Skeletal elements within teleost eyes and a discussion of their homology

Scleral ossicles and scleral cartilages form part of the craniofacial skeleton of many vertebrates. Some vertebrates, including all birds and most reptiles, but excluding most mammals, have scleral cartilages as well as scleral ossicles supporting their eyes. The teleost equivalent of these elements has received little attention in the literature. From radiographic and whole-mount analyses of over 400 individuals from 376 teleost species, we conclude that the teleost scleral skeletal elements (ossicles and cartilage) differ significantly from those of reptiles (including birds). Scleral ossicles in teleosts have different developmental origins, different positions within the eyeball, and different relationships with the scleral cartilaginous element than those in reptiles. From whole-mount staining of a growth series of four species of teleost (Danio rerio, Salmo salar, Esox lucius, and Alosa pseudoharengus), we interpret the development of these elements and show that they arise from within an Alcian blue-staining cartilaginous ring that develops around the eye earlier in development. We present possible scenarios on the evolution of these scleral skeletal elements from a common gnathostome ancestor, and consider that teleost scleral skeletal elements may not be homologous to those in reptiles. Our study indicates that homology cannot be assumed for these elements, despite the fact that they share the same name, scleral ossicles.     

Effects of different kinds of cranial deformation on the incidence of wormian bones

Researchers have debated whether the presence and frequency of wormian bones (sutural bones, supernumerary bones, and ossicles) are attributable to genetic factors, environmental factors, or both. This research examines the effects of many different kinds of cranial deformation on the incidence of wormian bones. A sample of 127 deformed and undeformed crania from New World archaeological sites was examined. An undeformed cranial sample (n=35) was compared to the following cranially deformed groups: 1) occipital, 2) lambdoid, 3) annular, 4) fronto-vertico-occipital, 5) parallelo-fronto-occipital, and 6) sagittal synostosis. Three levels of degree of cultural cranial deformation were qualitatively determined. Type and number of wormian bones along each major suture were recorded for each cranium. Group means were analyzed using Kruskal-Wallis one-way ANOVA statistical tests to test the null hypothesis that cranial deformation does not have an effect on wormian bone incidence. Results indicate that all forms of cranial deformation affect the frequency of some types of wormian bones. In particular, all cranially deformed groups exhibited significantly greater frequencies of lambdoid ossicles. Apical, parieto-mastoid, and occipito-mastoid wormian bones also appeared with greater frequency in some groups of culturally deformed crania. Further, varying degrees of cultural deformation all had more lambdoid wormian bones than the undeformed group. These results suggest that wormian bone development in posteriorly placed sutures may be affected more by environmental forces than are their anteriorly placed counterparts.     

The ear ossicles and the evolution of the primate ear: A biomechanical approach

From a morphometric viewpoint the variability of human and other primate ear ossicles appears to be suitable for the study of taxonomic and phylogenetic distinction among Primates. It may also be of interest to determine whether they are useful to show differences in the perception of sound from the environment and from conspecifics. The energy transmitted through the ossicles is mantained by the action of different leverages. These modify the action of the ossicles from relatively wide, low energy, movements of the hammer to the smaller, high energy, movements of the stirrup. It seems that the pongid type of ossicle leverage combination saves more energy, possibly with a certain loss of subtle information, but this may be more useful in the wild than decoding voice modulation. The human type leverage, being less demultiplied, may produce a major loss of energy but, perhaps, a more precise conservation of sound information useful for speech communication.     

