Osteology,myology and feeding mechanism of Astronotus ocellatus (Pisces,Perciformes)

Summary Astronotus ocellatus captures its prey by creating a negative pressure in the buccal cavity which is caused by its quick expansion. Once the prey has been accommodated, the buccal cavity undergoes a compression which may propel the prey towards the pharyngeal jaws for mastication. The motion picture recordings indicate retracted premaxillae at the beginning of food intake followed by a maximum attainment of mouth gape and then mastication. During the maximum opening of the mouth the premaxillae are protruded and dentaries are at maximum depression. These events are followed by activities such as buccopharyngeal cavity expansion, bulging on the ventral surface of the head, and prominent curvature on the ventral surface anterior to the urohyal, caused by the upward movement of the glossohyal. Based on the cinematographic results, it may be inferred that the maximum mouth gape is caused by the sternohyoid-hyoid-interopercular-mandible coupling, and not by the opercular apparatus-mandible coupling, as the latter acts after the full descent of the lower jaw. Impression of the expanded buccopharyngeal cavity has been made by a paraffin mold technique, which confirms the displacement of the buccopharyngeal elements during expansion of the cavity.     

The functions and mechanisms of the protrusible upper jaws of two species of cyprinid fish

Photographs have been taken of Idus and Gobio feeding. Idus , a primitive cyprinid, protrudes its premaxillae very little when it opens its mouth to take in food, but closes the mouth by further protrusion of the premaxillae. This enables it to close its mouth while keeping its buccal cavity fully expanded. Gobio , a more advanced cyprinid, protrudes its premaxillae strongly as it opens its mouth and is thereby able to apply the mouth closely to a surface from which it is taking food. The mechanisms of the movements are described.     

Suctorial feeding in a deep-sea octopus as a means of niche partitioning (Cephalopoda: Octopodidae)

Molecular phylogenetic analyses indicate that two clades of deep-sea octopuses evolved at opposite ends of the earth to become globally sympatric. Coexistence of these overtly similar, but phylogenetically distinct octopuses requires some means of niche partitioning. To investigate details of feeding, the buccal complexes of specimens of each clade, Muusoctopus and Graneledone, were sectioned at 90° to the radular ribbon. The buccal complex of Muusoctopus is the same as reported in Octopus; the radula and its bolsters extend the length of the buccal complex. In Graneledone, however, the radula and its bolsters are restricted to anterior half of the buccal complex. Posterior to the radular sac, a vertically oriented muscle, named here the buccal abductor, extends from the floor of the mouth to the base of the buccal complex. In Muusoctopus, the bolsters extend the radula to bring food into the mouth; the palps propel it to the esophagus. In Graneledone, although the bolsters extend the radula, contraction of the buccal abductor to expand the posterior mouth may be the primary food mover. The negative pressure differential created draws food into the mouth and to the entry to the esophagus. The buccal abductor may permit the ingestion of larger pieces of prey, as gut contents show. Its evolution may represent a key innovation that heightens deep-sea octopus diversity.     

Functional analysis of a specialized prey processing behavior: winnowing by surfperches (Teleostei: Embiotocidae).

Several surfperches (Embiotocidae), including the black surfperch, Embiotoca jacksoni, exhibit a specialized prey handling behavior known as winnowing, in which ingested food and non-nutritive debris are separated within the oropharyngeal cavity. Prey items are swallowed, and unpalatable material is ejected from the mouth. Winnowing is believed to play an important role in the partitioning of food resources among sympatric embiotocids. We present a mechanistic model for this separative prey processing based on high-speed video analysis, cineradiography, electromyography, and buccal and opercular cavity pressure transducer recording. Winnowing by embiotocids is characterized by premaxillary protrusions repeated cyclically with reduced oral gape. Protrusion is accompanied by depression of the hyoid apparatus and adduction of the opercula. Alternating expansion and contraction of the buccal and opercular cavities generate regular pressure waveforms that indicate bidirectional water flow during processing. Separation of food from debris by Embiotoca jacksoni occurs in three phases. The prey-debris bolus is transported anteriorly and posteriorly within the oropharyngeal cavity and is then sheared by the pharyngeal jaws. Mechanical processing is complemented by the rinsing action of water currents during hydraulic prey transport. The feeding apparatus of Embiotoca jacksoni is functionally versatile, although not obviously specialized relative to that of nonwinnowing surfperches. Protrusion of the premaxillae and depression of the hyoid apparatus are critical to both prey capture and subsequent prey processing. The pharyngeal jaws exhibit kinematic patterns during separation of food from debris distinct from those observed during mastication of uncontaminated prey. This behavioral flexibility facilitates resource partitioning and the coexistence of E. jacksoni in sympatric embiotocid assemblages.     

