Dr. Pletser’s experience includes 30 years of working with the European Space Agency as a Senior Physicist/Engineer and coordinating their parabolic flight campaigns, and he is the Guinness World Record holder for the most number of aircraft flown (12) in parabolas, personally logging more than 7,300 parabolas.
\\n\\n
Seeing the 5,000th book published makes us at the same time proud, happy, humble, and grateful. This is a great opportunity to stop and celebrate what we have done so far, but is also an opportunity to engage even more, grow, and succeed. It wouldn't be possible to get here without the synergy of team members’ hard work and authors and editors who devote time and their expertise into Open Access book publishing with us.
\\n\\n
Over these years, we have gone from pioneering the scientific Open Access book publishing field to being the world’s largest Open Access book publisher. Nonetheless, our vision has remained the same: to meet the challenges of making relevant knowledge available to the worldwide community under the Open Access model.
\\n\\n
We are excited about the present, and we look forward to sharing many more successes in the future.
\\n\\n
Thank you all for being part of the journey. 5,000 times thank you!
\\n\\n
Now with 5,000 titles available Open Access, which one will you read next?
Preparation of Space Experiments edited by international leading expert Dr. Vladimir Pletser, Director of Space Training Operations at Blue Abyss is the 5,000th Open Access book published by IntechOpen and our milestone publication!
\n\n
"This book presents some of the current trends in space microgravity research. The eleven chapters introduce various facets of space research in physical sciences, human physiology and technology developed using the microgravity environment not only to improve our fundamental understanding in these domains but also to adapt this new knowledge for application on earth." says the editor. Listen what else Dr. Pletser has to say...
\n\n\n\n
Dr. Pletser’s experience includes 30 years of working with the European Space Agency as a Senior Physicist/Engineer and coordinating their parabolic flight campaigns, and he is the Guinness World Record holder for the most number of aircraft flown (12) in parabolas, personally logging more than 7,300 parabolas.
\n\n
Seeing the 5,000th book published makes us at the same time proud, happy, humble, and grateful. This is a great opportunity to stop and celebrate what we have done so far, but is also an opportunity to engage even more, grow, and succeed. It wouldn't be possible to get here without the synergy of team members’ hard work and authors and editors who devote time and their expertise into Open Access book publishing with us.
\n\n
Over these years, we have gone from pioneering the scientific Open Access book publishing field to being the world’s largest Open Access book publisher. Nonetheless, our vision has remained the same: to meet the challenges of making relevant knowledge available to the worldwide community under the Open Access model.
\n\n
We are excited about the present, and we look forward to sharing many more successes in the future.
\n\n
Thank you all for being part of the journey. 5,000 times thank you!
\n\n
Now with 5,000 titles available Open Access, which one will you read next?
\n'}],latestNews:[{slug:"stanford-university-identifies-top-2-scientists-over-1-000-are-intechopen-authors-and-editors-20210122",title:"Stanford University Identifies Top 2% Scientists, Over 1,000 are IntechOpen Authors and Editors"},{slug:"intechopen-authors-included-in-the-highly-cited-researchers-list-for-2020-20210121",title:"IntechOpen Authors Included in the Highly Cited Researchers List for 2020"},{slug:"intechopen-maintains-position-as-the-world-s-largest-oa-book-publisher-20201218",title:"IntechOpen Maintains Position as the World’s Largest OA Book Publisher"},{slug:"all-intechopen-books-available-on-perlego-20201215",title:"All IntechOpen Books Available on Perlego"},{slug:"oiv-awards-recognizes-intechopen-s-editors-20201127",title:"OIV Awards Recognizes IntechOpen's Editors"},{slug:"intechopen-joins-crossref-s-initiative-for-open-abstracts-i4oa-to-boost-the-discovery-of-research-20201005",title:"IntechOpen joins Crossref's Initiative for Open Abstracts (I4OA) to Boost the Discovery of Research"},{slug:"intechopen-hits-milestone-5-000-open-access-books-published-20200908",title:"IntechOpen hits milestone: 5,000 Open Access books published!"},{slug:"intechopen-books-hosted-on-the-mathworks-book-program-20200819",title:"IntechOpen Books Hosted on the MathWorks Book Program"}]},book:{item:{type:"book",id:"3797",leadTitle:null,fullTitle:"Industrial Robotics: Programming, Simulation and Applications",title:"Industrial Robotics",subtitle:"Programming, Simulation and Applications",reviewType:"peer-reviewed",abstract:"This book covers a wide range of topics relating to advanced industrial robotics, sensors and automation technologies. Although being highly technical and complex in nature, the papers presented in this book represent some of the latest cutting edge technologies and advancements in industrial robotics technology.\r\nThis book covers topics such as networking, properties of manipulators, forward and inverse robot arm kinematics, motion path-planning, machine vision and many other practical topics too numerous to list here.\r\nThe authors and editor of this book wish to inspire people, especially young ones, to get involved with robotic and mechatronic engineering technology and to develop new and exciting practical applications, perhaps using the ideas and concepts presented herein.",isbn:null,printIsbn:"3-86611-286-6",pdfIsbn:"978-953-51-5808-0",doi:"10.5772/40",price:159,priceEur:175,priceUsd:205,slug:"industrial_robotics_programming_simulation_and_applications",numberOfPages:702,isOpenForSubmission:!1,isInWos:1,hash:"1e417b44016323de5c12e77210148af6",bookSignature:"Low Kin Huat",publishedDate:"December 1st 2006",coverURL:"https://cdn.intechopen.com/books/images_new/3797.jpg",numberOfDownloads:149315,numberOfWosCitations:140,numberOfCrossrefCitations:92,numberOfDimensionsCitations:185,hasAltmetrics:0,numberOfTotalCitations:417,isAvailableForWebshopOrdering:!0,dateEndFirstStepPublish:"September 9th 2013",dateEndSecondStepPublish:"September 30th 2013",dateEndThirdStepPublish:"January 4th 2014",dateEndFourthStepPublish:"April 4th 2014",dateEndFifthStepPublish:"May 4th 2014",currentStepOfPublishingProcess:5,indexedIn:"1,2,3,4,5,6,7",editedByType:"Edited by",kuFlag:!1,editors:[{id:"134111",title:"Dr.",name:"Kin Huat",middleName:null,surname:"Low",slug:"kin-huat-low",fullName:"Kin Huat Low",profilePictureURL:"https://mts.intechopen.com/storage/users/134111/images/system/134111.jpg",biography:"Dr. Low Kin Huat is currently a professor in the School of Mechanical and Aerospace Engineering, Nanyang Technological University in Singapore. 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\n
1. Introduction
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The exploitation of small ruminants (goat and sheep) has always been linked to the development of human civilizations, where they have mainly fed on their derived products such as milk and meat. Currently, the sheep population is around 1 billion head concentrated above 50% in three countries, China, Australia, and New Zealand, contrary to goats with around 720 million heads, distributed mainly in Asia, Africa, and South America. Both species have similar characteristics in some anatomical aspects (a pair of nipples), gestation period (150 days), and presence of seasonal anestrus, differing in terms of magnitude and depth and presence of the male effect. However, they are completely different in feeding habits, nutrient needs, and grazing systems, with differences in terms of the female’s reproductive tract, among other characteristics [1]. Currently the study of reproduction has intensified over the years in the goats and its counterpart that is the buck. Therefore, in the following topics, the importance of global reproduction of the goat will be discussed, considering that progress has been made today in the application of third generation reproductive techniques and that today they are already consolidated and developed in the bovine species [2].
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\n
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2. Reproduction in goats
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In most areas of the world, goats are mated once yearly in the fall, during their natural mating season, for spring kidding [3, 4, 5]. Animals bred at this time are more likely to get pregnant and have multiple kids. A longer breeding season allows for flexibility in breeding and kidding dates to times when the climate is more favorable, and forage is available for the lactating doe. In addition, dates of ethnic/alternative markets should also be considered in the decision about when to breed females. How long the males are kept in with females for mating determines how long kidding will last, but a 40 to 45-day breeding season will guarantee that each doe has had at least two opportunities to come into heat. The male-to-female ratio in this breeding system is approximately 1 male per 30–40 females, but in synchronized breeding, this ratio should be 1 male with 20 or less females.
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Likewise, under range conditions, bucks are often maintained with the doe herd throughout the year for continuous breeding. In such a system, proper health management is difficult and only limited supervision can be provided during kidding [1]. Care is also required to routinely remove offspring from the herd to avoid mother/son and father/daughter mating’s. Although buck exposure is continuous, kidding under continuous mating will eventually follow seasonal breeding patterns, depending on the location of the farm and the breed of goat used.
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However, globally, in intensive milk production systems, the use of basic reproductive techniques has been applied more extensively, for example the estrus synchronization techniques, artificial insemination, (AI), is being used more commonly by goat producers [6]. Artificial insemination makes it possible to obtain or transfer genetic material domestically and internationally. Many goat producers, both meat and dairy, utilize AI to produce animals that are more desired by markets and consumers as well as animals that will do well at local, state and national livestock shows.
