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The Lung-Swimbladder Issue: A Simple Case of Homology – Or Not?

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... (Darwin, 1859, chapter Simpson (1953) and by Mayr in his article on novelty (1960) Stephen Jay Gould has stressed that Darwin's explanation is "not only wrong, but backwards" (Gould, 2002(Gould, , p. 1224) The hypothesis favoured by Gould (that lung is the ancestral and swimbladder the derived condition) was actually already the dominant one in the Modern Synthesis era, as noted by Mayr and Simpson (Simpson, 1953, p. 192, note 11;Mayr, 1960, p. 352). The case is still debated today The main competing hypothesis is the independent derivation of both organs from a respiratory pharynx (Lambertz and Perry, 2015;Perry and Sander, 2004 the subsidiary function gradually becomes the chief function, the total function becomes quite different, and the consequence of the whole process is the transformation of the organ (Dohrn 1875, p60 cited by Russel, 1916). ...
Thesis
Full text available on request or in open access at https://ore.exeter.ac.uk/repository/handle/10871/35377# Evolutionary novelty, the origin of new characters such as the turtle shell or the flower, is a fundamental problem for an evolutionary view of life. Accordingly, it is a central research topic in contemporary biology involving input from several biological disciplines and explanations at several levels of organization. I study the evolution of research on novelty from the 1950s to the present. The problem of novelty has recently been appropriated by evolutionary developmental biology or evo-devo, a synthesis of evolutionary and developmental biology that started emerging in the 1980s following technological advances and discoveries in developmental genetics. I focus instead on three neglected dimensions of the problem of novelty: the functional-historical approach to the problem, research on novelty in the late Modern Synthesis era (1950-1980) and novelty in plants. My argument runs against the view of some scientists and historians, often tied to evo-devo, who oppose structuralist and functionalist approaches in biology and who claim that the origin of novelty is a structuralist problem. I advocate an approach to novelty that ties together structural and functional dimensions and show how some research programs of the last eighty years implemented different versions of this approach.
... Although parsimony is a key tool in phylogenetic reconstructions , it alone ultimately is in and of itself not a definitive indicator for the homology of traits, and additional organismic considerations are required (see e.g. Wagner, 2014; Lambertz and Perry, 2015 ). Indeed, the ossification sequence and mode of the sternum formation in enantiornithine birds, the dominant clade of Mesozoic birds (e.g., Zhou, 2004 ) here represented by Eopengornis , Longipteryx and Bohaiornis, is different from that of its sister taxon, the ornithuromorphs, which also include the modern birds (Zheng et al., 2012). ...
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This paper summarizes the main morphological tracts exhibited by lungs and gas bladders in fishes. The origin and organ location, the presence of a glottal region, the inner architecture, the characteristics of the exchange barrier and the presence of pulmonary arteries have been reviewed in the two types of air-breathing organs. With the exception of the dorsal (bladders) or ventral (lungs) origin from the posterior pharynx, none of the morphological traits analyzed can be considered specific for either lungs or gas bladders. This is exemplified by analysis of the morphology of the lung of the Dipnoii and Polypteriformes and of the bladder of the Lepisosteiformes. All of them are obligate air-breathers and show a lung-like (pulmonoid) air-breathing organ. However, while the lungfish lung and the bladder of the Lepisosteiformes occupy a dorsal position and are highly trabeculated, the polypterid lung occupies a ventral position and shows a smooth inner surface. Structural and ultrastructural differences are also highlighted. Noticeably, a large part of the inner surface area of the lung of the Australian lungfish is covered by a ciliated epithelium. A restricted respiratory surface area may help to explain the incapability of this species to aestivate. The respiratory bladder of basal teleosts displays a more complex morphology than that observed in more primitive species. The bladder of basal teleosts may appear divided into respiratory and non-respiratory portions, exhibit intricate shapes, invade adjacent structures and gain additional functions. The increase in morphological and functional complexity appears to prelude the loss of the respiratory functions.
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The coelacanth, Latimeria chalumnae Smith, 1939 [1] (Sarcopterygii: Actinistia), together with the closely related L. menadoensis Pouyaud et al., 1999 [2], remains the only living representative of one of the most basally-branching primary radiations of lobe-finned fishes (Sarcopterygii). Even though extant species cannot be considered 'primitive' due to the inherent logic of phylogenetic theory, the coelacanth nonetheless is invaluable for understanding evolutionary transformations in basal sarcopterygians as it can help in the determination of character polarity. The appearance of one novelty during early vertebrate evolution that had major implications for the success of a huge number of species is the origin of lungs. The conventional interpretation is that lungs evolved in basal bony fishes (Osteichthyes or Osteognathostomata), were maintained in the lobe-finned fishes, and eventually were transformed into a swimbladder among the ray-finned fishes (Actinopterygii) (e.g. [3]). However, the currently available data do not rule out separate origins of lungs and swimbladders from a common 'respiratory pharynx', even though this would require a slightly less parsimonious course of evolution [4,5]. The coelacanth is a key species in addressing this question and for this reason the data recently provided by Cupello and colleagues [6] are a very welcome addition to the discussion. Here, I would like to add a few points pertinent to lung evolution that appear to be a consequence of these exciting data. One of the most interesting aspects of the coelacanth is that it apparently exhibits an unpaired structure of putative homology with lungs [6-8]. In the Polypteriformes (bichir and reed fish), the lungs are paired [5,9,10], as are those of the lungfishes (Dipnoi) [11], except the Australian lungfish, Neoceratodus forsteri (Krefft, 1870)….
