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Given the right circumstances, and enough time, evolution leads to the emergence of new [[species]]. Scientists have struggled to find a precise and all-inclusive definition of ''species''. [[Ernst Mayr]] (1904–2005) defined a species as a population or group of populations whose members have the potential to interbreed naturally with one another to produce [[viable]], fertile offspring. Also, the members of a species cannot produce viable fertile offspring with members of other species.<ref name= >{{cite book |title= What Evolution Is |last= Mayer |first= Ernst |authorlink= |coauthors= |year=2001 |edition = |publisher= Basic Books |location= New York |isbn= 0465044255 }}</ref> Mayr's definition has gained wide acceptance among biologists, but does not apply to organisms such as [[bacteria]], which reproduce without sex.
Given the right circumstances, and enough time, evolution leads to the emergence of new [[species]]. Scientists have struggled to find a precise and all-inclusive definition of ''species''. [[Ernst Mayr]] (1904–2005) defined a species as a population or group of populations whose members have the potential to interbreed naturally with one another to produce [[viable]], fertile offspring. Also, the members of a species cannot produce viable fertile offspring with members of other species.<ref name= >{{cite book |title= What Evolution Is |last= Mayer |first= Ernst |authorlink= |coauthors= |year=2001 |edition = |publisher= Basic Books |location= New York |isbn= 0465044255 }}</ref> Mayr's definition has gained wide acceptance among biologists, but does not apply to organisms such as [[bacteria]], which reproduce without sex.


[[Speciation]] is the lineage-splitting event that results in two separate species forming from a single common ancestral population. The most widely accepted method of speciation is called ''[[allopatric speciation]]''. This requires the geographic separation of a population, such as the emergence of mountain ranges or the formation of canyons. For speciation to occur, separation must be substantial, so that genetic exchange between the two populations is completely disrupted. In their separate environments, the genetically isolated groups follow their own unique evolutionary pathways. Each group will accumulate different mutations as well as be subjected to different selective pressures. The accumulated genetic changes may result in separated populations that can no longer interbreed if they are reunited. <ref> {{cite web |url=http://ngm.nationalgeographic.com/ngm/0411/feature1/fulltext.html#top |title=Was Darwin Wrong? |accessdate=2007-12-23 |last=Quammen |first=David |coauthors= |date=2004 |work= National Geographic Magazine |publisher=National Geographic </ref> Barriers that prevent interbreeding are either ''prezygotic'' (prevent mating or fertilization) or ''postzygotic'' (barriers that occur after fertilization). If interbreeding is no longer possible, then they will be considered different species.<ref>{{cite web |url= http://www.smithsonianmagazine.com/issues/2005/december/darwin.php?page=2 |title=The Evolution of Charles Darwin |accessdate=2007-08-31 |last= Sulloway |first= Frank J |coauthors= |date=2005 |work= Smithsonian Magazine |publisher= Smithsonian Institute }}</ref>
[[Speciation]] is the lineage-splitting event that results in two separate species forming from a single common ancestral population. The most widely accepted method of speciation is called ''[[allopatric speciation]]''. This requires the geographic separation of a population, such as the emergence of mountain ranges or the formation of canyons. For speciation to occur, separation must be substantial, so that genetic exchange between the two populations is completely disrupted. In their separate environments, the genetically isolated groups follow their own unique evolutionary pathways. Each group will accumulate different mutations as well as be subjected to different selective pressures. The accumulated genetic changes may result in separated populations that can no longer interbreed if they are reunited. <ref> {{cite web |url=http://ngm.nationalgeographic.com/ngm/0411/feature1/fulltext.html |title=Was Darwin Wrong? |accessdate=2007-12-23 |last=Quammen |first=David |coauthors= |date=2004 |work= National Geographic Magazine |publisher=National Geographic </ref> Barriers that prevent interbreeding are either ''prezygotic'' (prevent mating or fertilization) or ''postzygotic'' (barriers that occur after fertilization). If interbreeding is no longer possible, then they will be considered different species.<ref>{{cite web |url= http://www.smithsonianmagazine.com/issues/2005/december/darwin.php?page=2 |title=The Evolution of Charles Darwin |accessdate=2007-08-31 |last= Sulloway |first= Frank J |coauthors= |date=2005 |work= Smithsonian Magazine |publisher= Smithsonian Institute }}</ref>
Those who reject evolution as a viable theory often claim that speciation has never been observed. However, speciation has been observed in several groups of organisms, including bacteria, round worms, insects, and fish, as well as in several groups of plants. In addition, past speciation events are recorded in fossils.<ref>{{cite journal |author=Jiggins CD, Bridle JR |title=Speciation in the apple maggot fly: a blend of vintages? |journal=Trends Ecol. Evol. (Amst.) |volume=19 |issue=3 |pages=111&ndash;4 |year=2004 |pmid=16701238}}<br />*{{cite web|author=Boxhorn, J|date=1995|url=http://www.talkorigins.org/faqs/faq-speciation.html|title=Observed Instances of Speciation|publisher=The TalkOrigins Archive|accessdate=2007-05-10}}<br />*{{cite journal |author=Weinberg JR, Starczak VR, Jorg, D |title=Evidence for Rapid Speciation Following a Founder Event in the Laboratory |journal=Evolution |volume=46 |issue=4 |pages=1214&ndash;20 |year=1992 |doi=10.2307/2409766}}</ref> For example, scientists have documented the formation of five new species of cichlid fishes from a single common ancestor that was isolated fewer than 4000 years ago from the parent stock in Lake Nagubago. The evidence for speciation in this case was morphology (physical appearance) and lack of natural interbreeding. These fish have complex mating rituals and a variety of colorations; the slight modifications introduced in the new species have changed the mate selection process and the five forms that arose could not be convinced to interbreed.<ref name= >{{cite book |title= Populations, Species, and Evolution |last= Mayr |first= Ernst |authorlink= |coauthors= |year= 1970 |edition = |publisher= Belknap Press of Harvard University Press |location= Cambridge, MA |isbn= 0674690109 }}</ref>
Those who reject evolution as a viable theory often claim that speciation has never been observed. However, speciation has been observed in several groups of organisms, including bacteria, round worms, insects, and fish, as well as in several groups of plants. In addition, past speciation events are recorded in fossils.<ref>{{cite journal |author=Jiggins CD, Bridle JR |title=Speciation in the apple maggot fly: a blend of vintages? |journal=Trends Ecol. Evol. (Amst.) |volume=19 |issue=3 |pages=111&ndash;4 |year=2004 |pmid=16701238}}<br />*{{cite web|author=Boxhorn, J|date=1995|url=http://www.talkorigins.org/faqs/faq-speciation.html|title=Observed Instances of Speciation|publisher=The TalkOrigins Archive|accessdate=2007-05-10}}<br />*{{cite journal |author=Weinberg JR, Starczak VR, Jorg, D |title=Evidence for Rapid Speciation Following a Founder Event in the Laboratory |journal=Evolution |volume=46 |issue=4 |pages=1214&ndash;20 |year=1992 |doi=10.2307/2409766}}</ref> For example, scientists have documented the formation of five new species of cichlid fishes from a single common ancestor that was isolated fewer than 4000 years ago from the parent stock in Lake Nagubago. The evidence for speciation in this case was morphology (physical appearance) and lack of natural interbreeding. These fish have complex mating rituals and a variety of colorations; the slight modifications introduced in the new species have changed the mate selection process and the five forms that arose could not be convinced to interbreed.<ref name= >{{cite book |title= Populations, Species, and Evolution |last= Mayr |first= Ernst |authorlink= |coauthors= |year= 1970 |edition = |publisher= Belknap Press of Harvard University Press |location= Cambridge, MA |isbn= 0674690109 }}</ref>