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2.1 Perspectives and advances in the study of the estrous cycle of the goat
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Currently the estrous cycle is being studied from a perspective of hormonal changes according to the ovarian structures that are present during each of the phases that occur (follicular and luteal) [7, 8]. The above is with the objective of evaluating the size of structures and correlating them with hormonal profiles. Considering that by understanding the physiology and anatomy and the perspective of manipulating the oestrus cycle, we can advance or achieve higher gestation rates [9]. It has been stabilized with the application of hormonal products and/or the male effect to have an oestrus presence of 100%. However, pregnancy percentages vary greatly according to a large number of factors (see Figure 1), where each of them affects the final result cross-sectionally, which is pregnancy.
\n
Figure 1.
Factors that affect the reproductive response in goats.
\n
Estrous induction began to develop in goats and sheep for more than 50 years, where injected progesterone began to be used daily, until today with the use of two types of vaginal devices: vaginal sponge and delivery device. Controlled (CIDR), each having its advantages and disadvantages [10, 11]. The response in each of the devices has been accompanied by secondary hormones of intramuscular application that favor the development of the follicles, the synchronization of them for their ovulation and that these become corpus luteum with adequate size and with a sustainable production of progesterone. It is well known that low LH levels during the progestogen synchronization protocol will affect the fate of large follicles. However, these follicles require LH for their maintenance and development, so they will present atresia and new ovulatory follicles appear that will grow. In long estrous synchronization protocols (above 10 days), when the vaginal devices are removed, they release little progestogen and do not completely suppress LH. With the above, an abnormal follicular development occurs, which become persistent, leading to low fertility and therefore gestation.
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2.2 Will it be possible to improve the parameters of presence of estrus and pregnancy using hormones in the coming years?
\n
Changes or results in estrous synchronization programs have been modified over the years depending on the duration of insertion of the sponge or the device in the goat, however, the use of hormones to regulate goat reproduction has been maintained over the years [12, 13], with changes especially in the higher use of nonsteroidal hormones, such as those derived from prostaglandins, gonadotropin-releasing hormones, and hormones of follicular growth and development such as equine chorionic gonadotropin; being the most frequent use in the European community for health reasons. The use of steroid hormones such as progestogens continue to be used globally [14], but under the premise of using short protocols (5 to 7 days). In the present and in future years the use of short protocols of 5–7 days will be used more and more because it has a series of advantages compared to short protocols; these being the decrease in the presence of vaginitis in animals; in the case of CIDR devices, reuse them up to twice more with an effectiveness of up to 90% of estrous in goats. However, the health risk must be considered as it can contaminate bacteria, viruses from one animal to another.
\n
The important thing is to be able to develop vaginal devices with a lower concentration of progesterone and avoid being reused to avoid this type of infection.
\n
On the other hand, the use of estrous synchronization protocols in goats using nonsteroidal hormones in combination with the male effect has been developing more intensively in recent years. For example, the administration of double doses of PGF2α is recommended to synchronize estrous in cycling goats, with an interval of 10–14 days (appointment), which ensures that most does will present the mid luteal phase, when applying the second dose, and that all will respond with the behavior of estrus and ovulation (appointment). However, their response may vary depending on the insemination technique, the dose to be applied and the interval between doses. Besides, it should be considered that only the goat that is cycling with the presence of an active luteal body, would work this protocol. Currently, the male effect is used, so that an estrus occurs, a CL is formed, and the protocol based on prostaglandins is started.
\n
In goats, PGF2α and its analogs are effective luteolytic agents, where very small doses (1.25 mg) of PGF2α are currently required, with the corpus luteum being more sensitive compared to cows. Likewise, responses to low doses of its analogues, such as cloprostenol, have been observed; 125 μg doses have been used in goats, but even a 26 μg dose has been shown to be effective [15, 16]. As in sheep, the age of the corpus luteum and, therefore, the day of the cycle in which PGF2α is administered determines the degree of synchronization obtained and the time required for the heat to appear, the LH peak and the ovulation [17]. Several studies indicate that goats treated on day 6 of the cycle go into heat and show an LH peak much earlier than those treated on day 12 [18, 19].
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\n
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2.3 Advances and use of the male effect, a case study until today!
\n
The use of the male effect (Figure 2) has been a case study up to nowadays at a global level [20, 21], where different alternatives have been evaluated in order to understand its way of acting under different scenarios of a goat production system and achieve further efficiency in reproduction in the goat [22]. The sudden introduction of the goat increases the release of LH in goats [23], where the first estrus is not silent [24], so the goat effect produces a high degree of estrus synchronization [25]. Also, short cycles of 5–6 days or 10–12 days may appear after introducing the male, in these cases fertility is lower than in normal cycles [26]. Over the years, different scenarios of the male effect have been validated, modified or compared [27]; for example, [28] determined that the male-female ratio does not decrease the ability of sexually active males to induce sexual activity in anovulatory goats, but it does delay the response to the male effect. Likewise, [29] determined that the separation of the goats from the male goats is not necessary as it was thought in previous years to be able to stimulate the sexual activity of goats subjected to the male effect. Followed by another investigation where they verified that the bleating (vocalizations) of the goat were not sufficient to stimulate the presence of estrus and ovulation, therefore, the frequency of pulses of the LH was not increased [30]. Likewise, there are studies where the introduction of estrogenized females when introducing the buck can stimulate the estrous activity of anovulatory goats [31]. Delgadillo et al. [32] reviewing the male effect on goats, mention that in previous years it was mentioned that the male should be in permanent contact with the goats, their studies elucidated that it is not necessary and with a minimum contact of 4–16 hours, percentages of estrus can be reached in goats subjected to the male effect, the same as in groups that are in permanent contact with the males.
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Figure 2.
Sequence of sexual behavior in bucks.
\n
However, despite the advantages of using the male effect in goats, even today in large goat populations its use has been limited to continue with the natural breeds according to the time of year. Perhaps the lack of basic infrastructure to install and separate the bucks who are going to have the light programs have made their practical application until today still limited.
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\n
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2.4 Male social hierarchy and its impact on reproduction
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One of the key aspects to improve the performance of the herd is the proper evaluation of the reproductive capacity of the male, performing both a general physical examination, a specific examination of the reproductive system, a seminal quality examination and another of their libido and ability to mount. [33], with the aim of ensuring an adequate selection of males that contribute to improving the efficiency and profitability of the reproductive unit (Figure 3).
\n
Figure 3.
Factors influencing the presentation of the male effect in goats.
\n
Previous studies have evaluated the social hierarchy in rams raised in pairs, identifying that the dominant males exhibit a greater sexual precocity and a greater reproductive capacity compared to the subordinate males. A negative influence on testosterone production has also been reported, due to the stress of the grouping of bucks [34, 35, 36].
\n
In goats housed in herds with different densities, the social interactions registered between them were evaluated, as well as the levels of cortisol in blood to determine if the levels of said hormone vary depending on the size of the herd, identifying that the size of the pen and the size of the herd influences the increase in stress due to clustering, negatively impacting the weight gain of the bucks and the productivity of the herd [37, 38].
\n
Ivasere et al. [39] recorded behavioral changes by intensifying production systems and their effect on productive aspects such as nutrition, reproduction and diseases. They observed that the social structure is of great importance in the physiological and ethological development in bucks, modifying the frequency of courtship, copulation and the stress level in bucks grouped in herds of different densities. So far there is little information in the literature about the effect of regrouping previously raised male goats in pairs and regarding how serum cortisol and testosterone concentrations, seminal quality and sexual behavior are affected after such grouping.
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2.4.1 Cortisol and stress physiology in bucks
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Within the management of goats, efforts have been made to develop strategies to improve the quality and efficiency within the herds. At an intensive level, the management carried out ranges from supplying drugs, palpation, semen extraction, and pen cleaning. These management activities, in conjunction with other factors, such as the size of the herd, overcrowding, feeding or the immune system of animals, influence the body’s physiological response to various stressful situations [40]. The most common stressors in goat production are mainly those caused by environmental heat and the increase in body temperature, deprivation or lack of access to food or water, as well as modifications in the hierarchical structure of the herd or change of habitat [41].
\n
Among the responses at the physiological level present in male goats in stressful situations is the secretion of glucocorticoids (GC), which exert a negative feedback effect on the hypothalamic-pituitary-gonadal axis, reducing the synthesis of GnRH and together thereby inhibiting the synthesis of gonadotropins and sex steroid hormones [34, 39]. This endocrine mechanism aims to stimulate the body to respond to stressors, such as loss of appetite, suppression of the immune system, energy mobilization, vasoconstriction, and loss of erection and receptive sexual behaviors [42].
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2.4.2 Social factors and sexual behavior in males
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The grouping of bucks is a widespread practice mainly in stable and mixed management systems around the country [17, 43].
\n
The study of behavior has shown that the establishment of hierarchical ranks and social organization influence sexual behavior that will be exhibited by a male under grouping conditions [44]. This dominance is related in turn to the live weight of the animal and its age, mainly as part of a display of reproductive competition, which guides producers as a key criterion for selecting males [45, 46].
\n
By remaining in coexistence conditions, one of the males tends to monopolize access to the females in estrus in order to ensure their reproductive success, being this considered the dominant male [47], which initiates a display of dominance behaviors, such as competition due to access to food, increased physical activity and hoarding of better resting places, while the subordinate male at the hierarchical level initiates evasive or submissive behaviors and sexual behavior is characterized by opportunistic-type strategies [48]. The dominant male is characterized by having a more aggressive behavior compared to the rest of the males and is also the one with the highest sexual activity [49].
