Nasal Capsule Research Articles

Morphology and evolution of the jaw suspension in lamniform sharks

The morphology of the jaw suspension and jaw protrusion mechanism in lamniform sharks is described and mapped onto a cladogram to investigate how changes in jaw suspension and protrusion have evolved. This has revealed that several evolutionary modifications in the musculoskeletal apparatus of the jaws have taken place among lamniform sharks. Galeomorph sharks (Carcharhiniformes, Lamniformes, Orectolobiformes, and Heterodontiformes) have paired ethmopalatine ligaments connecting the ethmoid process of the upper jaw to the ethmoid region of the cranium. Basal lamniform sharks also acquired a novel single palatonasal ligament connecting the symphysis of the upper jaw to the cranium mid-ventral to the nasal capsule. Sharks in the family Lamnidae subsequently lost the original paired ethmopalatine ligament while retaining the novel palatonasal ligament. Thus, basal lamniform taxa (Mitsukurina owstoni, Carcharius taurus, Alopias vulpinnis) have increased ligamentous support of the lateral region of the upper jaw while derived species (Lamnidae) have lost this lateral support but gained anterior support. In previous studies the morphology of the jaw suspension has been shown to play a major role in the mechanism of upper jaw protrusion in elasmobranchs. The preorbitalis is the primary muscle effecting upper jaw protrusion in squalean (sister group to galeomorphs) and carcharhiniform (sister group to lamniforms) sharks. The preorbitalis originates from the quadratomandibularis muscle and inserts onto the nasal capsule in squalean and carcharhiniform sharks. Carcharhiniform sharks have evolved a subdivided preorbitalis muscle with the new division inserting near the ethmoid process of the palatoquadrate (upper jaw). Alopid sharks have also independently evolved a partially subdivided preorbitalis with the new division inserting at the base of the ethmoid process and surrounding connective tissue. Lamnid sharks have retained the two preorbitalis divisions but have modified both of the insertion points. The original ventral preorbitalis division now inserts onto the connective tissue surrounding the mid-region of the upper jaw, while the new dorsal preorbitalis division inserts onto the surrounding connective tissue and skin at a more posterior position on the upper jaw. The retractor muscle of the jaws, the levator hyomandibularis, has also been modified during the evolution of lamniform sharks. In most sharks, including basal lamniforms, the levator hyomandibularis inserts onto the hyomandibula and functions to retract the jaws after protrusion. In alopid and lamnid sharks the levator hyomandibularis inserts primarily onto the upper and lower jaws around the jaw joint and is a more direct route for retracting the jaws. Thus, there has been at least one instance of character loss (ethmopalatine ligament), acquisition (palatonasal ligament), subdivision (preorbitalis), and modification (ventral preorbitalis, dorsal preorbitalis, and levator hyomandibularis) in the ligaments and muscles associated with the jaw suspension and jaw protrusion mechanism in lamniform sharks. While derived lamniform sharks (Lamna nasus, Carcharodon carcharius, and Isurus oxyrinchus) lost the ancestral passive lateral support of the ethmoid articulation of the upper jaw, they simultaneously acquired muscular support by way of the levator hyomandibularis, which provides a dynamic mechanism for lateral support. The evolution of multiple divisions of preorbitalis insertions onto the palatoquadrate and modification of the levator hyomandibularis insertion directly onto the jaws provides an active mechanism for multiple protractions and retractions of the upper jaw, which is advantageous in those sharks that gouge or saw pieces from large oversized prey items.

Respiratory cooling in rattlesnakes

We used infrared thermography to study respiratory cooling in the rattlesnakes (Viperidae: Crotalinae) and to partition the effects of air temperature, humidity, and activity levels on head–body temperature differences. We observed a single, cooled region centered around the mouth and nasal capsule that extended across the pit membrane at air temperatures above 20 °C. Both head and body temperatures of rattlesnakes increased linearly with air temperature. Head–body temperature differentials also increased with air temperature, but declined significantly at higher relative humidities. Rattling rattlesnakes exhibited significantly greater head–body temperature differentials than did resting rattlesnakes. We suggest that respiratory cooling may provide a thermal buffer for the thermoreceptive pit organs at high air temperatures, but caution that this adaptive hypothesis must be tested with direct neural or behavioral assays.

