Assessing morphology and function of the semicircular duct system: introducing new in-situ visualization and software toolbox.

The semicircular duct system is part of the sensory organ of balance and essential for navigation and spatial awareness in vertebrates. Its function in detecting head rotations has been modelled with increasing sophistication, but the biomechanics of actual semicircular duct systems has rarely been analyzed, foremost because the fragile membranous structures in the inner ear are hard to visualize undistorted and in full. Here we present a new, easy-to-apply and non-invasive method for three-dimensional in-situ visualization and quantification of the semicircular duct system, using X-ray micro tomography and tissue staining with phosphotungstic acid. Moreover, we introduce Ariadne, a software toolbox which provides comprehensive and improved morphological and functional analysis of any visualized duct system. We demonstrate the potential of these methods by presenting results for the duct system of humans, the squirrel monkey and the rhesus macaque, making comparisons with past results from neurophysiological, oculometric and biomechanical studies. Ariadne is freely available at http://www.earbank.org.

Implications of new early Homo fossils from Ileret, east of Lake Turkana, Kenya.

Sites in eastern Africa have shed light on the emergence and early evolution of the genus Homo. The best known early hominin species, H. habilis and H. erectus, have often been interpreted as time-successive segments of a single anagenetic evolutionary lineage. The case for this was strengthened by the discovery of small early Pleistocene hominin crania from Dmanisi in Georgia that apparently provide evidence of morphological continuity between the two taxa. Here we describe two new cranial fossils from the Koobi Fora Formation, east of Lake Turkana in Kenya, that have bearing on the relationship between species of early Homo. A partial maxilla assigned to H. habilis reliably demonstrates that this species survived until later than previously recognized, making an anagenetic relationship with H. erectus unlikely. The discovery of a particularly small calvaria of H. erectus indicates that this taxon overlapped in size with H. habilis, and may have shown marked sexual dimorphism. The new fossils confirm the distinctiveness of H. habilis and H. erectus, independently of overall cranial size, and suggest that these two early taxa were living broadly sympatrically in the same lake basin for almost half a million years.

Vestibular evidence for the evolution of aquatic behaviour in early cetaceans.

Early cetaceans evolved from terrestrial quadrupeds to obligate swimmers, a change that is traditionally studied by functional analysis of the postcranial skeleton. Here we assess the evolution of cetacean locomotor behaviour from an independent perspective by looking at the semicircular canal system, one of the main sense organs involved in neural control of locomotion. Extant cetaceans are found to be unique in that their canal arc size, corrected for body mass, is approximately three times smaller than in other mammals. This reduces the sensitivity of the canal system, most plausibly to match the fast body rotations that characterize cetacean behaviour. Eocene fossils show that the new sensory regime, incompatible with terrestrial competence, developed quickly and early in cetacean evolution, as soon as the taxa are associated with marine environments. Dedicated agile swimming of cetaceans thus appeared to have originated as a rapid and fundamental shift in locomotion rather than as the gradual transition suggested by postcranial evidence. We hypothesize that the unparalleled modification of the semicircular canal system represented a key 'point of no return' event in early cetacean evolution, leading to full independence from life on land.

New hominin genus from eastern Africa shows diverse middle Pliocene lineages.

Most interpretations of early hominin phylogeny recognize a single early to middle Pliocene ancestral lineage, best represented by Australopithecus afarensis, which gave rise to a radiation of taxa in the late Pliocene. Here we report on new fossils discovered west of Lake Turkana, Kenya, which differ markedly from those of contemporary A. afarensis, indicating that hominin taxonomic diversity extended back, well into the middle Pliocene. A 3.5 Myr-old cranium, showing a unique combination of derived facial and primitive neurocranial features, is assigned to a new genus of hominin. These findings point to an early diet-driven adaptive radiation, provide new insight on the association of hominin craniodental features, and have implications for our understanding of Plio-Pleistocene hominin phylogeny.

Using diagnostic radiology in human evolutionary studies.

This paper reviews the application of medical imaging and associated computer graphics techniques to the study of human evolutionary history, with an emphasis on basic concepts and on the advantages and limitations of each method. Following a short discussion of plain film radiography and pluridirectional tomography, the principles of computed tomography (CT) and magnetic resonance imaging (MRI) and their role in the investigation of extant and fossil morphology are considered in more detail. The second half of the paper deals with techniques of 3-dimensional visualisation based on CT and MRI and with quantitative analysis of digital images.

Rare temporal bone pathology of the Singa calvaria from Sudan.

