Overview of Evolution

Evolution is the process of change that has given us the Reality we have today. It has painted Life in the size, shape, colors, population frequency and activities that we find in every part of the World. It is not something of the past, but it is more of the present that leads to the future. It is the inherited traits of a population of organisms through successive generations. This change results from interactions between processes that introduce variation into a population, and other processes that remove it.

MUTATION is a major cause of genetic changes. These hereditary changes are passed on through reproduction and may give rise to alternative traits in organisms. Mutation may also be caused by variations in genetic recombination, which shuffles the genes into new combinations that can result in organisms exhibiting different traits.

NATURAL SELECTION – Traits that aid survival and reproduction become more common, while traits that hinder survival and reproduction become rarer. Natural selection occurs because only a small proportion of individuals in each generation will survive and reproduce, since resources are limited and organisms produce many more offspring than their environment can support.

ADAPTATION is a process that adjusts traits so they become better suited to an organism’s environment. Over many generations, heritable variation in traits is filtered by natural selection and the beneficial changes are successively retained through differential survival and reproduction which is Adaptation.

GENETIC DRIFT is another cause of evolution which leads to random changes in how common traits are in a population. Genetic drift is most important when traits do not strongly influence survival—particularly so in small populations, in which chance plays a disproportionate role in the frequency of traits passed on to the next generation.

SPECIATION is a key process in evolution in which a single ancestral species splits and diversifies into multiple new species. Ultimately, all living (and extinct) species are descended from a common ancestor via a long series of speciation events.


Evolution in organisms occurs through changes in heritable traits – particular characteristics of an organism.

The complete set of observable traits that make up the structure and behavior of an organism is called its phenotype. These traits come from the interaction of its genotype with the environment. As a result, many aspects of an organism’s phenotype are not inherited. For example, suntanned skin comes from the interaction between a person’s genotype and sunlight; thus, suntans are not passed on to people’s children.


An individual organism’s phenotype results from both its genotype and the influence from the environment it has lived in. A substantial part of the variation in phenotypes in a population is caused by the differences between their genotypes. The modern evolutionary synthesis defines evolution as the change over time in this genetic variation.

Variation comes from mutations in genetic material, migration between populations(gene flow), and the reshuffling of genes through sexual reproduction. Variation also comes from exchanges of genes between different species; for example, through horizontal gene transfer in bacteria, and hybridization in plants. However, even relatively small changes in genotype can lead to dramatic changes in phenotype: for example, chimpanzees and humans differ in only about 5% of their genomes.

Random mutations constantly occur in the genomes of organisms; these mutations create genetic variation. Mutations are changes in the DNA sequence of a cell’s genome and are caused by radiation, viruses, transposons and mutagenic chemicals, as well as errors that occur during meiosis or DNA replication.
Due to the damaging effects that mutations can have on cells, organisms have evolved mechanisms such as DNA repair to remove mutations. Therefore, the optimal mutation rate for a species is a trade-off between costs of a high mutation rate, such as deleterious mutations, and the metabolic costs of maintaining systems to reduce the mutation rate, such as DNA repair enzymes.  Viruses that use RNA as their genetic material have rapid mutation rates, which can be an advantage since these viruses will evolve constantly and rapidly and thus evade the defensive responses of e.g. the human immune system.

Sex and recombination 
In asexual organisms, genes are inherited together, or linked, as they cannot mix with genes of other organisms during reproduction. In contrast, the offspring of sexual organisms contain random mixtures of their parents’ chromosomes that are produced through independent assortment. In a related process called homologous recombination, sexual organisms exchange DNA between two matching chromosomes. Sex usually increases genetic variation and may increase the rate of evolution. Here, asexuality might allow the two sets of alleles in their genome to diverge and gain different functions.

Population genetics

White peppered moth                            Black morph in peppered moth evolution

A single gene in this population may have several alternate forms, which account for variations between the phenotypes of the organisms. An example might be a gene for coloration in moths that has two alleles: black and white.
A population is a localized group of individuals belonging to the same species.

To understand the mechanisms that cause a population to evolve, it is useful to consider what conditions are required for a population not to evolve. The Hardy-Weinberg principlestates that the frequencies of alleles (variations in a gene) in a sufficiently large population will remain constant if the only forces acting on that population are the random reshuffling of alleles during the formation of the sperm or egg, and the random combination of the alleles in these sex cells during fertilization. Such a population is said to be in Hardy-Weinberg equilibrium; it is not evolving.

