Taste bud form and distribution on lips and in the oropharyngeal cavity of cardinal fish species (Apogonidae, Teleostei), with remarks on their dentition

The oral dentition and type and number of taste buds (TB) on the lips and in the oropharyngeal cavity were compared by means of SEM in 11 species of cardinal fishes (Apogonidae) belonging to five genera. The occurrence of a dense cover of skin papillae on the lips of some species (e.g., Apogon frenatus), as well as differences in structure of vomer, tongue, and palatinum, expose additional morphological characters important for clarification of the taxonomy of this group of fishes. Differences are also revealed in the type of dentition, such as on the vomer and epi-hypopharyngeal bones. Strong and dense dentition of the anterior part of the oral cavity and a high number of TB on this site in species feeding on larger prey (e.g., Cheilodipterus spp) is compared to the relatively feeble jaw armor and richness of TB on the more pharyngeal site in species feeding on smaller prey (e.g., Apogon angustatus, A. frenatus). In addition to the three types of TB (Types I-III) previously described from various teleost fish, a fourth type (Type IV), comprising very small buds, was found in some cardinal fish (Apogon angustatus, A. frenatus). The various TB are distributed from the lips to the pharyngeal bones, on the breathing valves, tongue, palatinum, and pharyngeal bones; their number and type on the various sites differ in the different species. In all species studied the Types I and II TB, elevated above the surrounding epithelium, dominated the lips and anterior part of mouth, while Types III and IV, which end apically at the level with the epithelium, dominated the more posterior pharyngeal region. The highest number of TB, around 24,600, were found in Fowleria variegata, a typical nocturnal species, and the lowest in the diurnal and crepuscular Apogon cyanosoma (1,660) and Cheilodipterus quinquestriatus (2,400). Differences are also revealed in the type of dentition, such as on the vomer and epi-hypopharyngeal bones. The number of TB increased with growth of the fishes. The differences in the total number of TB and their distribution in the oropharyngeal cavity in the various species indicates possible different mechanisms of foraging and food-recognition.     

Potential roles of smell and taste in the orientation behaviour of coral-reef fish larvae: insights from morphology

An ontogenetic analysis of the olfactory organ and the number and distribution of internal taste buds was carried out in two neon gobies (Elacatinus lori and Elacatinus colini) with the goal of revealing morphological trends that might inform an understanding of the roles of olfaction and taste in larval orientation behaviour. The pattern of development of the olfactory organ is unremarkable and enclosure of the olfactory epithelium occurs concurrently with metamorphosis and settlement in both species. Like other gobies, juvenile and adult E. lori and E. colini lack complex olfactory lamellae, and lack the accessory nasal sacs present in some adult gobies that could facilitate active olfactory ventilation (i.e., sniffing). A small number of internal taste buds are present at hatch with most found in the caudal region of the buccal cavity (on gill arches, roof of buccal cavity). As taste bud number increases, they demonstrate an anterior spread to the lips, buccal valves and tongue (i.e., tissue covering the basihyal). In the absence of an active ventilatory mechanism for the olfactory organs, the water that moves through the buccal cavity with cyclic gill ventilation may provide chemical cues allowing the internal taste buds to play a role in chemical-mediated orientation and reef-seeking behavior in pelagic larval fishes.     

Comparison of the oropharyngeal cavity in the Starksiini (Teleostei: Blenniiformes: Labrisomidae): Taste buds and teeth,including a comparison with closely‐related genera

