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4-27-05 Muscle and Development Diverse body movements require variation in muscle activity • An individual muscle cell either contracts completely or not all.(muscle fiber) • Individual muscles, composed of many individual muscle fibers, that can contract to varying degrees. – One way variation is accomplished is by varying the frequency of action potentials reaching the muscle from a single motor neuron. – Graded muscle contraction can also be controlled by regulating the number of motor units involved in the contraction. Fig. 49.38 Copyright © 2002 Pearson Education, Inc., publishing as Benjamin Cummings – Recruitment of motor neurons increases the number of muscle cells involved in a contraction. – Some muscles, such as those involved in posture, are always at least partially contracted. • Fatigue is avoided by rotating among motor units. Copyright © 2002 Pearson Education, Inc., publishing as Benjamin Cummings • Fast and Slow Muscle Fibers. – Fast muscle fibers are adapted for rapid, powerful contractions. • Fatigue relatively quickly. Copyright © 2002 Pearson Education, Inc., publishing as Benjamin Cummings – Slow muscle fibers are adapted for sustained contraction. • Relative to fast fibers, slow fibers have. – Less SR Ca2+ remains in the cytosol longer. – More mitochondria, a better blood supply, and myoglobin. Copyright © 2002 Pearson Education, Inc., publishing as Benjamin Cummings • Other Types of Muscle. – In addition to skeletal muscle, vertebrates have cardiac and smooth muscle. – Cardiac muscle: similar to skeletal muscle. • Intercalated discs facilitate the coordinated contraction of cardiac muscle cells. • Can generate there own action potentials (even single cells in cell culture). • Action potentials of long duration. Copyright © 2002 Pearson Education, Inc., publishing as Benjamin Cummings Fig. 40.4 Copyright © 2002 Pearson Education, Inc., publishing as Benjamin Cummings – Smooth muscle: lacks the striations seen in both skeletal and cardiac muscle. • Contracts with less tension, but over a greater range of lengths, than skeletal muscle. • No T tubules and no SR. • Ca2+ enters the cytosol from via the plasma membrane. • Slow contractions, with more control over contraction strength than with skeletal muscle. • Found lining the walls of hollow organs. Copyright © 2002 Pearson Education, Inc., publishing as Benjamin Cummings • Invertebrate muscle cells are similar to vertebrate skeletal and smooth muscle cells. Copyright © 2002 Pearson Education, Inc., publishing as Benjamin Cummings Relevant pages in Campbell for reproduction and development • 975-980 • 998-1010 CHAPTER 46 ANIMAL REPRODUCTION Overview of Animal Reproduction 1. Both asexual and sexual reproduction occur in the animal kingdom 2. Diverse mechanisms of asexual reproduction enable animals to produce identical offspring rapidly 3. Reproductive cycles and patterns vary extensively among mammals Copyright © 2002 Pearson Education, Inc., publishing as Benjamin Cummings Both asexual and sexual reproduction occur in the animal kingdom Asexual reproduction involves the formation of individuals whose genes all come from one parent. There is no fusion of sperm and egg. Sexual reproduction is the formation of offspring by the fusion of haploid gametes. Ovum: female gamete - usually large and nonmotile. Spermatozooan: male gamete - usually small and motile. (the Echinoderm, an urchin) Sexual reproduction increases genetic variation among offspring (allows for possible survival if environment changes). 2. Diverse mechanisms of asexual reproduction enable animals to produce identical offspring rapidly Invertebrates: Fission: asexual reproduction in which a parent separates into two or more approximately equal sized individuals. Budding: asexual reproduction in which new individuals split off from existing ones (phylum Cindaria (hydrozoans, hydra, budding of new polyps from colonial polyps). Gemmules of sponges are an example of a type of asexual reproduction that involves the release of specialized cells that can grow into new individuals. Fragmentation: the breaking of the body into several pieces, some or all of which develop into complete adults. Requires regeneration of lost body parts (associated with lower phyla, not possible in higher vertebrates. Advantages of asexual reproduction: Can reproduce without needing to find a mate Can have numerous offspring in a short period of time In stable environments, allows for the perpetuation of successful genotypes. Copyright © 2002 Pearson Education, Inc., publishing as Benjamin Cummings 3. Reproductive cycles and patterns vary extensively among animals Reproductive cycles are controlled by both environmental and hormonal cues. Animals may be solely asexual or sexual. Or they may alternate between the two modes depending on environmental conditions. Daphnia (an Arthropod crustacean) reproduce by parthenogenesis under favorable conditions abundant food, absence of predators and sexually during times of environmental stress. Copyright © 2002 Pearson Education, Inc., publishing