The relative efficacy of functional and developmental cranial modules for reconstructing global human population history

This study tests the relative efficacy of human cranial modules, defined on the basis of developmental and functional criteria, for reconstructing neutral genetic population history. Specifically, two hypotheses were tested: 1) The "basicranial hypothesis" predicts that the endochondrally ossifying basicranium will be more reliable for reconstructing population history than intramembranously ossifying regions of the human cranium. This is based on the assumption that early ossification of the basicranium and its distinct functional constraints produce a cranial structure that is relatively immune to non-neutral evolutionary forces. 2) The "single function hypothesis" predicts that cranial regions associated with a single (sensory) function are less reliable indicators of neutral genetic history. Here the prediction is based on the logic that complex, multi-functional, integrated cranial regions are less likely toexhibit homoplasy and, therefore, provide a more accurate morphological proxy for genetic relationships. The congruence between craniometric affinity matrices and neutral genetic population matrices based on autosomal microsatellite and classical markers was assessed using a series of Mantel and Dow-Cheverud tests. The results did not support the predictions of the "basicranial hypothesis," as the endochondrally ossifying basicranium was not significantly more congruent with the genetic data than intramembraneously ossifying modules. Moreover, although the results provided some support for the "single function hypothesis," defining cranial modules on the basis of anatomical or functional complexity did not provide a consistent means of predicting their phylogenetic efficacy. These results have important implications for building an accurate inference model of cranial evolution in the human fossil record.     

Early development of the cephalic skeleton in the turbot

At hatching Scophthalmus maximus shows no cartilaginous and no bony structure. Mecke?s cartilages appear when the fry are 1 day old, followed on day 2, by formation of the trabecular bars, fused at the outset to form a trabecula communis. Concurrently, the palatoquadrates complete the mandibular arch, and the first two pairs of ceratobranchials, associated with a pair of hyoid bars, form the beginnings of the hyobranchial system. By day 3, the parachordals have fused with the trabecular bars, the hyosymplectics have linked to the hyoid bars by interhyals, and the first four pairs of ceratobranchials have appeared. The first bony structures appear: the preoperculars. On day 8, the frontals develop above the orbits and the maxillaries and dentaries appear. On day 10, the primordia of the taeniae marginales appear, the palatoquadrates bear a pterygoid process, and to the branchial basket have been added the fifth pair of ceratobranchials and the four pairs of epibranchials. On day 12, both pairs of posterior pharyngobranchials are present. The premaxillaries develop in front of the maxillaires, and retroarticulars and the angulars complete the lower jaws. On day 13, a thin parasphenoid contributes to the floor of the neurocranium, and ectopterygoids and entopterygoids to the splanchnocranium. The set of opercular bones is complete. On day 15, the tectum synoticum closes the braincase posteriorly. The splanchnocranium possesses a basihyal and the pharyngobranchials of the first epibranchials. On day 18, the tectum posterius completes the dome of the braincase. The rear end and lateral walls of the skull are formed by the basioccipital, the exoccipitals, the pterotics, and the parietals. The suspensorium is nearly complete. From day 10, the first resorptions begin in parallel with the construction of the chondrocranium. Mecke?s cartilages each split in two, then the posterior part of the trabecular bars disappears. On day 23, the right taenia marginalis separates from the lamina orbitonasalis and curves towards the centre. Simultaneously, the right eye begins its migration to the left. This is the only metamorphosis-linked asymmetry to appear during the development of the chondrocranium. On day 25, many more bony structures appear, a characteristic of this stage: the nasals, lateral ethmoids, mesethmoid, sphenotics, prootics, pleurosphenoids, epiotics, and supraoccipital. From this stage on, the bony structures continue to develop, while the front of the neurocranium and the jaws undergo a deep remodelling due to metamorphosis. The left taenia marginalis does not appear reduced until day 29. By day 45, there remain only a few small elements of the cartilaginous skull.     

