RESUMO
Premaxillary protrusion and the performance advantages it confers are implicated in the success of diverse lineages of teleost fishes, such as Cypriniformes and Acanthomorpha. Although premaxillary protrusion has evolved independently at least five times within bony fishes, much of the functional work investigating this kinesis relates to mechanisms found only in these two clades. Few studies have characterized feeding mechanisms in less-diverse premaxilla-protruding lineages and fewer yet have investigated the distinctive anatomy underlying jaw kinesis in these lineages. Here, we integrated dissection, clearing and staining, histology, micro-CT, and high-speed videography to investigate an isolated and independent origin of jaw protrusion in the hingemouth, Phractolaemus ansorgii, which employs a complex arrangement of bones, musculature, and connective tissues to feed on benthic detritus via a deployable proboscis. Our goals were to provide an integrative account of the underlying architecture of P. ansorgii's feeding apparatus and to assess the functional consequences of this drastic deviation from the more typical teleost condition. Phractolaemus ansorgii's cranial anatomy is distinct from all other fishes in that its adducted lower jaw is caudally oriented, and it possesses a mouth at the terminal end of an elongated, tube-like proboscis that is unique in its lack of skeletal support from the oral jaws. Instead, its mouth is supported primarily by hyaline-cell cartilage and other rigid connective tissues, and features highly flexible lips that are covered in rows of keratinous unculi. Concomitant changes to the adductor musculature likely allow for the flexibility to protrude the mouth dorsally and ventrally as observed during different feeding behaviors, while the intrinsic compliance of the lips allows for more effective scraping of irregular surfaces. From our feeding videos, we find that P. ansorgii is capable of modulating the distance of protrusion, with maximum anterior protrusion exceeding 30% of head length. This represents a previously undescribed example of extreme jaw protrusion on par with many acanthomorph species. Protrusion is much slower in P. ansorgii-reaching an average speed of 2.74 cm/s-compared to acanthomorphs feeding on elusive prey or even benthivorous cypriniforms. However, this reorganization of cranial anatomy may reflect a greater need for dexterity to forage more precisely in multiple directions and on a wide variety of surface textures. Although this highly modified mechanism may have limited versatility over evolutionary timescales, it has persisted in solitude within Gonorynchiformes, representing a novel functional solution for benthic feeding in tropical West African rivers.
Assuntos
Comportamento Alimentar , Arcada Osseodentária , Animais , Arcada Osseodentária/anatomia & histologia , Arcada Osseodentária/fisiologia , Fenômenos Biomecânicos , Comportamento Alimentar/fisiologia , Peixes/anatomia & histologia , Peixes/fisiologia , Microtomografia por Raio-XRESUMO
Epibranchial organs (EBOs), found in at least five of the eight otomorphan families, are used to aggregate small prey inside the buccopharyngeal cavity and range in morphological complexity from a singular, small slit on the pharyngeal roof to several, elongated soft tissue tubes. Despite broad phylogenetic representation, little is known about the origin, development, or evolution of EBOs. We hypothesize that both heterochronic and heterotopic changes throughout the evolution of EBOs are at the root of their morphological diversity. Heterochrony is a foundational explanation in developmental studies, however, heterotopy, a developmental change in spatial or topographical relationships, can have even more profound effects on a given structure but has received relatively little attention. Here, we investigate how developmental mechanisms may drive morphological diversity of EBOs within otomorphan fishes. We compare early pharyngeal development in three species, Anchoa mitchilli (Engraulidae) which has the most basic EBO, B. tyrannus (Clupeidae) which has a more complex EBO, and Hypophthalmichthys molitrix (Cyprinidae) which has the most complex EBO yet described. Using branchial arch growth rates and morphological analyses, we illustrate how both heterochronic and heterotopic mechanisms are responsible for some of the phenotypic diversity seen in otomorphan EBOs. Importantly, we also identify conserved developmental patterns that further our understanding of how EBOs may have first originated and evolved across actinopterygian fishes.
