Pakicetus inachus and the origin of whales and dolphins (Mammalia: Cetacea)

The present paper is concerned with the comparative morphology of the archeocete and odontocete skull. Among the archeocetes, the recently described lower Eocene Pakicetus inachus obviously represents an early stage of adaptation to aquatic life. The morphology of the incomplete cranial remains, however, gives no evidence that Pakicetus was an amphibious intermediate stage. The evolution of advanced archeocetes and odontocetes is characterized by the successive acquirement of new morphological devices related to the emission and perception of ultrasound under water. The formation of a sonar system in odontocetes obviously not only helped to compensate for the loss of the peripheral olfactory system but moreover was a substantial factor in the evolution of the exceptional dolphin brain.     

Computed tomography in the diagnosis of exudative otitis media

Temporal bone computed tomography (CT) was used to examine 37 patients aged 2 to 55 years who had exudative otitis media; in 27 patients of them, a pathological process was bilateral. An analysis of 58 temporal bone CT scans identified the CT signs of chronic exudative otitis media. These included a partial or complete block of the osseous foramen of the auditory tube; impaired pneumatization of the tympanic cavity, mastoid process fenestrae, and antrum; pathological drawing-in of the tympanic membrane. The preservation of the auditory ossicles and the absence of destructive changes in the walls of the cavities of the middle ear were observed in most cases. Repeated temporal bone CT study was performed in 10 patients (14 temporal bones) in different periods (from 2 months to 3 years) after surgery. The results of tympanostomy were visually assessed. These included recovered pneumatization of middle ear cavities (7 temporal bones), a cicatricial process in the tympanic cavity (5 temporal bones), recurrence of the CT manifestations of exudative otitis media (2 temporal bones).     

The mechanism of olfactory organ ventilation in Periophthalmus barbarus (Gobiidae, Oxudercinae)

Periophthalmus barbarus Linnaeus, 1766 has many adaptations for amphibious life as a consequence of tidal zone occupation. One of them is the ability to keep a little amount of water and air in mouth while on land or in hypoxic water, correlated with closing a gill lid for gas exchange improvement. It causes that mechanisms of olfactory organ ventilation described in other species of actinopterygians (compression of accessory nasal sac(s) by the skull and jaw elements while mouth and gill lid moving) are not in operation. There is a specific mechanism of olfactory organ ventilation independent on jaw and skull elements movements. Compression of accessory nasal sacs is possible by a0 contraction and it is a movement effect on bones combined by ligaments. This process can be observed on P. barbarus as lifting the rostral part of the head.     

Bapx1 regulates patterning in the middle ear: altered regulatory role in the transition from the proximal jaw during vertebrate evolution

The middle ear apparatus is composed of three endochondrial ossicles (the stapes, incus and malleus) and two membranous bones, the tympanic ring and the gonium, which act as structural components to anchor the ossicles to the skull. Except for the stapes, these skeletal elements are unique to mammals and are derived from the first and second branchial arches. We show that, in combination with goosecoid (Gsc), the Bapx1 gene defines the structural components of the murine middle ear. During embryogenesis, Bapx1 is expressed in a discrete domain within the mandibular component of the first branchial arch and later in the primordia of middle ear-associated bones, the gonium and tympanic ring. Consistent with the expression pattern of Bapx1, mouse embryos deficient for Bapx1 lack a gonium and display hypoplasia of the anterior end of the tympanic ring. At E10.5, expression of Bapx1 partially overlaps that of Gsc and although Gsc is required for development of the entire tympanic ring, the role of Bapx1 is restricted to the specification of the gonium and the anterior tympanic ring. Thus, simple overlapping expression of these two genes appears to account for the patterning of the elements that compose the structural components of the middle ear and suggests that they act in concert. In addition, Bapx1 is expressed both within and surrounding the incus and the malleus. Examination of the malleus shows that the width, but not the length, of this ossicle is decreased in the mutant mice. In non-mammalian jawed vertebrates, the bones homologous to the mammalian middle ear ossicles compose the proximal jaw bones that form the jaw articulation (primary jaw joint). In fish, Bapx1 is responsible for the formation of the joint between the quadrate and articular (homologues of the malleus and incus, respectively) enabling an evolutionary comparison of the role of a regulatory gene in the transition of the proximal jawbones to middle ear ossicles. Contrary to expectations, murine Bapx1 does not affect the articulation of the malleus and incus. We show that this change in role of Bapx1 following the transition to the mammalian ossicle configuration is not due to a change in expression pattern but results from an inability to regulate Gdf5 and Gdf6, two genes predicted to be essential in joint formation.     