\n
Among the activities carried out mainly by the dominant male, the increase in vocalizations, head movements, tapping, lunges and displacement and protection of the female in heat (tending) from other males stands out [42], thus reaching inhibit the sexual behavior of subordinate males [36].
\n
According to Mainguy et al. [41], the establishment of the dominance position is accentuated with the secondary sexual characteristics, which is also related to the age and body weight of the animal [50], helping to strengthen the male’s hierarchical position and social structure within the herd. As they reach sexual maturity, the frequency of mounting with ejaculate, the performance of riding and the production of semen in dominant bucks compared to subordinate’s increases [36].
\n
\n
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2.4.3 Measurement of sexual behavior in bucks
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To determine the potential as a possible male, it is necessary to establish tests that allow the identification and categorization of males according to their score, taking into account comprehensively both their physical characteristics, such as weight, body condition, and sexual behavior [1].
\n
Assessments to determine mount efficiency in bucks typically consist of exposing a male to a female in estrus, for a period of time ranging from 15 to 20 minutes to 1 hour in a pen without distractors [51]. During this period, an observer keeps track of the amount of sexual behavior. They are rapid, practical and inexpensive tests that allow identifying the willingness of the male to serve the female and together with this, discard males with unsuitable profiles within a reproductive program in natural mating in the shortest possible time [52].
\n
In tests of reproductive capacity, motivation is linked to the animal’s libido and for this reason some authors recommend the use of more than one female in estrus [49]. Other factors that influence the performance of males in the evaluation of reproductive capacity are the breed of the animal, season of the year, age, the sexual experience they have and the hierarchical position that the male occupies [33].
\n
For the evaluation of sexual behavior in bucks there are different strategies that can be implemented to determine acts of courtship and mating acts. The reaction time test allows us to identify the time it takes for the male to achieve the first mount with ejaculate and thus have an estimate of the libido of the evaluated male [11].
\n
The service ability test is one of the most widely used tests. It consists of placing the male before one or several females in heat for a certain period, usually between 15 to 60 minutes in a pen. During this period an observer counts the number of interactions between the male and the female (s), which can be optionally rated or, only indicate the frequency with which the courtship acts, the mounts or the ejaculations occur as the case may be [42].
\n
Observations can be made individually or in groups. Individual observations should be made in the absence of other males, while group observations should take into account that they must be of similar ages to give accuracy to the test, seeking to carry out at least three tests to estimate service capacity [25, 53, 54].
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\n\n',keywords:"bucks, testosterone, sexual behavior, reproduction in goats, nutrition",chapterPDFUrl:"https://cdn.intechopen.com/pdfs/73371.pdf",chapterXML:"https://mts.intechopen.com/source/xml/73371.xml",downloadPdfUrl:"/chapter/pdf-download/73371",previewPdfUrl:"/chapter/pdf-preview/73371",totalDownloads:137,totalViews:0,totalCrossrefCites:0,totalDimensionsCites:0,hasAltmetrics:0,dateSubmitted:"January 27th 2020",dateReviewed:"July 27th 2020",datePrePublished:"October 22nd 2020",datePublished:"January 20th 2021",dateFinished:"September 29th 2020",readingETA:"0",abstract:"The exploitation of small ruminants (goat and sheep) has always been linked to the development of human civilizations, where they have mainly fed on their derived products such as milk and meat. Currently, the sheep population is around 1 billion head concentrated above 50% in three countries, China, Australia, and New Zealand, contrary to goats with around 720 million heads, distributed mainly in Asia, Africa, and South America. Both species have similar characteristics in some anatomical aspects (a pair of nipples), gestation period (150 days), and presence of seasonal anestrus, differing in terms of magnitude and depth and presence of the male effect. However, they are completely different in feeding habits, nutrient needs, and grazing systems, with differences in terms of the female’s reproductive tract, among other characteristics. Currently, the study of reproduction has intensified over the years in the goats and its counterpart that is the buck. Therefore, in the following topics, the importance of global reproduction of the goat will be discussed, considering that progress has been made today in the application of third generation reproductive techniques and that today they are already consolidated and developed in the bovine species.",reviewType:"peer-reviewed",bibtexUrl:"/chapter/bibtex/73371",risUrl:"/chapter/ris/73371",book:{slug:"animal-reproduction-in-veterinary-medicine"},signatures:"Fernando Sánchez Dávila and Gerardo Pérez Muñoz",authors:[{id:"201830",title:"Dr.",name:"Fernando",middleName:"Sanchez",surname:"Davila",fullName:"Fernando Davila",slug:"fernando-davila",email:"fernando_sd3@hotmail.com",position:null,institution:{name:"Universidad Autónoma de Nuevo León",institutionURL:null,country:{name:"Mexico"}}},{id:"327566",title:"MSc.",name:"Gerardo",middleName:null,surname:"Perez-Muñoz",fullName:"Gerardo Perez-Muñoz",slug:"gerardo-perez-munoz",email:"gerardopemu@gmail.com",position:null,institution:{name:"Universidad Autónoma de Nuevo León",institutionURL:null,country:{name:"Mexico"}}}],sections:[{id:"sec_1",title:"1. Introduction",level:"1"},{id:"sec_2",title:"2. Reproduction in goats",level:"1"},{id:"sec_2_2",title:"2.1 Perspectives and advances in the study of the estrous cycle of the goat",level:"2"},{id:"sec_3_2",title:"2.2 Will it be possible to improve the parameters of presence of estrus and pregnancy using hormones in the coming years?",level:"2"},{id:"sec_4_2",title:"2.3 Advances and use of the male effect, a case study until today!",level:"2"},{id:"sec_5_2",title:"2.4 Male social hierarchy and its impact on reproduction",level:"2"},{id:"sec_5_3",title:"2.4.1 Cortisol and stress physiology in bucks",level:"3"},{id:"sec_6_3",title:"2.4.2 Social factors and sexual behavior in males",level:"3"},{id:"sec_7_3",title:"2.4.3 Measurement of sexual behavior in bucks",level:"3"}],chapterReferences:[{id:"B1",body:'\nMellado M. Técnicas para el manejo reproductivo de las cabras en agostadero. Tropical and Subtropical Agroecosytem. 2008;9:47-63\n'},{id:"B2",body:'\nGrizelj J, Špoljarić B, Dobranić T, Lojkić M, Dávila FS, Samardžija M, et al. Efficiency analysis of standard and day 0 superovulatory protocols in Boer breed goats. Veterinarski arhiv. 2017;87:473-486\n'},{id:"B3",body:'\nAbecia JA, Forcada F, González-Bulnes A. Pharmaceutical control of reproduction in sheep and goats. Veterinary Clinics: Food Animal Practice. 2011;27:67-79\n'},{id:"B4",body:'\nDias JCO, Veloso CM, Santos MCDR, Oliveira CTSAMD, Silveira CO, Iglesias E, et al. Seasonal variation in the reproductive activity of male goats raised under tropical climate conditions. Revista Brasileira de Zootecnia. 2017;46:192-201\n'},{id:"B5",body:'\nMendieta ES, Delgadillo JA, Flores JA, Flores MJ, Nandayapa E, Vélez LI, et al. Subtropical goats ovulate in response to the male effect after a prolonged treatment of artificial long days to stimulate their milk yield. Reproduction in Domestic Animals. 2018;53:955-962\n'},{id:"B6",body:'\nArredondo AJG, Gómez AG, Vázquez-Armijo JF, Ledezma-Torres RA, Bernal-Barragán H, Sánchez-Dávila F. Status and implementation of reproductive technologies in goats in emerging countries. African Journal of Biotechnology. 2015;14:719-727\n'},{id:"B7",body:'\nGonzalez-Bulnes A, Menchaca A, Martin GB, Martinez-Ríos P. Seventy years of progestagen treatments for management of the sheep oestrous cycle: Where we are and where we should go. Reproduction, Fertility, and Development. 2020;32:441-452\n'},{id:"B8",body:'\nMenchaca A, Neto CDS, Cuadro F. Estrous synchronization treatments in sheep: Brief update. Revista Brasileira de Reproducción Animal. 2017;41:340-344\n'},{id:"B9",body:'\nKhan UM, Khan AM, Khan UM, Selamoğlu Z. Effects of seasonal factorsin the goats’ reproductive efficiency. Turkish Journal of Agriculture-Food Science and Technology. 2019;7:1937-1940\n'},{id:"B10",body:'\nMontes-Quiroz GL, Sánchez-Dávila F, Domínguez-Díaz D, Vázquez-Armijo JF, Grizelj J, Ledezma-Torres RA, et al. Influence of eCG and breed on the number of oocytes collected and the production of in vitro embryos of young goats during the reproductive season. Tropical Animal Health and Production. 2019;51:2521-2527\n'},{id:"B11",body:'\nMuñoz GP, Barragán HB, Torres RAL, Morón RU, Dávila FS. Dominancia social sobre comportamiento sexual y calidad seminal en machos cabríos jóvenes criados en parejas durante la estación reproductiva. Revista Academica de Ciencia Animal. 2019;17(Suppl 1):323-326\n'},{id:"B12",body:'\nBrunet AG, Santiago-Moreno J, Toledano-Diaz A, Lopez-Sebastian A. Reproductive seasonality and its control in Spanish sheep and goats. Tropical and Subtropical Agroecosystems. 