Retinal neural progenitors express topographic map markers

Transplantation of neural stem cells for replacing neurons after neurodegeneration requires that the transplanted stem cells accurately reestablish the lost neural circuits in order to restore function. Retinal ganglion cell axons project to visual centers of the brain forming circuits in precise topographic order. In chick, dorsal retinal neurons project to ventral optic tectum, ventral neurons to dorsal tectum, anterior neurons to posterior tectum and posterior neurons to anterior tectum; forming a continuous point-to-point map of retinal cell position in the tectal projection. We found that when stem cells derived from ventral retina were implanted in dorsal host retina, the stem cells that became ganglion cells projected to dorsal tectum, appropriate for their site of origin in retina but not appropriate for their site of implant in retina. This led us to ask if retinal progenitors exhibit topographic markers of cell position in retina. Indeed, retinal neural progenitors express topographic markers: dorsal stem cells expressed more Ephrin B2 than ventral stem cells and, conversely, ventral stem cells expressed more Pax-2 and Ventroptin than dorsal stem cells. The fact that neural progenitors express topographic markers has pertinent implications in using neural stem cells in cell replacement therapy for replacing projecting neurons that express topographic order, e.g., analogous neurons of the visual, auditory, somatosensory and motor systems.     

The craniofacial skeleton in anencephalic human fetuses. I. Cranial floor

Twelve anencephalic and four normal fetuses 26 to 40 weeks gestational age were compared by anatomic, radiographic and histologic methods in order to gain information concerning morphogenesis. In the anencephalics, alterations located within the body of the sphenoid bone led to a reduced cranial floor angle and a more vertical clivus. The reduced lateral extension of the lesser and greater wings of the sphenoid constricted the anterior and middle cranial fossae respectively. The posterior cranial fossa tended to have an increased transverse dimension related to the supraoccipital and exoccipital bone orientation. The increased anterior and inferior position of the lateral end of the petrous temporal ridge was positively correlated with the degree of dorsal schisis in the anencephalics. Alterations in the size, form, or duration of the neural functional matrix are suggested as the cause of changes in the cranial floor.     

Skeletal development of the direct-developing caecilian <Emphasis Type="Italic">Gegeneophis ramaswamii</Emphasis> (Amphibia: Gymnophiona: Caeciliidae)

A few previous studies of skeletal and especially skull development in Gymnophiona often provided contradictory results. We studied the development of the skull and vertebral column of Gegeneophis ramaswamii, a direct-developing Indian caeciliid, based on 13 specimens. The chondrocranium forms at (Brauer in Zool Jahrb Anat 12:477-508,1899) stage 38. First dermal and perichondral ossifications occur at stage 40. The first dermal bones to form are the mentomeckelian, dentary, angular, vomer, and premaxillary. These are followed by the coronoid, palatine, pterygoid, maxillary, and the skull-roofing bones. The last occurring dermal ossifications are the parasphenoid and the squamosal. We present evidence for the occurrence of a lacrimal bone. No ectopterygoid, basioccipital, supraoccipital, pleurosphenoid, postorbital, or supratemporal elements were found. We assess the homology of the bones constituting the caecilian skull and discuss the above-mentioned terminologies. The phylogenetic implications of our findings are briefly discussed and we conclude that the evidence from developmental morphology is at present consistent with a monophyletic Lissamphibia of temnospondyl origin.     

Osteological development of the lophiid anglerfish,lophius gastrophysus

The osteological development of the head skeleton and dorsal, pectoral, and anal fin supports, are described from cleared and stained specimens ofLophius gastrophysus larvae, ranging from 4.6 to 21.8 mm NL; the results are compared with those of juvenile (79.8 mm SL) and adult (398 mm SL) specimens. Tiny conical teeth are present on the premaxillary, dentary, palatine and vomer since early stage. The first three dorsal fin spines are initially positioned on the midline of body posterior to the supraoccipital, but they migrate forward with growth and become cephalic in juveniles. The forward movement of the dorsal spines is produced by the forward extension of the cartilaginous basal inside the subepidermal space. During the planktonic larval stage the pectoral fins are on the sides of body as in ordinary fishes, but they move ventrad and become leg-like in bottom living juveniles and adults. Ossification of the caudal complex ofL. gastrophysus larvae proceeds very slowly and only the 21.8 mm NL larva has an almost completely ossified caudal complex. Eight principal caudal rays are loosely attached on the posterior edge of the hypurals and no procurrent rays are present. Larvae have well developed parhypurapophysis at the mid-portion of the urostyle which transforms into keel-like structure in juveniles and adults.     

Projections to the midbrain tectum in Salamandra salamandra L.

