Pectoral fin size in a fish species with paternal care: a condition-dependent sexual trait revealing infection status

1. The three-spined stickleback, Gasterosteus aculeatus L., is a territorial fish with exclusive male parental care. Males oxygenate the eggs with fanning movements of their pectoral fins. The present authors investigated whether the apparent sexual differences in the functional demands of the pectoral fins have resulted in sexual differences in fin size. If males have relatively larger pectoral fins, females may use this as a signal to aid their mate choice for good fathers. Therefore, further objectives were to study the condition-dependency of relative pectoral fin size in males and the relationship with male parasite load. 2. Reproductively active males possessed relatively larger pectoral fins than females in both wild-caught and laboratory-bred fish. 3. In the field, caring males with relatively large pectoral fins were in better physical condition and had more food in their stomachs. 4. Relatively small pectoral fins and poor body condition were associated with infection by the intestinal parasite Pomphorhynchus laevis (Acanthocephala), the prevalent parasite species in the study population.     

Functional Morphology of Aquatic Flight in Fishes: Kinematics, Electromyography, and Mechanical Modeling of Labriform Locomotion

Labriform locomotion is the primary swimming mode for many fishesthat use the pectoral fins to generate thrust across a broadrange of speeds. A review of the literature on hydrodynamics,kinematics, and morphology of pectoral fin mechanisms in fishesreveals that we lack several kinds of morphological and kinematicdata that are critical for understanding thrust generation inthis mode, particularly at higher velocities. Several needsinclude detailed three-dimensional kinematic data on speciesthat are pectoral fin swimmers across a broad range of speeds,data on the motor patterns of pectoral fin muscles, and thedevelopment of a mechanical model of pectoral fin functionalmorphology. New data are presented here on pectoral fin locomotionin Gomphosus varius, a labrid fish that uses the pectoral finsat speeds of 1 –6 total body lengths per second. Three-dimensionalkinematic data for the pectoral fins of G. varius show thata typical "drag-based" mechanism is not used in this species.Instead, the thrust mechanics of this fish are dominated bylift forces and acceleration reaction forces. The fin is twistedlike a propeller during the fin stroke, so that angles of attackare variable along the fin length. Electromyographic data onsix fin muscles indicate the sequence of muscle activity thatproduces antagonistic fin abduction and adduction and controlsthe leading edge of the fin. EMG activity in abductors and adductorsis synchronous with the start of abduction and adduction, respectively,so that muscle mechanics actuate the fin with positive work.A mechanical model of the pectoral fin is proposed in whichfin morphometrics and computer simulations allow predictionsof fin kinematics in three dimensions. The transmission of forceand motion to the leading edge of the fin depends on the mechanicaladvantage of fin ray levers. An integrative program of researchis suggested that will synthesize data on morphology, physiology,kinematics, and hydrodynamics to understand the mechanics ofpectoral fin swimming.     

A Three-Dimensional Kinematics Analysis of a Koi Carp Pectoral Fin by Digital Image Processing

Pectoral fins fascinate researchers for their important role in fish maneuvers. By possessing a complicated flexible structure with several fin rays made by a thin film, the fin exhibits a three-dimensional (3D) motion. The complex 3D fin kinematics makes it challenging to study the performance of pectoral fin. Nevertheless, a detailed study on the 3D motion pattern of pectoral fins is necessary to the design and control of a bio-inspired fin rays. Therefore, a highspeed photography system is introduced in this paper to study the 3D motion of a Koi Carp by analyzing the two views of its pectoral fin simultaneously. The key motions of the pectoral fins are first captured in both hovering and retreating. Next, the 3D configuration of the pectoral fins is reconstructed by digital image processing, in which the movement of fin rays during fish retreating and hovering is obtained. Furthermore, the method of Singular Value Decomposition (SVD) is adopted to extract the basic motion patterns of pectoral fins from extensive image sequences, i.e. expansion, bending, cupping, and undulation. It is believed that the movement of the fin rays and the basic patterns of the pectoral fins obtained in the present work can provide a good foundation for the development and control of bionic flexible pectoral fins for underwater propeller.     

Does body and fin form affect the maneuverability of fish traversing vertical and horizontal slits?

Synopsis The purpose of this study was to determine if body and fin form affected the maneuverability of teleostean fishes as measured by their ability to negotiate simple obstacles. Obstacles were vertical and horizontal rectangular slits of different widths, for which width was defined as the minimum dimension of a slit irrespective of slit orientation. Performance was measured as the smallest slit width traversed. Three species with different body and fin patterns were induced to swim through slits. Species tested were; goldfish Carassius auratus with a fusiform body, anterio-ventral pectoral fins and posterio-ventral pelvic fins; silver dollars Metynnis hypsauchen with the same fin configurations but a gibbose body; angelfish Pterophyllum scalare with a gibbose body and anterio-lateral pectoral fins. Minimum slit widths negotiated were normalized with the length of various body dimensions: total length, maximum width, span at the pectoral fins, and volume^1/3 (numerically equal to mass^1/3). Goldfish had the poorest performance, requiring the largest slit widths relative to these body dimensions. No consistent patterns in performance were found for silver dollars vs. angelfish. There were no differences among species in the ratio of minimum vertical slit width negotiated to that for horizontal slits, indicating fish were equally able to control posture while swimming on their sides. There were also no consistent patterns in the times taken to transit slits. Although the deep-bodied fish were able to maneuver through smaller slits, the most striking result is the similarity of minimum slit widths traversed in spite of the large variation in body form. Body form and fin plan may be more important for maneuvering and posture control during sub-maximum routine activities.     

