CHAPTER 17 Darwin and Evolution History of Evolutionary Thought a. Pre-Darwinian world-view was determined by intractable theological beliefs.

1) The earth is young. 2) Each species was specially created and did not change over time. 3) Variations are imperfections varying from a perfectly-adapted creation. 4) Observations are to substantiate the prevailing worldview. A. Mid-Eighteenth-Century Contributions 1. Carolus Linnaeus and Taxonomy a. Taxonomy is the science of classifying organisms; taxonomy had been a main concern of biology. b. Carolus Linnaeus was a Swedish taxonomist. 1) Linnaeus developed a binomial system of nomenclature (two-part names for each species [Homo sapiens]). 2) Like other taxonomists of his time, Linnaeus believed in the ideas of a) special creation-each species had an "ideal" structure and function; and b) fixity of species-each species had a place in the scala naturae, a ladder of life. B. Late Eighteenth-/Early-Nineteenth Century Contributions 1. Cuvier and Catastrophism a. George Cuvier (1769-1832), a French zoologist, was the first to use comparative anatomy to develop a system of classifying animals. b. He founded paleontology-the study of fossils-and said that a single fossil bone was all he needed to deduce the anatomy of an animal. c. To explain the fossil record, Cuvier proposed that a whole series of catastrophes and repopulations from other regions had occurred. d. Catastrophism is Cuvier's explanation of fossil history: the belief that catastrophic extinctions occurred, after which repopulation of surviving species occurred, giving an appearance of change through time. 2. Lamarck's Acquired Characteristics a. Lamarck was the first to state that descent with modification occurs and that organisms become adapted to their environments. b. Lamarck, an invertebrate zoologist, held ideas at odds with Cuvier's. c. Lamarck mistakenly saw "a desire for perfection" as inherent in all living things. d. Inheritance of acquired characteristics was belief that organisms adapt to their environment in their life and pass these adaptations on Darwin's Theory of Evolution B. Geology and Fossils 1. Observed massive geological changes were caused by slow, continuous processes. In contrast to catastrophists, Hutton and Lyell proposed that the earth was subject to slow but continuous geological processes that occur at a uniform rate, a theory called uniformitarianism. C. Biogeography 1. Biogeography is the study of the geographic distribution of life forms on earth. 2. Comparison of the animals of South America and the Galpagos Islands caused Darwin to conclude that adaptation to the environment can cause diversification, including origin of new species. 3. The Galpagos Islands b. Island species varied from the mainland species, and from island-to-island. c. Each island had a variation of tortoise; long and short necked tortoises correlated with different vegetation. d. Darwin's Finches 1) Finches on the Galpagos Islands resembled a mainland finch but there were more types. 2) The variation in finches posed questions to Darwin: did they descend from one mainland ancestor or did islands allow isolated populations to evolve independently, and could present-day species have resulted from changes occurring in each isolated population? D. Natural Selection and Adaptation 1. Darwin decided that adaptations develop over time; he sought a mechanism by which adaptations might arise. 2. Natural selection was proposed by both Alfred Russel Wallace and Darwin as a driving mechanism of evolution caused by environmental selection of organisms most fit to reproduce, resulting in adaptation. 3. Because the environment is always changing, there is no perfectly-adapted organism. 4. There are three preconditions for natural selection. a. The members of a population have random but heritable variations. b. In a population, many more individuals are produced each generation than the environment can support. c. Some individuals have adaptive characteristics that enable them to survive and reproduce better. 5. There are two consequences of natural selection. a. An increasing proportion of individuals in succeeding generations will have the adaptive characteristics. b. The result of natural selection is a population adapted to its local environment. E. Organisms Have Variations 1. In contrast to the previous worldview where imperfections were to be ignored, variations were essential in natural selection. 2. Darwin suspected, but did not have today's evidence, that the occurrence of variation is completely random. 3. Variations that make adaptation possible are those that are passed on from generation to generation. F. Organisms Struggle to Exist 1. Malthus proposed that human populations outgrow food supply and death and famine were inevitable. 2. Darwin applied this to all organisms; resources were not sufficient for all members to survive. 3. Therefore, there is a constant struggle for existence; only certain members survive and reproduce.