2011;15:847-870\n'},{id:"B13",body:'\nCamacho M. Control of estrous cycle and superovulation in goats [dissertation Ph.D.]. Gottingen, Germany: George August Universitaet; 2020. p. 96\n'},{id:"B14",body:'\nYu XJ, Wang J, Bai YY. Estrous synchronization in ewes: The use of progestogens and prostaglandins. Acta Agriculturae Scandinavica Section A Animal Science. 2018;68:219-230\n'},{id:"B15",body:'\nOmontese BO, Rekwot PI, Ate IU, Ayo JO, Kawu MU, Rwuaan JS, et al. An update on oestrus synchronisation of goats in Nigeria. Asian Pacific Journal of Reproduction. 2016;5:96-101\n'},{id:"B16",body:'\nSimões J. Recent advances on synchronization of ovulation in goats, out of season, for a more sustainablen production. Asian Pacific Journal of Reproduction. 2015;4:157-165\n'},{id:"B17",body:'\nMerlos-Brito M, Martínez-Rojero R, Torres-Hernández R, Mastache-Lagunas A, Gallegos-Sánchez J. Evaluación de características productivas en cabritos Boer x local, Nubia x local y locales en trópico seco de Guerrero, México. Veterinaria México. 2008;33:323-333\n'},{id:"B18",body:'\nMeza-Herrera CA, Romero-Rodríguez CA, Nevárez-Dominguez A, Flores-Hernández A, Cano-Villegas O, Macías-Cruz U, et al. The Opuntia effect and the reactivation of ovarian function and blood metabolite concentrations of anestrous goats exposed to active males. Animals. 2019;9:1-11\n'},{id:"B19",body:'\nMontes-Quiroz GL, Sánchez-Dávila F, Grizelj J, Bernal-Barragán H, Vazquez-Armijo JF, Bosque-González ASD, et al. The reinsertion of controlled internal drug release devices in goats does not increase the pregnancy rate after short oestrus synchronization protocol at the beginning of the breeding season. Journal of Applied Animal Research. 2018;46:714-719\n'},{id:"B20",body:'\nBedos M, Muñoz AL, Orihuela A, Delgadillo JA. The sexual behavior of male goats exposed to long days is as intense as during their breeding season. Applied Animal Behaviour Science. 2016;184:35-40\n'},{id:"B21",body:'\nDelgadillo JA, Vélez LI, Flores JA. Continuous light after a long-day treatment is equivalent to melatonin implants to stimulate testosterone secretion in Alpine male goats. Animal. 2016;10:649-654\n'},{id:"B22",body:'\nAraya J, Bedos M, Duarte G, Hernández H, Keller M, Chemineau P, et al. Maintaining bucks over 35 days after a male effect improves pregnancy rate in goats. Animal Production Science. 2017;57:2066-2071\n'},{id:"B23",body:'\nEspinoza-Flores LA, Andrade-Esparza JD, Hernández H, Zarazaga LA, Abecia JA, Chemineau P, et al. Male effect using photostimulated bucks and nutritional supplementation advance puberty in goats under semi-extensive management. Theriogenology. 2020;143:82-87\n'},{id:"B24",body:'\nZarazaga LA, Gatica MC, Hernández H, Keller M, Chemineau P, Delgadillo JA, et al. The reproductive response to the male effect of 7-or 10-month-old female goats is improved when photostimulated males are used. Animal. 2019;13:1658-1166\n'},{id:"B25",body:'\nMadrid-Bury E, González-Stagnaro C, Aranguren-Méndez J, Yanez F, Quintero-Moreno A. Sexual behavior of “Criollo Limonero” bulls. Revista Facultad de Agronomáía. 2011;28:505-513\n'},{id:"B26",body:'\nChemineau P, Bodin L, Migaud M, Thiéry JC, Malpaux B. Neuroendocrine and genetic control of seasonal reproduction in sheep and goats. Reproduction in Domestic Animals. 2010;45:42-49\n'},{id:"B27",body:'\nZarazaga LA, Gatica MC, Hernández H, Gallego-Calvo L, Delgadillo JA, Guzmán JL. The isolation of females from males to promote a later male effect is unnecessary if the bucks used are sexually active. Theriogenology. 2017;95:42-47\n'},{id:"B28",body:'\nCarrillo E, Véliz FG, Flores JA, Delgadillo JA. El decremento en la proporción macho-hembras no disminuye la capacidad para inducir la actividad estral de cabras anovulatorias. Vol. 45. Técnica Pecuaria en México; 2007. pp. 319-328\n'},{id:"B29",body:'\nVéliz-Deras FG, Monroy LV, Cabrera JF, Moreno GD, Massot PP, Malpaux B, et al. La presencia del macho en un grupo de cabras anestricas no impide su respuesta estral a la introducción de un nuevo macho. Veterinaria Mexico. 2004;35:169-178\n'},{id:"B30",body:'\nVielma J, Terrazas A, Véliz FG, Flores JA, Hernandez H, Duarte G, et al. Las vocalizaciones de machos cabríos no estimulan la secreción de la LH ni la ovulación en las cabras anovulatorias. Revista Mexicana de Ciencias Pecuarias. 2008;46:25-36\n'},{id:"B31",body:'\nSantiago-Miramontes D, de los Ángeles M, Marcelino-León S, Luna-Orozco JR, Rivas-Muñoz R, Rodríguez-Martínez R, et al. La presencia de hembras estrogenizadas al momento del efecto macho induce la actividad estral de cabras en el semidesierto mexicano. Revista Chapingo Serie Ciencias Forestales y del Ambiente. 2011;17:77-85\n'},{id:"B32",body:'\nDelgadillo JA, Gelez H, Ungerfeld R, Hawken PA, Martin GB. The ‘male effect’ in sheep and goats—Revisiting the dogmas. Behavioural Brain Research. 2009;200:304-314\n'},{id:"B33",body:'\nOrihuela A. Ram’s sexual behavior. Review. Revista Mexicana de Ciencias Pecuarias. 2014;5:49-89\n'},{id:"B34",body:'\nGiriboni J, Lacuesta L, Damián JP, Ungerfeld R. Grouping previously unknown bucks is a stressor with negative effects on reproduction. Tropical Animal Health and Production. 2015;47:317-322\n'},{id:"B35",body:'\nLacuesta L, Ungerfeld R. Sexual performance and stress response of previously unknown rams after grouping them in dyads. Animal Reproduction Science. 2012;134:158-163\n'},{id:"B36",body:'\nSánchez-Dávila F, Barragán HB, del Bosque-González AS, Ungerfeld R. Social dominance affects the development of sexual behaviour but not semen output in yearling bucks. Theriogenology. 2018;110:168-174\n'},{id:"B37",body:'\nKikusui T, Winslow JT, Mori Y. Social buffering: Relief from stress and anxiety. Philosophical Transactions of the Royal Society B. 2006;361:2215-2228\n'},{id:"B38",body:'\nVas J, Chojnacki R, Kjoren M, Lingwa C, Andersen I. Social interactions, cortisol and reproductive success of domestic goats (Capra hircus) subjected to different animal densities during pregnancy. Applied Animal Behavior Science. 2013;147:117-126\n'},{id:"B39",body:'\nIyasere O, James I, Williams T, Daramola J, Lawal K, Oke O, et al. Behavioural and physiological responses of West African Dwarf Goat dams and kids subjected to short-term separation. Tropical and Subtropical Agroecosytem. 2018;51:5-11\n'},{id:"B40",body:'\nSolano J, Galindo F, Orihuela A, Galina CS. The effect of social rank on the physiological response during repeated stressful handling in Zebu cattle (Bos indicus). Physiology of Behaviour. 2004;82:679-683\n'},{id:"B41",body:'\nMainguy J, Côté SD, Cardinal E, Houle M. Mating tactics and mate choice in relation to age and social rank in male mountain goats. Journal of Mammalogy. 2008;89:626-635\n'},{id:"B42",body:'\nChenoweth PJ. Sexual behavior of the bull: A review. Journal of Dairy Science. 1983;66:173-179\n'},{id:"B43",body:'\nHernández Z. La caprinocultura en el marco de la ganadería poblana (México): Contribución de la especie caprina y sistemas de producción. Archivos de Zootecnia. 2000;49:341-352\n'},{id:"B44",body:'\nMiranda-de la Lama G, Mattielo S. The importance of social behaviour for goat welfare in livestock farming. Small Ruminant Research. 2010;90:1-10\n'},{id:"B45",body:'\nClutton-Brock TH, Huchard E. Social competition and selection in males and females. Philosophical Transactions of the Royal Society B. 2013;368:1-15\n'},{id:"B46",body:'\nClutton-Brock T. Reproductive competition and sexual selection. Philosophical Transactions of the Royal Society B. 2017;372:1-10\n'},{id:"B47",body:'\nUngerfeld R. Managing reproductive seasonality in small ruminants. Archivos Latinoamericanos de Producción Animal. 2016;24:111-116\n'},{id:"B48",body:'\nMainguy K, Côté SD. Age and state-dependent reproductive effortin male mountain goats (Oreamnos americanus). Behavior Ecology Society. 2007;62:935-943\n'},{id:"B49",body:'\nKatz LS. Variation in male sexual behavior. Animal Reproduction Science. 2008;105:64-71\n'},{id:"B50",body:'\nPelletier F, Festa-Bianchet M. Sexual selection and social rank in bighorn rams. Animal Behavior. 2006;71:649-655\n'},{id:"B51",body:'\nGiriboni J, Lacuesta L, Ungerfeld R. Continuous contact with females in estrus throughout the year enhances testicular activity and improves seminal traits of male goats. Theriogenology. 2017;87:284-289\n'},{id:"B52",body:'\nImwalle DB, Katz LS. Development of sexual behavior over several serving capacity tests in male goats. Applied Animal Behaviour Science. 2004;89:315-319\n'},{id:"B53",body:'\nVejarano OA, Sanabria R, Trujillo G. Diagnostic of bulls reproduction capability from three livestock farms of the upper Magdalena. Revista MVZ Córdoba. 2005;10:648-662\n'},{id:"B54",body:'\nSaunders FC, McElligott AG, Safi K, Hayden TJ. Mating tactics of male feral goats (Capra hircus): Risks and benefits. Acta Ethologica. 2005;8:103-110\n'}],footnotes:[],contributors:[{corresp:"yes",contributorFullName:"Fernando Sánchez Dávila",address:"fernando_sd3@hotmail.com",affiliation:'
Laboratorio de Reproducción Animal, Unidad Académica “Marín”, Facultad de Agronomía, Universidad Autónoma de Nuevo León, México
Estudiante de la maestría del posgrado conjunto de la Facultad de Agronomía-Facultad de Medicina Veterinaria y Zootecnia, UANL, México