Morphological and histochemical characterization of the pectoral fin muscle of batoids

Batoids differ from other elasmobranch fishes in that they possess dorsoventrally flattened bodies with enlarged muscled pectoral fins. Most batoids also swim using either of two modes of locomotion: undulation or oscillation of the pectoral fins. In other elasmobranchs (e.g., sharks), the main locomotory muscle is located in the axial myotome; in contrast, the main locomotory muscle in batoids is found in the enlarged pectoral fins. The pectoral fin muscles of sharks have a simple structure, confined to the base of the fin; however, little to no data are available on the more complex musculature within the pectoral fins of batoids. Understanding the types of fibers and their arrangement within the pectoral fins may elucidate how batoid fishes are able to utilize such unique swimming modes. In the present study, histochemical methods including succinate dehydrogenase (SDH) and immunofluoresence were used to determine the different fiber types comprising these muscles in three batoid species: Atlantic stingray (Dasyatis sabina), ocellate river stingray (Potamotrygon motoro) and cownose ray (Rhinoptera bonasus). All three species had muscles comprised of two muscle fiber types (slow-red and fast-white). The undulatory species, D. sabina and P. motoro, had a larger proportion of fast-white muscle fibers compared to the oscillatory species, R. bonasus. The muscle fiber sizes were similar between each species, though generally smaller compared to the axial musculature in other elasmobranch fishes. These results suggest that batoid locomotion can be distinguished using muscle fiber type proportions. Undulatory species are more benthic with fast-white fibers allowing them to contract their muscles quickly, as a possible means of escape from potential predators. Oscillatory species are pelagic and are known to migrate long distances with muscles using slow-red fibers to aid in sustained swimming.     

miR-196 regulates axial patterning and pectoral appendage initiation

Vertebrate Hox clusters contain protein-coding genes that regulate body axis development and microRNA (miRNA) genes whose functions are not yet well understood. We overexpressed the Hox cluster microRNA miR-196 in zebrafish embryos and found four specific, viable phenotypes: failure of pectoral fin bud initiation, deletion of the 6th pharyngeal arch, homeotic aberration and loss of rostral vertebrae, and reduced number of ribs and somites. Reciprocally, miR-196 knockdown evoked an extra pharyngeal arch, extra ribs, and extra somites, confirming endogenous roles of miR-196. miR-196 injection altered expression of hox genes and the signaling of retinoic acid through the retinoic acid receptor gene rarab. Knocking down rarab mimicked the pectoral fin phenotype of miR-196 overexpression, and reporter constructs tested in tissue culture and in embryos showed that the rarab 3′UTR is a miR-196 target for pectoral fin bud initiation. These results show that a Hox cluster microRNA modulates development of axial patterning similar to nearby protein-coding Hox genes, and acts on appendicular patterning at least in part by modulating retinoic acid signaling.     

A comparison of pectoral fin ray morphology and its impact on fin ray flexural stiffness in labriform swimmers

The organization of tissues in appendages often affects their mechanical properties and function. In the fish family Labridae, swimming behavior is associated with pectoral fin flexural stiffness and morphology, where fins range on a continuum from stiff to relatively flexible fins. Across this diversity, pectoral fin flexural stiffness decreases exponentially along the length of any given fin ray, and ray stiffness decreases along the chord of the fin from the leading to trailing edge. In this study, we examine the morphological properties of fin rays, including the effective modulus in bending (E), second moment of area (I), segmentation, and branching patterns, and their impact on fin ray stiffness. We quantify intrinsic pectoral fin ray stiffness in similarly sized fins of two closely related species that employ fins of divergent mechanics, the flapping Gomphosus varius and the rowing Halichoeres bivittatus. While segmentation patterns and E were similar between species, measurements of I and the number of fin ray branch nodes were greater in G. varius than in H. bivittatus. A multiple regression model found that of these variables, I was always significantly correlated with fin ray flexural stiffness and that variation in I always explained the majority of the variation in flexural stiffness. Thus, while most of the morphological variables quantified in this study correlate with fin ray flexural stiffness, second moment of area is the greatest factor contributing to variation in flexural stiffness. Further, interspecific variation in fin ray branching pattern could be used as a means of tuning the effective stiffness of the fin webbing to differences in swimming behavior and hydrodynamics. The comparison of these results to other systems begins to unveil fundamental morphological features of biological beams and yields insight into the role of mechanical properties in fin deformation for aquatic locomotion.     

Spot-fixed fin digging behavior in foraging of the benthophagous maiden goby, Pterogobius virgo (Perciformes: Gobiidae)

 Several patterns of feeding behaviors have been documented in benthophagous fishes. The foraging behavior of the maiden goby, Pterogobius virgo, was studied at Kurahashi Island in the Seto Inland Sea, Japan. