Take Home Message The two types of reproductive cycles are the estrous cycle and the menstrual cycle. Re- productive cyclicity provides females with repeated chances for pregnancy. An estrous cycle consists of the physiologic events that occur between successive peri= ods of sexual receptivity (estrus or heat) and/or ovulations. The length of cycle varies from about four days in rodents to as long as 14-16 weeks in elephants. Each cycle consists of a follicular phase and a luteal phase. The follicular phase is dominated by estradiol secreted ‘by ovarian follicles, Estradiol causes marked changes in the female tract and initiates sexual receptivity. The luteal phase is dominated by progesterone from the corpus luteum that Prepares the reproductive tract for pregnancy. Periods of time when estrous cycles cease are called anestrus. Anestrus is caused by pregnancy, season of the year, lactation, certain forms of stress and pathology. ‘A menstrual cycle consists of the physiological events that occur between successive menstrual periods (about 28 days). At the conclusion of the luteal phase in the menstrual cycle, the endometrium is sloughed to the exterior (menstruation). No endometrial slough- ing occurs in animals with estrous cycles. Each menstrual cycle consists of 3 distinct phases that reflect the condition of the uterine endometrium. The cycle starts with menses (about a 4-6 day period) where the endometrium is sloughed to the exterior. The second phase (about 9 days) is the proliferative phase in which follicles develop and secrete estradiol. The endometrium begins to grow and increase in thickness. The final phase, the secretory phase (14 days), is dominated by the corpus luteum that secretes progesterone and estradiol. The endometrium grows and continues to increase in thickness as a function of progesterone. Atthe end of this 28 day period the endometrium begins to slough again if the woman is not pregnant and a new cycle begins. Amenorrhea refers to the lack of menstrual periods and is caused by many of the same factors that cause anestrus. This chapter will provide you with fundamen- {al knowledge about female reproductive cyclicity. Among mammals, reproductive eyclicity consists of the estrous cycle and the menstrual cycle. Both types of cycles provide the female with repeated opportuni- ties to become pregnant. The fundamental differences between these types of reproductive cycles will be pre- sented in the two sections that follow entitled, Estrous Cycles and the Menstrual Cycle. There are species exceptions to some of the principles described. Most of these exceptions will be described in later chapters especially Chapters 8 and 9 that deal specifically with the follicular and luteal phases. THE ESTROUS CYCLE After puberty, the female enters a period of reproductive cyclicity that continues throughout most of her life. Estrous cycles consist ofa series of Predictable reproductive events beginning at estrus (heat) and ending at the subsequent estrus. They continue throughout the adult female’s life and are interrupted by pregnancy, nursing and by season of the year in some species. Cyclicity may also cease if nutrition is inadequate or environmental conditions are unusually stressful. Pathologic conditions of the reproductive tract, such as uterine infection, persistent corpora lutea or a mummified fetus may also cause anestrus (a period when cyclicity stops). Estrous cycles provide females with repeated opportunities to copulate and become pregnant. Sexual receptiv- ity and copulation are the primary behavioral events that occur during estrus. Copulation generally occurs prior to ovulation. If conception (pregnancy) does not occur, another estrous cycle begins, providing the female with another opportunity to mate and conceive. When pregnancy occurs, the female enters a period of anestrus that ends after parturition (giving birth), uterine involution (acquisition of normal uterine size and function) and Inctation. Author's Note: Afier years of teaching about reproductive cyclicity and listening to repeated student feedback, I have concluded that developing a thorough understanding of the estrous cycle in animals makes tmderstanding the menstrual cycle easy. The reverse is not necessarily true. Escaneado con CamScannerLlevar A Casa Mensaje Los dos tipos de ciclos reproductivos son el ciclo estro ye! ciclo menstrual. La ciclided reproductiva proporciona a las hembras repetidas oportunidades de embarazo. Un ciclo estro consiste en los acontecimientos fisiolégicos que se producen entre peri-odos sucesivos de receptividad sexual (estrus 0 calor) y/o ovaciones. La duracién del ciclo varia de unos cuatro dias en roedores hasta entre 14 y 16 semanas en elefantes. Cada ciclo consiste en una fase folicular y una fase litea. La fase folicular esta dominada por el estradiol secretado por 198 foliculos ovéricos, El estradiol provoca cambios marcados en el tracto femenino e inicia la receptividad sexual. La fase hitea esté dominada por la progesterona del cuerpo hiteo que prepara el tracto reproductivo para el embarazo. Los periodos de tiempo en que cesan los ciclos estros se denominan anestrus. El anestrus es causado par el embarazo, la estaciGn del afo, la lactancia, ciertas formas de ‘estrés y patologia. Un ciclo menstrual consiste en los acontecimientos fisioldgicos que se producen entre petiodos menstruales sucesivas (unos 28 dias). Al final dela fasehitea det ciclo menstrual, el endometrio se desprende hacia el exterior (menstruacién). No se produce desprendimiento endometrial en animales con ciclos estos. Cada ciclo menstrual consta de 3 fases distintas que reflejan la condicién del endometrio uterino. El ciclo comienza con menses (alededor de 446 dias) donde el endometrio se desprende hacia el exterior. La segunda fase (unos 9 dias) es la fase proliferativa en la que los foliculos se desarrollan y secretan estradiol. El endometrio comienza a crecer y aumentar de grosor La fase final, la fase secretora (14 4ias), estd dominada por el corpus luteum que secreta progesterona y estradiol. El endomettio crece ¥ continua aumentando de groser en funcién de la progesterona. Ai final de