. Its cytoplasm presents spherical mitochondria with tubular ridges, often associated with “nüages” (Figure 3A–D).
Oocytes present in the cysts are also rounded, with nuclei containing chromatin in different forms of organization according to the stage of the prophase in which they are found (Figure 3G–M), but their cytoplasm does not differ them, always remaining slightly acidophilus and scarce. Initially, the prophase oocytes have a more basophilic nucleus than the oogonia, and there is a decrease in the amount of “nüage” in the cytoplasm. The leptotene oocyte shows a strongly basophilic nucleus, with at least one nucleolus quite evident. With the progression of the prophase, the oocyte gradually lost nuclear basophilia. The zygotene oocyte presents greater chromosome condensation, giving the nucleus a granular aspect. Formation of the synaptonemic complexes begins, allowing the pairing of the homologous chromosomes. In pachytene, the synaptonemic complexes are totally formed. In the nucleus of the oocyte, there is a strong basophilia next to the nuclear envelope. These stages are illustrated below.
Now, the germline cysts containing diplotene oocytes are invaded by somatic epithelial cells—the pre-follicle cells (Figure 3I–K). Pre-follicle cells strongly united by numerous desmosomes complete and gradually involve each oocyte which separates from the cyst, giving rise to an ovarian follicle (Figure 3L). During this process, known as folliculogenesis, pre-follicle cells begin to form the basement membrane, after differentiating into follicle cells (Figure 3M and N). Gradually, the basement membrane is synthesized, individualizing each ovarian follicle. After the oocyte enter and remain in diplotene stage, the lampbrush chromosomes become visible. The cytoplasm of the oocytes increases, becoming gradually more basophilic and initiating the primary growth while within the germline cysts. Now, the diplotene oocyte isolated in the ovarian follicle, and in primary growth, presents a nucleus with a variable number of nucleoli, which, initially located in the central region of the nucleus (oocytes with multiple nuclei), later become peripheral (perinucleolar oocytes).
In
Parasagittal histological section of the female gonads in
In
Parasagittal histological section of the female gonads in
In
Parasagittal histological section of the female gonads in
Parasagittal histological section of the female gonads in
At this stage of ovarian differentiation, the gonad is still compact in all the species here analyzed (Figures 4–7).
In most Teleostei fish, the ovaries are even saculiform organs, presenting a cavity in their interior. This type of ovarian organization is unique among vertebrates and is known as cystovarian ovary [32]. In this type, the ovaries are cavitary organs and present the germinal compartment in the form of lamellae, which protrude from the capsule toward the lumen of the organ. In this case, the ovarian cavity is continuous with the gonoducts [33], which merge caudally and flow into the urogenital papillae [3, 33].
The species utilized herein as representatives of Teleostei have this type of ovarian organization. The constitution of the ovary as a cavitary organ, and therefore the formation of the ovarian lumen, precedes the complete formation of the ovigerous lamellae in all of them, and, depending on the species, it may be concomitant to the constitution of the germinal epithelium. In all cases, the closure of the organ is gradual and can be followed through cross histological sections. Variations of the involved events can occur among the species studied.
In these species, the process of formation of the ovarian cavity follows what has been reported for most of fish [13] and is a result of the proliferation of somatic cells in the periphery of the ovary. This proliferation is responsible for the formation of the laminar tissues, which expand laterally and fuse, enclosing the forming ovary in a cavity—the before-known coelomatic cavity—now, the ovarian lumen.
Despite the similarities found during the cystovarian formation between the groups of fish analyzed, a single divergence could be observed, namely, the location of the somatic cell proliferation regions in the ovary and the direction (ventral or dorsal) of the laminar tissue toward the coelomic cavity.
In the Cypriniformes
Histological cross sections of the female gonads in
Histological cross section of the female gonads in
In
In both species, concomitant to the entrance of the oocyte in primary growth, the gonadal tissue, still compact, presents lateral tissue projections from the proliferation of somatic cells in the periphery of the gonadal tissue. Through the cross sections, the growth of the laminar tissue, on both sides of the ovary, can be traced toward the dorsal (
In
In all the species analyzed here, the gonadal tissue is still compact in the stage that precedes the formation of the ovigerous lamellae, even though the ovarian cavity is already formed. In the gonadal tissue, the developing ovarian follicles are gradually surrounded by a basement membrane (Figure 10), remaining immersed in the gonadal tissue, along with germline cysts of oogonia, of prophase oocytes and isolated oogonia [16].