Simulation of the power transmission of bone-conducted sound in a finite-element model of the human head

Bone conduction (BC) sound is the perception of sound transmitted in the skull bones and surrounding tissues. To better understand BC sound perception and the interaction with surrounding tissues, the power transmission of BC sound is investigated in a three-dimensional finite-element model of a whole human head. BC sound transmission was simulated in the FE model and the power dissipation as well as the power flow following a mechanical vibration at the mastoid process behind the ear was analyzed. The results of the simulations show that the skull bone (comprises the cortical bone and diploë) has the highest BC power flow and thereby provide most power transmission for BC sound. The soft tissues was the second most important media for BC sound power transmission, while the least BC power transmission is through the brain and the surrounding cerebrospinal fluid (CSF) inside the cranial vault. The vibrations transmitted in the skull are mainly concentrated at the skull base when the stimulation is at the mastoid. Other vibration transmission pathways of importance are located at the occipital bone at the posterior side of the head while the transmission of sound power through the face, forehead and vertex is minor. The power flow between the skull bone and skull interior indicate that some BC power is transmitted to and from the skull interior but the transmission of sound power through the brain seem to be minimal and only local to the brain–bone interface.     

The craniofacial air sac system of Mesozoic birds (Aves)

Birds are characterized by pneumatization of their skeletons by epithelial diverticula from larger, air—filled cavities. The diverticula—or 'air sacs'—that invade the postcranium result from outgrowths of the lungs; postcranial pneumaticity has been very well studied. Much more poorly understood are the air sacs that pneumatize the skull. Study of craniofacial pneumaticity in modern birds (Neornithes) indicates the presence of two separate systems: nasal pneumaticity and tympanic pneumaticity. The lacrimal and maxillary bones are pneumatized by diverticula of the main paranasal cavity, the antorbital sinus. There are five tympanic diverticula in neornithines that pneumatic the quadrate, articulare and the bones of the braincase. The pneumatic features of the following six genera of Mesozoic birds are examined: Archaeopteryx, Enaliornis, Baptornis, Parahesperornis, Hesperornis and Ichthyornis. Despite the 'archaic' aspect of most of these birds, many of the pneumatic features of neornithines are found in Mesozoic birds and are considered primitive for Aves. The phylogenetic levels at which most of the avian pneumatic features arose within Archosauria are uncertain. Until the phylogenetic levels at which homologous pneumatic features arose are determined, it is unwise to use most pneumatic characters in the discussion of avian origins. Within avian phylogeny, Ornithurae and Neornithes are well–supported by pneumatic synapomorphies. There is a trend towards reduction of craniofacial pneumaticity within Hesperornithiformes. Within Neornithes, four derived pneumatic characters suggest that the Palaeognathae (ratites and tinamous) is monophyletic.     

Middle ear structure and bone conduction in Spalax,Eospalax, and Tachyoryctes mole‐rats (Rodentia: Spalacidae)

There is evidence that spalacine, tachyoryctine, and myospalacine mole‐rats all communicate with conspecifics through a form of seismic signaling, but the route for the detection of these signals is disputed. It has been proposed that two unusual anatomical adaptations in Spalax allow jaw vibrations to pass to the inner ear via the incus and stapes: a pseudoglenoid (=postglenoid) fossa which accomodates the condylar process of the mandible, and a bony cup, supported by a periotic lamina, through which the incus articulates with the skull. In this study, a combination of dissection and computed tomography was used to examine the ear region in more detail in both Spalax and its subterranean relatives Tachyoryctes and Eospalax, about which much less is known. Tachyoryctes was found to lack a pseudoglenoid fossa, while Eospalax lacks a periotic lamina and bony cup. This shows that these structures need not simultaneously be present for the detection of ground vibrations in mole‐rats. Based on the observed anatomy, three hypothetical modes of bone conduction are argued to represent more likely mechanisms through which mole‐rats can detect ground vibrations: ossicular inertial bone conduction, a pathway involving sound radiation into the external auditory meatus, and a newly‐described fluid pathway between pseudoglenoid fossa and cranial cavity. The caudolateral extension of the tympanic cavity and the presence of a bony cup might represent synapomorphies uniting Spalax and Tachyoryctes, while the loss of the tensor tympani muscle in Spalax and Eospalax may be convergently derived. J. Morphol., 2010. © 2009 Wiley‐Liss, Inc.     

