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.     

Development of the cartilaginous skull in solea solea: trends in pleuronectiforms

The postembryonic development of the cartilaginous cephalic skeleton in Solea solea (Linné) was observed from hatching to the juvenile stage or postmetamorphic larva. Fry were trypsin-cleared and alcian-blue-stained. At hatching, Solea solea displays no cartilaginous structure. On day 3, the fry exhibit trabecular bars, Meckel's cartilages, palatoquadrates, hyosymplectics, hyoid bars, the first two ceratobranchials, and basibranchials 1–3. On day 4, ceratobranchials 3 and 4 form. On day 12, short taeniae marginales are formed on the well-developed otic capsules and the branchial basket is nearly complete. On day 14, a first cartilage reduction begins as the posterior part of the trabecula communis separates from the otic capsules. On day 18, metamorphosis begins. The tectum posterius completes posteriorly the braincase dome. On day 20, the left eye moves to the median crest of the head and the right eye shifts downwards slightly. Consequently, displacement of the right eye induces a deformation extending to the right pterygoid process. On day 23, both eyes are located on the same side. On day 29, cartilage reduction affects the hyosymplectics; it continues with the first two ceratobranchials on day 32 and with the otic capsules on day 40. On day 50, cartilage reduction has reached the lamina precerebralis. Only a few small lumps remain at the level of the neurocranium and the splanchnocranium.     

Postembryonic ontogeny and functional anatomy of the ligamentum mandibulo-hyoideum and the ligamentum interoperculo-mandibulare,with notes on the opercular bones and some other cranial elements in Salmo gairdneri Richardson, 1836 (Teleostei: Salmonidae)

This study deals with the development of the opercular bones the ligamentum mandibulo-hyoideum, the ligamentum interoperculo-mandibulare and with some aspects of the development of the ceratohyale and the cartilago meckeli during postembryonic ontogeny of Salmo gairdneri (rainbow trout). It is stated that the ligamentum mandibulo-hyoideum is present from hatching onward, while the development of the ligamentum interoperculo-mandibulare appears during the period of active feeding. Some functional reasons are proposed to explain these facts. It is further suggested that the opercular bones develop under influence of the opercular muscles.     

Early development of the cephalic skeleton of Barbus barbus (Teleostei, Cyprinidae)

The inception, and development of the cephalic skeleton of Barbus barbus from hatching to 24 days passes through periods of fast and slow growth; these rates are not the same in different parts of the skull. Trabeculae, parachordal plates, Meckelian cartilages and hyposymplectics are present at hatching. Then the cartilaginous floor of the neurocranium develops, the pars quadrata, the hyoid bars and branchial arches elements appear shortly before the first movable dermal bones, the dentaries, maxillae and opercles. The first bone of the braincase to appear is the parasphenoid; other bones develop subsequently and at the same time: the angular, quadrate, interopercle and fifth ceratobranchial. Later the splanchnocranium continues to develop at a relatively fast rate while the neurocranium shows little growth. The braincase does not begin to close before the 24th day, nor do the first bones of the skull roof appear, while the bucco-pharyngeal apparatus is complete, having the adult shape. The early constitution of the latter structures seems to be linked with the mechanical demands of biological functions such as breathing and feeding.     

Embryonic development of the sea bass Dicentrarchus labrax

The embryonic development of the sea bass Dicentrarchus labrax during the endotrophic period is discussed. An 8 cells stage, not reported for other studied species, results from two rapid successive cleavages. Blastula occurs at the eighth division when the embryo is made of 128 cells. During gastrulation, the infolded blastoderm creates the endomesoblastic layer. The Kupffer??s vesicle is reported to drive the left/right patterning of brain, heart and digestive tract. Heart formation starts at 8 pairs of somites, differentiation of myotomes and sclerotomes starts at the stage 18 pairs of somites; main parts of the digestive tract are entirely formed at 25 pairs of somites. At 28 pairs of somites, a rectal region is detected, however, the digestive tube is closed at both ends, the jaw appears the fourth day after hatching, but the mouth is not opened before the fifth day. Although cardiac beating and blood circulation are observed, gills are not reported in newly hatched individuals; eye melanization appears concomitant with exotrophic behavior.     