este periodo de 28 dias, el endometrio comienza a desprenderse de nuevo sila mujer no esta embarazada y comienza un nuevo ciclo, La amenorrea se refiere a la falta de periodos menstruales y es causada por muchos de los mismos factores que causan el anestrus. Este capitulo le proporcionard un conocimiento (calor)y terminando en el stro posterior Contindan ‘fundamental sobre la ciclidad reproductva femenina, Entre los mamiferos, a ciclidad reproductiva consiste «encl ciclo estoy el ciclo menstrual. Ambos tpos de «acs proporcionan ala hembra tazos de oportunidad repetidos| para quedar embarazada Las diferencias lundamentales entre estos tipos de ciclos reproductivos se enviarin ‘eevee ent ds scores aguetestuadas Ces Estresy Cio Menstrual Existen excepciones de especies a algunos delos pincpios descritos. La mayoria de ‘estas excepciones se deserbirdn en captuos posterores, especialmente enlos captulosB y9, quetratanespecifcaente las fases folicular y lutea. EL CICLO ESTROS Despues dela pubertad, a hembra entra enun periodo de ciclidad reproductiva que continua durante la ‘mayor parte de su vida, Los ccls estros consisten en una serie de-eventos reproductivos predecibles que comienzan en celo - + Nota del autor Despuds de afos de ensenar sobre los estudiantes, he llegado a la conclu ién de que decarrollat u ofotorgo de la vida dela herb adullay 109 nerrump das por el embarazo, la lactancia y por estacién del afio en algunas especies. La ciclidad también puede cesar sila rnutricién e¢ inadecuada 0 si las condiciones ambientales son inusualmente estresantes, Las afecciones patoldgicas deltractorepraductiva, coma infeccién uterina, corpora lutea persistente o un feta momificade también pueden causar anestius (un petiado en el que se detieneta cia) Los oclos estros brindan alas hembras repetidas ‘oportuidades de copuiaty queda embarazadas. Lareceptivdad serualy la copu'a son los principales eventos conductuales que ocuren durante el estius. La copulacion se produce ener almerte antes de la ovulacion. Sila euncepeidn (erivarazo) no veut, comenza vie ccl esta, proporcionande ‘la hembra ote oportunidad de apavearse y convebit. Cuando se produce el embarazo, la hembra entra en un perioda be anestrus que termina despuds dela patwicidn (da al), tainvercion vievina(adauiticidn dol tomato y uncién ‘ering normates) ya lsctancia Ciclidad reproductiva y escuchar repetidos comentanos de ‘comprensién exhaustiva del ciclo estr en animales, + faciita la comprensién de|ciclo menstrual, Locontrano no es necesariamente cierto. Escaneado con CamScanner442 Reproductive Cyclicity Terminology Describing Reproductive Cyclicity can be Confusing ‘The words used to describe the estrous cycle are spelled similarly, but have sublly different mean- ings. The proper use of the words estrus and estrous must be understood to prevent confusion. The word ‘estrus isa noun, while estrous is an adjective. Oestrus and oestrous are the preferred spellings in British and European literature, Estrual is also an adjective and is used to identify a condition related to estrus. For example, an estrual female is a female in estrus. An estrous eycle is the period between one estrus and the next. Estrus is the period of sexual receptivity. Estrus is commonly referred to as heat. The term estrus (oestrus) originated from a Greek word mean- ing “gadfy, sting or frenzy”. This word (oestrus) was used to describe a family of parasitic biting insects, (Oestridae). These insects caused cattle to stampede with their tails ailing in the air as the insect buzzed around them. The behavior occurring in females in estrus was deemed similar to that observed during these insect attacks. Thus, the term oestrus or estrus was applied to the period of sexual receptivity in mammalian females. Another common term used to describe a reproductive pattem is season. This refers to several estrous cycles that may occur during a certain season of the year. For example, a mare “coming into season” begins to show cyclicity and visible signs of estrus. She will cycle several times during her “season” (See Figure 7-1). ESTRUS isa noun. “The cow is displaying estrus,” ESTROUS is an adjective. “The length of the estrous cycle in ‘the pig is 21 days.” Figure 7-1. Types of Estrous Cycles as Described by Annual Estradiol (E,) Profiles POLYESTRUS (Mare) (Ewe, doe, el, nanny) Relative blood concentrations of Ez MONOESTRUS (Dog ©, wolf, fox, bear) (Cow, queen, pig, rodents) SEASONAL POLYESTRUS (Long Day) Spring breeding season SEASONAL POLYESTRUS (Short Day) Aucumn breeding season Escaneado con CamScanner142 Ciclcidad Reproductiva ‘Terminology Describing Reproductive Cyclicty puede ser confusa Las palabras tlizadas para desonirel clo estre sedelevean de manera sar per tenensigniicados sutimente diferentes. El uso adecuado de las palabras estro y estros debe entenderse para evtarconfusiones. La palabra esto es un sustantivo, mientras que el estros es un ajetvo. Elestroy el estro sons grafias preferidas ena ineratura btanica y europea. Elestrual es también un adjetivay se tliza para identficar na condicion relaconada con el celo. Por ejemplo, una hembra estruai es una hembra en estro Un ciclo estros es e! periodo entre un estro y el siguiente Elestrus es el periodo dela receptividad sexual. Elestrus ‘se conoce cominmente como calor. El término estro (oestrus) se oriqind de una palabra griega que siqnifica “gadfly, aguién o tenes? Esta palabra (oestrus) se utiizo para descrbr una familia de insectos patésits picadores (Oestidae)Estosinsecos hicieron que el ganado se estampiea con us colas agtando en el aire mientras el insecto zumbaba atrededot de ellos. El comportamiento ccurrido enhembras en celo se considerd similar al observado durante estos ataqus de insets Asi émino esto o esto se aplicé al periodo de receptividad sexual en hembras deraneres Oto temeacomun tad pore 1 escribir un patron reproductiva es temporada. Esto se i refiere a vans ciclos estos que pueden ocurir durante na delerminadaestacién del afo, Por ejemplo, unayequa | que ‘entra en temporada’ comlenza a mostrar ciclismo y signos visibles de celo. Ella pedaleard varias veces durante su temporada’ (Ver Figura 7-1), ESTRUS