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Ch.16 Evolution of Populations How Common is Genetic Variation? • Genetic variation is studied within populations. • Because members within populations interbreed, they share a common group of genes called a gene pool. • A gene pool is made up of all of the genes (including all of the alleles) that are present in a population. • The relative frequency of an allele is the number of times that allele shows up, compared to the number of times other alleles for that same gene occur. • In genetic terms, evolution is any change in the relative frequency of alleles in a population. • (change from 10-20% allele frequency= evolution) Evolution Versus Genetic Equilibrium • Genetic Equilibrium is when allele frequencies remain constant (there is no evolution or change in the population) • The Hardy –Weinberg Principle states that allele frequencies in a population will remain constant as long as the following five conditions are met: • • • • • Random mating- no preference in mate selection Large population size- so that small changes will not be significant No migration- no gene flow: no new alleles brought into the population No mutations- no new alleles added to the population No Natural selection- all organisms are reproductively successful therefore no genes are favored • If any of the above are not met, then the population will evolve The Process of Speciation • Recall that a species is defined as a group of organisms that look alike and can interbreed to produce fertile offspring in nature. • The evolution of new species is called speciation. • Reproductive isolation occurs when formerly interbreeding organisms can no longer mate and produce fertile offspring. • There are three types of reproductive isolation: behavioral, temporal, and geographic. Types of Reproductive Isolation Behavioral Isolation • Two organisms are capable of interbreeding, but they have different courtship rituals. Ex: different mating songs of birds eastern and western meadowlarks Temporal Isolation • Two or more species reproduce at different times. Ex: Pollen released at different times from orchids. Geographic Isolation • Geographic isolation occurs whenever a physical barrier divides a population . Ex: tree frogs Geographic Isolation • When geographic isolation divides a population of tree frogs, the individuals no longer mate across populations. • Tree frogs are a single population. • The formation of a river may divide the frogs into two populations. • Over time, the divided populations may become two species that may no longer interbreed, even if reunited. Reproductive Isolation Flowchart Reproductive Isolation results from Isolating mechanisms which include Behavioral isolation Geographic isolation Temporal isolation produced by produced by produced by Behavioral differences Physical separation Different mating times which result in Independently evolving populations which result in Formation of new species Testing Natural Selection in Nature • • Peter and Rosemary Grant have worked for the past twenty years to show that Darwin’s hypothesis is correct. They realized that Darwin relied on two assumptions: 1. In order for beak size and shape to evolve in finches, there must have been many varieties in those traits for natural selection to work on. 2. Differences in beak size and shape must produce differences in fitness that cause natural selection to occur. Variation • The Grants found that the finches of the Galapagos Islands had a great variety of heritable traits. • Many of the characteristics appeared in a bell-shaped distributions typical of polygenic traits. • This change in beak size is an example of directional selection operating on an anatomical trait. Rapid Evolution • The Grants found that natural selection takes place frequently—and sometimes very rapidly. • Changes in the food supply caused measurable fluctuations in the finch populations over a period of only decades. • This is very different from the slow, gradual evolution that Darwin envisioned. Speciation in Darwin’s Finches Founders Arrive • A few finches travel from South America to one of the islands, there they survive and reproduce. Separation of populations • Some birds from species A cross to a second island. The two populations no longer share a gene pool. Changes in the gene pool • Seed sizes on the second island favor birds with larger beaks. The population on the second island evolves into a population, B, with larger beaks. Eventually, population A and B will evolve into a separate species. Reproductive Isolation • Even if a few birds from population B move back to the first island, they will not breed with the birds from population A. • The differences in beak size and mating behavior will lead to reproductive selection. Ecological Competition • The two species will compete with each other for seeds when they live together. Continued Evolution • The process of isolation on different islands, genetic change, and reproductive isolation will continue to repeat itself. • Thirteen different species of finches live on the Galapagos islands today. Studying Evolution since Darwin Why is understanding evolution so important? • Evolution continues today. • For example, bacteria and viruses are evolving resistance to drugs. • Insects are evolving resistance to pesticides. • Evolutionary theory can help us understand and respond to these changes in ways that can improve human life. There are still many unanswered questions that need to be addressed. Chapter 17 The History of Life Origins: The Early Idea • In the past, the ideas that decaying meat produced maggots, mud produced fishes, and grain produced mice were reasonable explanations for what people observed occurring in their environment. • Such observations led people to believe in spontaneous generation—the idea that nonliving material can produce life. Spontaneous generation is disproved • In 1668, an Italian physician, Francesco Redi, disproved a commonly held belief at the time—the idea that decaying meat produced maggots, which are immature flies. • However, during Redi’s time, scientists began to use the latest tool in biology—the microscope. • Although Redi had