The present study describes the distribution of taste buds and teeth in the oropharyngeal cavity of 13 species of adult (18–60 mm SL) Starksiini fishes inhabiting subtidal waters of the Neotropical region. Four types of taste buds described previously in other fish groups were observed within the oropharyngeal cavity, of which type I, situated on prominent protruding papillae, is the most common. The number of taste buds in this cavity varies considerably, ranging from ca. 202 in Starksia lepicoelia to ca. 770 in S. sluiteri. In all the studied species, taste buds are more numerous on the posterior (160–396) than on the anterior (42–294) part of the oropharyngeal cavity. The presence of different numbers of taste buds in different Starksiini species of the same standard length suggests that numbers of taste buds are not directly correlated with size and may be species‐specific. Teeth are found on the premaxilla, dentary, vomer, palatine (in some species) and the upper and lower pharyngeal jaws (third pharyngobranchials and fifth ceratobranchials, respectively); the form and number of teeth and taste buds on each of these sites differs among the various species of Starksiini and between them and closely related species of the labrisomid tribes Labrisomini, Mnierpini, and Paraclinini. The results thus suggest potential systematic value in certain features of the oropharyngeal cavity for blenniiform fishes. It is also shown that benthic‐feeding omnivorous fishes have higher densities of taste buds than piscivorous fishes. A possible correlation among numbers of taste buds, their positions in the oropharyngeal cavity, and other parameters is discussed. J. Morphol., 2012. © 2012 Wiley Periodicals, Inc.     

Evidence of putative sensory receptors from snout and tongue in an upstream amphihaline migratory fish hilsa <Emphasis Type="Italic">Tenualosa ilisha</Emphasis>

Epidermal sensory structures of adults and juveniles of amphihaline migratory fish hilsa Tenualosa ilisha were studied from two habitats, i.e., freshwater (FW) and marine water (MW). Every year, adults and sexually mature hilsa migrate upstream from marine habitat to riverine freshwater habitat for breeding. This report provides evidences of chemoreception on their upstream migration through several characteristic features on their body, especially on the head and oral cavity. Scanning electron microscopy (SEM) reveals that freshwater adult hilsa (FH) has abundant solitary chemosensory cells (SCCs) on the snout epidermis (around the openings of the epidermal pit) and upper lip, whereas marine water adult hilsa (MH) moderately possesses such sensory structures. The juveniles returning to marine water completely lack SCCs. Immunohistochemical studies revealed the expression of PLC β2 on the snout of FH and tongue of both FH and MH. Further analysis (immunofluorescence, immunoblot and densitometry) of the epidermis confirms the presence of chemosensory structures through strong expression and localization of G-proteins (Gαq and Gα s/olf) from the snout as well as tongue in freshwater hilsa. The SEM also confirms the presence of two types of taste buds in FH, viz. type I (TB I) and type III (TB III). Whereas TB I and TB III are observed on the upper palatine and lips, most of the TB III are located on the tongue region of freshwater and marine hilsa. The juvenile hilsa are devoid of such structures. The presence of dense and rich SCCs and taste sensory cells in adults could be a characteristic feature for strong sensory reception to recognize odour and food-related environmental cues from habitats where they often migrate.     

Structural Organization of the Taste Apparatus in Characins (Characidae,Teleostei)

The morphology and distribution of taste buds in the outer integument of the body and in the oral cavity of two forms (blind cave and sighted terrestrial ones) of the astyanax Astyanax fasciatus and in intact and blinded individuals of the Buenos Aires tetra Hyphessobrycon anisitsi have been studied using electronic scanning and light microscopy. In sighted individuals of both species, the morphometric parameters of the taste apparatus and the distribution of taste receptors are similar; the taste apparatus in the oral cavity is more developed than in the outer covers. Morphologically different taste zones were found in the oral cavity of characins. In blind fish, the taste apparatus of the maxillary zones is distinguished by smaller taste buds and a greater density of their distribution. The sensory field of taste buds in blind and sighted individuals of astyanax and tetra has a similar ultrastructure; it is formed by taste cells of three types. In blind astyanaxes and blinded individuals of tetra, numerous modified epidermal cells were found for the first time in the epithelium of the taste zones and in contact with taste buds, which are regarded as tactile receptors and a constituent element of polysensory taste-tactile complexes localized in blind fish in mainly ventral sensory zones.     