as Benjamin Cummings Parthenogenesis is the process by which an unfertilized egg develops into (often) a haploid adult. Parthenogenesis plays a role in the social organization of species of bees, wasps, and ants. Male honeybees are haploid and female honeybees are diploid. Several genera of fishes, amphibians, and lizards produce by a form of parthenogenesis that produces diploid zygotes. Some lizards require the simulation of intercourse without fertilization to activate their eggs. Other member can be a female. Copyright © 2002 Pearson Education, Inc., publishing as Benjamin Cummings Mechanisms of Sexual Reproduction 1. Internal and external fertilization both depend on mechanisms ensuring that mature sperm encounter fertile eggs of the same species 2. Species with internal fertilization usually produce fewer zygotes but provide more parental protection than species with external fertilization 3. Complex reproductive systems have evolved in many animal phyla Copyright © 2002 Pearson Education, Inc., publishing as Benjamin Cummings 1. Internal and external fertilization both depend on mechanisms ensuring that mature sperm encounter fertile eggs of the same species Internal fertilization requires cooperative behavior that leads to copulation. Example of delayed implantation in seals and mustelids (weasels). Females congregate in a colony to have pups so are accessible to males at that time. Other times they are ranging far and wide for food. Copyright © 2002 Pearson Education, Inc., publishing as Benjamin Cummings External fertilization requires a moist habitat that will protect a developing egg from desiccation and heat stress Example: amphibians,annelids molluscs and fishes. – Specific mating behaviors assure that sperm and egg will be in the same place at the same time (Frogs calling in the spring). Fig. 46.4 Copyright © 2002 Pearson Education, Inc., publishing as Benjamin Cummings Pheromones: chemical signals released by one organism that influence the behavior of other individuals of the same species. In mammals signals the female is ready to mate and is ovulating a egg that can be fertilized. Many act as male attractants. Copyright © 2002 Pearson Education, Inc., publishing as Benjamin Cummings 2. Species with internal fertilization usually produce fewer zygotes but provide more parental protection than species with external fertilization Internal fertilization usually results in the production of fewer zygotes than does internal fertilization. However, the survival rate is lower for external fertilization than it is for internal fertilization. Fishes as an example. Less than 1% survive to establish a new cohort for that year. If conditions are very favorable more than 1% survive and when mature there is a bountiful harvest. Especially important for the fishing industry to understand reproductive strategies. Cod fisheries collapsed. Copyright © 2002 Pearson Education, Inc., publishing as Benjamin Cummings The externally fertilized eggs of fishes and amphibians are surrounded by a gelatinous coat. The internally fertilized amniote eggs of birds, reptiles, and monotremes are protected by calcium and protein shells and have large yolks as sources of building blocks (amino acid, lipids nucleic acids) and energy resources. In mammals the embryo is retained within the females reproductive tract and nourished there. Copyright © 2002 Pearson Education, Inc., publishing as Benjamin Cummings – Parental care of offspring may occur regardless of whether fertilization is external or internal. Example in this picture is a male water bug with fertilized eggs on its back. Fish, birds, some reptiles and mammals. Fig. 46.5 Copyright © 2002 Pearson Education, Inc., publishing as Benjamin Cummings 3. Complex reproductive systems have evolved in many animal phyla • Insects storing sperm and getting rid of it sometimes. • Delayed implantation. • Sequential hermaphrodites female first mal 2nd or vice versa. Large female can produce more eggs or large male can guard a harem. Copyright © 2002 Pearson Education, Inc., publishing as Benjamin Cummings Most insects have separate sexes with complex reproductive systems. In many species the female reproductive system includes a spermatheca, a sac in which sperm may be stored for a year or more. Fig. 46.7 Copyright © 2002 Pearson Education, Inc., publishing as Benjamin Cummings The basic plan of all vertebrate reproductive systems are very similar. However, there are variations. In many non-mammalian vertebrates the digestive, excretory, and reproductive systems share a common opening to the outside, the cloaca (birds,reptiles and monotremes egg laying mammals). Mammals have separate opening for the digestive and reproductive systems. Female mammals also have separate openings for the excretory and reproductive system.s. Copyright © 2002 Pearson Education, Inc., publishing as Benjamin Cummings CHAPTER 47 ANIMAL DEVELOPMENT The Stages of Early Embryonic Development 1. From egg to organism, an animal’s form develops gradually: the concept of epigenesis 2. Fertilization activates the egg and bring