Assuntos
Evolução Biológica , Peixes , Animais , Região Branquial , FilogeniaRESUMO
Some fishes rely on large regions of the dorsal (epaxial) and ventral (hypaxial) body muscles to power suction feeding. Epaxial and hypaxial muscles are known to act as motors, powering rapid mouth expansion by shortening to elevate the neurocranium and retract the pectoral girdle, respectively. However, some species, like catfishes, use little cranial elevation. Are these fishes instead using the epaxial muscles to forcefully anchor the head, and if so, are they limited to lower-power strikes? We used X-ray imaging to measure epaxial and hypaxial length dynamics (fluoromicrometry) and associated skeletal motions (XROMM) during 24 suction feeding strikes from three channel catfish (Ictalurus punctatus). We also estimated the power required for suction feeding from oral pressure and dynamic endocast volume measurements. Cranial elevation relative to the body was small (<5 deg) and the epaxial muscles did not shorten during peak expansion power. In contrast, the hypaxial muscles consistently shortened by 4-8% to rotate the pectoral girdle 6-11 deg relative to the body. Despite only the hypaxial muscles generating power, catfish strikes were similar in power to those of other species, such as largemouth bass (Micropterus salmoides), that use epaxial and hypaxial muscles to power mouth expansion. These results show that the epaxial muscles are not used as motors in catfish, but suggest they position and stabilize the cranium while the hypaxial muscles power mouth expansion ventrally. Thus, axial muscles can serve fundamentally different mechanical roles in generating and controlling cranial motion during suction feeding in fishes.
Assuntos
Bass , Músculo Esquelético , Animais , Fenômenos Biomecânicos , Comportamento Alimentar , SucçãoRESUMO
Silver carp regularly consume and digest particles of food as small as 5 µm. This ability drives their efficient consumption of phytoplankton and because they feed low on the food chain they have an important place in aquaculture worldwide. In North America, where they are considered invasive, silver carp deplete food resources for native species and in so doing occupy increased niche space. Here, we determine the ontogenetic stage and size at which silver carp are morphologically capable of primarily feeding on particles <10 µm. Ecological studies on this species have shown that there is an ontogenetic shift in diet as predominantly zooplanktivorous juveniles later switch to eating much smaller phytoplankton. The occupation of this new trophic niche presents both a metabolic and a mechanical challenge to these fish, since it is unclear how they can efficiently feed on such small particles. We hypothesize that the epibranchial organ (EBO) in silver carp is essential in aggregating these small particles of food, allowing the species to consume mass quantities of tiny particles, thus mitigating metabolic constraints. In this study, we investigate early ontogeny of the EBO in silver carp to determine when this structure achieves the requisite morphology to become functional. We find that at around 80 mm standard length (SL) the EBOs are consistently filled with food, demonstrating that this accumulating organ has become functional. This size corresponds with previous ecological data documenting important shifts in the type of food consumed. While the basic bauplan of the EBO is established very early in ontogeny (by 15 mm SL), multiple waves of histological maturation of muscle, cartilage, gill rakers and epithelium ultimately form the functional structure.
Assuntos
Carpas/crescimento & desenvolvimento , Brânquias/crescimento & desenvolvimento , Animais , Carpas/anatomia & histologia , Comportamento Alimentar/fisiologia , Brânquias/anatomia & histologia , FitoplânctonRESUMO
Tooth replacement in piranhas is unusual: all teeth on one side of the head are lost as a unit, then replaced simultaneously. We used histology and microCT to examine tooth-replacement modes across carnivorous piranhas and their herbivorous pacu cousins (Serrasalmidae) and then mapped replacement patterns onto a molecular phylogeny. Pacu teeth develop and are replaced in a manner like piranhas. For serrasalmids, unilateral tooth replacement is not an "all or nothing" phenomenon; we demonstrate that both sides of the jaws have developing tooth rows within them, albeit with one side more mineralized than the other. All serrasalmids (except one) share unilateral tooth replacement, so this is not an adaptation for carnivory. All serrasalmids have interlocking teeth; piranhas interdigitate lateral tooth cusps with adjacent teeth, forming a singular saw-like blade, whereas lateral cusps in pacus clasp together. For serrasalmids to have an interlocking dentition, their teeth need to develop and erupt at the same time. We propose that interlocking mechanisms prevent tooth loss and ensure continued functionality of the feeding apparatus. Serrasalmid dentitions are ubiquitously heterodont, having incisiform and molariform dentitions reminiscent of mammals. Finally, we propose that simultaneous tooth replacement be considered as a synapomorphy for the family.