Development and evolution of craniofacial patterning is mediated by eye-dependent and -independent processes in the cavefish Astyanax

We studied the development and evolution of craniofacial features in the teleost fish, Astyanax mexicanus. This species has an eyed surface dwelling form (surface fish) and many different cave dwelling forms (cavefish) with various degrees of reduced eyes and pigmentation. The craniofacial features we examined are the tooth-bearing maxillary bones, the nasal and antorbital bones, the circumorbital bones, and the opercular bones, all of which show evolutionary modifications in different cavefish populations. Manipulations of eye formation by transplantation of the embryonic lens, by lentectomy, or by removing the optic vesicle showed that eye-dependent and -independent processes change both the surface fish and cavefish craniofacial skeletons. The size of the olfactory pits, which the nasal and antorbital bones define, and the size and positioning of the circumorbital bones were found to correlate with eye development. For the six suborbital bones (SO1-6), the relationship with the developing eye appears to be due to ossification initiated from foci in the suborbital canal of cranial neuromasts, whose patterning is also highly correlated with the presence or absence of an eye. By contrast, we found that the number of maxillary teeth, the number of SO3 bone elements, the positioning of SO4-6 with respect to the opercular bone, and the shape of the opercular bone are not dependent on eye formation and vary among different cavefish populations. The results suggest that evolution of the cavefish craniofacial skeleton is controlled by multiple developmental events, some a direct consequence of eye degeneration and others unrelated to loss of the eye.     

The presence of an embryonic opercular flap in amniotes

The operculum is a large flap consisting of several flat bones found on the side of the head of bony fish. During development, the opercular bones form within the second pharyngeal arch, which expands posteriorly and comes to cover the gill-bearing arches. With the evolution of the tetrapods and the assumption of a terrestrial lifestyle, it was believed that the operculum was lost. Here, we demonstrate that an embryonic operculum persists in amniotes and that its early development is homologous with that of teleosts. As in zebrafish, the second pharyngeal arch of the chick embryo grows disproportionately and comes to cover the posterior arches. We show that the developing second pharyngeal arch in both chick and zebrafish embryos express orthologous genes and require shh signalling for caudal expansion. In amniotes, however, the caudal edge of the expanded second arch fuses to the surface of the neck. We have detailed how this process occurs and also demonstrated a requirement for thyroid signalling here. Our results thus demonstrate the persistence of an embryonic opercular flap in amniotes, that its fusion mirrors aspects of amphibian metamorphosis and gives insights into the origin of branchial cleft anomalies in humans.     

Morphological and behavioural development of halibut, Hippoglossus hippoglossus (L.) larvae

Live yolk-sac halibut, Hippoglossus hippoglossus (L.) larvae from rearing experiments at Austevoll Aquaculture Station, Norway, were examined from hatching to past first feeding for developmental morphology and behaviour. The findings include development of the respiratory and circulatory organs, eye pigmentation, mouth formation, organs of the digestive system and the process of yolk absorption, as well as swimming speed and activity levels.
A stomodeum is not present at hatching although drinking is possible through a pair of branchial pits which gradually develop into the operculum and gill basket. The mouth normally opens slowly, the gape being restricted by a transverse septum until bones are formed. The amount of time spent swimming varies from less than 15% of the observation period during the first 2 weeks after hatching to between 70 and 100% around the seventh week after hatching, when individual differences become more apparent. Larvae generally react with a burst of swimming when two come into contact. Speed and duration of swimming seems to be correlated with development of eye pigment, heart size and fin formation. The yolk-sac period is divided into four stages.     

The mode of hatching of the monogenean Entobdella soleae, a skin parasite of the common sole (Solea solea).