The oral sucker of Gyrinocheilus aymonieri (Teleostei: Cypriniformes)

The sucker was studied in young and mature fish by light microscopy, histochemistry, transmission and scanning electron microscopy, X-ray probe microanalysis, dissection, staining preparations of whole skeletons, and watching the animals in aquaria. The fleshy lips are supported by highly flexible, chondroid tissues, the structure and histochemistry of which differ substantially from those of cartilage. They allow the sucker to evert when the fish attaches to a stone or aquarium wall and are connected to the maxillae, premaxillae and dentaries. Lining the inside of the lips are two horny rasps, each with several regular rows of small hooks. The scraping blades of these hooks are keratinized and point towards the mouth. They increase the coefficient of friction for adhesion and enable the fish to feed on encrusting algae. Between the posterior rasp and the -anterior margin of the mandible are two invaginations of the lower lip that extend the sucker chamber beneath large hollows in the dentaries. The anterior margin itself contacts the outer surface of the maxillary oral valve when the mouth is closed, and isolates the sucker chamber from the rest of the buccal and pharyngeal cavities. Contrary to previous views, it is thought that a true vacuum is produced, and that attached fish spend long periods without taking water in through the mouth. The attachments of the principal jaw muscles are described and their role in sucker action discussed. There are similarities with the jaw mechanism of catostomids.     

Feeding behavior and kinematics of the lesser electric ray, Narcine brasiliensis (Elasmobranchii: Batoidea)

Jaw protrusion is a major functional motif in fish feeding and can occur during mouth opening or closing. This temporal variation impacts the role that jaw protrusion plays in prey apprehension and processing. The lesser electric ray Narcine brasiliensis is a benthic elasmobranch (Batoidea: Torpediniformes) with an extreme and unique method of prey capture. The feeding kinematics of this species were investigated using high-speed videography and pressure transduction. The ray captures its food by protruding its jaws up to 100% of head length (approximately 20% of disc width) beneath the substrate and generating negative oral pressures (< or = 31 kPa) to suck worms into its mouth. Food is further winnowed from ingested sediment by repeated, often asymmetrical protrusions of the jaws (> 70 degrees deviation from the midline) while sand is expelled from the spiracles, gills and mouth. The pronounced ram contribution of capture (jaw protrusion) brings the mouth close enough to the food to allow suction feeding. Due to the anatomical coupling of the jaws, upper jaw protrusion occurs in the expansive phase (unlike most elasmobranchs and similar to bony fishes), and also exhibits a biphasic (slow-open, fast-open) movement similar to tetrapod feeding. The morphological restrictions that permit this unique protrusion mechanism, including coupled jaws and a narrow gape, may increase suction performance, but also likely strongly constrain dietary breadth.     

The mechanism of breathing in the South Asmerican lungfish, Lepidosiren paradoxa; radiological study

In filling the lungs by means of the buccal "force-pump", air drawn into the buccal cavity by the lowering of the floor of the mouth is forced into the lungs when the mouth is firmly closed and the buccal floor raised. This mechanism depends on the ability of the fish to prevent air being forced out of the mouth through the lips and it is shown that the tongue is pressed against the roof of the mouth to form a seal. Radiologically it has been shown that the hyoid apparatus ("ceratohyal" of some authors) plays a large part in the movements necessary to draw air into the mouth, seal it in and force it into the lungs. Another skeletal element involved is the pectoral girdle which makes considerable movements in association with the filling of the lungs. Expiration of the air from the lungs, which as shown by radiographs may be remarkably complete, is brought about by the elasticity of the lung tissue.     

