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Diff. Biology Stud y Guide: Evolu tion Key Terms 1. Biological evolution- Evolution of life 2. Jean Baptist de Lamarck- -established early principles associated with evolution by comparing current species to fossils 1. Use and Disuse- Useful organs are enhanced and non-useful organs are discarded. (Giraffe) 2. Inheritance of acquired characteristics- Useful traits that are enhanced are passed to offspring -natural inheritance- Giraffe -man-influenced inheritance- evolution of dog 3. inheritance of acquired characteristics4. James Hutton- his most famous work, "Theory of the Earth". This work was interpreted and used by many as the basis for geological theory. He desired to trace back the origin of the various minerals and rocks, and thus to arrive at some clear understanding of the history of the earth. For many years he continued to study the subject. At last, in the spring of the year 1785, he communicated his views to the recently established Royal Society of Edinburgh in a paper entitled Theory of the Earth, or an Investigation of the Laws Observable in the Composition, Dissolution and Restoration of Land upon the Globe. In this remarkable work the doctrine is expounded that geology is not cosmogony, but must confine itself to the study of the materials of the earth; that everywhere evidence may be seen that the present rocks of the earth's surface have been in great part formed out of the waste of older rocks; that these materials having been laid down under the sea were there consolidated under great pressure, and were subsequently disrupted and upheaved by the expansive power of subterranean heat; that during these convulsions veins and masses of molten rock were injected into the rents of the dislocated strata; that every portion of the upraised land, as soon as exposed to the atmosphere, is subject to decay; and that this decay must tend to advance until the whole of the land has been worn away and laid down on the sea-floor, whence future upheavals will once more raise the consolidated sediments into new land. This is the premise to plate tectonics & continental drift. The problem with the traditional rock cycle is that it implies that rocks just cycle endlessly from one to the other. The drawing above expresses this cyclical nature (rock cycle illustrations come in endless variety; this is a very simple stripped down version.) It also expresses the view of the 19th century Uniformitarian school of thinking, captured best by James Hutton when he said the earth has "no vestige of a beginning, and no prospect of an end." He envisioned earth processes going round and round but never getting anywhere, never evolving. 5. catastrophism- Past Cataclysmic Activity Catastrophism is the idea that many of Earth’s crustal features (strata layers, erosion, polystrate fossils, etc) formed as a result of past cataclysmic activity. In other words, the Earth’s surface has been scarred by catastrophic natural disasters. Empirical Evidence Catastrophism is supported by actual, recorded history. Nearly 300 ancient flood legends have survived the ravishment of time. Legends of a worldwide deluge, commonly known as the "Noachian Flood," are found in Europe, Asia, Africa, Australia, North American and South America. Furthermore, earth's sedimentary layers with the fossil record seem to suggest a past marine cataclysm. Sedimentary rock (sandstone, siltstone, shale, limestone, etc) is the result of moving water, laid down layer upon layer by hydrologic sorting. Animals whose fossil remains are found within those layers must have been caught in this running water to have been buried and preserved. The remains, as well as the rocks, would be sorted according to density. Otherwise, the carcasses would rot or be scavenged. Approximately 95% of all earth's fossil remains discovered thus far are marine invertebrates. Of the remainder, approximately 4.74% are plant fossils, 0.25% are land invertebrates (including insects), and 0.0125% are vertebrates (the majority of which are fish). Roughly 95% of all land vertebrates discovered and recorded to date consist of less than one bone. 6.uniformitarianism - "The Present is the Key to the Past" Uniformitarianism is a geological doctrine. It states that current geologic processes, occurring at the same rates observed today, in the same manner, account for all of Earth's geological features. Thus, it assumes that geological processes are essentially unchanged today from those of the unobservable past, and that there have been no cataclysmic events in earth's history. As present processes are thought to explain all past events, the Uniformitarian slogan is, "the present is the key to the past." James Hutton and Sir Charles Lyell The doctrine of Uniformitarianism was significantly advanced by James Hutton (1726-1797) in his publication, Theory of the Earth (1785). Hutton influenced Sir Charles Lyell (1797-1875), who is acclaimed as the father of modern geology with his work, Principles of Geology (18301833, a three volume work). Lyell, in turn, influenced Charles Darwin, who later wrote The Origin of Species (1859). Lyell is responsible for the general acceptance of Uniformitarianism among geologists for the past 150 years 7.Charles Lyell - Sir Charles Lyell (1797-1875) was a British geologist. In his Principles of Geology (3 volumes, 1830-33), Lyell conclusively showed that the earth was very old and had changed its form slowly, mainly from conditions such as erosion. Lyell was able to date the ages of rocks by using fossils embedded in the stone as time indicators. Charles Darwin made use of Lyell's data on fossils for his theory of evolution. Lyell himself had believed that the various species of plants and animals had remained unchanged since they were created. When confronted with Darwin's findings, he admitted "I now realize I have been looking down the wrong road." He became one of Darwin's strongest supporters. Lyell was born in Scotland. He studied geology at Oxford University and traveled on several geological expeditions in Europe and North America. 8.Alfred Russel Wallace. (January 8, 1823 — November 7, 1913) was a British naturalist, geographer, anthropologist and biologist. Wallace's independent proposal of a theory of evolution by natural selection prompted Charles Darwin to reveal his own more developed and researched, but unpublished, theory sooner than he had intended. In 1855, Wallace published a paper, "On the Law Which has Regulated the Introduction of Species" (1855) (http://www.wku.edu/%7Esmithch/wallace/S020.htm), in which he gathers and enumerates general observations regarding the geographic and geologic distribution of species, and concludes that "Every species has come into existence coincident both in space and time with a closely allied species." The paper was a foreshadowing of the momentous paper he would write three years hence. Wallace had once briefly met Darwin, and was one of Darwin's numerous correspondents from around the world, whose observations Darwin used to support his theories. Wallace knew that he was interested in the question of how species originate, and trusted his opinion on the matter. Thus, he sent him his essay, "On the Tendency of Varieties to Depart Indefinitely From the Original Type" (1858) (http://www.wku.edu/%7Esmithch/wallace/S043.htm), and asked him to review it. In it, Wallace describes a novel theory of what is now known as "natural selection," and proposes that it explains the diversity of life. It was essentially the same as the theory that Darwin had worked on for twenty years, but had yet to publish. Darwin wrote in a letter to Charles Lyell: "he could not have made a better short abstract! Even his terms now stand as heads of my chapters!" Although Wallace had not requested that his essay be published, Charles Lyell and Joseph Hooker decided to present the essay, together with excerpts from a paper that Darwin had written in 1844, and kept confidential, to the Linnean Society of London on 1 July 1858, highlighting Darwin's priority. 9.Charles Darwin - (12 February 1809 – 19 April 1882) was an English naturalist whose revolutionary theory laid the foundation for both the modern theory of evolution and the principle of common descent by proposing natural selection as a mechanism. He published this proposal in 1859 in the book The Origin of Species, which remains his most famous work. A worldwide sea voyage aboard HMS Beagle and observations on the Galapagos Islands in particular provided inspiration and much of the data on which he based his theory. 10. Natural Selection - is the primary mechanism within the scientific theory of evolution, i.e. it alters the frequency of alleles within a population. After a century of obscure and vague preliminary formulations, it was proposed as the main mechanism of evolution by Charles Darwin and Alfred Russel Wallace in 1858. Natural selection can be subdivided into two types; ecological selection, and sexual selection. Natural selection is distinguished from artificial selection by humans. Other mechanisms of evolution include genetic drift and gene flow. Mutations create the genetic variation on which natural selection acts. It is important to note that the term "natural selection" is often used in the inaccurate yet fairly harmless metaphorical sense as having causal status. To be precise, natural selection is not truly a "mechanism" in itself, as opposed to something like gravity. Instead, natural selection is the result of genetic and environmental forces acting upon an organism. 