Gene Flow

When they mature, male lions leave the pride where they were born and take over a new pride to mate, causing gene flow between prides.

Gene flow is the exchange of genes between populations, which are usually of the same species. Examples of gene flow within a species include the migration and then breeding of organisms, or the exchange of pollen. Gene transfer between species includes the formation of hybrid organisms and horizontal gene transfer.


Migration into or out of a population can change allele frequencies, as well as introducing genetic variation into a population. Immigration may add new genetic material to the established gene pool of a population. Conversely, emigration may remove genetic material. As barriers to reproduction between two diverging populations are required for the populations to become new species, gene flow may slow this process by spreading genetic differences between the populations. Gene flow is hindered by mountain ranges, oceans and deserts or even man-made structures such as the Great Wall of China, which has hindered the flow of plant genes.
Hybridization is, however, an important means of speciation in plants, since polyploidy (having more than two copies of each chromosome) is tolerated in plants more readily than in animals. Polyploidy is important in hybrids as it allows reproduction, with the two different sets of chromosomes each being able to pair with an identical partner during meiosis. Polyploids also have more genetic diversity, which allows them to avoid inbreeding depression in small populations.

Horizontal gene transfer is the transfer of genetic material from one organism to another organism that is not its offspring; this is most common among bacteria. In medicine, this contributes to the spread of antibiotic resistance, as when one bacteria acquires resistance genes it can rapidly transfer them to other species. Large-scale gene transfer has also occurred between the ancestors of eukaryotic cells and prokaryotes, during the acquisition of chloroplasts and mitochondria.


The two main mechanisms that produce evolution are natural selection and genetic drift. Natural selection is the process which favors genes that aid survival and reproduction. Genetic drift is the random change in the frequency of alleles, caused by the random sampling of a generation’s genes during reproduction. Natural selection usually predominates in large populations, whereas genetic drift dominates in small populations. The dominance of genetic drift in small populations can even lead to the fixation of slightly deleterious mutations. As a result, changing population size can dramatically influence the course of evolution. Population bottlenecks, where the population shrinks temporarily and therefore loses genetic variation, result in a more uniform population.

Natural Selection
Natural selection is the process by which genetic mutations that enhance reproduction become, and remain, more common in successive generations of a population. It has often been called a “self-evident” mechanism because it necessarily follows from three simple facts:

  • Heritable variation exists within populations of organisms.
  • Organisms produce more offspring than can survive.
  • These offspring vary in their ability to survive and reproduce.

These conditions produce competition between organisms for survival and reproduction. Consequently, organisms with traits that give them an advantage over their competitors pass these advantageous traits on, while traits that do not confer an advantage are not passed on to the next generation.

The central concept of natural selection is the evolutionary fitness of an organism. Fitness is measured by an organism’s ability to survive and reproduce, which determines the size of its genetic contribution to the next generation.  However, fitness is not the same as the total number of offspring: instead fitness is indicated by the proportion of subsequent generations that carry an organism’s genes. For example, if an organism could survive well and reproduce rapidly, but its offspring were all too small and weak to survive, this organism would make little genetic contribution to future generations and would thus have low fitness.

Natural selection within a population for a trait that can vary across a range of values, such as height, can be categorized into three different types. The first is directional selection, which is a shift in the average value of a trait over time — for example, organisms slowly getting taller.  Secondly, disruptive selection is selection for extreme trait values and often results in two different values becoming most common, with selection against the average value. This would be when either short or tall organisms had an advantage, but not those of medium height. Finally, in stabilizing selection there is selection against extreme trait values on both ends, which causes a decrease in variance around the average value and less diversity.  This would, for example, cause organisms to slowly become all the same height.

A special case of natural selection is sexual selection, which is selection for any trait that increases mating success by increasing the attractiveness of an organism to potential mates. Traits that evolved through sexual selection are particularly prominent in males of some animal species.

