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Chapter Opener 28 © 2013 Pearson Education, Inc. Figure 28.1 Diagrams showing the approximate size of a human conceptus from fertilization to the early fetal stage. Embryo Fertilization 1-week conceptus 3-week embryo (3 mm) 5-week embryo (10 mm) 8-week embryo (22 mm) 12-week fetus (90 mm) © 2013 Pearson Education, Inc. Figure 28.2 Sperm use acrosomal enzymes and receptors to approach, bind, and enter the oocyte. The cortical reaction prevents further sperm entry, ensuring that only two copies of each chromosome are present in the fertilized ovum. 5 Block of polyspermy. 4 Fusion. Sperm, delivered to the vagina and capacitated in the female reproductive tract, stream toward a secondary oocyte. 2 Acrosomal reaction. Binding of the sperm to sperm-binding receptors in the zona pellucida causes the Ca2+ levels within the sperm to rise, triggering the acrosomal reaction. Acrosomal enzymes from many sperm digest holes through the zona pellucida, clearing a path to the oocyte membrane. 3 Binding. The sperm’s membrane binds to the oocyte’s Sperm-binding receptors. The sperm and oocyte plasma membranes fuse, allowing sperm contents to enter the oocyte. 1 Approach. Aided by enzymes on its surface, a sperm cell weaves its way past granulosa cells of the corona radiata. Entry of the sperm’s contents causes Ca2+ levels in the oocyte’s cytoplasm to rise, triggering the cortical reaction (exocytosis of cortical granules). As a result, the zona pellucida hardens and the sperm receptors are clipped off (slow block to polyspermy). Extracellular space Sperm Sperm Zona pellucida Polar body Oocyte nucleus arrested in meiotic metaphase II Granulosa cells of corona radiata Zona pellucida Sperm-binding receptors Oocyte spermbinding membrane receptors Cortical granules Microtubules from sperm flagellum Mitochondria Zona pellucida Extracellular space Oocyte plasma membrane © 2013 Pearson Education, Inc. Sperm nucleus Figure 28.2 Sperm use acrosomal enzymes and receptors to approach, bind, and enter the oocyte. The cortical reaction prevents further sperm entry, ensuring that only two copies of each chromosome are present in the fertilized ovum. (1 of 2) Sperm, delivered to the vagina and capacitated in the female reproductive tract, stream toward a secondary oocyte. 1 Approach. Aided by enzymes on its surface, a sperm cell weaves its way past granulosa cells of the corona radiata. Extracellular space Sperm Sperm Zona pellucida Polar body Granulosa cells of corona radiata Oocyte nucleus arrested in Meiotic metaphase II Zona pellucida Extracellular space Oocyte plasma membrane © 2013 Pearson Education, Inc. Figure 28.2 Sperm use acrosomal enzymes and receptors to approach, bind, and enter the oocyte. The cortical reaction prevents further sperm entry, ensuring that only two copies of each chromosome are present in the fertilized ovum. (2 of 2) 4 Fusion. The 5 Block of 2 Acrosomal reaction. Binding of the sperm to spermbinding receptors in the zona pellucida causes the Ca2+ levels within the sperm to rise, triggering the acrosomal reaction. Acrosomal enzymes from many sperm digest holes through the zona pellucida, clearing a path to the oocyte membrane. Zona pellucida Sperm-binding receptors © 2013 Pearson Education, Inc. sperm and oocyte plasma 3 Binding. membranes The sperm’s membrane binds fuse, allowing to the oocyte’s sperm contents sperm-binding to enter the oocyte. receptors. Oocyte sperm-binding membrane receptors Cortical granules polyspermy. Entry of the sperm’s contents causes Ca2+ levels in the oocyte’s cytoplasm to rise, triggering the cortical reaction (exocytosis of cortical granules). As a result, the zona pellucida hardens and the sperm receptors are clipped off (slow block to polyspermy). Microtubules from sperm flagellum Sperm Mitochondria nucleus Figure 28.3 Events of fertilization. Extracellular space Corona radiata Zona pellucida Second meiotic division of