Gustatory apparatus of the oropharyngeal cavity in juvenile rainbow trout <Emphasis Type="Italic">Parassalmo mykiss</Emphasis>

Study of the structural organization of gustatory apparatus in rainbow trout Parasalmo mykiss performed using electron scanning microscopy demonstrated that external taste buds are absent in this species in skin covers of the head and in the circumoral region. In the oropharyngeal cavity (oral and gill cavities and pharynx) of the rainbow trout, a well-developed gustatory receptor apparatus was found. In correspondence with specific features of morphology and anatomy of the skull, taste buds form seven gustatory zones. Morphometric analysis demonstrated differences between gustatory zones in the pattern and density of distribution of taste buds, as well as in average sizes of their sensory field. Zones of similar innervation have many common features in morphology. Morphologically similar zones form three regions in the oropharyngeal cavity: rostral, central, and caudal. A tendency for a decrease in the concentration of taste buds in the rostrocaudal direction common for all sensory zones was revealed. The highest concentration of taste buds was recorded at papillae of rostral regions near big teeth. A typical feature of taste buds in rainbow trout is irregular shape of the taste pore. Analysis of ultrastructural specific features of apical processes of taste cells allows us to distinguish five cell shapes in the composition of taste buds. The numeric ratio of cell shapes varies in buds of different localization. The quantitative distribution of taste buds over sensory zones, specific features of morphology and sizes of their sensory field are discussed in relation to the feeding pattern of the species.     

齐口裂腹鱼味蕾及上皮细胞的扫描电镜观察

本文报道扫描电镜对齐口裂腹鱼的味蕾在须、口咽腔、食道的分布和表面形态的观察。结果表明在须、唇、颌、口腔顶壁、咽、舌、食道均有味蕾分布,且在表面形态上呈现多态性。上皮细胞表面有微脊。     

Surface architecture of the oropharyngeal cavity and the digestive tract of Mulloidichthys flavolineatus from the red sea,Egypt: A scanning electron microscope study

Surface architecture of the buccal cavity and the surface organization of the luminal mucosa of the oesophagus, stomach, and intestine of the carnivorous fish M. flavolineatus from the Red Sea were studied by using SEM. The results revealed that M. flavolineatus has four kinds of teeth; curved-blunt, wedge-shaped, flattened crowns, molariform and papilliform. Three types of taste buds (type I, II and III) were recorded in the oropharyngeal cavity. It was observed that taste buds and teeth are co-located in the pharyngeal region. Characteristic patterns of microridges of the surface cells in the oral cavity and oesophagus were observed. Mucous cells are distributed in the lining of the mouth cavity, oesophagus, stomach, and intestine. Characteristic patterns of mucosal folds throughout the alimentary canal, concerning oesophagus, stomach, and intestine were revealed. Numerous gastric pits, which represents the emergence of the gastric glands, were recorded in the anterior and middle regions of the stomach. Complex patterns of the folds and mucous cells were recorded in the intestinal mucosa. These results were discussed with other teleost fishes.     

Analysis of the embryonic lineage of vertebrate taste buds

In all vertebrates, taste buds are the last sensory receptorsto appear late in embryonic development. They are thought toarise locally from the oropharyngeal epithelium, although thishypothesis has not been tested experimentally. Alternatively,taste buds have been proposed to arise from neurocctodermalcells that migrate from peripheral neurogenic sources to theoropharyngeal epithelium and give rise to taste bud precursorcells. In order to determine the exact embryonic lineage ofthe cells of vertebrate taste buds, we have employed a combinationof endogenous and exogenous cell marking techniques to followneuroectodermal and endodermal cells through development. Wefind, in the ambystomatid salamander used in our studies, tastebuds arise locally within the endodermally-derived epitheliumlining the oropharyngeal cavity, and do not receive a contributionfrom neuroectodermal sources, i.e. ectodermal placodes or cephalicneural crest.     

Effects of season, sex and body size on the feeding ecology of turtle-headed sea snakes (Emydocephalus annulatus) on IndoPacific inshore coral reefs