Revision as of 22:48, 23 December 2007

To see a brief description of evolution in simpler language, visit the Simple Wikipedia article on evolution.

Evolution is the accumulation of changes through succeeding generations of organisms that results in the emergence of new species. Since the origin of life, evolution has transformed the first species (the common ancestor of all living things) into a large number of different species.

Evolutionary biology, the scientific study of evolution, has helped provide a clear understanding of this process. For example, Gregor Mendel's work with plants demonstrated the hereditary patterns of genetics that are the source of variation between organisms.[1] . The discovery of the molecular structure of DNA combined with advances in the field of population genetics has led to a better understanding of how new species develop from ancestral forms, an important component of evolution known as speciation.

The theory of evolution is the foundation of nearly all research conducted in biology, including molecular biology, paleontology, and taxonomy. Substantiated by a large quantity of reliable scientific evidence, evolution is supported by 99.9% of the scientific community.[2] However, while the general theory is agreed upon, uncertainty remains regarding the detailed mechanisms of change.

Darwin's idea: evolution by natural selection

Charles Darwin by George Richmond (late-1830s)

In 1859, Charles Darwin (1809–1882) published On the Origin of Species, which articulated the first full-fledged theory of evolution by natural selection. Darwin viewed the history of life like a tree, each fork in the tree’s limbs representing a shared ancestry. The tips of the limbs represented modern species and the branches represented the common ancestors shared amongst species. To explain these relationships, Darwin contended that all living things were related and descended from a few forms, or even from a single common ancestor,in a process he described as "descent with modification".[3]

Darwin's view was controversial because humans did not receive special consideration in this evolutionary tree: they were merely one of its many branches. Though he did not make this explicit at first, his friend and supporter Thomas Henry Huxley soon presented evidence that humans and apes shared a common ancestor. The popular press of the day misinterpreted this as an assertion that humans were descended from monkeys.

Darwin's explanation of the mechanism of evolution relied on his theory of natural selection, a theory developed from the following observations:[4]

  1. If all the individuals of a species reproduced successfully, the population of that species would increase exponentially.
  2. Except for seasonal fluctuations, populations tend to remain stable in size.
  3. Environmental resources are limited.
  4. The traits found in a population vary extensively. No two individuals in a given species are exactly alike.
  5. Many of the variations found in a population can be passed on to offspring.

From these observations, Darwin deduced that the production of more offspring than the environment can support leads to a struggle for existence, with only a small percentage of individuals surviving in each generation. He noted that the chance for surviving this struggle is not random, but depends on how well-adapted each individual is to its environment. Well-adapted, or "fit" individuals will more likely leave a greater number of offspring than their less well-adapted competitors. Darwin concluded that the unequal ability of individuals to survive and reproduce leads to gradual changes in the population as the traits which help the organism survive and reproduce accumulate over generations and those that inhibit its survival and reproduction are lost. Darwin used the term natural selection to describe this process.

It is now known that the variations in a population arise by chance mutations in DNA, but natural selection is not a process of chance: the environment determines the probability of reproductive success. The end products of natural selection are organisms that are adapted to their present environments.

In orchids Darwin observed a relationship that has developed with insects that insures successful pollination of the plant. He noted that orchids have developed a variety of elaborate structures to attract insects and guarantee that the pollen sticks to their bodies so that it may be transported to the female flower of another orchid. Despite the appearance of complexity, the flower parts in the orchid are derived from ordinary flower parts that usually perform different functions. Darwin proposed that the orchids do not represent the work of an ideal engineer, but that they were “rigged” from a pre-existing parts.[5]

Darwin noted that orchids exhibited a variety of complex adaptations to insure pollination; all derived from basic floral parts.