Observations have been made on the hatching behaviour of the larva of the monogenean Entobdella soleae and on the effects of various enzymes on the opercular cement of the egg. These observations indicate that hatching is brought about by a proteolytic hatching fluid which is produced by two pairs of ventral head glands and spread over the inner surface of the opercular joint by rotation of the larva about its longitudinal axis. After hatching there is no trace of the ventral head glands. Similar glands are present before hatching in the oncomiracidium of E. hippoglossi.     

Why is grouper larval rearing difficult?: an approach from the development of the feeding apparatus in early stage larvae of the grouper,Epinephelus coioides

The osteological development of elements forming the oral cavity was examined in early stage larvae of the grouper,Epinephelus coioides, from hatching to 242.5 hours after hatching. By the time of initial mouth opening, at 54 hours after hatching, the fundamental elements, composed of the trabecula, some components of the lower branchial and hyoid arches, the quadrate and symplectic-hyomandibular cartilages, maxilla and Meckel's cartilage, had appeared. No further elements were observed until 165 hours after initial mouth opening, except some components in the lower branchial arch and head region. The appearance of new elements and initial ossification of existing cartilage occurred thereafter, but all elements related to feeding either had not appeared or had not started ossifying until 188.5 hours after initial mouth opening. Based on the morphology and developmental modes of these elements, the feeding mode of grouper larvae was considered to be “sucking/grasping.” However, the appearance and ossification of elements occurred slowly, with no transitional phase from sucking to grasping modes of feeding being observed during the study; such delayed development of the feeding-related bony elements was considered to be a cause of the difficulty in rearing early stage grouper larvae.     

The structure and function of the dermal pectoral girdle in bony fishes with particular reference to ostariophysines

The structure of the dermal pectoral girdle of teleostean fishes is analyzed in relation to its functions. In bony fishes the vertebral column, with a horizontal axis, and the pectoral girdle, with a basically vertical axis, form the only skeletal links between the head and the body. The individual bones of the dermal girdle are considered as supporting units joined by a series of articulations that permit differential movement between adjacent bones. The movements mediated by this linkage system are: lateral swinging of the head relative to the body, expansion of the distance between the central areas of the two pectoral girdles to permit passage of large food items, and fore-and-aft movements of the anteroventral ends of the cleithra relative to the skull. Among other factors affecting the structure of the dermal pectoral girdle are the provision for the support of the pectoral fin base and the requirement for the effective operation of a sleeve valve between the girdle and the opercular cover.
Modifications of the dermal pectoral girdle in ostariophysine fishes are discussed. A brief history of the bony fish girdle in terms of its functional components is postulated.     

Age validation,and comparison of otolith,vertebra and opercular bone for estimating age of <Emphasis Type="BoldItalic">Schizothorax o’connori</Emphasis> in the Yarlung Tsangpo River,Tibet

A collection of 514 Schizothorax o’connori was made between August 2008 and August 2009 from Yarlung Tsangpo River to assess the suitability of three bony structures for age estimation. The annulus characteristics of otolith, vertebra and opercular bone were described. Location of the first annulus was validated by daily growth increment (DGI) analysis in the otoliths. Annual periodicity was verified by marginal increment ratio (MIR) analysis in otoliths and edge analysis in vertebrae and opercular bones. Annuli formed, once a year, between March and May for all three bony structures. Otoliths, vertebrae and opercular bones were examined to determine which structure produced the most precise and accurate age estimates in S. o’connori. Vertebrae and otoliths matched closely for the first 21 years of life, while opercular bones appeared to underestimate age. For older fish, the counts diverged and otoliths consistently providing higher age estimates. Sectioned otoliths proved to be the most precise and accurate structure for age estimation. The oldest observed schizothoracine fish was 50, more than twice the longevity previously accepted in S. o’connori.     

Electron microscopic study of larval eye development in Turbellaria Polycladida

The appearance and development of the embryonic and larval eyes of the polyclad turbellarian Stylochus mediterraneus were studied. In the embryo, the left epidermal eye appears first. Subsequently, the right epidermal eye appears, and within hours it sinks into the parenchyma and turns into a cerebral eye. Newly hatched Götte's larvae possess both the left epidermal and the right cerebral eye. Three days after hatching, an incomplete eye appears adjacent to the left epidermal eye. The left cerebral eye then originates from this incomplete eye as it sinks into the parenchyma. This third eye is believed to originate through a process of induction.     