In terrestrial snakes, many cases of intraspecific shifts in dietary habits as a function of predator sex and body size are driven by gape limitation and hence are most common in species that feed on relatively large prey and exhibit a wide body-size range. Our data on sea snakes reveal an alternative mechanism for intraspecific niche partitioning, based on sex-specific seasonal anorexia induced by reproductive activities. Turtle-headed sea snakes (Emydocephalus annulatus) on coral reefs in the New Caledonian Lagoon feed entirely on the eggs of demersal-spawning fishes. DNA sequence data (cytochrome b gene) on eggs that we palpated from stomachs of 37 snakes showed that despite this ontogenetic stage specialization, the prey comes from a taxonomically diverse array of species including damselfish (41 % of samples, at least 5 species), blennies (41 %, 4 species) and gobies (19 %, 5 species). The composition of snake diets shifted seasonally (with damselfish dominating in winter but not summer), presumably reflecting seasonality of fish reproduction. That seasonal shift affects male and female snakes differently, because reproduction is incompatible with foraging. Adult female sea snakes ceased feeding when they became heavily distended with developing embryos in late summer, and males ceased feeding while they were mate searching in winter. The sex divergence in foraging habits may be amplified by sexual size dimorphism; females grow larger than males, and larger snakes (of both sexes) feed more on damselfish (which often lay their eggs in exposed sites) than on blennies and gobies (whose eggs are hidden within narrow crevices). Specific features of reproductive biology of coral reef fish (seasonality and nest type) have generated intraspecific niche partitioning in these sea snakes, by mechanisms different from those that apply to terrestrial snakes.     

Parental defence on the reef: antipredator tactics of coral‐reef fishes against egg‐eating seasnakes

On coral reefs in New Caledonia, the eggs of demersal‐spawning fishes are consumed by turtle‐headed seasnakes (Emydocephalus annulatus). Fish repel nest‐raiding snakes by a series of tactics. We recorded 232 cases (involving 22 fish species) of antipredator behaviour towards snakes on a reef near Noumea. Blennies and gobies focused their attacks on snakes entering their nests, whereas damselfish (Pomacentridae) attacked passing snakes, as well as nest‐raiders (reflecting territorial defence). Biting the snake was the most common form of attack, although damselfish and blennies also slapped snakes with the tail, or (blennies only) plugged the nest entrance with the parent fish's body. Gobies rarely defended the nest, although they sometimes bit or threw sand at the snake. A snake was more likely to flee if it was attacked before it began feeding rather than after it found the eggs (82% versus 3% repelled) and if bitten on the head rather than the body (68% versus 53%). Tail‐slaps were not effective, although plugging the burrow and throwing sand often caused snakes to flee. These strong patterns reflect phylogenetic variation in fish behaviour (e.g. damselfish detect a snake approach sooner than do substrate‐dwelling blennies and gobies) coupled with intraspecific variation in snake diets. © 2014 The Linnean Society of London, Biological Journal of the Linnean Society, 2015, 114 , 415–425.     

SEM study of the oral cavity of members of the Kyphosidae and Girellidae (Pisces,Teleostei), with remarks on Crenidens (Sparidae), focusing on teeth and taste bud numbers and distribution

The present study compares dental morphology and taste bud distribution in the oral cavity of four species of the teleost family Kyphosidae, five species of Girellidae and one species of Sparidae. Some of these species are predominantly herbivorous, while others are omnivorous with a high portion of invertebrates in their diets. All the kyphosids feature one row of frontal cusped teeth on their jaws and areas of miniature teeth on the tongue and palate; the girellid species feature 2–5 rows of denticulate teeth, and no teeth inside the oral cavity. The total number of taste buds in the oral cavity is higher in Girella spp. than in the kyphosids, and their number is species-specific, not correlated with fish size. For example, Kyphosus bigibbus of 500 mm standard length bears 1780 taste buds, while the kyphosid Neoscorpis lithophilus of 80 mm bears 3460. The maximum number of taste buds, 7900, is found in Girella punctata of 60 mm standard length, and the minimum number, 1320, in Kyphosus vaigiensis of 175 mm. The higher number of taste buds appears to characterize species that possess a more diversified diet. The present study contributes to our understanding of the differences between the two families Kyphosidae and Girellidae. In particular, it relates the eco-morphological adaptations to the type of diet consumed by the various species.     