Natural selection does not involve progress towards an ultimate goal. Evolution does not necessarily strive for more advanced, more intelligent, or more sophisticated life forms. For example, fleas (wingless parasites) are descended from a winged, ancestral scorpionfly,[6] and snakes are lizards that no longer require limbs. Organisms are merely the outcome of variations that succeed or fail, dependent upon the environmental conditions at the time. In reality, when the environment changes, most species fail to adapt and become extinct.

Mendel’s contribution: heredity

File:Mendel.png
Gregor Mendel's work on the inheritance of traits laid the foundation for genetics.

Darwin’s theory of natural selection laid the groundwork for modern evolutionary theory. However, Darwin lacked an accurate explanation for the source of variations within populations. Like many of his predecessors, Darwin incorrectly deduced that heritable traits were a product of the environment, assuming that characteristics acquired during an organism's lifetime could be passed on to its offspring (e.g. giraffes stretching for leaves on higher branches would give birth to offspring with longer necks). This misconception (of the inheritance of acquired characters) became known as Lamarckism, after one of its primary supporters, Jean-Baptiste Lamarck (1744–1829). However, the suggestion that frequently used body structures become better developed, while unused structures deteriorate, is not supported by evidence.[7]

The missing information necessary to help explain the emergence of new traits in offspring was provided by the pioneering genetics work of Gregor Mendel (1822–1884). Mendel’s experiments with breeding pea plants demonstrated that heredity works by reshuffling and recombining factors (what we now know as genes) during sexual reproduction. Genes are the basic units of heredity in living organisms. They contain the biological information that directs the physical development and behavior of the organism.[8] Later research by Thomas Hunt Morgan showed that genes are linked in a series on chromosomes and it is the reshuffling of these chromosomes that allow for unique combinations in the offspring.

The merging of Darwin's theory with an understanding of heredity led to a clear understanding of the mechanisms that cause evolution.[9]

Evidence for evolution

During the voyage of the Beagle, naturalist Charles Darwin collected fossils in South America, and found fragments of armor like giant versions of the scales on the modern armadillos living nearby. On his return, the anatomist Richard Owen showed that the fragments were from gigantic extinct glyptodons, related to the armadillos. This was one of the patterns of distribution that helped Darwin to develop his theory.[10]

Science has discovered a wide range of evidence for evolution, including fossils, homologous structures, and molecular similarities between species' DNA.

The fossil record

Research in the field of paleontology, the study of fossils, supports the idea that all living creatures are related. Fossils provide evidence that accumulated changes in organisms over long periods of time have led to the diverse forms of life we see today. A fossil itself reveals the organism's structure and the relationships between present and extinct species, allowing paleontologists to construct a family tree for all of the life forms on earth. [11]

Modern paleontology began with the work of Georges Cuvier (1769–1832). Cuvier noted that, in sedimentary rock, each layer contained a specific group of fossils. The deeper layers, which he conjectured to be older, contained simpler life forms. He also noted that many forms of life from the past are no longer present today. Cuvier proposed the idea of catastrophism, which explained the fossil record in the light of the theological views of his time. He proposed that catastrophes had occurred in localized areas throughout the earth’s history. Such areas were then repopulated by species that migrated from nearby locations.[12]

A very large number of fossils have now been discovered and identified. These fossils serve as a chronological record of evolution. The fossil record also provides examples of transitional species that demonstrate ancestral links between past and present life forms.[11] One such transitional fossil is Archaeopteryx, an ancient creature that had the distinct characteristics of a reptile, yet also had the feathers of a bird. The implication from such a find is that modern reptiles and birds arose from a common ancestor.[13] [5]

Comparative anatomy

Taxonomy

Homologous structures. Note how the same basic structure appears repeatedly in different types of forelimbs of different species.

Taxonomy is the branch of biology that names and classifies all living things. Scientists use morphological and genetic similarities to assist them in categorizing life forms based on ancestral relationships. For example, orangutans, gorillas, chimpanzees, and humans all belong to the same taxonomic grouping referred to as a family – in this case the family called Hominidae. These animals are grouped together because of similarities in morphology that come from common ancestry (called homology).[14]

Strong evidence for evolution comes from analysis of homologous structures in different species that no longer perform the same task. One example is the forelimbs of mammals. The forelimbs of a human, cat, whale, and bat all have strikingly similar bone structures. However, each of these four species' forelimbs performs a different task. The same bones that construct a bird's wings, which are used for flight, also construct a whale's flippers, which are used for swimming. Such a "design" makes little sense if they are unrelated and uniquely constructed for their particular tasks. The theory of evolution explains these homologous structures: all four animals shared a common ancestor, and each has undergone change over many generations. These changes in structure have produced forelimbs adapted for different tasks. Darwin described such changes in morphology as descent with modification.[15]

Embryology
In some cases, anatomical comparison of structures in the embryos of two or more species provides evidence for a shared ancestor that may not be obvious in the adult forms. Such homologies might be lost or take on different functions as the embryo develops. For example, part of the basis of classifying the vertebrate group (which includes humans), is the presence of a tail (extending beyond the anus) and pharyngeal gill slits; both appear during some stage of development but are not always obvious in the adult form. In humans, for example, the pharyngeal gill slit and the tail degenerates during embryonic development.[16]

Because of the morphological similarities present in embryos of different species during development, it was once assumed that organisms re-enact their evolutionary history as an embryo – for example, human embryos passed through an amphibian then a reptilian stage before completing their development as mammals. Such a re-enactment, called ontogeny recapitulates phylogeny, is not supported by scientific evidence. What does occur, however, is that the first stages are similar in broad groups of organisms [17] – for instance, a hollow ball of cells in animals, which then differentiate. At the pharyngula stage, for instance, all vertebrates are extremely similar, but do not exactly resemble any ancestral species. As development continues, more and more of the features typical of the species emerge from the basic pattern.