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Borrego",slug:"juan-j.-borrego"},{id:"71146",title:"Dr.",name:"Alejandro",middleName:null,surname:"Labella",fullName:"Alejandro Labella",slug:"alejandro-labella"},{id:"71148",title:"Dr.",name:"Concepcion",middleName:null,surname:"Berbel",fullName:"Concepcion Berbel",slug:"concepcion-berbel"},{id:"71149",title:"Dr.",name:"Manuel",middleName:null,surname:"Manchado",fullName:"Manuel Manchado",slug:"manuel-manchado"},{id:"71151",title:"Dr.",name:"Dolores",middleName:null,surname:"Castro",fullName:"Dolores Castro",slug:"dolores-castro"}]},{id:"24079",title:"Bacteria Isolated From Diseased Wild and Farmed Marine Fish in Greece",slug:"bacteria-isolated-from-diseased-wild-and-farmed-marine-fish-in-greece",signatures:"Mary Yiagnisis and Fotini Athanassopoulou",authors:[{id:"71057",title:"Dr.",name:"Mary",middleName:null,surname:"Yiagnisis",fullName:"Mary Yiagnisis",slug:"mary-yiagnisis"}]},{id:"24080",title:"The Immune System Drugs in Fish: Immune Function, Immunoassay, Drugs",slug:"the-immune-system-drugs-in-fish-immune-function-immunoassay-drugs",signatures:"Cavit Kum and Selim Sekkin",authors:[{id:"68075",title:"Dr.",name:"Cavit",middleName:null,surname:"Kum",fullName:"Cavit Kum",slug:"cavit-kum"}]},{id:"24081",title:"Antibacterial Drugs in Fish Farms: Application and Its Effects",slug:"antibacterial-drugs-in-fish-farms-application-and-its-effects",signatures:"Selim Sekkin and Cavit Kum",authors:[{id:"68279",title:"Dr.",name:"Selim",middleName:null,surname:"Sekkin",fullName:"Selim Sekkin",slug:"selim-sekkin"}]}]}]},onlineFirst:{chapter:{type:"chapter",id:"67314",title:"Pathogenesis of Periodontal Disease",doi:"10.5772/intechopen.86548",slug:"pathogenesis-of-periodontal-disease",body:'\n
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1. Introduction
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Periodontitis is a globally widespread pathology of the human oral cavity. Approximately 10% of the global adult population is highly vulnerable to severe periodontitis, and 10–15% appears to be completely resistant to it, while the remainder varies between these two situations [1]. Periodontitis is a major public health problem due to its high prevalence, as well as because it may lead to tooth loss and disability, negatively affect chewing function and aesthetics, be a source of social inequality, and impair the quality of life. Periodontitis accounts for a substantial proportion of edentulism and masticatory dysfunction, results in significant dental care costs, and has a plausible negative impact on general health [2].
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2. Periodontal support tissues
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The periodontium is a complex of tissues with blood vessels, nerves, and bundles of fibers, which provide nutrition and sensibility, supporting and investing the tooth. The periodontium has the potential for regeneration and remodeling throughout life, which allows the primary dentition to be transient and to be replaced by the permanent dentition [3, 4]. It is important to understand that each of the periodontal tissues has a very specialized structure and that these structural characteristics directly define the function. In fact, the proper functioning of the periodontium is only achieved through the structural integrity and interaction between its components [4].
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The periodontium is one of the morphofunctional components of the stomatognathic system, and its “design” not only responds to intrinsic functions related to nutrition or the subjection of the tooth but also to functions integrated within the physiology of the stomatognathic system [5]. The main function of the periodontium is to join the tooth to the bone tissue and maintain integrity on the surface of the masticatory mucosa of the oral cavity [6]. The periodontium includes four tissues located near the teeth: (1) the alveolar bone (AB), (2) root cementum (CR), (3) periodontal ligament (PL), and (4) gingiva (G) (e.g., Figure 1a) [4, 7, 8].
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Figure 1.
Periodontal tissues. (a) Tissues that support the tooth include the alveolar bone (AB), root cementum (RC), periodontal ligament (PL), and gingiva (G). (b) Forms of cementum: acellular afibrillar cementum (AAC), acellular extrinsic fibers cementum (AEFC), cellular mixed stratified cementum (CMSC), and cellular intrinsic fibers cementum (CIFC). (c) Bundles of collagen fibers: crestal alveolar fibers (CAF), horizontal fibers (HF), oblique fibers (OF), and apical fibers (AF). (d) Parts of the gingiva: free gingiva (FG), interdental gingiva (IG), and attached gingiva (AG).
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2.1 Alveolar bone
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The alveolar bone, together with the root cementum and the periodontal ligament, constitutes the tooth insertion apparatus, whose main function is to distribute the forces generated by chewing and other contacts [6]. The maxilla and mandible of the adult human can be subdivided into two parts: (a) the alveolar process that involves in housing the roots of the erupted teeth and (b) the basal body that does not involve in housing the roots [8].
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The alveolar process is the bone of the jaws that contain the sockets (alveoli) of the teeth. It consists of outer cortical plates (buccal, lingual, and palatal) of the compact bone, a central spongiosa, and bone lining the alveolus (alveolar bone) [4]. The alveolar process is dependent on the teeth as they develop and remodel with their formation and eruption. Therefore, the shape, location, and function of the teeth determine its morphology [8]. The periodontal ligament contains progenitor cells that can differentiate into osteoblasts for the maintenance and repair of the alveolar bone. However, in the absence of the tooth, it is lost. These characteristics suggest that the regulatory mechanisms are important for the alveolar bone, so there is an interdependence of the periodontal tissues, which work together as a unit [4].
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2.2 Root cementum
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The root cementum is an avascular mineralized connective tissue covering the entire root surface, forming the interface between the root dentine and the periodontal ligament [6, 7]. In addition, the root cementum plays important roles in nourishing the tooth as well as in stabilizing the tooth via the attachment to the periodontal ligament. This enables the tooth to maintain its relationship to adjacent and opposing teeth [3]. Unlike the bone, the root cementum does not contain blood or lymphatic vessels, lacks innervation, and does not undergo remodeling or physiological resorption, but it is characterized by the fact that continues to be deposited throughout life [6]. The composition of cementum contains about 50% mineral (substituted apatite) and 50% organic matrix. Type I collagen is the predominant organic component, constituting up to 90% of the organic matrix. Other collagens associated with cementum include type I, III, V, VI, XII, and XIV [4].
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Cementum performs different functions: it fixes the main fibers of the periodontal ligament to the root and contributes in the repair process when the root surface has been damaged [6]. Cementum has been classified as cellular and acellular cementum depending on the presence and absence of cementocytes, further grouped into intrinsic and extrinsic fiber cementum depending on the presence of collagen fibers formed by cementoblasts or by fibroblasts, respectively [8]. There are different forms of cementum (e.g., Figure 1b): (1) acellular afibrillar cementum (AAC), (2) acellular extrinsic fiber cementum (AEFC), (3) cellular mixed stratified cementum (CMSC), and (4) cellular intrinsic fibers cementum (CIFC) [4, 6].
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2.3 Periodontal ligament
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The periodontal ligament is the soft and specialized connective tissue situated between the cementum covering the root of the tooth and the bone forming the socket wall (alveolodental ligament) [4]. The periodontal ligament consists of cells and an extracellular compartment comprising collagenous and noncollagenous matrix constituents. The cells include osteoblasts and osteoclasts, fibroblasts, epithelial cell rests of Malassez, monocytes and macrophages, undifferentiated mesenchymal cells, and cementoblasts and odontoclasts. The extracellular compartment consists mainly of well-defined collagen fiber bundles embedded in an amorphous background material, known as ground substance [4, 8]. These bundles of collagen fibers can be classified into the following groups, according to their disposition (e.g., Figure 1c): crestal alveolar fibers (CAF), horizontal fibers (HF), oblique fibers (OF), and apical fibers (AF) [6].
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The main function of the periodontal ligament is to support the teeth in their sockets and at the same time allow them to withstand the considerable forces of mastication. In addition, the periodontal ligament has the capacity to act as a sensory receptor necessary for the proper positioning of the jaws during mastication, and, very importantly, it is a cell reservoir for tissue homeostasis, regeneration, and repair [4].
