Morphogenesis of the nasal apparatus of the red deer (Cervus elaphus L.)

Three stages of morphogenesis of the nasal apparatus of the red deer (Cervus elaphus L.) were studied. Many ancestral traits reminiscent of relationships in other mammals and even in reptiles were found, including a cart, ectochoanalis, paraseptal cartilages, the septum nasi and its ventral trabecular enlargement, a lamina transversalis ant., clear separation of the cart, parietotectalis and cart, paranasalis from each other and a crista semicircularis. A maxilloturbinale, was present, but not a nasoturbinale. The main specific features were a completely rostrally localized, peculiar cartilaginous structure in the preseptal space, for which there is as yet no morphological explanation, and pronounced bulging of the cartilaginous wall of the nasal capsule in a ventrolateral direction, level with the rostral region of the olfactory labyrinth (caudally to the aboral end of the maxilloturbinale). In the early stages of morphogenesis, it was found that the ethmoturbinalia might be formed by fusion of the edges of the anlagen of the paranasal cartilage and the lamina orbitonasalis. The structure of the olfactory labyrinth was reminiscent of its organization in the sheep embryo; the recessus frontalis was completed by a series of frontoturbinal recesses and frontoturbinalia, which are poorly developed in the red deer, however. The floor of the caudal part of the nasal capsule was very little developed and there was no cart, paraseptalis post.     

An acoustic valve within the nose of sperm whales Physeter macrocephalus

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Histologic studies of cartilaginous structures in the anterior region of the head of the sperm whale Physeter macrocephalus

Recently discovered cartilaginous structures in the forehead of the sperm whale (Behrmann and Klima 1985) were investigated histologically. The largest and most important of these structures is the nasal roof cartilage which can be derived from the tectum nasi, a part of the embryonic nasal capsule (Klima et al. 1986). In the investigated sperm whale fetuses, this structure consists of embryonic hyaline cartilage which is well suited for morphogenetic processes and fast growth. In the investigated adult sperm whale, the originally hyaline cartilage has been transformed into a special kind of elastic cartilage. The arrangement of cells, territories, and extracellular substance resembles hyaline cartilage. This component represents an adaptation to pressure load. The appearance and arrangement of elastic fibres resembles elastic cartilage. This component is an adaptation to distortion forces. Obviously, pressure and distortion are the strongest mechanical strains that the nasal roof cartilage is exposed. We see the function of this cartilage structure therein that, being a pressure-elastic skeletal support and following the left nasal meatus along its whole extension through the massive and soft forehead, it secures the only direct respiratory passage. Additionally, fibre bundles of transversely striated muscles are anchored in the perichondrium of the nasal roof cartilage. The function of this delicately interwoven muscle system is seen by us in the fine tuning of contraction and dilatation of the respiratory passage. Moreover, a possible function as a sound conducting cartilaginous structure serving the echolocation system is considered (c.f. Pilleri et al. 1983).     

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.     

Aspects of hadrosaurian cranial anatomy

Supraorbital bones in Saurolophus angustirostris are described, and their presence in all hadrosaurs is suggested. Frontal-nasal and premaxillar-nasal fontanellae are distinguished in hadrosaurs; their presence is explained as connected with growth and considered to he responsible for the variability of crest structures. New data indicating the presence of a cartilaginous diverticulum nasi within the circumnarial depression in Saurobphus ongustirostris are presented. A physiological (respiratory and/or thermoregulatory) function of the nasal diverticulum is proposed.     

Contribution to knowledge of the morphogenesis of the nasal apparatus of the fallow deer (Dama dama L.)

As in the red deer, in the fallow deer embryo we found a number of ancestral structures reminiscent of relationships in other mammals, such as paraseptal cartilages, a septum nasi with trabecular widening, a lamina transversalis ant., a cart. ectochoanalis, a capsule wall with a roof and a lateral wall formed of a clearly distinguishable cart. parietotectalis and cart. paranasalis, an ethmoturbinale I projecting a long way rostrally and additionally, in the fallow deer, cart. paraseptales posteriores. I regard the relationship of the cart. alaris inf. to the parietotectal cartilage (or "marginoturbinale") as relatively "primitive"; this may mean that the term "atrioturbinale" is also justified in mammals and that the relevant structure is homologous with the one known by the same name in birds. The specializations found during study of the morphogenesis of the nasal apparatus in the red deer (Slaby 1990b) are accentuated in the fallow deer. The chief ones are the specific rostral processes of the anlage of the nasal septum, which are a significant part of reinforcement of the nostril, the marked widening of the nasal capsule in a lateral direction (so that even the paranasal cartilages have a largely horizontal course), the striking ventrolateral bulge in the nasal capsule at the beginning of the olfactory region and the final resultant decrease in the height (i. e. flattening) of the capsule. This leads to reduction of the frontoturbinalia and their corresponding recesses, which - where they are developed - are oriented more horizontally. The structure of ethmoturbinale I, together with its insertion, is also simplified. As in the corresponding red deer embryo, the paranasal cartilage zone in the anterior part of the olfactory region is strikingly thickened; the frontoturbinalia do not, however, originate (in our stage) by the formation of cavities in the cartilage, but develop as simple processes. A crista semicircularis and foramen epiphaniale and also, as distinct from the red deer embryo, cart. paraseptales posteriores, are clearly discernible. In conclusion, it can therefore be claimed that the morphogenesis of specialized cervid features is accentuated in Dama more than in Cervus and that relationships in the fallow deer represent a further step in specialization, or - if we are speaking of the development of radiations - specialization here has progressed further.     