Vestigial structures

Homology also includes a unique group of shared structures referred to as vestigial structures. Vestigial refers to anatomical parts that are of minimal, if any, value to the organism that possesses them. These apparently illogical structures are remnants of organs that played an important role in ancestral forms. For example, whales have small vestigial leg bones that appear to be remnants of the legs that their ancestors used to walk on land.[18]

Humans also have many vestigial structures, including the ear muscles, the wisdom teeth, the appendix, the tail bone, body hair (including goose bumps), and the semilunar fold in the corner of the eye.

Convergent evolution

However, anatomical comparisons can also be misleading; not all anatomical similarities indicate a close relationship. Organisms that share similar environments will often develop similar physical features; a process known as convergent evolution. For example, both sharks and dolphins have similar body forms, yet are only distantly related – sharks are fish and dolphins mammals. Such similarities are a result of both populations being exposed to the same selective pressures. Within both groups, changes that aid swimming would be favored. Thus, over time, they develop similar morphology, even though they are not closely related.[4]

Artificial selection

The results of artificial selection: Great Danes and Chihuahuas

Artificial selection is the controlled breeding of domestic plants and animals. In controlled breeding, humans determine which animals will reproduce, and to some degree, which alleles will be passed on to future generations. The process of artificial selection has had a significant impact on the evolution of domestic animals. For example, people have produced different types of dogs by controlled breeding. The differences between the Chihuahua and the Great Dane are the result of artificial selection. Despite their dramatically different physical appearance, they and all other dogs evolved from a few wolves domesticated by humans in what is now China fewer than 15,000 years ago.[19]

Artificial selection has also produced a wide variety of plants. In the case of maize (corn), recent genetic evidence suggests that domestication occurred 10,000 years ago in central Mexico.[20] Prior to domestication, the edible portion of the wild form was small and difficult to collect. Today The Maize Genetics Cooperation • Stock Center maintains a collection of more than 10,000 genetic variations of maize that have arisen by random mutations and chromosomal variations from the original wild type.[21]

Darwin drew much of his support for natural selection from observing the outcomes of artificial selection. Much of his book On the Origin of Species was based on his observations of the diversity in domestic pigeons arising from artificial selection. Darwin proposed that if dramatic changes in domestic plants and animals could be achieved by humans in short periods, then natural selection, given millions of years, could produce the differences between living things today. In fact, there is no real difference in the genetic processes underlying artificial and natural selection. As in natural selection, the variations are a result of random mutations; the only difference is that in artificial selection, humans select which organisms will be allowed to breed.[4]


A section of DNA

Molecular biology

Every living organism contains molecules of DNA, RNA, and protein. If two organisms are closely related, these molecules will be very similar. On the other hand, the more distantly related two organisms are, the more molecular differences they will have. For example, two brothers will be very closely related and will have very similar DNA, while distant cousins will have more differences in their DNA. Comparing these molecules is extremely useful when studying species that are very closely related. The extent of their relationship is shown by how similar these molecules are. For example, comparing the DNA of chimpanzees with that of gorillas and humans showed that there is as much as a 96% similarity between the DNA of humans and chimps,[22] suggesting humans and chimpanzees are more closely related to each other than to gorillas.[23]

Scientists have made great strides in analyzing these molecules, particularly the DNA that makes up organisms' genes. Genes are the pieces of DNA that carry information and they influence the properties of an organism. For example, your genes decide your general appearance and the color of your hair and eyes – since close relatives have similar genes they tend to look alike. The exact form of the genes in an organism is called the organism's genotype and this set of genes influences the properties (or phenotype) of an organism.[24] The field of molecular systematics focuses on measuring the similarities in these molecules and using this information to work out how different types of organisms are related through evolution. These comparisons have allowed biologists to build a relationship tree of the evolution of life on earth.[25] They have even allowed scientists to unravel the relationships of organisms whose common ancestors lived such a long time ago that no real similarities remain in the appearance of the organisms.

Co-evolution

Co-evolution is a process in which two or more species influence the evolution of each other. All organisms are influenced by life around them; however, in "co-evolution", there is evidence that genetically determined traits in each species directly resulted from the interaction between the two organisms.[4]

An extensively documented example of co-evolution is the relationship between Pseudomyrmex, a type of ant, and the acacia, a plant that the ant uses for food and shelter. The relationship between the two is so intimate that it has led to the evolution of special structures and behaviors in both organisms. The ant defends the acacia against herbivores and clears the forest floor of the seeds from competing plants. In response, the plant has evolved swollen thorns that the ants use as shelter and special flower parts that the ants eat.<[26] Such co-evolution does not imply that the ants and the tree choose to behave in an altruistic manner. Rather, across a population small genetic changes in both ant and tree benefited each and the benefit gave a slightly higher chance of the characteristic being passed on to the next generation, where it gave the increasingly interdependent trees and ant colonies a greater chance of survival. Over time, successive mutations created the relationship we observe today.

