CHAPTER 32 INTRODUCTION TO ANIMAL EVOLUTION. Section A: What is an animal?

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1 CHAPTER 32 INTRODUCTION TO ANIMAL EVOLUTION Section A: What is an animal? 1. Structure, nutrition, and life history define animals 2. The animal kingdom probably evolved from a colonial, flagellated protist 3. Some animals can do some amazing things. And then this

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3 1. Structure, nutrition and life history define animals Five criteria, when taken together, create a reasonable definition. (1) Animals are all multicellular, all heterotrophic eukaryotes.

4 (2) Animal cells lack cell walls that provide structural supports for plants and fungi. The multicellular bodies of animals are held together with extracellular proteins, especially collagen. In addition, other structural proteins create several types of intercellular junctions, including tight junctions, desmosomes, and gap junctions, that hold tissues together. (3) Animals have two unique types of tissues: nervous tissue for impulse conduction and muscle tissue for movement.

5 (4) Most animals reproduce sexually, with the diploid stage usually dominating the life cycle. In most species, a small flagellated sperm fertilizes a larger, nonmotile egg. The zygote undergoes cleavage, a succession of mitotic cell divisions, leading to the formation of a multicellular animal. Fig. 32.1

6 (5. Animals have Hox genes. What are they, again? Many of these Hox genes contain common modules of DNA sequences, called homeoboxes. Only animals possess genes that are both homeoboxcontaining in structure and homeotic in function. All animals, from sponges to the most complex insects and vertebrates have Hox genes, with the number of Hox genes correlated with complexity of the animal s anatomy. OK, then, how do you explain this????

7 2. Animals evolved from protists Most systematists now agree that the animal kingdom is monophyletic, meaning all animals have one common ancestor. That ancestor was most likely a colonial flagellated protist that lived over 700 million years ago in the Precambrian era. Evidence for multi-cellularity evolving?

8 One hypothesis for the origin of animals from a flagellated protist suggests that a colony of identical cells evolved into a hollow sphere. The cells of this sphere then specialized, creating two or more layers of cells. Fig. 32.3

9 Introduction Trying to classify animals is a mess. Different comparisons, like DNA vs. embryological development, suggest different versions of how the history of life has played out. Luckily for you, you can stay clear of the mess and just learn a few basics about the different solutions to the challenges of survival the evolutionary process has produced. Let s look at some of the big events in the adaptive radiation of animals from their protist ancestor.

10 Fig. 32.4

11 Fig. 32.5

12 Linked with bilateral symmetry is cephalization, the concentration of sensory equipment on the anterior end. Cephalization also includes the development of a central nervous system concentrated in the head and extending toward the tail as a longitudinal nerve cord. Animals that move actively are bilateral, such that the head end is usually first to encounter food, danger, and other stimuli. Usually this helps keep the animal out of harm s way, but no system is perfect

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14 The basic organization of germ layers, concentric layers of embryonic tissue that form various tissues and organs, differs between radiata and bilateria. The radiata are said to be diploblastic because they have two germ layers. The ectoderm, covering the surface of the embryo, gives rise to the outer covering and the central nervous system. The endoderm, the innermost layer, lines the developing digestive tube, or archenteron, and gives rise to the lining of the digestive tract and the organs derived from it, such as the liver and lungs of vertebrates.

15 The bilateria are triploblastic. The third germ layer, the mesoderm lies between the endoderm and ectoderm. The mesoderm develops into the muscles and most other organs between the digestive tube and the outer covering of the animal.

16 (3) The Bilateria can be divided by the presence or absence of a body cavity (a fluid-filled space separating the digestive tract from the outer body wall) and by the structure of the body cavity. Acoelomates (the phylum Platyhelminthes) have a solid body and lack a body cavity. Fig. 32.6a

17 Coelomates are organisms with a true coelom, a fluid-filled body cavity completely lined by mesoderm. The inner and outer layers of tissue that surround the cavity connect dorsally and ventrally to form mesenteries, which suspend the internal organs. Fig. 32.6b

18 A body cavity has many adaptive functions. Its fluid cushions the internal organs, helping to prevent internal injury. The noncompressible fluid of the body cavity can function as a hydrostatic skeleton against which muscles can work. The presence of the cavity enables the internal organs to grow and move independently of the outer body wall.