G. Organisms Differ in Fitness 1. Organisms whose traits enable them to reproduce to a greater degree have a greater fitness. b. Many crop plant varieties can be traced to a single ancestor. d. Evolution by artificial or natural selection occurs when more fit organisms reproduce and leave more offspring H. Organisms Become Adapted 1. An adaptation is a trait that helps an organism be more suited to its environment. 2. Unrelated organisms living in the same environment often display similar characteristics. 3. Because of differential reproduction, adaptive traits increase in each succeeding generation. 17.3 The Evidence of Evolution A. Fossils Evidence 1. The fossil record is the history of life recorded by remains from the past. Transitional forms reveal links between groups. 1. Biogeography studies the distribution of plants and animals worldwide. 2. Distribution of organisms is explained by related forms evolving in one locale and spreading to other accessible areas. 3. Physical factors, such as the location of continents, determine where a population can spread. B. Anatomical Evidence 1. Organisms have anatomical similarities when they are closely related because of common descent. a. Homologous structures in different organisms are inherited from a common ancestor. b. Analogous structures are inherited from unique ancestors and have come to resemble each other because they serve a similar function. 2. Vestigial structures are remains of a structure that was functional in some ancestors but is no longer functional in the organism in question. 3. Embryological development C. Biochemical Evidence 1. Almost all living organisms use the same basic biochemical molecules, DNA, ATP, enzymes. 2. Organisms utilize the same DNA triplet code and the same 20 amino acids in their proteins. D. Because it is supported by so many lines of evidence, evolution is no longer considered a hypothesis. 1. Evolution is one of the great unifying theories of biology, similar in status to the germ theory of disease in medicine. 2. In science, a theory is supported by a large number of observations or a large amount of experimental evidence E. Pace of Evolution 1. Phyletic gradualism - slow process with many transitional forms 2. Punctuated equilibrium - speciation occurs rapidly, transitional links not evident, explains lack of fossils- Living fossils supports this

CHAPTER 18 PROCESS OF EVOLUTION

18.1 Microevolution 2. A population is all of the members of a single species occupying a certain area at the same time. 3. Evolution that occurs within a population is called microevolution. 4. Population genetics studies the variation in alleles in a gene pool. 5. The gene pool is the total of all the alleles in a population; it is described in terms of gene frequencies. 6. Industrial Melanism a. The case of the peppered moths provides a case study in a shift in phenotype frequencies under selection. b. With birds acting as a selective agent, dark-colored moths were better adapted to survive on the darkened trees A. Causes of Microevolution 1. Genetic Mutations a. Many traits in organisms are polymorphic, i.e., two or more distinct phenotypes are present in the population due to mutated genes. b. In humans, freckles are an example of polymorphism, as are the ABO blood types. c. Mutations can be beneficial, neutral, or harmful. Gene Flow 1. Gene flow (gene migration) is the movement of alleles among populations by migration of breeding individuals. 2. Gene flow can increase variation within a population by introducing novel alleles, but continued decreases diversity, causing similar gene pools 3. Gene flow among populations can prevent speciation from occurring. C. Nonrandom Mating 1. Assortative mating occurs when individuals mate with those that have the same phenotype. 2. Sexual selection occurs when males compete for the right to reproduce and the female selects males of a particular phenotype. (guppies, lions) D. Genetic Drift 1. Genetic drift refers to changes in allele frequencies of a gene pool due to chance, more often in small populations 2. Genetic drift occurs when founders start a new population, or after a genetic bottleneck with interbreeding. 1) The bottleneck effect is caused by a reduction in population size due to a natural disaster, predation, or habitat reduction= reduce diversity b. The founder effect is a genetic drift where rare alleles occur in higher frequency in a population isolated from the general population. 1) This is due to founding individuals containing a fraction of total genetic diversity of the original population. 18.2 Natural Selection - the process that results in adaptation of a population to the environment. 1. Natural selection requires a. variation (the members of a population differ from one another), b. inheritance (many of the differences between individuals in a population are heritable genetic differences), c. differential adaptedness (some differences affect how well an organism is adapted to its environment), and d. differential reproduction (better adapted individuals are more likely to reproduce). 2. Fitness is the extent to which an individual contributes fertile offspring to the next generation. A. Types of Selection. 1. Directional selection occurs when an extreme phenotype is favored; the distribution curve shifts that direction.