Assuntos
Evolução Biológica , Caraciformes/crescimento & desenvolvimento , Dentição , Odontogênese , Animais , FilogeniaRESUMO
When animals move they must coordinate motion among multiple parts of the musculoskeletal system. Different behaviours exhibit different patterns of coordination, however, it remains unclear what general principles determine the coordination pattern for a particular behaviour. One hypothesis is that speed determines coordination patterns as a result of differences in voluntary versus involuntary control. An alternative hypothesis is that the nature of the behavioural task determines patterns of coordination. Suction-feeding fishes have highly kinetic skulls and must coordinate the motions of over a dozen skeletal elements to draw fluid and prey into the mouth. We used a dataset of intracranial motions at five cranial joints in channel catfish ( Ictalurus punctatus), collected using X-ray reconstruction of moving morphology, to test whether speed or task best explained patterns of coordination. We found that motions were significantly more coordinated (by 20-29%) during prey capture than during prey transport, supporting the hypothesis that the nature of the task determines coordination patterns. We found no significant difference in coordination between low- and high-speed motions. We speculate that capture is more coordinated to create a single fluid flow into the mouth while transport is less coordinated so that the cranial elements can independently generate multiple flows to reposition prey. Our results demonstrate the benefits of both higher and lower coordination in animal behaviours and the potential of motion analysis to elucidate motor tasks.
Assuntos
Deglutição , Ictaluridae/fisiologia , Comportamento Predatório , Animais , Fenômenos Biomecânicos , Crânio/anatomia & histologiaRESUMO
Invasive silver carp are thriving within eutrophic environments in the United States, in part because of their highly efficient filter-feeding mechanism. Silver carp utilize modified gill rakers to capture a specific range of food; however, their greatly modified filtering morphology allows them to feed on phytoplankton and zooplankton ranging in size from 4 to 85â µm. The filtering apparatus of silver carp comprises rigid filtering plates where the outer anatomy of these plates is characterized by long parallel channels that change in orientation along the length of the plate. Here, we investigate the underlying morphology and concomitant hydrodynamics that support the filtration mechanisms of silver and bighead carp. Bighead carp are also invasive filter feeders, but their filtering apparatus is morphologically distinct from that of silver carp. Using 3D particle image velocimetry, we determined how particles and fluid interact with the surface of the gill rakers/plates. Filtering plates in silver carp induce strong directed vortical flow, whereas the filtering apparatus of bighead carp resulted in a type of haphazard cross-flow filtration. The organized vortical flow established by silver carp likely increased the number of interactions that the particle-filled water had with the filtering membrane. This strong vortical organization is maintained only at 0.75â bodyâ lengths s-1, and vortical flow is poorly developed and maintained at slower and faster speeds. Moreover, we found that absolute vorticity magnitude in silver carp is an order of magnitude greater than in bighead carp.
Assuntos
Carpas/anatomia & histologia , Carpas/fisiologia , Comportamento Alimentar , Hidrodinâmica , Animais , Modelos Biológicos , Boca/diagnóstico por imagem , Reologia , Movimentos da Água , Microtomografia por Raio-X/veterináriaRESUMO
To expand the buccal cavity, many suction-feeding fishes rely on a considerable contribution from dorsal rotation of the dorsal part of the head including the brains, eyes, and several bones forming the braincase and skull roof (jointly referred to as the neurocranium). As the neurocranium takes up a large part of the total mass of the head, this rotation may incur a considerable inertial cost. If so, this would suggest a significant selective pressure on the kinematics and mass distribution of the neurocranium of suction feeders. Here, an inverse dynamic model is formulated to calculate the instantaneous power required to rotate the neurocranium, approximated by a quarter ellipsoid volume of homogeneous density, as well as to calculate the instantaneous suction power based on intra-oral pressure and head volume quantifications. We applied this model to largemouth bass (Micropterus salmoides) and found that the power required to rotate the neurocranium accounts for only about 4% of the power required to suck water into the mouth. Furthermore, recovery of kinetic energy from the rotating neurocranium converted into suction work may be possible during the phase of neurocranial deceleration. Thus, we suggest that only a negligible proportion of the power output of the feeding muscles is lost as inertial costs in the largemouth bass. Consequently, the feeding performance of piscivorous suction feeders with generalised morphology, comparable to our model species, is not limited by neurocranial motion during head expansion. This suggests that it is thus not likely to be a factor of importance in the evolution of cranial shape and size.