Auditory anatomy of beaked whales and other odontocetes: Potential for cochlear stimulation via a “vibroacoustic duct mechanism”

Computed tomography (CT) and microcomputed tomography (microCT) were used to examine the structures involved in cochlear stimulation in odontocetes and terrestrial mammals. Cranial CT examined the osseous attachment of the skull to the tympanoperiotic complex (TPC) and the path of the endocranial foramen of the vestibulocochlear nerve (EFVN), which was assumed to contain the perilymphatic duct. Additional CTs of TPC were taken postextraction to examine the gross morphology of this structure. MicroCT was used to examine the acoustic windows of the cochlea, including the round and oval windows and the apertures of the cochlear and vestibular aqueducts. Cranial CT scans demonstrated an osseous connection between the skull and TPC in beaked whales and Physeter macrocephalus. EFVN traveled through a greater length of cranial bone and communicated more closely with the periotic bone in beaked whales than in other species. Ziphius cavirostris was observed to have a reduced medial sulcus of the mallear ridge (MSMR) and tympanic plate and an enlarged aperture of the cochlear aqueduct, respectively. The potential significance of these findings, including the role of the perilymphatic duct as a novel route of cochlear stimulation referred to as the “vibroacoustic duct mechanism,” are discussed.     

The ethmoid and presphenoid of cetaceans

A cribriform plate, a perpendicular plate, and two lateral masses are major components of the ethmoid bone of mammals. Notwithstanding the noticeable bone, virtually sitting in the center of the skull, extensive modifications of the skull of modern cetaceans, especially odontocetes (toothed whales), and the lack of clarity as to what characteristics delimit each element of the ethmoid has made the problem of the nature of the cetacean ethmoid more complicated and elusive than in other, less modified mammals. Furthermore, contention as to whether a perpendicular plate of the ethmoid, or the mesethmoid, exists in all mammals including cetaceans has remained unsettled. In odontocetes, the mesethmoid has been variably identified not only as the osseous nasal septum but also as the mediodorsal region of the posterior wall of the nasal passage below the nasals, as a mass of bone encased by the vomer in front of the osseous nasal cavity at the base of the rostrum, and as a combination of some portions mentioned above. The presence or absence of the mesethmoid in various groups of mammals has attracted the attention of some biologists, and here, I demonstrate that cetaceans have no mesethmoid. The close inspection of the ontogenetic changes of the basicranial elements in cetaceans reveals that a mass of bone ensheathed by the vomer in front, or at the level of the osseous nasal cavity is actually the presphenoid. It is highly likely that in odontocetes the posterior wall of the nasal passages below the nasals consists of the combination of the frontal, the imperforated cribriform plate, the paired ectethmoids, and the vomer, the latter three of which partially concealing the presphenoid dorsally and laterally as the ontogeny proceeds. In contrast, mysticetes clearly display ethmoturbinates and a cribriform plate, which are morphologically similar to those in terrestrial mammals. J. Morphol. 277:1661–1674, 2016. © 2016 Wiley Periodicals, Inc.     

Computed tomography of the temporal bones in children with developmental anomalies of the external and middle ear]

Computed polypositional high-resolution tomography (CT) was provided to 45 children with dysplasia of the external acoustic meatus (EAM). The computed tomograms were analyzed in accordance with the scheme which included the evaluation of the following structures: atresia or stenosis area, pneumatized mastoid process, tympanic cavity, ant rum, osteal opening of the Eustachian tube, chain of auditory ossicles, windows of the labyrinth, internal ear, facial nerve canal, and large vessels. Preoperative CT of the temporal bones allows for the evaluation of the above-listed structures, which is of paramount importance for the planning of the patient management policy. Also, CT enables the assessment of the risk of surgical intervention associated with a potential injury to the facial nerve, large vessels, and temporomandibular articulation. CT is to be an indispensable diagnostic modality for examination of children presenting with EAM dysplasia.     

Congenital anomalies of the ear resulting from cyclophosphamide treatment in the rat

A single dose of cyclophosphamide (20 mg/kg) administered on day 12 of gestation to CF rats resulted in microtia in 97.5% of fetuses at term. The ears of affected fetuses were low set and dorsally placed. Histological examination revealed persistence of meatal plug, branching of the primordium of external acoustic meatus, periotic hemorrhages, narrowing of tympanic cavity, presence of only one to two primordia of middle ear ossicles, hypoplasia of stapedial artery, a maximum of two turns of cochlea and under differentiation of the organ of Corti and the semicircular ducts. Hemorrhages in and around the region of the ear were extensive. Bilaterally symmetrical distribution of microtia and consistency of associated anomalies suggest that this is a good animal model for further investigation into the pathogenetic mechanisms of these ear anomalies.     