Natural selection most generally makes nature the measure against which individuals, and individual traits, are more or less likely to survive. “Nature” in this sense refers to an ecosystem, that is, a system in which organisms interact with every other element, physical as well as biological, in their local environment. Eugene Odum, a founder of ecology, defined an ecosystem as: “Any unit that includes all of the organisms…in a given area interacting with the physical environment so that a flow of energy leads to clearly defined trophic structure, biotic diversity, and material cycles (i.e.: exchange of materials between living and nonliving parts) within the system.”  Each population within an ecosystem occupies a distinct niche, or position, with distinct relationships to other parts of the system. These relationships involve the life history of the organism, its position in the food chain, and its geographic range. This broad understanding of nature enables scientists to delineate specific forces which, together, comprise natural selection.

Genetic drift

Simulation of genetic drift of 20 unlinked alleles in populations of 10 (top) and 100 (bottom). Drift to fixation is more rapid in the smaller population.

Genetic drift is the change in allele frequency from one generation to the next that occurs because alleles in offspring are a random sample of those in the parents, as well as from the role that chance plays in determining whether a given individual will survive and reproduce. In mathematical terms, alleles are subject to sampling error. As a result, when selective forces are absent or relatively weak, allele frequencies tend to “drift” upward or downward randomly (in a random walk). This drift halts when an allele eventually becomes fixed, either by disappearing from the population, or replacing the other alleles entirely. Genetic drift may therefore eliminate some alleles from a population due to chance alone. Even in the absence of selective forces, genetic drift can cause two separate populations that began with the same genetic structure to drift apart into two divergent populations with different sets of alleles.


Evolution influences every aspect of the form and behavior of organisms. Most prominent are the specific behavioral and physical adaptations that are the outcome of natural selection. These adaptations increase fitness by aiding activities such as finding food, avoiding predators or attracting mates. Organisms can also respond to selection by co-operating with each other, usually by aiding their relatives or engaging in mutually beneficial symbiosis. In the longer term, evolution produces new species through splitting ancestral populations of organisms into new groups that cannot or will not interbreed.

These outcomes of evolution are sometimes divided into macroevolution, which is evolution that occurs at or above the level of species, such as extinction and speciation, and microevolution, which is smaller evolutionary changes, such as adaptations, within a species or population.  In general, macroevolution is regarded as the outcome of long periods of microevolution. Thus, the distinction between micro- and macroevolution is not a fundamental one – the difference is simply the time involved.   In this sense, microevolution and macroevolution might involve selection at different levels – with microevolution acting on genes and organisms, versus macro evolutionary processes such as species selection acting on entire species and affecting their rates of speciation and extinction.

A common misconception is that evolution has goals or long-term plans; realistically however, evolution has no long-term goal and does not necessarily produce greater complexity. Although complex species have evolved, they occur as a side effect of the overall number of organisms increasing, and simple forms of life still remain more common in the biosphere. For example, the overwhelming majority of species are microscopic prokaryotes, which form about half the world’s biomass despite their small size, and constitute the vast majority of Earth’s biodiversity. Simple organisms have therefore been the dominant form of life on Earth throughout its history and continue to be the main form of life up to the present day, with complex life only appearing more diverse because it is more noticeable. Indeed, the evolution of microorganisms is particularly important to modern evolutionary research, since their rapid reproduction allows the study of experimental evolution and the observation of evolution and adaptation in real time.

Major evolutionary changes have occurred due to farming techniques that have resulted in MASS EXTINCTION of large quantities of simple biomass. Tilling of the soil has resulted in environmental destruction, most particularly dehydration of the soil and subsequent prokaryote death.

Adaptation is one of the basic phenomena of biology, and is the process whereby an organism becomes better suited to its habitat.  Also, the term adaptation may refer to a trait that is important for an organism’s survival. For example, the adaptation of horses’ teeth to the grinding of grass, or the ability of horses to run fast and escape predators. By using the term adaptation for the evolutionary process, and adaptive trait for the product (the bodily part or function), the two senses of the word may be distinguished. Adaptations are produced by natural selection The following definitions are due to Theodosius Dobzhansky.

  1. Adaptation is the evolutionary process whereby an organism becomes better able to live in its habitat or habitats.
  2. Adaptedness is the state of being adapted: the degree to which an organism is able to live and reproduce in a given set of habitats.
  3. An adaptive trait is an aspect of the developmental pattern of the organism which enables or enhances the probability of that organism surviving and reproducing.