Assuntos
Encéfalo/anatomia & histologia , Encéfalo/fisiologia , Comportamento Alimentar/fisiologia , Crânio/anatomia & histologia , Crânio/fisiologia , Animais , Bass , Fenômenos Biomecânicos , Hidrodinâmica , Modelos Biológicos , Boca/fisiologia , Movimento , Comportamento Predatório/fisiologia , Pressão , Rotação , SucçãoRESUMO
A protrusible upper jaw has independently evolved multiple times within teleosts and has been implicated in the success of two groups in particular: Acanthomorpha and Cypriniformes. We use digital particle image velocimetry (DPIV) to compare suction feeding flow dynamics in a representative of each of these clades: goldfish and bluegill. Using DPIV, we contrast the spatial pattern of flow, the temporal relationship between flow and head kinematics, and the contribution of jaw protrusion to the forces exerted on prey. As expected, the spatial patterns of flow were similar in the two species. However, goldfish were slower to reach maximal kinematic excursions, and were more flexible in the relative timing of jaw protrusion, other jaw movements and suction flows. Goldfish were also able to sustain flow speeds for a prolonged period of time as compared with bluegill, in part because goldfish generate lower peak flow speeds. In both species, jaw protrusion increased the force exerted on the prey. However, slower jaw protrusion in goldfish resulted in less augmentation of suction forces. This difference in force exerted on prey corresponds with differences in trophic niches and feeding behavior of the two species. The bluegill uses powerful suction to capture insect larvae whereas the goldfish uses winnowing to sort through detritus and sediment. The kinethmoid of goldfish may permit jaw protrusion that is independent of lower jaw movement, which could explain the ability of goldfish to decouple suction flows (due to buccal expansion) from upper jaw protrusion. Nevertheless, our results show that jaw protrusion allows both species to augment the force exerted on prey, suggesting that this is a fundamental benefit of jaw protrusion to suction feeders.
Assuntos
Cipriniformes/fisiologia , Arcada Osseodentária/fisiologia , Animais , Evolução Biológica , Fenômenos Biomecânicos , Comportamento Alimentar/fisiologia , Carpa Dourada/fisiologia , Hidrodinâmica , Modelos Anatômicos , Modelos Biológicos , Modelos Estatísticos , Perciformes/fisiologia , Filogenia , Comportamento Predatório/fisiologia , Prognatismo , Especificidade da Espécie , Estresse Mecânico , Fatores de TempoRESUMO
Premaxillary protrusion in cypriniform fishes involves rotation of the kinethmoid, an unpaired skeletal element in the dorsal midline of the rostrum. No muscles insert directly onto the kinethmoid, so its rotation must be caused by the movement of other bones. In turn, the kinethmoid is thought to push on the ascending processes of the premaxillae, effecting protrusion. To determine the causes and effects of kinethmoid motion, we used XROMM (x-ray reconstruction of moving morphology) to measure the kinematics of cranial bones in common carp, Cyprinus carpio. Mean kinethmoid rotation was 83 deg during premaxillary protrusion (18 events in 3 individuals). The kinethmoid rotates in a coordinated way with ventral translation of the maxillary bridge, and this ventral translation is likely driven primarily by the A1ß muscle. Analyses of flexibility (variability between behaviors) and coordination (correlation between bones within a behavior) indicate that motion of the maxillary bridge, not the lower jaw, drives premaxillary protrusion. Thus, upper jaw protrusion is decoupled from lower jaw depression, allowing for two separate modes of protrusion, open mouth and closed mouth. These behaviors serve different functions: to procure food and to sort food, respectively. Variation in starting posture of the maxilla alone dictates which type of protrusion is performed; downstream motions are invariant. For closed mouth protrusion, a ventrally displaced maxillary starting posture causes kinethmoid rotation to produce more ventrally directed premaxillary protrusion. This flexibility, bestowed by the kinethmoid-maxillary bridge-A1ß mechanism, one of several evolutionary novelties in the cypriniform feeding mechanism, may have contributed to the impressive trophic diversity that characterizes this speciose lineage.
