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CAMPBELL BIOLOGY IN FOCUS Urry • Cain • Wasserman • Minorsky • Jackson • Reece 36 Reproduction and Development Lecture Presentations by Kathleen Fitzpatrick and Nicole Tunbridge © 2014 Pearson Education, Inc. Overview: Pairing Up for Sexual Reproduction Each sea slug produces sperm and eggs and thus acts as both mother and father to the next generation Animal reproduction takes many forms A population outlives its members only by reproduction, the generation of new individuals from existing ones © 2014 Pearson Education, Inc. Figure 36.1 © 2014 Pearson Education, Inc. Concept 36.1: Both asexual and sexual reproduction occur in the animal kingdom Sexual reproduction is the creation of an offspring by fusion of a male gamete (sperm) and female gamete (egg) to form a zygote Asexual reproduction is the creation of offspring without the fusion of egg and sperm © 2014 Pearson Education, Inc. Mechanisms of Asexual Reproduction Many invertebrates reproduce asexually by budding, in which new individuals arise from outgrowths of existing ones Also common among invertebrates is fission, the separation of a parent organism into two individuals of approximately equal size Video: Hydra Budding © 2014 Pearson Education, Inc. Figure 36.2 © 2014 Pearson Education, Inc. Fragmentation is breaking of the body into pieces, some or all of which develop into adults Fragmentation must be accompanied by regeneration, regrowth of lost body parts Parthenogenesis is the development of a new individual from an unfertilized egg © 2014 Pearson Education, Inc. Sexual Reproduction: An Evolutionary Enigma Sexual females have half as many daughters as asexual females; this is the “twofold cost” of sexual reproduction Despite this, almost all eukaryotic species reproduce sexually © 2014 Pearson Education, Inc. Figure 36.3-1 Sexual reproduction Asexual reproduction Female Generation 1 Female Generation 2 Male © 2014 Pearson Education, Inc. Figure 36.3-2 Sexual reproduction Asexual reproduction Female Generation 1 Female Generation 2 Male Generation 3 © 2014 Pearson Education, Inc. Figure 36.3-3 Sexual reproduction Asexual reproduction Female Generation 1 Female Generation 2 Male Generation 3 Generation 4 © 2014 Pearson Education, Inc. Sexual reproduction results in genetic recombination The resulting increased variation among offspring may enhance the reproductive success of parents in changing environments © 2014 Pearson Education, Inc. Reproductive Cycles Most animals exhibit reproductive cycles related to changing seasons Reproductive cycles are controlled by hormones and environmental cues Because seasonal temperature is often an important cue in reproduction, climate change can decrease reproductive success © 2014 Pearson Education, Inc. Figure 36.4 © 2014 Pearson Education, Inc. In some species of whiptail lizards, the members of breeding pairs alternate roles Reproduction is exclusively asexual, and there are no males However, courtship and mating behavior still take place, with one female of each pair mimicking male mating behavior The two alternate roles two or three times during the breeding season © 2014 Pearson Education, Inc. Hormone level Ovary size Figure 36.5 Estradiol Ovulation Ovulation Progesterone Behavior Time (a) A. uniparens females © 2014 Pearson Education, Inc. Female Male- Female like (b) The sexual behavior of A. uniparens is correlated with the cycle of ovulation. Malelike Figure 36.5a (a) A. uniparens females © 2014 Pearson Education, Inc. Hormone level Ovary size Figure 36.5b Estradiol Ovulation Ovulation Progesterone Behavior Time Female Male- Female like (b) The sexual behavior of A. uniparens is correlated with the cycle of ovulation. © 2014 Pearson Education, Inc. Malelike Variation in Patterns of Sexual Reproduction For many animals, finding a partner for sexual reproduction may be challenging One solution is hermaphroditism, in which each individual has male and female reproductive systems Any two hermaphrodites can mate, and some hermaphrodites can self-fertilize © 2014 Pearson Education, Inc. Individuals of some species undergo sex reversals For example, in a coral reef fish, the bluehead wrasse, a lone male defends a group of females If the male dies, the largest female in the group transforms into a male Within a few weeks, this individual can begin to produce sperm instead of eggs © 2014 Pearson Education, Inc. External and Internal Fertilization Sexual reproduction requires fertilization, the union of sperm and eggs In external