es un sustantivo ‘La vaca esta mostando est’ ESTROUS es un acjetivo “La duracion del ciclo estrus en el cerdo es de 21 dias’. Figura 7-1. Tipos de ciclos estros descritos por los perfiles ~anuales de Estradiol (£2) POLYESTRUS (vaca, ina, cerdo, roedores) | (Mare) BZ |! rmenenSRAAL POLYESTUS (0a bygo) Twoan arg MIND = rare sans pron MONOESTRUS (Perr0, kt, za 0:0), RaFF ES © verge Se S SPFSEB Escaneado con CamScannerReproductive Cycllcity 443 Examples of other words that can lead to confusion in spelling and usage are: anestrous vs. anestrus and polyestrous vs. polyestrus. Ifthe word isused as an adjective, itis spelled aus. Forexample, “polyestrous females have repeated estrous cycles.” If the word is used as a noun, it is spelled -us. For ‘example, “the female is experiencing anestrus.” The three types of estrous cyclicity are: + polyestrus + seasonally polyestrus + monoestrus Estrous cycles are categorized according to the frequency of occurrence throughout the year. ‘These classifications are polyestrus, seasonally polyestrus and monoestrus (See Figure 7-1). Poly- estrous females, such as caitle, swine and rodents, are characterized as having a uniform distribution of estrous cycles throughout the entire year. Polyestrous females can become pregnant throughout the year without regard to season. Seasonally polyestrous females (sheep, goats, mares, deer and elk) display “clusters” of estrous cycles that occur only during a certain season of the year. For example, sheep and goats are short-day breeders because they begin to eycleas day length decreases in autumn, In contrast, the mare isa long-day breeder because she initiates eyclicity as day length increases in the spring. ‘Monoestrous females are defined as having only one cycle per year, Dogs, wolves, foxes and bears are animals that are characterized as having a single estrous cycle per year, Domestic canids typi- cally have three estrous cycles every two years but they are generally classified as monoestrus. In general, monoestrous females have periods of estrus that last for several days. Such a prolonged period of estrus inereases the probability that mating and pregnancy can occur. Each type of eycle pattern is represented in Figure 7-1. The Estrous Cycle Consists of ‘Two Major Phases The estrous cycle can be divided into two distinct phases that are named after the dominant structure present of the ovary during each phase of the cycle. These divisions of the estrous cycle are the follicular phase and the luteal phase. The follicular phase is the period from the regression of corpora lutea to ovulation, In general, the follicular phase is relatively short, encompassing about 20% of the estrous cycle (See Figure 7-2). During the follicular phase, the primary ovarian structures are large grow- ing follicles that secrete the primary reproductive hormone, estradiol. During the follicular phase: * large antral follicles = the primary ovarian structure + estradiol (secreted by follicles) = the primary hormone The luteal phase is the period from ovula- tion until corpora lutea regression. The luteal phase is much longer than the follicular phase and, in most mammals, occupies about 80% of the estrous cycle (See Figure 7-2). During this phase, the dominant ovarian structures are the corpora lutea (CL) and the primary reproductive hormone is progesterone. Even though the luteal phase is dominated by progesterone from the CL, follicles continue to grow and regress during this phase but they do not produce high con- centrations of estradiol, Details of follicular growth are presented in Chapter 8. During the luteal phase: * corpora lutea = the primary ovarian structures + progesterone (secreted by corpora lutea) = the primary hormone The Estrous Cycle can Also be Divided into Four Stages The four stages of an estrous cycle are proestrus, estrus, metestrus and diestrus, Each of these stages is a subdivision of the follicular and luteal phases of the eycle. For example, the follicular phase includes proestrus and estrus. The luteal phase includes metestrus and diestrus. Follicular phase = Proestrus + Estrus Luteal phase = Metestrus + Diestrus Escaneado con CamScannerEjemplos deotras palabras que puedenlievar = ® confusion en la ortografiay el uso son: anestros vs anestrus y polestos vs. polestus. ils palabra se | usa como adjetivo, se escribe ous, Por eempla, ‘las hembras Polesttes tienen ciclos estros repetidos”.Silapalsbra Se utiliza como sustantivo, se deletrea -nosotros. Por /emplo, la hembra esta experimentando anestrus" Los tres tipos de ciclidad estral son: * Poliestro * monoestrus * estacional Los ciclos estros se clasifican de acuerdo con la frecuencia de ocurencia alo largo del afo. Estas Clasificaciones son poliestro, poliestro estacional y monoestro (wéase la fiqura 7-1) Las hembras poliestras, como el ganado vacuno, los cerdosy ls roedores, se caracterizan por tener una distnbucién uniforme de los ccios festros durante todo el ao. Lashembras polestres pueden 4 quedar embarazadas durante todo el aio sin tener en Cuentala estacin Las hembras polestraisestacionales 1 (ovas, cabs yequas, eros y aces) presentan ' “qupos' de ciclos estos que sola se producen durante una determinada temporada del ao Por ejemplo, la ovejas ylas cabras son ctiadoras de dia corto porque comienzan a Citeuar a medida que dsminuye la durecindel dia enotofo Encontraste, a yegua es una criadora de larga duraciin Porque inicislacicidad a medida que aumertala =. duracién del ia ena primavera Las hembas monoestrosas se definen como tener solo un ciclo al afio. Los pros lobos,zaxosy 0808 son animales que Caracterizan por tener un inco ciclo esti al ao. Los canidos cdomiésticos tipo cally tienen tres ciclos estros cada dos af0s, pero generalmente se clasfican como manoestrus. En general las hembras monoestres tienen perodos de estrus que | duran varios dias. Un periodo tan prolongade de celo umentaa probabiidad de que se produzcanapareamventoy embarazo Cada tipo de pation de ciclo est representado en laFigua7.1 El ciclo estrastico consiste en os fases principales Eloico estos se puede divs en dos fases distintas que llevan el nombre deta estructura dominante