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320  Veterinary Histology of Domestic Mammals and Birds



                   9, ‘Endocrine System’). Neurosecretory cells located in   In the  oestrous phase, the follicle develops to full
       VetBooks.ir  hypothalamic nuclei periodically secrete releasing and inhib-  maturity and blood oestrogen levels peak. Levels of LH
                   iting hormones that are transported by the hypophyseal  rise rapidly while FSH secretion begins to decline. In most
                   portal system to the adenohypophysis, where they stimulate  species, a surge in LH is followed by ovulation (spontane-
                   or inhibit the release of gonadotrophic hormones.  ous ovulation). Ovulation in cats occurs in response to
                      Ovarian function is regulated by the gonadotropins  mating (induced ovulation). Depending on species, ovula-
                   follicle-stimulating hormone (FSH), luteinising hormone  tion occurs at the end of oestrus or at various timepoints
                   (LH) and prolactin. Luteinising hormone stimulates the  during metoestrus.
                   production of androgens (testosterone, androstenedione)   In  metoestrus, the corpus haemorrhagicum trans-
                   by thecal cells. Follicle-stimulating hormone activates the  forms into the corpus luteum, which begins to synthesise
                   aromatase enzyme system of the granulosa cells for con-  progesterone.
                   version of androgens to oestrogens. The ratio of FSH:LH   The corpus luteum reaches maximum activity in dioes-
                   determines the timing of ovulation. Production of pro-  trus. Regression of the corpus luteum is triggered by
                   gesterone by the corpus luteum is mediated by prolactin.  prostaglandins (PGF ) produced by the uterine mucosa
                                                                                   2α
                   Feedback loops regulate the release of gonadotropins from  (corpus luteum cyclicum). If conception occurs, progester-
                   the adenohypophysis.                           one production is maintained (corpus luteum graviditatis)
                      The oestrous cycle is divided into the following phases:  and prostaglandin-induced retrogression of the corpus
                                                                  luteum is delayed until the end of pregnancy. Regression
                     ·  pro-oestrus,                              is followed by development of the corpus albicans.
                     ·  oestrus,
                     ·  metoestrus and                            Hormonal effects on the uterine tubes
                     ·  dioestrus.                                The uterine tubes are quiescent in pro-oestrus. During
                                                                  oestrus, rising oestrogen levels lead to increased activity
                   In theriogenology, metoestrus, dioestrus and pro-oestrus  of the secretory epithelial cells.
                   are also combined under the term interoestrus, repre-  The greatest volumes of secretion are produced during
                   senting the period between oestrous phases of the cycle.  metoestrus when the oocyte is in the ampulla tubae uteri-
                   Anoestrus, the period in which oestrus does not occur, is  nae. This is accompanied by increased beating of the cilia.
                   observed in monoestrous species (bitch, see below) at the   In the subsequent phase, dioestrus, the activity of the
                   end of a prolonged metoestrus, or after conception.  uterine tube epithelium rapidly declines. The cells become
                      Division of the oestrous cycle into phases is based on  shorter and partly degenerate.
                   structural changes in the reproductive organs and associ-
                   ated fluctuations in circulating hormone concentrations.  Cyclic changes in the uterus
                   Cyclic changes in FSH, oestrogens, progesterone, LH and  During pro-oestrus, the uterine epithelium grows, the
                   prolactin vary among species. Hormone levels measured  glands enlarge but remain straight and vascularisation
                   in the blood do not necessarily indicate the sensitivity of  increases (proliferative phase).
                   hormone receptors on the target cells (refer to veterinary   Uterine changes reach their peak during oestrus. Under
                   endocrinology and physiology texts).           the influence of oestrogen, epithelial cells and uterine
                      The length of the oestrous cycle varies considerably  glands release secretions (uterine milk), the spinocellular
                   among domestic mammals. Based on cycle frequency, a  subepithelial connective tissue thickens and intercellular
                   distinction is made between:                   spaces become expanded and oedematous.
                                                                     Hyperplasia of the uterine glands continues into
                     ·  polyoestrous species (e.g. mare, cow, sow) and  metoestrus. During this phase, the connective tissue layer
                     ·  monoestrous species (e.g. bitch).         thins and oedema decreases.
                                                                     In dioestrus, the glands become extensively coiled and
                   In monoestrous animals, an extended period of metoes-  shortened. Progesterone brings about maximal glandu-
                   trus (50–70 days) is followed by anoestrus. In polyoestrous  lar secretion, which is sustained if pregnancy occurs. If
                   species, the corpus luteum regresses and the animal enters  implantation does not take place, secretory activity gradu-

                   pro-oestrus and then oestrus (see endocrinology and  ally decreases, the blood supply decreases and the glands
                   theriogenology texts).                         revert to their tubular morphology (involution).
                                                                     Changes in the cervix are closely linked with those
                   Cyclic changes in the ovary                    occurring in the uterus. Under the influence of oestrogens,
                   During pro-oestrus, release of FSH from the adenohy-  the cervical epithelium secretes increasing amounts of
                   pophysis brings about follicular maturation. Oestrogen  thin, clear mucus. During dioestrus, the mucus becomes
                   secretion increases.                           more viscous.









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