fertilization, eggs shed by the female are fertilized by sperm in the external environment In internal fertilization, sperm are deposited in or near the female reproductive tract, and fertilization occurs within the tract © 2014 Pearson Education, Inc. A moist habitat is almost always required for external fertilization to prevent gametes from drying out and to allow sperm to swim to eggs Many aquatic invertebrates simply shed gametes into the surrounding water In this case, timing of release of gametes is crucial to ensure that sperm and egg encounter one another If external fertilization is not synchronous across a population, courtship behaviors might lead to fertilization © 2014 Pearson Education, Inc. Figure 36.6 © 2014 Pearson Education, Inc. However fertilization occurs, the mating animals may use pheromones, chemicals released by one organism that can influence physiology and behavior of another individual of the same species © 2014 Pearson Education, Inc. Ensuring the Survival of Offspring Internal fertilization is typically associated with production of fewer gametes but the survival of a higher fraction of zygotes Internal fertilization is also often associated with mechanisms to provide protection of embryos and parental care of young © 2014 Pearson Education, Inc. The embryos of some terrestrial animals develop in eggs with calcium- and protein-containing shells and several internal membranes Some other animals retain the embryo, which develops inside the female In many animals, parental care helps ensure survival of offspring © 2014 Pearson Education, Inc. Figure 36.7 © 2014 Pearson Education, Inc. Concept 36.2: Reproductive organs produce and transport gametes Sexual reproduction in animals relies on sets of cells that are precursors for eggs and sperm © 2014 Pearson Education, Inc. Variation in Reproductive Systems Many (but not all) animals have gonads, organs that produce gametes Some simple systems do not have gonads, but gametes form from undifferentiated tissue More elaborate systems include sets of accessory tubes and glands that carry, nourish, and protect gametes and sometimes developing embryos © 2014 Pearson Education, Inc. Most insects have separate sexes with complex reproductive systems In many insects, the female has a spermatheca in which sperm is stored during copulation © 2014 Pearson Education, Inc. A cloaca is a common opening between the external environment and the digestive, excretory, and reproductive systems A cloaca is common in nonmammalian vertebrates; mammals usually have a separate opening to the digestive tract © 2014 Pearson Education, Inc. Human Male Reproductive Anatomy The male’s external reproductive organs are the scrotum and penis The internal organs are gonads that produce sperm and reproductive hormones, accessory glands that secrete products essential to sperm movement, and ducts that carry sperm and glandular secretions Animation: Male Reproductive Anatomy © 2014 Pearson Education, Inc. Figure 36.8 (Urinary bladder) Seminal vesicle (Urinary duct) (Rectum) (Pubic bone) Vas deferens Ejaculatory duct Prostate gland Bulbourethral gland © 2014 Pearson Education, Inc. Erectile tissue Urethra Vas deferens Epididymis Testis Scrotum Glans Prepuce Penis Testes The male gonads, or testes, produce sperm in highly coiled tubes called seminiferous tubules Production of normal sperm cannot occur at the body temperatures of most mammals The scrotum, a fold of the body wall, maintains testis temperature at about 2oC below the core body temperature © 2014 Pearson Education, Inc. Ducts From the seminiferous tubules of a testis, sperm pass into the coiled duct of the epididymis During ejaculation, sperm are propelled through the muscular vas deferens and the ejaculatory duct and then exit the penis through the urethra © 2014 Pearson Education, Inc. Accessory Glands Semen is composed of sperm plus secretions from three sets of accessory glands The two seminal vesicles contribute about 60% of the total volume of semen The prostate gland secretes its products directly into the urethra through several small ducts The bulbourethral glands secrete a clear mucus before ejaculation that neutralizes acidic urine remaining in the urethra © 2014 Pearson Education, Inc. Penis The human penis is composed of three cylinders of spongy erectile tissue During sexual arousal, erectile tissue fills with blood from the arteries, causing an erection The head of the penis, or glans, has a thinner skin covering than the shaft The prepuce, or foreskin, is a fold of skin