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A. DEFINITION The word homology, coined in about 1656, derives from the Greek homologos, where homo = agreeing, equivalent, same + logos = relation. In biology, two things are homologous if they bear the same relationship to one another, such as a certain bone in various forms of the "hand." In 1843, Richard Owen defined homology as "the same organ in different animals under every variety of form and function". Organs as different as a bat's wing, a seal's flipper, a cat's paw and a human hand have a common underlying structure of bones and muscles. Owen reasoned that there must be a common structural plan for all vertebrates, as well as for each class of vertebrates (Anonymousa. 2012). Richard Owen (1848) introduced the term homology to refer to structural similarities among organisms. To Owen, these similarities indicated that organisms were created following a common plan or archetype. That is, although each species is unique, the plans for each might share many features. Nevertheless, if every organism were created independently, it is unclear why there would be so many homologies among certain organisms, while so few among others. It is also hard to make sense of the fact that homologous structures can be inefficient or even useless (Anonymousb. 2012). Homology began as a philosophical concept that certain physical attributes share similarity in form and position among species in a natural group. Within the recognized vertebrate groups, early comparative anatomists had noted that individual bones of vertebrate skeletons showed positional correspondence among specific elements, e.g. the forelimb of reptiles, chickens, dogs and humans. Such congruent configurations are classified as homologous and the corresponding parts as homologues (Figure 1a). In the case of complex structures composed of repetitive elements, e.g. the vertebrae of the spinal column, such

elements are termed serially homologous. With the introduction of the principle of descent with modification (Darwin, 1859), the concept evolved beyond the simple one of topological similarity from an idealized type into an understanding that such similarities arise through inheritance and common ancestry (Figure 1b).

Figure 1. Concepts of homology. (a) Owens homology was a concept of corresponding structures between organisms as similar or homologous based on their relationship to the archetype. (b) After Darwin, the similarity in structure is understood to arise from ancestor/descendant relationships. Owen (1843: 379) defined homology simply as the same organ in different animals under every variety of form and function. He also distinguished homology from analogy, which he defined as a part or organ in one animal which has the same function as another part or organ in a different animal (Owen, 1843: 374). Here, homology is recognized as an innate property

of a group of organisms that is associated with a specific archetype, or idealized form. Without understanding of genetics or modes of inheritance, Owen and his predecessors were cognizant of fundamental groups of organisms that shared characteristics (Figure 1a ). Such information was used in classification of organisms into natural groups and was a forerunner to later revelations of evolution and natural selection (Anonymousc. 2012). Homology and analogy are argued by Owen to not be mutually exclusive terms (Panchen, 1994), i.e. homologues may have analogous functions or may evolve wholly different functions. The adaptation of homologous structures into new uses is the root of animal diversity and comparative biology. An understanding of the full variation of homologues in nature provides insight into the broad adaptive response of organisms possessing that homologue. Classic examples of such modifications are the development of ear ossicles from homologous structures found in the hinge of the reptilian jaw and the modification into flowers of complex arrays of leaves (de Beer, 1971) (Anonymousc. 2012). B. Developmental Genetics and Homology In the early 1800s, it was understood that similar animals shared patterns in development as surely as they shared patterns in adult morphology. These similarities in developmental pattern led vonBaer (1828) to outline four principles of embryology. Paraphrased, they are: (1) general features of a group appear earlier in the embryo than do the specialized features, (2) less general characters are developed from the more general, (3) the embryo of a given species does not share features with adults of other species but instead shares embryonic features that diverge in the two adults, and (4) therefore, the early embryo of a higher animal isnot like an adult of a lower animal, but they share features (homology) in early development. (Anonymousc. 2012). Characteristics are shared among organisms because of the common developmental pattern of natural groups, not from the transmutation of lower

animals into higherones. The discovery of homologous larval forms of unique animals has increased our understanding of evolution in these groups. Barnacles have a nauplius larva that relates them to crustaceans that possess a similar larva, instead of to molluscs with which they were once considered to have affinities. Sea squirts, or ascidians, were discovered to have a notochord in their larvae, homologous to that of other chordates, even though no vestige of it remains in the adult animal. Similar patterns are seen in other vertebrates: snakes possess pelvic girdles and baleen whales possess tooth rudiments. Even humans possess a blastodisc embryo similar to other amniotes, because our ancestors were constrained to develop on the surface of a large, yolky egg. Such shared homology in development continues to be at the core of evolutionary comparative development and embryology (Anonymousc. 2012). However, developmental pattern alone is not evidence of homology or the lack thereof. Several examples of pitfalls are outlined by de Beer (1971). The two most straightforward examples involve the formation of vertebrate alimentary canals and the lens formation in the eye. The origin of the alimentary canal varies from the dorsal surface of the embryonic gut in sharks, the ventral surface in lampreys, dorsal or ventral surface in different frogs, or from the lower blastodermal layer in reptiles and birds. Does this make the alimentary canal nonhomologous among these taxa? Lens formation in the closely related frogs Rana fusca and R. esculenta occurs differently. In R. fusca, the optic cup induces the lens to form. A small invagination of the overlying ectoderm of the eye is induced by the optic cup coming in close proximity. However, in developmental experiments with R. esculenta, the lens forms in a similar process whether the optic cup is present or not. Are the lenses nonhomologous? Clearly, the answer is no for both cases. While the source tissues that produce or induce structures can vary among taxa, the homologous structures that result from them are homologous among the descendant taxa due to common ancestry. The origin of a homologous structure or its developmental mode of induction can change even