disproved the spontaneous generation of Control group large organisms, many scientists thought that microorganisms were so numerous and widespread that they must arise spontaneouslyprobably from a vital force in the air. Time Time Experimental group Spontaneous generation is disproved • In the mid-1800s, Louis Pasteur designed an experiment that disproved the spontaneous generation of microorganisms. • Pasteur set up an experiment in which air, but no microorganisms, was allowed to contact a broth that contained nutrients. • Pasteur’s experiment showed that microorganisms do not simply arise in broth, even in the presence of air. • From that time on, biogenesis, the idea that living organisms come only from other living organisms, became a cornerstone of biology. • The cell theory states that cells can arise from preexisting cells. Broth is boiled. Broth is free of microorganisms for a year. Curved neck is removed. Broth is teeming with microorganisms. The Origin of Life • Francesco Redi and Louis Pasteur designed controlled experiments to disprove spontaneous generation. • Their experiments and others like them convinced scientists to accept biogenesis. • But if life comes only from life, then how did life on Earth first begin? • No one has yet proven scientifically how life on Earth began. • However, scientists have developed theories about the origin of life on Earth from testing scientific hypotheses about conditions on early Earth. Origins: The Modern Ideas 1. Early Earth was hot; atmosphere contained poisonous gases. – Some scientists suggest that it was probably very hot. The energy from colliding meteorites could have heated its surface, while both the compression of minerals and the decay of radioactive materials heated its interior. – Volcanoes might have frequently spewed lava and gases, relieving some of the pressure in Earth’s hot interior. These gases helped form Earth’s early atmosphere. Earth’s early atmosphere probably contained hydrogen cyanide, carbon dioxide, carbon monoxide, nitrogen, hydrogen sulfide, and water. Origins: The Modern Ideas 2. Earth cooled and oceans condensed. • About 4.4 billion years ago, Earth might have cooled enough for the water in its atmosphere to condense. This might have led to millions of years of rainstorms with lightning—enough rain to fill depressions that became Earth’s oceans. Origins: The Modern Ideas 3. Simple organic molecules may have formed in the oceans. – In the 1930s, a Russian scientist, Alexander Oparin, hypothesized that life began in the oceans that formed on early Earth. – He suggested that energy from the sun, lightning, and Earth’s heat triggered chemical reactions to produce small organic molecules from the substances present in the atmosphere – Then, rain probably washed the molecules into the oceans to form what is often called a primordial soup. – In 1953, two American scientists, Stanley Miller and Harold Urey, tested Oparin’s hypothesis by simulating the conditions of early Earth in the laboratory Origins: The Modern Ideas 4. Small sequences of RNA may have formed and replicated. • Scientists hypothesize that two developments must have preceded the appearance of life on Earth. – First, simple organic molecules, or molecules that contain carbon, must have formed. – Then these molecules must have become organized into complex organic molecules such as proteins, carbohydrates, and nucleic acids that are essential to life. 5. First prokaryotes may have formed when RNA or DNA was enclosed in microspheres. • The first forms of life may have been prokaryotic forms that evolved from a protocell. A protocell is a large, ordered structure, enclosed by a membrane, that carries out some life activities, such as growth and division. – Because Earth’s atmosphere lacked oxygen, these organisms were most likely anaerobic. – For food, the first prokaryotes probably used some of the organic molecules in oceans. – Over time, these heterotrophs would have used up the food supply. • However, organisms that could make food had probably evolved by the time the food was gone. – These first autotrophs were probably similar to present-day archaebacteria. – Archaebacteria are prokaryotic and live in harsh environments, such as deep-sea vents and hot springs. – The earliest autotrophs probably made glucose by chemosynthesis rather than by photosynthesis. – In chemosynthesis, autotrophs release the energy of inorganic compounds, such as sulfur compounds, in their environment to make their food. • 6. Later prokaryotes were photosynthetic and produced oxygen. – Photosynthesizing prokaryotes might have been the next type of organism to evolve. – As the first photosynthetic organisms increased in number, the concentration of oxygen in Earth’s atmosphere began to increase. – Organisms that could respire aerobically would have evolved and thrived. – The presence of oxygen in Earth’s atmosphere probably affected life on Earth in another important way. • 7. An oxygenated atmosphere capped by the ozone layer protected Earth. – The sun’s rays would have converted much of the oxygen into ozone molecules that would then have formed a layer that contained more ozone than the rest of the atmosphere. • 8. First eukaryotes may have been communities of prokaryotes. – Complex eukaryotic cells probably evolved from prokaryotic cells. – The endosymbiont theory, proposed by American biologist Lynn Margulis explains how eukaryotic cells may have arisen. The theory proposes that eukaryotes evolved through a symbiotic relationship between ancient prokaryotes. • New evidence from scientific research supports this theory and has shown that chloroplasts and mitochondria have their own ribosomes that are similar to the ribosomes in prokaryotes. • In addition, both chloroplasts and mitochondria reproduce independently of the cells that contain them. • The fact that some modern prokaryotes live in close association with eukaryotes also supports the theory. Chloroplast Aerobic bacteria Plants and plantlike protists Ancient Prokaryotes Nuclear envelope evolving Photosynthetic bacteria Mitochondrion Primitive Photosynthetic Eukaryote Animals, fungi, and non-plantlike protists Ancient Anaerobic Prokaryote