Assuntos
Carpas/anatomia & histologia , Carpas/fisiologia , Arcada Osseodentária/anatomia & histologia , Arcada Osseodentária/fisiologia , Animais , Evolução Biológica , Fenômenos Biomecânicos , Comportamento Alimentar , Movimento , PosturaRESUMO
Physical principles and laws determine the set of possible organismal phenotypes. Constraints arising from development, the environment, and evolutionary history then yield workable, integrated phenotypes. We propose a theoretical and practical framework that considers the role of changing environments. This 'ecomechanical approach' integrates functional organismal traits with the ecological variables. This approach informs our ability to predict species shifts in survival and distribution and provides critical insights into phenotypic diversity. We outline how to use the ecomechanical paradigm using drag-induced bending in trees as an example. Our approach can be incorporated into existing research and help build interdisciplinary bridges. Finally, we identify key factors needed for mass data collection, analysis, and the dissemination of models relevant to this framework.
Assuntos
Evolução Biológica , Ecossistema , Fenótipo , ÁrvoresRESUMO
Remoras are fishes that piggyback onto larger marine fauna via an adhesive disc to increase locomotor efficiency, likelihood of finding mates and access to prey. Attaching rapidly to a large, fast-moving host is no easy task, and while research to date has focused on how the disc supports adhesion, no attention has been paid to how or if remoras are able to sense attachment. We identified push-rod-like mechanoreceptor complexes embedded in the soft lip of the remora adhesive disc that are known in other organisms to respond to touch and shear forces. This is, to our knowledge, the first time such mechanoreceptor complexes are described in fishes as they were only known previously in monotremes. The presence of push-rod-like mechanoreceptor complexes suggests not only that fishes may be able to sense their environment in ways not heretofore described but that specialized tactile mechanoreceptor complexes may be a more basal vertebrate feature than previously thought.
RESUMO
The Weberian apparatus, a complex assemblage of greatly modified vertebral elements, significantly enhances hearing within Otophysi. Ultimately we are interested in investigating the genetic mechanisms responsible for the origin, development and morphological diversification of these vertebral elements in the Weberian apparatus of otophysan fishes. However, a necessary first step involves identifying changes in growth of this region as compared with the vertebrae from which these modified elements purportedly derive. Using an ontogenetic series of the zebrafish, Danio rerio, we collected growth data for specific elements within the Weberian apparatus, including neural arches, ribs, and parapophyses. These data are compared to both serially homologous structures in posterior thoracic vertebrae (which act as internal controls) and vertebral elements from the same axial levels in three other non-otophysan teleosts. Significant differences in growth rate were found among serially homologous structures, as well as at equivalent axial levels in different species. Uniform changes in growth rates (in which all structures derived from a specific somite were equally affected) were not found, suggesting precise targeting of morphological change to specific structures. The variation in growth of anterior vertebrae in and among species was greater than expected. This variation in growth rates created developmental patterns unique to each species. Such patterns of growth may help illuminate the specific heterochronic mechanisms required for the origin and subsequent morphological diversification of the Weberian apparatus. This morphological diversity is exemplified by the multitude of forms seen in the cypriniform Weberian apparatus. Understanding patterns of growth in discrete elements of the Weberian apparatus allows us to hypothesize as to the specific developmental changes, likely constituting differences in gene expression in pathways involved in bone and cartilage differentiation, responsible for this morphological diversity.