covering the glans; it is removed when a male is circumcised © 2014 Pearson Education, Inc. Human Female Reproductive Anatomy The female external reproductive structures include the clitoris and two sets of labia The internal organs are a pair of gonads and a system of ducts and chambers that carry gametes and house the embryo and fetus Animation: Female Reproductive Anatomy © 2014 Pearson Education, Inc. Figure 36.9 Oviduct Ovary Uterus (Urinary bladder) (Pubic bone) (Rectum) (Urethra) Cervix Vagina Body Clitoris Glans Prepuce Major vestibular gland Labia minora Vaginal opening © 2014 Pearson Education, Inc. Labia majora Ovaries The female gonads, a pair of ovaries, lie in the abdominal cavity Each ovary contains many follicles, which consist of a partially developed egg, called an oocyte, surrounded by support cells © 2014 Pearson Education, Inc. Oviducts and Uterus Upon ovulation, a mature egg cell is released It travels from the ovary to the uterus via an oviduct Cilia in the oviduct convey the egg to the uterus, also called the womb The uterus lining, the endometrium, has many blood vessels The neck of the uterus is the cervix, which opens into the vagina © 2014 Pearson Education, Inc. Vagina and Vulva The vagina is a muscular but elastic chamber that is the repository for sperm during copulation and serves as the birth canal The vagina opens to the outside at the vulva, which consists of the labia majora, labia minora, hymen, and clitoris © 2014 Pearson Education, Inc. Mammary Glands The mammary glands are not part of the reproductive system but are important to mammalian reproduction Within the glands, small sacs of epithelial tissue secrete milk © 2014 Pearson Education, Inc. Gametogenesis Gametogenesis is the production of gametes There is a close relationship between the structure and function of the gonads Video: Flagella in Sperm Video: Sperm Flagellum © 2014 Pearson Education, Inc. Figure 36.10a Epididymis Seminiferous tubule Testis Primordial germ cell in embryo Cross section of seminiferous tubule Mitotic divisions Spermatogonial stem cell Sertoli cell nucleus 2n Mitotic divisions Spermatogonium 2n Mitotic divisions Primary spermatocyte 2n Meiosis I Secondary spermatocyte Lumen of seminiferous tubule Tail Plasma membrane Neck Midpiece n Meiosis II Spermatids (two stages) Early spermatid n n n n Differentiation (Sertoli cells provide nutrients) Head Acrosome Nucleus Mitochondria © 2014 Pearson Education, Inc. n Sperm cell n n n n Figure 36.10aa Epididymis Seminiferous tubule Sertoli cell nucleus Testis Spermatogonium Primary spermatocyte Secondary spermatocyte Cross section of seminiferous tubule Spermatids (two stages) Sperm cell Lumen of seminiferous tubule © 2014 Pearson Education, Inc. Figure 36.10ab Primordial germ cell in embryo Mitotic divisions Spermatogonial stem cell 2n Mitotic divisions Spermatogonium 2n Mitotic divisions Primary spermatocyte 2n Meiosis I Secondary spermatocyte n n Meiosis II Early spermatid Sperm cell © 2014 Pearson Education, Inc. n n n n Differentiation (Sertoli cells provide nutrients) n n n n Figure 36.10ac Tail Plasma membrane Neck Midpiece Head Acrosome Nucleus Mitochondria © 2014 Pearson Education, Inc. Figure 36.10b Ovary Primary oocyte within follicle In embryo Primordial germ cell Mitotic divisions 2n Oogonium Growing follicle Mature follicle Mitotic divisions Primary oocyte (present at birth), arrested in prophase of meiosis I 2n Completion of meiosis I and onset of meiosis II First polar n body n Secondary oocyte, arrested at metaphase of meiosis II Ruptured follicle Ovulated secondary oocyte Ovulation, sperm entry Completion of meiosis II Second polar n body n © 2014 Pearson Education, Inc. Fertilized egg Corpus luteum Degenerating corpus luteum Figure 36.10ba Ovary Primary oocyte within follicle Growing follicle Ruptured follicle Ovulated secondary oocyte Mature follicle Corpus luteum Degenerating corpus luteum © 2014 Pearson Education, Inc. Figure 36.10bb In embryo Primordial germ cell Mitotic divisions 2n Oogonium Mitotic divisions 2n © 2014 Pearson Education, Inc. Primary oocyte (present at birth), arrested in prophase of meiosis I Figure 36.10bc Completion of meiosis I and onset of meiosis II First polar n body n Secondary oocyte, arrested at metaphase of meiosis II Ovulation, sperm entry Completion of meiosis II Second polar n body n © 2014 Pearson Education, Inc. Fertilized egg Spermatogenesis, the development of sperm, is continuous and prolific (millions of sperm are produced per day); each sperm takes about 7 weeks to develop