11. survival of the fittest - is a phrase which is a shorthand for a concept relating to competition for survival or predominance. It codifies the observance of zoologists and animal breeders that individuals having the highest level of fitness for a particular environment tend to survive longer in it. Moveover, the longer they survive, the more territory they dominate and the more offspring they have -- tending to crowd out competitors. The phrase is essentially a metaphor and is often felt to be unhelpful: it is very close to a tautology, since if "fitness" is measured in the obvious objective terms - survival - the phrase becomes "survival of the survivors". The phrase "survival of the fittest" was first used by Herbert Spencer in his 1851 work Social Statics. It was used by Charles Darwin in the 6th edition of The Origin of Species, in a secondary header of Chapter 4 about natural selection [1] (http://www.bartleby.com/11/4001.html) and at several places in the text, mostly using the phrase "Natural Selection, or the Survival of the Fittest". He gave full credit to Spencer, writing "I have called this principle, by which each slight variation, if useful, is preserved, by the term natural selection, in order to mark its relation to man's power of selection. But the expression often used by Mr. Herbert Spencer, of the Survival of the Fittest, is more accurate, and is sometimes equally convenient.". At this time the word "fittest" would have primarily meant "most suitable" or "most appropriate" rather than "in the best physical shape". In the first five editions of The Origin of Species, Darwin used the phrase natural selection [2] 12. Thomas Malthus - (February, 1766 – December 23, 1834), who preferred to be known as "Robert Malthus", was an English demographer and political economist best known for his pessimistic but highly influential views. Malthus's views were largely developed in reaction to the optimistic views of his father and his associates, notably Rousseau and William Godwin. In An Essay on the Principle of Population, published in 1798, Malthus predicted population would outrun food supply, leading to a decrease in food per person. This prediction was based on the idea that population if unchecked increases at a geometric rate whereas the food supply grows at an arithmetic rate. (See Malthusian catastrophe for more information.) Only misery, moral restraint and vice (which for Malthus included contraception) could check excessive population growth. Malthus favoured "moral restraint" (including late marriage and sexual abstinence) as a check on population growth. 13. variation - The gene pool of a species or a population is the complete set of unique alleles that would be found by inspecting the genetic material of every living member of that species or population. A large gene pool indicates a large genetic diversity, which is associated with robust populations that can survive bouts of intense selection. Meanwhile, low genetic diversity (see inbreeding and population bottlenecks) can cause reduced fitness and an increased chance of extinction. The phenotypes associated with multiple alleles creates a range of expressions. This helps ensure certain phenotypic/genotypic progeny are able to adapt to changing environmental stresses and thus survive. 14. excessive reproduction- all populations have the potential to overpopulate their environment. 15. evidence for evolution- tools used by biologists to provide evidence for the theory of evolution. 16. Geologic- utilizing sediment formations and remnants within these layers to address chronological evolution. 17. Radioactive decay- most elements have unstable, radioactive isotopes which decay at a specific rate (half-life). The ratios of radioactive elements to their stable decomposed elements provides a relative time for the existence of an earlier organism. 18. Fossils- Fossils are the mineralized remains of animals or plants or other artifacts such as footprints. The totality of fossils and their placement in rock formations and sedimentary layers (strata) is known as the fossil record. The study of fossils is called paleontology. 19. Paleontology- study of life of the past using fossil remnants 20. Biogeography- the study of land forms of the earth and the organisms that inhabit these locations. 21. Comparative anatomy- comparing anatomical structures of living or nonliving organisms. 22. homologous- Two or more structures are said to be homologous if they are alike because of shared ancestry. This could be evolutionary ancestry, meaning that the structures evolved from some structure in a common ancestor (the wings of bats and the arms of humans are homologous in this sense) or developmental ancestry, meaning that the structures arose from the same tissue in embryonal development (the ovaries of females and the testicles of males are homologous in this sense). 23. analogous- Two structures in biology are said to be analogous if they perform a similar function by a simlar mechanism, but did not arise from a common ancestor performing that function: for example, the wings of insects and the wings of birds. These similar structures most likely evolved through different pathways, a process known as convergent evolution. 24. vestigial- A vestigial organ is an organ whose original function has been lost during evolution. In 1893, Robert Wiedersheim published a list of 86 human organs that had no known function. Theorizing that they were vestiges of evolution, he called them "vestigial". Today, the list of human vestigial organs is much smaller, and hotly debated. It still includes the appendix, and coccyx. Many people maintain that the coccyx is a remnant of a lost tail. Wisdom teeth are vestigial as well. Also, the plica semilunaris, the small fold of tissue on the inside corner of the eye, is the vestigial remnant of the nictitating membrane (the third eyelid) in other animals. The formation of goose bumps in humans under emotional stress is a vestigial reflex; its purpose in our evolutionary ancestors was to raise hair to make the animal appear bigger and scare off enemies. 25. Domestication- Domesticated animals, plants, and other organisms are those whose collective behavior, life cycle, or physiology has been altered as a result of their breeding and living conditions being under human control for multiple generations. Humans have brought these populations under their care for a wide range of reasons: for help with various types of work, to produce food or valuable commodities (such as wool, cotton, or silk), and to enjoy as pets or ornamental plants. 26. Embryology- Embryology is the subdivision of developmental biology that studies embryos and their development. The formation and development of an embryo is known as embryogenesis. 27. Biochemistry- study of similiarities & differences in biomolecules. 28. Molecular Genetics29. Modern Synthesis- The modern evolutionary synthesis (often referred to simply as the modern synthesis), neo-Darwinian synthesis or neo-Darwinism, brings together Charles Darwin's theory of the evolution of species by natural selection with Gregor Mendel's theory of genetics as the basis for biological inheritance. Major figures in the development of the modern synthesis include Ronald Fisher, Theodosius Dobzhansky, J.B.S. Haldane, Sewall Wright, Julian Huxley, Ernst Mayr, George Gaylord Simpson and G. Ledyard Stebbins. Essentially, the modern synthesis (or neo-Darwinism) introduced the connection between two important discoveries; the units of evolution (genes) with the mechanism of evolution (selection). George John Romanes coined the term neo-Darwinism to refer to the theory of evolution preferred by Alfred Russel Wallace et al. Wallace rejected the Lamarckian idea of inheritance of acquired characteristics, something that Darwin, Huxley et al wouldn't rule out. The mechanism of inheritance wasn't discovered in Darwin or Wallace's time, however, so the debate was never settled. Mendelian genetics was rediscovered in 1900. However, there were differences of opinion as to what was the variation that natural selection acted upon. The biometric school, led by Karl Pearson followed Darwin's idea that small differences were important for evolution. The Mendelian school, led by William Bateson, however thought that Mendel's work gave an evolutionary mechanism with large differences. This issue was finally resolved by Ronald Fisher, who in 1918 produced a paper entititled "The Correlation Between Relatives on the Supposition of Mendelian Inheritance", which showed using a model how continuous variation could be the result of the action of many discrete loci. This is generally regarded as the starting point of the synthesis. According to the modern synthesis as established in the 1930s and 1940s, genetic variation in populations arises by chance through mutation (this is now known to be due to mistakes in DNA replication) and recombination (crossing over of homologous chromosomes during meiosis). Evolution consists primarily of changes in the frequencies of alleles between one generation and another as a result of genetic drift, gene flow and natural selection. Speciation occurs gradually when populations are reproductively isolated by geographic barriers. The modern evolutionary synthesis continued to be developed and refined after the initial establishment in the 1930s and 1940s. The most notable paradigm shift was the so-called Williams revolution, after George C. Williams presented a gene-centric view of evolution in the 1960s. The synthesis as it exists now has extended the scope of the Darwinian idea of natural selection, specifically to include subsequent scientific discoveries and concepts unknown to Darwin such as DNA and genetics that allow rigorous, in many cases mathematical, analyses of phenomena such as kin selection, altruism, and speciation. A particular interpretation of neoDarwinism most commonly associated with Richard Dawkins asserts that the gene is the only true unit of selection. Dawkins further extended the Darwinian idea to include non-biological systems exhibiting the same type of selective behavior of the 'fittest' such as memes in culture. 