19 Fig. 32.7

20 The third difference centers on the fate of the blastopore, the opening of the archenteron. In protostomes, the blastopore develops into the mouth and a second opening at the opposite end of the gastrula develops into the anus. Proto means First, and stome means the mouth. In deuterostomes, the blastopore usually develops into the anus and the mouth is derived from the secondary opening. Deutero means second. Let s watch gastrulation 1:00

21 1. Most animal phyla originated in a relatively brief span of geological time Data from molecular systematics suggest an animal origin about a billion years ago. Nearly all the major animal body plans appear in Cambrian rocks from 543 to 525 million years ago. During this relatively short time, a burst of animal origins, the Cambrian explosion, left a rich fossil assemblage. It includes the first animals with hard, mineralized skeletons

22 Much of the diversity in body form among animal phyla is associated with variations in the spatial and temporal expression of Hox genes within the embryo. A reasonable hypothesis is that the diversification of animals was associated with the evolution of the Hox regulatory genes, which led to variation in morphology during development. Biologists investigating evo-devo, the new synthesis of evolutionary biology and developmental biology, may provide insights into the Cambrian explosion.

23 Introduction All animals except sponges belong to the Eumetazoa, the animals with true tissues. The oldest eumetazoan clade is the Radiata, animals with radial symmetry and diploblastic (only ectoderm and endoderm) embryos. In fact, remember, this now is thought to be the oldest of all animal groups.

24 1. Phylum Cnidaria: Cnidarians have radial symmetry, a gastrovascular cavity, and cnidocytes The cnidarians (nigh-dare-ee-uns - hydras, jellies, sea anemones, and coral animals) have a relatively simple body construction. The basic cnidarian body plan is a sac with a central digestive compartment, the gastrovascular cavity.

25 This basic body plan has two variations: the sessile polyp and the floating medusa. The cylindrical polyps, such as hydras and sea anemones, adhere to the substratum by the aboral end and extend their tentacles, waiting for prey. Medusas (also called jellies) are flattened, mouth-down versions of polyps that move by drifting passively and by contacting their bell-shaped bodies. Fig. 33.4

26 Cnidarians are carnivores that use tentacles arranged in a ring around the mouth to capture prey and push the food into the gastrovascular chamber for digestion. Batteries of cells called cnidocytes on the tentacles defend the animal or capture prey. The cnidocytes have organelles called nematocysts evert a thread that can inject poison into the prey, or stick to or entangle the target. Watch 1:00

27 Fig. 33.5

28 Muscles and nerves exist in their simplest forms in cnidarians. Cells of the epidermis and gastrodermis have bundles of microfilaments arranged into contractile fibers. True muscle tissue appears first in triploblastic animals. When the animal closes its mouth, the gastrovascular cavity acts as a hydrostatic skeleton against which the contractile cells can work. Movements are controlled by a noncentralized nerve net associated with simple sensory receptors that are distributed radially around the body.

29 The three cnidarian classes show variations on the same body theme of polyp and medusa. Fig. 33.6

30 For one jelly, this is truly a cycle. See this amazing discovery:

31 Anemones and corals belong to the class Anthozoa. They occur only as polyps. Coral animals live as solitary or colonial forms and secrete a hard external skeleton of calcium carbonate. Each polyp generation builds on the skeletal remains of earlier generations to form skeletons that we call coral. In tropical seas, coral reefs provide habitat for a great diversity of invertebrates and fishes. Coral reefs in many parts of the world are currently being damaged by environmental changes - global warming is one suspect. Watch here 3:00 Great example of how environmental change can affect the distribution of organisms.

32 1. Phylum Platyhelminthes: Flatworms are acoelomates with gastrovascular cavities There are about 20,000 species of flatworms living in marine, freshwater, and damp terrestrial habitats. They also include many parasitic species, such as the flukes and tapeworms. Flatworms have thin bodies, ranging in size from nearly microscopic to tapeworms over 20 m long. Flatworms and other bilaterians are triploblastic, with a middle embryonic tissue layer, mesoderm, which contributes to more complex organs and organs systems and to true muscle tissue.

33 While flatworms are structurally more complex than cnidarians or ctenophores, they are simpler than other bilaterans. Like cnidarians and ctenophores, flatworms have a gastrovascular cavity with only one opening (and tapeworms lack a digestive system entirely and absorb nutrients across their body surface). Unlike other bilaterians, flatworms lack a coelom.

34 Planarians and other flatworms lack organs specialized for gas exchange and circulation. Their flat shape places all cells close to the surrounding water, and the digestive system is highly branched to distribute food throughout the animal. Nitrogenous wastes are removed by diffusion and simple ciliated flame cells help maintain osmotic balance. Fig

35 A planarian has a head with a pair of eyespots to detect light and lateral flaps that function mainly for smell. The planarian nervous system is more complex and centralized than the nerve net of cnidarians. Planarians can learn to modify their responses to stimuli.

36 Planarians can reproduce asexually through regeneration. The parent constricts in the middle, and each half regenerates the missing end. Watch :30 Planarians can also reproduce sexually. These hermaphrodites crossfertilize.