Oogenesis, the development of a mature egg, is a prolonged process Immature eggs form in the female embryo but do not complete their development until years or decades later © 2014 Pearson Education, Inc. Spermatogenesis differs from oogenesis in three ways All four products of meiosis develop into sperm, while only one of the four becomes an egg Spermatogenesis occurs throughout adolescence and adulthood Sperm are produced continuously without the prolonged interruptions in oogenesis © 2014 Pearson Education, Inc. Concept 36.3: The interplay of tropic and sex hormones regulates reproduction in mammals Human reproduction is coordinated by hormones from the hypothalamus, anterior pituitary, and gonads Gonadotropin-releasing hormone (GnRH) is secreted by the hypothalamus and directs the release of FSH (follicle-stimulating hormone) and LH (luteinizing hormone) from the anterior pituitary © 2014 Pearson Education, Inc. FSH and LH regulate processes in the gonads and the production of sex hormones The major androgen is testosterone and major estrogens are estradiol and progesterone Both males and females produce androgens and estrogens but differ in their blood concentrations of particular hormones For example, males have testosterone levels about 10 times higher than females © 2014 Pearson Education, Inc. Sex hormones regulate gamete production both directly and indirectly They serve many additional functions including sexual behavior and the development of primary and secondary sex characteristics Androgens are responsible for male vocalizations and courtship displays © 2014 Pearson Education, Inc. Figure 36.11 © 2014 Pearson Education, Inc. Hormonal Control of the Male Reproductive System FSH and LH act on different cells in the testes to direct spermatogenesis Sertoli cells, found in the seminiferous tubules, respond to FSH by nourishing developing sperm Leydig cells, connective tissue between the tubules, respond to LH by secreting testosterone and other androgens, which promote spermatogenesis Animation: Male Hormones © 2014 Pearson Education, Inc. Figure 36.12 Hypothalamus GnRH FSH LH Leydig cells Sertoli cells Inhibin Spermatogenesis Testis © 2014 Pearson Education, Inc. Testosterone Negative feedback Negative feedback Anterior pituitary Two negative feedback mechanisms control sex hormone production in males Testosterone regulates the production of GnRH, FSH, and LH through negative feedback mechanisms Sertoli cells secrete the hormone inhibin, which reduces FSH secretion from the anterior pituitary © 2014 Pearson Education, Inc. Hormonal Control of Female Reproductive Cycles Females produce eggs in cycles Prior to ovulation, the endometrium thickens with blood vessels in preparation for embryo implantation If an embryo does not implant in the endometrium, the endometrium is shed in a process called menstruation © 2014 Pearson Education, Inc. Hormones closely link the two cycles of female reproduction Changes in the uterus define the menstrual cycle (also called the uterine cycle) Changes in the ovaries define the ovarian cycle Animation: Ovulation Animation: Post-Ovulation © 2014 Pearson Education, Inc. Figure 36.13 (a) Control by hypothalamus Inhibited by combination of estradiol and progesterone Hypothalamus Stimulated by high levels of estradiol GnRH 1 Anterior pituitary 2 FSH Inhibited by low levels of estradiol LH Pituitary gonadotropins in blood (b) 6 LH FSH FSH and LH stimulate follicle to grow 3 (c) Ovarian cycle 7 Growing follicle Maturing follicle 8 Follicular phase Corpus luteum Ovulation Degenerating corpus luteum Luteal phase Estradiol secreted by growing follicle in increasing amounts 4 (d) LH surge triggers ovulation Progesterone and estradiol secreted by corpus luteum Ovarian hormones in blood Peak causes LH surge (see 6 ) 5 10 9 Estradiol Progesterone Estradiol level very low (e) Progesterone and estradiol promote thickening of endometrium Uterine (menstrual) cycle Endometrium Days Menstrual flow phase © 2014 Pearson Education, Inc. 0 5 Proliferative phase 10 14 15 Secretory phase 20 25 28 Figure 36.13a (a) Control by hypothalamus Inhibited by combination of estradiol and progesterone Hypothalamus Stimulated by high levels of estradiol GnRH 1 Anterior pituitary 2 FSH Inhibited by low levels of estradiol LH Pituitary gonadotropins in blood (b) 6 LH FSH Days 3 0 © 2014 Pearson Education, Inc. FSH and LH stimulate follicle to grow 5 10 LH surge triggers ovulation 14 15 20 25 28 Figure 36.13b (b) Pituitary gonadotropins in blood 6 LH FSH 3 Days (c) FSH and LH stimulate