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Basic Structure and Function of the Cervix Tc3ec01 6/29/06 10:45 Page 2 Tc3ec01 6/29/06 10:45 Page 3 TC3eC01 6/29/06 10:45 Page 1 PART 1 Basic structure and function of the cervix TC3eC01 6/29/06 10:45 Page 2 TC3eC01 6/29/06 10:45 Page 3 CHAPTER 1 Morphogenesis and differentiation of the cervicovaginal epithelium John M. McLean Generally, the crown–rump length of larger embryos and all INTRODUCTION fetuses should be stated in preference to, or at least in addition Although the study of prenatal life dates back to antiquity, to, the supposed age (O’Rahilly and Muller, 1987) but, in this the first attempt to describe human prenatal development, account, the reference point will be the postovulatory age, utilising only human material, was the Manual of Human i.e. the length of time since the last ovulation, related, when Embryology published in two volumes by Kiebel and Mall in appropriate, to the Carnegie stage. As ovulation and fertilisa- 1910 and 1912. The Carnegie collection, established by Mall tion are closely related in time, the postovulatory interval is in 1915 with 813 human embryos, is now, with its several an adequate measure of embryonic age. Embryonic age, thousand specimens, the most important and most thoroughly length and stage are all interrelated. Age, however, conveys studied collection of human embryos in the world. an immediate meaning as it is a familiar yardstick, but it must As neonates of placental mammals must be capable of sus- be recognised that prenatal ages are only as useful as postnatal taining extra-uterine life, all essential organs are developed and ages, because they are reference points for the usual pattern or functional at birth. This prerequisite for survival is achieved range of developmental events. during markedly different periods of gestation. An interval of Much of the available information on human embryogenesis 14–16 days between fertilisation and parturition is sufficient is derived from traditional embryological studies. However, for the hamster, 20–23 days suffice for mice and rats while the with the advent of assisted conception and the establishment comparable period for elephants is 22 months and for humans of the Human Fertilisation and Embryology Authority, direct is approximately 38 weeks. The first 8 weeks of prenatal observation of the human embryo in vitro has provided an human life constitute the embryonic phase and the remaining additional source of information on the preimplantation phase 30 weeks the fetal phase. During embryogenesis, all essential of human embryogenesis. Furthermore, the use of sophist- organ systems develop and, with the exception of the lungs, icated computer technology in association with specifically are functional while the recognisable form of the human infant designed three-dimensional transvaginal probes allows more is established. The majority of congenital anomalies arise as a detailed in vivo three-dimensional ultrasound reconstructions result of defective morphogenesis during the later part of of embryos and early fetuses. The results obtained using these embryogenesis and early part of the fetal period. techniques correspond well to those obtained from classical Because development is a continuous process, various means human embryology (Blaas et al., 1998). have been used to identify and tabulate the progression of events during normal human embryogenesis. The founder of EARLY EMBRYOGENESIS the Carnegie collection, Mall, was the first to introduce stag- ing into human embryology. Mall’s observations, together The embryo, at fertilisation, initiates, sustains, directs and with those of his successor, Streeter, form the basis upon controls its own development. Embryogenesis begins with which the first 8 weeks of human development are described in fertilisation, Carnegie stage 1, and is followed by a phase 23 Carnegie stages. Using the Carnegie collection and other of preimplantation development, Carnegie stages 2 and 3. human embryos, including some fertilised in vitro, O’Rahilly Implantation begins on day 6 and is completed on day 12, (1973) produced the first comprehensive and authoritative Carnegie stages 4 and 5. During these few days, the embryo account of the prenatal period from fertilisation to the end of enters into an intimate vascular relationship with the mother the third week of gestation (Carnegie stages 1–9). In 1987, and becomes dependent on her for continued existence. Signi- with Muller as co-author, the 1973 publication was revised ficant events are also taking place within the embryo during and extended to cover the whole of embryogenesis (Carnegie this period: continuous cell division and differentiation; the stages 1–23). formation of the blastocyst within which is the embryonic 3 TC3eC01 6/29/06 10:45 Page 4 4 Basic structure and function of the cervix