Parallel phenotypic evolution of skull-bone structures and head measurements of Arctic charr morphs in two subarctic lakes

Interspecific morph variations in trophic morphology related to skull-bones and head traits is associated to ecological segregation of Arctic charr morphs (genus Salvelinus) in two sub-arctic lakes (Fjellfrøsvatn and Skogsfjordvatn, Norway). The replicated morph pair, the profundal spawning benthivorous PB-morph and the littoral spawning omnivorous LO-morph of Arctic charr, diverge along the shallow-deep-water resource axis. In Skogsfjordvatn there is also a profundal spawning piscivorous PP-morph. The PB-morphs from both lakes have similar skull-bone traits and head morphology such as elongated jaw-bones, small opercular bones and relatively longer heads. The PP-morph also has an elongated head, relatively small opercular bones as well as larger jaw-bones. In contrast, the LO-morphs in both lakes have shorter jaw-bones, larger opercular bones in addition to relatively small heads. However, some small non-parallel differences exist among the morphs from the two lakes. Overall, all profundal morphs (PB and PP) have relatively similar skull-bone structures, suggesting adaptations to the deep-water environment but also to their separated dietary niches. There is strong evidence for parallel evolution with some local adaptations in skull-bones and head morphology of the PB-morph and the LO-morph from separate lakes.     

Feeding behaviour in Cylindrophis and its bearing on the evolution of alethinophidian snakes

Cylindrophis ruffus ingests prey using two distinct mechanisms. During initial phases of prey transport, lateral movements of the rear of the braincase combine with small unilateral movements of the toothed bones of each side; prey is usually constricted during this phase to permit the snake to push its head over the prey. Once transport has carried the leading part of the prey into the anterior oesophagus, Cylindrophis begins to use bilaterally synchronized movements of the jaw apparatus combined with low-amplitude, short wave-length flexions of the anterior vertebral column. Transport of prey is many times faster during the bilateral phase than during the unilateral phase.
Radiographic and cinematographic evidence indicates that the mandibular tips of Cylindrophis do not separate more than 1.5–2.0 times the resting distance between the dentary tips. Although this limits potential gape size, the intramandibular joint is highly mobile, allowing the mandibles to conform to a variety of prey shapes. Manipulations of anaesthetized and fresh, dead specimens revealed that the palatomaxillary arches are tightly attached to the ventral bones of the snout, movements of each arch being reflected in equivalent movements of the ipsilateral elements of the snout.
Cylindrophis represents a functional stage intermediate between most lizards with limited palatomaxillary kinesis and advanced snakes with considerable palatomaxillary mobility. Contrary to previous hypotheses, however, upper jaw liberation in Cylindrophis is due to liberation of the ventral snout, not to reduction of attachments to the braincase and snout. This suggests that the nose played a crucial role in the evolution of the feeding apparatus in alethinophidian snakes.     

A Turkish case of subcortical/subependymal heterotopia associated with corpus callosum dysgenesis, craniofacial dysmorphism, severe eye abnormalities, and growth-mental retardation