Assuntos
Evolução Biológica , Desenvolvimento Ósseo/genética , Audição/fisiologia , Coluna Vertebral/crescimento & desenvolvimento , Peixe-Zebra/crescimento & desenvolvimento , Peixe-Zebra/genética , Animais , Regulação da Expressão Gênica no Desenvolvimento , Audição/genética , Especificidade da Espécie , Peixe-Zebra/anatomia & histologiaRESUMO
Upper jaw protrusion is hypothesized to improve feeding performance in teleost fishes by enhancing suction production and stealth of the feeding event. However, many cyprinodontiform fishes (mid-water feeders, such as mosquitofish, killifish, swordtails, mollies and pupfish) use upper jaw protrusion for "picking" prey out of the water column or off the substrate; this feeding mode may require improved jaw dexterity, but does not necessarily require increased stealth and/or suction production. We describe functional aspects of the bones, muscles and ligaments of the anterior jaws in three cyprinodontiform genera: Fundulus (Fundulidae), Gambusia and Poecilia (Poeciliidae). All three genera possess a premaxillomandibular ligament that connects the premaxilla of the upper jaw to the mandible. The architecture of this ligament is markedly different from the upper-lower jaw connections previously described for basal atherinomorphs or other teleosts, and this loose ligamentous connection allows for more pronounced premaxillary protrusion in this group relative to closely related outgroup taxa. Within poeciliids, a novel insertion of the second division of the adductor mandibulae (A2) onto the premaxilla has also evolved, which allows this jaw adductor to actively retract the premaxilla during mouth closing. This movement is in contrast with most other teleosts, where the upper jaw is retracted passively via pressure applied by the adduction of the lower jaw. We postulate that this mechanism of premaxillary protrusion mediates the cyprinodontiforms' ability to selectively pick specific food items from the water column, surface or bottom, as a picking-based feeding mechanism requires controlled and coordinated "forceps-like" movements of the upper and lower jaws. This mechanism is further refined in some poeciliids, where direct muscular control of the premaxillae may facilitate picking and/or scraping material from the substrate.
Assuntos
Ciprinodontiformes/anatomia & histologia , Comportamento Alimentar/fisiologia , Maxila/anatomia & histologia , Animais , Ciprinodontiformes/classificação , Ligamentos/anatomia & histologia , Sistema Musculoesquelético/anatomia & histologia , FilogeniaRESUMO
Premaxillary protrusion is hypothesized to confer a number of feeding advantages to teleost fishes; however, most proposed advantages relate to enhanced stealth or suction production during prey capture. Cyprinodontiformes exhibit an unusual form of premaxillary protrusion where the descending process of the premaxilla does not rotate anteriorly to occlude the sides of the open mouth during prey capture. Instead, the premaxilla is protruded such that it gives the impression of a beak during prey capture. We quantified premaxillary kinematics during feeding in four cyprinodontiform taxa and compared them with three percomorph taxa to identify any performance consequences of this protrusion mechanism. Individual prey capture events were recorded using digital high-speed video at 250-500 frames per second (n >or= 4 individuals, >or= 4 strikes per individual). Species differed in the timing of movement and the maximum displacement of the premaxilla during the gape cycle and in the contribution of the premaxilla to jaw closing. Cyprinodontiform taxa produced less premaxillary protrusion than the percomorph taxa, and were consistently slower in the time to maximum gape. Further, it appears cyprinodontiforms can alter the contribution of the premaxilla to mouth closure on an event-specific basis. We were able to demonstrate that, within at least one species, this variability is associated with the location of the prey (bottom vs. water column). Cyprinodontiform upper jaw movements likely reflect increased dexterity associated with a foraging ecology where prey items are "picked" from a variety of locations: the bottom, water column, or surface. We postulate that dexterity requires slow, precisely controlled jaw movements; thus, may be traded off for some aspects of suction-feeding performance, such as protrusion distance and speed.
