18.1: Understanding Evolution

18.1.1: What is Evolution?

Learning Objective

Describe the historical influences on Darwin's theory of evolution

Key Points

• Ancient Greeks expressed ideas about evolution, which were reintroduced in the eighteenth century by Georges-Louis Leclerc Comte de Buffon who observed different environments had different plant and animal populations.
• James Hutton proposed that geological changes occur gradually over time via the accumulation of small changes rather than through large catastrophic events.
• Charles Lyell popularized James Hutton's theory; this theory of incremental change influenced Darwin's theory of evolution.
• Jean-Baptiste Lamarck proposed the theory of the inheritance of acquired characterstics; this theory has now been discredited, but it served as an important influence on the theory of evolution.

Key Terms

evolution

the change in the genetic composition of a population over successive generations

inheritance of acquired characteristics

hypothesis that physiological changes acquired over the life of an organism may be transmitted to its offspring

Introduction: Evolution

All species of living organisms, including bacteria and chimpanzees, evolved at some point from a different species. Although it may seem that living things today stay the same, this is not the case: evolution is a gradual and ongoing process.

The theory of evolution is the unifying theory of biology, meaning it is the framework within which biologists ask questions about the living world. The Ukrainian-born American geneticist Theodosius Dobzhansky famously wrote that "nothing makes sense in biology except in the light of evolution." The tenet that all species have evolved and diversified from a common ancestor is the foundation from which we approach all questions in biology. It provides a direction for predictions about living things, which has been validated through extensive scientific experimentation.

Evolution by natural selection describes a mechanism for the change of species over time . Well before Darwin began to explore the concept of evolution, the idea that species change over time had already been suggested and debated. The view that species are static and unchanging was grounded in the writings of Plato, yet there were also ancient Greeks who expressed ideas about evolution. During the eighteenth century, ideas about the evolution of animals were reintroduced by the naturalist Georges-Louis Leclerc Comte de Buffon who observed that various geographic regions have different plant and animal populations, even when the environments are similar. It was also accepted that there are extinct species.

Evolution by Natural Selection

All organisms are products of evolution adapted to their environment. (a) Saguaro (Carnegiea gigantea) can soak up 750 liters of water in a single rain storm, enabling these cacti to survive the dry conditions of the Sonoran desert in Mexico and the Southwestern United States. (b) The Andean semiaquatic lizard (Potamites montanicola), discovered in Peru in 2010, lives between 1,570 to 2,100 meters in elevation and, unlike most lizards, is nocturnal and swims. Scientists still do not know how these cold-blood animals are able to move in the cold (10 to 15°C) temperatures of the Andean night.

During this time, a Scottish naturalist named James Hutton proposed that geological change occurs gradually by the accumulation of small changes over long periods of time. This theory contrasted with the predominant view of the time: that the geology of the planet is a consequence of catastrophic events that occurred during a relatively brief past. During the nineteenth century, Hutton's views were popularized by the geologist Charles Lyell, who was a friend of Charles Darwin. Lyell's ideas, in turn, influenced Darwin's concept of evolution. The greater age of the earth proposed by Lyell supported the gradual evolution that Darwin proposed, and the slow process of geological change provided an analogy for the gradual change in species.

In the early nineteenth century, Jean-Baptiste Lamarck published a book that detailed a different mechanism for evolutionary change. This mechanism is now referred to as an inheritance of acquired characteristics. This idea states that modifications in an individual are caused by its environment, or the use or disuse of a structure during its lifetime, and that these changes can be inherited by its offspring, bringing about change in a species. While this mechanism for evolutionary change was discredited, Lamarck's ideas were an important influence on the concept of evolution.