The patient is a 12-year-old boy with a history of learning disability, growth retardation, and strabismus. Weight, height and head circumference were below the 3rd percentile. A café-au-lait spot, 1x1 cm a diameter, on the back region and pectus excavatum deformity were diagnosed. He had facial asymmetry, a broad nose, sparse eyebrows and eyelashes, a rudimentary frontal sinus, deviation of the nasal septum, and bilateral small maxillary bones. The left orbital fossa was also mildly rudimentary. On eye examination the movements of the left globe to the upward and lateral side were limited and internal strabismus was noted at this side. Visual acuity was 1/10, bilaterally. Bilateral choroid coloboma, glaucoma, vertical and horizontal nystagmus were diagnosed. Fundoscopic examination revealed bilateral optic atrophy and macular and paramacular granulation tissues on the left side. Intelligence quotient was 46. Electroencephalography revealed bilateral frontal slow-wave activity. Visual evoked potential revealed prolonged p100 wave latencies bilaterally. Magnetic resonance imaging of the brain demonstrated corpus callosum dysgenesis, bilateral subcortical heterotopia in the frontal lobes and subependymal heterotopia in the posterior horn of the left ventricle. Chromosomal analysis revealed a normal male karyotype, 46, XY. Although several cases of heterotopia in association with mental retardation, craniofacial dysmorphism, cerebral, and eye abnormalities have been described the combination of abnormalities diagnosed in our case has not previously been reported. We hypothesize that the combination of subcortical/subependymal heterotopia, corpus callosum dysgenesis, craniofacial dysmorphism, severe eye abnormalities, and growth-mental retardation may be a new syndrome.     

The energetics of feeding strikes in larval carp, Cyprinus carpio

The prey intake of larval carp is described from high-speed (200–1250 frames s⁻¹) films with synchronous lateral and ventral views. Even in first-feeding carp larvae, the operculars are functional in sealing effectively the opercular slit until the moment of prey intake, and the maxillaries close off the corners of the mouth, preventing leak flow. In reducing the distance between larva and prey during attack, the relative importance of sucking the prey towards the mouth and swimming forward is variable; overall they are about equally important. The volume and the velocity of the water sucked into the mouth cavity during prey uptake are calculated. The energy costs of suction, i.e., accelerating the water sucked into the mouth cavity, during prey intake are estimated from these values. The energy costs of suction and swimming are in the same order of magnitude. Together they form only a fraction of 1% of the energetic content of the prey, so considerations about energy expenditure seem unimportant in a strategy to optimize the prey attack. During searching, however, they will be important. Power requirements during attack may also be important.     

Embryonic development and organogenesis in the snail Marisa cornuarietis (Mesogastropoda: Ampullariidae). V. Development of the nervous system.

The nervous system is ectodermal in origin. All nerve ganglia arise separately by proliferation and later delamination from the ectoderm, not by invagination. They become secondarily connected to one another by commissures and connectives developing as extensions from the peripheral layer of ganglionic nerve cells. Rudiments of the cerebral, pedal, pleural and intestinal (parietal) ganglia arise almost simultaneously at a relatively early stage (Stage V). The cerebral ganglia develop from the ectoderm of the head plates. Rudiments of the pedal and pleural ganglia are separate at their inception. They later fuse (Stage VI) to form a pleuro-pedal ganglionic mass on each side. The 2 intestinal ganglia are symmetrical at the beginning, but they soon lose their symmetry as a result of torsion. The right ganglion crosses to the left over the gut and persists as the supraintestinal ganglion. The left or subintestinal ganglion shifts to the right and forward, and fuses with the right pleural ganglion (Stage VIII), thus obscuring the chiastoneury. The paired buccal and single visceral (abdominal) ganglia start differentiating in Stage VII. The former develop from the ectodermal wall of the stomodaeum, while the visceral ganglion delaminates from the right wall of the visceral sac, then shifts to the left during torsion. The statocysts develop early (Stage V) from 2 ectodermal invaginations on either side of the rudimentary foot. They later separate from the overlying ectoderm and statoconi appear in their lumina. Contrary to earlier reports on related ampullariids, the osphradium proved to be ontogenetically older than the mantle and mantle cavity. It starts differentiating as a thickened ectodermal plate in the right wall of the visceral sac (Stage V). During torsion, it becomes engulfed in the mantle cavity and shifts to the left side, then is carried forward as the mantlegrow. The eyes develop late (Stage IX) as ectodermal invaginations which rapidly separate from the ectoderm to form closed vesicles. Their cells start differentiating before hatching to form the retina, in which pigment is deposited, and the inner cornea. The lens is secreted in the lumen of the eye and grows by addition of concentric layers of secretion.