oocyte Second meiotic division of first polar body 1 After the sperm penetrates the secondary oocyte, the oocyte completes meiosis II, forming the ovum and second polar body. Male pro-nucleus Female pro-nucleus (swollen ovum nucleus) Polar bodies 2 Sperm and ovum nuclei swell, forming pronuclei. Male pronucleus Mitotic spindle Centriole Female pronucleus Zygote Sperm nucleus 3 Pronuclei approach each other and mitotic spindle forms between them. 4 Chromosomes of the pronuclei intermix. Fertilization is accomplished. Then, the DNA replicates in preparation for the first cleavage division. Male and female pronuclei Polar bodies © 2013 Pearson Education, Inc. Figure 28.3a Events of fertilization. (1 of 4) Sperm nucleus Extracellular space Corona radiata Zona pellucida Second meiotic division of oocyte Second meiotic division of first polar body © 2013 Pearson Education, Inc. 1 After the sperm penetrates the secondary oocyte, The oocyte Completes meiosis II, forming the ovum and second polar body. Figure 28.3a Events of fertilization. (2 of 4) Male pronucleus Female pronucleus (swollen ovum nucleus) Polar bodies © 2013 Pearson Education, Inc. 2 Sperm and ovum nuclei swell, forming pronuclei. Figure 28.3a Events of fertilization. (3 of 4) Male pronucleus Mitotic spindle Centriole Female pronucleus © 2013 Pearson Education, Inc. 3 Pronuclei approach each other and mitotic spindle forms between them. Figure 28.3a Events of fertilization. (4 of 4) Zygote © 2013 Pearson Education, Inc. 4 Chromoomes of the pronuclei intermix. Fertilization is accomplished. Then, the DNA replicates in preparation for the first cleavage division. Figure 28.3b Events of fertilization. Male and female pronuclei Polar bodies © 2013 Pearson Education, Inc. Figure 28.4 Cleavage: From zygote to blastocyst. 4-cell stage 2 days Zygote (fertilized egg) Morula (a solid ball of blastomeres). 3 days Zona pellucida Degenerating zona pellucida Sperm Blastocyst cavity Uterine tube Fertilization (sperm meets and enters egg) Early blastocyst (Morula hollows out, fills with fluid, and “hatches” from the zona pellucida). 4 days Implanting blastocyst (Consists of a sphere of trophoblast cells and an eccentric cell cluster called the inner cell mass). 7 days Ovary Oocyte (egg) Trophoblast Ovulation Uterus Endometrium Cavity of uterus © 2013 Pearson Education, Inc. Blastocyst cavity Inner cell mass Zygote (fertilized egg) Figure 28.4a Cleavage: From zygote to blastocyst. Zona pellucida © 2013 Pearson Education, Inc. Figure 28.4b Cleavage: From zygote to blastocyst. 4-cell stage 2 days © 2013 Pearson Education, Inc. Morula (a solid ball of blastomeres). 3 days Figure 28.4c Cleavage: From zygote to blastocyst. © 2013 Pearson Education, Inc. Figure 28.4d Cleavage: From zygote to blastocyst. Early blastocyst (Morula hollows out, fills with fluid, and “hatches” from the zona pellucida). 4 days Degenerating zona pellucida Blastocyst cavity © 2013 Pearson Education, Inc. Figure 28.4e Cleavage: From zygote to blastocyst. Implanting blastocyst (Consists of a sphere of trophoblast cells and an eccentric cell cluster called the inner cell mass). 7 days Inner cell mass Trophoblast © 2013 Pearson Education, Inc. Blastocyst cavity Figure 28.5 Implantation of the blastocyst. Endometrium Uterine endometrial epithelium Inner cell mass Trophoblast Blastocyst cavity Lumen of uterus Endometrial stroma with blood vessels and glands Syncytiotrophoblast Cytotrophoblast Blastocyst cavity Lumen of uterus © 2013 Pearson Education, Inc. Figure 28.5a Implantation of the blastocyst. Endometrium Uterine endometrial epithelium Inner cell mass Trophoblast Blastocyst cavity Lumen of uterus © 2013 Pearson Education, Inc. Figure 28.5b Implantation of the blastocyst. Endometrium Uterine endometrial epithelium Inner cell mass Trophoblast © 2013 Pearson Education, Inc. Figure 28.5c Implantation of the blastocyst. Endometrial stroma with blood vessels and glands Syncytiotrophoblast Cytotrophoblast