Population genetics

Some definitions

From a genetic viewpoint, evolution is a generation-to-generation change in the frequencies of alleles within a population that shares a common gene pool. A population is a localized group of individuals belonging to the same species. For example, all the moths of the same species living in an isolated forest are a population. An allele is one specific form of a gene; for example, a gene for coloration in moths may have two alleles, black and white. A gene pool is the complete set of alleles in a single population. Each allele occurs a certain number of times in the gene pool. The fraction of genes that belong to a given allele is called the allele frequency. Therefore, if half of the body-color genes in a population of moths are genes for black-bodies, then the black-body allele frequency is 0.50 or 50%.[27] Evolution occurs when there are changes in the frequencies of alleles within a population of interbreeding organisms.

Hardy-Weinberg equilibrium

The Hardy-Weinberg principle states that the frequencies of alleles in a sufficiently large population will remain constant if the only forces acting on the population are the random reshuffling of alleles during the formation of the gametes and random combination of the genes in these sex cells during fertilization.[28] A population in which the frequencies of alleles are constant is not experiencing evolution.

Examples

White and black mice

Suppose a group of mice inhabit a barn. In this population, there are only two versions of the gene that controls fur color. One allele produces black fur and accounts for 75% of the genes, the other produces white fur and makes up the remaining 25% of the genes. If an allele’s chance of being passed on to the next generation is due entirely to random processes (the shuffling and combining that takes place in the formation of sex cells and fertilization), the allele frequencies will stay the same and the composition of the gene pool remains 75% black-coding genes and 25% white-coding genes. Since there is no change in the allelic frequencies, there is no evolutionary change in fur color. This population is in Hardy-Weinberg equilibrium or is non-evolving.[4] It is very rare for natural populations to experience no change in the frequency of alleles from generation to generation. Therefore, frequencies of alleles in a gene pool are typically changing, resulting in evolution of populations over successive generations. Several forces may change the composition of the gene pool.

One process that can lead to changes is the gene pool is the exchange of alleles between members of different populations. Exchanges occur when new members join the population or when others leave. Such migration between populations is called gene flow.[4] Mutations can also alter the gene pool by creating new alleles and thus change the frequency of the pre-existing alleles.[4] Mutations immediately change the gene pool by substituting one allele for another. Mutations have very little impact on the overall allele frequency in a large population.[4] In addition, preference for any particular allele during mate selection will effect the frequency of alleles. Perhaps the most dramatic effect on a gene pool results from natural selection which can reduce or increase the frequency of a specific allele in a gene pool.[4] The chance that each allele has for survival and reproduction must be the same if the frequency of alleles is to remain constant.

Small populations are very susceptible to chance fluctuations in the number of individuals, a condition known as genetic drift.[4] Two common situations may arise that affect the genetic makeup of a small population as a result of genetic drift. The first situation that can lead to genetic drift is the The Bottleneck Effect. This occurs because random chance events, such as fires or floods, significantly reduce the number of individuals in a population.[4] The remaining members may not accurately represent the original gene pool. The second situation that can lead to genetic drift is theThe Founder Effect. This occurs when only a few individuals colonize a new habitat. The smaller the founding party, the less likely its gene pool will represent the gene pool of the original population.[4]

Species

There are numerous species of cichlids that demonstrate dramatic variations in morphology.

Given the right circumstances, and enough time, evolution leads to the emergence of new species. Scientists have struggled to find a precise and all-inclusive definition of species. Ernst Mayr (1904–2005) defined a species as a population or group of populations whose members have the potential to interbreed naturally with one another to produce viable, fertile offspring. Also, the members of a species cannot produce viable fertile offspring with members of other species.[29] Mayr's definition has gained wide acceptance among biologists, but does not apply to organisms such as bacteria, which reproduce without sex.

Speciation is the lineage-splitting event that results in two separate species forming from a single common ancestral population. The most widely accepted method of speciation is called allopatric speciation. This requires the geographic separation of a population, such as the emergence of mountain ranges or the formation of canyons. For speciation to occur, separation must be substantial, so that genetic exchange between the two populations is completely disrupted. In their separate environments, the genetically isolated groups follow their own unique evolutionary pathways. Each group will accumulate different mutations as well as be subjected to different selective pressures. The accumulated genetic changes may result in separated populations that can no longer interbreed if they are reunited. [30] Barriers that prevent interbreeding are either prezygotic (prevent mating or fertilization) or postzygotic (barriers that occur after fertilization). If interbreeding is no longer possible, then they will be considered different species.[31] Those who reject evolution as a viable theory often claim that speciation has never been observed. However, speciation has been observed in several groups of organisms, including bacteria, round worms, insects, and fish, as well as in several groups of plants. In addition, past speciation events are recorded in fossils.[32] For example, scientists have documented the formation of five new species of cichlid fishes from a single common ancestor that was isolated fewer than 4000 years ago from the parent stock in Lake Nagubago. The evidence for speciation in this case was morphology (physical appearance) and lack of natural interbreeding. These fish have complex mating rituals and a variety of colorations; the slight modifications introduced in the new species have changed the mate selection process and the five forms that arose could not be convinced to interbreed.[33]

Barriers to breeding between species

Reproductive barriers that prevent interbreeding can be classified as either prezygotic barriers or postzygotic barriers.[4] The distinction between the two lies in whether the barrier prevents the generation of offspring before fertilization of the egg or following such fertilization.