follicle to grow Ovarian cycle 7 Growing follicle Maturing follicle LH surge triggers ovulation 8 Corpus luteum Follicular phase Ovulation Estradiol secreted 4 by growing follicle in increasing amounts 0 © 2014 Pearson Education, Inc. 5 10 14 15 Degenerating corpus luteum Luteal phase Progesterone and estradiol secreted by corpus luteum 20 25 28 Figure 36.13c (c) Ovarian cycle 7 Growing follicle Maturing follicle 8 Follicular phase Ovulation Ovarian hormones in blood 5 Degenerating corpus luteum Luteal phase Estradiol secreted by growing follicle in increasing amounts 4 (d) Corpus luteum Progesterone and estradiol secreted by corpus luteum Peak causes LH surge (see 6 ) 10 9 Estradiol Progesterone Progesterone and estradiol promote thickening of endometrium Days Estradiol level very low 0 © 2014 Pearson Education, Inc. 5 10 14 15 20 25 28 Figure 36.13d (d) Ovarian hormones in blood Peak causes LH surge (see 6 ) 5 10 9 Estradiol Progesterone Estradiol level very low (e) Progesterone and estradiol promote thickening of endometrium Uterine (menstrual) cycle Endometrium Days Menstrual flow phase 0 © 2014 Pearson Education, Inc. 5 Proliferative phase 10 14 15 Secretory phase 20 25 28 The Ovarian Cycle The sequential release of GnRH then FSH and LH stimulates follicle growth Follicle and oocyte growth and an increase in the hormone estradiol characterize the follicular phase of the ovarian cycle Estradiol causes GnRH, FSH, and LH levels to rise through positive feedback The final result is the maturation of the follicle © 2014 Pearson Education, Inc. At the end of the follicular phase, the follicle releases the secondary oocyte The follicular tissue left behind transforms into the corpus luteum in response to LH; this is the luteal phase The corpus luteum disintegrates, and ovarian steroid hormones decrease by negative feedback mechanisms © 2014 Pearson Education, Inc. The Uterine (Menstrual) Cycle Hormones coordinate the uterine cycle with the ovarian cycle Thickening of the endometrium during the proliferative phase coordinates with the follicular phase Secretion of nutrients during the secretory phase coordinates with the luteal phase Shedding of the endometrium during the menstrual flow phase coordinates with the growth of new ovarian follicles © 2014 Pearson Education, Inc. Menopause After about 500 cycles, human females undergo menopause, the cessation of ovulation and menstruation Menopause is very unusual among animals Menopause might have evolved to allow a mother to provide better care for her children and grandchildren © 2014 Pearson Education, Inc. Menstrual Versus Estrous Cycles Menstrual cycles are characteristic only of humans and some other primates The endometrium is shed from the uterus in a bleeding called menstruation Sexual receptivity is not limited to a time frame © 2014 Pearson Education, Inc. Estrous cycles are characteristic of most mammals The endometrium is reabsorbed by the uterus Sexual receptivity is limited to a “heat” period The length and frequency of estrus cycles vary from species to species © 2014 Pearson Education, Inc. Human Sexual Response Two reactions predominate in both sexes Vasocongestion, the filling of tissue with blood Myotonia, increased muscle tension The sexual response cycle has four phases: excitement, plateau, orgasm, and resolution Excitement prepares the penis and vagina for coitus (sexual intercourse) © 2014 Pearson Education, Inc. Direct stimulation of genitalia maintains the plateau phase and prepares the vagina for receipt of sperm Orgasm is characterized by rhythmic contractions of reproductive structures In males, semen is first released into the urethra and then ejaculated from the urethra In females, the uterus and outer vagina contract © 2014 Pearson Education, Inc. During the resolution phase, organs return to their normal state and muscles relax © 2014 Pearson Education, Inc. Concept 36.4: Fertilization, cleavage, and gastrulation initiate embryonic development Across animal species, embryonic development involves common stages occurring in a set order First is fertilization, which forms a zygote During the cleavage stage, a series of mitoses divide the zygote into a many-celled embryo The resulting blastula then undergoes rearrangements into a three-layered embryo called a gastrula © 2014 Pearson Education, Inc. Figure 36.14 EMBRYONIC DEVELOPMENT Sperm Zygote Adult frog Egg Blastula Metamorphosis Larval stages Gastrula Tail-bud embryo © 2014 Pearson Education, Inc. Fertilization Molecules and events at the egg surface play