presente en el ovario durante cada fase del ciclo. Estas divisiones del ciclo estro sonla fase foiculary la faseliitea. La fase folicular es el periodo desde la regresion de la corpora Ciclicidad reproductiva 143 lutea ala ovulacion. En general, la fase folicular es, ‘elatvamente covta y abarca aproximadamente 20% det Ciclo esto (vase ta figura 7-2) Durante la fase folie ula, 'as estructuras ovaricas primarias son grandes foliculos ‘en crecimiento que secretan Ia hormona reproductiva Prmana ot exraaot Durante la fase folicular: Foliculos antorales grandes = la ‘estructura ovérlca primaria esti adio! (secretada por toiculos) = la hormona primaria Lata litea es el periodo desde ls ovulacin hasta a regresion fa corpora tea, Lafase tea 68 ‘mucho mds larga qu lafasefoicuay ena ayia delos mamilerns, ccupa alededr del RP del clo estro (vease la figuta 7-2). Durante esta tase, las scstuas eves dinates sn ta corporate (CL) y ls hormonareproducta primana es la progesterona ‘Aunque la fae ttea esté dominadsa pola proge sterona el CL, tos foliculos continuan erecienda y regresanda ‘ucante esta tase pero no producen atas concentracones de estradiol, Los detalles del crecimiento folicuar se presentan en el Capitulo 8 [a Durante la fase litea:b corpora lutea = las estructuras. radroowincas primarias: Laprogesterona seeretada por ‘bal corpora lutea) * la hormona even El ciclo estrus también se puede dividir en custo etapas Las cuatro etapas deun Ciclo esto, estrus, metro y diestrus. Cada ‘una de estas etapas es una subdivision de las fases folicular y lites del ciclo, Por ejemplo, la fase folicular incluye proestro y estro. La fase lea nye metetoy destus Fase folicular-Proestrus+ Estrus Luteal phase = Metestrus + Diestrus Escaneado con CamScanner144 Reproductive Cyclicity Figure 7-2. Phases of the Estrous Cycle > 28 g8 a5 ee 8 |e Luteal Phase SSATTACHITIASETOINUABMI WITH %MO1II4 SS Day of Cycle is designated as day 0. ‘The follicular phase begins after luteolysis that causes the decline in progesterone. Gonadotropins (FSH and LH) are therefore secreted that cause follicles to secrete estradiol E,). The folicular phase Is dominated by estradiol secreted by ovarian follicles. The follicular phase ends at ovulation. Estrus ‘The luteal phase begins after ovulation and includes the development of corpora lutea that secrele progesterone (P.). The luteal phase also includes luteolysis that is accompanied by arapid drop in progesterone. Luteolysis is brought about by prostaglandin Fae. Proestrus is the Period Immediately Preceding Estrus Proestrus begins when progesterone declines as a result of luteolysis (destruction of the corpus lu- teum) and terminatesat the onset of estrus. Proestrus lasts from 2 to 5 days depending on species and is characterized by a major endocrine transition, from 1 period of progesterone dominance to a period of estradiol dominance (See Figure 7-3). The pituitary gonadotropins, FSH and LH, are the primary hor- mones responsible for this transition. It is during proestrus that antral follicles mature for ovulation and the female reproductive system prepares for the onset of estrus and mating. Estrus is the Period During Which the Female Allows Copulation Estrus is the most recognizable stage of the estrous cycle because it is characterized by visible behavioral symptoms such as sexual receptivity and mating, Estradiol is the dominant hormone during this stage of the estrous cycle, Estradiol not only induces profound behavioral alterations, but causes major physiologic changes in the reproductive tract. When a female enters estrus, she does so gradually and is not sexually receptive at first, She may display behavioral characteristics that are indicative of her appronching sexual receptivity. Proestrus = Formation of ovulatory follicles + Ey secretion Estrus = Sexual receptivity + peak E; secretion Metestrus = CL formation + beginning of P, secretion Diestrus = Sustained luteal secretion of P, Escaneado con CamScanner444 CiidadReproduetva Figura 7-2. Fases del ciclo estrastico SSADIATI DIAS ETAIMUNBHEBTBHMO1II4ASE Dia del Ciclo fase folcularcomiena despuds dea dlisis que provoca la disminucién de la progesterona Por lo tanto, se secretan gonadotropinas (F5H1y LH) que hacen que los ftcuos seteten estradiol (€2), La fase oluaresté dominada por el estradiol secretado por los foliculos ovaricos. La fase folicular termina en Ia ovulacion. Estrus ‘se designa como el dia 0. 1s@ lutea comienza despues de la ovulacion luye el desarrolo de corpora lutea que eecreta, progesterone (P4) La fase hitea tambien incuyela tls que se acompana de una pda disminucion de la progesterona. La luidlisis es provocada por la prostaylandina F2- 3. SSS Proestrus es el periodo inmediatamente Elestrus es el periodo durante el cual anterior al Estrus lahembra permite la copulacién Proestrus comienza cuando la progesterona di smunuye. Estrus es la etapa mas reconocible del ciclo como resultado dela ltSisis(destruccén del cuerpo li estrus porque se caracteriza por sintomas conductuales eum) y termina al inicio del estrus El proestrus dura de 2 visbes como la receptividad sexual y el apareamiento Et a5 dias dependiendo de la especie y se caractenza por estradiol es la hormona dominante durante esta fase del tuna transicién endocrina importante, de un periodo de Ciclo estre.E estradiol no solo induce profundas elteraciones ddominacion de ta progesterona a un periodo de domninacién el compurtamento, sino que wrovoca cambios fisickigioos del etal véesla gs 73) Las goradtopeas importantes en el tracto reproductive. Cuando hipofisarias, FSH y LH, son las principales hor mones "una hembra entra en e celo, lo hace responsables de esta ransicién, Es durante el proestrus graduaimeite yno es sexualmente receptival prneipio Puede Aue os fotos atrales maduan para a ovlacén ye sistema ‘mostrar caracteristicas de comportamiento que son ‘eproductor femienino se prepara para la aparicién det indcativas de su aproximacién dela receptvidad sexual. celalo yl apreamenta. Proestrus Formacion de foliculos ovulatorios + €, secrecién tonila Estrus Receptividad sexual + pico E, secrecién Metestrus CL formacién + inicio de P, secrecién Diestrus