18.1.2: Charles Darwin and Natural Selection

Learning Objective

Explain how natural selection can lead to evolution

Key Points

• Wallace traveled to Brazil to collect and observe insects from the Amazon rainforest.
• Darwin observed that finches in the Galápagos Islands had different beaks than finches in South America; these adaptations equiped the birds to acquire specific food sources.
• Wallace and Darwin observed similar patterns in the variation of organisms and independently developed the same explanation for how such variations could occur over time, a mechanism Darwin called natural selection.
• According to natural selection, also known as "survival of the fittest," individuals with traits that enable them to survive are more reproductively successful; this leads to those traits becoming predominant within a population.
• Natural selection is an inevitable outcome of three principles: most characteristics are inherited, more offspring are produced than are able to survive, and offspring with more favorable characteristics will survive and have more offspring than those individuals with less favorable traits.

Key Terms

natural selection

a process in which individual organisms or phenotypes that possess favorable traits are more likely to survive and reproduce

descent with modification

change in populations over generations

Example

• One of the best examples of evolution by natural selection has been demonstrated in the birds that helped to inspire Darwin's theory: Galápagos finches. Peter and Rosemary Grant and their colleagues have studied Galápagos finch populations every year since 1976. The Grants found changes from one generation to the next in the distribution of beak shapes with the medium ground finch on the Galápagos island of Daphne Major. The birds have inherited variation in the bill shape with some birds having wide deep bills and others having thinner bills. During a period in which rainfall was higher than normal because of an El Niño, the large hard seeds that large-billed birds ate were reduced in number; however, there was an abundance of the small soft seeds which the small-billed birds ate. Therefore, survival and reproduction were much better in the following years for the small-billed birds. In the years following this El Niño, the Grants measured beak sizes in the population and found that the average bill size was smaller. Since bill size is an inherited trait, parents with smaller bills had more offspring and the size of bills had evolved to be smaller. As conditions improved in 1987 and larger seeds became more available, the trend toward smaller average bill size ceased.

Charles Darwin and Natural Selection

In the mid-nineteenth century, the mechanism for evolution was independently conceived of and described by two naturalists: Charles Darwin and Alfred Russel Wallace. Importantly, each naturalist spent time exploring the natural world on expeditions to the tropics. From 1831 to 1836, Darwin traveled around the world to places like South America, Australia, and the southern tip of Africa. Wallace traveled to Brazil to collect insects in the Amazon rainforest from 1848 to 1852 and to the Malay Archipelago from 1854 to 1862. Darwin's journey, as with Wallace's later journeys to the Malay Archipelago, included stops at several island chains, the last being the Galápagos Islands west of Ecuador. On these islands, Darwin observed that species of organisms on different islands were clearly similar, yet had distinct differences. For example, the ground finches inhabiting the Galápagos Islands comprised several species with a unique beak shape . The species on the islands had a graded series of beak sizes and shapes with very small differences between the most similar. He observed that these finches closely resembled another finch species on the mainland of South America. Darwin imagined that the island species might be modified from one of the original mainland species. Upon further study, he realized that the varied beaks of each finch helped the birds acquire a specific type of food. For example, seed-eating finches had stronger, thicker beaks for breaking seeds, while insect-eating finches had spear-like beaks for stabbing their prey.

Beak Shape Among Finch Species

Darwin observed that beak shape varies among finch species. He postulated that the beak of an ancestral species had adapted over time to equip the finches to acquire different food sources.

Natural Selection

Wallace and Darwin observed similar patterns in other organisms and independently developed the same explanation for how and why such changes could take place. Darwin called this mechanism natural selection. Natural selection, also known as "survival of the fittest," is the more prolific reproduction of individuals with favorable traits that survive environmental change because of those traits. This leads to evolutionary change, the trait becoming predominant within a population. For example, Darwin observed that a population of giant tortoises found in the Galapagos Archipelago have longer necks than those that lived on other islands with dry lowlands. These tortoises were "selected" because they could reach more leaves and access more food than those with short necks. In times of drought, when fewer leaves would be available, those that could reach more leaves had a better chance to eat and survive than those that could not reach the food source. Consequently, long-necked tortoises would more probably be reproductively successful and pass the long-necked trait to their offspring. Over time, only long-necked tortoises would be present in the population.