Barriers that prevent fertilization

Prezygotic barriers prevent mating between species or prevent the fertilization of the egg if the species attempt to mate. Some examples are:

Different species of fireflies do not recognize each others' mating signals, and as a result do not generally interbreed.
  • Temporal isolation – Occurs when species mate at different times. Populations of the western spotted skunk (Spilogale gracilis) overlap with the eastern spotted skunk (Spilogale putorius) yet remain separate species because the former mates in summer and the latter in late winter.[4]
  • Behavioral isolation – Signals that elicit a mating response may be sufficiently different to prevent a desire to interbreed. The rhythmic flashing in male fireflies is species-specific and thus serves as a prezygotic barrier.[34]
  • Mechanical isolation – Anatomical differences in reproductive structures may prevent interbreeding. This is especially true in flowering plants that have evolved specific structures adapted to certain pollinators. Mechanical barriers often contribute to reproductive isolation of flowers that are pollinated by insects.
  • Gametic isolation – The gametes of the two species are chemically incompatible, thus preventing fertilization. Gamete recognition may be based on specific molecules on the surface of the egg that attach only to complementary molecules on the sperm. Such mechanisms are common in fish species.[4]
  • Geographic/habitat isolationGeographic: The two species are separated by large-scale physical barriers, such as a mountain or large body of water, and therefore cannot mate with each other. This is illustrated in two separate species of antelope squirrels, genus Ammospermophilus, which inhabit opposite sides of the Grand Canyon. Habitat: The two species prefer different habitats, even if they live in the same general area, and therefore do not encounter each other. For example, two different species of garter snakes in the genus Thamnophis occur in the same area but one prefers the water while the other prefers dry land.[4] ====Barriers acting after fertilization====
File:100 0637.jpg
The mule is a hybrid of a female horse and a male donkey, and is usually infertile.

Postzygotic barriers occur after fertilization, usually resulting in the formation of a hybrid zygote[35] that is either not viable[36] or not fertile. This is typically a result of incompatible chromosomes in the zygote. Examples include:

  • Reduced hybrid viability – A barrier between species occurs after the formation of the zygote, resulting in incomplete development and death of the offspring.
  • Reduced hybrid fertility – Even if two different species successfully mate, the offspring produced may be infertile. Crosses of horse species within the genus Equus tend to produce viable but sterile offspring. For example, crosses of zebra x horse and zebra x donkey produce sterile zorses and zedonks. Horse-donkey crosses produce sterile mules. Very rarely, a female mule may be fertile.[37]
  • Hybrid breakdown – Some hybrids are fertile for a single generation but then become weak or inviable.


Different perspectives on the mechanism of evolution

The theory of evolution is widely accepted among the scientific community, serving to link the diverse specialty areas of biology. Evolution provides the field of biology with a solid scientific base. The significance of evolutionary theory is best described by the title of a paper by Theodosius Dobzhansky (1900–1975), published in American Biology Teacher; "Nothing in Biology Makes Sense Except in the Light of Evolution."

Nevertheless, the theory of evolution is not static. In fact, there is much discussion within the scientific community concerning the mechanisms behind the evolutionary process. For example, the rate at which evolution occurs is still under discussion. In addition, the primary unit of evolutionary change, the organism or the genes, is not agreed on.

Rate of change

Two views exist concerning the rate of evolutionary change. Darwin and his contemporaries viewed evolution as a slow and gradual process. Evolutionary trees are based on the idea that profound differences in species are the result of many small changes that accumulate over long periods.

The view that evolution is gradual had its basis in the works of the geologist James Hutton (1726–1797) and his theory called "gradualism". Hutton's theory suggests that profound geological change is the cumulative product of slow, continuous processes. A similar perspective was adopted for biological changes. Such a view fails to explain the fossil record, which shows evidence of new species appearing suddenly, then persisting in that form for long periods. The paleontologist Stephen Jay Gould (1940–2002) developed a model that suggests that evolution experiences periods of rapid change alternating with periods of relative stability, a model called "punctuated equilibrium".[38]

Unit of change

It is generally accepted amongst biologists that the unit of selection in evolution is the organism, and that natural selection serves to either enhance or reduce the reproductive potential of an individual. Reproductive success, therefore, can be measured by the volume of an organism's surviving offspring. The organism view has been challenged by a variety of biologists, as well as philosophers. Richard Dawkins, for example, proposes that much insight can be gained if we look at evolution from the gene's point of view, and view natural selection as selecting amongst genes, in addition to organisms[39]. In his book The Selfish Gene, he explains:

Individuals are not stable things, they are fleeting. Chromosomes too are shuffled to oblivion, like hands of cards soon after they are dealt. But the cards themselves survive the shuffling. The cards are the genes. The genes are not destroyed by crossing-over; they merely change partners and march on. Of course they march on. That is their business. They are the replicators and we are their survival machines. When we have served our purpose we are cast aside. But genes are denizens of geological time: genes are forever.

Others view selection working on many levels, not just at a single level of organism or gene; for example, Stephen Jay Gould called for a hierarchical perspective on selection.[40]

Summary

Evolution in popular culture
The language of evolution became pervasive in Victorian Britain as Darwin's work spread and became better known:
"Survival of the fittest" – used by Herbert Spencer in Principles of Biology (1864)
"Nature, red in tooth and claw" – from Alfred Lord Tennyson's In Memoriam A.H.H. (1849)[41]
It even merited a song in Gilbert and Sullivan's 1884 opera, Princess Ida, which concludes:

"Darwinian man, though well behaved,
at best is only a monkey shaved!"

The theory of evolution, which explains the variations in biological species, is founded on several basic observations. The first is that there are genetic variations within a population. Some offspring, by chance, have features that allow them to survive and thrive better than others. The offspring that survive will be more likely to have offspring of their own. Some of these useful features are then passed along to new generations.

Evolution is not a random process for creating new life forms. Mutations are (partly) random, but natural selection is far from random. Therefore, evolution is an inevitable result of imperfectly copying, self-replicating machines reproducing over billions of years under the selection pressure of the environment.