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A recessive mutation was identified in a family of transgenic mice that resulted in a reversal of left-right polarity (situs inversus) in 100 percent of the homozygous transgenic mice tested. Sequences that flanked the transgenic integration site were cloned and mapped to mouse chromosome 4, between the Tsha and Hxb loci. During early embryonic development, the direction of postimplantation turning, one of the earliest manifestations of left-right asymmetry, was reversed in homozygous transgenic embryos. This insertional mutation identifies a gene that controls embryonic turning and visceral left-right polarity.
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Lungs are the characteristic air-filled organs (AO) of the Polypteriformes, lungfish and tetrapods, whereas the swimbladder is ancestral in all other bony fish. Lungs are paired ventral derivatives of the pharynx posterior to the gills. Their respiratory blood supply is the sixth branchial artery and the venous outflow enters the heart separately from systemic and portal blood at the sinus venosus (Polypteriformes) or the atrium (lungfish), or is delivered to a separate left atrium (tetrapods). The swimbladder, on the other hand, is unpaired, and arises dorsally from the posterior pharynx. It is employed in breathing in Ginglymodi (gars), Halecomorphi (bowfin) and in basal teleosts. In most cases, its respiratory blood supply is homologous to that of the lung, but the vein drains to the cardinal veins. Separate intercardiac channels for oxygenated and deoxygenated blood are lacking. The question of the homology of lungs and swimbladders and of breathing mechanisms remains open. On the whole, air ventilatory mechanisms in the actinopterygian lineage are similar among different groups, including Polypteriformes, but are distinct from those of lungfish and tetrapods. However, there is extreme variation within this apparent dichotomy. Furthermore, the possible separate origin of air breathing in actinopterygian and 'sarcopterygian' lines is in conflict with the postulated much more ancient origin of vertebrate air-breathing organs. New studies on the isolated brainstem preparation of the gar (Lepisosteus osseus) show a pattern of efferent activity associated with a glottal opening that is remarkably similar to that seen in the in-vitro brainstem preparation of frogs and tadpoles. Given the complete lack of evidence for AO in chondrichthyans, and the isolated position of placoderms for which buoyancy organs of uncertain homology have been demonstrated, it is likely that homologous pharyngeal AO arose in the ancestors of early bony fish, and was pre-dated by behavioral mechanisms for surface (water) breathing. The primitive AO may have been the posterior gill pouches or even the modified gills themselves, served by the sixth branchial artery. Further development of the dorsal part may have led to the respiratory swimbladder, whereas the paired ventral parts evolved into lungs.
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Convergence is an important evolutionary phenomenon often attributed solely to natural selection acting in similar environments. The frequency of mutation and number of ways a phenotypic trait can be generated genetically, however, may also affect the probability of convergence. Here we report both a high frequency of loss of gas bladder (swim bladder) mutations in zebrafish and widespread convergent loss of gas bladders among teleost fishes. The phenotypes of 22 of 27 recessive lethal mutations, carried by a sample of 26 wild-caught zebrafish, involve loss or noninflation of the gas bladder. Nine of these bladderless mutations showed no other obvious phenotypic abnormalities other than the lack of an inflated gas bladder. At least 19 of the 22 bladderless mutations are genetically distinct, as shown by unique morphology or complementation. Although we were not able to obtain eggs for all 21 required crosses to demonstrate definitively that the remaining three mutations are different from all other bladderless mutations, all available evidence suggests that these mutants are also distinct. At least 79 of 425 families of extant teleosts include one or more species lacking a gas bladder as adults. Analysis of the trait's phylogenetic distribution shows that the gas bladder has been lost at least 30-32 times independently. Although adaptive explanations for gas bladder loss are convincing, a developmental bias toward bladderless phenotypes may also have contributed to the widespread convergence of this trait among teleosts. If gas bladder development in teleosts is as vulnerable to genetic perturbation as it is in zebrafish, then perhaps a supply of bladderless phenotypes has been readily available to natural selection under conditions for which it is advantageous not to have a gas bladder. In this way, developmental bias and selection can work together to produce widespread convergence.
Article
Homology is an essential idea of biology, referring to the historical continuity of characters, but it is also conceptually highly elusive. The main difficulty is the apparently loose relationship between morphological characters and their genetic basis. Here I propose that it is the historical continuity of gene regulatory networks rather than the expression of individual homologous genes that underlies the homology of morphological characters. These networks, here referred to as 'character identity networks', enable the execution of a character-specific developmental programme.
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Mutations in the DNAH11 (axonemal heavy chain dynein type 11) gene cause one form of situs inversus totalis and most likely primary ciliary dyskinesia
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Bartolini, L., J.-L. Blouin, Y. Pan, C. Gehrig, A.K. Maiti, N. Scamuffa, C. Rossier, M. Jorissen, M. Armengot, M. Meeks, H.M. Mitchison, E.M.K. Chung, C.D. Delozier-Blanchet, W.J. Craigen and S.E. Antonarakis. 2002. Mutations in the DNAH11 (axonemal heavy chain dynein type 11) gene cause one form of situs inversus totalis and most likely primary ciliary dyskinesia. Proc. Natl. Acad. Sci. USA 99: 10282-10286.
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