Mechanics of the feeding action of a cyprinid fish

The pressures in the buccal cavities of Golden orfe (Idus idus) have been recorded as the fish took food from a tube connected to a pressure transducer. Pressures 50–105 cm water below the pressure of the surrounding water were recorded as the buccal cavity expanded to suck the food in. Calculations based on the dimensions of the muscles lead to the conclusion that the levatores hyoidei, sternohyoideus, rectus abdominis, epaxial trunk muscles and obliquus internus probably all had to exert almost their maximum isometric tensions, to produce the largest reductions of pressure. These muscles seem nicely balanced in strength for their functions in feeding.     

Functional morphology of the feeding system in the ruff — Gymnocephalus cernua (L. 1758) — (Teleostei,Percidae)

The ruff, Gymnocephalus cernua, is a European freshwater fish that feeds by sucking up small invertebrates from the bottom of ponds and slow flowing rivers. The feeding movements have been studied by simultaneous electromyography of seventeen muscles of the head and cinematographic techniques. A theoretical model of movements imposes the functional demands of suction upon an abstraction of the form of a teleost head. Three phases in the feeding act, a preparatory phase, a suction phase and a transport phase, could be correlated with the observed movements and EMGs. Differences between the predicted and the actual movement are discussed. Two different types of feeding occur. The direction, magnitude and duration of the suction forces during feeding are modified, according to the position of the prey. A mechanism preventing early mandibular depression allows sudden and strong suction. Retardation of the suspensorial abduction during the overall expansion of the buccal cavity is ascribed to kinetic interrelations with the hyoid arch. Protrusion of the upper jaws also permits an earlier closure of the mouth and directs the food-containing waterflow posteriorly. When the fish is feeding on sinking prey, protrusion occurs later in the sequence of movements than when it is feeding from the bottom. As the protruded jaws produce a downwardly pointed mouth this retardation aims the suction force.     

Dictyocaulus capreolus n. sp. (Nematoda: Trichostrongyloidea) from roe deer,Capreolus capreolus and moose,Alces alces in Sweden

Dictyocaulus capreolus n. sp. recovered from roe deer, Capreolus capreolus and moose, Alces alces in Sweden is described and figured. Morphological studies revealed the new species to be closest to D. eckerti and D. africanus on the basis of mouth shape, all three species having an elongate mouth opening. The other species of the genus, including D. viviparus, all have a circular to oval mouth opening. Dictyocaulus capreolus n. sp. can be distinguished from D. eckerti and D. africanus on the basis of the morphology of the buccal capsule and the bursa. These morphological studies support earlier evidence of the presence of a new species of Dictyocaulus in roe deer and moose that could be distinguished from D. eckerti and D. viviparus using either a PCR-linked hybridization assay or image analysis software to study the dimensions of the buccal capsule.     

Feeding and the control of volume within the tests of regular sea-urchins

The rigid, fluid filled, echinoid test presents problems of internal volume control when the vital functions of the animal involve the introduction of material, from outside the test, into the internal fluid systems. The volume of food in the guts of echinoids can be shown to vary considerably in the short term, to an extent which demands considerable capabilities for internal volume compensation. This is achieved by adjusting the quantity of fluid held within the gut so that, although the volume of food may vary widely, the total volume of the gut remains constant within narrow limits. The volume of gut fluid varies inversely with the quantity of food, and ingestion of food through the mouth is compensated by ejection of an equal volume of fluid from the anus. Defaecation is compensated by intake of fluid into the gut. On starvation, the gut becomes completely fluid filled and free from food.
The food of regular urchins is habitually compacted into regularly sized subpherical pellets covered with a tough mucus coating. The formation of coated pellets takes place in the buccal cavity and pharynx. The pellets remain discrete throughout their passage along the gut. The mucus coating resists digestion and remains intact even after defaecation. The functional morphology of the buccal cavity and pharynx is described in detail.
It is suggested that the functional significance of the pellets is associated with the gut fluid, volume compensation mechanism, which requires free flow of fluid throughout the length of the gut. It is also suggested that the siphon may be implicated in volume control, where fine adjustment of gut volume can be achieved by balancing ejection of fluid from the anus with intake of water through the mouth, but that the water intake is by the siphonal route, thus avoiding the necessity of flushing and diluting the enzyme rich digestive stomach region.     