Secrecion luteal sostenida de P , Escaneado con CamScannerReproductive Cycitcity 145 Figure 7-3. Stages of the Estrous Cycle Relative Hormone Concentrations (blood) SSMTDIOTTIAS ETOP MUAHKEETMMMOT TITS ES Day of Cycle Proestrus is cheracter- lzed by a significant rise in estradiol (E;) secreted by maturing follicles. ‘When estradiol reach- es a certain level, the female shows behav- ioral estrus and then ovulates. Following ovulation, cells of the follicle are transformed into luteal calls that form the cor- us luteum (CL) during metestrus. Diestrus is characterized by a fully functional CL and high progesterone (Pa). ‘These include increased locomotion, phonation (vo- cal expression), nervousness and attempts to mount other animals, However, during this early period she Will not accept the male for mating. As the period of estrus progresses, so docs the female's willingness to accept the male for mating. This willingness is referred to as standing estrus. It is during the time of estrus that the female displays a characteristic ‘mating posture known as lordosis, so named because of a characteristic arching of the back in prepara for mating. Standing behavior (lordosis) is easily observed and is used as a diagnostic tool to identi the appropriate time to inseminate the female arti- ficially or to expose her to the breeding male. The average duration of estrus is characteristic for each species. However, the range in the duration of estrus can be quite large even within species (See Table 7-1), Understanding and appreciating the magnitude of these ranges is important because it allows one to Predict cyclic events with a degree of uecuracy. Metestrus is the Transition from Estradiol Dominance to Progesterone Dominance Metestrus is the period between ovulation and the formation of functional corpora lutea. During arly metestrus both estradiol and progesterone are relatively low (See Figure 7-3). The newly ovulated follicle undergoes cellular and structural remodeling resulting in the formation of an intraovarian endocrine gland called the corpus luteum. This cellular trans- formation is called luteinization (See Chapter 9). Progesterone secretion begins in metestrus and is detectable soon after ovulation. However, two to five days are usually required after ovulation before the newly formed corpora lutea produce significant quanti- ties of progesterone (See Figure 7-3). Diestrus is the Period of Maximum Luteal Function Diestrus is the longest stage of the estrous cycle and is the period of time when the corpus luteum is fully functional and progesterone secretion is high. Escaneado con CamScannerEspanol Ciclicidad reproductiva 145 Figura 7-3. Etapas del ciclo estristico Diestrus Ta TTIOTIIeF Ae TwUMHNEM TR Después de (Ouando Wear seancaunceto the lahembra muestra CL eomple ato nivel de progesterona Taowiacin as cbllasdel oliculo se transforman en céllas Tpoeanseecaretes For un aumento signet de rte finecsoral yun ‘tail (€2) secetado polos folcus enmaduacién coropus ten (CL) diveereste (Pa. Estos incluyen aumento de lalocomoci,fonacin (expres vecal nevis: einenog de ments eres ‘animes Sm erbargo, drat este enodo termpanono ecetara almacho para el apareamiento Amedida que avanae el petiodo de cela, también lohace la voluntad dela hembra «de aceptar al macho paraelapareamienta Feta dispasicén se conoce como esti dep. Es durante la Bpuca del colo que la hombra muesta una postta de apareamiento caractersticaconncda como lordoas famada asi debido ‘aunatco caacteristica dela espaldaenpreparacin para laparearvento Elcomportamientodepie(erdoss} se hse fdcimerty se ti como heraianta de dagndston para identficar el momento apropiado para seminar ala hembra artficialmente o para exponerla al macho teproducto La duracon mevda de actuz es calactersicn Ja especie. Sin embargo, elrango en a duracién del cence puede cer bastante grande incluso dento de las species (vate taba 7-1) Comp uf de estos rangos es importante porque pemite predeci eventos cilicas concierto grado de precisén rider y aes Metestrus esta transcion de Estradiol Dominance ‘a Progesterona Dominance Metestus es 1 penodo entre fa ovulacion y la formacion ‘e corpora lutea funevinal urate el metro tempeang, tomo elestradiol come ia progesterona sonrelatwamerte bajos (weasel figura 7.3) Elfoteulaecién avulada 3e comete ‘una remodelacion cer y estructural que resuta ents foracidn deus pn rin ineaoai ima cmpo Luteo, Esta formacidn tans celular se Laenizacén (vease el Capitulo 9) La secrecién de ragesterona comienza en el melo y es detectable poco despues dela a eerbargo, ge cinco dias despues dela ovulacion antes de que la corpora itearecién fnmada produrca avs cuantatvs signfativos do progesterona (wae ta fgua 7.3) vente se requere de os a Diestruses el period a6 ‘uci teal maxima Diestus ests etapa mis 79a cece esto yes el periaia de tiempo en que el cuerpo kiteo es carmgetamant funcanal yl seein 6 progesona eta Escaneado con CamScanneril 146 Reproductive Cyclicity Itends when the corpus luteum is destroyed (luteoly- sis). High progesterone prompts the uterus to prepare suitable environment for early embryo development and eventual attachment of the conceptus to the endo- metrium. Diestrus usually lasts about 10 to 14 days in most large mammals, The duration of diestrus is directly related to the length of time that the corpus luteum remains functional (ie. secretes progesterone). Females in diestrus do not display estrous behavior. The Estrous Cycle of the Bitch and Queen Varies from Patterns Previously Described The estrous cycle of the domestic bitch has a different stage sequence than other mammals. The cycle consists of anestrus, proestrus, estrus and di- estrus. Anestrus usually lasts for about 20 weeks in the nonpregnant bitch. The long anestrus (5 months) causes the the bitch to display two estrous periods in three years. However, wild canids (wolf, coyote, Australian dingo) display only one estrous period per year and these periods are usually seasonal. Figure 7-4 illustrates the stages, sequence, relative timeline and the endocrine profiles of the cycle in the bitch. The onset of proestrus is usually considered to be the beginning ofthe