Histological section of the female gonads in
In
Parasagittal histological section of the female gonads in
Thus, the lamellae gradually increase in size and project into the ovarian cavity. In the region of projection and formation of the ovigerous lamellae, the somatic cells peripheral to the gonadal tissue reorganize forming an epithelium that borders the newly formed lamellae. This newly formed epithelium isolates the germ cells from the interlamellar lumen. This mechanism, at the same time, forms the ovigerous lamellae and originates the germinal epithelium that borders the lamellae (Figure 12) [16].
Parasagittal histological section of the female gonads in
In
Parasagittal (A–H), longitudinal (I), and cross (J) histological section of female gonads in
With the distancing of several longitudinal parallel rows of somatic cells to provide the ovarian lumen formation, the gonadal tissue is separated longitudinally in its central-medial region, becoming pleated. Thus, several parallel longitudinal pleats are formed, each one delimited by the somatic cells that originated them, composing the primordium of the ovigerous lamellae (Figure 13I and J).
Within each newly formed ovarian lamellae, the oogonia and germline cysts reorganize and migrate to the lamellar periphery, associating with the epithelial somatic cells that compose the border of each ovigerous lamellae, thus constituting the female germinal epithelium in
By a mechanism opposite to the other Ostariophysi (the Cypriniformes
There are few reports on the different mechanisms that can lead to the formation of the ovigerous lamellae. Therefore, these mechanisms in other Teleostei are still quite unknown, making it impossible for an in-depth comparison along the evolutionary scale.
Thus, even though the formation of ovigerous lamellae is different in
During the formation of the female germinal epithelium, the somatic epithelial cells, originated from specific regions according to each species, interpose among the germ cells, interconnecting them, after migrating through the compact gonadal tissue. The germ cells, from the beginning of the formation of the gonad, are segregated from other tissue components by the pre-follicle cells. These, in their turn, are supported on a basement membrane. Thus, the germinal epithelium, when formed, will be separated from the ovarian stroma by the basement membrane [16, 21].
In all the species analyzed here, along the female gonadal tissue, there are other cellular components which are interposed to the ovarian follicles already formed and to the cysts of oogonia and/or oocytes. Among these cellular components, small spaces arise and expand gradually giving rise to extravascular spaces. The extravascular spaces are filled by fluids rich in glycoproteins and polysaccharides. In adult animals, they originate from extravasation of blood plasma, which leaves the circulatory system between the endothelial cells and begin to fill regions within the gonadal tissue [15]. It is assumed that the extravascular spaces, in animals with gonadal differentiation, are formed in the same way [16].
Now, this fluid is disposed between the cellular components, moving them apart. Concomitantly, among the cellular components, star-shaped cells with mesenchymal characteristics interconnect progressively, forming a loose network that gives rise to an interstitial compartment [16].
This compartment corresponds to the ovarian stroma, in which new cellular components will differentiate, remaining isolated from the germinal compartment by a basement membrane. The ovarian stroma in the fish is usually formed by a loose connective tissue, in which the extravascular spaces are larger and the amount of collagen fibers is small [16, 28].
From the newly formed stroma, the mesenchymal cells emit cytoplasmic projections which interact with the ovarian follicles and respond from now on by the formation of constituents of the theca. Since the follicles already have a totally formed basement membrane, the mesenchymal cells contacting the follicle, supported by their basement membrane, differentiate into a pre-theca cell and later into theca cells (Figure 14) [16].
Histological section of the ovary in
The ovarian stroma may be more or less developed, depending on the species. In
With the differentiation of the theca cells, the ovarian follicle becomes the follicle complex. The follicle complex is formed by the diplotene oocyte, surrounded by follicle cells, sustained by a basement membrane, and by two layers of theca cells [28, 33–36]. Thus, now the gonad presents two distinct compartments—the germinal epithelium of the ovigerous lamellae and the ovarian stroma [33, 35, 36]—separated by the basement membrane that becomes totally continuous (Figure 15).