37 The blood fluke Schistosoma infects 200 million people, leading to body pains, anemia, and dysentery. Fig

38 Tapeworms (class Cestoidea) are also parasitic. The adults live mostly in vertebrates, including humans. Suckers and hooks on the head or scolex anchor the worm in the digestive tract of the host. A long series of proglottids, sacs of sex organs, lie posterior to the scolex. Tapeworms absorb food particles from their hosts; no need for a digestive system. Watch 2:00 Fig

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40 5. Phylum Mollusca The phylum Mollusca includes snails and slugs, oysters and clams, and octopuses and squids. Most mollusks are marine, though some inhabit fresh water, and some snails and slugs live on land. Mollusks are soft-bodied animals, but most are protected by a hard shell of calcium carbonate. Slugs, squids, and octopuses have reduced or lost their shells completely during their evolution.

41 Many mollusks feed by using a straplike rasping organ, a radula, to scrape up food. Watch :35 The visceral mass includes a complete, one-way digestive tract (gut) (alimentary canal), unlike sponges, cnidarians and platyhelminthes.

42 Most gastropods use their radula to graze on algae or plant material. Some species are predators. In these species, the radula is modified to bore holes in the shells of other organisms or to tear apart tough animal tissues. Seen shells with holes in them?? In the tropical marine cone snails, teeth on the radula form separate poison darts, which penetrate and stun their prey, including fishes. See here and here.

43 Lightning whelk eating a scallop

44 The class Bivalvia includes clams, oysters, mussels, and scallops. Bivalves have shells divided into two halves. The two parts are hinged at the mid-dorsal line, and powerful adductor muscles close the shell tightly to protect the animal. When the shell is open, the bivalve may extend its hatchet-shaped foot for digging or anchoring. Fig

45 Watch these scallops respond to an environmental stimulus predator juice!! Neat stuff from FSU Marine Lab.

46 Cephalopods use rapid movements to dart toward their prey which they capture with several long tentacles. Water from the excurrent siphon provides the propulsion. Squids and octopuses use beaklike jaws to bite their prey and then inject poison to immobilize the victim. A mantle covers the visceral mass, but the shell is reduced and internal in squids, missing in many octopuses, and exists externally only in nautiluses. Fig

47 Cephalopods have an active, predaceous lifestyle. Unique among mollusks, cephalopods have a closed circulatory system to facilitate the movements of gases, fuels, and wastes through the body. They have a well-developed nervous system with a complex brain and welldeveloped sense organs. This supports learning and complex behavior. Cool 1:00 and cooler 2:30

48 6. Phylum Annelida: Annelids are segmented worms All annelids ( little rings ) have segmented bodies. There are about 15,000 species ranging in length from less than 1 mm to 3 m for the giant Australian earthworm. Annelids live in the sea, most freshwater habitats, and damp soil. They are protostomes, like the mollusks.

49 The coelom of the earthworm, a typical annelid, is partitioned by septa, but the digestive tract, longitudinal blood vessels, and nerve cords penetrate the septa and run the animal s length. Fig

50 The digestive system consists of a pharynx, an esophagus, crop, gizzard, and intestine. The closed circulatory system carries blood with oxygen-carrying hemoglobin through dorsal and ventral vessels connected by segmental vessels. The dorsal vessel and five pairs of esophageal vessels act as muscular pumps to distribute blood. In each segment is a pair of excretory tubes, metanephridia, that remove wastes from the blood and coelomic fluid. Analagous to? Wastes are discharged through exterior pores.

51 A brainlike pair of cerebral ganglia lie above and in front of the pharynx. Earthworms are cross-fertilizing hermaphrodites. Two earthworms exchange sperm and then separate. The received sperm are stored while a special organ, the clitellum, secretes a mucus cocoon. As the cocoon slides along the body, it picks up eggs and stored sperm and slides off the body into the soil. Some earthworms can also reproduce asexually by fragmentation followed by regeneration.

52 The majority of leeches inhabit fresh water, but land leeches move through moist vegetation. Many leeches feed on other invertebrates, but some bloodsucking parasites feed by attaching temporarily to other animals, including humans.

53 Until this century, leeches were frequently used by physicians for bloodletting. Leeches are still used for treating bruised tissues and for stimulating the circulation of blood to fingers or toes that have been sewn back to hands or feet after accidents. Watch 2:45 Fig d

54 Introduction The primary evidence for defining the clade Ecdysozoa is data from molecular systematics. All members of this group share the phenomenon of ecdysis, the shedding of an exoskeleton outgrown by the animal. These are Nematodes and Arthropods

55 1. Phylum Nematoda Roundworms are found in most aquatic habitats, wet soil, moist tissues of plants, and the body fluids and tissues of animals. They range in length from less than 1 mm to more than a meter.