2. Stabilizing selection occurs when extreme phenotypes are eliminated and the intermediate phenotype is favored. 3. Disruptive selection occurs when extreme phenotypes are favored and can lead to more than one distinct form. B. Maintenance of Variations 1. Populations always show some genotypic variation; populations that lack variation may not be able to adapt to new conditions. 2. The following forces promote genetic variation. a. Mutation creates new alleles and genetic recombination still combines these alleles. b. Gene flow among small populations introduces new alleles. 18.3 Macroevolution Macroevolution refers to any evolutionary change at or above the species level. Speciation is the splitting of one species into two or more species or the transformation of one species into a new species over time; speciation is the final result of changes in gene pool allele and genotypic frequencies. A. What is a Species? 1. Linnaeus separated species based on morphology; Darwin saw that similar species are related by common descent. 2. biological species concept: a species is a group of potentially interbreeding populations that are reproductively isolated from other such groups. B. Reproductive Isolating Mechanisms 1. For two species to be separate, gene flow must not occur between them. 2. A reproductive isolating mechanism is any structural, functional, or behavioral characteristic that prevents reproduction from occurring. 3. Prezygotic isolating mechanisms are anatomical or behavioral differences between the members of two species that prevent mating or make it unlikely fertilization will take place if mating occurs. Occurs after postzygotic isolation. a. Habitat isolation occurs when two species occupy different habitats, even within the same geographic range b. Temporal isolation occurs when two species live in the same location, but each reproduces at a different time of year c. Behavioral isolation results from differences in mating behavior between two species. d. Mechanical isolation is the result of differences between two species in reproductive structures or other body parts e. Gamete isolation includes incompatibility of gametes of two different species so they cannot fuse to form a zygote 4. Postzygotic isolating mechanisms prevent development of a hybrid after mating has taken place. First to occur a. Zygote mortality is when hybrids (offspring of parents of two different species) do not live to reproduce. b. Hybrid sterility occurs when the hybrid offspring are sterile (mules). c. In F2 fitness, the offspring are fertile but the F2 generation is sterile. C. Modes of Speciation 1. Allopatric speciation occurs when new species result from populations being separated by a geographical barrier that prevents their members from reproducing with each other. a. While geographically isolated, variations accumulate until the populations are reproductively isolated. 2. Sympatric speciation would occur when members of a single population develop a genetic difference (e.g., chromosome number) that prevents them from reproducing with the parent type. D. Adaptive Radiation 1. Adaptive radiation is a rapid development from a single ancestral species of many new species. 2. The case of Darwin's finches illustrates the adaptive radiation of 13 species from one founder mainland finch.

Chapter Outline............................19.1 Origin of Life

A. The Early Earth 1. The Earth came into being about 4.6 BYA (BYA). 2. Heat from gravitation and radioactivity formed the Earth in several layers with iron and nickel in a liquid core, silicate minerals in a semi-liquid mantle, and upwellings of volcanic lava forming the first crust. 3. The Earth's mass provides a gravitational field strong enough to hold an atmosphere. 4. Early Earth's atmosphere differed from the current atmosphere, consisting of: water vapor, nitrogen, CO2, small amt of H, methane, ammonia,CO 5. The early atmosphere was formed by volcanic out-gassing characteristic of the young Earth. 6. The early atmosphere contained little free oxygen and was probably a reducing atmosphere=allows formation of complex organic molecules. B. Monomers Evolve 3. Oparin/Haldane Hypothesis (1920s) a. Oparin/Haldane independently suggested organic molecules could be formed in the presence of outside energy sources using atmospheric gases. b. Experiments performed by Miller and Urey (1953) showed experimentally that these gases (methane, ammonia, hydrogen, water) reacted with one another to produce small organic molecules (amino acids, organic acids). 4. Lack of oxidation and decay allowed organic molecules to form a thick, warm organic soup. C. Polymers Evolve -Newly formed organic molecules polymerized to produce larger molecules. 2. Protein-first Hypothesis a. Sidney Fox demonstrated amino acids polymerize abiotically if exposed to dry heat. b. Amino acids collected in shallow puddles along the rocky shore; heat of the sun caused them to form proteinoids c. When proteinoids are returned to water, they form cell-like microspheres composed of protein. d. This assumes DNA genes came after protein enzymes; DNA replication needs protein enzymes. 3. The Clay Hypothesis a. Graham Cairns-Smith suggests that amino acids polymerize in clay, with radioactivity providing energy. b. Clay attracts small organic molecules and contains iron and zinc atoms serving as inorganic catalysts for polypeptide formation. c. Clay collects energy from radioactive decay and discharges it if temperature or humidity changes. d. If RNA nucleotides and amino acids became associated so polypeptides were ordered by and helped synthesize RNA, then polypeptides and RNA arose at the same time.