Adaptation may cause either the gain of a new feature, or the loss of an ancestral feature. An example that shows both types of change is bacterial adaptation to antibiotic selection, with genetic changes causing antibiotic resistance by both modifying the target of the drug, and increasing the activity of transporters that pump the drug out of the cell. Other striking examples are the bacteria Escherichia coli evolving the ability to use citric acid as a nutrient in a long-term laboratory experiment, Flavobacterium evolving a novel enzyme that allows these bacteria to grow on the by-products of nylon manufacturing, and the soil bacterium Sphingobium evolving an entirely new metabolic pathway that degrades the synthetic pesticide pentachlorophenol. An interesting but still controversial idea is that some adaptations might increase the ability of organisms to generate genetic diversity and adapt by natural selection (increasing organisms’ evolvability).

A baleen whale skeleton, a and b label flipper bones, which were adapted from front leg bones: while c indicates vestigial leg bones, suggesting an adaptation from land to sea.

Adaptation occurs through the gradual modification of existing structures. Consequently, structures with similar internal organization may have different functions in related organisms. This is the result of a single ancestral structure being adapted to function in different ways. The bones within bat wings, for example, are very similar to those in mice feet and primate hands, due to the descent of all these structures from a common mammalian ancestor. However, since all living organisms are related to some extent, even organs that appear to have little or no structural similarity, such as arthropod, squid and vertebrate eyes, or the limbs and wings of arthropods and vertebrates, can depend on a common set of homologous genes that control their assembly and function; this is called deep homology.

During adaptation, some structures may lose their original function and become vestigial structures.  Such structures may have little or no function in a current species, yet have a clear function in ancestral species, or other closely related species. Examples include pseudo genes, the non-functional remains of eyes in blind cave-dwelling fish, wings in flightless birds, and the presence of hip bones in whales and snakes.  Examples of vestigial structures in humans include wisdom teeth, the coccyx, the vermiform appendix, and other behavioral vestiges such as goose bumps, and primitive reflexes.

However, many traits that appear to be simple adaptations are in fact exaptations: structures originally adapted for one function, but which coincidentally became somewhat useful for some other function in the process. One example is the African lizard Holaspis guentheri, which developed an extremely flat head for hiding in crevices, as can be seen by looking at its near relatives. However, in this species, the head has become so flattened that it assists in gliding from tree to tree—an exaptation.  Within cells, molecular machines such as the bacterial flagella and protein sorting machinery evolved by the recruitment of several pre-existing proteins that previously had different functions.  Another example is the recruitment of enzymes from glycolysis and xenobiotic metabolism to serve as structural proteins called crystallins within the lenses of organisms’ eyes.

A critical principle of ecology is that of competitive exclusion: no two species can occupy the same niche in the same environment for a long time.  Consequently, natural selection will tend to force species to adapt to different ecological niches. This may mean that, for example, two species of cichlid fish adapt to live in different habitats, which will minimize the competition between them for food.

An area of current investigation in evolutionary developmental biology is the developmental basis of adaptations and exaptations.   This research addresses the origin and evolution of embryonic development and how modifications of development and developmental processes produce novel features.  These studies have shown that evolution can alter development to create new structures, such as embryonic bone structures that develop into the jaw in other animals instead forming part of the middle ear in mammals.  It is also possible for structures that have been lost in evolution to reappear due to changes in developmental genes, such as a mutation in chickens causing embryos to grow teeth similar to those of crocodiles.   It is now becoming clear that most alterations in the form of organisms are due to changes in a small set of conserved genes.


Common garter snake (Thamnophis sirtalis sirtalis) which has evolved resistance to tetrodotoxin in its amphibian prey.

Interactions between organisms can produce both conflict and co-operation. When the interaction is between pairs of species, such as a pathogen and a host, or a predator and its prey, these species can develop matched sets of adaptations. Here, the evolution of one species causes adaptations in a second species. These changes in the second species then, in turn, cause new adaptations in the first species. This cycle of selection and response is called co-evolution.  An example is the production of tetrodotoxin in the rough-skinned newt and the evolution of tetrodotoxin resistance in its predator, the common garter snake. In this predator-prey pair, an evolutionary arms race has produced high levels of toxin in the newt and correspondingly high levels of toxin resistance in the snake.