Specialization for underwater hearing by the tympanic middle ear of the turtle, Trachemys scripta elegans

Turtles, like other amphibious animals, face a trade-off between terrestrial and aquatic hearing. We used laser vibrometry and auditory brainstem responses to measure their sensitivity to vibration stimuli and to airborne versus underwater sound. Turtles are most sensitive to sound underwater, and their sensitivity depends on the large middle ear, which has a compliant tympanic disc attached to the columella. Behind the disc, the middle ear is a large air-filled cavity with a volume of approximately 0.5 ml and a resonance frequency of approximately 500 Hz underwater. Laser vibrometry measurements underwater showed peak vibrations at 500-600 Hz with a maximum of 300 μm s(-1) Pa(-1), approximately 100 times more than the surrounding water. In air, the auditory brainstem response audiogram showed a best sensitivity to sound of 300-500 Hz. Audiograms before and after removing the skin covering reveal that the cartilaginous tympanic disc shows unchanged sensitivity, indicating that the tympanic disc, and not the overlying skin, is the key sound receiver. If air and water thresholds are compared in terms of sound intensity, thresholds in water are approximately 20-30 dB lower than in air. Therefore, this tympanic ear is specialized for underwater hearing, most probably because sound-induced pulsations of the air in the middle ear cavity drive the tympanic disc.     

Origin of the human stapedius muscle

More than 100 human embryos and fetuses, beginning from 6 mm (25-26 days) up to newborns have been investigated by means of histological serial sections, as well as by means of macro- micropreparation of the tympanic cavity. The m. stapedius is not independent, but takes origin from muscle fasciculi of the musculi digastrii posterior belly. The latter approaches the eminence of the future mastoid process, gives a part of muscle fasciculi into the tympanic cavity; they get through the stilomastoid cleft and run to the hammer neck, forming the m. stapedius. That is why both muscles are innervated from the facial nerve.     

Trait independence primes convergent trait loss

The repeated, independent evolution of traits (convergent evolution) is often attributed to shared environmental selection pressures. However, developmental dependencies among traits can limit the phenotypic variation available to selection and bias evolutionary outcomes. Here, we determine how changes in developmentally correlated traits may impact convergent loss of the tympanic middle ear, a highly labile trait within toads that currently lack adaptive explanation. The middle ear's lability could reflect evolutionary trade‐offs with other skull features under selection, or the middle ear may evolve independently of the rest of the skull, allowing it to be modified by active or passive processes without pleiotropic trade‐offs with other skull features. We compare the skulls of 55 species (39 eared, 16 earless) within the family Bufonidae, spanning six hypothesized independent middle ear transitions. We test whether shared or lineage‐specific changes in skull shape distinguish earless species from eared species and whether earless skulls lack other late‐forming skull bones. We find no evidence for pleiotropic trade‐offs between the middle ear and other skull structures. Instead, middle ear loss in anurans may provide a rare example of developmental independence contributing to evolutionary lability of a sensory system.     

Anatomy and physics of the exceptional sensitivity of dolphin hearing (Odontoceti: Cetacea)

During the past 50 years, the high acoustic sensitivity and the echolocation behavior of dolphins and other small odontocetes have been studied thoroughly. However, understanding has been scarce as to how the dolphin cochlea is stimulated by high frequency echoes, and likewise regarding the ear mechanics affecting dolphin audiograms. The characteristic impedance of mammalian soft tissues is similar to that of water, and thus no radical refractions of sound, nor reflections of sound, can be expected at the water/soft tissue interfaces. Consequently, a sound-collecting terrestrial pinna and an outer ear canal serve little purpose in underwater hearing. Additionally, compared to terrestrial mammals whose middle ear performs an impedance match from air to the cochlea, the impedance match performed by the odontocete middle ear needs to be reversed to perform an opposite match from water to the cochlea. In this paper, we discuss anatomical adaptations of dolphins: a lower jaw collecting sound, thus replacing the terrestrial outer ear pinna, and a thin and large tympanic bone plate replacing the tympanic membrane of terrestrial mammals. The paper describes the lower jaw anatomy and hypothetical middle ear mechanisms explaining both the high sensitivity and the converted acoustic impedance match.     