Natural selection, Darwin argued, was an inevitable outcome of three principles that operated in nature. First, most characteristics of organisms are inherited, or passed from parent to offspring, although how traits were inherited was unknown. Second, more offspring are produced than are able to survive. The capacity for reproduction in all organisms outstrips the availability of resources to support their numbers. Thus, there is competition for those resources in each generation. Both Darwin and Wallace were influenced by an essay written by economist Thomas Malthus who discussed this principle in relation to human populations. Third, Darwin and Wallace reasoned that offspring with the inherited characteristics that allow them to best compete for limited resources will survive and have more offspring than those individuals with variations that are less able to compete. Because characteristics are inherited, these traits will be better represented in the next generation. This will lead to change in populations over successive generations in a process that Darwin called descent with modification. Ultimately, natural selection leads to greater adaptation of the population to its local environment; it is the only mechanism known for adaptive evolution.

Papers by Darwin and Wallace presenting the idea of natural selection were read together in 1858 before the Linnean Society in London. The following year, Darwin's book, On the Origin of Species, was published. His book outlined his arguments for evolution by natural selection.

Charles Darwin and Alfred Wallace

Both (a) Charles Darwin and (b) Alfred Wallace wrote scientific papers on natural selection that were presented together before the Linnean Society in 1858.

18.1.3: The Galapagos Finches and Natural Selection

Learning Objective

Describe how finches provide visible evidence of evolution

Key Points

• Darwin observed the Galapagos finches had a graded series of beak sizes and shapes and predicted these species were modified from one original mainland species.
• Darwin called differences among species natural selection, which is caused by the inheritance of traits, competition between individuals, and the variation of traits.
• Offspring with inherited characteristics that allow them to best compete will survive and have more offspring than those individuals with variations that are less able to compete.
• Large-billed finches feed more efficiently on large, hard seeds, whereas smaller billed finches feed more efficiently on small, soft seeds.
• When small, soft seeds become rare, large-billed finches will survive better, and there will be more larger-billed birds in the following generation; when large, hard seeds become rare, the opposite will occur.

Key Terms

evolution

the change in the genetic composition of a population over successive generations

natural selection

a process in which individual organisms or phenotypes that possess favorable traits are more likely to survive and reproduce

Visible Evidence of Ongoing Evolution: Darwin's Finches

From 1831 to 1836, Darwin traveled around the world, observing animals on different continents and islands. On the Galapagos Islands, Darwin observed several species of finches with unique beak shapes. He observed these finches closely resembled another finch species on the mainland of South America and that the group of species in the Galápagos formed a graded series of beak sizes and shapes, with very small differences between the most similar . Darwin imagined that the island species might be all species modified from one original mainland species. In 1860, he wrote, "seeing this gradation and diversity of structure in one small, intimately related group of birds, one might really fancy that from an original paucity of birds in this archipelago, one species had been taken and modified for different ends."

Darwin's Finches

Darwin observed that beak shape varies among finch species. He postulated that the beak of an ancestral species had adapted over time to equip the finches to acquire different food sources. This illustration shows the beak shapes for four species of ground finch: 1. Geospiza magnirostris (the large ground finch), 2. G. fortis (the medium ground finch), 3. G. parvula (the small tree finch), and 4. Certhidea olivacea (the green-warbler finch).

Natural Selection

Darwin called this mechanism of change natural selection. Natural selection, Darwin argued, was an inevitable outcome of three principles that operated in nature. First, the characteristics of organisms are inherited, or passed from parent to offspring. Second, more offspring are produced than are able to survive; in other words, resources for survival and reproduction are limited. The capacity for reproduction in all organisms exceeds the availability of resources to support their numbers. Thus, there is a competition for those resources in each generation. Third, offspring vary among each other in regard to their characteristics and those variations are inherited. Out of these three principles, Darwin reasoned that offspring with inherited characteristics that allow them to best compete for limited resources will survive and have more offspring than those individuals with variations that are less able to compete. Because characteristics are inherited, these traits will be better represented in the next generation. This will lead to change in populations over generations in a process that Darwin called "descent with modification," or evolution.