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2.4 Gingiva
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Gingiva is a portion of the oral mucosa covering the tooth-carrying part of the alveolar bone and the cervical neck of the tooth. Three parts of the gingiva can be distinguished (e.g., Figure 1d): (1) free gingiva (FG), (2) interdental gingiva (IG), and (3) attached or inserted gingiva (AG) [6]. Histologically, the epithelial component of the gingiva shows regional morphological variations that reflect the adaptation of the tissue to the tooth and alveolar bone. These include the epithelium that covers a connective tissue, chorion, or lamina propria. A keratinized stratified squamous epithelium protects the lamina propria of the gingiva on its masticatory surfaces and a nonkeratinized epithelium protects the lamina propria on its crevicular and junctional surfaces [6, 9]. The junctional epithelium plays a crucial role since it essentially seals off periodontal tissues from the oral environment. Its integrity is thus essential for maintaining a healthy periodontium. Periodontal disease sets in when the structure of the junctional epithelium starts to fail, an excellent example of how structure determines function [4].
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During pathological conditions, such as inflammation, the periodontal connective tissues, including the gingiva, undergo many changes. Clinically detected gingival overgrowth is one of the alterations that occurs in chronic periodontitis. It is caused by a variety of etiological factors and is exacerbated by local bacterial biofilm accumulation, because the periodontopathogen products act on the gingival tissues activating cellular events that induce the alteration of connective tissue homeostasis and the destruction of the alveolar bone [9]. Likewise, junctional epithelial cells differ considerably from those of the gingival epithelium. There are polymorphonuclear leukocytes and monocytes that pass from the subepithelial connective tissue through the junctional epithelium and into the gingival sulcus. The mononuclear cells, together with molecules they secrete and others originating from junctional epithelial cells, blood and tissue fluid, represent the first line of defense in the control of the perpetual microbial challenge [4].
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3. Periodontal pathogenesis
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Periodontitis is a chronic multifactorial disease characterized by an inflammation of the periodontal tissue mediated by the host, which is associated with dysbiotic plaque biofilms, resulting in the progressive destruction of the tooth-supporting apparatus and loss of periodontal attachment [1, 10]. The bacterial biofilm formation initiates gingival inflammation; however, periodontitis initiation and progression depend on dysbiotic ecological changes in the microbiome in response to nutrients from gingival inflammatory and tissue breakdown products and anti-bacterial mechanisms that attempt to contain the microbial challenge with in the gingival sulcus area once inflammation has initiated. This leads to the activation of several key molecular pathways, which ultimately activate host-derived proteinases that enable loss of marginal periodontal ligament fibers, apical migration of the junctional epithelium, and allows apical spread of the bacterial biofilm along the root surface [1]. Therefore, the primary features of periodontitis include the loss of periodontal tissue support, manifested through clinical attachment loss and radiographically assessed alveolar bone loss, presence of periodontal pocketing, and gingival bleeding [10].
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3.1 Periodontal histopathology
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The development of gingivitis and periodontitis can be divided into a series of stages: initial, early, established, and advanced lesions (e.g., Figure 2) [11, 12]. The initial lesion begins 2–4 days after the accumulation of the microbial plaque. During the initial lesion, an acute exudative vasculitis in the plexus of the venules lateral to the junctional epithelium, migration of polymorphonuclear (PMN) cells through the junctional epithelium into the gingival sulcus, co-exudation of fluid from the sulcus, and the loss of perivascular collagen were observed. The early injury develops within 4–10 days. This lesion is characterized by a dense infiltrate of T lymphocytes and other mononuclear cells, as well as by the pathological alteration of the fibroblasts [6, 11, 12, 13].
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Figure 2.
Histopathological lesions of periodontal pathogenesis. Initial, early, established, and advanced lesions of the development of gingivitis and periodontitis.
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Subsequently, the established lesion develops within 2–3 weeks. This lesion is dominated by activated B cells (plasma cells) and accompanied by further loss of the marginal gingival connective tissue matrix, but no bone loss is yet detectable. Several PMN continue to migrate through the junctional epithelium, and the gingival pocket is gradually established. Finally, in the advanced lesion, plasma cells continue to predominate as the architecture of the gingival tissue is disturbed, together with the destruction of the alveolar bone and periodontal ligament. It is characterized by a conversion of junctional epithelium to the pocket epithelium, formation of denser inflammatory infiltrate composed of plasma cells and macrophages, loss of collagen attachment to the root surface, and resorption of the alveolar bone [6, 11, 12, 13].
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3.2 Immune responses in the pathogenesis of periodontal disease
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In normal health conditions, periodontal tissues are capable of coping with the presence of bacteria through several mechanisms of the host immune system (e.g., Figure 3) [14]. However, when the balance between the infection control mechanisms and the subgingival biofilm is lost [15], which includes Porphyromonas gingivalis, Aggregatibacter actinomycetemcomitans, Tannerella forsythia, and Treponema denticola [16], innate, inflammatory, and adaptive reactions are triggered. These processes result in the destruction of the tissues that surround and support the teeth, and eventually in the loss of tissues, bones and finally, of the teeth [17].
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Figure 3.
Innate and adaptive immune response during periodontal disease (description in the text).
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3.2.1 Innate immune response in periodontal disease
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The most important characteristic of periodontitis is the inflammatory reabsorption of the tooth-supporting alveolar bone due to the uncontrolled host immune response against periodontal infection, since the destructive events, which lead to the irreversible phenotype of periodontal disease, are the result of the persistence of a chronic and exacerbated inflammatory immune response [18]. Inflammation is a process of the innate immune system activation, in response to exogenous and endogenous factors, such as infection by microorganisms, tissue stress, and injuries. Inflammation is a protective response, characterized by its cardinal signs, such as redness, swelling, heat, pain, and disrupted function [19]. The inflammatory response consists of four main components: (1) endogenous or exogenous factors, such as molecular patterns associated with pathogens (PAMP) and damage (DAMP), which are derived from bacteria, viruses, fungi, parasites, and cell damage, as well as toxic cellular components or any other harmful condition; (2) cellular receptors that recognize these molecular patterns (PRR), for example, Toll-like receptors (TLR); (3) proinflammatory mediators, such as cytokines, chemokines, the complement system, etc.; and (4) target cells and tissues, where these proinflammatory mediators act [20, 21]. The inflammatory response is mainly characterized by four successive phases: (1) silent phase, where the cells synthesize and release the first proinflammatory mediators; (2) vascular phase, characterized by an increase in vascular permeability and dilatation; (3) cellular phase, characterized by the infiltration of inflammatory cells at the site of injury; and (4) the resolution of the inflammatory response [22].
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The inflammatory immune response is triggered by the interaction of resident cells with the bacterial biofilm attached to the tooth surface. Bacterial biofilm attaches to the tooth surface, making it impossible for the immune system to eradicate the infecting microorganisms efficiently, perpetuating the insult to the periodontal tissues [18]. The junctional epithelium is the first periodontal structure to face the bacterial challenge [23]. Bacteria are capable to cross the junctional epithelium and pass to the gingival conjunctive tissue, where they stimulate the gingival epithelial cells and fibroblasts to trigger the initial inflammatory responses [24]. These resident periodontal cells detect bacterial PAMP, such as lipopolysaccharide (LPS) [25], which binds to the Toll-like receptors (TLR4/2), triggering the recruitment of several protein kinases in the cytoplasmic end of the receptors, ultimately causing the activation of proinflammatory transcription factors, such as nuclear factor kappa B (NFκB) and activator protein 1 (AP-1) [26], which induces the synthesis and release of mediators to trigger the inflammatory response. Likewise, the gingival fibroblasts and the periodontal ligament are responsible for the destruction and disorganization of the fibrous component of the extracellular matrix of the periodontal tissue by increasing the local production and the activity of the matrix metalloproteinases (MMPs) [27, 28].
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The periodontal lesion is initiated as acute inflammation characterized by increased numbers of neutrophils migrating into the gingival crevice through the junctional epithelium, which have the de novo biosynthetic capacity for chemokines and cytokines with proinflammatory, anti-inflammatory, or immunoregulatory properties. Neutrophils, through the release of chemokines, can induce the recruitment of interleukin-17-producing CD4-positive T-helper 17 cells to sites of infection or inflammation. In addition, they can promote the survival, proliferation, and development of B cells into antibody-secreting plasma cells. Likewise, it was shown that activated neutrophils express membrane-bound receptor activator of nuclear factor kappa Β ligand (RANKL), a key osteoclastogenic cytokine and, thereby able of inducing osteoclastic bone resorption [29]. These recent concepts suggest that neutrophils could contribute to periodontitis not only by initiating the lesion but also by participating in its progression, by recruiting T-helper 17 cells or promoting the accumulation of B cells and plasma cells in the established and advanced lesions.