30. species- In biology, a species is, loosely speaking, a group of related organisms that share a more or less distinctive form and are capable of interbreeding. As defined by Ernst Mayr, species are "groups of actually or potentially interbreeding natural populations which are reproductively isolated from other such groups" (however, see definitions of species below). 31. gene pool- The gene pool of a species or a population is the complete set of unique alleles that would be found by inspecting the genetic material of every living member of that species or population. A large gene pool indicates a large genetic diversity, which is associated with robust populations that can survive bouts of intense selection. Meanwhile, low genetic diversity (see inbreeding and population bottlenecks) can cause reduced fitness and an increased chance of extinction. When many alleles exist for a given gene or locus, a population is said to be polymorphic with respect to that gene or locus. When no variation exists, it is labelled monomorphic. 32. allelic frequency- a term of population genetics that is used in characterizing the genetic diversity of a species population, or equivalently the richness of its gene pool. Allele frequency is defined as follows. Given a) a particular chromosome locus, b) a gene occupying that locus, c) a population of individuals carrying n loci in each of their somatic cells (e.g. two loci in the cells of diploid species, which contain two sets of chromosomes) and finally d) a variant or allele of the gene, then the allele frequency of that allele is the fraction or percentage of loci that the allele occupies within the population. Note that for diploid genes, however, the proportion of individuals that carry this allele may be up to two in five. If the allele distributes randomly, then the binomial theorem will apply: 32% of the population will be heterozygous for the allele (i.e. carry one copy of that allele and one copy of another in each somatic cell) and 4% will be homozygous (carrying two copies of the allele). So all together 36% of diploid individuals would be expected to carry an allele that has a frequency of 20%. However, alleles distribute randomly only in the absence of selection and under other assumptions. When these conditions apply, a population is said to be in Hardy-Weinberg equilibrium. The frequencies of all the alleles of a given gene often are graphed together as an allele frequency distribution histogram. Population genetics studies the different "forces" that might lead to changes in the distribution and frequencies of alleles -- in other words, to evolution. Besides selection, these forces include genetic drift, mutation and migration. 33. genetic equilibrium- the frequency of alleles, genotypes & phenotypes are relatively stable within a population exposed to consistent environmental factors 34. genotype frequencies- the frequency of a certain genotypes (homozygous dominant, homozygous recessive, or heterozygous) within a population. 35. Hardy-Weinberg Law- The Hardy — Weinberg principle (HWP) (also Hardy — Weinberg equilibrium (HWE), or Hardy — Weinberg law) states that, under certain conditions, after one generation of random mating, the genotype frequencies at a single gene locus will become fixed at a particular equilibrium value. It also specifies that those equilibrium frequencies can be represented as a simple function of the allele frequencies at that locus. 36. genetic drift-Genetic drift is a mechanism of evolution that acts in concert with natural selection to change the characteristics of species over time. It is a stochastic effect that arises from the role of random sampling in the production of offspring. Like selection, it acts on populations, altering the frequency of alleles and the predominance of traits amongst members of a population, and changing the diversity of the group. Drift is observed most strongly in small populations and results in changes that need not be adaptive. 37. neutral selection- the idea that many of the changes seen at the molecular level are selectively. Prior to the 1960's little was known about the details of the molecular basis of heredity or about the extent of natural variation in proteins and DNA. With the development of techniques to visualize isozyme variation and to determine the primary sequence of proteins, population biologists began to examine the amount of variation in proteins both within and between populations. The amount of variation and the rate at which it was accumulating was surprisingly high. Based upon population genetic theory, which had been developed during the first half of the 20th century, the amount of variation was difficult to account for if all of the variation was under selection and was being maintained by selection. 38. mutation pressure 39. gene flow- (also known as gene migration) is the transfer of genes from one population to another. Migration into or out of a population may be responsible for a marked change in gene pool frequencies (the number of individual members with a particular trait). Immigration may result in the addition of new genetic material to the established gene pool of a particular species or population, and conversely emigration results in the removal of genetic material. There are a number of factors that affect the rate of gene flow between different populations. One of the most significant factors is mobility, and animals tend to be more mobile than plants. Greater mobility of an individual tends to give it greater migratory potential. 40. bottleneck effect- A population bottleneck (or genetic bottleneck) is an evolutionary event in which a significant percentage of a population or species is killed or otherwise prevented from reproducing, and the population is reduced by 50% or more, often by several orders of magnitude. A graph of this change resembles the neck of a bottle, from wide to narrow; hence the name. Population bottlenecks increase genetic drift, as the rate of drift is inversely proportional to the population size. It also changes the relationship of natural selection (see: inbreeding). 41. founder effect- The founder effect is an evolutionary phenomenon. Founder effects arise when a new and isolated environment is invaded by only a few members of a species, which then multiply rapidly. In the extreme case, a single fertilised female might arrive in a new environment. It is a type of population bottleneck. The result of the small number of founders is that there is a sharp loss of genetic variation compared with the parent population. As a result, the new population may be distinctively different, genetically and phenotypically, from the parent population it derived from. In addition, there is a raised probability of inbreeding, resulting in an unusual number of defects due to recessive genes. Founder effects are common in island ecology, but the isolation need not be geographical. For example, the Amish populations in the United States, which have grown from a very few founders but have not recruited newcomers, and tend to marry within the community, exhibit founder effects: phenomena such as polydactyly (extra fingers and toes, a symptom of Ellis-van Creveld syndrome), though still rare absolutely, are more common in Amish communities than in the US population at large. In extreme cases, founder effects may lead to the speciation and subsequent evolution of new species. 42. selection pressure- an environmental factor which influences genotypes/phenotypes in a population. 