Phylogenetic analysis of the Brachyura (Crustacea, Decapoda) based on characters of the foregut with establishment of a new taxon

The Brachyura, within the decapod crustaceans, is one of the most species-rich taxa with up to 10 000 species. However, its phylogenetic history, evolution and fossil record remain subjects of controversy. In our study, we examined the phylogenetic relationships of the Brachyura based on morphological characters of the foregut. The cladistic analysis supports a monophyletic Brachyura including the Dromiidae and Raninidae. A clade comprising Dromiidae and Dynomenidae forms the most basal assemblage within the Brachyura, followed by the Homolidae and Latreilliidae. As a result, neither Podotremata nor Archaeobrachyura form a clade. In contrast, foregut data suggest that the classical taxon Oxystomata, comprising Calappidae, Parthenopidae, Dorippidae, Leucosiidae, Cymonomidae and Raninidae, is monophyletic. This makes the Heterotremata paraphyletic or polyphyletic. A newly established taxon, Neobrachyura, embraces some representatives of the Heterotremata and the monophyletic Thoracotremata.     

Study of age-related changes in Middle ear transfer function

Abstract

Osteoporosis (OP) is common with advancing age. Several studies have shown a strong correlation between OP and otosclerosis. However, no studies have investigated OP of the malleus, incus or stapes in the human middle ear, its effect on middle ear transfer function. Here, we investigate whether these three ossicles develop OP, and how this affects middle ear transfer function. The effect of OP on middle ear transfer function was investigated in simulations based on a finite element (FE) method. First, the FE model used in our previous study was refined, and optimized by introducing viscoelastic properties to selected soft tissues of the middle ear. Then, the FE model was used to simulate OP of the three ossicles and assess its influence on middle ear transfer function. Other possible age-related changes, such as stiffness of the joints or ligaments in the middle ear, were also investigated. The results indicated that OP of the ossicles could increase the high frequency displacement of both the umbo and stapes footplate (FP). However, the stiffness of the middle ear soft tissue can lead to the decrease of middle ear gain at lower frequencies. Furthermore, loosening of these joints or ligaments could increase displacement of the umbo and stapes FP. In conclusion, although age-related hearing loss is most commonly conceived of as sensorineural hearing loss (SNHL), we found that age-related changes may also include OP and changes in joint stiffness, but these will have little effect on middle ear transfer function in elderly people.     

Organs of hearing and equilibrium during ontogenesis of mammals

The peripheral part of acoustic analyzer was studied during pre-and postnatal development of mammals. The main trends of structural evolution of the outer, middle, and inner ear were followed in representatives of different ecological groups during postnatal development. The use of ecologomorphological approach made it possible to establish that specific structural features of the hearing organ in different mammals are determined by adaptation to specific acoustic properties of the environment. It was shown that morphofunctional adaptations directed at optimization of acoustic sensitivity in different environmental conditions were leading in the hearing organ evolution. Comparative-embryological studies of the peripheral part of acoustic system made it possible to determine the stages of formation of individual structures and establish general patterns of prenatal development of the organs of hearing and equilibrium in different mammals.     

Ontogenetic and phylogenetic transformations of the ear ossicles in marsupial mammals

This study is based on the examination of histological sections of specimens of different ages and of adult ossicles from macerated skulls representing a wide range of taxa and aims at addressing several issues concerning the evolution of the ear ossicles in marsupials. Three-dimensional reconstructions of the ear ossicles based on histological series were done for one or more stages of Monodelphis domestica, Caluromys philander, Sminthopsis virginiae, Trichosurus vulpecula, and Macropus rufogriseus. Several common trends were found. Portions of the ossicles that are phylogenetically older develop earlier than portions representing more recent evolutionary inventions (manubrium of the malleus, crus longum of the incus). The onset of endochondral ossification in the taxa in which this was examined followed the sequence; first malleus, then incus, and finally stapes. In M. domestica and C. philander at birth the yet precartilaginous ossicles form a supportive strut between the lower jaw and the braincase. The cartilage of Paauw develops relatively late in comparison with the ear ossicles and in close association to the tendon of the stapedial muscle. A feeble artery traverses the stapedial foramen of the stapes in the youngest stages of M. domestica, C. philander, and Sminthopsis virginiae examined. Presence of a large stapedial foramen is reconstructed in the groundplan of the Didelphidae and of Marsupialia. The stapedial foramen is absent in all adult caenolestids, dasyurids, Myrmecobius, Notoryctes, peramelids, vombatids, and phascolarctids. Pouch young of Perameles sp. and Dasyurus viverrinus show a bicrurate stapes with a sizeable stapedial foramen. Some didelphids examined to date show a double insertion of the Tensor tympani muscle. Some differences exist between M. domestica and C. philander in adult ossicle form, including the relative length of the incudal crus breve and of the stapes. Several differences exist between the malleus of didelphids and that of some phalangeriforms, the latter showing a short neck, absence of the lamina, and a ventrally directed manubrium. Hearing starts in M. domestica at an age in which the external auditory meatus has not yet fully developed, the ossicles are not fully ossified, and the middle ear space is partially filled with loose mesenchyme. The ontogenetic changes in hearing abilities in M. domestica between postnatal days 30 and 40 may be at least partially related to changes in middle ear structures.     