THE SPERM WHALE'S NOSE: SEXUAL SELECTION ON A GRAND SCALE?1

The world's largest nose belongs to the sperm whale, yet its functional significance remains equivocal. In order to help shed light on its function, the head of a postmortem neonate sperm whale was subjected to CT scanning. Geometric comparisons between homologous cephalic structures in sperm whales and dolphins (normalized for body size) show extreme hypertrophy and size sexual dimorphism in the sperm whale's lipid spermaceti organ. Anatomic geometry, energetics, and behavior suggest that this immense nasal apparatus is a bioacoustical machine. Sexual selection via an acoustic display is suggested as an explanation for the size and continuous (physiologically isolated) energy investment in the construction and maintenance of the male's spermaceti organ.     

Sound transmission in the nose of the sperm whale Physeter catodon. A post mortem study

During a sperm whale stranding at R?m?, the Wadden Sea, Denmark, on 4 December 1997, we were notified in time to start acoustic transmission measurements in the spermaceti complex 1 h after the specimen was seen alive. Frequency-modulated sound pulses, sweeping from 30 kHz to 10 kHz in 25 ms, were injected at the frontal surface at two positions: at the distal sac, and at the center of the junk (a compartmentalized structure below the spermaceti organ). A hydrophone next to the projector served as receiver. The analyses of the recordings show a repetitive, decaying reflection pattern at both projection sites, reminiscent of the multi-pulse click peculiar to sperm whales, although with minor differences in the duration of the intra-click intervals. This experimental evidence supports the Norris and Harvey (1972) theory of click generation in the spermaceti organ. Accordingly, the click is composed of a primary event, followed by a train of reflected pulses, spaced by the time required for the event to travel back and forth between air sacs (reflectors) at each end of the organ. The results also show that the junk readily transmits sound and probably is in acoustic contact with the spermaceti organ.     

Morphology of the interorbital region of Saimiri sciureus

The skull of the platyrrhine primate Saimiri sciureus is distinguished by a large interorbital fenestra. Juvenile skulls still show a bony interorbital septum with some small gaps. A morphogenetic study was undertaken to better understand the structures of the interorbital region, which represents a linkage between the base of the braincase and the nasal skeleton. Already in early ontogenetic stages a reduction of the posterior portion of the nasal capsule and of the cartilaginous interorbital septum are observed, resulting in the formation of a primary interorbital fenestra. A bony interorbital septum is mainly formed in perinatal age stages by ossification of the presphenoid and by medial fusion of the frontals; the primary interorbital fenestra is retained as a small opening. It only occurs in late juvenile stages when the definitive interorbital fenestra develops by by secondary transformation of bone into a membrane of dense connective tissue; this process is most probably caused by mechanical friction of the very closely approximated eyes of both sides.     

Dlx5 and Dlx6 expression in the anterior neural fold is essential for patterning the dorsal nasal capsule

Morphogenesis of the vertebrate facial skeleton depends upon inductive interactions between cephalic neural crest cells (CNCCs) and cephalic epithelia. The nasal capsule is a CNCC-derived cartilaginous structure comprising a ventral midline bar (mesethmoid) overlaid by a dorsal capsule (ectethmoid). Although Shh signalling from the anterior-most region of the endoderm (EZ-I) patterns the mesethmoid, the cues involved in ectethmoid induction are still undefined. Here, we show that ectethmoid formation depends upon Dlx5 and Dlx6 expression in a restricted ectodermal territory of the anterior neural folds, which we name NF-ZA. In both chick and mouse neurulas, Dlx5 and Dlx6 expression is mostly restricted to NF-ZA. Simultaneous Dlx5 and Dlx6 inactivation in the mouse precludes ectethmoid formation, while the mesethmoid is still present. Consistently, siRNA-mediated downregulation of Dlx5 and Dlx6 in the cephalic region of the early avian neurula specifically prevents ectethmoid formation, whereas other CNCC-derived structures, including the mesethmoid, are not affected. Similarly, NF-ZA surgical removal in chick neurulas averts ectethmoid development, whereas grafting a supernumerary NF-ZA results in an ectopic ectethmoid. Simultaneous ablation or grafting of both NF-ZA and EZ-I result, respectively, in the absence or duplication of both dorsal and ventral nasal capsule components. The present work shows that early ectodermal and endodermal signals instruct different contingents of CNCCs to form the ectethmoid and the mesethmoid, which then assemble to form a complete nasal capsule.     