Histological section of the ovary in
Within the follicle complexes, the oocyte development proceeds. Microvilli arise in the oocyte plasma membrane and in the membrane of the apical region of the follicle cells. In this region, oocyte and follicle cells contact, and the formation of the zona pellucida begins [16, 29].
Once the germinal epithelium is fully established, it will become permanently active. In the epithelium, the oogonia proliferate forming clusters, denominated nests (Figure 16A and B). In these, the oogonia associate to the somatic cells of epithelial origin, differentiate, and form a new germline cyst (Figure 16C–E) [30]. Within the cyst, oogonia proliferate or enter into meiosis giving rise to germline cysts of prophase oocytes (Figure 16F–K). Isolated oogonia, oogonia inside cysts, cysts containing oocytes, and pre-follicle cells start occupying the inside of the same nest [16, 30].
Folliculogenesis in a totally differentiated ovary of
After the formation of the ovarian follicle (Figure 16L), the oocyte follows its growth (Figure 16M), remaining connected to the germinal epithelium through a certain extension of the basement membrane shared between the follicle cells and the epithelial cells (Figure 16N) [16, 30, 33].
Once the ovarian follicle is formed, i.e., the folliculogenesis process is complete, the oocyte effectively initiates its primary growth [36–38]. From here, the oocytes will be ready to respond to the stimuli that lead to the incorporation of the yolk, and therefore they undergo maturation and subsequent ovulation or spawning [35–38].
The species
When analyzing different representatives of Teleostei, it can conclude that the processes involved in female gonadal differentiation are quite similar and it is possible to differentiate supposedly female and male gonads, even in the early development stages, independent on being a basal or derived species.
In the three species analyzed here, representatives of basal and derived taxa in Teleostei, the beginning of the female gonadal differentiation is marked by the entrance of the oogonia into meiosis, in early stages of the gonadal development, when the gonad is still a compact tissue. Thus, the formation of the ovarian cavity occurs only after the entrance into meiosis of the oocytes, preceding the formation of ovigerous lamellae in
Thus, although there are differences in the chronology of the differentiation among species of Teleostei, the processes involved are quite similar and culminate in the formation of analogous structures in the different fishes. Therefore, these data showed here can be applied to the most different groups of Teleostei fish.
Methodology used: Larvae and juveniles were obtained from spawns of adult of the three species. After hatching, part of the brood was sampled periodically covering the period of histologically discernible sex differentiation. The specimens were anesthetized with 0.1% benzocaine and killed according to the institutional animal care protocols and approval (175/2009-CEEA-IBB/UNESP). The gonadal tissues were fixed by immersion in 2% glutaraldehyde and 4% paraformaldehyde in Sorensen’s phosphate buffer (0.1 M, pH 7.2) for at least 24 h.
For light microscopy, the gonadal tissue from larvae and juveniles was embedded in historesin (Leica HistoResin). Serial sections (3μm) were stained with periodic acid Schiff (PAS) + hematoxylin + metanil yellow [39] and with the reticulin method that enhances the basement membranes. Gonadal tissues were evaluated by using a computerized image analyzer (Leica Qwin 2.5). The reticulin stain [40] uses an oxidizing agent, potassium permanganate, to oxidize aldehyde groups. Subsequently, the oxidized aldehyde groups are detected by the deposition of positive silver ions followed by their reduction using formalin. The result is a black hue of the reticulin fibers. As reticulin fibers are part of basement membranes, the method clearly detects basement membranes.
For electron microscopy, the gonadal tissue from larvae and juveniles was postfixed for 2 h in the dark in 1% osmium tetroxide (in the same buffer). To highlight the cellular structures, block-staining was carried out using an aqueous solution of 5% uranyl acetate for 2 h. Subsequently, the specimens were dehydrated and embedded in Araldite, sectioned, and post-stained with a saturated solution of uranyl acetate in 50% ethanol and 0.2% lead citrate in NaOH (1 N). Electron micrographs were obtained using a Tecnai Spirit Fei Company Transmission Electron Microscope.
We would like to thank Brazilian Agencies CNPq (Conselho Nacional de Desenvolvimento Científico e Tecnológico), CAPES/PROAP (Coordenação de Aperfeiçoamento de Pessoal de Nível Superior), and FAPESP (Fundação de Amparo à Pesquisa do Estado de São Paulo).