Scaling of lunge feeding in rorqual whales: An integrated model of engulfment duration

Rorqual whales (Balaenopteridae) obtain their food by lunge feeding, a dynamic process that involves the intermittent engulfment and filtering of large amounts of water and prey. During a lunge, whales accelerate to high speed and open their mouth wide, thereby exposing a highly distensible buccal cavity to the flow and facilitating its inflation. Unsteady hydrodynamic models suggest that the muscles associated with the ventral groove blubber undergo eccentric contraction in order to stiffen and control the inflation of the buccal cavity; in doing so the engulfed water mass is accelerated forward as the whale’s body slows down. Although the basic mechanics of lunge feeding are relatively well known, the scaling of this process remains poorly understood, particularly with regards to its duration (from mouth opening to closure). Here we formulate a new theory of engulfment time which integrates prey escape behavior with the mechanics of the whale’s body, including lunge speed and acceleration, gape angle dynamics, and the controlled inflation of the buccal cavity. Given that the complex interaction between these factors must be highly coordinated in order to maximize engulfment volume, the proposed formulation rests on the scenario of Synchronized Engulfment, whereby the filling of the cavity (posterior to the temporomandibular joint) coincides with the moment of maximum gape. When formulated specifically for large rorquals feeding on krill, our analysis predicts that engulfment time increases with body size, but in amounts dictated by the specifics of krill escape and avoidance kinematics. The predictions generated by the model are corroborated by limited empirical data on a species-specific basis, particularly for humpback and blue whales chasing krill. A sensitivity analysis applied to all possible sized fin whales also suggests that engulfment duration and lunge speed will increase intra-specifically with body size under a wide range of predator-prey scenarios. This study provides the theoretical framework required to estimate the scaling of the mass-specific drag being generated during engulfment, as well as the energy expenditures incurred.     

Linking cranial kinematics,buccal pressure,and suction feeding performance in largemouth bass

The rate and magnitude of buccal expansion are thought to determine the pattern of water flow and the change in buccal pressure during suction feeding. Feeding events that generate higher flow rates should induce stronger suction pressure and allow predators to draw prey from further away. We tested these expectations by measuring the effects of prey capture kinematics on suction pressure and the effects of the latter on the distance from which prey were drawn-termed suction distance. We simultaneously, but not synchronously, recorded 500-Hz video and buccal pressure from 199 sequences of four largemouth bass, Micropterus salmoides, feeding on goldfish. From the video, we quantified several kinematic variables associated with the head and jaws of the feeding bass that were hypothesized to affect pressure. In a multiple regression, kinematic data accounted for 79.7% of the variation among strikes in minimum pressure. Faster mouth opening and hyoid depression were correlated with lower pressures, a larger area under the pressure curve, and a faster rate of pressure reduction. In contrast, buccal pressure variables explained only 16.5% of the variation in suction distance, and no single pressure variable had a significant relationship with suction distance. Thus, although expected relationships between head kinematics and buccal pressure were confirmed, suction distance was only weakly related to buccal pressure. Three explanations are considered. First, bass may not attempt to maximize the distance from which prey are drawn. Second, the response of prey items to suction-induced flow depends on prey behavior and orientation and is, therefore, subject to considerable variation. Third, previous theoretical work indicates that water velocity decays exponentially with distance from the predator's mouth, indicating that variation among strikes in flow at the mouth opening is compressed away from the mouth. These findings are consistent with other recent data and suggest that suction distance is a poor metric of suction feeding performance.     