estrous cycle. The drop in blood FSH that occurs, during proestrus is presumably due to negative feedback on FSH by inhibin secreted from developing follicles. ‘The bitch becomes receptive to the male during decreas- ing estradiol and rising progesterone concentrations. Ovulation occurs 2-3 days after the LH surge. Fertiliza- tion generally lakes place 48-72 hours after ovulation. This delay between ovulation and fertilization allows for superfecundation to occur frequently in canids. Superfecundation occurs when multiple ovulations produce multiple oocytes during a single estrus period that are fertilized by spermatozoa from different males. Therefore, bitches that are allowed to “roam free” dur- ing estrus have a high probability of delivering litters with multiple breeds of puppies. Figure 7-4. The Annual Reproductive Cycle of the Bitch (Modified from Johnston, Root Kustritz and Olson, 2001. Canine and Feline Theriogenology) ‘ANESTRUS 5Mo Relative Hormone Concentrations (blood) PROESTRUS oletemanees [ Luteal Phase | Latter Wein A 2Mo ---¢-Fertitation complete Troasera DHOMEEMGANOT Tes 4s SETS SAITT OL AIA ET ED Weeks Days Days Weeks estrus, Proestrus Estrus Diestrus A period of reproduc- | | Proestrus is considered the} | Shortly after peak estradiol, | | Both pregnant and open behavioral estrus begins. | | bitches are considered to beginning of the cycle and is characterized by the appear- ance ofa blood-tinged vaginal] discharge. It ends when the! bitch coputates with the male, Estraciol gradually Increases} ‘and peaks slightly before the| onset of estrus. tive quiescence. This long anestrus period is responsible for a cyclic profile of three cycles in two years. beindiestrus. Pregnancy status does not alter the length of diestrus. Pro- gesterone peaks at about 15 days then decreases gradually. Bitches that do not become pregnant are often considered to be pseudopregnant, Both LH and FSH peak in early estrus. Ovulation is completed at about the third day of estrus and ertlization is completed at about the sixth day. Pro- gesterone increases dur- ing the later part of estrus signifying luteinization, Escaneado con CamScanner146 Ciclidad Reproductiva Termina cuando se destruye el corpus hte (teoy sis) Un alto nivel de progesterona incita al utero a preparar ‘un entorno adecuado para el desarrollo embrionario temprano y a eventual unién del concepto al endometro. estrus suele durar de 10 214 dias en la mayoria de los mamiferos grandes. La duacién de diestrus esté <érectamenterelaconada con el tempo que el cuerpo luteo ‘permanece funcronal (es dece, secreta progesterona) Las hhembras en destrusno muestran comportamiento estos. El ciclo estros de la perra y la reina varia a partir de patrones descritos anteriormente El ciclo estral de la perra doméstica tiene luna secuencia de etapas diferente ala de otros mamiferos. EI ciclo consiste en anestrus, presto, estioy di-estro. E] anestrus suele durar unas 20 semanas en la perra no embarazada. El anestrus largo (5 meses) hace ue la perra muestre dos periodos estros en tres afios. | Sin embargo, los cénidos silvestres (lobo, coyote, 5 dingo austatano) solo muestra un période esto al aio y st periodos suelen ser estacionales La figura 7-4iustra las etapas la secvencia la linea de tempo relativa y los perfies endocrinos del ciclo ena pera. La aparicén de proestro se suele considerar el comienzo del ciclo estro. La caida enla sane FSH que se produce areal proestusse ce presumemente 41a retroalmentacion negativa sobre ia FSH por el inhibxdor seeretado de fos foicuos en desarrollo. La perra se vuelve ‘eceptiva al macho durante la disminucdén del estradily el ‘aumento de las concentraciones de progesterona, La owiacién se produce 2:3 dias despues de acleada de LM La feriizacén generalmente tiene gar 48-72 horas después de la ovulacién Este retraso entre le ovulaciény la fertizacié permite que a superfecundacién ocurra con frecuencia enos canidos, La superfecundacién se produce cuando mihiples ovlaciones producen multiples ovocitos durante un solo periodo de estrsode que on feteacot por espematenos cesfereres mactos, Por lo tanto, las perras alas que se les permite ‘vagar be’ deestro duo tenen una ata probabidad de dara luz camadas ‘con multiples azas de cachorros, Figura 7-4. El ciclo reproductivo anual de la perra Pacur Peas Ce) ah) Unperiods de regroduccstn | | Proestrus se considera el tive quiescencia, Este | | comenzo del octoy es period largo de anestus.es | } caracterizada por la aparienciar responsable den ccico | | ance de un vain edo oe sangre perfil de tres ciclos Descarga, Termina cuando el en dos afos, perra copula con el macho. estradiol aumenta gradualmente y picos ligeramente antes no delasis Pomdepusdetpenderstatss || se eonstera qu tant las comenzadestrusconductual | | sevosrneanrsi cmos Tanto LH como FSH aleanzan | | ableras estinen diestrus supunto ménimo enel estrus} | estdo de etaraza allen tengrano Laonisensecoreets | | la duacién de diestrus. La sproximadamemtealtercer || proqestona acanza supueto @aceecmutylateizaconse || masimoatos 15 dasy ompetaaproumatimeesiuets | | Iuego arsminuye dia La progesteona umens |} gradusiment. Las paras la dtona pane oe errviiogue | | wie avdanwnoarasadan 4 signticalaMenszacin »—) | rversernpemmrat, Escaneado con CamScannerReproductive Cyclicity 447 ‘As you can see from Figure 7-4, the bitch does not have a defined metestrus as in other species. The initial development of luteal tissue occurs during estrus shortly after ovulation as in other mammals. In the queen, stages of the estrous cycle include proestrus, estrus, postestrus, diestrus and anestrus. There is little evidence for seasonality in queens and they tend to be polyestrus. However, as photoperiod increases, the length of estrus Felids are induced ovulators and copulation | srequired for induction of the LH surge. Postestrus is a term used to describe an inter- estrus period that follows estrus in a queen that has not been induced to ovulate by copulation (See Figure 7-5). In queens that have not copulated, no ovulation occurs and no corpora lutea form, Therefore, neither metestrus (CL formation) nor diestrus occurs. As in most induced