a crucial role in each step of fertilization Sperm penetrate the protective layer around the egg Receptors on the egg surface bind to molecules on the sperm surface Changes at the egg surface prevent polyspermy, the entry of multiple sperm nuclei into the egg © 2014 Pearson Education, Inc. Video: Cortical Granule Video: Calcium Release of Egg Video: Calcium Wave of Egg © 2014 Pearson Education, Inc. Figure 35.15-1 Basal body (centriole) Sperm head Acrosome Jelly coat Spermbinding receptors © 2014 Pearson Education, Inc. Vitelline layer Egg plasma membrane Figure 35.15-2 Basal body (centriole) Sperm head Acrosome Jelly coat Spermbinding receptors © 2014 Pearson Education, Inc. Hydrolytic enzymes Vitelline layer Egg plasma membrane Figure 35.15-3 Sperm nucleus Acrosomal process Basal body (centriole) Sperm head Acrosome Jelly coat Spermbinding receptors © 2014 Pearson Education, Inc. Actin filament Hydrolytic enzymes Vitelline layer Egg plasma membrane Figure 35.15-4 Sperm plasma membrane Sperm nucleus Acrosomal process Basal body (centriole) Sperm head Actin filament Fused plasma membranes Acrosome Jelly coat Spermbinding receptors © 2014 Pearson Education, Inc. Hydrolytic enzymes Vitelline layer Egg plasma membrane Figure 35.15-5 Sperm plasma membrane Sperm nucleus Acrosomal process Basal body (centriole) Sperm head Acrosome Jelly coat Spermbinding receptors © 2014 Pearson Education, Inc. Actin filament Cortical Fused granule plasma membranes Perivitelline Hydrolytic enzymes space Fertilization Vitelline layer envelope Egg plasma membrane The events of fertilization also initiate metabolic reactions that trigger the onset of embryonic development, thus “activating” the egg Activation leads to events such as increased protein synthesis that precede the formation of a diploid nucleus Sperm entry triggers a release of Ca2+, which activates the egg and triggers the cortical reaction, the slow block to polyspermy © 2014 Pearson Education, Inc. Figure 36.16 Seconds 1 2 3 4 6 8 10 20 30 40 50 1 Binding of sperm to egg Acrosomal reaction: plasma membrane depolarization (fast block to polyspermy) Increased intracellular calcium level Cortical reaction (slow block to polyspermy) Formation of fertilization envelope complete Minutes 2 3 4 5 Increased protein synthesis 10 20 30 40 60 90 © 2014 Pearson Education, Inc. Fusion of egg and sperm nuclei complete Onset of DNA synthesis First cell division Cleavage and Gastrulation Fertilization is followed by cleavage, a period of rapid cell division without growth Cleavage partitions the cytoplasm of one large cell into many smaller cells The blastula is a ball of cells with a fluid-filled cavity called a blastocoel The blastula is produced after about five to seven cleavage divisions Video: Cleavage of Egg © 2014 Pearson Education, Inc. Figure 36.17 50 m (a) Fertilized egg © 2014 Pearson Education, Inc. (b) Four-cell stage (c) Early blastula (d) Later blastula Figure 36.17a 50 m (a) Fertilized egg © 2014 Pearson Education, Inc. Figure 36.17b 50 m (b) Four-cell stage © 2014 Pearson Education, Inc. Figure 36.17c 50 m (c) Early blastula © 2014 Pearson Education, Inc. Figure 36.17d 50 m (d) Later blastula © 2014 Pearson Education, Inc. After cleavage, the rate of cell division slows The remaining stages of embryonic development are responsible for morphogenesis, the cellular and tissue-based processes by which the animal body takes shape © 2014 Pearson Education, Inc. During gastrulation, a set of cells at or near the surface of the blastula moves to an interior location, cell layers are established, and a primitive digestive tube forms The hollow blastula is reorganized into a two- or three-layered embryo called a gastrula © 2014 Pearson Education, Inc. The cell layers produced by gastrulation are called germ layers The ectoderm forms the outer layer and the endoderm the inner layer In vertebrates and other animals with bilateral symmetry, a third germ layer, the mesoderm, forms between the endoderm and ectoderm © 2014 Pearson Education, Inc. Figure 36.18 Animal pole Blastocoel Mesenchyme cells Vegetal plate Vegetal pole Blastocoel Filopodia Archenteron Mesenchyme cells Blastopore Key Future ectoderm Future mesoderm Future endoderm 50 m Blastocoel Ectoderm Mouth Mesenchyme (mesoderm forms future skeleton) © 2014 Pearson Education, Inc. Archenteron Blastopore Digestive tube (endoderm) Anus (from blastopore) Figure 36.18a-1 Animal pole Blastocoel Mesenchyme cells Vegetal plate Vegetal pole © 2014 Pearson Education, Inc. Future ectoderm Future mesoderm Future