Studies of Natural Selection After Darwin

Demonstrations of evolution by natural selection can be time consuming. Peter and Rosemary Grant and their colleagues have studied Galápagos finch populations every year since 1976 and have provided important demonstrations of the operation of natural selection. The Grants found changes from one generation to the next in the beak shapes of the medium ground finches on the Galápagos island of Daphne Major.

The medium ground finch feeds on seeds. The birds have inherited variation in the bill shape with some individuals having wide, deep bills and others having thinner bills. Large-billed birds feed more efficiently on large, hard seeds, whereas smaller billed birds feed more efficiently on small, soft seeds. During 1977, a drought period altered vegetation on the island. After this period, the number of seeds declined dramatically; the decline in small, soft seeds was greater than the decline in large, hard seeds. The large-billed birds were able to survive better than the small-billed birds the following year.

The year following the drought when the Grants measured beak sizes in the much-reduced population, they found that the average bill size was larger . This was clear evidence for natural selection of bill size caused by the availability of seeds. The Grants had studied the inheritance of bill sizes and knew that the surviving large-billed birds would tend to produce offspring with larger bills, so the selection would lead to evolution of bill size. Subsequent studies by the Grants have demonstrated selection on and evolution of bill size in this species in response to other changing conditions on the island. The evolution has occurred both to larger bills, as in this case, and to smaller bills when large seeds became rare.

Finches of Daphne Major

A drought on the Galápagos island of Daphne Major in 1977 reduced the number of small seeds available to finches, causing many of the small-beaked finches to die. This caused an increase in the finches' average beak size between 1976 and 1978.

18.1.4: Processes and Patterns of Evolution

Learning Objective

Explain why only heritable variation can be acted upon by natural selection

Key Points

• Genetic variation within a population is a result of mutations and sexual reproduction.
• A mutation may be neutral, reduce an organism's fitness, or increase an organism's fitness.
• An adaptation is a heritable trait that increases the survival and rate of reproduction of an organism in its present environment.
• Divergent evolution describes the process in which two species evolve in diverse directions from a common point.
• Convergent evolution is the process in which similar traits evolve independently in species that do not share a recent common ancestry.

Key Terms

divergent evolution

the process by which a species with similar traits become groups that are tremendously different from each other over many generations

adaptation

modification of something or its parts that makes it more fit for existence under the conditions of its current environment

convergent evolution

a trait of evolution in which species not of similar recent origin acquire similar properties due to natural selection

Processes and Patterns of Evolution

Variation

Natural selection can only take place if there is variation, or differences, among individuals in a population. Importantly, these differences must have some genetic basis; otherwise, the selection will not lead to change in the next generation. This is critical because variation among individuals can be caused by non-genetic reasons, such as an individual being taller due to better nutrition rather than different genes.

Genetic diversity within a population comes from two main mechanisms: mutation and sexual reproduction. Mutation, a change in the DNA sequence, is the ultimate source of new alleles, or new genetic variation in any population. The genetic changes caused by mutation can have one of three outcomes:

• Many mutations will have no effect on the fitness of the phenotype; these are called neutral mutations.
• A mutation may affect the phenotype of the organism in a way that gives it reduced fitness (a lower likelihood of survival or fewer offspring).
• A mutation may produce a phenotype with a beneficial effect on fitness. Different mutations will have a range of effects on the fitness of an organism that expresses them in their phenotype, from a small effect to a great effect.

Sexual reproduction also leads to genetic diversity: when two parents reproduce, unique combinations of alleles assemble to produce the unique genotypes and thus phenotypes in each of the offspring. However, sexual reproduction can not lead to new genes, but rather provides a new combination of genes in a given individual.

Adaptations

A heritable trait that aids the survival and reproduction of an organism in its present environment is called an adaptation. Scientists describe groups of organisms becoming adapted to their environment when a change in the range of genetic variation occurs over time that increases or maintains the "fitness" of the population to its environment. The webbed feet of platypuses are an adaptation for swimming. The snow leopards' thick fur is an adaptation for living in the cold. The cheetahs' fast speed is an adaptation for catching prey.