Histology and structural integration of the major morphologies of the Cypriniform Weberian apparatus

The Weberian apparatus, a diagnostic feature of otophysan fishes, is a novel hearing adaptation integrating several developmental and morphological systems (ear-vertebral column-swim bladder). Otophysan fishes are one of the largest and most successful freshwater clades, with over 10,000 species across most continents. The largest otophysan order, Cypriniformes, dominates the freshwaters of Asia, Europe, North America, and Africa. Spanning such a wide variety of environments, the Weberian apparatus undergoes morphological modifications to maintain functionality. Within Cypriniformes, we propose three distinct morphological classes of the Weberian apparatus based on the level of skeletal expansion around the swim bladder: simple (typical of most Cyprinidae), anterior plate (found in families such as Gyrinocheilidae, Catostomidae, and Botiidae), and encapsulated (either single-capsule as found, e.g., in Gobionidae and Cobitidae, or double-capsule as found, e.g., in Nemacheilidae and Balitoridae). Little ontological or comparative data exists regarding the construction or integration of these different morphologies, and less is known about the tissue level integration and variation within these morphologies. We used paraffin histology to document the hard and soft tissue anatomy of the Weberian apparatus in six species representing all morphological classes. We found sites of similarity across the morphologies including size and structure of the saccule, aspects of ossicle ossification, and swim bladder tunica composition, indicating potential sites of developmental and functional constraint. In contrast, we found differences across both auditory and nonauditory features in otic chamber size, ossification within ossicles and other vertebral elements, and composition of ligaments, indicating likely sites of adaptability. Some of these changes are likely evolutionary (taxonomic), but may be influenced by the environmental niche occupied by the clade. These results show a clear need for increased ontological and comparative study of the complete cypriniform Weberian apparatus, particularly histologically, as well as increased auditory studies across morphological types.     

Nocturnality in synapsids predates the origin of mammals by over 100 million years

Nocturnality is widespread among extant mammals and often considered the ancestral behavioural pattern for all mammals. However, mammals are nested within a larger clade, Synapsida, and non-mammalian synapsids comprise a rich phylogenetic, morphological and ecological diversity. Even though non-mammalian synapsids potentially could elucidate the early evolution of diel activity patterns and enrich the understanding of synapsid palaeobiology, data on their diel activity are currently unavailable. Using scleral ring and orbit dimensions, we demonstrate that nocturnal activity was not an innovation unique to mammals but a character that appeared much earlier in synapsid history, possibly several times independently. The 24 Carboniferous to Jurassic non-mammalian synapsid species in our sample featured eye morphologies consistent with all major diel activity patterns, with examples of nocturnality as old as the Late Carboniferous (ca 300 Ma). Carnivores such as Sphenacodon ferox and Dimetrodon milleri, but also the herbivorous cynodont Tritylodon longaevus were likely nocturnal, whereas most of the anomodont herbivores are reconstructed as diurnal. Recognizing the complexity of diel activity patterns in non-mammalian synapsids is an important step towards a more nuanced picture of the evolutionary history of behaviour in the synapsid clade.