Histology and growth pattern of the pachy-osteosclerotic premaxillae of the fossil beaked whale Aporotus recurvirostris (Mammalia, Cetacea, Odontoceti)

Beaked whales (Ziphiidae) often show highly specialized features, involving bone morphology or structure, in the rostral region of their skulls. Previous studies revealed an extremely derived and peculiar histological structure in the rostrum of the extant Mesoplodon densirostris. In order to assess if this structure is a general feature of ziphiids, the swollen premaxillae of Aporotus recurvirostris, a Miocene species from the North Sea, were studied histologically. These bones are pachyostotic and strongly osteosclerotic. However, their structural organization is entirely different from that of M. densirostris rostrum: they are basically made of a non-remodeled, laminar tissue that was cyclically deposited by the periosteum. As compared to the generalized structure of the premaxillae of toothed whales exemplified by the bottlenose dolphin, Tursiops truncatus, the pachyostotic condition of Aporotus premaxillae was obviously due to a particularly high and sustained growth-rate, occurring in a dorso-lateral direction. The osteosclerotic structure of these bones resulted from a complete lack of inner resorption activity. The histological features of Aporotus premaxillae indicate that these bones are not likely to have been hypermineralized, and thus, their physical properties must have differed from those of the M. densirostris rostrum. The possible functional involvements of rostral peculiarities in beaked whales are discussed with reference to the whole set of available comparative data.     

Studies on the tardigrades. I. Fine structure of the anterior foregut of Milnesium tardigradum Doyère

The fine structure of the anterior foregut of the tardigrade Milnesium tardigradum is presented. The oral region consists of a terminal mouth opening surrounded by six plate-like lips lying within a circlet of six prominent papillae. The buccal cavity is enclosed within a thick cuticular tube which possesses appendage structures, the stylet sheaths, stylet supports and paired protrusible stylets. Two large salivary glands envelop the buccal structures and contain voluminous amounts of secretory product. The arrangement, possible functions and phyletic significance of these structures are discussed.     

Non-respiratory podia of clypeasteroids (Echinodermata,Echinoides): II. Diversity

Summary There are five major types of non-respiratory podia in the Order Clypeasteroida: accessory, barrel-tipped, food groove, large food groove, and buccal. The anatomy of each type is intimately related to its function in the feeding mechanism of clypeasteroids. Accessory podia are found aborally and orally in some species, only aborally and ambitally in others. Accessory podia are largely sensory and manipulatory, but can be locomotory in the small fibulariids and juvenile sand dollars. Barrel-tipped podia have expanded disk muscles and connective tissue, and are usually found in two sizes, large and small. In species that have them they are usually restricted to the oral surface. These podia collect food and pass it towards the food grooves in the manner of a bucket brigade. Food groove podia are found only in species with food grooves. These podia are small, with reduced tip musculature and expanded secretory tissue for coating food with mucus. They transport food down the food grooves to the mouth. Large food groove podia are simply large versions of ordinary food groove podia. They help move the clumped food into the mouth area towards the buccals, and are found only in the Clypeasteridae and some scutellines. Buccal podia lack tip musculature, but possess tip support fibres and a single type of small secretory cell. They are sensory, and capable of manipulating particles into the mouth. Buccals are present in all families except the Clypeasteridae. Juvenile Echinarachnius less than 3 mm in diameter have only respiratory, accessory and buccal podia. Food groove and barrel-tipped podia start to differentiate from the accessories as the juvenile approaches a diameter of 4 to 5 mm. Clypeasteroid podial diversity increases the efficiency of the food collecting mechanism. The anatomy and distribution of podia on the oral surface of scutellines supports the fact that this surface is the prime food collecting area in all true sand dollars. The podia (not miliary spines) are the major source of mucus used during the feeding process and are the primary feeding appendages.     