Whether or not a trait is favorable depends on the environmental conditions at the time. The same traits are not always selected because environmental conditions can change. For example, consider a species of plant that grew in a moist climate and did not need to conserve water. Large leaves were selected because they allowed the plant to obtain more energy from the sun. Large leaves require more water to maintain than small leaves, and the moist environment provided favorable conditions to support large leaves. After thousands of years, the climate changed and the area no longer had excess water. The direction of natural selection shifted so that plants with small leaves were selected because those populations were able to conserve water to survive the new environmental conditions.

The evolution of species has resulted in enormous variation in form and function. Sometimes, evolution gives rise to groups of organisms that become tremendously different from each other. When two species evolve in diverse directions from a common point, it is called divergent evolution. Such divergent evolution can be seen in the forms of the reproductive organs of flowering plants which share the same basic anatomies; however, they can look very different as a result of selection in different physical environments and adaptation to different kinds of pollinators .

Flowering Plants

Flowering plants evolved from a common ancestor. Notice that the (a) dense blazing star (Liatrus spicata) and the (b) purple coneflower (Echinacea purpurea) vary in appearance, yet both share a similar basic morphology.

In other cases, similar phenotypes evolve independently in distantly-related species. For example, flight has evolved in both bats and insects; they both have structures we refer to as wings, which are adaptations to flight. However, the wings of bats and insects have evolved from very different original structures. This phenomenon is called convergent evolution, where similar traits evolve independently in species that do not share a recent common ancestry. The two species came to the same function, flying, but did so separately from each other.

These physical changes occur over enormous spans of time and help explain how evolution occurs. Natural selection acts on individual organisms, which in turn can shape an entire species. Although natural selection may work in a single generation on an individual, it can take thousands or even millions of years for the genotype of an entire species to evolve. It is over these large time spans that life on earth has changed and continues to change.

18.1.5: Evidence of Evolution

Learning Objective

Explain the development of the theory of evolution

Key Points

• Fossils serve to highlight the differences and similarities between current and extinct species, showing the evolution of form over time.
• Similar anatomy across different species highlights their common origin and can be seen in homologous and vestigial structures.
• Embryology provides evidence for evolution since the embryonic forms of divergent groups are extremely similar.
• The natural distribution of species across different continents supports evolution; species that evolved before the breakup of the supercontinent are distributed worldwide, whereas species that evolved more recently are more localized.
• Molecular biology indicates that the molecular basis for life evolved very early and has been maintained with little variation across all life on the planet.

Key Terms

vestigial structure

genetically determined structures or attributes that have apparently lost most or all of their ancestral function in a given species

biogeography

the study of the geographical distribution of living things

homologous structure

the traits of organisms that result from sharing a common ancestor; such traits often have similar embryological origins and development

Evidence of Evolution

The evidence for evolution is compelling and extensive. Looking at every level of organization in living systems, biologists see the signature of past and present evolution. Darwin dedicated a large portion of his book, On the Origin of Species, to identifying patterns in nature that were consistent with evolution. Since Darwin, our understanding has become clearer and broader.

Fossils, Anatomy, and Embryology

Fossils provide solid evidence that organisms from the past are not the same as those found today; they show a progression of evolution. Scientists calculate the age of fossils and categorize them to determine when the organisms lived relative to each other. The resulting fossil record tells the story of the past and shows the evolution of form over millions of years. For example, scientists have recovered highly-detailed records showing the evolution of humans and horses . The whale flipper shares a similar morphology to appendages of birds and mammals, indicating that these species share a common ancestor . Over time, evolution led to changes in the shapes and sizes of these bones in different species, but they have maintained the same overall layout. Scientists call these synonymous parts homologous structures.

Common Ancestors

The similar construction of these appendages indicates that these organisms share a common ancestor.

Evolution of Humans and Horses

(a) In this display, fossil hominids are arranged from oldest (bottom) to newest (top). As hominids evolved, the shape of the skull changed. (b) An artist's rendition of extinct species of the genus Equus reveals that these ancient species resembled the modern horse (Equus ferus), but varied in size.

Some structures exist in organisms that have no apparent function at all, appearing to be residual parts from a common ancestor. These unused structures (such as wings on flightless birds, leaves on some cacti, and hind leg bones in whales) are vestigial.