The functioning links between cancer and inflammation was approached by a great scientist and physician Rudolf Ludwing Carl Virchow in 1863 [1, 2, 3]. Thereafter, for long Virchow’s idea had almost been unevaluated and discussed insufficiently [1]. Balkwill and colleagues (2001) supported Virchow’s idea and stated that if molecular deregulation is the “match that lights the fire” of cancer, then some types of inflammation may act as the fuel that stimulates the flame. For instance, the inflammatory process act as a cofactor in malignancy in the bladder, cervical, ovarian, gastric, MALT (mucosa-associated lymphoid tissue) lymphoma, esophageal, colorectal, hepatocellular, bronchial, mesothelioma, and Kaposi sarcoma [3].
Currently, it is scientifically proven that inflammation promotes all stages of tumor formation as well as the development of cancer where chronic inflammation or non-resolving inflammation is playing a principal role in the initiation, promotion, malignant transformation, invasion and metastasis of cancer [4, 5, 6, 7, 8, 9]. Interestingly, cancer-related inflammation is representing its 7th position as a cancer hallmark and catching the current research attention in human cancer biology [5]. Basically, Inflammation act on cancer development by linking extrinsic and intrinsic pathway [9]. The extrinsic pathway develops inflammatory condition or microenvironment by inflammatory leukocytes particularly macrophages and soluble mediators (vasoactive amines such as histamine and serotonin, peptide such as bradykinin, and eicosanoids such as thromboxanes, leukotrienes, and prostaglandins) that raises cancer risk [9, 10, 11]. Intrinsic pathway is driven by genetic events (e.g. oncogenes, genetic aberrations) causing neoplastic transformation, initiate the expression of inflammation-related programs which guide the construction of an inflammatory microenvironment [11]. Inflammatory system involves the dynamic regulations of hundreds of genes and complex transcriptional program [12]. Moreover, the gene regulations by intron are often expressed in most of the cell through the effects of splicing or specific features [13, 14, 15]. Recently, several scientific reports claim that retained intron deregulates splicing machine in tumor transcriptoms [16, 17, 18]. Intron retention is considered as mis-splicing in which rather than being spliced out intron stays back and retained in mature mRNA [17]. However, the broad gap exists in monitoring of introns role in understanding of gene expression which could be a powerful tool in biotechnological and therapeutic applications. This chapter will cover intron’s role in the development of inflammation, intron retaining genes causing inflammation to cancer and finally unfolding of a hypothesis about CRISPR like machine to monitor introns function.
Inflammation is activated by leukocytes which make inflammatory mediators in the extrinsic pathway and this pathway is also triggered by various infections and toxic agents such as gastric acid reflux, autoimmune disease etc. [9]. Patients with inflammatory bowel diseases have an increased chance of getting colorectal cancer. As an example, about 43% of patients with ulcerative colitis develop colorectal cancer [19]. Moreover,
Intrinsic pathway is activated by genetic variation such as proto-oncogene activation, inactivated tumor suppressor genes, and chromosomal multiplication as well as mutation which develops neoplasia [22]. A gene coding protein namely tyrosine kinase RET shows rapid and ample genetic variation in human papillary thyroid carcinoma (PTC) and initiates the transcriptional program that links to the development of inflammation [33]. Transcriptome profile is activated by tyrosine kinase RET in human papillary carcinoma and comprises with colony-stimulating factors (CSFs), interleukin 1β (IL-1β), cyclo-oxygenase 2 (COX2), chemokines attacking monocytes and dendritic cells (CCL2 & CCL20), angiogenicchemokines (CXCL8), matrix-degrading enzymes and inhibitors, chemokine receptor (CXCR4) [34]. For example, patients with lymph nodes metastasis have shown an elevated level of tyrosine kinase RET activated inflammatory molecules in their biopsy results which demonstrate that genetic events have taken place in the pathogenesis of tumor and constructed an inflammatory environment [33, 34, 35]. Moreover,
Intron is defined as any intervening nucleotide sequence that formed splicing at the RNA level [39]. Intron was first discovered in 1970s with a traditional views that the coding region of eukaryotic genes are interrupted by introns which are spliced out from pre mature mRNA transcripts before the formation of mature mRNA [39, 40, 41]. After the elucidation of intron splicing mechanism, scientist became excited about its function on gene expression and speculated that may be introns carry out some function like regulation of splicing function, regulation of transcription, evolutionary function or coding capacity but there was no clear examples of their active functions on gene expression [41]. From the starting 21st century, many researches have claimed about the intron function on gene expression or intron mediated enhancement of gene expression [13, 42, 43, 44, 45, 46, 47]. However, a question remains unclear that within the genome, who is responsible to remember or decide which intron or parts of nucleotide sequences are necessary to stay within the mature mRNA stand for inflammatory gene expression rather than form splicing that result in cancer? Current chapter sheds light on the above question and hypothesize CRISPR like machine in perspective of inflammation mediated cancer development.