43. polymorphism- describes multiple possible states for a single property (it is said to be polymorphic). In biology multiple alleles of a gene within a population, usually expressing different phenotypes, are called polymorphism. For instance, human skin color is polymorphic. See polymorphism (biology). Polymorphism in biology can also refer to single nucleotide polymorphisms 44. heterozygote advantage- A heterozygote advantage (heterozygous advantage or overdominance) describes the case in which the heterozygote genotype has a higher relative fitness than either the homozygote dominant or homozygote recessive genotype. This selection favoring the heterozygote is one of the mechanisms that maintains polymorphism and helps to explain some kinds of genetic variability. There are several cases in which the heterozygote is conveyed certain advantages and some disadvantages while both versions of homozygotes are only at disadvantages. A well established case of heterozygote advantage is that of the gene involved in sickle cell anaemia. Often times the advantages and disadvantages conveyed are rather complicated, due to the fact that more than one gene exists on any given allele. Major genes almost always have multiple effects, which can simultaneously convey separate advantageous traits and disadvantageous traits upon the same organism. In this instance, the state of the organism’s environment will provide selection, with a net effect either favoring or working in opposition to the gene, until an environmentally-determined equilibrium is reached. This sort of selection can be seen in all kinds of populations human and non-human. In the fly Drosophila melanogaster, there is an autosomal, completely recessive gene that expresses ebony body-color. When there is a fly with two copies of the recessive allele, this homozygote expresses the dark ebony color, but is also terribly weakly, and is placed at a harsh reproductive disadvantage. If this were the only effect of the gene, and only conveyed disadvantages, we would expect selection to ‘weed out’ this gene until it became extinct from the population. However, the same gene also conveys some advantages, by providing improved viability. This advantage is dominant, and conveyed in the heterozygote. So in this case, the homozygote ‘ebony color’ gene fly will be at a distinct advantage due to weakness; the heterozygote will express none of the disadvantages of weakness, but will gain from his one allele that improves his viability; and the homozygote wild-type will be perfectly healthy, but does not possess the improved viability of the heterozygote, and will be at a disadvantage in survivorship and reproduction. Sickle cell anemia (SCA) is a genetic disorder that is caused by the presence of two incompletely recessive alleles. When a sufferer’s red blood cells are exposed to low oxygen conditions, the cells lose their healthy round shape and become sickle shaped. This deformation of the cells can cause them to become lodged in capillaries, depriving other parts of the body from precious oxygen. When untreated, a person with SCA may suffer from painful periodic bouts, often causing damage to internal organs, strokes, or anemia. Typically the disease results in premature death. However, there is convincing evidence indicating that in areas with persistent malaria outbreaks, individuals with the heterozygous state had a distinct advantage historically. Those with the benign sickle trait possess a resistance to malarial infection. The pathogen that causes the disease spends part of its cycle in the red blood cells, and those with sickle cells effectively stop the pathogen in its tracks, until the immune system destroys the foreign bodies. These individuals have a great immunity to infection and have a greater chance of surviving outbreaks and going on to reproduce. However, those with two alleles for SCA may survive malaria but will typically die from their genetic disease unless they have access to advanced medical care. The homozygous ‘normal’ or wild-type case will have a greater chance of passing on their genes successfully, in that there is no chance of their offspring suffering from SCA; yet, they are more susceptible to dying from malarial infection before they have a chance to pass on their genes. Cystic fibrosis, or CF, is an autosomal recessive hereditary disease of the lungs, sweat glands and digestive system. The disorder is caused by the malfunction of the CFTR protein, which controls inter-membrane transport of chloride ions, which is vital to maintaining equilibrium of water in the body. The malfunctioning protein causes viscous mucus to form in the lungs and intestinal tract. Historically, a child born with CF would only have a life expectancy of a few years, but modern medicine has made it possible for these people to life into adulthood. However, even in these individuals, male and female, CF typically causes sterility. It is the most common genetic disease among people of European descent. The possession of the CF disease can increase survivorship of people affected by diseases involving loss of body fluid, typically due to diarrhea. The most common of these maladies is cholera, which killed many Europeans historically. Those with cholera would often die of dehydration when they ingested water slower than their body passed it. Those with CF, or either carriers of the allele where much less likely to suffer this fate. Due to this increased resistance to disease, the CF allele in low frequencies was beneficial to the gene pool. Even today, approximately one in twenty-five of European descent is a carrier for the disease, and one in every 2500 children born is affected by cystic fibrosis. 45. directional selection- In population genetics, directional selection occurs when natural selection favors a single allele and therefore allele frequency continuously shift in one direction. It is in contradistinction to balancing selection where selection may favor multiple alleles. Directional selection is a particular mode or mechanism of natural selection. A common example is the peppered moth (Biston betularia). Before the industrial revolution in England (1740?), the peppered moth was mostly found in a light gray form with little black speckled spots. The allele for dark-bodied moths is dominant, while the allele for light-bodied moths is recessive. The light-bodied moths were able to blend in with the light colored lichens and tree bark. The less common black peppered moth was more likely to be eaten by birds. Therefore, the frequency of the dark allele was about 0.01%. During the industrial revolution in England, many of the light-bodied lichens died from sulphur dioxide emissons. The trees became covered with soot from the new coalburning factories. This led to an increase in bird predation for the light-colored moths (they no longer blended in as well). The dark-bodied moths, however, blended in very well with the trees. As a result, during reproduction, a lot of light-bodied moths were produced, and a few dark-bodied moths. Most of the light-bodied moths didn't survive, while the black-bodied moths continued to survive. Gradually, the allele frequency shifted towards the dominant allele, as more and more darkbodied moths survived to reproduce. 46. disruptive selection- Disruptive selection is a type of natural selection that simultaneously favors individuals at both extremes of the distribution. When disruptive selection operates, individuals at the extremes contribute more offspring than those in the center, producing two peaks in the distribution of a particular trait. A hypothetical example of this is where you have a population of rabbits. The dominant trait being black fur represented by “B” the recessive trait being white fur represented by “b”. A rabbit with the genotype of “BB” would have a phenotype of black fur, a genotype of “Bb” would have gray fur, and a geno type of “bb” would have a phenotype of white fur. Put this population of rabbits into an area that had very dark black rocks as the enviroment and also near by was an area of very white colored stone. The rabbits with the black fur would be able to hide from preditors amongst the black rocks and the white furred rabbits would be able to hide in the white rocks but the gray furred rabbits would stand out in both of the habitats and thus not survive. 47. stabilizing selection- Stabilizing selection favors the norm, the common, average traits in a population. Look at the Siberian Husky, a dog bred for working in the snow. The Siberian Husky is a medium dog, males weighing 16-27kg (35-60lbs). These dogs have strong pectoral and leg muscles, allowing it to move through dense snow. The Siberian Husky is well designed for working in the snow. If the Siberian Husky had heavier muscles, it would sink deeper into the snow, so they would move slower or would sink and get stuck in the snow. Yet if the Siberian Husky had lighter muscles, it would not be strong enough to pull sleds and equipment, so the dog would have little value as a working dog. So stabilizing selection has chosen a norm for the the size of the Siberian Husky. 