A new baleen whale from the Late Miocene of Denmark and early mysticete hearing

Abstract:  The extinct mysticete fauna of the North East Atlantic is primarily known from the abundant but fragmented Belgian specimens. Compared to the well-preserved contemporary mysticete fauna from deposits in North America, there are only few near complete European Miocene mysticete fossils. Presented here is a new, almost complete fossil baleen whale Uranocetus gramensis gen. et sp. nov. from the Upper Miocene Gram Formation in South West Denmark. It is the first stem-balaenopterid that has an initial stage of reduction in the mandibular cavity and a rostral configuration that is intermediate between that of other stem-balaenopterids and true balaenopterids. It is likely that Uranocetus used a gulp feeding technique that approaches that of balaenopterids. Details of the periotic and mandibular morphology place Uranocetus in the family Diorocetidae Steeman 2007. The large mandibular cavity in Uranocetus and most other extinct mysticetes, when compared to the reduced condition in recent mysticetes, is not an indication that early mysticetes used odontocete-like echolocation. In Uranocetus and a distantly related mysticete, high frequency sounds in the range odontocetes use for echolocation would suffer a significant volume loss across the lateral mandibular wall on the passage towards the inner ear. A reduction in the mandibular cavity in separate evolutionary lineages of mysticetes may be the result of a shift towards the use of low frequency sounds.     

Potentialities of temporal bone computed tomography in the identification of causes of hypoacusis of postinflammatory genesis

Temporal bone computed tomography (CT) was used to examine 64 patients with impaired hearing due to inflammatory diseases of the middle year. In 21 patients, the pathological process was bilateral. A total of 85 series of temporal bone CT scans were analyzed. The patients' age ranged from 2 to 66 years. CT verified adhesive otitis media in 62 cases, otosclerosis in 7, local malformation of the auditory ossicles and/or the labyrinthine fenestrae in 11. No CT changes were revealed in 5 cases. The CT symptoms of adhesive otitis media were identified. These included soft tissue bands and/or soft tissue-density portions that fix the auditory ossicles or block the niches of the labyrinthine fenestrae (31 temporal bones); sclerosis or ossification of the ligaments and tendons of the middle ear (7 temporal bones); calcification foci in the tympanic cavity (9 cases); osteosclerotic changes in the epitympanus (2 cases); cicatricial changes in the tympanic membrane (24 cases); destructive changes in the auditory ossicles (19 temporal bones). There has been evidence that CT may be used for the differential diagnosis of adhesive otitis media from otosclerosis and congenital malformations of the structures of the middle ear.     

Potentialities of temporal bone CT in the diagnosis of chronic purulent otitis media and its complications

Temporal bone CT was used to examine a group of 87 patients with chronic purulent otitis media (103 temporal bones). The patients' age ranged from 2 to 74 years. A scheme was developed and proposed to evaluate the temporal bone by CT. The CT signs of chronic purulent otitis media uncomplicated by cholesteatoma and those of cholesteatomic purulent otitis were identified. The CT symptomatology of chronic purulent otitis includes: sclerotic changes in the bone tissue of the mastoid process, impaired pneumatization of the cavities of the middle ear, including the tympanic cavity, destructive changes in auditory ossicles, carious changes in the walls of the cavities of the middle ear. The CT semiotics of cholesteatoma depends on its site and spread into the temporal bone and includes as follows: deformation of the epitympanum due to soft tissue mass-induced destruction of the lateral wall; the dilated entrance into the antrum; the presence of a cavity with the sclerosed walls in the antromastoid area; carious changes in the auditory ossicles; the displacement of a chain of ossicles medially or laterally in relation to the initial site of cholesteatoma. CT reflects carious changes in the walls of the cavities of the middle ear, including the roof and labyrinthine wall of the tympanum, which allows labyrinthine fistula and intracranial cholesteatomic complications. The study of the temporal bone by the proposed scheme may reveal anomalies and the specific features of its structure: the presentation of the sigmoid sinus, the high elevation of the bulb of the jugular vein, diverticulum of the latter, the low standing of the bottom of the ACH.     

The Evolution of the Tetrapod Middle Ear in the Rhipidistian-Amphibian Transition

From a survey ot the structure of the skull in rhipidistianfishes and early labylinthodont Amphibia and of the mechanismof hearing in these two groups, an account of the evolutionof the tetrapod middle ear is presented. The overall modificationof the otic region of the skull during the rhipidistian-amphibiantransition is analyzed in terms of changes in different organsystems in response to different selective pressures (affecting,for example, the feeding, respiratory, and locomotory mechanisms).These changes are seen to occur in a completely integrated pattern.Considerations of the different requirements for sound receptionunder water and in air, in connection with this correlated progressionof evolutionary change in the otic region of the head, revealthe manner in which the hyomandibular, spiracular diverticulum,and operculum of rhipidistian fishes became modified to formthe stapes, the tympanic cavity, and the outer portion of thetympanum, respectively, of tetrapods.