Prenatal growth of the interorbital septum in Macaca mulatta

The interorbital septum is the portion of the anterior cranial base directly contiguous with the eyes. It is considered to be a primitive trait that has evolved independently in various primate groups as a result of ocular and olfactory convergence with concomitant encephalization. This process is hypothesized to have reduced the size of the lateral nasal capsule exposing the anterior cranial base to the ocular apparatus and thus creating an interorbital septum. The purpose of this study was to determine whether differential growth trajectories occur among the chondrocranial elements corresponding to this hypothesis. Macaca mulatta embryos from the Zingeser Collection were selected for this analysis since this primate shows a well-developed interorbital septum throughout ontogeny. Embryos between 40 and 90 days of gestation were selected from the collection and coronal sections including the eye, anterior cranial base and lateral nasal capsule were subjected to video microscopy and computerized reconstruction using SURFdriver Software. Tissue volumes were computed for these tissues while chondrocytic growth attributes were measured utilizing stereological techniques. Results showed a strong correlation between the volume of the lateral nasal capsule and anterior cranial base and these two structures showed a consistent correlation with an increasing eye volume. Chondrocytic volume density and average diameter were larger in the lateral nasal capsule while shape, numerical density and average volume did not differ between the two tissues. These data suggest if differential growth does account for a reduction of the nasal capsule compared to the central cranial base stem, it does not appear to result from differential tissue size change. However, certain cellular growth activities leading to premature chondrocytic hypertrophy may be involved.

Ontogeny and homology of the paranasal sinuses in Platyrrhini (Mammalia: Primates)

The identity and taxonomic distribution of paranasal sinuses among living platyrrhines has remained a contentious issue (e.g., Cave [1967] Am J Phys Anthropol 26:277-288 vs. Hershkovitz [1977] Chicago: University of Chicago Press) largely because the ontogenetic data required for their detection and identification (e.g., Cave [1967]; Maier [2000] Cambridge, UK: Cambridge University Press, 99-132.) were not attainable without sacrificing valuable juvenile and subadult specimens. Non-invasive computed tomography (CT) scanning of ontogenetic series of skulls for 10 platyrrhine genera demonstrates the presence of maxillary and ethmoid sinuses, as well as homologs of the human sphenoid and frontal sinuses. Differences in the latter two sinuses between platyrrhines and hominoids highlight the need for early developmental data in establishing sinus homology. In particular, the identification of homologous recesses in the cartilaginous nasal capsule, from which sinuses later develop, emerges as the critical step. This developmental approach also reveals that the anterior and posterior ethmoid sinuses are each sets of serial homologs, a point which reconciles previous difficulties in establishing sinus homologies across mammalian orders (e.g., Paulli [1900] Gegenbaurs Morphol Jahrb 28:147-178, 179-251, 483-564).

Early development of the nose in human embryos: a stereomicroscopic and histologic analysis.

To analyze the morphologic features of the nose in the human embryo from the 4th to 8th developmental week according to Carnegie stage. Stereomicroscopic and histologic analysis of the morphology of the human embryo. A total of 27 cases of embryos, ranging from Carnegie stage 13 to 23, were analyzed. The external morphology was observed with a stereomicroscope, photographed, and analyzed. The histologic features were observed with a light microscope in the horizontally transected specimens stained with hematoxylin-eosin. The nasal placode was observed in stage 13, and it became flat or even concave in stage 14. In stage 15, the nasal pit was formed. In stage 16, the nasal sac and nasal fin were observed. In stage 17, the oronasal membrane was formed by thinning of the nasal fin. In stage 18, the primitive choana was established by a rupture of the oronasal membrane. In stage 19, the lateral palatine process projected vertically below the level of the tongue. The cartilaginous nasal capsule was formed in stage 20. In stage 21, the olfactory area was localized to the upper portion of the lateral nasal wall and the nasal septum. In stage 22, the lateral palatine process developed in a somewhat horizontal orientation. In stage 23, the premaxilla and primitive choana were formed. The development of the nose is most active from Carnegie stage 13 to 19, which corresponds to the end of the fourth embryonic week to the end of the seventh week. Thus, this period is considered to be the most important period in human nasal embryonic development.