Primitive Aerobic Eukaryote Origins: The Modern Ideas 9. Sexual reproduction increased genetic variability, hastening evolution. • increase in genetic variation greatly increases the chances of evolutionary change in a species due to natural selection 10. Multicellular eukaryotes evolved. • cells increased in diversity more rapidly Evolution of Life Review Evolution of Life Early Earth was hot; atmosphere contained poisonous gases. Earth cooled and oceans condensed. Simple organic molecules may have formed in the oceans.. Small sequences of RNA may have formed and replicated. First prokaryotes may have formed when RNA or DNA was enclosed in microspheres. Later prokaryotes were photosynthetic and produced oxygen. An oxygenated atmosphere capped by the ozone layer protected Earth. First eukaryotes may have been communities of prokaryotes. Sexual reproduction increased genetic variability, hastening evolution. Multicellular eukaryotes evolved. Patterns of Evolution • The history of life is the story of increasing complexity and diversity, but what does this history reveal about the process of evolution? • Biologists have observed different patterns of evolution that occur throughout the world in different natural environments. • Macroevolution refers to the large-scale evolutionary changes that take place over long periods of time. • Six important patterns of macroevolution are – – – – – – mass extinctions adaptive radiation convergent evolution coevolution punctuated equilibrium changes in developmental genes • These patterns support the idea that natural selection is an important agent for evolution. Mass Extinctions • • • • • • The term used to describe a species that has died out is extinct. More than 99 percent of all species that ever lived are now extinct. Huge numbers of species have disappeared in mass extinctions. The disappearance of so many species left many habitats open. For the survivors, there was a new world of ecological opportunity. Often, the result was a burst of evolution that produced an abundance of new species. We are the last Dodo birds on this planet, so I have put all of our eggs safely into this basket. Divergent Evolution • Divergent evolution occurs when populations change as they adapt to different environmental conditions, eventually resulting in new species. Species that were once similar to an ancestral species diverge, or become increasingly distinct. Extinct mamo Amakihi Possible Ancestral Lasan finch Crested honeycreeper Kauai Niihau • One type of divergent evolution is adaptive radiation. • Ex: Darwin’s Finches Molokai Oahu Maui Lanai Akialoa Kahoolawe Akepa Akiapolaau Akikiki Liwi Hawaii Apapane Maui parrotbill Palila Ou Grosbeak finch Adaptive Radiation • When an ancestral species evolves into an array of species to fit a number of diverse habitats, the result is called adaptive radiation. • Adaptive radiation in both plants and animals has occurred and continues to occur throughout the world and is common on islands. Coevolution • The process by which two species evolve in response to changes in each other over time is called coevolution. • The trees have evolved in response to their seed predators, we can observe geographic differences in pinecones. – Where there are squirrels, the pinecones are heavier with fewer seeds, but have thinner scales, like the pinecone on the left. – Where there are only crossbills, pinecones are lighter with more seeds, but have thick scales, like the one on the right. • The crossbills have evolved in response to the pine trees, we can observe geographic differences in birds. – Where the pinecones have thick scales, birds should have deeper, less curved bills (below right) than where the pinecones have thin scales (below left). Convergent Evolution • A pattern of evolution in which distantly related organisms evolve similar traits is called convergent evolution. • Convergent evolution occurs when unrelated species occupy similar environments in different parts of the world. • Each of these animals has a streamlined body and various appendages that enable it to move rapidly through water. Yet, the shark is a fish, the penguin is a bird, and the dolphin is a mammal. • They have evolved similar traits because they share similar environmental pressures. Gradualism vs. Punctuated Equilibrium • Scientists once argued that evolution occurs at a slow, steady rate, with small, adaptive changes gradually accumulating over time in populations. • In 1972, scientists proposed a different hypothesis known as punctuated equilibrium, which argues that speciation occurs relatively quickly, in rapid bursts, with long periods of genetic equilibrium in between. • Environmental changes, such as higher temperatures or the introduction of a competitive species, lead to rapid changes in a small population’s gene pool that is reproductively isolated from the main population. • Speciation happens quickly—in about 10,000 years or less. Loxodonta africana Elephas maximus 0 1 2 Elephas 3 4 Mammuthus primigenius Loxodonta Mammuthus 5 Primelephas 6 Ancestral species about 55 million years ago Gradualism vs. Punctuated Equilibrium • Biologists generally agree that both gradualism and punctuated equilibrium can result in speciation, depending on the circumstances. • Gradualism involves a slow, steady change in a particular line of descent. • Punctuated equilibrium involves stable periods interrupted by rapid changes involving many different lines of descent. Developmental Genes and Body Plans • Hox genes are called “master control genes,” because they control growth as an embryo develops. • Some hox genes determine which parts become front, rear, top, and bottom. • Others control the growth of body parts such as arms, legs, and wings. • The timing of genetic control during embryonic development can make the difference between various traits. Patterns of Evolution Flow Chart Species that are Unrelated form Related in under under in in Inter-relationships Similar environments Intense environmental pressure Small populations Different environments can undergo can undergo can undergo can undergo can undergo Co evolution Convergent evolution Extinction Punctuated equilibrium Adaptive radiation