After the discovery of alternative splicing (AS), the transcriptomic and proteomic complexity has increases significantly [47, 48]. Recent breakthrough studies in high-throughput sequencing have explored a pivotal role of AS in normal biology that more than 95% of human multi exonic genes are subject to AS and produce at least two alternative isoforms [49, 50]. Moreover, Braunschweig and colleague compared 11 vertebrate species and observed that about 50–75% of multi-exonic genes are affected by intron retention (IR) which is one kind of AS [47, 48, 51]. While another study showed that IR affects near about 80% of protein coding genes in humans [52]. Some scientific reports also considered IR as a harmful process for the body by slowing down splicing kinetics and delaying the onset of gene expression, by raising pre-mRNA degradation in the nucleus through nuclear exosomes and finally by enhancing cytoplasmic pre-mRNA degradation through nonsense-mediated decay [51, 53, 54]. This statement is also supported by the Green and colleague’s research as the genes that encoded the regulators of macrophage transcription, signaling inflammation, and phagocytosis has increased their expression when the IR events decreased [55]. As it is known, that intron retention (IR) is the process where instead of typically being spliced out, the introns remain intact in the mature mRNAs and thus whole process of IR supposedly has numerous physiological drawbacks resulting in different diseases [47, 48].Currently, many researchers strongly claim that IR is a key mechanism to control gene expression during the development, differentiation and activation of several types of mammalian cell [56, 57, 58, 59, 60, 61, 62, 63]. A recent study by Green and colleague claimed that intron retention affected the expression of key genes
Gene name | Protein name | General function | Types of cancer that is developed by IR gene | Abnormal function | References |
---|---|---|---|---|---|
TGFB Induced Factor Homeobox 2 | TGF-β-induced factor homeobox 2 ( | Ovarian cancer | Over expressed | [64, 65] | |
Epstein–Barr nuclear antigen 3 | This gene plays a principle role for activation and immortalization of human B-cells. Represses transcription of viral promoters TP1 and Cp interact with RBPJ and also inhibits the | Lymphoma and Epithelial cancers | Amplication | [66, 67] | |
Apolipoprotein E4 | This gene gives instructions to make a protein called apolipoprotein E. This protein combines with fats (lipids) in the human body to form molecules called lipoproteins. Lipoproteins in body package cholesterol and other fats and transfer them by the bloodstream. | Breast cancer | Increased frequency | [68, 69] | |
Epidermal growth factor receptor | Receptor tyrosine kinase binding ligands of the EGF family. They activate a lot of signaling cascades to transform them extracellular cues into appropriate cellular responses. | Lung cancer | Mutation or cell damage | [70, 71] | |
Receptor Tyrosine Kinase | Ovarian cancer, cholangiocarcinoma, inflammatory myofibroblastic tumor, colorectal, and angiosarcoma | Mutation | [72, 73] | ||
RUNX Family Transcription Factor 1 | The protein encoded by this gene represents the alpha subunit of CBF(Core binding factor) and is thought to be involved in the development of normal hematopoiesis. Chromosomal translocations involving by this gene are well-documented and have been associated with several types of leukemia. Three transcript variants encoding different isoforms have been found for this gene | Myeloid and lymphoid | Mutation | [74, 75] | |
Tumor Protein 53 | Breast cancer, bone and soft tissue sarcomas, brain tumors and adrenocortical carcinomas (ADC), leukemia, stomach cancer and colorectal cancer | Gene Alternation or Deletion or Mutation | [76, 77] |
List of intron retention genes and their functions.