Embryology, the study of the development of the anatomy of an organism to its adult form, provides evidence for evolution as embryo formation in widely-divergent groups of organisms tends to be conserved. Structures that are absent in the adults of some groups often appear in their embryonic forms, disappearing by the time the adult or juvenile form is reached. For example, all vertebrate embryos, including humans, exhibit gill slits and tails at some point in their early development. These disappear in the adults of terrestrial groups, but are maintained in adults of aquatic groups, such as fish and some amphibians. Great ape embryos, including humans, have a tail structure during their development that is lost by birth.

Another form of evidence of evolution is the convergence of form in organisms that share similar environments. For example, species of unrelated animals, such as the arctic fox and ptarmigan living in the arctic region, have been selected for seasonal white phenotypes during winter to blend with the snow and ice . These similarities occur not because of common ancestry, but because of similar selection pressures: the benefits of not being seen by predators.

Adaptations: Winter Coats

The white winter coat of the (a) arctic fox and the (b) ptarmigan's plumage are adaptations to their environments.

Biogeography

The geographic distribution of organisms on the planet follows patterns that are best explained by evolution in conjunction with the movement of tectonic plates over geological time. Broad groups that evolved before the breakup of the supercontinent Pangaea (about 200 million years ago) are distributed worldwide. Groups that evolved since the breakup appear uniquely in regions of the planet, such as the unique flora and fauna of northern continents that formed from the supercontinent Laurasia compared to that of the southern continents that formed from the supercontinent Gondwana.

The great diversification of marsupials in Australia and the absence of other mammals reflect Australia's long isolation. Australia has an abundance of endemic species (those found nowhere else) which is typical of islands whose isolation by expanses of water prevents species from migrating. Over time, these species diverge evolutionarily into new species that look very different from their ancestors that may exist on the mainland. The marsupials of Australia, the finches on the Galápagos, and many species on the Hawaiian Islands are all unique to their one point of origin, yet they display distant relationships to ancestral species on mainlands.

Molecular Biology

Like anatomical structures, the structures of the molecules of life reflect descent with modification. Evidence of a common ancestor for all of life is reflected in the universality of DNA as the genetic material, in the near universality of the genetic code, and in the machinery of DNA replication and expression. In general, the relatedness of groups of organisms is reflected in the similarity of their DNA sequences. This is exactly the pattern that would be expected from descent and diversification from a common ancestor.

DNA sequences have also shed light on some of the mechanisms of evolution. For example, it is clear that the evolution of new functions for proteins commonly occurs after gene duplications that allow the free modification of one copy by mutation, selection, or drift (changes in a population's gene pool resulting from chance), while the second copy continues to produce a functional protein.

18.1.6: Misconceptions of Evolution

Learning Objective

Discuss misconceptions about the theory of evolution

Key Points

• Attacks on the theory of evolution sometimes take issue with the word "theory", which in the vernacular means a guess or suggested explanation. In scientific language, "theory" indicates a body of thoroughly-tested and verified explanations for a set of observations of the natural world.
• Evolution does not take place on an individual level; evolution is the average change of a characteristic within an entire population.
• Evolution does not explain the origin of life; the theory of evolution instead explains how populations change over time and how traits are selected in order to increase the fitness of a population.
• Favorable traits do not arise as a result of the environment as these traits are already present; individuals with favorable traits are more likely to survive and, thus, will have greater fitness than individuals with less desirable traits.
• Evolution and natural selection are not synonymous. Natural selection is just one mechanism by which evolution occurs.

Key Term

theory

a well-substantiated explanation of some aspect of the natural world based on knowledge that has been repeatedly confirmed through observation and experimentation

Example

• Lamarckian inheritance is the idea that an organism can pass on characteristics that it acquired during its lifetime to its offspring (also known as heritability of acquired characteristics or soft inheritance). It is named after the French biologist Jean-Baptiste Lamarck (1744–1829), who incorporated the action of soft inheritance into his evolutionary theories. Mendelian genetics supplanted the notion of inheritance of acquired traits, eventually leading to the development of the modern evolutionary synthesis and the general abandonment of the Lamarckian theory of evolution in biology. Despite this abandonment, interest in Lamarckism has continued as studies in the field of epigenetics have highlighted the possible inheritance of behavioral traits acquired by the previous generation.