Mechanisms of control of alae nasi muscle activity.

Human upper airway dilator muscles are clearly influenced by chemical stimuli such as hypoxia and hypercapnia. Whether in humans there are upper airway receptors capable of modifying the activity of such muscles is unclear. We studied alae nasi electromyography (EMG) in normal men in an attempt to determine 1) whether increasing negative intraluminal pressure influences the activity of the alae nasi muscle, 2) whether nasal airway feedback mechanisms modify the activity of this muscle, and 3) if so, whether these receptor mechanisms are responding to mucosal temperature/pressure changes or to airway deformation. Alae nasi EMG was recorded in 10 normal men under the following conditions: 1) nasal breathing (all potential nasal receptors exposed), 2) oral breathing (nasal receptors not exposed), 3) nasal breathing with splints (airway deformation prevented), and 4) nasal breathing after nasal anesthesia (mucosal receptors anesthetized). In addition, in a separate group, the combined effects of anesthesia and nasal splints were assessed. Under each condition, EMG activity was monitored during basal breathing, progressive hypercapnia, and inspiratory resistive loading. Under all four conditions, both load and hypercapnia produced a significant increase in alae nasi EMG, with hypercapnia producing a similar increment in EMG regardless of nasal receptor exposure. On the other hand, loading produced greater increments in EMG during nasal than during oral breathing, with combined anesthesia plus splinting producing a load response similar to that observed during oral respiration. These observations suggest that nasal airway receptors have little effect on the alae nasi response to hypercapnia but appear to mediate the alae nasi response to loading or negative airway pressure.(ABSTRACT TRUNCATED AT 250 WORDS)     

On the origin of the so-called Meckelian ossicles in the nasal skull of odontocetes

Although Cave (1987) accepts the theory that the Meckelian ossicles originate from the maxilloturbinals, evidence given in his study in fact supports the opinion of Klima and van Bree (1985) that the Meckelian ossicles arise from elements of the nasal floor, solum nasi, of the embryonic nasal capsule, in particular from the lamina transversalis anterior and the cartilago paraseptalis.     

Extracellular matrix in development of the intervertebral disc.

Intervertebral discs allow bending and twisting of the spine whilst resisting compression from gravity and muscle action, and are composite structures of the peripheral annulus fibrosus enclosing the nucleus pulposus. Their development is complex, involving several different connective tissue types, yet little is known of the developing extracellular matrix (ECM). We report the ECM composition of foetal rat discs from their first appearance to birth. The earliest collagen detected was type III, which was subsequently replaced by type II in the cartilaginous inner annulus and joined by type I in the fibrous outer annulus. Type IV collagen appeared in outer annulus, associated with myofibroblast-like cells of the orienting collagenous lamellae. Laminin and fibronectin co-distributed here in later stages, although overall they had a wider distribution. Aggrecan occurred in early nucleus pulposus and then appeared in the inner annulus, in association with cartilage differentiation. Versican appeared later in the inner annulus, and also in the dorsal region of the outer annulus. Comparisons of glycosaminoglycan and proteoglycan label allowed extrapolations to be made as to likely glycosaminoglycan components of the large proteoglycans, and of other proteoglycans that may be present - thus differential distribution of aggrecan and keratan sulfate label suggested the presence of fibromodulin and/or lumican. Functionally aggrecan would confer compression resistance to cartilaginous structures. Versican may also contribute, but along with the small proteoglycans is likely to be associated with various stages of control of cell differentiation, tissue morphogenesis and collagen fibre formation in the assembly of the annulus fibrosus.     

Dynamic rhinoplasty for the plunging nasal tip: functional unity of the inferior third of the nose

To achieve permanent results for the correction of a drooping nasal tip, it is important to understand the mechanism responsible for the caudal rotation of the tip when a person speaks or smiles. This mechanism can be considered to depend on a "functional unity" formed by three components: (1) the cartilaginous framework (alar cartilages and accessories acting as a single structure); (2) muscular motors (m. levator labii superioris alaeque nasi and depressor septi nasi); and (3) gliding areas (apertura piriformis, the valvular mechanism between the upper lateral cartilages and alar cartilages, the lax tissue of the nasal dorsum, and the membranous septum). We describe a new anatomical and functional concept responsible for the plunging of the nasal tip. When a person smiles, the functional unit is activated by a combination of two forces acting simultaneously in opposite directions that rotate the tip caudally and elevate the nasal base. The levator moves the alar base upward and the depressor pulls the tip caudally. To correct the drooping tip, the transcartilaginous incision is extended laterally, and the lateral portion of the alar arch is dissected free from the skin and the mucosa, thus exposing the accessory cartilages. The arch is then severed at the level of the accessories to allow the cephalad rotation of the domes. The muscle insertions are dissected free from the accessories and a section of the muscle and, if necessary, the accessory cartilages, is removed. From January of 1991 onward, 312 patients have had this ancillary procedure performed in addition to the basic rhinoplasty technique.     