4. RNA-first Hypothesis a. Only the macromolecule RNA was needed at the beginning to lead to the first cell. b. Thomas Cech and Sidney Altman discovered that RNA can be both a substrate and an enzyme. c. RNA would carry out processes of life associated with DNA (in genes) and protein enzymes. D. A Protocell Evolves 1. Before the first true cell arose, there would have been a protocell or protobiont. 2. A protocell would have a lipid-protein membrane and carry on energy metabolism. 3. Fox showed that if lipids are made available to microspheres, lipids become associated with microspheres producing a lipid-protein membrane. 4. Oparin demonstrated a protocell could have developed from coacervate droplets. a. Coacervate droplets are complex spherical units that spontaneously form when concentrated mixtures of macromolecules are held in the right temperature, ionic composition, and pH. b. Coacervate droplets absorb and incorporate various substances from the surrounding solution. c. In a liquid environment, phospholipid molecules spontaneously form liposomes, spheres surrounded by a layer of phospholipids; this is called the "membrane-first" hypothesis. d. A protocell could have contained only RNA to function as both genetic material and enzymes. 5. If a protocell was a heterotrophic fermenter living on the organic molecules in the organic soup that was its environment, this would indicate heterotrophs preceded autotrophs. 6. If the protocell evolved at hydrothermal vents, it would be chemosynthetic and autotrophs would have preceded heterotrophs. 7. The first protocells may have used preformed ATP, but as supplies dwindled, natural selection would favor cells that could extract energy from carbohydrates to transform ADP to ATP. 8. Since glycolysis is a common metabolic pathway in living things, it evolved early in the history of life. 9. As there was no free O2, it is assumed that protocells carried on a form of fermentation. 10. The first protocells had a limited ability to break down organic molecules 11. Fox has shown that a microsphere has some catalytic ability; Oparin found that coacervates incorporate enzymes if they are available in the medium. E. A Self-Replication System Evolves b. Ribozymes are RNA that acts as enzymes. c. Some viruses contain RNA genes with a protein enzyme called reverse transcriptase that uses RNA as a template to form DNA; this could have given rise to the first DNA. 3. The protein-first hypothesis contends that proteins or at least polypeptides were the first to arise. a. Only after the protocell develops complex enzymes could it form nucleic acids from small molecules. b. Because a nucleic acid is complicated, the chance that it arose on its own is minimal. c. Therefore, enzymes are needed to guide the synthesis of nucleotides and then nucleic acids. 4. Cairns-Smith suggests that polypeptides and RNA evolved simultaneously. a. The first true cell would contain RNA genes that replicated because of the presence of proteins; they become associated in clay in such a way that the polypeptides catalyzed RNA formation; both events happen at the same time. 5. Once the protocell was capable of reproduction, it became a true cell and biological evolution began. a. After DNA formed, the genetic code still had to evolve to store information. b. Because the current code is subject to fewer errors than other possible codes, and because it minimizes mutations, it likely underwent a natural selection process. a. Abiotic synthesis of organic molecules such as amino acids occurred in the atmosphere or at hydrothermal vents. b. Monomers joined together to form polymers at seaside rocks or clay, or at vents; the first polymers could have been proteins or RNA or both. c. Polymers aggregated inside a plasma membrane to make a protocell that had limited ability to grow; if it developed in the ocean it was a heterotroph, if at a hydrothermal vent, a chemoautotroph. d. Once the protocell contained DNA genes or RNA molecules, it was a true cell. 19.2 History of Life A. Fossils Tell a Story 1. A fossil is the remains or traces of past life, usually preserved in sedimentary rock. 4. Sedimentation has been going on since the Earth was formed; it is an accumulation of particles forming a stratum, a recognizable layer in a stratigraphic sequence laid down on land or in water. 5. The sequence indicates the age of fossils; a stratum is older than the one above it and younger than the one below it. B. Relative Dating of Fossils 1. Strata of the same age in England and Russia may have different sediments. 2. geologists discovered that strata of the same age contain the same fossils, or index fossils. fossils can be used for the relative dating of strata. 4. A particular species of fossil ammonite is found over a wide range and for a limited time period; therefore, all strata in the world that contain this ammonite are of the same age. C. Absolute Dating of Fossils 1. Absolute dating relies on radioactive dating to determine the actual age of fossils. 2. Radioactive isotopes have a half-life, the time it takes for half of a radioactive isotope to change into a stable element. 3. Carbon 14 (14C) is a radioactive isotope contained within organic matter. a. Half of the carbon 14 (14C) will change to nitrogen 14 (14N) every 5,730 years. b. Comparing 14C radioactivity of a fossil to modern organic matter calculates the age of the fossil. c. After 50,000 years, the 14C radioactivity is so low it cannot be used to measure age accurately. 4. Also potassium 40 (40K) and argon 40 to date rocks and infer the age of a fossil.