Adaptation of swallowing hyo-laryngeal kinematics is distinct in oral vs. pharyngeal sensory processing

Before a bolus is pushed into the pharynx, oral sensory processing is critical for planning movements of the subsequent pharyngeal swallow, including hyoid bone and laryngeal (hyo-laryngeal) kinematics. However, oral and pharyngeal sensory processing for hyo-laryngeal kinematics is not fully understood. In 11 healthy adults, we examined changes in kinematics with sensory adaptation, sensitivity shifting, with oropharyngeal swallows vs. pharyngeal swallows (no oral processing), and with various bolus volumes and tastes. Only pharyngeal swallows showed sensory adaptation (gradual changes in kinematics with repeated exposure to the same bolus). Conversely, only oropharyngeal swallows distinguished volume differences, whereas pharyngeal swallows did not. No taste effects were observed for either swallow type. The hyo-laryngeal kinematics were very similar between oropharyngeal swallows and pharyngeal swallows with a comparable bolus. Sensitivity shifting (changing sensory threshold for a small bolus when it immediately follows several very large boluses) was not observed in pharyngeal or oropharyngeal swallowing. These findings indicate that once oral sensory processing has set a motor program for a specific kind of bolus (i.e., 5 ml water), hyo-laryngeal movements are already highly standardized and optimized, showing no shifting or adaptation regardless of repeated exposure (sensory adaptation) or previous sensory experiences (sensitivity shifting). Also, the oral cavity is highly specialized for differentiating certain properties of a bolus (volume) that might require a specific motor plan to ensure swallowing safety, whereas the pharyngeal cavity does not make the same distinctions. Pharyngeal sensory processing might not be able to adjust motor plans created by the oral cavity once the swallow has already been triggered.     

Ultrastructure and distribution of the taste buds in the buccal cavity in relation to the food and feeding habit of a herbivorous fish: Oreochromis niloticus

The buccal cavity of an herbivorous fish Oreochromis niloticus was investigated by means of scanning electron microscopy. The buccal cavity distinguished into the roof and the floor. Three different types of taste buds (type I, II and III) are distributed in the buccal cavity. The proximal part of the buccal cavity bears relatively high epidermal papillae in which type I TBs was found. The middle region of the buccal cavity is mostly occupied by low epidermal papillae containing type II TBs. Type III TBs which are present within the metabranchial buccal cavity; never rise above the normal level of the epithelium.The different types of TBs are useful for ensuring full utilization of the gustatory ability of the fish. It is postulated that the TBs serve different functions: TBs type I and II may act as chemoreceptors and mechanoreceptors. TBs type III acts predominantly as a chemoreceptors. TBs of each type show two kinds of receptor villi within their receptor areas: tall villi and small villi. The surface of the lining epithelial cells shows a delicate microridge system. These structures protect against physical abrasion potentially caused during food maneuvering and swallowing. Furthermore, protection of the epithelium from abrasion is enhanced with goblet cells secretion.     

Epithelium of lips and associated structures of the Indian major carp,Catla catla

Structural organization of the epithelium of the lips and associated structures of the Indian major carp,Catla catla, is described. The upper lip is thin and is associated on its dorsal side with a membranous fold of skin and the rostral cap. In contrast, the lower lip is thick and very conspicuous. It is associated on its ventral side with a fold of skin between it and the ventral head skin. The lower lip is divided into a non-projectile portion, a projectile portion and an intermediate groove region. The projectile portion remains folded covering a part of the ventral head skin when the mouth is closed. Their role in relation to the formation of the characteristic feeding tube is discussed. The epithelium of the lips and associated structures is stratified in nature and is composed of the epithelial cells, mucous cells, club cells, lymphocytes and the taste buds. The mucous cells are small, few or even absent and do not appear to secrete profusely at the surfaces of the upper and the lower lips. This suggests that the lips inCatla catla, which feeds on micro-organisms, do not need extra lubrication for protection against abrasion during feeding. In the epithelium at the folds of skin, the voluminous mucous cells secrete profusely and provide extra lubrication to their surface. This reduces the resistance to surface drag during stretching and enables the jaws to protrude with increasing efficiency and swiftness. The club cells are developed additionally to complement the mucous cells in the rostral cap and the upper lip epithelium. Their primary function appears protective in some way, which needs further confirmation. The taste buds, though few in the lower lip, are located in a good number in the upper lip on the characteristic epithelial papillae-like projections, and are projected at the surface. These have been associated with the acute gustatory sense of the fish. The taste buds are absent on the folds of skin where they may not be of much significance.     

