For other uses, see Homology (disambiguation).
A common example of homologous structures is the forelimbs of vertebrates, where the wings of bats and birds, the arms of primates, the front flippers of whales and the forelegs of four-legged vertebrates like dogs and crocodiles are all derived from the same ancestral tetrapod structure.
The term was first applied to biology in a non-evolutionary context by the anatomist Richard Owen in 1843.
In developmental biology, organs that developed in the embryo in the same manner and from similar origins, such as from matching primordia in successive segments of the same animal, are serially homologous.
Homology among proteins or DNA is inferred from their sequence similarity.
Significant similarity is strong evidence that two sequences are related by divergent evolution from a common ancestor.
Alignments of multiple sequences are used to discover the homologous regions.
In the German Naturphilosophie tradition, homology was of special interest as demonstrating unity in nature.
The serial homology of limbs was described late in the 18th century.
The French zoologist Etienne Geoffroy Saint-Hilaire showed in 1818 in his theorie d'analogue ("theory of homologues") that structures were shared between fishes, reptiles, birds, and mammals.
Geoffroy stated the principle of connections, namely that what is important is the relative position of different structures and their connections to each other.
The Estonian embryologist Karl Ernst von Baer stated what are now called von Baer's laws in 1828, noting that related animals begin their development as similar embryos and then diverge: thus, animals in the same family are more closely related and diverge later than animals which are only in the same order and have fewer homologies.
von Baer's theory recognises that each taxon (such as a family) has distinctive shared features, and that embryonic development parallels the taxonomic hierarchy: not the same as recapitulation theory.
The term "homology" was first used in biology by the anatomist Richard Owen in 1843 when studying the similarities of vertebrate fins and limbs, defining it as the "same organ in different animals under every variety of form and function", and contrasting it with the matching term "analogy" which he used to describe different structures with the same function.
Owen codified 3 main criteria for determining if features were homologous: position, development, and composition.
The word homology, coined in about 1656, is derived from the Greek ὁμόλογος homologos from ὁμός homos "same" and λόγος logos "relation".
Homology thus implies divergent evolution.
Similarly, the forelimbs of ancestral vertebrates have evolved into the front flippers of whales, the wings of birds, the running forelegs of dogs, deer, and horses, the short forelegs of frogs and lizards, and the grasping hands of primates including humans.
Homology vs analogy
Further information: Convergent evolution
Similarly, the wings of a sycamore maple seed and the wings of a bird are analogous but not homologous, as they develop from quite different structures.
A structure can be homologous at one level, but only analogous at another.
Pterosaur, bird and bat wings are analogous as wings, but homologous as forelimbs because the organ served as a forearm (not a wing) in the last common ancestor of tetrapods, and evolved in different ways in the three groups.
Thus, in the pterosaurs, the "wing" involves both the forelimb and the hindlimb.
Further information: Cladistics
Specialised terms are used in taxonomic research.
Primary homology is a researcher's initial hypothesis based on similar structure or anatomical connections, suggesting that a character state in two or more taxa share is shared due to common ancestry.
Primary homology may be conceptually broken down further: we may consider all of the states of the same character as "homologous" parts of a single, unspecified, transformation series.
This has been referred to as topographical correspondence.
For example, in an aligned DNA sequence matrix, all of the A, G, C, T or implied gaps at a given nucleotide site are homologous in this way.
Character state identity is the hypothesis that the particular condition in two or more taxa is "the same" as far as our character coding scheme is concerned.
Thus, two Adenines at the same aligned nucleotide site are hypothesized to be homologous unless that hypothesis is subsequently contradicted by other evidence.
Secondary homology is implied by parsimony analysis, where a character state that arises only once on a tree is taken to be homologous.
Shared ancestral character states, symplesiomorphies, represent either synapomorphies of a more inclusive group, or complementary states (often absences) that unite no natural group of organisms.
For example, the presence of wings is a synapomorphy for pterygote insects, but a symplesiomorphy for holometabolous insects.
Absence of wings in non-pterygote insects and other organisms is a complementary symplesiomorphy that unites no group (for example, absence of wings provides no evidence of common ancestry of silverfish, spiders and annelid worms).
On the other hand, absence (or secondary loss) of wings is a synapomorphy for fleas.
Patterns such as these lead many cladists to consider the concept of homology and the concept of synapomorphy to be equivalent.
Some cladists follow the pre-cladistic definition of homology of Haas and Simpson, and view both synapomorphies and symplesiomorphies as homologous character states
In different taxa
Homologies provide the fundamental basis for all biological classification, although some may be highly counter-intuitive.