56 The cylindrical bodies of roundworms are covered with a tough exoskeleton, the cuticle. As the worm grows, it periodically sheds its old cuticle and secretes a new, larger one. Ecdysis. They have a complete digestive tract and use the fluid in their pseudocoelom to transport nutrients since they lack a circulatory system. Fig d

57 The nematodes also include many species that are important plant pests that attack roots. Other species parasitize animals. 2:10 Over 50 nematode species, including various pinworms and hookworms, parasitize humans. Trichinella spiralis causes trichinosis when the nematode worms encyst in a variety of human organs, including skeletal muscle. They are acquired by eating undercooked meat that has juvenile worms encysted in the muscle tissue. Fig b

58 2. Arthropods are segmented coelomates with exoskeletons and jointed appendages The world arthropod population has been estimated at a billion billion (10 18 ) individuals. Nearly a million arthropod species have been described - two out of every three organisms known are arthropods. This phylum is represented in nearly all habitats in the biosphere. On the criteria of species diversity, distribution, and sheer numbers, arthropods must be regarded as the most successful animal phylum.

59 The diversity and success of arthropods is largely due to three features: body segmentation, a hard exoskeleton, and jointed appendages. Groups of segments and their appendages have become specialized for a variety of functions, permitting efficient division of labor among regions. Fig

60 The body of an arthropod is completely covered by the cuticle, an exoskeleton constructed from layers of protein and chitin. The exoskeleton protects the animal and provides points of attachment for the muscles that move appendages. The exoskeleton of arthropods is strong and relatively impermeable to water. In order to grow, and mate, an arthropod must molt (ecdysis) its old exoskeleton and secrete a larger one, a process that leaves the animal temporarily vulnerable to predators and other dangers. Watch the blue crab 2:15

61 Arthropods have well-developed sense organs, including eyes for vision, olfactory receptors for smell, and antennae for touch and smell. Most sense organs are located at the anterior end of the animal, showing extensive cephalization. Arthropods have an open circulatory system in which hemolymph fluid is propelled by a heart through short arteries into sinuses (the hemocoel) surrounding tissues and organs. Hemolymph returns to the heart through valved pores. The true coelom is much reduced in most species.

62 The move onto land by several groups of arthropods (insects, millipedes, centipedes, some chelicerates, and few crustaceans) was made possible, in part, by the exoskeleton. While it initially evolved for protection and locomotion, on land the exoskeleton also solved problems of water loss because the cuticle is relatively impermeable to water, helping prevent desiccation. The firm exoskeleton also provided support when arthropods left the relative buoyancy of water.

63 Several hypotheses have been proposed for the evolution of wings. In one hypothesis, wings first evolved as extensions of the cuticle that helped the insect absorb heat and were later modified for flight. A second hypothesis argues that wings allowed animals to glide from vegetation to the ground. Alternatively, wings may have served as gills in aquatic insects. Still another hypothesis proposes that insect wings functioned for swimming before they functioned for flight.

64 The internal anatomy of an insect includes several complex organ systems. Metabolic wastes are removed from the hemolymph by Malpighian tubules, outpockets of the digestive tract. Respiration is accomplished by a branched, chitin-lined tracheal system that carries O 2 from the spiracles (openings) directly to the cells. Let s watch the Insect DVD. Regulating the spiracles (similar to how a plant does its stomata) helps control breathing and drying out.

65 Fig

66 Metamorphosis is central to insect development. In incomplete metamorphosis (seen in grasshoppers and some other orders), the young resemble adults but are smaller and have different body proportions. Through a series of molts, the young look more and more like adults until it reaches full size. In complete metamorphosis, larval stages specialized for eating and growing change morphology completely during the pupal stage and emerge as adults. Fig

67 Introduction At first glance, sea stars and other echinoderms would seem to have little in common with the phylum Chordata, which includes the vertebrates. However, these animals share the deuterostome characteristics of radial cleavage, development of the coelom from the archenteron, and the formation of the anus from the blastopore. These developmental features that define the Deuterostomia are supported by molecular systematics.

68 1. Phylum Echinodermata: Echinoderms have a water vascular system and secondary radial symmetry Sea stars and most other echinoderms are sessile, or slow-moving animals with pentaradial symmetry. The internal and external parts of the animal radiate from the center, often as five spokes. A thin skin covers an endoskeleton of hard calcareous plates. Most echinoderms are prickly from skeletal bumps and spines that have various functions.

69 Unique to echinoderms is the water vascular system, a network of hydraulic canals branching into extensions called tube feet. These function in locomotion, feeding, and gas exchange.

70 Sea stars (class Asteroidea) have five arms (sometimes more) radiating from a central disk. The undersides of arms have rows of tube feet. Each can act like a suction disk that is controlled by hydraulic and muscular action. Fig

71 Sea stars and some other echinoderms can regenerate lost arms and, in a few cases, even regrow an entire body from a single arm. Fig a

72 And so how about a tribute To David Attenborough

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