Following unilateral iontophoretic application of HRP into the optic tectum of Salamandra salamandra, retrogradely HRP-filled cells were found bilaterally in the pretectum, tegmentum isthmi, the reticular formation, pars medialis, and in the nucleus vestibularis magnocellularis. The area octavo-lateralis projects only to the caudal part of the tectum. Ipsilateral projections were noted from the dorsal gray columns of the cervical spinal cord, the dorsal tegmentum, the thalamus dorsalis pars medialis, thalamus dorsalis, pars anterior (to the rostral one-third of the tectum), the thalamus ventralis (in its entire rostro-caudal extent), and the preoptico-hypothalamic complex. Retrogradely filled cells were identified in deeper layers of the contralateral tectum. There are two telencephalic nuclei projecting ipsilaterally to the tectum via the lateral forebrain: the ventral part of the lateral pallium, and the posterior strioamygdalar complex.     

Convergence of multisensory inputs in Xenopus tadpole tectum

The integration of multisensory information takes place in the optic tectum where visual and auditory/mechanosensory inputs converge and regulate motor outputs. The circuits that integrate multisensory information are poorly understood. In an effort to identify the basic components of a multisensory integrative circuit, we determined the projections of the mechanosensory input from the periphery to the optic tectum and compared their distribution to the retinotectal inputs in Xenopus laevis tadpoles using dye‐labeling methods. The peripheral ganglia of the lateral line system project to the ipsilateral hindbrain and the axons representing mechanosensory inputs along the anterior/posterior body axis are mapped along the ventrodorsal axis in the axon tract in the dorsal column of the hindbrain. Hindbrain neurons project axons to the contralateral optic tectum. The neurons from anterior and posterior hindbrain regions project axons to the dorsal and ventral tectum, respectively. While the retinotectal axons project to a superficial lamina in the tectal neuropil, the hindbrain axons project to a deep neuropil layer. Calcium imaging showed that multimodal inputs converge on tectal neurons. The layer‐specific projections of the hindbrain and retinal axons suggest a functional segregation of sensory inputs to proximal and distal tectal cell dendrites, respectively. © 2009 Wiley Periodicals, Inc. Develop Neurobiol, 2009     

Functional anatomy of the luring apparatus of the deep-sea ceratioid anglerfish <Emphasis Type="Italic">Cryptopsaras couesii</Emphasis> (Lophiiformes: Ceratiidae)

The osteology and myology of the illicial apparatus of Cryptopsaras couesii were examined in an effort to elucidate function. The apparatus consists of two bones, a tiny illicial bone, completely enveloped by tissue of the esca, and an extremely long supporting pterygiophore that lies within a deep groove on the dorsal surface of the head. The anterior end of the pterygiophore emerges on the tip of the snout from between the frontal bones, while the posterior end, encased in a dermal sheath when the illicial apparatus is retracted, emerges on the back just anterior to the dorsal caruncles. Five pairs of muscles control movement of the illicial apparatus: small, short erectors and depressors control movement of the illicial bone, while extremely long inclinators, protractors, and retractors control movement of the pterygiophore. The protractors and retractors of C. couesii are more robust and much longer than those of other lophiiforms, indicating that the pterygiophore of this species has an exceptionally wide range of movement in the anterior and posterior plane. Moreover, these muscles wind around the pterygiophore in opposite directions, a unique anatomy that suggests that C. couesii extends and retracts the pterygiophore by rotation.     

湘西北志留纪胴甲鱼化石

本文记述了采自湘西北澧县石门水库附近秀山组上段隶属云南鱼目的 Shimenolepis graniferus gen. et sp. nov. 和 QuJinolepidae gen. et sp. indet. 化石,对含鱼层的时代也进行了讨论.     

Electrophysiological analysis of cortico-tectal connections in the turtle

Two types of evoked potentials are recorded in the tectum mesencephali in response to electrical stimulation of the forebrain surface of the turtleEmys orbicularis. The results of a layer-by-layer analysis show that evoked potentials of type I in response to stimulation of the hippocampal and piriform cortex are generated outside the tectum. Evoked potentials of type II, consisting of two surface-negative components, are recorded in the tectum in response to stimulation of the rostro-central surface of the forebrain. The first component appeared after a latent period of 20 msec and lasted 40–60 msec; the second component appeared after 80–100 msec and lasted 100–300 msec. Layer-by-layer and pharmacological analysis showed that the first component of the type II evoked potential is generated in the tegmental structures of the mesencephalon, whereas the second (long-latency) is generated in the tectum. The tectal origin of the second component is confirmed by its interaction with the tectal response to photic stimulation or to electrical stimulation of the optic nerve, evidence that these evoked potentials are generated by common structures. The efferent pathway from the dorsal cortex to the primary visual center is unilateral and has features of polysynaptic projections (long latent period, low lability).     