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Macrophages are an important source of proinflammatory and potentially destructive molecules for tissues, such as interleukin-1 (IL-1), tumor necrosis factor alpha (TNF-α), MMP, and prostaglandin E2 [30], which play an important role and are elevated in the gingival tissue and in the gingival crevicular fluid of patients with chronic periodontitis [28]. Therefore, studies have shown a direct correlation of macrophage infiltration with the severity of periodontal disease [31], contributing greatly to the intensification of the degradation of the collagen matrix in the connective periodontal tissue [32, 33]. These macrophages may undergo a classical (M1) or alternative (M2) activation. M1 macrophages are induced by microbial agents (e.g., LPS) or by Th1 cytokines and show high phagocytic capacity and an increased expression of proinflammatory cytokines, costimulatory, and antimicrobial molecules. In contrast, M2 macrophages are induced by Th2 cytokines and secrete high levels of IL-10 and transforming growth factor beta 1 (TGF-β1). Therefore, they have immunoregulatory properties and promote cell proliferation and tissue regeneration [29, 34]. In periodontal inflammation models, macrophages share properties of both M1 and M2. However, M1 macrophages show a predominance over M2 macrophages, suggesting that M1 macrophages probably represent a subset associated with periodontitis [35, 36, 37].
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3.2.2 Adaptive immune response in periodontal disease
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When the inflammatory response becomes chronic, the lymphocytes of the adaptive immune system invade the periodontal tissues releasing inflammatory and immune molecular mediators, which alter the balance of bone metabolism, marking the transition from gingivitis to periodontitis [29]. The activation of lymphocytes requires two types of signals: a signal induced by the antigen receptor itself when recognizing its related antigen and a costimulatory signal by professional antigen-presenting cells (APCs) [22]. In gingivitis, the predominant APCs are CD14+ and CD83+ dendritic cells. While in the periodontitis, the predominant APCs are CD19+ and CD83+ B lymphocytes [38]. Therefore, the activation of adaptive immunity has a great influence on the bone loss in periodontitis, associated with B and T lymphocytes, since several studies have shown that these cells are the main cellular sources of activator of the κB ligand receptor of the nuclear factor (RANKL) during periodontal inflammation [39].
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RANKL is a cytokine member of the TNF family that can be bound or secreted to the membrane and stimulates the differentiation of osteoclasts, cell fusion, and activation that leads to bone resorption [40, 41]. Osteoblasts and stromal cells of the bone marrow predominantly express RANKL bound to the membrane, which induces osteoclastogenesis through cell contact with osteoclast precursors. Likewise, activated T and B cells produce both the membrane-bound and soluble RANKL forms. Soluble RANKL can induce osteoclastogenesis independently of direct contact between infiltrating lymphocytes and osteoclast precursors on the bone surface. However, 17 T-helper cells expressing RANKL, but not T-helper 1 cells, activate osteoclasts also by direct cell-cell contact [42]. In the alveolar bone, the RANKL/OPG/RANK system controls the balance of the bone metabolism [43]. RANKL is the osteoclasts activator and the molecular signal directly responsible for the bone resorption, which interacts with its associated receptor RANK on the surface of osteoclast and osteoclast precursors, which triggers its recruitment on the bone surface and cell fusion and activation [44]. Osteoprotegerin (OPG) is a soluble protein that has the ability to block the biological functions of RANKL by competitive inhibition [45]. In periodontitis, the increase in RANKL/OPG promotes the recruitment of osteoclast precursors, their fusion, and subsequent activation, leading to bone resorption [46].
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On the other hand, Th1 lymphocytes have a fundamental role in the establishment and progression of periodontitis, through the increase of IFN-γ levels [18]. Studies have shown that mice IFN-γ-deficient showed low levels of inflammatory cytokines and chemokines, as well as macrophages infiltrated in periodontal tissue, developing a less severe phenotype of alveolar bone destruction [47]. IL-1β and TNF-α are cytokines secreted by Th1 lymphocytes. TNF-α and IL-1β produce vasodilation, stimulate the activation of endothelial cells to increase the recruitment of immune cells, increase the chemokines production in most cell types, participate in the activation of neutrophils, and stimulate secretion and tissue activation of MMPs, among other functions. Although neither IL-1β nor TNF-α is directly involved in the stimulation of bone resorption, they indirectly promote bone destruction by stimulating sustained inflammation of the periodontal tissue [48]. Th2 lymphocytes are the main cellular source of IL-4, which promotes the change of class to the secretion of IgE in B cells and favors the alternative activation of macrophages in an IFN-γ-independent pathway. These effector functions of the Th2 lymphocytes negatively regulate the inflammatory and Th1 lymphocyte responses, so that the polarization of a Th2-type immune response in periodontitis may represent a damaged adaptive immune response [18, 49]. Finally, RANKL can also be secreted by Th17 lymphocytes, which in cooperation with inflammatory cytokines derived from Th1 lymphocytes are capable to tilt bone metabolism favoring bone resorption [50].
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4. Conclusion
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The main etiological factor of periodontal disease is the bacteria, which are capable of activating the innate immune response of the host inducing an inflammatory response. The evolution of this inflammatory response culminates in the destruction of periodontal tissues. For this reason, it is important to understand the different molecular and cellular mechanisms of the pathogenesis of periodontal disease, with the purpose of making an opportune diagnosis and appropriate treatment and prognosis.
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Acknowledgments
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Thanks to the authors who collaborated in the writing of this chapter: Dr. en C. José Luis Muñoz-Carrillo, Dra. Viridiana Elizabeth Hernández-Reyes, Dr. Oscar Eduardo García-Huerta, Dra. Karla Mariana Chávez-Ruvalcaba, Dra. en C. Francisca Chávez-Ruvalcaba, Dra. en C. María Isabel Chávez-Ruvalcaba, and Dra. Lizbeth Díaz-Alfaro, as well as the Universities involved, Cuauhtémoc University Aguascalientes, Autonomous University of Zacatecas, and Autonomous University of Aguascalientes for financial support for chapter publication.
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Conflict of interest
We have no conflict of interest related to this work.
\n',keywords:"periodontal tissues, biofilm, inflammatory response, innate and adaptive immunity, periodontitis",chapterPDFUrl:"https://cdn.intechopen.com/pdfs/67314.pdf",chapterXML:"https://mts.intechopen.com/source/xml/67314.xml",downloadPdfUrl:"/chapter/pdf-download/67314",previewPdfUrl:"/chapter/pdf-preview/67314",totalDownloads:1749,totalViews:0,totalCrossrefCites:1,dateSubmitted:"November 23rd 2018",dateReviewed:"April 26th 2019",datePrePublished:"June 6th 2019",datePublished:null,dateFinished:null,readingETA:"0",abstract:"Inflammation is a physiological response of the innate immune system against several endogenous or exogenous stimuli. Inflammation begins with an acute pattern; however, it can become chronic by activating the adaptive immune response through cellular and noncellular mechanisms. The main etiologic factor of periodontal disease is bacteria which substantially harbor the human oral cavity. The most common periodontal diseases are gingivitis and periodontitis, whose main characteristic is inflammation. The knowledge of how immune mechanisms and inflammatory responses are regulated is fundamental to understanding the pathogenesis of periodontal disease. The purpose of this chapter is to show the current panorama of the immunological mechanisms involved in the pathogenesis of periodontal disease.",reviewType:"peer-reviewed",bibtexUrl:"/chapter/bibtex/67314",risUrl:"/chapter/ris/67314",signatures:"José Luis Muñoz-Carrillo, Viridiana Elizabeth Hernández-Reyes, Oscar Eduardo García-Huerta, Francisca Chávez-Ruvalcaba, María Isabel Chávez-Ruvalcaba, Karla Mariana Chávez-Ruvalcaba and Lizbeth Díaz-Alfaro",book:{id:"8202",title:"Periodontal Disease",subtitle:"Diagnostic and Adjunctive Non-surgical Considerations",fullTitle:"Periodontal Disease - Diagnostic and Adjunctive Non-surgical Considerations",slug:"periodontal-disease-diagnostic-and-adjunctive-non-surgical-considerations",publishedDate:"February 5th 2020",bookSignature:"Nermin Mohammed Ahmed Yussif",coverURL:"https://cdn.intechopen.com/books/images_new/8202.jpg",licenceType:"CC