pole; the appearance of the amniotic cavity within the embry- Ectoderm of dorsal onic pole on day 7 and, with it, the formation of two basic cell surface of body types, ectoderm and endoderm, establishing the bilaminar Neural tube embryo; the appearance of the primitive umbilical cord on day Intraembryonic 11; and, during Carnegie stage 6, days 13–15, the formation mesoderm of the primitive streak and the conversion of the embryo into a Foregut trilaminar structure of ectoderm, intraembryonic mesoderm Midgut and endoderm. Hindgut Pericardial cavity The central nervous system and the skin, together with its appendages, arise from ectoderm; endoderm gives origin to Allantoic diverticulum the gastrointestinal tract, the respiratory tract and the urogen- ital sinus, while the remaining body structures are formed Vitellointestinal duct from intraembryonic mesoderm, within which the pericardial, Ventral yolk sac pleural and peritoneal cavities also develop. The flat trilaminar Body stalk or ventral early embryo, however, remains bilaminar at two sites, the (a) surface of body buccopharyngeal membrane at the rostral end of the embryo and the cloacal membrane at its caudal end. Paraxial mesoderm FLEXION Somite Intermediate mesoderm The neural tube, derived from ectoderm, is enclosed within Somatopleuric mesoderm the intraembryonic mesoderm during Carnegie stages 9–13, Peritoneal cavity days 19–28. The rapid growth of the neural tube causes the Splanchnopleuric flat trilaminar embryo to fold along its longitudinal and trans- mesoderm verse axes, creating a marked dorsal convexity and ventral Intraembryonic coelom continuous with concavity and forming head, tail and lateral folds (Fig. 1.1a extraembryonic coelom and b). (b) Body stalk The endoderm is drawn into the ventral concavity of the embryo and is subdivided into foregut, midgut and hindgut. Fig. 1.1 (a) A midline section of the embryo after formation of the head The hindgut is caudal to the rostral limit of the allantoic diver- and tail folds. (b) A transverse section of the midembryo region after ticulum and also dorsal and rostral to the cloacal membrane formation of the lateral folds. (Fig. 1.1a). The intraembryonic mesoderm in the midembryo region (Fig. 1.1b) is subdivided into paraxial mesoderm, lat- Intermediate eral mesoderm and intermediate mesoderm. The paraxial mesoderm mesoderm surrounds the neural tube and is the site of somite Spinal cord formation. The lateral mesoderm, so named because of its Dorsal mesentery Paraxial mesoderm Peritoneal cavity location in the flat trilaminar embryo, is carried ventrally by Mesonephric region the formation of the lateral folds. The lateral mesoderm Body stalk accommodates the primitive peritoneal cavity, which divides it Gonadal ridge into splanchnopleuric mesoderm, associated with endoderm Growing urorectal and destined to form the visceral muscle of the gut tube and septum (ventral bladder, and somatopleuric mesoderm, associated with ecto- mesoderm of derm and destined to participate in the formation of the body terminal hindgut) wall. Cloacal membrane The intermediate mesoderm extends the length of the Fig. 1.2 The caudal half of the embryo showing the gonadal ridge and body cavity. It is lateral and ventral to the paraxial mesoderm mesonephric region of the intermediate mesoderm which, at its caudal and adjacent to the midline, dorsal mesentery of the gut limit, is continuous with the mesoderm investing the hindgut. tube. At the caudal end of the primitive peritoneal cavity, the intermediate mesoderm is in continuity with the mesoderm The urorectal septum investing the terminal portion of the hindgut and, ventral to the hindgut, with the urorectal septum (Fig. 1.2). Within the The hindgut, established during the process of flexion, is the intermediate mesoderm, structures involved in the morpho- endoderm enclosed within the tail fold of the embryo. It lies genesis of the urinary and reproductive systems develop. caudal to the rostral limit of the allantoic diverticulum and TC3eC01 6/29/06 10:45 Page 5 Morphogenesis and differentiation of the cervicovaginal epithelium 5 Allantoic diverticulum Rostral Hindgut Ventral Dorsal Cloacal membrane Caudal Amniotic cavity (a) Fig. 1.3 (a) The primitive hindgut is enclosed within the embryonic tail fold. Urogenital sinus (b) The developing urorectal septum grows dorsally and caudally from the rostral limit Urorectal septum of the allantoic diverticulum. (c) The fusion Rectum of the urorectal septum with the cloacal membrane divides the hindgut into (b) Developing (c) urogenital sinus and rectum. urorectal septum Spinal cord Intermediate mesoderm Gut tube Peritoneal cavity Fig. 1.4 A transverse section through Urorectal septum the embryonic pelvis demonstrates the Urogenital sinus continuity of the intermediate mesoderm with the mesoderm
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