The idea can be hard to grasp, so there are many misconceptions about evolution, and for various reasons people raise objections to the theory of evolution. In fact, the theory of evolution is supported by a tremendous amount of evidence. This evidence includes direct observation in the laboratory and field studies. Also, domesticated animals have evolved as we selectively breed them for certain traits. Finally, the records of past evolution are found in fossils as well as in our fundamentally similar genetic codes, demonstrating common ancestry of all organisms, both surviving and extinct.

Evolution is one of the most successful scientific theories ever produced and is used and universally accepted in every field of biology. An understanding of evolution underlies all biological sciences and much of medicine.

Notes

  1. ^ Smith, John Maynard (1998). Evolutionary Genetics. New York: Oxford University Press. ISBN 0198502311. {{cite book}}: Cite has empty unknown parameter: |coauthors= (help)
  2. ^ Delgado, Cynthia (2006). "Finding the Evolution in Medicine". NIH Record (National Institutes of Health). Retrieved 2007-12-21. {{cite web}}: Cite has empty unknown parameter: |coauthors= (help)
  3. ^ Wyhe, John van (2002). "Charles Darwin: gentleman naturalist". The Complete Work of Charles Darwin Online. University of Cambridge. Retrieved 2007-12-21. {{cite web}}: Cite has empty unknown parameters: |1= and |coauthors= (help)
  4. ^ a b c d e f g h i j k l m n o p Campbell, Neil (2002). (6th Edition ed.). Benjamin Cummings. ISBN 0-8053-6624-5. {{cite book}}: |edition= has extra text (help); Missing or empty |title= (help); Unknown parameter |coauthors= ignored (|author= suggested) (help) Cite error: The named reference "Campbell" was defined multiple times with different content (see the help page).
  5. ^ a b Gould, Stephen Jay (1981). The Panda's Thumb: More Reflections in Natural History. New York: W.W, Norton & Company. ISBN 0393308197. {{cite book}}: Cite has empty unknown parameter: |coauthors= (help)
  6. ^ Hutchinson, Robert (1999). "Fleas". Veterinary Entomology. Retrieved 2007-09-03. {{cite web}}: Cite has empty unknown parameter: |coauthors= (help)
  7. ^ Waggoner, Ben (1996). "Jean-Baptiste LaMarck". Understanding Evolution. University of California Museum of Paleontology. Retrieved 2007-12-23. {{cite web}}: Cite has empty unknown parameter: |coauthors= (help)
  8. ^ Genes are composed of DNA, a long molecule that has the form of a "double helix". It resembles a ladder that has been twisted. The rungs of the ladder are formed by chemicals called nucleotides. There are four types of nucleotides, and the sequence of nucleotides carries the information in the DNA. There are short segments of the DNA called genes. The genes are like sentences built up of the "letters" of the nucleotide alphabet. Chromosomes are packages for carrying the DNA in the cells.
  9. ^ Caroll, Sean (2005). Endless Forms Most Beautiful. New York: W.W, Norton & Company. ISBN 0393060160. {{cite book}}: Cite has empty unknown parameter: |coauthors= (help)
  10. ^ Template:Harvard reference Retrieved on 2006-12-15
  11. ^ a b "The Fossil Record - Life's Epic". The Virtual Fossil Museum. Retrieved 2007-08-31. {{cite web}}: Cite has empty unknown parameter: |coauthors= (help)
  12. ^ Tattersall, Ian (1995). The Fossil Trail: How We Know What We Think We Know About Human Evolution. New York: Oxford University Press. ISBN 0195061012. {{cite book}}: Cite has empty unknown parameter: |coauthors= (help)
  13. ^ Gould, Stephen Jay (1995). Dinosaur in a Haystack. New York: Harmony Books. ISBN 0517703939. {{cite book}}: Cite has empty unknown parameter: |coauthors= (help)
  14. ^ Diamond, Jared (1992). The Third Chimpanzee: the evolution and future of the human animal. New York: HarperCollins. ISBN 0060183071. {{cite book}}: Cite has empty unknown parameter: |coauthors= (help)
  15. ^ Mayer, Ernst (2001). What Evolution Is. New York: Basic Books. ISBN 0465044255. {{cite book}}: Cite has empty unknown parameter: |coauthors= (help)
  16. ^ Weichert &, Charles (1975). Elements of Chordate Anatomy. New York: McGraw-Hill. ISBN 0070690081. {{cite book}}: Unknown parameter |coauthors= ignored (|author= suggested) (help)
  17. ^ Miller, Kenneth (1997). "Haeckel and his Embryos". Evolution Resources. Retrieved 2007-09-02. {{cite web}}: Cite has empty unknown parameter: |coauthors= (help)
  18. ^ Pagel, Mark (2002). Vestigial Organs and Structures. In Encyclopedia of Evolution. Oxford University Press. ISBN 0195122003. {{cite book}}: Cite has empty unknown parameter: |coauthors= (help)
  19. ^ McGourty, Christine (2002). "Origin of dogs traced". BBC News. Retrieved 2007-12-14. {{cite web}}: Cite has empty unknown parameter: |coauthors= (help)
  20. ^ Hall, Hardy. "Transgene Escape: Are Traditioanl Corn Varieties In Mexico Threatened by Transgenic Corn Crops". Scientific Creative Quarterly. Retrieved 2007-12-14. {{cite web}}: Cite has empty unknown parameter: |coauthors= (help)