together the nuclei of sperm and egg 3. Cleavage partitions the zygote into many smaller cells Copyright © 2002 Pearson Education, Inc., publishing as Benjamin Cummings 1. From egg to organism, an animal’s form develops gradually: the concept of epigenesis • Preformation: the egg or sperm contains an embryo that is a preformed miniature adult. • Epigenesis: the form of an animal emerges from a relatively formless egg. Aristotle proposed that 2000 years ago, but preformation hypothesis only died a couple hundred years ago. • An organism’s development is primarily determined by the genome of the zygote and the organization of the egg cytoplasm. Figure 47.1 A “homunculus” inside the head of a human sperm 2. Fertilization activates the egg and bring together the nuclei of sperm and egg • Sea urchins are models for the study of the early development of deuterostomes(Echinoderms and Chordates). – Sea urchin eggs are fertilized externally. – Sea urchin eggs are surrounded by a jelly coat. Copyright © 2002 Pearson Education, Inc., publishing as Benjamin Cummings Inject with 0.5M KCl to Induce spawning Acrosomal tip binds vitellin membrane receptor Copyright © 2002 Pearson Education, Inc., publishing as Benjamin Cummings • The Acrosomal Reaction. – Acrosomal reaction: when exposed to the jelly coat the sperm’s acrosome discharges it contents by exocytosis. • Hydrolytic enzymes enable the acrosomal process to penetrate the egg’s jelly coat. • The tip of the acrosomal process adheres to the vitelline layer just external to the egg’s plasma membrane. Binds to a receptor on the vitelline layer. Copyright © 2002 Pearson Education, Inc., publishing as Benjamin Cummings – The sperm and egg plasma membranes fuse and a single sperm nucleus enter the egg’s cytoplasm. • Na+ channels in the egg’s plasma membrane opens. – Na+ flows into the egg and the membrane depolarizes: fast block to polyspermy. Copyright © 2002 Pearson Education, Inc., publishing as Benjamin Cummings • The Cortical Reaction. – Fusion of egg and sperm plasma membranes also triggers a signal-transduction pathway. • Ca2+ from the eggs ER is released into the cytosol and propagates as a wave across the fertilized egg inositol triphosphate (IP3) and diacylglycerol (DAG) are produced. – IP3 opens ligand-gated channels in the ER and the Ca2+ released stimulates the opening of other channels. Copyright © 2002 Pearson Education, Inc., publishing as Benjamin Cummings – High concentrations of Ca2+ cause cortical granules to fuse with the plasma membrane and release their contents into the perivitelline space. • The vitelline layer separates from the plasma membrane. • An osmotic gradient draws water into the perivitelline space, swelling it and pushing it away from the plasma membrane. • The vitelline layer hardens into the fertilization envelope: a component of the slow block to polyspermy. • The plasma membrane returns to normal and the fast block to polyspermy no longer functions. Copyright © 2002 Pearson Education, Inc., publishing as Benjamin Cummings • Activation of the Egg, – High concentrations of Ca2+ in the egg stimulates an increase in the rates of cellular respiration and proteins synthesis. – In sea urchins, DAG activates a protein that transports H+ out of the egg. • The reduced pH may be indirectly responsible for the egg’s metabolic responses to fertilization. Copyright © 2002 Pearson Education, Inc., publishing as Benjamin Cummings – In the meantime, back at the sperm nucleus... • The sperm nucleus swells and merges with the egg nucleus diploid nucleus of the zygote. – DNA synthesis begins and the first cell division occurs. Copyright © 2002 Pearson Education, Inc., publishing as Benjamin Cummings • Fertilization in Mammals (early steps same as urchin). • Capacitation, a function of the female reproductive system, enhances sperm function. – A capacitated sperm migrates through a layer of follicle cells before it reaches the zona pellucida. – Binding of the sperm cell induces an acrosomal reaction similar to that seen in the sea urchin. Copyright © 2002 Pearson Education, Inc., publishing as Benjamin Cummings Fig. 47.5 • Enzymes from the acrosome enable the sperm cell to penetrate the zona pellucida and fuse with the egg’s plasma membrane. – The entire sperm enters the egg. – The egg membrane depolarizes: functions as a fast block to polyspermy. – A cortical reaction occurs. • Enzymes from cortical granules catalyze alterations to the zona pellucida: functions as a slow block to polyspermy like in the urchin vitelline membrane. Copyright © 2002 Pearson Education, Inc., publishing as Benjamin Cummings – The envelopes of both the egg and sperm nuclei disperse. • The chromosomes from the two gametes share a common spindle apparatus during the first mitotic division of the zygote. Copyright © 2002 Pearson Education, Inc., publishing as Benjamin Cummings 3. Cleavage partitions the zygote into many smaller cells • Cleavage follows fertilization. (sea urchin) – The zygote is partitioned into blastomeres. • Each blastomere contains different regions of the undivided cytoplasm and