PCR technique revealed two alternate splice forms of
The expression of the
It was claimed that 2340 and 1422 genes show tumor-specific and normal tissue-specific retention events respectively [89, 90]. For example,
Intron retention frequency may be responsible to inactive tumor suppressor genes in many cancers cell [93]. Study revealed that in cancerous condition, retained intron transcript exits in NMD pathway and inactivates the
The evolutionary history and relationship of an organism or group of species are named phylogeny. Phylogeny depicts the connection of an organism. Phylogenetic relationships give information on shared common ancestry but not obligatory how organisms are similar or different. Phylogenic tree analysis of all genes those are performing intron retention scenario, PcGs and Ras oncogenes (Figures 1–3) has done to find out the relation among the genes. MEGA X software has been used to construct the phylogenic tree to understand the relation among all the cancers genes (Tables 1 and 2).
Phylogenetic tree of intron retaining (IR) genes.
Phylogenic tree between
Phylogenic tree among IR gene, RAS oncogenes and PcGs.
Gene name | Protein name | General function | Types of cancer that is developed by IR gene | Abnormal function | References |
---|---|---|---|---|---|
B-lymphoma Moloney murine leukemia virus insertion region-1 | This gene functions through chromatin remodeling as a principle epigenetic repressor of numerous regulatory genes involved in embryonic development and self-renewal in somatic stem cell and also plays a median role in DNA damage repair. It is an oncogene and abnormal expression is related with multiple cancers and resistance to certain chemotherapies. | Gastric, ovarian, breast, head and neck, pancreatic and lung cancer, primary hepatocellular carcinoma (HCC) and endometrial carcinoma | Over expression | [95, 96] | |
Chromobox proteins 7 | This gene encodes a protein that comprises the CHROMO (CHRomatin Organization MOdifier) domain. It is thought to control the lifespan of several normal human cells. | Breast, Thyroid, Colorectal, Pancreas, Lung carcinoma and Glioblastoma | Down regulation | [97, 98] | |
Phenylalanine Hydroxylase | This gene gives instructions for making an enzyme called phenylalanine hydroxylase. This enzyme is responsible for the primary step in processing phenylalanine, which is a building block of proteins (an amino acid) obtained through the diet. | Liver cancer | Down regulation | [99, 100] | |
Ring Finger 1A | This gene encodes proteins characterized by a RING domain, a zinc-binding motif related to the zinc finger domain. The gene product can bind DNA and can act as a transcriptional repressor. It is related with the multimericpolycomb group protein complex. | Hepatocellular and colorectal carcinomas | Down regulation | [101, 102] | |
Polycomb group ring finger 2 | Breast cancer, Prostate cancer | Loss of expression and down regulation | [103, 104] | ||
Enhancer of zeste homolog-2 | The | Breast cancer, Colorectal cancer, Endometrial cancer, Gastric cancer, Liver cancer, Lung cancer | Over expression | [105, 106] | |
Embryonic ectoderm development | This gene encodes a member of the Polycomb-group family. It maintains the transcriptional repressive state of genes over successive cell generations. This protein interacts with enhancer of zeste 2, the cytoplasmic tail of integrin beta7, immunodeficiency virus type 1 (HIV-1) MA protein, and histone deacetylase proteins. This protein mediates repression of gene activity through histone deacetylation, and may act as a specific regulator of integrin function. Two transcript variants encoding distinct isoforms have been identified for this gene. | Colorectal Cancer, acute myeloid leukemia and diffuse large B cell lymphoma | High expression | [107, 108, 109] | |
suppressor of zeste 12 homolog | This zinc finger gene has been detected at the breakpoints of a recurrent chromosomal translocation reported in endometrial stromal sarcoma. Recombination of these breakpoints results in the fusion of this gene and | Colorectal, ovarian and non-small lung cancer, head and neck squamous cell carcinoma | Over expression | [110, 111] | |
PHD fing protein 19 | Hepatocellular carcinoma, glioma, and ovarian cancers, glioblastoma progression, prostate cancer | Over expression | [112, 113] | ||
TumourProtein 53 | Functions are the same as discussed in Table 1 | Types of cancers are the same as discussed in Table 1 | Gene alternation or deletion or mutation | [76, 77] | |
Kirsten rat sarcoma viral oncogene | The | Non-small cell lung cancer, colorectal cancer, and pancreatic cancer | Mutation | [114, 115] |
List of PcGs genes and their functions.