Relationships of Lithogenes villosus Eigenmann, 1909 (Siluriformes, Loricariidae): Evidence from High-Resolution Computed Microtomography

The unusual neotropical freshwater catfish Lithogenes villosus Eigenmann, 1909 was described from a single specimen collected in 1908 at Aruataima Falls, a cataract of the upper Potaro River of Guyana, as a new genus and species of the armored catfish family Loricariidae. There are no subsequent published reports of additional specimens. Lithogenes villosus is unusual among the armored loricariid catfishes in being nearly naked, or unplated. The dermal plates which typically encase the head and body of other representatives of the Loricariidae are restricted in Lithogenes villosus to three paired series of small plates on the posterior portion of the trunk, plus a set of three plates on either side of the lateral cheek. Its taxonomy has been debated largely because it is intermediate in morphology between the armored loricariids and the naked astroblepids; these two families are presently regarded as sister groups among the larger assemblage of neotropical loricarioid catfishes. Nijssen and Isbrücker (1987, Revue Française d'Aquariologie et Herpétologie 13: 93–98.) transferred Lithogenes villosus, without comment or justification, from the Loricariidae to the Astroblepidae, and several recent presentations have renewed the controversy surrounding Lithogenes relationships by asserting that Lithogenes is well supported as the sister group of the Astroblepidae. Lithogenes has not been subjected to a thorough systematic investigation because it is thus far known only from the holotype, a situation that precludes rigorous comparative anatomical survey via traditional, destructive methods. Given its unusual morphology and putative basal position among the loricariids, ultimate resolution of Lithogenes relationships is crucial to resolving several issues in siluriform systematics. In this study, three-dimensional reconstructions of computed tomography (CT) scans of the holotype made at 13-μm resolution revealed details of internal anatomy informative of its relationships. Parsimony analysis of 41 characters obtained from the CT reconstructions, gross examination of external features, and previously published characters surveyed among loricarioid and outgroup catfishes indicated that Lithogenes shares a rich suite of derived characters with loricariids, most notably the presence of a mesethmoid condyle, posterior placement of the nasal capsule, closed pterotic aperture, lateral ethmoid lamina, and metapterygoid contacting the lateral ethmoid, and is the sister group of all other loricariids, exclusive of astroblepids. Evaluation of this dataset revealed that support for alternative relationships was considerably worse. A sister-group relationship between Lithogenes and astroblepids occurred in less than 1% of 10,000 bootstrap replicate trees and required 7 additional steps relative to the shortest, most parsimonious tree. A sister-group relationship between astroblepids and loricariids, exclusive of Lithogenes villosus, was not observed among the bootstrap replicates; forcing this topological constraint required nine additional steps relative to the shortest tree. Although Lithogenes villosus shares several morphological similarities with astroblepids, most of these represent symplesiomorphy and are therefore not indicative of relationships.

Chondrogenic differentiation during midfacial development in the mouse: in vivo and in vitro studies

Because the mouse is now the main model for developmental research of all types, it is important to understand the basic developmental pattern of various organs. The first aim of the present study was to establish normal prenatal developmental standards of the cartilaginous nasal capsule during embryonic development of the mouse. For this purpose we have performed sagittal and coronal sections ranging from E12.5 to E18.5 in gestation age. The primordia of the nasal septal cartilage is recognizable around the 14th embryonic day as demonstrated by the metachromatic toluidine blue staining and by immunostaining of type II collagen. Northern blot analysis of the transcription factors Cart-1 and Sox-9 indicated maximum mRNA levels at E12.5 then a decreased expression during the following days of gestation. Type II collagen and aggrecan mRNA levels are constant during the embryonic period. In the second part of this study, we have established a primary culture system where chondrocytes were isolated from E.18 mouse embryo nasal septum. The purpose of this second part was to assess if chondrocytes could further differentiate in vitro until the hypertrophic phase and matrix mineralization. After the condensation phase, the cells synthesize an extracellular matrix including type II collagen and aggrecan. Progressively, typical cartilaginous nodules composed of clusters of round cells are visible, then increase in size and finally mineralize at day 12 of culture. Cart-1 and Sox-9 mRNA levels remain constant throughout the cultures, whereas type II collagen and aggrecan gradually decrease. Ultrastructural observations of the nodules show typical chondrocytes embedded in a dense network of fibers with matrix vesicles and mineralized foci. Other ultrathin sections revealed the presence of chondrons, typical of hyaline cartilage. Results from this study provide useful tools to further investigate morphogenesis and differentiation of the cartilaginous nasal capsule, and could in the future serve as a basic developmental standard.