However, not all interactions between species involve conflict.  Many cases of mutually beneficial interactions have evolved. For instance, an extreme cooperation exists between plants and the mycorrhizal fungi that grow on their roots and aid the plant in absorbing nutrients from the soil.  This is a reciprocal relationship as the plants provide the fungi with sugars from photosynthesis. Here, the fungi actually grow inside plant cells, allowing them to exchange nutrients with their hosts, while sending signals that suppress the plant immune system.
Coalitions between organisms of the same species have also evolved. An extreme case is the eusociality found in social insects, such as bees, termites and ants, where sterile insects feed and guard the small number of organisms in a colony that are able to reproduce.  .
Such cooperation within species may have evolved through the process of kin selection, which is where one organism acts to help raise a relative’s offspring.  This activity is selected for because if the helping individual contains alleles which promote the helping activity, it is likely that its kin will also contain these alleles and thus those alleles will be passed on.   Other processes that may promote cooperation include group selection, where cooperation provides benefits to a group of organisms.


The four mechanisms of speciation.

Speciation is the process where a species diverges into two or more descendant species. Evolutionary biologists view species as statistical phenomena and not categories or types. This view is counterintuitive since the classical idea of species is still widely held, with a species seen as a class of organisms exemplified by a “type specimen” that bears all the traits common to this species. Instead, a species is now defined as a separately evolving lineage that forms a single gene pool. Although properties such as genetics and morphology are used to help separate closely related lineages, this definition has fuzzy boundaries. Indeed, the exact definition of the term “species” is still controversial, particularly in prokaryotes, and this is called the species problem.  Biologists have proposed a range of more precise definitions, but the definition used is a pragmatic choice that depends on the particularities of the species concerned. Typically the actual focus on biological study is the population, an observable interacting group of organisms, rather than a species, an observable similar group of individuals.

There are four mechanisms for speciation. The most common in animals is allopatric speciation, which occurs in populations initially isolated geographically, such as by habitat fragmentation or migration. Selection under these conditions can produce very rapid changes in the appearance and behavior of organisms. As selection and drift act independently on populations isolated from the rest of their species, separation may eventually produce organisms that cannot interbreed.

The second mechanism of speciation is peripatric speciation, which occurs when small populations of organisms become isolated in a new environment. This differs from allopatric speciation in that the isolated populations are numerically much smaller than the parental population. Here, the founder effect causes rapid speciation through both rapid genetic drift and selection on a small gene pool.
The third mechanism of speciation is parapatric speciation. This is similar to peripatric speciation in that a small population enters a new habitat, but differs in that there is no physical separation between these two populations. Instead, speciation results from the evolution of mechanisms that reduce gene flow between the two populations. Generally this occurs when there has been a drastic change in the environment within the parental species’ habitat.

Geographical isolation of finches on the Galápagos Islands produced over a dozen new species.

Finally, in sympatric speciation species diverge without geographic isolation or changes in habitat. This form is rare since even a small amount of gene flow may remove genetic differences between parts of a population. Generally, sympatric speciation in animals requires the evolution of both genetic differences and non-random mating, to allow reproductive isolation to evolve.

One type of sympatric speciation involves cross-breeding of two related species to produce a new hybrid species. This is not common in animals as animal hybrids are usually sterile. This is because during meiosis the homologous chromosomes from each parent are from different species and cannot successfully pair. However, it is more common in plants because plants often double their number of chromosomes, to form polyploids. This allows the chromosomes from each parental species to form matching pairs during meiosis, since each parent’s chromosomes are represented by a pair already. An example of such a speciation event is when the plant species Arabidopsis thaliana and Arabidopsis arenosa cross-bred to give the new species Arabidopsis suecica. This happened about 20,000 years ago, and the speciation process has been repeated in the laboratory, which allows the study of the genetic mechanisms involved in this process.

Speciation events are important in the theory of punctuated equilibrium, which accounts for the pattern in the fossil record of short “bursts” of evolution interspersed with relatively long periods of stasis, where species remain relatively unchanged.  In this theory, speciation and rapid evolution are linked, with natural selection and genetic drift acting most strongly on organisms undergoing speciation in novel habitats or small populations. As a result, the periods of stasis in the fossil record correspond to the parental population, and the organisms undergoing speciation and rapid evolution are found in small populations or geographically restricted habitats, and therefore rarely being preserved as fossil.  This process may have resulted in the introduction of new species including Man.