Figure 1 demonstrates that the vertical line on the left side is the common ancestor or the root of the seven gene sequences from which the genes have been evolved in a period. The evolution period can be explained through the horizontal lines near the sequences. The small length of lines before sequences means the sequences evolved in a short period whereas the long length of lines means longer time was needed for the sequences to be evolved. The common ancestor or root has been divided into two branch points. From the first branch point total of five sequences have been modified which were
Figure 2 depicts that both type of cancer genes (
Figure 3 demonstrates that all the genes were evolved from a common ancestor. The IR gene
CRISPR-Cas9 is also known as a genome-editing tool where CRISPR stands for “Clustered Regularly Interspaced Short Palindromic Repeats and Cas9 is an enzyme that cuts foreign DNA [116]. In the 21st century, CRISPR-Cas9 is being used immensely in medical technology to edit, remove or add a gene to correct genetic defects [116]. CRISPR has conformed from the natural defense mechanism of bacteria, archaea and developed an immune system by CRISPR loci [116]. CRISPR and Cas9 enzyme serve as an immune guard and provide safety against bacteriophage, viruses, and foreign invaders [117, 118, 119]. The immunization process after invading foreign genetic elements, a small fragments of foreign DNA are integrated into the CRISPR repeat-spacer array within the host chromosome as new spacers. Thus, a genetic record of prior infection will save into the host body that enables to prevent future invasion of the same invader [120, 121] (Figure 4).
CRISPR biology.
The nucleotide repeats and spacers are main two component of CRISPR. Repeated sequence of nucleotide is distributed in the CRISPR region and Spacers are a small portion of DNA present in the CRISPR region. Destruction of foreign invading DNA or RNA occurs by Cas9 enzyme. If in future, the foreign body again attacks the organism, they fight it off as they have the virus or foreign invader’s DNA from beforehand and thus they recognize it and kill it [121]. So in CRISPR biology, spacer acts as a responsible sequence to remember or decide which foreign body needs to be killed where Cas9 enzyme plays a pivotal role. According to our hypothesis, intron network might work as CRISPR like functioning model.
According to the section 3, it is confirmed that IR have the ability to influence inflammation mediated cancer development. From the CRISPR function it is clear that according to the demand of the cell, CRISPR can get activated. It might be possible that according to the demand of the abnormal cells, intron retention may form to confirm inflammation mediated malignancy state (Figure 5). Moreover, our phylogenetic tree analysis depicts that PcGs,
Illustration of retained intron in the gene causing mutation which leads to inflammation and tumorigenesis resulting in development of cancer.
Cancer is a genetic disease and is one of the leading causes of death around the world. As a genetic event, the intron retention causes inflammation as well as the development of cancer cells. So far, it is clear that the intron is spliced away during gene expression while exons remain and express the genes. However, the retention of introns is an unlikely phenomenon that differs from the common hypothesis and appears anomalous. The genes involved in retaining intron have a carcinogenic effect. It may be speculated that some nucleotide sequence whether it is coding or non-coding region could function as a memory sequence, hence are able to remember intronic sequence as like spacer responsibility of CRISPR system and would only be functioning according to the demand of the cell. Future in depth analysis on intron retaining genes is required to explore their effect on inflammation and cancer.
We gratefully acknowledge Asian Network of Research on Antidiabetic Plants (ANRAP) and Bangladesh University of Health Sciences (BUHS) for providing all types of logistic facilities to conduct this work.
Vascular endothelium growth factor Papillary thyroid carcinoma colony-stimulating factors Interleukin 1β Cyclo-oxygenase 2 Polycomb complex target genes Alzheimer’s disease Clustered Regularly Interspaced Short Palindromic Repeats Alternative splicing Intron retention Chronic obstructive pulmonary disease Mucosa-associated lymphoid tissue
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