Blastocyst cavity Lumen of uterus © 2013 Pearson Education, Inc. Figure 28.5d Implantation of the blastocyst. Endometrial stroma with blood vessels and glands Syncytiotrophoblast Cytotrophoblast Lumen of uterus © 2013 Pearson Education, Inc. Figure 28.6 Hormonal changes during pregnancy. Relative blood levels Human chorionic gonadotropin Estrogens Progesterone 0 4 8 Ovulation and fertilization © 2013 Pearson Education, Inc. 12 16 24 20 28 Gestation (weeks) 32 36 Birth Figure 28.7 Events of placentation, early embryonic development, and extraembryonic membrane formation. Endometrium Lacuna (intervillous space) containing maternal blood Maternal blood vessels Amniotic cavity Primary germ layers Chorionic villus • Ectoderm Chorion • Mesoderm Amnion • Endoderm Proliferating syncytiotrophoblast Forming umbilical cord Allantois Cytotrophoblast Amniotic cavity Yolk sac Bilayered embryonic disc • Epiblast • Hypoblast Endometrial epithelium Implanting 71/2 -day blastocyst. The syncytiotrophoblast is eroding the endometrium. Cells of the embryonic disc are now separated from the amnion by a fluid-filled space. Extraembryonic mesoderm Chorion being formed Lumen of uterus 12-day blastocyst. Implantation is complete. Extraembryonic mesoderm is forming a discrete layer beneath the cytotrophoblast. Extraembryonic coelom 16-day embryo. Cytotrophoblast and associated mesoderm have become the chorion, and chorionic villi are elaborating. The embryo exhibits all three germ layers, a yolk sac, and an allantois, which forms the basis of the umbilical cord. Placenta Decidua basalis Decidua basalis Maternal blood Chorionic villi Chorionic villus Yolk sac Umbilical blood vessels in umbilical cord Amnion Amniotic cavity Amnion Amniotic cavity Yolk sac Umbilical cord Extraembryonic coelom Lumen of uterus Chorion Decidua capsularis Decidua capsularis Extraembryonic 41/2 -week embryo. The decidua capsularis, decidua basalis, amnion, and coelom yolk sac are well formed. The chorionic villi lie in blood-filled intervillous spaces within the endometrium. The embryo is nourished via the umbilical 13-week fetus. vessels that connect it (through the umbilical cord) to the placenta. © 2013 Pearson Education, Inc. Uterus Lumen of uterus Figure 28.7a–c Events of placentation, early embryonic development, and extraembryonic membrane formation. Endometrium Lacuna (intervillous space) containing maternal blood Maternal blood vessels Proliferating syncytiotrophoblast Chorionic villus • Ectoderm Chorion • Mesoderm Amnion • Endoderm Cytotrophoblast Amniotic cavity Yolk sac Implanting 71/2 -day blastocyst. The syncytiotrophoblast is eroding the endometrium. Cells of the embryonic disc are now separated from the amnion by a fluid-filled space. © 2013 Pearson Education, Inc. Forming umbilical cord Allantois Bilayered embryonic disc • Epiblast • Hypoblast Endometrial epithelium Amniotic cavity Primary germ layers Extraembryonic mesoderm Chorion being formed Lumen of uterus 12-day blastocyst. Implantation is complete. Extraembryonic mesoderm is forming a discrete layer beneath the cytotrophoblast. Extraembryonic coelom 16-day embryo. Cytotrophoblast and associated mesoderm have become the chorion, and chorionic villi are elaborating. The embryo exhibits all three germ layers, a yolk sac, and an allantois, which forms the basis of the umbilical cord. Figure 28.7d Events of placentation, early embryonic development, and extraembryonic membrane formation. Decidua basalis Maternal blood Chorionic villus Umbilical blood vessels in umbilical cord Amnion Amniotic cavity Yolk sac Extraembryonic coelom Lumen of uterus Chorion Decidua capsularis 41/2 -week embryo. The decidua capsularis, decidua basalis, amnion, and yolk sac are well formed. The chorionic villi lie in blood-filled intervillous spaces within the endometrium. The embryo is nourished via the umbilical vessels that connect it (through the umbilical cord) to the placenta. © 2013 Pearson