Evolutionary History of Life

Origin of life 
The origin of life is a necessary precursor for biological evolution, but understanding that evolution occurred once organisms appeared and investigating how this happens does not depend on understanding exactly how life began.  The current scientific consensus is that the complex biochemistry that makes up life came from simpler chemical reactions, but it is unclear how this occurred.  Not much is certain about the earliest developments in life, the structure of the first living things, or the identity and nature of any last universal common ancestor or ancestral gene pool.  Consequently, there is no scientific consensus on how life began, but proposals include self-replicating molecules such as RNA, and the assembly of simple cells.

Common descent 

The hominoids are descendants of a common ancestor.

All organisms on Earth are descended from a common ancestor or ancestral gene pool.  Current species are a stage in the process of evolution, with their diversity the product of a long series of speciation and extinction events.  The common descent of organisms was first deduced from four simple facts about organisms: First, they have geographic distributions that cannot be explained by local adaptation. Second, the diversity of life is not a set of completely unique organisms, but organisms that share morphological similarities. Third, vestigial traits with no clear purpose resemble functional ancestral traits, and finally, that organisms can be classified using these similarities into a hierarchy of nested groups – similar to a family tree. However, modern research has suggested that, due to horizontal gene transfer, this “tree of life” may be more complicated than a simple branching tree since some genes have spread independently between distantly related species.
Past species have also left records of their evolutionary history. Fossils, along with the comparative anatomy of present-day organisms, constitute the morphological, or anatomical, record.  By comparing the anatomies of both modern and extinct species, paleontologists can infer the lineages of those species. However, this approach is most successful for organisms that had hard body parts, such as shells, bones or teeth
More recently, evidence for common descent has come from the study of biochemical similarities between organisms. For example, all living cells use the same basic set of nucleotides and amino acids.  The development of molecular genetics has revealed the record of evolution left in organisms’ genomes: dating when species diverged through the molecular clock produced by mutations.  For example, these DNA sequence comparisons have revealed that humans and chimpanzees share 96% of their genomes and analyzing the few areas where they differ helps shed light on when the common ancestor of these species existed.

Evolution of life

Evolutionary tree showing the divergence of modern species from their common ancestor in the center.  The three domains are colored, with bacteria blue, archaea green, and eukaryotes red.

Despite the uncertainty on how life began, it is generally accepted that prokaryotes inhabited the Earth from approximately 3–4 billion years ago.  No obvious changes in morphology or cellular organization occurred in these organisms over the next few billion years.

The eukaryotes were the next major change in cell structure. These came from ancient bacteria being engulfed by the ancestors of eukaryotic cells, in a cooperative association called endosymbiosis.  The engulfed bacteria and the host cell then underwent co-evolution, with the bacteria evolving into either mitochondria or hydrogenosomes.  An independent second engulfment of cyanobacterial-like organisms led to the formation of chloroplasts in algae and plants.   It is unknown when the first eukaryotic cells appeared though they first emerged between 1.6 – 2.7 billion years ago.

The history of life was that of the unicellular eukaryotes, prokaryotes, and archaea until about 610 million years ago when multicellular organisms began to appear in the oceans in the Ediacaran period.  The evolution of multicellularity occurred in multiple independent events, in organisms as diverse as sponges, brown algae, cyanobacteria, slime moulds and myxobacteria.

Soon after the emergence of these first multicellular organisms, a remarkable amount of biological diversity appeared over approximately 10 million years, in an event called the Cambrian explosion. Here, the majority of types of modern animals appeared in the fossil record, as well as unique lineages that subsequently became extinct. Various triggers for the Cambrian explosion have been proposed, including the accumulation of oxygen in the atmosphere from photosynthesis.   About 500 million years ago, plants and fungi colonized the land, and were soon followed by arthropods and other animals.   Insects were particularly successful and even today make up the majority of animal species.   Amphibians first appeared around 300 million years ago, followed by early amniotes, then mammals around 200 million years ago and birds around 100 million years ago (both from “reptile”-like lineages). However, despite the evolution of these large animals, smaller organisms similar to the types that evolved early in this process continue to be highly successful and dominate the Earth, with the majority of both biomass and species being prokaryotes.