Biomechanical investigations on the skulls of reptiles and mammals

The skulls of reptiles and mammals can be loaded mechanically in three ways: the weight of the head acting downward, perhaps reinforced by a prey or bunch of food lifted from the ground or water surface; by forces acting in the plane of the tooth row, created by movements of the prey in relation to the head or by a movement of the head in relation to a fixed food object; and by the adduction of the mandible, which leads to reaction forces in the skull. While the former two evoke stress patterns comparable to that in a beam which is supported at its rear end (by the occipital condyle(s) and the neck muscles), the latter evoke stress patterns comparable to a beam supported at both ends. Its anterior bearing are the teeth which transmit a reaction force from the seized prey, the adductor muscles of the mandible move the intermediate part of the skull downward, and the posterior bearing is provided by the mandibular joint. Three-dimensional FEM-analysis of the flow of stresses within solid, homogeneous bodies under loads like those described above have been made. As a result, the stress flows have been found to correspond closely to the arrangements of bony material in the akinetic skulls of Crocodiles, Lacertilia, Sphenodon. Except crocodiles and chelonians, reptilian skulls often show large gaps between the load-bearing plates and rods. These gaps correspond to little stressed areas between the stress-bearing parts. One of the stress-bearing rods is the small braincase. In long, slender jaws like those in crocodiles the stresses are concentrated on the periphery, with more or less stress-free areas in the center of the cross sections. In many mammals (shrews, primates includingHomo), however, the very large bony nasal capsule and braincase lead to a distribution of the forces over large areas like in thin-walled shell structures, which are strong enough to sustain the existing forces, without reinforcing superstructures. Even the zygomatic arch can be dispensible. The decisive a priori factors which determine the development of either a rod- or a shell-like structure in a FEM model are 1st the relative shape and length of the toothrow and its position in relation to the posterior part of the skull, especially the braincase, and 2nd the size of the nasal capsule and the braincase. We conclude that the exact form of the skull in both classes of animals is determined by 1st the shape and length of the jaws and 2nd by the space requirements of the olfactory and the optical sense organs, and the braincase. The second factor is an expression of the overall evolutionary level. The literature contains plausible biological arguments to explain the high selective influence of lifestyle characteristics on the first factor. These arguments usually cover also the position of the eye openings, the nasal opening and the relative height and length of the whole skull. If these factors are given, the exact morphology of the bony structure turns out to correspond completely to the pattern of stresses, and no other reasons behind skull shape must be searched. The arrangement of the muscles seems to follow in all cases the principle to distribute the force created at the origines on a large surface or on many individual bony elements.

The Neurocranium of the Stereospondyl Mastodonsaurus Giganteus

New and rich material of the stereospondyl amphibian Mastodonsaurus giganteus from Kupferzell in southern Germany allows for the first time a detailed study of the neurocranium of this species. Both sections and uncrushed specimens preserved in three dimensions were examined. The sphenethmoid ranges from the sella turcica region far to the anterior almost towards the nasal capsules, which remained unossified. The basisphenoid persisted as cartilage, whose morphology can be traced from imprints in surrounding bones. The otic is a single but complex element which underwent major ontogenetic changes. During development, the posterior braincase became increasingly ossified and finally formed a single, compound unit. In addition, otic and sphenethmoid are broadly co‐ossified in large specimens. The basioccipital is present but rudimentary, wedged in between the exoccipital and parasphenoid and without contribution to the occipital condyles. The course of the optic, trigeminal, and facial nerves is studied in detail. The homology of the passages is assessed by means of phylogenetic arguments and comparative anatomical data, based on observations on nearest living crown groups.

Ventilation foetale et développement cranio-maxillaire

Data acquired by means of color Doppler ultrasound very explicitly suggest what the role of the fetal ventilation and nasal capsules in the morphogenesis of the maxillary prognathism, turbinates, nasal valves and nasopharynx could be. Furthermore, the dysmorphologies observed in Apert or Crouzon craniosynostosis, achondroplasia or unilateral cleft lip would also testify that the influence of the fetal ventilatory dynamics goes beyond the limits of the face and extends to the cranial base and the cranium. The wealth of raised hypothesis thanks to the contribution of this imaging system could question the validity of some conceptions of the fetal craniomaxillary morphogenesis.

Development of the facial midline.