Education, Inc. Figure 28.7e Events of placentation, early embryonic development, and extraembryonic membrane formation. Placenta Decidua basalis Chorionic villi Yolk sac Amnion Amniotic cavity Umbilical cord Decidua capsularis Extraembryonic coelom 13-week fetus. © 2013 Pearson Education, Inc. Uterus Lumen of uterus Figure 28.8 Detailed anatomy of the vascular relationships in the mature decidua basalis. Placenta Chorionic villi Decidua basalis Maternal arteries Umbilical cord Decidua capsularis Uterus Lumen of uterus Chorionic villus containing fetal capillaries Maternal blood in lacuna (intervillous space) Fetal arteriole Fetal venule Amnion Umbilical cord © 2013 Pearson Education, Inc. Maternal veins Myometrium Stratum basalis of endometrium Maternal portion of placenta (decidua basalis) Fetal portion of placenta (chorion) Umbilical arteries Umbilical vein Connection to yolk sac Figure 28.9 Formation of the three primary germ layers. Amnion Bilayered embryonic disc Head end of bilayered embryonic disc Yolk sac Frontal section 3-D view Section view in (e) Primitive streak Head end Cut edge of amnion Epiblast Yolk sac (cut edge) Right Left 14-15 days Hypoblast Endoderm Ectoderm Primitive streak Tail end Bilayered embryonic disc, superior view © 2013 Pearson Education, Inc. 16 days Mesoderm Endoderm Figure 28.10 Folding of the embryonic body, lateral views. Head Tail Amnion Yolk sac Ectoderm Mesoderm Endoderm Trilaminar embryonic disc Future gut (digestive tube) Lateral fold Somites (seen through ectoderm) Tail fold Head fold Yolk sac Neural tube Notochord Primitive gut Hindgut © 2013 Pearson Education, Inc. Yolk sac Foregut Figure 28.10a Folding of the embryonic body, lateral views. Head Tail Amnion Yolk sac Ectoderm Mesoderm Endoderm © 2013 Pearson Education, Inc. Trilaminar embryonic disc Figure 28.10b Folding of the embryonic body, lateral views. Lateral fold © 2013 Pearson Education, Inc. Future gut (digestive tube) Figure 28.10c Folding of the embryonic body, lateral views. Somites (seen through ectoderm) Tail fold Head fold Yolk sac © 2013 Pearson Education, Inc. Figure 28.10d Folding of the embryonic body, lateral views. Neural tube Notochord Primitive gut Hindgut © 2013 Pearson Education, Inc. Yolk sac Foregut Figure 28.11 Endodermal differentiation. Pharynx Parathyroid glands and thymus Thyroid gland Esophagus Trachea Connection to yolk sac Right and left lungs Stomach Liver Umbilical cord Pancreas Gallbladder Small intestine Allantois Large intestine 5-week embryo © 2013 Pearson Education, Inc. Figure 28.12 Neurulation and early mesodermal differentiation. Head Amnion Amniotic cavity Left Right Neural plate Primitive streak Cut edge of amnion Tail 17 days. The flat three-layered embryo has completed gastrulation. Notochord and neural plate are present. Ectoderm Mesoderm Notochord Endoderm Yolk sac Neural crest Neural groove Neural fold Somite Intermediate mesoderm Lateral plate mesoderm Coelom 20 days. The neural folds form by folding of the neural plate, which then deepens, producing the neural groove. Three mesodermal aggregates form on each side of the notochord (somite, intermediate mesoderm, and lateral plate mesoderm). Surface ectoderm 22 days. The neural folds have closed, forming the neural tube which has detached from the surface ectoderm and lies between the surface ectoderm and the notochord. Embryonic body is beginning to undercut. Neural crest Neural tube Somite Notochord Neural tube (ectoderm) Somite Dermatome Myotome Sclerotome Kidney and gonads (intermediate mesoderm) Epidermis (ectoderm) Gut lining (endoderm) Lateral plate mesoderm • Limb bud • Smooth muscle of gut • Visceral serosa Peritoneal cavity (coelom) © 2013 Pearson Education, Inc. • Parietal serosa • Dermis End of week 4. Embryo undercutting is complete. Somites have subdivided into sclerotome, myotome, and dermatome, which form the vertebrae, skeletal muscles, and dermis respectively. Body coelom present. Figure 28.12a