Assuntos
Ciprinodontiformes/anatomia & histologia , Ciprinodontiformes/fisiologia , Comportamento Alimentar/fisiologia , Maxila/anatomia & histologia , Maxila/fisiologia , Comportamento Predatório/fisiologia , Animais , Ciprinodontiformes/classificação , FilogeniaRESUMO
Hypophthalmichthys molitrix, silver carp, is an invasive Asian carp that has become increasingly widespread and ecologically destructive within the upper Mississippi River Basin. Its complex trophic anatomy may help explain the apparent efficiency with which they consume phytoplankton, outcompeting native filter feeders. This cypriniform species is characterized by trophic synapomorphies that include a palatal organ, loss of upper pharyngeal jaws, and a hypertrophied lower pharyngeal jaw. However, in silver carp these structures have become greatly modified and diverge from the more basal condition that characterizes species such as goldfish. The trophic apparatus of silver carp is composed of discrete structures that are functionally coupled: filtering plates, paired epibranchial organs (EBO), a modified palatal organ composed of large muscular folds that interdigitate with the filtering plates, and hypertrophied lower pharyngeal jaws and teeth. The filtering plates fill a significant portion of the buccal cavity, especially since the distal parts of these filtering plates make up a key component of the EBOs. EBOs, food aggregating structures found in many teleosts, are thought to have independently evolved at least six times. Ranging in complexity from small slits on the dorsal wall of the pharyngeal cavity to exceedingly intricate spiraling structures, EBOs are morphologically diverse among filter-feeding fishes. Despite this morphological diversity and broad taxonomic distribution, little is known regarding the functional anatomy of the EBO. Moreover, the EBO in silver carp is distinct from the organs previously described in other species, being created by four independent pharyngeal involutions (instead of the more typical one or two) that form spiral-shaped pharyngeal tubes surrounded by circumferential muscle. On each side of the head greatly hypertrophied hyomandibulae and opercles are connected to the anterior cartilaginous caps of the bilateral EBOs via enlarged muscles. Given that these fish are pump filter feeders we hypothesize that the opercula may compress and expand the EBOs during pumping causing food to be moved posteriorly toward the pharyngeal jaws.
Assuntos
Estruturas Animais/anatomia & histologia , Carpas/anatomia & histologia , Estruturas Animais/citologia , Estruturas Animais/ultraestrutura , Animais , Brânquias/anatomia & histologia , Músculos/anatomia & histologia , Músculos/diagnóstico por imagem , Faringe/anatomia & histologia , Microtomografia por Raio-XRESUMO
Filter feeding fishes possess several morphological adaptations necessary to capture and concentrate small particulate matter from the water column. Filter feeding teleosts typically employ elongated and tightly packed gill rakers with secondary bony or epithelial modifications that increase filtering efficiency. The gill rakers of Hypophthalmichthys molitrix, silver carp, are anatomically distinct from and more complex than the filtering apparatus of other teleostean fishes. The silver carp filtering apparatus is composed of biserial, fused filtering plates used to capture particles ranging in size from 4 to 80 µm. Early in ontogeny, at 15-25 mm standard length (SL), silver carp gill rakers are reminiscent of other more stereotypical teleostean rakers, characterized by individual lanceolate rakers that are tightly packed along the entirety of the branchial arches. At 30 mm SL, secondary epithelial projections and concomitant dermal ossification begin to stitch together individual gill rakers. During later juvenile stages, dermal bone further modifies the individual gill rakers and creates a bony scaffold that supports the now fully fused and porous epithelium. By adulthood, the stitching of bone and complete fusion of the overlying epithelium creates rigid filtering plates with morphologically distinct faces. The inner face of the plates is organized into a net-like matrix while the outer face has a sponge-like appearance comprised of differently sized pores. Here, we present morphological data from an ontogenetic series of the filtering apparatus within silver carp. These data inform hypotheses regarding both how these gill raker plates may have evolved from a more basal condition, as well as how this novel architecture allows this species to feed on exceedingly small phytoplankton, particles that represent a greater filtering challenge to the typical anatomy of the gill rakers of fishes.