"Intellectual excellence lies in having faith in the observation of apparently nontranscendental and unimportant facts. To observe an anatomic element calmly, with an open, analytical spirit, and with spiritual freedom, can lead to an explosive vortex of new knowledge."-Miguel Orticochea, M.D.(1) Traditional descriptive embryology based upon the interaction of frontonasal, lateral nasal, and medial nasal prominences is incapable of explaining the three-dimensional development of the facial midline. The internal structure of the nose and that of the oronasal midline can best be explained by the presence of paired A fields originating from the prechordal mesendoderm, associated with the nasal and optic placodes, supplied by the internal carotid artery, and sharing a common genetic coding with the prosomeres of the forebrain. Mesial drift of these fields leads to fusion of their medial walls; this in turn provides bilateral functional matrics within which form the orbits ethmoids, lacrimals, turbinates, premaxillae, vomerine bones, and the cartilages of the nose. This two-part paper reports six lines of evidence supporting the field theory model of facial development: (1) An apparent watershed exists in the midline of the base between the territories of the internal and external carotid systems. Isolation of the ICA in injected fetal specimens confirmed that the demarcation was distinct and restricted to the embryonic nasal capsule. (2) Field theory explains the developmental anatomy of the contents of the nasal capsule. (3) The neuromeric model of CNS development provides a genetic basis for the anatomy and behavior of fields. (4) Mutants for the Dlx5 gene demonstrate A field deletion patterns. These experiments relate the nasal placode to the structures of the A fields. (5) Separate regions of the original nasal placodes give rise to neurons, which are dedicated to separate sensory and endocrine systems. The A fields constitute the pathways by which these neurons reach the brain. (6) Non-cleft lip-related cleft palate, holoprosencephaly, and the Kallmann syndrome are clinical models that demonstrate the effects of anatomic disturbances within the A fields.

The braincase, mandible and dental structures of the Early Devonian lungfish Dipnorhynchus kurikae from Wee Jasper, New South Wales

Dipnorhynchus kurikae Campbell & Barwick was described from the Receptaculites Limestone Member at Wee Jasper. Our new specimens include material of the braincase, the snout, the nasal capsules, the palate and the mandible which were not previously available. The new specimens give new data on the pores in the snout, the arrangement of the soft tissues in the nasal capsule, the growth of the hard tissues on the surface of the dentary, the formation of the hard surface on the palate and the prearticulars, the arrangement of sacculus and the nerves VII, IX and X in the otic region, and the interpretation of the palatoquadrate. The lateral lines on the mandible have been exposed by excavation. Thin sections show that the bone on the overgrowth of the dentary is of the same kind as occurs in the cores of the ridges on the tuberosities and marginal ridges on the palate and prearticular. A thin layer of dentine-like columns which contain no dentine tubules overlie the bony cores. This, and the mode of growth, demonstrate that the dental plates—being made of a bony core—have different patterns of deposition from tooth plates in most later Palaeozoic genera, which are made of dentine. This throws light on the way tooth plates evolved from a dipnorhynchid pattern through a speonesydrionid pattern to true tooth plates. The complex tooth plates of Diabolepis are considered to be independently developed. Ventrally the parasphenoid of Dipnorhynchus kurikae does not divide the pterygoids, but stops short at the posterior end of the pterygoids. The semicircular canals and the superior sinus of the saccular cavity are described from the well-preserved material. In comparison with D. sussmilchi, this species has a number of distinctive features, and is easily recognised as a separate species.

Immunoidentification of CRF-like material in the intermaxillary glands during Bufo arenarum development

The presence of corticotropin-releasing factor-like material in the intermaxillary glands was studied by immunocytochemical techniques during the metamorphosis of Bufo arenarum. The intermaxillary glands appeared at stage XV (midprometamorphosis) with CRF-like material slightly immunoreactive. These glands are located posterior to the premaxillae and between the nasal capsules in the roof of the mouth and are formed of alveoli or tubules. During metamorphic climax, corticotropin-releasing factor-like material was identified strongly immunostained at the apices of the secretory cells. It was observed that collecting ducts of the gland open to the anterior palatal surface suggesting that the secretion could be ingested by tadpoles. Our results clearly showed that ir-CRF-like material present in the intermaxillary glands is ingested by tadpoles during metamorphosis and could play an important role during amphibian development.