Development of the chondrocranium in the suckermouth armored catfish Ancistrus cf. triradiatus (Loricariidae, Siluriformes)

The chondrocranium of the suckermouth armored catfish Ancistrus cf. triradiatus was studied. Its development is described based on specimens ranging from small prehatching stages with no cartilage visible, to larger posthatching stages where the chondrocranium is reducing. Cleared and stained specimens, as well as serial sections, revealed a cartilaginous skeleton with many features common for Siluriformes, yet several aspects of A. cf. triradiatus are not seen as such in other catfishes, or to a lesser extent. The skull is platybasic, but the acrochordal cartilage is very small and variably present, leaving the notochord protruding into the hypophyseal fenestra in the earlier stages. The ethmoid region is slender, with a rudimentary solum nasi. A lateral commissure and myodomes are present. The larger posterior myodome is roofed by a prootic bridge. The maxillary barbel is supported by a conspicuous cartilaginous rod from early prehatching stages. The ceratohyal has four prominent lateral processes. Infrapharyngobranchials I-II do not develop. During ontogeny, the skull lengthens, with an elongated ethmoid, pointing ventrally, and a long and bar-shaped hyosymplectic-pterygoquadrate plate. Meckel's cartilages point medially instead of rostrally.     

The ethmoidal region of Dibamus taylori (Squamata: Dibamidae), with a phylogenetic hypothesis on dibamid relationships within Squamata

The nasal capsule and nasal sac of Dibamus taylori are described based on a plate reconstruction of serially sectioned material and are compared with several other squamate taxa. The ethmoidal region of D. taylori is characterized by a short nasal vestibule, a well developed concha, a much reduced and nearly functionless Jacobson's organ, an incomplete nasal roof, and a true nasopharyngeal duct with a complicated secondary palate. Cartilaginous and epithelial structures of the ethmoidal region provide many informative and useful characters for a phylogenetic analysis. Different anatomical features of D. taylori are compared with other burrowing forms ( Acontias meleagris, Annietta, pulchra Voetzkowia mira, Feylinia curron ) and discussed functionally and phylogenetically. Phylogenetic analysis of all higher squamate taxa using 144 available data derived from many regions of the body indicates relative relationships as (((Dibamidae, Amphisbaenia) Serpentes) Scleroglossa).     

Allometric patterns of fetal head growth in mysticetes and odontocetes: Comparison of Balaena mysticetus and Stenella attenuata

Unlike other mammals, odontocetes and mysticetes have highly derived craniofacial bones. A growth process referred to as “telescoping” is partly responsible for this morphology. Here, we explore how changes in facial morphology during fetal growth relate to differences in telescoping between the adult odontocete Stenella attenuata and the mysticete Balaena mysticetus. We conclude that in both Stenella and Balaena head size increases allometrically. Similarly, odontocete nasal length and mysticete mouth size have strong positive allometry compared to total body length. However, the differences between odontocetes and mysticetes in telescoping are not directly associated with their fetal growth patterns. Our results suggest that cranial changes related to echolocation and feeding between odontocetes and mysticetes, respectively, begin during ontogeny before telescoping is initiated.     

孤雌生殖长角血蜱的哈氏器超微结构与发育

为阐明长角血蜱Haemaphysalis longicornis孤雌生殖种群的哈氏器结构及发育特征, 用扫描电镜对其各虫期哈氏器进行了观察, 分析了血餐对哈氏器发育的影响。结果表明： 该种群幼蜱、 若蜱和成蜱哈氏器形态结构基本相同, 均由前窝和后囊构成。幼蜱前窝感毛6根, 位于同一基盘; 若蜱和成蜱哈氏器相似, 前窝感毛7根, 其中1根孔毛位于外侧基盘, 另6根感毛位于内侧基盘。各虫期饱血后哈氏器大小均比饥饿状态下显著增大（P<0.05）。幼蜱前窝与后囊面积比值与若蜱相比无显著差异（P>0.05）, 若蜱前窝与后囊面积比值与成蜱相比差异显著（P<0.05）。各虫期哈氏器均在发育, 且血餐对哈氏器发育有重要影响。幼蜱至若蜱期哈氏器前窝与后囊的发育速度相似, 若蜱至成蜱期哈氏器前窝发育快于后囊。本研究结果在一定程度上揭示了孤雌生殖长角血蜱的哈氏器发育规律。