48. balanced polymorphism - A balanced polymorphism is a situation in which natural selection within a population is able to maintain stable frequencies of two or more phenotypic forms. There are two major mechanisms by which natural selection preserves this variation and consequently produces a balanced polymorphism. The first mechanism is known as heterozygote advantage; in which an individual who is heterozygous at a particular gene locus has a greater reproductive success than a homozygous individual. This can be seen in human populations with the locus for a certain protein present in hemoglobin (an important component in blood). Individuals who are homozygous for the recessive allele at this locus are inflicted with sickle-cell disease,a disease in which blood cells are grossly misshapen and which often results in a reduced lifespan. An individual heterozygous at this locus will not suffer from sickle-cell disease but because of slightly irregularly shaped blood cells they are resistant to malaria. This resistance is favored by natural selection in tropical regions where malaria is present and so the heterozygote has an evolutionary edge. It is in this way that natural selection preserves stable frequencies of both the heterozygote and the homozygote dominant phenotypes. The second important mechanism by which natural selection can preserve two or more phenotypic forms is known as frequency-dependent selection. Frequency-dependent selection is a form of selection in which the relative "fitness" of a specific phenotype declines if the frequency of that phenotype becomes too high. An example of this type of selection is between parasites and their hosts. An example follows: suppose that a certain parasite can recognize one of two receptors in its host, receptor Alpha or receptor Beta, if many parasites with receptor Alpha exist then hosts with receptor Beta will be selected for, and this will subsequently increase the selective pressure on parasites which use receptor Beta and this relationship will continue rocking back and forth. 49. adaptation- Adaptation in biology, an anatomical structure, physiological process or behavioral trait that has evolved over a period of time by the process of natural selection that increases the likelihood of producing larger numbers of offspring or its reproductive success. 50. fitness- Fitness in biology refers to individual's ability to propagate its genes. 51. sexual selection- Sexual selection is the theory that competition for mates between individuals of the same sex drives the evolution of certain traits. It is distinct from ecological selection which is the competition for food within the species' ecological niche. Many traits, e.g. smooth skin or fur, strong muscles, fluid motions, appear not only to enable hunting or gathering but also to be important sexual attractors, especially in the more intelligent species. For these, ecological and sexual selection both operate on a trait. Traits amenable to sexual selection, which give an individual an advantage over its rivals in courtship without being directly involved in reproduction, are called secondary sex characteristics. Sex differences directly related to reproduction and serving no direct purpose in courtship are called primary sex characteristics. These traits are caused by secondary sexual characteristics. Secondary sexual characteristics distinguish the sexes of a species, yet have no direct role in reproduction. As mentioned above, these dimorphic traits sometimes become overdeveloped relative to other members of a species. Sometimes these exaggerated traits prove to be a hindrance to the animal, thereby lowering their fitness. These animals are still selected sexually and are therefore able to reproduce. Through this process, the exaggerated trait is passed on to future generations. Large antlers such as that of a moose are bulky and heavy and slow the creature down when running from a predator as well as run the risk of becoming entangled in low hanging tree branches and shrubs. No doubt this trait has lead to the demise of a creature in more than one instance. Bright colorations, such as those seen in many male birds, capture both the eye of a female and that of a predator; when a male peacock spreads it tail, it is beautiful, but very obvious. These traits also represent a costly energetic investment for the animal that bears them. 52. species- In biology, a species is, loosely speaking, a group of related organisms that share a more or less distinctive form and are capable of interbreeding. As defined by Ernst Mayr, species are "groups of actually or potentially interbreeding natural populations which are reproductively isolated from other such groups" (however, see definitions of species below). The definition of a species given above as taken from Mayr, is somewhat idealistic. Since it assumes sexual reproduction, it leaves the term undefined for a large class of organisms that reproduce asexually. Biologists frequently do not know whether two morphologically similar groups of organisms are "potentially" capable of interbreeding. Further, there is considerable variation in the degree to which hybridization may succeed under natural and experimental conditions, or even in the degree to which some organisms use sexual reproduction between individuals to breed. Consequently, several lines of thought in the definition of species exist: •A morphological species is a group of organisms that have a distinctive form: for example, we can distinguish between a chicken and a duck because they have different shaped bills and the duck has webbed feet. Species have been defined in this way since well before the beginning of recorded history. Although much criticised, the concept of morphological species remains the single most widely used species concept in everyday life, and still retains an important place within the biological sciences, particularly in the case of plants. •The biological species or isolation species concept identifies a species as a set of actually or potentially interbreeding organisms. This is generally the most useful formulation for scientists working with living examples of the higher taxa like mammals, fish, and birds, but meaningless for organisms that do not reproduce sexually. It distinguishes between the theoretical possibility of interbreeding and the actual likelihood of gene flow between populations. For example, it is possible to cross a horse with a donkey and produce offspring, however they remain separate species—in this case for two different reasons: first because horses and donkeys do not normally interbreed in the wild, and second because the fruit of the union is rarely fertile. The key to defining a biological species is that there is no significant cross-flow of genetic material between the two populations. •A mate-recognition species is defined as a group of organisms that are known to recognise one another as potential mates. Like the isolation species concept above, it applies only to organisms that reproduce sexually. •A phylogenetic or evolutionary or Darwinian species is a group of organisms that shares an ancestor; a lineage that maintains its integrity with respect to other lineages through both time and space. At some point in the progress of such a group, members may diverge from one another: when such a divergence becomes sufficiently clear, the two populations are regarded as separate species. 53. subspecies- In taxonomy, a subspecies is the taxon immediately subordinate to a species. Members of one subspecies differ morphologically but sometimes only genetically (disputed — see talk page) from members of other subspecies of the species. Subspecies are defined in relation to species. It is not possible to understand the concept of a subspecies without first grasping what a species is. In the context of large living organisms like trees, flowers, birds, fish and humans, a species can be defined as a distinct and recognisable group that satisfies two conditions: 1. Members of the group are reliably distinguishable from members of other groups. The distinction can be made in any of a wide number of ways, such as: differently shaped leaves, a different number of primary wing feathers, a particular ritual breeding behaviour, relative size of certain bones, different DNA sequences, and so on. There is no set minimum 'amount of difference': the only criterion is that the difference be reliably discernable. In practice, however, very small differences tend to be ignored. 2. The flow of genetic material between the group and other groups is small and can be expected to remain so because even if the two groups were to be placed together they would not interbreed to any great extent. Note the key qualifier above: to be regarded as different groups rather than as a single varied group, the difference must be distinct, not simply a matter of continuously varying degree. If, for example, the population in question is a type of frog and the distinction between two groups is that individuals living upstream are generally white, while those found in the lowlands are black, then they are classified as different groups if the frogs in the intermediate area tend to be either black or white, but a single, varied group if the intermediate population becomes gradually darker as one moves downstream.This is not an arbitrary condition. A gradual change, called a cline, is clear evidence of substantial gene flow between two populations. A sharp boundary between black and white, or a relatively small and stable hybrid zone, on the other hand, shows that the two populations do not interbreed to any great extent and are indeed separate species. Their classification as separate species or as subspecies, however, depends on why they do not interbreed. 