Evidence has recently accumulated that the Singa calvaria from Sudan probably dates from Oxygen Isotope Stage 6 (>130 ka). Morphological studies have indicated a mixture of archaic and more modern human traits, but such analyses are complicated by the possibility that the vault is pathologically deformed, although the exact etiology has not been established. Now computed tomography (CT) has revealed that the right temporal bone lacks the structures of the bony labyrinth. The most likely cause of this rare pathological condition appears to be labyrinthine ossification, in which newly deposited bone obliterates the inner ear spaces following an infectious disease or occlusion of the labyrinthine blood supply. A possible cause of vascular compromise could have been the presence of an expanding acoustic neuroma in the internal acoustic meatus, which is suggested by a significantly wider right meatus compared with the left side. Interestingly, labyrinthine ossification is also consistent with the controversial diagnosis that an anemia caused the characteristic diploic widening at the parietal bosses, because prime etiological factors of ossification are among the common complications of some of these blood diseases. CT examination of the vault and a review of the literature suggest that a blood disorder may well have caused the unusual parietal morphology. Given the nature of these pathological conditions, the Singa individual must have experienced a period of considerable disability. The morphological evidence from the normal bony labyrinth on the left side and from the CT evaluation of the vault is consistent with the interpretation of Singa as a late archaic hominid or an early representative of Homo sapiens drawn from a population which might be directly ancestral to modern humans.

Comparative review of the human bony labyrinth.

The bony labyrinth inside the petrous part of the temporal bone houses the organs of hearing and balance. Being functionally linked with sensory control of body movements and located in a part of the basicranium that is closely associated with the brain, this structure is of great interest in the study of human evolutionary history. However, few aspects of its morphology have been studied in nonhuman primates. This review compares the bony labyrinth of humans with that of the great apes and 37 other primate species based on data newly acquired with computed tomography combined with previous descriptions. With body mass taken into account, consistent differences are found between the size of the semicircular canals in humans, the great apes, and other primates. In particular, the arcs of the anterior and posterior canals are larger in humans than in the African apes. The functional implications of semicircular canal dimensions for registering angular head motion are evaluated in relation to locomotor behavior. Biophysical models, comparative studies, and some neurophysiological experiments all support a link between semicircular canal size and agility, or more specifically the frequency contents of natural head movements, but the evidence on the exact nature of this link is ambiguous. It is concluded that any link between the characteristic dimensions of the human canals and locomotion will be more complex than a simple association with the broad categories of quadrupedal vs. bipedal behavior. The functionally important planar orientations of the semicircular canals are similar in humans and the African apes as well as in many other species. In contrast, other aspects of the human labyrinth differ markedly in shape, following a pattern that seems to reflect the characteristic architecture of the human basicranium. Indeed, it is found that labyrinthine and basicranial shape are interspecifically correlated in the sample, and in most respects the human morphology is consistent with the general trend among primate species. Differences in brain growth and development are proposed as the predominant factor underlying both the unique shape of the human labyrinth as well as the interspecific labyrintho-basicranial correlations.

A late Neanderthal associated with Upper Palaeolithic artefacts.

The French site of Arcy-sur-Cure is a key locality in documenting the Middle-Upper Palaeolithic transition in Europe. Reliable attribution of the fragmentary hominid fossils associated with its early Upper Palaeolithic Châtelperronian industry has not been possible. Here we report the first conclusive identification of one of these fossils as Neanderthal on the basis of newly discovered derived features of the bony labyrinth. Dated at about thirty-four thousand years (34 kyr) ago, the fossil is representative of the youngest known Neanderthal populations, and its archaeological context indicates that these hominids used a rich bone industry as well as personal ornaments. The evidence supports the hypothesis of a long term coexistence with technocultural interactions between the first modern humans and the last Neanderthals in Europe. However, the complete absence of the derived Neanderthal traits in labyrinths of modern Upper Palaeolithic specimens from western Europe argues against phylogenetic continuity between the two populations in this region.

Morphometry of the primate bony labyrinth: a new method based on high-resolution computed tomography.

A method is described for taking accurate measurements of the bony labyrinth of humans and other primates using high-resolution computed tomography (CT). The measurements comprise 8 dimensions, 14 orientations and 2 indices of the labyrinth, as well as 7 orientations of related structures of the petrous pyramid. Comparison of the measurements taken from CT scans with those taken from subsequently made casts and cryosections demonstrates that the method is sufficiently accurate to permit the morphometric analysis of labyrinthine size and shape. Since the CT method is nondestructive, fast and easy to perform, it is applicable to large samples and to rare or precious anthropological specimens.

Implications of early hominid labyrinthine morphology for evolution of human bipedal locomotion.

The upright posture and obligatory bipedalism of modern humans are unique among living primates. The evolutionary history of this behaviour has traditionally been pursued by functional analysis of the postcranial skeleton and the preserved footprint trails of fossil hominids. Here we report a systematic attempt to reconstruct the locomotor behaviour of early hominids by looking at a major component of the mechanism for the unconscious perception of movement, namely by examining the vestibular system of living primates and early hominids. High-resolution computed tomography was used to generate cross-sectional images of the bony labyrinth. Among the fossil hominids the earliest species to demonstrate the modern human morphology is Homo erectus. In contrast, the semicircular canal dimensions in crania from southern Africa attributed to Australopithecus and Paranthropus resemble those of the extant great apes. Among early Homo specimens, the canal dimensions of Stw 53 are unlike those seen in any of the hominids or great apes, whereas those of SK 847 are modern-human-like.