A fish that uses its hydrodynamic tongue to feed on land

To capture and swallow food on land, a sticky tongue supported by the hyoid and gill arch skeleton has evolved in land vertebrates from aquatic ancestors that used mouth-cavity-expanding actions of the hyoid to suck food into the mouth. However, the evolutionary pathway bridging this drastic shift in feeding mechanism and associated hyoid motions remains unknown. Modern fish that feed on land may help to unravel the physical constraints and biomechanical solutions that led to terrestrialization of fish-feeding systems. Here, we show that the mudskipper emerges onto land with its mouth cavity filled with water, which it uses as a protruding and retracting ‘hydrodynamic tongue’ during the initial capture and subsequent intra-oral transport of food. Our analyses link this hydrodynamic action of the intra-oral water to a sequence of compressive and expansive cranial motions that diverge from the general pattern known for suction feeding in fishes. However, the hyoid motion pattern showed a remarkable resemblance to newts during tongue prehension. Consequently, although alternative scenarios cannot be excluded, hydrodynamic tongue usage may be a transitional step onto which the evolution of adhesive mucosa and intrinsic lingual muscles can be added to gain further independence from water for terrestrial foraging.     

Phylogenetic analysis of the pearlfish tribe Carapini (Pisces: Carapidae)

Fishes of the tribe Carapini (Encheliophis and Carapus) share a noteworthy peculiarity: they shelter in holothurian echinoderms or bivalve hosts. Some species are considered parasitic, others commensal. This study focuses on the phylogeny of the tribe, using two other Carapidae species as an outgroup (Snyderidia canina and Onuxodon fowleri). Insofar as possible, the selected anatomical and behavioural characters where chosen in an ecomorphological perspective, as features that could be responses to various lifestyle‐related constraints. Our character selection also took into account the fact that some features are (presumably) linked. Such features were grouped together as a single trait to avoid their overvaluation. This methodology enabled commensals to be separated from parasites, the former belonging to Carapus and the latter to Encheliophis. Carapus species reflect in their morphology the constraints imposed by a diet of hard, mobile, elusive prey, showing predator‐type features: a strong dentition, a wide mouth opening, a robust food intake apparatus. On the other hand, the endoparasitic Encheliophis species show a generally weaker buccal apparatus and narrow mouth opening, in relation to the different constraints of their lifestyle where the diet constraints are less pronounced: they eat body parts of their host. Changes in both generic diagnoses are proposed and three species are transferred from Encheliophis to Carapus.     

Structure of the mouth opening and pharyngeal apparatus in three species of Baikal Cottoidei in relation to their feeding

The structure of the mouth opening and pharyngeal apparatus was studied, and its relation to the feeding pattern in three coastal species of Baikal Cottidae fish was analyzed. Analysis of data demonstrated that the general pattern of structure of the mouth opening and pharyngeal apparatus is similar in the species under study since their food is represented by bottom organisms, mainly by amphipods. Interspecies differences are manifested in an increase in relative values of morphological features and the size of consumed prey.     

Stomatogenesis during sexual and asexual reproduction in an amicronucleate strain of Paramecium caudatum.

Amicronucleate cells of Paramecium caudatum, whose micronuclei have been artifically removed by micropipetting, are characterized by the appearance of a deciliated area at the posterior part of the buccal opening. These cells form food vacuoles at a slightly lower rate than micronucleate cells. Their mean interfission time is longer than that in micronucleates. The exconjugants of amicronucleate cells can not form food vacuoles and eventually die witout fission, though conjugation proceeds normally in them as well as in their micronucleate mate. The oral apparatus of amicronucleate exconjugants seems to be shallower than that of micronucleates. The membranellar cilia, therefore, can be seen through the buccal overture by scanning electron microscope. The results obtained from the cross of micronucleate and amicronucleate strains and from the induction of autogamy in amicronucleate strains suggest that the micronucleus has a primary role in developing the normal oral apparatus after nuclear reorganization.