endoderm Figure 36.18a-2 Animal pole Blastocoel Mesenchyme cells Vegetal plate Vegetal pole Future ectoderm Future mesoderm Future endoderm Filopodia Archenteron © 2014 Pearson Education, Inc. Figure 36.18a-3 Animal pole Blastocoel Mesenchyme cells Vegetal plate Vegetal pole Future ectoderm Future mesoderm Future endoderm Filopodia Archenteron Blastocoel Archenteron Blastopore © 2014 Pearson Education, Inc. Figure 36.18a-4 Animal pole Blastocoel Mesenchyme cells Vegetal plate Vegetal pole Future ectoderm Future mesoderm Future endoderm Filopodia Archenteron Blastocoel Ectoderm Mouth Mesenchyme (mesoderm forms future skeleton) © 2014 Pearson Education, Inc. Archenteron Blastopore Digestive tube (endoderm) Anus (from blastopore) Figure 36.18b Blastocoel Filopodia Archenteron Mesenchyme cells Blastopore © 2014 Pearson Education, Inc. 50 m Cell movements and interactions that form the germ layers vary among species One distinction is whether the mouth develops at the first opening that forms in the embryo (protostomes) or the second (deuterostomes) Sea urchins and other echinoderms are deuterostomes, as are chordates © 2014 Pearson Education, Inc. Each germ layer contributes to a distinct set of structures in the adult animal Some organs and many organ systems derive from more than one germ layer © 2014 Pearson Education, Inc. Figure 36.19 ECTODERM (outer layer of embryo) • Epidermis of skin and its derivatives (including sweat glands, hair follicles) • Nervous and sensory systems • Pituitary gland, adrenal medulla • Jaws and teeth • Germ cells MESODERM (middle layer of embryo) • Skeletal and muscular systems • Circulatory and lymphatic systems • Excretory and reproductive systems (except germ cells) • Dermis of skin • Adrenal cortex ENDODERM (inner layer of embryo) • Epithelial lining of digestive tract and associated organs (liver, pancreas) • Epithelial lining of respiratory, excretory, and reproductive tracts and ducts • Thymus, thyroid, and parathyroid glands © 2014 Pearson Education, Inc. Human Conception, Embryonic Development, and Birth Conception, fertilization of an egg by a sperm, occurs in the oviduct The resulting zygote begins cleavage about 24 hours after fertilization and produces a blastocyst after 4 more days A few days later, the embryo implants into the endometrium of the uterus © 2014 Pearson Education, Inc. The condition of carrying one or more embryos in the uterus is called gestation, or pregnancy Human pregnancy averages 38 weeks from fertilization Gestation varies among mammals, from 21 days in many rodents to more than 600 days in elephants Human gestation is divided into three trimesters of about three months each © 2014 Pearson Education, Inc. Figure 36.20 3 Cleavage 4 Cleavage continues. Ovary 2 Fertilization Uterus 1 Ovulation 5 Implantation Endometrium (a) From ovulation to implantation Endometrium Inner cell mass Cavity Blastocyst (b) Implantation of blastocyst © 2014 Pearson Education, Inc. Trophoblast During the first trimester, the embryo secretes hormones that signal its presence and regulate the mother’s reproductive system Human chorionic gonadotropin (hCG) acts to maintain secretion of progesterone and estrogens by the corpus luteum through the first few months of pregnancy © 2014 Pearson Education, Inc. Occasionally an embryo splits during the first month of development, resulting in identical or monozygotic twins Fraternal, or dizygotic, twins arise when two eggs are fertilized and implant as two genetically distinct embryos Some embryos cannot complete development due to chromosomal or other abnormalities These spontaneously abort, often before a woman is aware of her pregnancy © 2014 Pearson Education, Inc. During its first 2 to 4 weeks, the embryo obtains nutrients directly from the endometrium Meanwhile, the outer layer of the blastocyst, called the trophoblast, mingles with the endometrium and eventually forms the placenta Materials diffuse between maternal and embryonic circulation to supply the embryo with nutrients, immune protection, and metabolic waste disposal © 2014 Pearson Education, Inc. The first trimester is the main period of organogenesis, development of the body organs All the major structures are present by 8 weeks, and the embryo is called a fetus Video: Ultrasound of Fetus © 2014 Pearson Education, Inc. Figure 36.21 (a) 5 weeks © 2014 Pearson Education, Inc. (b) 14 weeks (c) 20 weeks Figure 36.21a (a) 5 weeks © 2014 Pearson Education, Inc. Figure 36.21b (b) 14 weeks © 2014 Pearson Education, Inc. Figure 36.21c (c) 20 weeks © 2014 