Misconceptions of Evolution

Although the theory of evolution generated controversy when it was first proposed, it was almost universally accepted by biologists within 20 years of the publication of On the Origin of Species. Nevertheless, the theory of evolution is a difficult concept and misconceptions about it abound.

Evolution is Just a Theory

Critics of the theory of evolution dismiss its importance by purposefully confounding the everyday usage of the word "theory" with the way scientists use the word. In science, a "theory" is understood to be a body of thoroughly-tested and verified explanations for a set of observations of the natural world. Scientists have a theory of the atom, a theory of gravity, and the theory of relativity, each of which describes understood facts about the world. In the same way, the theory of evolution describes facts about the living world. A theory in science has also survived significant efforts to discredit it by scientists. In contrast, a "theory" in common vernacular is a word meaning a guess or suggested explanation; this meaning is more akin to the scientific concept of "hypothesis. " When critics of evolution say evolution is "just a theory," they are implying that there is little evidence supporting it and that it is still in the process of being rigorously tested. This is a mis-characterization.

Individuals Evolve

Evolution is the change in genetic composition of a population over time, specifically over generations, resulting from differential reproduction of individuals with certain alleles. Individuals do change over their lifetime, obviously, but this is called development and involves changes programmed by the set of genes the individual acquired at birth in coordination with the individual's environment. When thinking about the evolution of a characteristic, it is probably best to think about the change of the average value of the characteristic in the population over time. For example, when natural selection leads to bill-size change in medium-ground finches in the Galápagos, this does not mean that individual bills on the finches are changing. If one measures the average bill size among all individuals in the population at one time and then measures the average bill size in that population several years later, this average value of the population will be different as a result of evolution.

Evolution Explains the Origin of Life

It is a common misunderstanding that evolution includes an explanation of life's origins. The theory of evolution explains how populations change over time. It does not shed light on the beginnings of life, including the origins of the first cells, which is how life is defined. The mechanisms of the origin of life on earth are a particularly difficult problem because it occurred a very long time ago and, presumably, it occurred just once. However, while evolution does not explain the origin of life, it may have something to say about some of the processes operating once pre-living entities acquired certain properties. Once a mechanism of inheritance was in place in the form of a molecule like DNA, either within a cell or pre-cell, these entities would be subject to the principle of natural selection. More effective reproducers would increase in frequency at the expense of inefficient reproducers.

Organisms Evolve on Purpose

Statements such as "organisms evolve in response to a change in an environment" may lead to the misunderstanding that evolution is somehow intentional. A changed environment results in some individuals in the population, those with particular phenotypes, benefiting and, therefore, producing proportionately more offspring than other phenotypes. This results in change in the population if the characteristics are genetically determined.

It is important to understand that the variation that natural selection works on is already present in a population and does not arise in response to an environmental change. For example, applying antibiotics to a population of bacteria will, over time, select a population of bacteria that are resistant to antibiotics. The resistance, which is caused by a gene, did not arise by mutation because of the application of the antibiotic. The gene for resistance was already present in the gene pool of the bacteria, probably at a low frequency. The antibiotic, which kills the bacterial cells without the resistance gene, strongly selects individuals that are resistant, since these would be the only ones that survived and divided. Experiments have demonstrated that mutations for antibiotic resistance do not arise as a result of antibiotics.

In a larger sense, evolution is not goal directed. Species do not become "better" over time; they track their changing environment with adaptations that maximize their reproduction. The characteristics that evolve in a species are a function of preexisting variation and the environment, both of which are constantly changing non-directionally. A trait that is fit in one environment at one time may also be fatal at some point in the future.