中华蟾蜍中脑组织学研究

采用HE染色和Holmes银染法对蟾蜍中脑的显微结构进行了研究.中脑背侧,视叶可分为顶盖和被盖,顶盖从外侧到内侧依次分为:带状层、外灰质层、浅白质层、中灰质层、中白质层、深灰质层、深白质层和中央灰质.被盖前端分层与顶盖相同,后端分层不明显.中脑腹侧包括被盖和大脑脚,HE染色和Holmes银染法显示,大脑脚从外向内颜色由浅变深,存在大量纵向神经纤维束,两脚底分界处有横向交错的神经纤维.被盖外侧细胞不分层,聚集形成核团.被盖内侧,细胞和纤维以中脑水管为中心,呈同心圆环分8层.通过比较蟾蜍中脑背腹差异程度,了解背腹功能不同.同时对中华蟾蜍中脑同其他脊椎动物的进行了比较.     

Axonal guidance in the chick visual system: posterior tectal membranes induce collapse of growth cones from the temporal retina

Membranes from posterior and anterior thirds of the chick optic tectum were added to explants from nasal and temporal retina. Posterior membranes, and to a lesser extent anterior membranes, cause temporal growth cones to collapse and their axonal processes to retract. Neither tectal source has an effect on nasal growth cones. We interpret these results to mean that there is a tectal activity, stronger in the posterior than the anterior region of the tectum, which helps guide growth cones during the development of the retinotectal map. We believe that in vivo this activity helps to steer temporal growth cones away from the posterior tectum. Nasal growth cones, which must map to the posterior tectum, are resistant to it. In vitro, when posterior membranes contact temporal growth cones over their surface, filopodia and lamellipodia withdraw rapidly. This leads to loss of contact between the growth cone and the substrate, followed by collapse.     

Anatomy of the fully formed chondrocranium of Podocnemis unifilis (Pleurodira: Podocnemididae)

This study describes the anatomy of the chondrocranium of Podocnemis unifilis (Pleurodira, Podocnemididae), based on recently hatched specimens, and cleared and double‐stained specimens. The orbitotemporal region is dramatically different from those observed for other species of turtles in that the: (1) planum supraseptale is greatly reduced and present only as tiny projections on the posterodorsal margin of the interorbital septum, (2) pila metoptica is free from all neighbouring structures and bifurcates distally, (3) pila antotica is greatly reduced, (4) foramina for optic nerve, ophthalmic artery and oculomotor nerves are open dorsally by virtue of this species lacking the taenia marginalis and taenia medialis, and (5) tectum synoticum is present and invested dorsally by the supraoccipital, despite the fact that this bone forms by replacement of the supraoccipital. The unique morphology of the pila metoptica is explained either as de novo formation of processes on the terminus of this cartilage or by retention of portions of the taenia medialis (anteriorly) and pila antotica or pila accessoria (posteriorly). Variation in the orbitotemporal region presented here is discussed for two other pleurodiran turtles (Phrynops hilarii and Emydura subglobosa) and briefly compared with the anatomy observed in Cryptodira.     

Comparative forefoot trabecular bone architecture in extant hominids

The appearance of a forefoot push-off mechanism in the hominin lineage has been difficult to identify, partially because researchers disagree over the use of the external skeletal morphology to differentiate metatarsophalangeal joint functional differences in extant great apes and humans. In this study, we approach the problem by quantifying properties of internal bone architecture that may reflect different loading patterns in metatarsophalangeal joints in humans and great apes. High-resolution x-ray computed tomography data were collected for first and second metatarsal heads of Homo sapiens (n = 26), Pan paniscus (n = 17), Pan troglodytes (n = 19), Gorilla gorilla (n = 16), and Pongo pygmaeus (n = 20). Trabecular bone fabric structure was analyzed in three regions of each metatarsal head. While bone volume fraction did not significantly differentiate human and great ape trabecular bone structure, human metatarsal heads generally show significantly more anisotropic trabecular bone architectures, especially in the dorsal regions compared to the corresponding areas of the great ape metatarsal heads. The differences in anisotropy between humans and great apes support the hypothesis that trabecular architecture in the dorsal regions of the human metatarsals are indicative of a forefoot habitually used for propulsion during gait. This study provides a potential route for predicting forefoot function and gait in fossil hominins from metatarsal head trabecular bone architecture.     