thus different cytoplasmic determinants. Copyright © 2002 Pearson Education, Inc., publishing as Benjamin Cummings Fig. 47.6 – Except for mammals, most animals have both eggs and zygotes with a definite polarity. • Thus, the planes of division follow a specific pattern relative to the poles of the zygote. • Polarity is defined by the heterogeneous distribution of substances such as mRNA, proteins, and yolk. – Yolk is most concentrated at the vegetal pole and least concentrated at the animal pole. • In some animals, the animal pole defines the anterior end of the animal. Copyright © 2002 Pearson Education, Inc., publishing as Benjamin Cummings • A blastocoel forms within the morula blastula Frog Fig. 47.8d Copyright © 2002 Pearson Education, Inc., publishing as Benjamin Cummings • In birds the yolk is so plentiful that it restricts cleavage to the animal pole: meroblastic cleavage. • In animals with less yolk there is complete division of the egg: holoblastic cleavage. Copyright © 2002 Pearson Education, Inc., publishing as Benjamin Cummings Figure 47.6x Sea urchin development, from single cell to larva Pluteus Sea urchin gastrulation Gastrulation rearranges the embryonic blastula into a triploblastic gastrula with three embryonic germ layers that are the ectoderm endoderm and mesoderm Fig. 47.9 Copyright © 2002 Pearson Education, Inc., publishing as Benjamin Cummings • Sea urchin gastrulation. – Begins at the vegetal pole where individual cells enter the blastocoel as mesenchyme cells. • The remaining cells flatten and buckle inwards: invagination. – Cells rearrange to form the archenteron. • The open end, the blastopore, will become the anus. • An opening at the other end of the archenteron will form the mouth of the digestive tube. Copyright © 2002 Pearson Education, Inc., publishing as Benjamin Cummings 5. In organogenesis, the organs of the animal body form from the three embryonic germ layers • The derivatives of the ectoderm germ layer are: – – – – Epidermis of skin, and its derivatives Epithelial lining of the mouth and rectum. Cornea and lens of the eyes. The nervous system; adrenal medulla; tooth enamel; epithelium of the pineal and pituitary glands. Copyright © 2002 Pearson Education, Inc., publishing as Benjamin Cummings • The endoderm germ layer contributes to: – The epithelial lining of the digestive tract (except the mouth and rectum). – The epithelial lining of the respiratory system. – The pancreas; thyroid; parathyroids; thymus; the lining of the urethra, urinary bladder, and reproductive systems. Copyright © 2002 Pearson Education, Inc., publishing as Benjamin Cummings • Derivatives of the mesoderm germ layer are: – – – – – – The notochord. The skeletal and muscular systems. The circulatory and lymphatic systems. The excretory system. The reproductive system (except germ cells). And the dermis of skin; lining of the body cavity; and adrenal cortex. Copyright © 2002 Pearson Education, Inc., publishing as Benjamin Cummings Neural crest found only in vertebrates Dorsal Hollow nerve tube Fig. 47.11 Copyright © 2002 Pearson Education, Inc., publishing as Benjamin Cummings 6. Amniote embryos develop in a fluid-filled sac within a shell or uterus • The amniote embryo is the solution to reproduction in a dry environment. – Shelled eggs of reptiles and birds. – Uterus of placental mammals. Copyright © 2002 Pearson Education, Inc., publishing as Benjamin Cummings • Avian Development. • Cleavage is meroblastic, or incomplete. • Cell division is restricted to a small cap of cytoplasm at the animal pole. • Produces a blastodisc, which becomes arranged into the epiblast and hypoblast that bound the blastocoel, the avian version of a blastula. Fig, 47.12 (1) Copyright © 2002 Pearson Education, Inc., publishing as Benjamin Cummings • During gastrulation some cells of the epiblast migrate (arrows) towards the interior of the embryo through the primitive streak. • Some of these cells move laterally to form the mesoderm, while others move downward to form the endoderm. Fig, 47.12 (2) Copyright © 2002 Pearson Education, Inc., publishing as Benjamin Cummings • In early organogenesis the archentreron is formed as lateral folds pinch the embryo away from the yolk. • The yolk stalk (formed mostly by hypoblast cells) will keep the embryo attached to the yolk. • The notochord, neural tube, and somites form as they do in frogs. • The three germ layers and hypoblast cells contribute to the extraembyonic membrane system. Fig, 47.12 (3) Copyright © 2002 Pearson Education, Inc., publishing as Benjamin Cummings • The four extraembryonic membranes are the yolk sac, amnion, chorion, and allantois. – Cells of the yolk sac digest yolk providing nutrients to the embryo. – The amnion encloses the embryo in a fluid-filled amniotic sac which protects the embryo from drying out. – The chorion cushions the embryo against mechanical shocks. – The allantois functions as a disposal sac for uric acid. Copyright © 2002 Pearson Education, Inc., publishing as Benjamin Cummings Fig. 47.14 Copyright © 2002 Pearson Education, Inc., publishing as Benjamin Cummings