54. anagenesis- Anagenesis is the progressive evolution of species involving a change in gene frequency in an entire population rather than a cladogenetic branching event. When enough mutations reach fixation in a population to significantly differentiate from an ancestral population a new species name may be assigned. A key point is that the entire population is different from the ancestral population so that the ancestral population can be considered extinct. It is easy to see from the preceding definition how controversy can arise among taxonomists regarding when the differences are significant enough to warrant a new species classification. Anagenesis may also be referred to as phyletic evolution 55. cladogenesis- Cladogenesis is an evolutionary splitting event in which each branch and its smaller branches is a "clade"; an evolutionary mechanism and a process of adaptive evolution that leads to the development of a greater variety of animals or plants. 56. cline- In population genetics, a cline is a gradual change of a character or feature (phenotype) in a species over a geographical area. The change in phenotype does not result in different species as long as the geographically spread populations can interbreed with one another. This meaning of cline was introduced by Sir Julian Huxley 57. subspecies- see 53 58. races- A race is a distinct population of humans distinguished in some way from other humans. The most widely observed races are those based on skin color, facial features, ancestry, and genetics. Conceptions of race, as well as specific racial groupings, are often controversial due to their political and sociological uses and implications. Since the 1940s, evolutionary scientists have rejected the view of race according to which a number of finite lists of essential (e.g., Platonic) characteristics could be used to determine a like number of races. By the 1960s, data and models from population genetics called into question taxonomic understandings of race, and many have turned from conceptualizing and analyzing human variation in terms of race to doing so in terms of populations and clines instead. That being said, many scientists still believe that race is a valid and useful concept. Moreover, since the 1990s, data and models from genomics and cladistics have resulted in a revolution in our understanding of human evolution, which has led some to propose a new "lineage" definition of race. These scientists have made related arguments that races are valid when understood as fuzzy sets, clusters, or extended families. Currently, opinions differ substantially within and among academic disciplines. 59. speciation - refers to the appearance of a new species of life on earth, particularly as seen in the fossil record. There are many ideas about the process leading to the creation of new species, each typically based on any of the Darwinian theories of biological evolution. Most of these ideas share the hypothesis that speciation occurs when a parent species (also referred to as a common ancestor) splits into two (or more) reproductively-isolated populations, each of which then accumulates changes from sexual reproduction and/or random mutation (in addition to any other various contributors to genetic change) until the populations are no longer capable of interbreeding (cladogenesis). (This is one definition of what a species is; see species.) If a single population of a species changes enough over time to be designated a new species while the old species dies out, we have a process called anagenesis. Among simpler forms of life, such as bacteria, single mutations can cause drastic changes (called "saltation") that result in speciation in a very short time. Speciation is also related to a process known as adaptive radiation. Ernst Mayr proposed a speciation mechanism called allopatry. Allopatry begins when subpopulations of a species become isolated geographically (for example, by habitat fragmentation or migration). The isolated populations are then liable to diverge evolutionarily over many generations as a) they become subjected to dissimilar selective pressures and b) they independently undergo genetic drift; particularly when one of the subpopulations is small (a scenario that leads to the "founder effect"). This kind of speciation is evident in many vertebrates' taxa. See also the issue of Ring species. Another proposed mechanism of speciation is sympatry, by which new species emerge alongside the old. This might occur if, say, subpopulations become dependent on different plants in the same area; and if variations in mating lead one subpopulation to become reproductively isolated from the other. Many examples of this kind of speciation are found in the invertebrates, especially the insects. Polyploidy is also a very common cause of sympatric speciation. Polyploidy is seen as a mode of speciation in many plants, a well studied example being that of the wheat species. In asexually reproducing organisms, there are other modes that become prominent including horizontal gene transfer by viruses and mutations. A further mechanism is parapatry, where the zones of two species abut but do not overlap. There is only partial separation afforded by geography, so individuals of each species may come in contact or cross the barrier from time to time. 60. microevolution- Microevolution is the occurrence of small-scale changes in gene frequencies in a population over a few generations, also known as change at or below the species level. These changes may be due to several processes: mutation, gene flow, genetic drift, as well as natural selection. Population genetics is the branch of biology that provides the mathematical structure for the study of the process of microevolution. Biologists distinguish between microevolution and macroevolution, which is the occurrence of large-scale changes in gene frequencies in a population over a long period of time (and may culminate in the evolution of new species). 61. macroevolution- Macroevolution is the concept that evolution of species and higher taxa is the result of large-scale changes in gene-frequencies over time. 62. phylogeny- A phylogeny (or phylogenesis) is the origin and evolution of a set of organisms, usually of a species. A major task of systematics is to determine the ancestral relationships among known species (both living and extinct), and the most commonly used methods to infer phylogenies include cladistics, phenetics, maximum likelihood, and Bayesian. During the late 19th century, the theory of recapitulation, or Haeckel's biogenetic law, was widely accepted. This theory was often expressed as "ontogeny recapitulates phylogeny", i.e. that the development of an organism exactly mirrors the evolutionary development of the species. The early version of this hypothesis has since been rejected as being oversimplified and misleading. However, modern biology recognizes numerous connections between ontogeny and phylogeny, explains them using evolutionary theory, and views them as supporting evidence for that theory. See the article on ontogeny and phylogeny 63. prezygotic isolating mechanisms- A type of reproductive isolation that occurs before the formation of a zygote can take place. In most cases mating does not even occur. Forms of prezygotic isolation include spatial, behavioral, mechanical and temporal isolation. (See 64-69) 64. geographic isolation -Geographic isolation is the physical separation of members of a population. Populations may be physically separated when their original habitat becomes divided, as, for example, when new land or water barriers form. Also, when part of a population colonizes anew, remote area such as an island, the colonizers geographically isolated from other populations of the species. For example, when a group of American finches colonized the Hawaiian islands, the group became geographically isolated from other populations of the species. these finches eventually gave rise to the 23 species of Hawaiian honeycreepers. Geographic isolation of a population may occur as a result of physical changes in an environment. When a river changes course or even when a highway is built across a field, populations may become geographically isolated. An example in which geographic isolation may have led to speciation. The desert of Death Valley, California, has a number of isolated ponds a formed by springs. Each pond contains a species of fish that lives only in that pond. Scientists suggest that these species arose through geographic isolation. Geologic evidence from a study of wave patterns in sedimentary rocks indicates that most of Death Valley was covered by a huge lake during the last ice age. When the ice age ended, the region became dry. Only small, spring fed ponds remained. Members of a fish species that previously formed a single population in the lake may have become isolated in different ponds. The environments of the isolated ponds differed enough that natural selection and perhaps genetic drift acted on the separate populations. Eventually the fish in the different ponds may have diverged so much genetically that they could no longer interbreed even if brought together. In this way geographic isolation of fishes in Death Valley probably led to the formation of new species. Geographic isolation, plus reproductive isolation, probably is the usual cause of the formation of new species. 