Neurulation and early mesodermal differentiation. Head Amnion Amniotic cavity Left Right Cut edge of amnion Primitive streak Tail Neural plate Ectoderm Mesoderm Notochord Endoderm Yolk sac © 2013 Pearson Education, Inc. 17 days. The flat three-layered embryo has completed gastrulation. Notochord and neural plate are present. Figure 28.12b Neurulation and early mesodermal differentiation. Neural crest Neural groove Neural fold Coelom © 2013 Pearson Education, Inc. Somite Intermediate mesoderm 20 days. The neural folds form by folding of the neural plate, which then deepens, producing the neural groove. Three mesodermal Lateral plate aggregates form on each side of the notochord (somite, mesoderm intermediate mesoderm, and lateral plate mesoderm). Figure 28.12c Neurulation and early mesodermal differentiation. Surface ectoderm Neural crest Neural tube Somite Notochord © 2013 Pearson Education, Inc. 22 days. The neural folds have closed, forming the neural tube which has detached from the surface ectoderm and lies between the surface ectoderm and the notochord. Embryonic body is beginning to undercut. Figure 28.12d Neurulation and early mesodermal differentiation. Neural tube (ectoderm) Somite Dermatome Myotome Sclerotome Kidney and gonads (intermediate mesoderm) Epidermis (ectoderm) Gut lining (endoderm) Lateral plate mesoderm • Limb bud • Smooth muscle of gut • Visceral serosa Peritoneal cavity (coelom) © 2013 Pearson Education, Inc. • Parietal serosa • Dermis End of week 4. Embryo undercutting is complete. Somites have subdivided into sclerotome, myotome, and dermatome, which form the vertebrae, skeletal muscles, and dermis respectively. Body coelom present. Figure 28.13 Flowchart showing major derivatives of the embryonic germ layers. Epiblast ECTODERM MESODERM Notochord Somite Intermediate mesoderm ENDODERM Lateral plate mesoderm Somatic mesoderm • Epidermis, hair, nails, glands of skin • Brain and spinal cord • Neural crest and derivatives (e.g., cranial, spinal, and sympathetic ganglia and associated nerves; chromaffin cells of the adrenal medulla; pigment cells of the skin) © 2013 Pearson Education, Inc. Nucleus pulposus of intervertebral discs • Sclerotome: vertebrae and ribs • Dermatome: dermis of dorsal body region • Myotome: trunk and limb musculature • Kidneys • Parietal serosa • Gonads • Dermis of ventral body region • Connective tissues of limbs (bones, joints, and ligaments) Splanchnic mesoderm • Wall of digestive and respiratory tracts (except epithelial lining) • Visceral serosa • Heart • Blood vessels Epithelial lining and glands of digestive and respiratory tracts Figure 28.14 Circulation in fetus and newborn. Newborn Fetus Aortic arch Superior vena cava Ductus arteriosus Ligamentum arteriosum Pulmonary artery Pulmonary veins Heart Lung Foramen ovale Fossa ovalis Liver Ductus venosus Ligamentum venosum Hepatic portal vein Umbilical vein Ligamentum teres Inferior vena cava Umbilicus Abdominal aorta Common iliac artery Umbilical arteries Medial umbilical ligaments Urinary bladder Umbilical cord Placenta High oxygenation Moderate oxygenation Low oxygenation Very low oxygenation © 2013 Pearson Education, Inc. Figure 28.14a Circulation in fetus and newborn. Fetus Aortic arch Superior vena cava Ductus arteriosus Ligamentum arteriosum Pulmonary artery Pulmonary veins Heart Lung Foramen ovale Fossa ovalis Liver Ductus venosus Ligamentum venosum Hepatic portal vein Umbilical vein Ligamentum teres Inferior vena cava Umbilicus Abdominal aorta Common iliac artery Umbilical arteries Medial umbilical ligaments Urinary bladder Umbilical cord Placenta © 2013 Pearson Education, Inc. High oxygenation Moderate oxygenation Low oxygenation Very low oxygenation Figure 28.14b Circulation in fetus and newborn. Aortic arch Superior vena cava Ductus arteriosus Newborn Ligamentum arteriosum Pulmonary artery Pulmonary veins Heart Lung Foramen ovale Fossa ovalis Liver Ductus venosus Ligamentum venosum Hepatic portal