Assuntos
Carpas/anatomia & histologia , Comportamento Alimentar , Brânquias/anatomia & histologia , Brânquias/fisiologia , Animais , Carpas/crescimento & desenvolvimento , Brânquias/crescimento & desenvolvimento , Brânquias/ultraestrutura , OsteogêneseRESUMO
Cypriniformes (which includes the minnows, carps, loaches, algae-eaters, stone loaches, and suckers) is a morphologically diverse and incredibly speciose order of teleosts. It has been suggested that a number of evolutionary innovations, key to improved hearing and feeding, have played an important role in cypriniform fishes' success. One such innovation, the Weberian apparatus, is a novel assemblage of vertebral elements and modified ribs that relay and amplify sound pressure changes from the gas bladder to the inner ear. The Weberian apparatus unites Cypriniformes with other major orders into an extremely species-rich group of fishes, the Otophysi. Together, otophysan fishes comprise one of the largest groups of fishes in the world, as well as the majority of freshwater fishes. Here we present a detailed comparison of the Weberian apparatus in a number of cypriniform families using cleared and stained specimens. We present data regarding inter- and intrafamilial morphological variation within Cypriniformes. With few, but evolutionarily important, exceptions we find that diagnostic features of the Weberian apparatus characterize each family. Interspecific variation within each of the families Balitoridae, Gyrinocheilidae, and Catostomidae is only slight, whereas variation among subfamilies within Cyprinidae and Cobitidae is far more significant. This comparative study identifies a number of distinct morphologies, some of which appear highly correlated with ecological niche. For example, inhabiting swift-moving waters appears to be a key factor in the encapsulation of the anterior gas bladder in some cobitids, balitorids, and gobionin cyprinids.
Assuntos
Sacos Aéreos/anatomia & histologia , Cipriniformes/anatomia & histologia , Variação Genética , Coluna Vertebral/anatomia & histologia , Animais , Cipriniformes/classificação , Orelha/anatomia & histologia , Audição , FilogeniaRESUMO
Cranial skeletal muscles underlie breathing, eating, and eye movements. In most animals, at least two types of muscle fibers underlie these critical functions: fast and slow muscle fibers. We describe here the anatomical distribution of slow and fast twitch muscle in the zebrafish (Danio rerio) head in the adult and at an early larval stage just after feeding has commenced. We found that all but one of the cranial muscles examined contain both slow and fast muscle fibers, but the relative proportion of slow muscle in each varies considerably. As in the trunk, slow muscle fibers are found only in an anatomically restricted zone of each muscle, usually on the periphery. The relative proportion of slow and fast muscle in each cranial muscle changes markedly with development, with a pronounced decrease in the proportion of slow muscle with ontogeny. We discuss our results in relation to the functional roles of each muscle in larval and adult life and compare findings among a variety of vertebrates.
Assuntos
Músculo Esquelético/embriologia , Peixe-Zebra/embriologia , Animais , Diferenciação Celular , Embrião não Mamífero , Larva , Fibras Musculares de Contração Rápida , Fibras Musculares de Contração LentaRESUMO
While much of the functional work on suction feeding has involved members of Acanthopterygii, an earlier cypriniform radiation led to over 3200 species filling nearly every freshwater trophic niche. Within the great majority of acanthomorph clades that have been investigated suction feeding and the underlying morphology responsible for the generation of rapid suction have been largely conserved. This conserved feeding-apparatus is often associated with increasing the force experienced by the prey item, thus making a strike on elusive prey more effective. Cypriniforms' trophic anatomy is comprised of a number of novelties used for benthic feeding, which characterized early members of this clade. The modified cypriniform structure of the oral jaws represents a situation in which a particular type of suction feeding allowed for probing the benthos with a more functionally maneuverable anatomy. Requisite evolutionary modifications included origin and elongation of a median kinethmoid, duplications of certain divisions of the muscles of the adductor mandibulae, and origin of a dorsal, intra-buccal muscular palatal organ used in winnowing detritus. The elongated kinethmoid (coupled with modified adductor muscles) allowed for a type of premaxillary protrusion that decoupled the upper and lower jaws, enabled premaxillary protrusions with a closed mouth, and facilitated benthic feeding by increasing functional flexibility. The resultant flow of fluid generated by cypriniforms is also quite flexible, with multiple instances of peak flow in a single feeding event. This greatly modified morphology allowed for a degree of kinematic maneuverability not seen within most acanthomorphs. Later cypriniform radiations into piscivorous, insectivorous, or planktivorous feeding guilds were associated with shortening of the kinethmoid and with simplified morphology of the adductor, likely involving an emphasis on ram feeding. Although this suite of morphological novelties seemingly originated within the context of benthic feeding, with minimal modifications these anatomical features were later coopted during radiations into different functional niches.