Evolution = Natural Selection

The terms “evolution” and “natural selection” are often conflated, as the two concepts are closely related. They are not, however, synonymous. Natural selection refers to the process by which organisms better suited for their environment are more likely to survive and produce offspring, thereby proliferating those favorable genetics in a population. Evolution is defined more broadly as any change in the genetic makeup of a population over time. As expounded by Darwin, natural selection is a major driving force of evolution, but it is not the only one. 

Genetic drift, for example, is another mechanism by which evolution may occurs. Genetic drift occurs when allelic frequency is altered due to random sampling. It is evolution by chance, and the smaller the population, the more significant the effects on genetic distribution due to sampling error. For example, a population bottleneck, which occurs when an event such as a natural disaster dramatically reduces the size of a population, can result in the elimination or significant reduction of a trait within a population, regardless of how beneficial that trait may be to survival or reproduction. Thus evolution can occur without natural selection.

Misconceptions of Evolution: Lamarckian evolution and giraffes

Lamarck's hypothesis for how evolution worked stated that organisms could pass on traits they had acquired during their lifetime to the progeny, such as giraffes gaining a long neck because their parents needed to reach high food.

Attributions

• What is Evolution?
  • "Boundless." http://www.boundless.com/. Boundless Learning CC BY-SA 3.0.
  • "inheritance of acquired characteristics." http://en.wikipedia.org/wiki/inheritance%20of%20acquired%20characteristics. Wikipedia CC BY-SA 3.0.
  • "evolution." http://en.wiktionary.org/wiki/evolution. Wiktionary CC BY-SA 3.0.
  • "OpenStax College, Biology. November 4, 2013." http://cnx.org/content/m44568/latest/?collection=col11448/latest. OpenStax CNX CC BY 3.0.
  • "OpenStax College, Biology. October 16, 2013." http://cnx.org/content/m44561/latest/?collection=col11448/latest. OpenStax CNX CC BY 3.0.
  • "OpenStax College, Introduction. October 16, 2013." http://cnx.org/content/m44561/latest/Figure_18_00_01ab.jpg. OpenStax CNX CC BY 3.0.
• Charles Darwin and Natural Selection
  • "Boundless." http://www.boundless.com/. Boundless Learning CC BY-SA 3.0.
  • "natural selection." http://en.wiktionary.org/wiki/natural_selection. Wiktionary CC BY-SA 3.0.
  • "OpenStax College, Biology. October 16, 2013." http://cnx.org/content/m44568/latest/?collection=col11448/latest. OpenStax CNX CC BY 3.0.
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  • "OpenStax College, Understanding Evolution. January 16, 2015." http://cnx.org/content/m44568/latest/. OpenStax CNX CC BY 3.0.
• The Galapagos Finches and Natural Selection
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  • "evolution." http://en.wiktionary.org/wiki/evolution. Wiktionary CC BY-SA 3.0.
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  • "Animal Behavior/Darwin's Finches." http://en.wikibooks.org/wiki/Animal_Behavior/Darwin's_Finches. Wikibooks CC BY-SA 3.0.
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• Processes and Patterns of Evolution
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  • "adaptation." http://en.wiktionary.org/wiki/adaptation. Wiktionary CC BY-SA 3.0.
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• Evidence of Evolution
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  • "homologous structure." http://en.wikipedia.org/wiki/homologous%20structure. Wikipedia CC BY-SA 3.0.
  • "vestigial structure." http://en.wiktionary.org/wiki/vestigial_structure. Wiktionary CC BY-SA 3.0.
  • "biogeography." http://en.wiktionary.org/wiki/biogeography. Wiktionary CC BY-SA 3.0.
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  • "OpenStax College, Understanding Evolution. October 16, 2013." http://cnx.org/content/m44568/latest/Figure_18_01_06.jpg. OpenStax CNX CC BY 3.0.
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  • "OpenStax College, Understanding Evolution. October 16, 2013." http://cnx.org/content/m44568/latest/Figure_18_01_07ab.jpg. OpenStax CNX CC BY 3.0.
• Misconceptions of Evolution
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  • "Lamarckism." http://en.wikipedia.org/wiki/Lamarckism. Wikipedia CC BY-SA 3.0.
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