Pearson Education, Inc. During the second trimester the fetus grows and is very active The mother may feel fetal movements as early as one month into the second trimester © 2014 Pearson Education, Inc. During the third trimester, the fetus grows and fills the space within the embryonic membranes © 2014 Pearson Education, Inc. Childbirth begins with labor, a series of strong, rhythmic contractions that push the fetus and placenta out of the body Once labor begins, local regulators, prostaglandins and hormones, induce and regulate further contractions of the uterus © 2014 Pearson Education, Inc. Lactation, the production of milk, is an aspect of maternal care unique to mammals Suckling stimulates the release of prolactin, which in turn stimulates mammary glands to produce milk, and the secretion of oxytocin, which triggers milk release from the glands © 2014 Pearson Education, Inc. Contraception Contraception, the deliberate prevention of pregnancy, can be achieved in a number of ways Contraceptive methods fall into three categories Preventing development or release of eggs and sperm Keeping sperm and egg apart Preventing implantation of an embryo © 2014 Pearson Education, Inc. A health-care provider should be consulted for complete information on the choice and risks of contraception methods © 2014 Pearson Education, Inc. Figure 36.22 Male Method Female Event Event Method Production of Production of sperm primary oocytes Vasectomy Sperm transport Oocyte down male development duct system and ovulation Abstinence Condom Coitus interruptus (very high failure rate) Combination birth control pill (or injection, patch, or vaginal ring) Abstinence Female condom Sperm deposited in vagina Capture of the oocyte by the oviduct Tubal ligation Sperm movement through female reproductive tract Transport of oocyte in oviduct Spermicides; diaphragm; progestin alone (as minipill or injection) Meeting of sperm and oocyte in oviduct Union of sperm and egg Implantation of blastocyst in endometrium © 2014 Pearson Education, Inc. Morning-after pill; intrauterine device (IUD) Figure 36.22a Male Method Female Event Event Production of Production of sperm primary oocytes Vasectomy Oocyte Sperm transport development down male duct system and ovulation Abstinence Condom Coitus interruptus (very high failure rate) Method Combination birth control pill (or injection, patch, or vaginal ring) Abstinence Female condom Sperm deposited in vagina Capture of the oocyte by the oviduct Tubal ligation Spermicides; diaphragm; progestin alone (as minipill or injection) © 2014 Pearson Education, Inc. Figure 36.22b Male Method Female Event Sperm movement through female reproductive tract Event Transport of oocyte in oviduct Method Meeting of sperm and oocyte in oviduct Union of sperm and egg Implantation of blastocyst in endometrium © 2014 Pearson Education, Inc. Morning-after pill; intrauterine device (IUD) The rhythm method, or natural family planning, is temporary abstinence when conception is most likely; it has a pregnancy rate of 10–20% Coitus interruptus, the withdrawal of the penis before ejaculation, is unreliable Barrier methods block fertilization with a pregnancy rate of less than 10% A condom fits over the penis A diaphragm is inserted into the vagina before intercourse © 2014 Pearson Education, Inc. Intrauterine devices (IUDs) are inserted into the uterus and interfere with fertilization and implantation; the pregnancy rate is less than 1% Female birth control pills are hormonal contraceptives with a pregnancy rate of less than 1% © 2014 Pearson Education, Inc. Infertility and In Vitro Fertilization Infertility, the inability to conceive offspring, affects about one in ten couples in the United States and worldwide The causes are varied and equally likely to affect men and women Sexually transmitted diseases (STDs) are the most significant preventable causes of infertility © 2014 Pearson Education, Inc. Some forms of infertility are treatable Hormone therapy can sometimes increase sperm or egg production In vitro fertilization (IVF) mixes oocytes with sperm in culture dishes and returns the embryo to the uterus at the eight-cell stage Sperm may be injected directly into an oocyte as well © 2014 Pearson Education, Inc. Figure 36.UN01 © 2014 Pearson Education, Inc. Figure 36.UN02 Human gametogenesis Spermatogenesis Oogenesis Primary spermatocyte 2n Primary oocyte 2n Polar n body n Secondary spermatocytes n n n n n Spermatids n n n n Sperm n Secondary oocyte n Polar body n © 2014 Pearson Education, Inc. Fertilized egg