The External Nasal Cartilages in Chiroptera: Significance for Intraordinal Relationships

Detailed description and comparison of the prenatal and postnatal morphology of the external nasal cartilages across bats and their relatives have identified morphological characters that contribute to the resolution of phylogenetic conflicts concerning the relationships among bat subgroups. In this study, I include observations on the morphology of the cupula nasi anterior and of the processus alaris superior in chiropteran species not previously examined, including members of the Rhinolophidae and Megadermatidae. Contrary to recent results, based on analysis of single-copy DNA hybridization and DNA sequence data from nuclear and mitochondrial genes, morphology of the nasal cartilages supports the traditional view that Microchiroptera is monophyletic and that Rhinolophoidea is more closely related to other microchiropteran families than it is to Pteropodidae. A dorsal apertura nasi externa diagnoses the grouping Megadermatidae + Rhinolophidae and Rhinopomatidae and the presence of a cartilago accessoria appears to be a putative synapomorphy of the clade Rhinolophidae + Megadermatidae. Most Microchiroptera share a commissura alicupularis and a shell-shaped processus alaris superior. The few extant Microchiroptera that lack the shell-shaped processus alaris superior show a very different form of the anterior nasal capsule and superior alaris process: the processus alaris superior is integrated as part of the cupular side wall (e.g., emballonurids, megadermatids), or has a half-tube shape (rhinolophids). Based on observations of a sickle-shaped superior alaris process in Megachiroptera and other potential relatives of the Chiroptera, it seems most likely that a shell-shaped processus alaris superior is a synapomorphy of the living microchiropteran bats.

Goosecoid acts cell autonomously in mesenchyme-derived tissues during craniofacial development.

Mice homozygous for a targeted deletion of the homeobox gene Goosecoid (Gsc) have multiple craniofacial defects. To understand the mechanisms responsible for these defects, the behavior of Gsc-null cells was examined in morula aggregation chimeras. In these chimeras, Gsc-null cells were marked with beta-galactosidase (beta-gal) activity using the ROSA26 lacZ allele. In addition, mice with a lacZ gene that had been introduced into the Gsc locus were used as a guide to visualize the location of Gsc-expressing cells. In Gsc-null<->wild-type chimeras, tissues that would normally not express Gsc were composed of both Gsc-null and wild-type cells that were well mixed, reflecting the overall genotypic composition of the chimeras. However, craniofacial tissues that would normally express Gsc were essentially devoid of Gsc-null cells. Furthermore, the nasal capsules and mandibles of the chimeras had defects similar to Gsc-null mice that varied in severity depending upon the proportion of Gsc-null cells. These results combined with the analysis of Gsc-null mice suggest that Gsc functions cell autonomously in mesenchyme-derived tissues of the head. A developmental analysis of the tympanic ring bone, a bone that is always absent in Gsc-null mice because of defects at the cell condensation stage, showed that Gsc-null cells had the capacity to form the tympanic ring condensation in the presence of wild-type cells. However, analysis of the tympanic ring bones of 18.5 d.p.c. chimeras suggests that Gsc-null cells were not maintained. The participation of Gsc-null cells in the tympanic ring condensation of chimeras may be an epigenetic phenomenon that results in a local environment in which more precursor cells are present. Thus, the skeletal defects observed in Gsc-null mice may reflect a regional reduction of precursor cells during embryonic development.

Dlx5 regulates regional development of the branchial arches and sensory capsules.

We report the generation and analysis of mice homozygous for a targeted deletion of the Dlx5 homeobox gene. Dlx5 mutant mice have multiple defects in craniofacial structures, including their ears, noses, mandibles and calvaria, and die shortly after birth. A subset (28%) exhibit exencephaly. Ectodermal expression of Dlx5 is required for the development of olfactory and otic placode-derived epithelia and surrounding capsules. The nasal capsules are hypoplastic (e.g. lacking turbinates) and, in most cases, the right side is more severely affected than the left. Dorsal otic vesicle derivatives (e. g. semicircular canals and endolymphatic duct) and the surrounding capsule, are more severely affected than ventral (cochlear) structures. Dlx5 is also required in mandibular arch ectomesenchyme, as the proximal mandibular arch skeleton is dysmorphic. Dlx5 may control craniofacial development in part through the regulation of the goosecoid homeobox gene. goosecoid expression is greatly reduced in Dlx5 mutants, and both goosecoid and Dlx5 mutants share a number of similar craniofacial malformations. Dlx5 may perform a general role in skeletal differentiation, as exemplified by hypomineralization within the calvaria. The distinct focal defects within the branchial arches of the Dlx1, Dlx2 and Dlx5 mutants, along with the nested expression of their RNAs, support a model in which these genes have both redundant and unique functions in the regulation of regional patterning of the craniofacial ectomesenchyme.