Are there efferent synapses in fish taste buds?

In fish, nerve fibers of taste buds are organized within the bud's nerve fiber plexus. It is located between the sensory epithelium consisting of light and dark elongated cells and the basal cells. It comprises the basal parts and processes of light and dark cells that intermingle with nerve fibers, which are the dendritic endings of the taste sensory neurons belonging to the cranial nerves VII, IX or X. Most of the synapses at the plexus are afferent; they have synaptic vesicles on the light (or dark) cells side, which is presynaptic. In contrast, the presumed efferent synapses may be rich in synaptic vesicles on the nerve fibers (presynaptic) side, whereas the cells (postsynaptic) side may contain a subsynaptic cistern; a flat compartment of the smooth endoplasmic reticulum. This structure is regarded as a prerequisite of a typical efferent synapse, as occurring in cochlear and vestibular hair cells. In fish taste buds, efferent synapses are rare and were found only in a few species that belong to different taxa. The significance of efferent synapses in fish taste buds is not well understood, because efferent connections between the gustatory nuclei of the medulla with taste buds are not yet proved.     

Sorting food from stones: the vagal taste system in Goldfish, Carassius auratus

The sense of taste, although a relatively undistinguished sensory modality in most mammals, is a highly developed sense in many fishes, e.g., catfish, gadids, and carps including goldfish. In these species, the amount of neural tissue devoted to this modality may approach 20% of the entire brain mass, reflecting an enormous number of taste buds scattered across the external surface of the animal as well as within the oral cavity. The primary sensory nuclei for taste form a longitudinal column of nuclei along the dorsomedial surface of the medulla. Within this column of gustatory nuclei, the sensory system is represented as a fine-grain somatotopic map, with external body parts being represented rostrally within the column, and oropharyngeal surfaces being represented caudally. Goldfish have a specialization of the oral cavity, the palatal organ, which enables them to sort food particles from particulate substrate material such as gravel. The palatal organ taste information reaches the large, vagal lobe with a complex laminar and columnar organization. This lobe also supports a radially-organized reflex system which activates the musculature of the palatal organ to effect the sorting operation. The stereotyped, laminated structure of this system in goldfish has facilitated studies of the circuitry and neurotransmitter systems underlying the goldfish’s ability to sort food from stones.     

Occurrence of non‐native fishes in the Danube east of Vienna (Austria) and potential interactions of invasive gobiids with native fishes

Non‐native fish species pose a major threat to local fish populations and aquatic ecosystems in general. Invasive gobies are a particular focus of research, but with partly inconsistent results. While some studies reported severe detrimental impacts on native species, others have concluded less serious or neutral effects. We provide results from a large‐scale, multi‐annual fish monitoring program on the occurrence and abundance of non‐native fishes in the main stem of a free‐flowing section of the Austrian Danube. Special emphasis was placed on identifying positive or negative interactions of invasive gobies with native species. Whereas most non‐native species occurred too sporadically or were too few in number to infer a direct threat on the local fish community, invasive gobies were among the most common fishes throughout all sampling years. Co‐occurrence analyses revealed species‐ and mesohabitat type‐specific associations of gobies with native species, which were primarily positive. Notably, native predators such as asp, burbot, or perch probably benefit from the ubiquitous gobies. Two characteristic fluvial fishes revealed negative associations with invasive gobies, namely barbel (Barbus barbus) and Danube whitefin gudgeon (Romanogobio vladykovi): they appear to avoid habitats occupied by gobies. Accordingly, high abundances of round and bighead goby most likely resulted in population losses of barbel and whitefin gudgeon, respectively. Overall, our results indicate a limited negative impact of non‐native species in the sampling area. This is because only two out of 51 occurring species were found to be adversely affected by gobies, the share of co‐occurrences with native species was high, and other non‐native species were generally rare. Nevertheless, invasions are highly dynamic, and new non‐native species are likely to occur in the Austrian Danube, calling for continued monitoring and awareness.