65. ecological(habitat) isolation- For habitat isolation to occur, populations do not need to be separated by great distance. Instead, they must occupy different habitats, even within the same area. For example, if two populations of flies exist in the same geographical area, but one group lives in the soil and another lives on the surface of the water, members of the two populations are very unlikely to meet and reproduce. 66. temporal(seasonal) isolation- Temporal isolation represents another way in which populations living in the same area can be prevented from mating. Different populations may be ready to mate at different times of the year. For example, two populations of plants may produce flowers in different seasons, making mating between the populations impossible. 67. behavioral isolation- In many animals, courtship displays and rituals are vital to reproduction. Such behaviors can be very specific, varying between closely related species. Male behaviors such as courtship calls, songs, and dances will only be recognized by females of the same species. Some species of crickets are morphologically identical, but can be distinguished by the fact that females will only respond to the mating songs of males of their own species. Males of other species are ignored. 68. mechanical isolation- Mechanical isolation occurs when mating is physically impossible. Often this occurs because the genitalia of different species are incompatible. Bush babies, a group of small arboreal primates, are divided into several species based on mechanical isolation. Each species has distinctly shaped genitalia that, like locks and keys, only fit with the genitalia of its own species. 69. gametic isolation- In some cases, there are no barriers to mating, but the gametes are unable to fuse and fertilization does not take place. Often this occurs because the female immune system recognizes sperm as foreign and attacks it. In other cases, the sperm or pollen may not be adapted to the environment of the female reproductive track. 70. postzygotic isolating mechanisms- In some cases, no barriers exist to mating between members of different species. In these cases, the zygote formed is called a hybrid. However, even after a hybrid zygote forms, reproduction may still not be successful. When that reproduction is successful, living hybrids are usually themselves unable to reproduce. The production of hybrid offspring is costly: the energy of mating and producing offspring has still been spent by the parents, with no future inheritance of their genetic material in return. 71. developmental isolation- While gametes from different species can sometimes fuse to produce a hybrid zygote, these zygotes are frequently abnormal. Most do not survive to birth or germination. Those that do often do not develop normally and never reach sexual maturity. In these cases, the mating of different species is said to be unsuccessful even though offspring were produced because those offspring are incapable of passing on their genes. 72. hybrid inviability- Unlike mules, most hybrid offspring that survive the zygote stage to be born are not healthy. Most die before reaching reproductive age. As with the production of hybrids that cannot mature into reproductive adults, reproduction that results in unhealthy hybrids that die before reaching sexual maturity is considered unsuccessful. 73. hybrid sterility- Among those hybrid offspring that do develop normally and reach sexual maturity, most are sterile, meaning they do not produce viable gametes. For example, mules result from the mating of a horse with a donkey. They are born and develop into normal healthy adult animals, but they cannot produce offspring of their own. 74. allopatric speciation- A type of speciation which occurs when a population becomes segregated into two populations by some sort of geographic barrier (also called geographic speciation). This phenomenon is presumed to have been the mechanism whereby many species of organisms evolved 75. sympatric speciation- one of three theoretical models for the phenomenon of speciation. In complete contrast to allopatry, species undergoing sympatric speciation are not geographically isolated by, for example, a mountain or a river. The speciating populations share the same territory. This speciation phenomenon most commonly occurs through polyploidy, in which an offspring or group of offspring will be produced with twice the normal number of chromosomes. Where a normal individual has two copies of each chromosome (diploidy), these offspring may have four copies (tetraploidy). A tetraploid individual cannot mate with a diploid individual, creating reproductive isolation. Sympatric speciation is rare. It occurs more often among plants than animals, since it is so much easier for plants to self-fertilize than it is for animals. A tetraploidy plant can fertilize itself and create offspring. For a tetraploidy animal to reproduce, it must find another animal of the same species but of opposite sex that has also randomly undergone polyploidy. A number of models have been proposed to account for this mode of speciation. The most popular, disruptive speciation, was first put forward by John Maynard Smith in 1962. Smith suggested that heterozygous individuals may, under particular environmental conditions, have a greater fitness than those with alleles homozygous for a certain trait. Under the mechanism of natural selection, therefore, heterozygosity would be favoured over homozygosity, eventually leading to speciation. 76. polyploidy- Polyploid (in Greek: multiple) cells or organisms contain more than two copies (ploidy) of their chromosomes. The polyploid types are names triploid (3x), tetraploid (4x), pentaploid (5x), hexaploid (6x) and so on. Where an organism is normally diploid, a monoploid (1x) may arise as a spontaneous abberation, monoploidy may also occur as a normal stage in an organisms life cycle. Polyploidy occurs in animals but is especially common among flowering plants, including both wild and cultivated species. Wheat, for example, after millennia of hybridization and modification by humans, has strains that are diploid (two sets of chromosomes), tetraploid (four sets of chromosomes) with the common name of durum or macaroni wheat, and hexaploid (six sets of chromosomes) with the common name of bread wheat. Polyploidy can be induced in cell culture by some chemicals: the best known is colchicine, which causes chromosome doubling. -Examples 77. evolutionary bottleneck- see #40 78. punctuated equilibrium- a theory of evolution which states that changes such as speciation can occur relatively quickly, with long periods of little change—equilibria—in between. This theory is one of the proposed explanations of the evolutionary patterns of species as observed in the fossil record, particularly the relatively sudden appearance of new species in a geologically short time period, and the perhaps typical lack of substantial change of species during their existence. 79. gradualism- Gradualism, in biology, holds that evolution occurs through the accumulation of slight modifications over a period of generations. This is in contrast to saltationism, which holds that evolution occurs through large changes, possibly in a single generation. Gradualism holds that every individual is the same species as its parents, and that there is no clear line of demarcation between the old species and the new species. It holds that the species is not a fixed type, and that it is the population, not the individual, that evolves. "Support for gradualism derives from the fact that many, probably most, evolutionary processes are indeed gradual and incremental....However, it is one thing to believe that gradual processes predominate in nature and quite another to hold that all evolutionary processes must be gradual. The issue is, after all, simply an empirical one; even if no nongradual changes were ever witnessed, one could never exclude the possibility that the next evolutionary process to be uncovered might be nongradual." Examples include sudden adaptation due to natural disaster, preadaptations and exaptations (Gould and Vrba 1982), and possibly, syntax and language, which being hierarchical could not have been gradual, "the difference between flat structure (beads on a wire) and hierarchyical structure is absolute" (Bickerton 2000, p.159). One example of preadaptation is insect flight as insects were originally exclusively terrestrial, some with fanlike structures for cooling which were selected for until they were large and efficient enough to allow flight (ibid, p.160). 