Morphology of the nasal capsule of Heloderma suspectum with comments on the systematic position of helodermatids (Squamata: Helodermatidae)

A serially sectioned embryonic head of Heloderma suspectum formed the basis for a three‐dimensional reconstruction of the cartilaginous nasal capsule and its membrane bones. Further, the soft parts of the nasal capsule were analysed microscopically. The embryonic nasal capsule is described and compared to the morphological conditions in other lizards, especially the varanids, which many recent authors believe to be their closest relatives.The results support the close affinities between the helodermatids and the varanids. Both groups have a well developed recessus lateralis in a cartilaginous capsule, entered by the nasal concha. Further, in both groups a fragmentation of the cartilago paraseptalis occurs in a comparable location and form. Heloderma has no double lacrimal duct, in contrast to previous accounts, but the two canaliculi at the origin of the duct fuse noteworthy far ahead (in other squamates they fuse in front of the bulbus oculi). In the varanids the two branches of the lacrimal duct remain separate and open separately.Some unique features of Heloderma are presented as well, above all the presence of a second concha.

Anatomy of the feeding apparatus of the nurse shark, Ginglymostoma cirratum

The anatomy of the feeding apparatus of the nurse shark, Ginglymostoma cirratum, was investigated by gross dissection and computer axial tomography. The labial cartilages, jaws, jaw suspension, muscles, and ligaments of the head are described. Palatoquadrate cartilages articulate with the chondrocranium caudally by short, laterally projecting hyomandibulae and rostrally by ethmoorbital articulations. Short orbital processes of the palatoquadrates are joined to the ethmoid region of the chondrocranium by short, thin ethmopalatine ligaments. In addition, various ligaments, muscles, and the integument contribute to the suspension of the jaws. When the mouth is closed and the palatoquadrate retracted, the palatine process of the palatoquadrate is braced against the ventral surface of the nasal capsule and the ascending process of the palatoquadrate is in contact with the rostrodorsal end of the suborbital shelf. When the mandible is depressed and the palatoquadrate protrudes slightly rostroventrally, the palatoquadrate moves away from the chondrocranium. A dual articulation of the quadratomandibular joint restricts lateral movement between the mandible and the palatoquadrate. The vertically oriented preorbitalis muscle spans the gape and is hypothesized to contribute to the generation of powerful crushing forces for its hard prey. The attachment of the preorbitalis to the prominent labial cartilages is also hypothesized to assist in the retraction of the labial cartilages during jaw closure. Separate levator palatoquadrati and spiracularis muscles, which are longitudinally oriented and attach the chondrocranium to the palatoquadrate, are hypothesized to assist in the retraction of the palatoquadrate during the recovery phase of feeding kinematics. Morphological specializations for suction feeding that contribute to large subambient suction pressures include hypertrophied coracohyoideus and coracobranchiales muscles to depress the hyoid and branchial arches, a small oral aperture with well-developed labial cartilages that occlude the gape laterally, and small teeth. J. Morphol. 241:33–60, 1999. © 1999 Wiley-Liss, Inc.

Anatomy of the feeding apparatus of the nurse shark, Ginglymostoma cirratum.