For example, deep homologies like the pax6 genes that control the development of the eyes of vertebrates and arthropods were unexpected, as the organs are anatomically dissimilar and appeared to have evolved entirely independently.
Further information: Arthropod leg
The embryonic body segments (somites) of different arthropod taxa have diverged from a simple body plan with many similar appendages which are serially homologous, into a variety of body plans with fewer segments equipped with specialised appendages.
|1||antennae||chelicerae (jaws and fangs)||antennae||antennae||1st antennae|
|2||1st legs||pedipalps||-||-||2nd antennae|
|3||2nd legs||1st legs||mandibles||mandibles||mandibles (jaws)|
|4||3rd legs||2nd legs||1st maxillae||1st maxillae||1st maxillae|
|5||4th legs||3rd legs||2nd maxillae||2nd maxillae||2nd maxillae|
|6||5th legs||4th legs||collum (no legs)||1st legs||1st legs|
|7||6th legs||-||1st legs||2nd legs||2nd legs|
|8||7th legs||-||2nd legs||3rd legs||3rd legs|
|9||8th legs||-||3rd legs||-||4th legs|
|10||9th legs||-||4th legs||-||5th legs|
Further information: Comparative anatomy
The malleus and incus develop in the embryo from structures that form jaw bones (the quadrate and the articular) in lizards, and in fossils of lizard-like ancestors of mammals.
Both lines of evidence show that these bones are homologous, sharing a common ancestor.
Rudimentary organs such as the human tailbone, now much reduced from their functional state, are readily understood as signs of evolution, the explanation being that they were cut down by natural selection from functioning organs when their functions were no longer needed, but make no sense at all if species are considered to be fixed.
The tailbone is homologous to the tails of other primates.
Leaves, stems, and roots
Leaves are variously modified from photosynthetic structures to form the insect-trapping pitchers of pitcher plants, the insect-trapping jaws of Venus flytrap, and the spines of cactuses, all homologous.
|Primary organs||Defensive structures||Storage structures|
|Leaves||Spines||Swollen leaves (e.g. succulents)|
|Stems||Thorns||Tubers (e.g. potato), rhizomes (e.g. ginger), fleshy stems (e.g. cacti)|
|Roots||-||Root tubers (e.g. sweet potato), taproot (e.g. carrot)|
Further information: ABC model of flower development
Each of the four types of flower parts is serially repeated in concentric whorls, controlled by a small number of genes acting in various combinations.
Thus, A genes working alone result in sepal formation; A and B together produce petals; B and C together create stamens; C alone produces carpels.
When none of the genes are active, leaves are formed.
Two more groups of genes, D to form ovules and E for the floral whorls, complete the model.
The genes are evidently ancient, as old as the flowering plants themselves.
For example, adult snakes have no legs, but their early embryos have limb-buds for hind legs, which are soon lost as the embryos develop.
The implication that the ancestors of snakes had hind legs is confirmed by fossil evidence: the Cretaceous snake Pachyrhachis problematicus had hind legs complete with hip bones (ilium, pubis, ischium), thigh bone (femur), leg bones (tibia, fibula) and foot bones (calcaneum, astragalus) as in tetrapods with legs today.
Main article: Sequence homology
Homology among proteins or DNA is typically inferred from their sequence similarity.
Significant similarity is strong evidence that two sequences are related by divergent evolution of a common ancestor.
Alignments of multiple sequences are used to indicate which regions of each sequence are homologous.
Homologous sequences are orthologous if they are descended from the same ancestral sequence separated by a speciation event: when a species diverges into two separate species, the copies of a single gene in the two resulting species are said to be orthologous.
Homologous sequences are paralogous if they were created by a duplication event within the genome.
For gene duplication events, if a gene in an organism is duplicated to occupy two different positions in the same genome, then the two copies are paralogous.
Paralogous genes often belong to the same species.
They can shape the structure of whole genomes and thus explain genome evolution to a large extent.
As a result, Hox genes in most vertebrates are spread across multiple chromosomes: the HoxA–D clusters are the best studied.
Main article: Homology (psychology)
It has been suggested that some behaviours might be homologous, based either on sharing across related taxa or on common origins of the behaviour in an individual's development; however, the notion of homologous behavior remains controversial, largely because behavior is more prone to multiple realizability than other biological traits.
As with morphological features or DNA, shared similarity in behavior provides evidence for common ancestry.
The hypothesis that a behavioral character is not homologous should be based on an incongruent distribution of that character with respect to other features that are presumed to reflect the true pattern of relationships.
This is an application of Willi Hennig's auxiliary principle.
Credits to the contents of this page go to the authors of the corresponding Wikipedia page: en.wikipedia.org/wiki/Homology (biology).