80. Geologic time chart- see geologic time 81. index fossils- Index fossils (or zone fossils) are fossils used to define and identify geologic periods (or faunal stages). They work on the premise that although different sediments may look different depending on the conditions under which they were laid down, they may include the remains of the same species of fossil. If the species concerned were short-lived (in geological terms, lasting a few hundred thousand years) then you can be sure that the sediments in question were deposited within that narrow time period. The shorter the lifespan of a species, the more precisely you can correlate different sediments, and so rapidly evolving types of fossils are particularly valuable. The best index fossils are common, easy-to-identify at species level, and have a broad distribution—otherwise the likelihood of finding and recognising one in the two sediments is low. Ammonites fit these demands well, and are the best-known fossils that have been widely used for this. Other important groups that provide index fossils are the corals, graptolites, brachiopods, trilobites, and echinoids (sea urchins). Conodonts may be identified by experts using light microscopy such that they can be used to index a given sample with good resolution. Terrestrial fossils, such as the teeth of mammals, have also been used. 82. radioactive dating- a technique used to date materials based on a knowledge of the decay rates of naturally occurring isotopes, and the current abundances. 83. phylogeny- A phylogeny (or phylogenesis) is the origin and evolution of a set of organisms, usually of a species. A major task of systematics is to determine the ancestral relationships among known species (both living and extinct), and the most commonly used methods to infer phylogenies include cladistics, phenetics, maximum likelihood, and Bayesian. A phylogenetic tree is a tree showing the evolutionary interrelationships among various species or other entities that are believed to have a common ancestor. A phylogenetic tree is a form of a cladogram. In a phylogenetic tree, each node with descendants represents the most recent common ancestor of the descendants, and edge lengths correspond to time estimates. Each node in a phylogenetic tree is called a taxonomic unit. Internal nodes are generally referred to as Hypothetical Taxonomic Units (HTUs) as they cannot be directly observed. A rooted phylogenetic tree is a directed tree with a unique node corresponding to the (usually imputed) most recent common ancestor of all the entities at the leaves of the tree. Figure 1 depicts a rooted phylogenetic tree, which has been colored according to the three-domain system (Woese 1998). An unrooted phylogenetic tree is, loosely speaking, a tree derived from a rooted phylogenetic tree by omitting the root. More precisely, it is a forest of rooted phylogenetic trees depicted so that the roots are all linked. Figure 2 depicts an unrooted phylogenetic tree¹ for myosin, a superfamily of proteins 84. systematics- Systematics is the study of the diversity of organism characteristics. In biology, systematists are the scientists who classify species and other taxa, which they do with the aim of defining how they relate evolutionarily. 85a. convergent evolution- In evolutionary biology, convergent evolution describes the process whereby organisms not closely related independently acquire similar characteristics while evolving in separate and sometimes varying ecosystems. An example of convergent evolution is the similar nature of the wings of insects, birds, and bats. All three serve the same function and are similar in structure, but each evolved independently. Eyes also evolved independently in various animals. Convergent evolution is a different phenomenon than evolutionary relay and parallel evolution. Similar to convergent evolution, evolutionary relay describes how independent species acquire similar characteristics through their evolution in similar ecosystems, but not at the same time (dorsal fins of extinct ichthyosaurs and sharks). Parallel evolution occurs when two independent species evolve together at the same time in the same ecospace and acquire similar characteristics (extinct browsing-horses and extinct paleotheres). Structures that are the result of convergent evolution are called analogous structures or homoplasies; they should be contrasted with homologous structures which have a common origin. 85b. divergent evolution- Pattern of evolution in which two closely related species gradually become more and more dissimilar. Compare with convergent evolution and parallel evolution. Divergent evolution, when organisms evolve to fill more diverse ecological niches than ancestral forms is probably much more common in the history of life than convergent evolution. In the broadest sense, all living things are the product of divergent evolution. Most scientists believe that all organisms are in fact descended from one population of early single celled organisms. However, in this essay we will begin to look at divergent evolution within the scope of smaller taxonomic groups, mostly orders and families. The way to recognize divergent evolution on any scale is that at the base taxonomic level the animals share characteristics that link them, but they possess superficial differences that allow them to exploit different ecological niches. This is most likely due to differential selection on a relatively short time scale. 86. DNA hybridization- The technique of DNA-DNA hybridization provides a way of comparing the total genome of two species. Let us examine the procedure as it might be used to assess the evolutionary relationship of species B to species A: The total DNA is extracted from the cells of each species and purified. For each, the DNA is heated so that it becomes denatured into single strands (ssDNA). The temperature is lowered just enough to allow the multiple short sequences of repetitive DNA to rehybridize back into double-stranded DNA (dsDNA). The mixture of ssDNA (representing single genes) and dsDNA (representing repetitive DNA) is passed over a column packed with hydroxyapatite. The dsDNA sticks to the hydroxyapatite; ssDNA does not and flows right through. The purpose of this step is to be able to compare the information-encoding portions of the genome — mostly genes present in a single copy — without having to worry about varying amounts of noninformative repetitive DNA. The ssDNA of species A is made radioactive. The radioactive ssDNA is then allowed to rehybridize with nonradioactive ssDNA of the same species (A) as well as — in a separate tube — the ssDNA of species B. After hybridization is complete, the mixtures (A/A) and (A/B) are individually heated in small (2°–3°C) increments. At each higher temperature, an aliquot is passed over hydroxyapatite. Any radioactive strands (A) that have separated from the DNA duplexes pass through the column, and the amount is measured from their radioactivity. A graph showing the percentage of ssDNA at each temperature is drawn. The temperature at which 50% of the DNA duplexes (dsDNA) have been denatured (T50H) is determined. As the figure shows, the curve for A/B is to the left of A/A, i.e., duplexes of A/B separated at a lower temperature than those of A/A. The sequences of A/A are precisely complementary so all the hydrogen bonds between complementary base pairs (A-T, C-G) must be broken in order to separate the strands. But where the gene sequences in B differ from those in A, no base pairing will have occurred and denaturation is easier. Thus DNA-DNA hybridization provides genetic comparisons integrated over the entire genome. Its use has cleared up several puzzling taxonomic relationships. 87. molecular clocks Questions 1. Explain Charles Darwin's contributions to evolutionary ideas. Give the basic assumptions upon which Darwin's theory rests. Indicate the types of evidence that Darwin used in formulating his theory. 2. List the processes that can lead to genetic variation. 3. Explain why changes in somatic cells cannot bring about evolution. 4. Briefly contrast Lamarck's theory of inheritance of acquired characteristics with Darwin's theory of natural selection. 5. Explain the concept of the gene pool. Given the frequencies of two alleles, be able calculate the ratios of the genotypes produced by them, using a Punnett square or algebraic method. 6. State the Hardy-Weinberg Law, and discuss its four conditions for maintenance of genetic equilibrium. Why are these conditions rarely met in nature? 7. Explain how natural selection in one generation can affect the genotype of the next generation. 8. Using diagrams contrast directional selection, stabilizing selection and disruptive selection. 9. Describe heterozygote superiority, explain how this process helps maintain deleterious recessive genes. (Note sickle cell anemia) 10. Give a biological definition of species. 11. Explain the geographic isolation model of speciation, in doing so take into account the roles of mutation, natural selection, and the gene pool. Distinguish between extrinsic and intrinsic isolating mechanisms. 12. Explain what is meant by adaptive radiation, and discuss the evidence for this phenomenon. 13. Compare the hypotheses of gradualism and punctuated equilibrium, and give an example supporting each hypothesis.