The anatomy of the feeding apparatus of the nurse shark, Ginglymostoma cirratum, was investigated by gross dissection and computer axial tomography. The labial cartilages, jaws, jaw suspension, muscles, and ligaments of the head are described. Palatoquadrate cartilages articulate with the chondrocranium caudally by short, laterally projecting hyomandibulae and rostrally by ethmoorbital articulations. Short orbital processes of the palatoquadrates are joined to the ethmoid region of the chondrocranium by short, thin ethmopalatine ligaments. In addition, various ligaments, muscles, and the integument contribute to the suspension of the jaws. When the mouth is closed and the palatoquadrate retracted, the palatine process of the palatoquadrate is braced against the ventral surface of the nasal capsule and the ascending process of the palatoquadrate is in contact with the rostrodorsal end of the suborbital shelf. When the mandible is depressed and the palatoquadrate protrudes slightly rostroventrally, the palatoquadrate moves away from the chondrocranium. A dual articulation of the quadratomandibular joint restricts lateral movement between the mandible and the palatoquadrate. The vertically oriented preorbitalis muscle spans the gape and is hypothesized to contribute to the generation of powerful crushing forces for its hard prey. The attachment of the preorbitalis to the prominent labial cartilages is also hypothesized to assist in the retraction of the labial cartilages during jaw closure. Separate levator palatoquadrati and spiracularis muscles, which are longitudinally oriented and attach the chondrocranium to the palatoquadrate, are hypothesized to assist in the retraction of the palatoquadrate during the recovery phase of feeding kinematics. Morphological specializations for suction feeding that contribute to large subambient suction pressures include hypertrophied coracohyoideus and coracobranchiales muscles to depress the hyoid and branchial arches, a small oral aperture with well-developed labial cartilages that occlude the gape laterally, and small teeth. Copyright 1999 Wiley-Liss, Inc.

Development of the cetacean nasal skull.

The adaptation of cetaceans to aquatic life habits is reflected, in their nasal region, in three marked changes from the original relations found in land mammals. These changes include (1) the loss of the sense of smell, (2) translocation of the nostrils from the tip of the rostrum to the vertex of the head, and (3) elongation of the anterior head to form a rostrum protruding far towards anterior. The morphogenetic processes taking place during embryogenesis of the nasal skull play a decisive part in the development of all these changes. The lateral parts of the embryonic nasal capsule, encompassing the nasal passages, change their position from horizontal to vertical. At the same time, the structures of the original nasal floor (the solum nasi) are shifted in front of the nasal passages towards the rostrum. The structures of the original nasal roof (the tectum nasi) and of the nasal side wall (the paries nasi) are translocated behind the nasal passages towards the neurocranium. The medial nasal septum (the septum nasi) mostly loses its connection to the nasal passages and is produced into a point protruding far towards anterior. The transformed embryonic nasal skull of the Cetacea can be divided into three sections: 1. The median structures. These include the cartilaginous structures, viz., the rostrum nasi, the septum interorbitale and the spina mesethmoidalis, which are accompanied by the dermal bones, the vomer and the praemaxillare. In adult cetaceans the rostrum nasi is mostly preserved as a robust cartilage of the skull, which may possibly serve as a sound transmitting structure of the sonar system, or it may be responsible for the sensing of water streams and vibrations. 2. The posterior side wall structures. These include the following cartilaginous structures that are mostly heavily reduced or mutually fused: the cupula nasi anterior, the tectum nasi, the lamina cribrosa, the paries nasi, the commissura orbitonasalis, the cupula nasi posterior, the processus paraseptalis posterior, the crista semicircularis, the frontoturbinale, the ethmoturbinale I and the maxilloturbinale. The cartilaginous structures are largely accompanied by the dermal bone, the maxillare. Of these embryonic elements, very little is preserved in adult cetaceans. The cartilages of the cupula nasi anterior form the variable skeleton around the nostrils. In Physeter the tectum nasi forms a very long cartilaginous bar that passes through the whole giant anterior head of the sperm whale as a structure accompanying the left nasal passage. 3. The anterior side wall structures. These include the cartilaginous structures, viz., the cartilago ductus nasopalatini, the cartilago paraseptalis, the processus lateralis ventralis and the lamina transversalis anterior, accompanied by the dermal bones, the praemaxillare and the vomer. These structures participate in the formation of the robust rostrum of the cetacean skull, and they are partly preserved even in adults in the form of the isolated ossa pararostralia (the Meckelian ossicles). The comparison of morphogeny of the nasal skull has also made it possible to draw certain conclusions on the phylogeny and systematics of Cetacea. Already the earliest embryonic stages permit us to discern weighty transformations of the original nasal skull of land mammals. These transformations are common to all embryos examined. This fact indicates a common origin of all Cetacea, which thus form a single monophyletic order. However, later embryonic stages show some different modifications of the nasal capsule according to which at least three major groups can be distinguished within the order Cetacea, probably ranking as superfamilies: Balaenopteroidea, Physeteroidea and Delphinoidea. Our observations, being in full accordance with other morphological, and embryological, as well as molecular biological results, suggest that the division of the order Cetacea into two suborders, Mysticeti and Odontoceti, is no longer tenable.