between closely related species, but it does not discount the homology of those structures (Anonymousc. 2012). C. Homology as the proof of evolution Called homology or the homology theory, since Darwin this view has been presented as a major evidence of macroevolution theory. An example of this reasoning is as follows: If you look at a 1953 Corvette and compare it to the latest model, only the most general resemblances are evident, but if you compare a 1953 and a 1954 Corvette, side by side, then a 1954 and a 1955 model, and so on, the descent with modification is overwhelmingly obvious. This is what paleontologists do with fossils, and the evidence is so solid and comprehensive that it cannot be denied by reasonable people [emphasis in original]. (Anonymousd. 2012). Homology is not merely a minor proof of evolution, but instead has been widely cited by evolutionists as one of the most compelling lines of evidence for their theory. Darwin concluded that homology was critically important evidence for common descent: Darwin reasoned that the members of the same class of animals resemble each other in the general plan of their design and, in his words, this resemblance is critical because of the fact that the hand of a man, formed for grasping, that of a mole for digging, the leg of the horse, the paddle of the porpoise and the wing of the bat are all constructed on the same pattern and include similar bones in the same relative positions is specifically what the theory of common descent would expect.5 An early example of how homology was used to argue for macroevolution is a 1928 biology text which, in answer to the question Why do the individuals in a species have all of their parts homologous?, said: The obvious answer is, that they all descended from the same ancestors. Biologists carry this answer a step further and say that since homology within the species is the result of common ancestry therefore all homology is due to common ancestry and the closeness of relationship

determines the number of homologous parts [emphasis in original]. (Anonymousd. 2012). The seven bones in the human neck correspond with the same seven, much larger, neckbones in the giraffe: they are homologues. The number of cervical vertebrae is a trait shared by creatures descended from a common ancestor. Related species share corresponding structures, though they may be modified in various ways (Anonymousd. 2012). A much more recent quote illustrates how this line of reasoning is still being used today to argue that the evidence of homology for the common ancestry of all life is very strong. Why is it that bats and whales have so much in common anatomically with mice and men? Why do virtually all vertebrate forelimbs have the same basic pentadactyl (five fingered) design? (This is one of numerous examples of homologous structures exhibited by related species.) The author concludes the answer is evolution. Barr lists homology as the first argument on his list of evidence for evolution. As the above quotes show, the same line of reasoning has been used to prove evolution for more than a century. However, Dobzhansky admitted that homology does not prove evolution, in the sense that nobody has actually witnessed the gradual changes in the millions of consecutive generations which led from a common ancestor to a bird on the one hand and to man on the other . But, he adds, homology strongly suggests evolution; the facts of homology make sense if they are supposed to be due to evolution of now-different organisms from a common stock. They do not make sense otherwise. (Anonymousd. 2012). The division is based on a distinction between similarity due to common ancestry, or homology, and resemblance which is due solely to similarity of function, called analogy. An example is the forelimbs of humans, horses, whales and birds which are judged homologous because they are all constructed on the same pattern, and include similar bones in the same relative positions because

these are all derived from the same ancestral bones. The wings of birds and insects, on the other hand, are analogous: they serve the same purpose, but do not constitute modified versions of a structure present in a common ancestor. The wings of birds and bats are homologous in skeletal structure because of descent from the forelimb of a common reptilian ancestor; but they are analogous in terms of their modification for flightfeathers in birds, skin membranes in bats. In other words, if a design similarity supports evolutionary assumptions, it is listed as an homology and is accepted as evidence for evolution. Conversely, if a design similarity does not support evolution, it is called analogy, and the conclusion is drawn that the similarity exists because a certain design is highly functional for a specific body part, and not because of a common ancestor. Many analogous structures are assumed to exist due to convergent evolution, which is defined as the separate evolution of similar structures because of similar environmental demands. Convergent evolution also is used to explain similar structures that have formed from different embryo structures or precursors. Many examples of homology are actually better explained by analogy, and the resemblance that exists is often due to similarity of function and/or design constraints. The forelimbs of humans, whales and birds are similar because they serve similar functions and have similar design constraints. The conclusion that two homologous bones are similar because they are putatively derived from the same ancestral bones (as Barr claims) is not based on direct evidence but instead on a priori conclusions demanded by macroevolution. Jones concluded that Convergent evolution has been hypothesized to explain the numerous examples of homology in which the available evidence suggested that the animals under consideration were not closely linked by descent. An example of such gratuitous hypothesizing is the following: Some similarities between distant species may be caused by adaptation to similar environments, which is known as convergent evolution. Development of streamlined fins in fish (teleosts) and flippers in dolphins (mammals) are

analogous: they function alike, but are very different in underlying structure. . . . Linnaeuss original classification of animals does not distinguish between analogous and homologous structures. Creatures were often put in the same groups by resemblances to an imagined divine plan or design. Since Darwin . . . species are classified to reflect the relative closeness or distance of their common ancestry. (Anonymousd. 2012).

DAFTAR PUSTAKA Anonymousa. 2012. Homologi. Diakses pada maret 2012.

Anonymousb. 2012. Homology Makes Sense Evolution. Diakses pada maret 2012. AnonymousC. 2012. Homology In Character Evolution. Diakses pada maret 2012. Anonymousd. 2012. Does Homology Provide Evidence of Evolutionary Naturalism? . Diakses pada maret 2012.

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:// KUSNADI/BUKU_SAKU_BIOLOGI_SMA,KUSNADI_dkk/Kelas_ XII/7._Evolusi/EVOLUSI_ZV.pdf 3. homology 4. homology make sense evolution 4 tm

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