ovulators, it would be appropriate to consider that the female would remain in a constant follicular phase until copulation occurs. After copula tion the female ovulates and only then do corpora lutea form. In this context induced ovulators constitute a special form of estrous cycle that does not have a true luteal phase: Anestrus Means “Without Estrus (Heat)” ‘Anestrus is a condition when the female does not exhibit estrous cycles. During anestrus the ovaries are relatively inactive and neither ovulatory follicles nor functional corpora lutea are present. Anestrus is the result of insufficient GnRH release from the hy- pothalamus to stimulate and maintain gonadotropin secretion by the pituitary. tis important to ‘anestrus caused by insufficient hormonal stimuli and ‘apparent anestrus caused by failure to detect estrus Figure 7-5. Reproductive Cyclicity Profile of Queens With and Without Copulation Queen in estrus - no mating Relative Hormone Concentrations (blood) Mating Weeks ‘A queen enters estrus (about 9 days) every 17 days. If copulation does not occur, the queen enters a postestrus phase and comes into estrus a few days later. Since the queen is an induced ovulator, when mating does not occur, ovulation does not occur and a CL is not formed. ‘When mating occurs during estrus, ovulation Is induced, fertilization occurs and pregnancy takes place. After ‘ovwlation corporaluteaare formed causing a marked elevation in progesterone. After a 60 day gestation period, parturition occurs and lactation ensues. Lactational anestrus does not occur in the cat because she will come into estrus while lactating. Escaneado con CamScannerCiclicidad reproductiva 147 ———- Keser Como se puede ver en la Figura 7-4, la Perrano tiene un mestre defido como en otras especies Eldesarolla cial del tejdo hteo se produce durante el estrus poco después de a ovulacion, como en otfos ‘mamiferos. En la reina, etapas del ciclo estros incluyen proestrus, estro, poststus, diestrus y anestrus. Hay poca evidencia de estacionalidad enlas reinas y tienden a ser poliesttus. Sin embargo, a medida que ‘aumenta el fotoperiodo, aumentalalongitud de cela. Los felidos son ovuladores inducdos y se requiete cépula parala induccién de la oleada de LH, Posesrusesuntemina itaado grader un periodo nterestro que sigue al estro en una eina que no ha sdonducida a ovular por copula (vase la figura ' 7-5) Enlas reinas que no han cépulado, no se produce ovulacién ni forma de corpora lutea. Por lo tanto, ni se producen el metro (formacién de) ri diestrus. Coma ena mayoria de los ovuladoresinducdes, seria aprop'ado ‘Considerar que la hermbra permaneceria en una constante fase folicular hasta que se produzcala copula. Después dela ‘cOpulacion las hembras ovulan y sélo entonces se forman corporalutea Eneste contestolos ovadoes indue-dos constitujen una forma especial de ciclo estio que ro tere una vetadea hei Anestrus significa ‘Sin Estrus (caler)* Aveatusesura conden en guts eer presenta ciclos estros. Durante el anestrus los ovarios estén ‘elatwamente inactwos y r¥ los foliculos ovy adores nla ‘Corpora lutea funcional estan presentes. El anestrus es el ‘resultado de una liberaciin insuficente de GnRH del hypothalamus: para estimular y mantener la secrecion de gonadotropina porta hipdfisis. Es importante dstinguit entre el anestrus verdadero Ceausado por estimulos hormonaesingufcientes y el anestrus aparente causado por la falta de deteccion del estrus. Figura 7:5, Perfil de ciclidad reproductiva de las reinas con y sin cépula Reina enestro- sin apareamuento (Una rea wtta en celo (unas 90s) cada 17 dias. Sino se produce la copa aver entra enna fue pts yrtraenetrouncs gua despues, Dao qual rena ex un cxuadat nda cuando no se produce el aparearienta nosepradaels oven nose formance Houeanets Cuando se produce el apateamiento durante eda, cory onuaoén wepodula ferhiizacion y se produce el embarazo, Detzuet de ts owson se forman corpora eas ausando una elevacion marcada de la ‘progesterona. Después de un periodo de ‘estan dee das vepoducelapatuceny Seared acne Elves tote durante lactarcia Escaneado con CamScanner450 Reproductive Cyclicity Onset of Seasonal Cyclicity Is Similar to the Onset of Puberty Seasonal anestrus is characterized by a reduc- tion in the frequency of hypothalamic GnRH secretion (as in the prepubertal female). Before the breeding season can begin, the hypothalamus must be able to secrete sufficient quantities of GnRH to elicit a response by the anterior lobe of the pituitary. The release of FSH and LH at levels capable of maintain- ing follicular development and causing ovulation is required. Seasonal breeders can be categorized as either long-day breeders or short-day breeders (Sce Figure 7-1), The mare is characterized as a long-day breeder because as the day length increases in the spring the mare begins to cycle. During the short days of the winter months, the mare is anestrus. Short-day breed ers are animals that begin to cycle during the shorter days of fll, Animals such as sheep, deer, elk and goats are categorized as short-day breeders. The duration of the breeding season varies among and within species. For example, in sheep, the Merino breed has a period of cyclicity that ranges from 200 to 260 days, while blackface breeds have shorter periods of eyclicity ranging from 100 to 140 days. The two primary fuctors that influence the onset of the breeding season are photoperiod and temperature, Photoperiod is by far the most impor- tant, It is well known that artificial manipulation of the photoperiod can alter the cyclicity of the seasonal breeder. Figure 7-7. Possible Role of Kisspeptin Neurons in the Regulation of Cyclicity in Long-Day and Short-Day Breeders Suprachlasmadie ‘nucleus Long photoperiods (shorter dark periods) \ Anterior lobe © 1 Daylength -- } excitation of retinal neurons © Retinal neurons synapse in suprachiasmatic nucleus, © Inhibitory neurons (black neuron) convert excitatory response to an inhibitory response © Postsynaptic adrenergic fiber —-] norepinephrine secretion @ | norepinephrine — | melatonin by pinealocyte @ | motstonin —+ | RFRP-3 from RFRP neuron @ | RFRP-3 — | Kiss-10 — | GnRH — 1 FSH & LH = 1 kiss-10— | GNRH — | FSH & LH Posterior se Pineal gland ° Low norepinephrine secretion ex Present | ° Low melatonin release RFRP neuron YN Superior cervical tRERP-: RFRP-3, ganglion af © et te Short-day Long-day r— | kiss neurons kiss neurons Inhibited stimulated ‘ t Low kiss-10 @ High kiss -10 Hypothalamus-| 1 { L— jcnRH @ tGnRH Pituitary} }FSH+LH @ tFSH+LH Nocycles @ Cyclicity ® @ Escaneado con CamScannertou ux \aaa Keprogucuva eee ee El inicio de la ciclidad estacional es similar al inicio dela pubertad El anestrus estaconal se caracterza por una reducoiin ena frecuencia de secrecién de GnRH hipotalamica (como enta hembra prepibera) Antes de que pueda comenzar la temporada de reproduccién, el hipotalamo debe ser ‘capaz de secretar cantidades suficientes de GaRH para provocar una respuesta del lobulo anterior de la hipéfis, ‘Se requiere la liberacion de FSH y LH aniveles capaces de mantener el desarrollo folicular y causar ovulacién. Los, cnadores estacionales pueden clasifcarse como criadores de dia largo 0 de dia corto (véase la figura 7-1). La yegua se caracteriza como un erador ce larga duracién porque a medida que aumenta la duracion del dia en ta primavera el Ja yegua empieza a cicvlar. Durante los dias cortos de los meses de inviemo, la yegua es anestrus. Las crias de dia corto son animaies que comeenzan a circular durante los dias, mis coos dl otofo Animales coma oes, crs, alces ycabras se dasfican como ciadores de dia corto. La duracién de la epoca de reproducciin varia ent las especies y dentro de ellas. Por ejemplo, en las ovejas, ta raza Merino tiene un periodo de ciclidad que oscila entre 200 y 260 dias, ‘mientras que las razas de cara neg tienen periados decicldad mas cortos que van de 100 a 140 dias. Los dos factores principales que influyen en eliinicio de la temporada de reproduceién son el {otoperiodo y la temperatua. El fotoperiodo es, con mucho, el mas importante, Es bien sabido que la manipulacion artificial del fotoperiodo puede alterar la cclidad del criador estaconal Figura 7-7. Posible papel de las neuronas de Kisspeptina en la regulacién de la ciclidad en criadores de dia largo y de dia corto Nicleo Fotonenodos laryos (pertava seo mas rts) © Longtuddeiss—+ Excitacionfarmacotdgica de las ‘pauronas tetmianes 2 Sinapsis en ef nucleo supraquiasmatic © Las neurcras whibidoras (neurona negta) conlesten!a resouesta enna enurarespuests rhea © tora adrenéigica postinsinaptica, secrecon de norepinetina © de norepinetrina metatonina por pinealocitos. 6 melatonina-1 RFRP-3 de la neurona @ RFRP 1 RFRP-3 Kiss:10-1 GnRH HSH & LH HCKiss-10 GNRH-FSH & LH Gionduiapreal ° Secrestnta de 7 Pinealocitos ° Lmeercentajaae rebar ‘ Neurona RERP san cervical suet ‘reaps TRFRP.3 ot to wonasde ems tree woo 1 1 Beso bajo-10 Alto beso-10 Hgotilame + fonnH = @_~—GNRH Pruitaria, [3 FSH+uH 7 FSH+ {| Snecos 7 Ciclitud ® © Escaneado con CamScanner‘A major question that must be answered in order to understand the influence of day length on the onset of reproductive activity is, “How is photoperiod translated into a physiologic signal?” ‘Approposed pathway for both the fong-day and short-day breeder is presented in Figure 7-7. During Jong photoperiods, the retina of the eye is stimulated by light. This results in elevated tonic excitation of retinal neurons. This excitation is transmitted by anerve tract, to a specific area of the hypothalamus known as the suprachiasmatic nucleus. From the suprachiasmatic nucleus a second nerve tract travels to the superior lion. The presynaptic neurons synapse itory neurons that convert an excitatory sig- nal into into an inhibitory response. As a result, the postsynaptic adrenergic fibers are inhibited and they reduce their secretion of norepinephrine, Reduced norepinephrine results in low melatonin secretion from the pineal gland, Low melatonin results in excitation of RFRP neurons and they increase secretion of their neurotransmitter, RFRP-3, The RFRP neuron’s name is derived from the following: a) the “RF” designation re- fers to “amide related proteins” that are small peptides secreted by the neurons; b) the second “R” refers to the amino acid arginine and c) the second “P" refers o the amino acid phenylalanine, The RF amide molecule has Reproductive Cyclicity 454 an arg-phe-NH, at the C terminus and is probably 10 amino acids in length. Elevated RFRP-3 has different effects in the short and long-day breeder. For example, in the long-day breeder, RFRP-3 stimulates groupings of kisspeptin neurons in the hypothalamus and they secrete high levels of kisspeptin-10. It is thought that kisspeptin-10 acts directly on GnRH neurons to stimulate the secretion of FSH and LH. As as conse- quence, the long-day female begins to eycle. In the short-day breeder, kisspeptin neurons are thought to be inhibited by RFRP-3 and thus kisspeptin-10 secre- tion is reduced and GnRH neurons do not stimulate the release of FSH and LH. In summary, its thought that the fundamental reason that differences between seasonal breeders ex- ists (short-day versus long-day) is related to genetic differences in the responsiveness of certain groups of kisspeptin neurons to RFRP-3. When days are short, melatonin increases, which in tum decreases the RFRP-3 inhibition on kisspeptin neurons. In short-day breeding females, this signal elevates levels, of GnRH and thus FSH and LH to initiate cyclicity. On the other hand, these conditions (high melatonin during short days) signal the long-day breeding female to reduce levels of GnRH and thus low FSH and LH terminates cyclicity. Figure 7-8, Influence of Suckling Frequency Upon Blood LH (a Direct Indication of GnRH Release) in Postpartum Beef Cows (Derived from the data of Dr. G.L. Williams, Texas A&M University, Beeville) When the number of suckling sessions Is between 3 and 20 per day, amplitude and pulse frequency of blood LH are quite low and the cow remains in anestrus. ‘When the number of suckling sessions is limited to two or less per day, the amplitude and pulse frequency of LH increases dramatically and the cow will begin to cycle. ANESTRUS Number of Suckling Sessions per Day Threshold Parturition Weeks Postpartum Escaneado con CamScanner