  21. ^ "The Maize Genetics Cooperation • Stock Center". National Plant Germplasm. U.S. Department of Agriculture. 2006. Retrieved 2007-12-19. {{cite web}}: Cite has empty unknown parameter: |coauthors= (help)
  22. ^ Lovgren, Stefan (2005). "Chimps, Humans 96 Percent the Same, Gene Study Finds". National Geographic News. National Geographic. Retrieved 2007-12-23. {{cite web}}: Cite has empty unknown parameter: |coauthors= (help)
  23. ^ Carroll SB, Grenier J, Weatherbee SD (2000). From DNA to Diversity: Molecular Genetics and the Evolution of Animal Design (2nd Edition ed.). Oxford: Blackwell Publishing. ISBN 1-4051-1950-0. {{cite book}}: |edition= has extra text (help); Cite has empty unknown parameter: |1= (help)CS1 maint: multiple names: authors list (link)
  24. ^ Graur, Dan (2000). Fundamentals of Molecular Evolution. Sunderland, MA: Sinauer Associates. ISBN 0878932666. {{cite book}}: Unknown parameter |coauthors= ignored (|author= suggested) (help)
  25. ^ Ciccarelli FD, Doerks T, von Mering C, Creevey CJ, Snel B, Bork P (2006). "Toward automatic reconstruction of a highly resolved tree of life". Science. 311 (5765): 1283–7. PMID 165139821. {{cite journal}}: Check |pmid= value (help)CS1 maint: multiple names: authors list (link)
  26. ^ Janzen, Daniel (1974). "Swollen-Thorn Acacias of Central America" (PDF). Smithsonian Contributions to Biology. Smithsonian Institute. Retrieved 2007-08-31. {{cite web}}: Cite has empty unknown parameter: |coauthors= (help)
  27. ^ Allele frequencies are often given as percentages. In this case, the frequency of black-body alleles is 50%.
  28. ^ Starr, Cecie (2001). Biology: The Unity and Diversity of Life. Australia: Brooks Cole. ISBN 0534575463. {{cite book}}: Unknown parameter |coauthors= ignored (|author= suggested) (help)
  29. ^ Mayer, Ernst (2001). What Evolution Is. New York: Basic Books. ISBN 0465044255. {{cite book}}: Cite has empty unknown parameter: |coauthors= (help)
  30. ^ {{cite web |url=http://ngm.nationalgeographic.com/ngm/0411/feature1/fulltext.html |title=Was Darwin Wrong? |accessdate=2007-12-23 |last=Quammen |first=David |coauthors= |date=2004 |work= National Geographic Magazine |publisher=National Geographic
  31. ^ Sulloway, Frank J (2005). "The Evolution of Charles Darwin". Smithsonian Magazine. Smithsonian Institute. Retrieved 2007-08-31. {{cite web}}: Cite has empty unknown parameter: |coauthors= (help)
  32. ^ Jiggins CD, Bridle JR (2004). "Speciation in the apple maggot fly: a blend of vintages?". Trends Ecol. Evol. (Amst.). 19 (3): 111–4. PMID 16701238.
    *Boxhorn, J (1995). "Observed Instances of Speciation". The TalkOrigins Archive. Retrieved 2007-05-10.
    *Weinberg JR, Starczak VR, Jorg, D (1992). "Evidence for Rapid Speciation Following a Founder Event in the Laboratory". Evolution. 46 (4): 1214–20. doi:10.2307/2409766.{{cite journal}}: CS1 maint: multiple names: authors list (link)
  33. ^ Mayr, Ernst (1970). Populations, Species, and Evolution. Cambridge, MA: Belknap Press of Harvard University Press. ISBN 0674690109. {{cite book}}: Cite has empty unknown parameter: |coauthors= (help)
  34. ^ Cratsley, Christopher K (2004). "Flash Signals, Nuptial Gifts and Female Preference in Photinus Fireflies". Integrative and Comparative Biology. bNet Research Center. Retrieved 2007-09-03. {{cite web}}: Cite has empty unknown parameter: |coauthors= (help)
  35. ^ A zygote is a fertilized egg before it divides, or the organism that results from this fertilized egg.
  36. ^ Viable is defined as: capable of life or normal growth and development
  37. ^ Wood (2001). "HybriDatabase: a computer repository of organismal hybridization data". Study Group Discontinuity: Understanding Biology in the Light of Creation. Baraminology. Retrieved 2007-05-10. {{cite web}}: Cite has empty unknown parameter: |coauthors= (help)
  38. ^ Gould, Stephen Jay (1991). "Opus 200". Stephen Jay Gould Archive. Natural History. Retrieved 2007-08-31. {{cite web}}: Cite has empty unknown parameter: |coauthors= (help)
  39. ^ Wright, Sewall (September, 1980). "Genic and Organismic Selection". Evolution. 34 (5): 825. Retrieved 2007-12-23. {{cite journal}}: Check date values in: |date= (help)
  40. ^ Gould, SJ & Lloyd, EA (1999). "Individuality and adaptation across levels of selection: how shall we name and generalize the unit of Darwinism?". [1] Proc. Natl. Acad. Sci. U.S.A. 96 (21): 11904–09. PMID 10518549
  41. ^ Of course, this poem preceded the publication of Darwin's work in 1859, but it came to represent evolution for both evolution detractors and supporters. This poem was influenced by the ideas of evolution presented in Vestiges of the Natural History of Creation which had been published in 1844 (Josef L. Altholz, Professor of History, University of Minnesota (1976). "The Warfare of Conscience with Theology". The Mind and Art of Victorian England. Victorian Web. Retrieved 2007-11-06. {{cite web}}: External link in |publisher= (help)CS1 maint: multiple names: authors list (link) ).

Further reading

Evolution websites

Videos about evolution

Printable Introduction to evolution

Genetics