vein Umbilical vein Ligamentum teres Inferior vena cava Umbilicus Abdominal aorta Common iliac artery Umbilical arteries Medial umbilical ligaments Urinary bladder © 2013 Pearson Education, Inc. High oxygenation Moderate oxygenation Low oxygenation Very low oxygenation Figure 28.15 Photographs of a developing fetus. Amniotic sac Umbilical cord Umbilical vein Chorionic villi Yolk sac Cut edge of chorion Embryo at week 7, about 17 mm long. © 2013 Pearson Education, Inc. Fetus in month 3, about 6 cm long. Fetus late in month 5, about 19 cm long. Figure 28.15a Photographs of a developing fetus. Amniotic sac Umbilical cord Umbilical vein Chorionic villi Yolk sac Cut edge of chorion Embryo at week 7, about 17 mm long. © 2013 Pearson Education, Inc. Figure 28.15b Photographs of a developing fetus. Fetus in month 3, about 6 cm long. © 2013 Pearson Education, Inc. Figure 28.15c Photographs of a developing fetus. © 2013 Pearson Education, Inc. Fetus late in month 5, about 19 cm long. Figure 28.16 Relative size of the uterus before conception and during pregnancy. Before conception (Uterus the size of a fist and resides in the pelvis.) © 2013 Pearson Education, Inc. 4 months (Fundus of the uterus is halfway between the pubic symphysis and the umbilicus.) 7 months (Fundus is well above the umbilicus.) 9 months (Fundus reaches the xiphoid process.) Table 28.1 Developmental Events of the Fetal Period (1 of 3) © 2013 Pearson Education, Inc. Table 28.1 Developmental Events of the Fetal Period (2 of 3) © 2013 Pearson Education, Inc. Table 28.1 Developmental Events of the Fetal Period (3 of 3) © 2013 Pearson Education, Inc. Figure 28.17 Hormonal induction of labor. Start Estrogen Oxytocin from placenta from fetus and mother's posterior pituitary Induces oxytocin receptors on uterus Stimulates uterus to contract Stimulates placenta to release (+) Prostaglandins Stimulate more vigorous contractions of uterus © 2013 Pearson Education, Inc. Positive feedback (+) Figure 28.18 Parturition. 1a Early dilation. Baby’s head engaged; widest dimension Is along left-right axis. 1b Late dilation. Baby’s head rotates so widest dimension is in anteroposterior axis (of pelvic outlet). Dilation nearly complete Umbilical cord Placenta Uterus Cervix Vagina Pubic symphysis Sacrum 2 Expulsion. Baby’s head extends as it is delivered 3 Placental stage. After baby is delivered, the placenta detaches and is removed. © 2013 Pearson Education, Inc. Perineum Uterus Placenta (detaching) Umbilical cord Figure 28.18 Parturition. (1 of 4) 1a Early dilation. Baby’s head engaged; widest dimension Is along left-right axis. Umbilical cord Placenta Uterus Cervix Vagina © 2013 Pearson Education, Inc. Figure 28.18 Parturition. (2 of 4) 1b Late dilation. Baby’s head rotates so widest dimension is in anteroposterior axis (of pelvic outlet). Dilation nearly complete Pubic symphysis Sacrum © 2013 Pearson Education, Inc. Figure 28.18 Parturition. (3 of 4) 2 Expulsion. Baby’s head extends as it is delivered Perineum © 2013 Pearson Education, Inc. Figure 28.18 Parturition. (4 of 4) 3 Placental stage. After baby is delivered, the placenta detaches and is removed © 2013 Pearson Education, Inc. Uterus Placenta (detaching) Umbilical cord Figure 28.19 Milk production and the positive feedback mechanism of the milk let-down reflex. Hypothalamus releases prolactin releasing factors (PRF) to portal circulation. Start Positive feedback Stimulation of mechanoreceptors in nipples by suckling infant sends afferent impulses to the hypothalamus. Hypothalamus sends efferent impulses to the posterior pituitary where oxytocin is stored. Anterior pituitary secretes prolactin to blood. Oxytocin is released from the posterior pituitary and stimulates myoepithelial cells of breasts to contract. Prolactin targets mammary glands of breasts. Let-down reflex. Milk is ejected through ducts of nipples. © 2013 Pearson Education, Inc. Milk production Closer Look 28.1 © 2013 Pearson Education, Inc.