A NEW TRISTICHOPTERID (SARCOPTERYGII, TETRAPODOMORPHA) FROM THE UPPER FAMENNIAN EVIEUX FORMATION (UPPER DEVONIAN) OF BELGIUM

Abstract:  Additional material of a large specimen of tristichopterid fish from the Upper Famennian Evieux Formation of Belgium is described. This large fish was previously assigned to Tristichopteridae gen. et sp. indet. due to the lack of diagnostic anatomical data. New available material consists of the internal surface of the parietal shield, vomers and anterior part of the parasphenoid, subopercular and submandibulo-branchiostegal bones, and an internal view of the anterior part of the mandible. A possible autapomorphy of the new form from Belgium, Langlieria socqueti gen. nov. et sp. nov., is the absence of marginal teeth on the vomer except on its most lateral part. Apart from these features, it only differs from the genus Mandageria from Australia in the absence of marginal teeth between the dentary fang and the mandibular symphysis, in the presence of a raised marginal crest lateral to the anterior coronoid fang, and in the presence of numerous small marginal teeth on the premaxilla. It differs from the cosmopolitan genus Eusthenodon in a number of respects: the supratemporal, tabular, and postparietal bones are superficially fused, as are the intertemporal and parietal bones, the dermal ornament is proportionally very fine, and the denticulated field of the parasphenoid stands proud rather than being recessed into the body of the bone.     

Osteology and systematic position of the Eocene salmonid †Eosalmo driftwoodensis Wilson from western North America

The fossil salmonid †Eosalmo driftwoodensis was originally described from fragmentary specimens. Study of new material of this fossil species confirms that it is a stem-group salmonine, with a mixture of primitive and derived salmonine features in its skull, but with its postcranial skeleton essentially of modern salmonine construction. Two autapomophies define the genus †Eosalmo: a long anterodorsal process of the subopercle meeting the dorsal edge of the bone at an angle of about 60^o, and a thin dermal basihyal plate apparently lacking teeth. Its salmonine relationship is supported by eight derived features: (1) posterior part of frontal widely expanded above autosphenotic, (2) hyomandibular fossa on pterotic long, (3) posterior part of endopterygoid extending posteriorly and broadly overlapped by both metapterygoid and quadrate, (4) premaxillary process of maxilla extending dorsally at an angle larger than 10^o, (5) infraorbitals 3 to 5 narrow and covering less than anterior half of hyomandibula, (6) presence of suprapreopercle, (7) anterior end of preopercular canal on horizontal arm distinctiy turning to anteroventral corner of preopercle, (8) first uroneural amplified into large fan-shaped stegural, and (9) scales small, with more fhan two lateral line scales per vertebral centrum. Salmonidae are a monophyletic family defined by at least three synapomorphies: posterior surface of epiotic with sulcus, peg-and-socket connection in caudal skeleton, and tetraploid karyotype. Within the Salmonidae, Thymallinae and Salmoninae form a clade based on features from premaxilla, supramaxilla, anguloarticular, and supraorbital.     

Computerized tomography of the otic capsule and otoliths in the oyster toadfish,Opsanus tau

The neurocranium of the toadfish (Opsanus tau) exhibits a distinct translucent region in the otic capsule (OC) that may have functional significance for the auditory pathway. This study used ultrahigh resolution computerized tomography (100 µm voxels) to compare the relative density of three sites along the OC (dorsolateral, midlateral, and ventromedial) and two reference sites (dorsal: supraoccipital crest; ventral: parasphenoid bone) in the neurocranium. Higher attenuation occurs where structural density is greater; thus, we compared the X‐ray attenuations measured, which provided a measure of relative density. The maximum attenuation value was recorded for each of the five sites (x and y) on consecutive sections throughout the OC and for each of the three calcareous otoliths associated with the sensory maculae (lagena, saccule, and utricle) in the OC. All three otoliths had higher attenuations than any sites in the neurocranium. Both dorsal and ventral reference sites (supraoccipital crest and parasphenoid bone, respectively) had attenuation levels consistent with calcified bone and had relatively small, irregular variations along the length of the OC in all individuals. The lowest relative attenuations (lowest densities) occurred consistently at the three sites along the OC. In addition, the lowest attenuations measured along the OC occurred at the ventromedial site around the saccular otolith for all seven fish. The decrease in bone density along the OC is consistent with the hypothesis that there is a low‐density channel in the skull to facilitate transmission of acoustic stimuli to the auditory endorgans of the ear. J. Morphol. 276:228–240, 2015. © 2014 Wiley Periodicals, Inc.