BY 3.0",editedByType:"Edited by",editors:[{id:"210472",title:"Dr.",name:"Nermin",middleName:"Mohammed Ahmed",surname:"Yussif",slug:"nermin-yussif",fullName:"Nermin Yussif"}],productType:{id:"1",title:"Edited Volume",chapterContentType:"chapter",authoredCaption:"Edited by"}},authors:null,sections:[{id:"sec_1",title:"1. Introduction",level:"1"},{id:"sec_2",title:"2. Periodontal support tissues",level:"1"},{id:"sec_2_2",title:"2.1 Alveolar bone",level:"2"},{id:"sec_3_2",title:"2.2 Root cementum",level:"2"},{id:"sec_4_2",title:"2.3 Periodontal ligament",level:"2"},{id:"sec_5_2",title:"2.4 Gingiva",level:"2"},{id:"sec_7",title:"3. Periodontal pathogenesis",level:"1"},{id:"sec_7_2",title:"3.1 Periodontal histopathology",level:"2"},{id:"sec_8_2",title:"3.2 Immune responses in the pathogenesis of periodontal disease",level:"2"},{id:"sec_8_3",title:"3.2.1 Innate immune response in periodontal disease",level:"3"},{id:"sec_9_3",title:"3.2.2 Adaptive immune response in periodontal disease",level:"3"},{id:"sec_12",title:"4. Conclusion",level:"1"},{id:"sec_13",title:"Acknowledgments",level:"1"},{id:"sec_16",title:"Conflict of interest",level:"1"}],chapterReferences:[{id:"B1",body:'Papapanou PN, Sanz M, Buduneli N, Dietrich T, Feres M, Fine DH, et al. Periodontitis: Consensus report of workgroup 2 of the 2017 world workshop on the classification of periodontal and peri-implant diseases and conditions. Journal of Periodontology. 2018;89(Suppl. 1):S173-S182. DOI: 10.1002/JPER.17-0721\n'},{id:"B2",body:'Kassebaum NJ, Bernabe E, Dahiya M, Bhandari B, Murray CJ, Marcenes W. Global burden of severe periodontitis in 1990-2010: A systematic review and meta-regression. Journal of Dental Research. 2014;93(11):1045-1053\n'},{id:"B3",body:'Ansari G, Golpayegani MV, Welbury R. Histology and embryology of the teeth and periodontium. In: Ansari G, Golpayegani MV, Welbury R, editors. Atlas of Pediatric Oral and Dental Developmental Anomalies. Chichester: Wiley; 2018. pp. 13-16. DOI: 10.1002/9781119380894.ch2\n'},{id:"B4",body:'Nanci A, Bosshardt DD. Structure of periodontal tissues in health and disease. Periodontology 2000. 2006;40(1):11-28. DOI: 10.1111/j.1600-0757.2005.00141.x\n'},{id:"B5",body:'Ramos MJ. Biomecánica de los tejidos periodontales. Kiru. 2013;10(1):75-82\n'},{id:"B6",body:'Lindhe J, Lang NP. Periodontología clínica e implantología odontológica. 6a Edición ed. Madrid, España: Editorial Medica Panamericana S. A de C.V; 2017. pp. 3-37\n'},{id:"B7",body:'Hassell TM. Tissues and cells of the periodontium. Periodontology 2000. 1993;3(1):9-38. DOI: 10.1111/j.1600-0757.1993.tb00230.x\n'},{id:"B8",body:'Cho MI, Garant PR. Development and general structure of the periodontium. Periodontology 2000. 2000;24(1):9-27. DOI: 10.1034/j.1600-0757.2000.2240102.x\n'},{id:"B9",body:'Pisoschi C, Stanciulescu C, Banita M. Growth factors and connective tissue homeostasis in periodontal disease. In: Buduneli N, editor. Pathogenesis and Treatment of Periodontitis. London: InTechOpen; 2012. pp. 55-80. DOI: 10.5772/33669\n'},{id:"B10",body:'Tonetti MS, Greenwell H, Kornman KS. Staging and grading of periodontitis: Framework and proposal of a new classification and case definition. 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The host response to the microbial challenge in periodontitis: Assembling the players. Periodontology 2000. 1997;14(1):33-53. DOI: 10.1111/j.1600-0757.1997.tb00191.x\n'},{id:"B16",body:'Mira A, Simon-Soro A, Curtis MA. Role of microbial communities in the pathogenesis of periodontal diseases and caries. Journal of Clinical Periodontology. 2017;44(Suppl. 18):S23-S38. DOI: 10.1111/jcpe.12671\n'},{id:"B17",body:'Silva N, Abusleme L, Bravo D, Dutzan N, Garcia-Sesnich J, Vernal R, et al. Host response mechanisms in periodontal diseases. Journal of Applied Oral Science. 2015;23(3):329-355. DOI: 10.1590/1678-775720140259\n'},{id:"B18",body:'Cavalla F, Biguetti CC, Garlet TP, Trombone APF, Garlet GP. Inflammatory pathways of bone resorption in periodontitis. In: Bostanci N, Belibasakis G, editors. Pathogenesis of Periodontal Diseases. Cham: Springer; 2018. pp. 59-85. DOI: 10.1007/978-3-319-53737-5_6\n'},{id:"B19",body:'Muñoz Carrillo JL, Castro García FP, Gutiérrez Coronado O, Moreno García MA, Contreras Cordero JF. Physiology and pathology of innate immune response against pathogens. In: Rezaei N, editor. Physiology and Pathology of Immunology. London: InTechOpen; 2017. pp. 99-134. DOI: 10.5772/intechopen.70556\n'},{id:"B20",body:'Muñoz-Carrillo JL, Ortega-Martín Del Campo J, Gutiérrez-Coronado O, Villalobos-Gutiérrez PT, Contreras-Cordero JF, Ventura-Juárez J. Adipose tissue and inflammation. In: Szablewski L, editor. Adipose Tissue. London: InTechOpen; 2018. pp. 93-121. DOI: 10.5772/intechopen.74227\n'},{id:"B21",body:'Muñoz-Carrillo JL, Contreras-Cordero JF, Gutiérrez-Coronado O, Villalobos-Gutiérrez PT, Ramos-Gracia LG, Hernández-Reyes VE. Cytokine Profiling Plays a Crucial Role in Activating Immune System to Clear Infectious Pathogens. London: IntechOpen. 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Revisiting the Page & Schroeder model: The good, the bad and the unknowns in the periodontal host response 40 years later. Periodontology 2000. 2017;75(1):116-151. DOI: 10.1111/prd.12181\n'},{id:"B30",body:'Cekici A, Kantarci A, Hasturk H, Van Dyke TE. Inflammatory and immune pathways in the pathogenesis of periodontal disease. Periodontology 2000. 2014;64(1):57-80. DOI: 10.1111/prd.12002\n'},{id:"B31",body:'Gupta M, Chaturvedi R, Jain A. Role of monocyte chemoattractant protein-1 (MCP-1) as an immune-diagnostic biomarker in the pathogenesis of chronic periodontal disease. Cytokine. 2013;61(3):892-897. DOI: 10.1016/j.cyto.2012.12.012\n'},{id:"B32",body:'Bodet C, Chandad F, Grenier D. Inflammatory responses of a macrophage/epithelial cell co-culture model to mono and mixed infections with Porphyromonas gingivalis, Treponema denticola, and Tannerella forsythia. Microbes and Infection. 2006;8(1):27-35. DOI: 10.1016/j.micinf.2005.05.015\n'},{id:"B33",body:'Kiili M, Cox SW, Chen HY, Wahlgren J, Maisi P, Eley BM, et al. Collagenase-2 (MMP-8) and collagenase-3 (MMP-13) in adult periodontitis: Molecular forms and levels in gingival crevicular fluid and immunolocalisation in gingival tissue. Journal of Clinical Periodontology. 2002;29(3):224-232. DOI: 10.1034/j.1600-051x.2002.290308.x\n'},{id:"B34",body:'Braga TT, Agudelo JS, Camara NO. Macrophages during the fibrotic process: M2 as friend and foe. Frontiers in Immunology. 2015;6:602. DOI: 10.3389/fimmu.2015.00602\n'},{id:"B35",body:'Gonzalez OA, Novak MJ, Kirakodu S, Stromberg A, Nagarajan R, Huang CB, et al. Differential gene expression profiles reflecting macrophage polarization in aging and periodontitis gingival tissues. Immunological Investigations. 2015;44(7):643-664. DOI: 10.3109/08820139.2015.107026\n'},{id:"B36",body:'Yu T, Zhao L, Huang X, Ma C, Wang Y, Zhang J, et al. Enhanced activity of the macrophage M1/M2 phenotypes and phenotypic switch to M1 in periodontal infection. Journal of Periodontology. 2016;87(9):1092-1102. DOI: 10.1902/jop.2016.160081\n'},{id:"B37",body:'Lam RS, O�Brien-Simpson NM, Lenzo JC, Holden JA, Brammar GC, Walsh KA, et al. Macrophage depletion abates Porphyromonas gingivalis-induced alveolar bone resorption in mice. Journal of Immunology. 2014;193(5):2349-2362. DOI: 10.4049/jimmunol.1400853\n'},{id:"B38",body:'Gemmell E, Seymour GJ. Cytokine profiles of cells extracted from humans with periodontal diseases. Journal of Dental Research. 1998;77(1):16-26. DOI: 10.1177/00220345980770010101\n'},{id:"B39",body:'Dutzan N, Gamonal J, Silva A, Sanz M, Vernal R. Over-expression of forkhead box P3 and its association with receptor activator of nuclear factor-kappa B ligand, interleukin (IL)-17, IL-10 and transforming growth factor-beta during the progression of chronic periodontitis. 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DOI: 10.1111/j.1399-302X.2008.00463.x\n'}],footnotes:[],contributors:[{corresp:"yes",contributorFullName:"José Luis Muñoz-Carrillo",address:"mcbjlmc@gmail.com",affiliation:'
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Faculty of Odontology, School of Biomedical Sciences, Cuauhtémoc University Aguascalientes, Mexico
Department of Stomatology, Autonomous University of Aguascalientes, Mexico
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We included 111 consecutive patients treated at the cathlab of the University Hospital Ostrava, Czech Republic, with ISR within 12 months after implantation of a bare-metal stent. The control group consisted of 111 matched patients with identical main demographic and clinical risk factors. To set the reference intervals for MMPs, we measured the blood concentrations of these analytes in a group of healthy volunteers (N = 180) with an average age of 40–50 years. The enzyme concentrations were measured by immunosorbent assay. Statistical analysis was performed using IBM SPSS Statistics version 22 and MedCalc Version 14.12. We found that increased levels of MMP-3 and 9 were associated with a significant increase in ISR risk. The MMP-9 cut-off value for ISR risk prediction was determined to be ≥64.8 ng/mL. 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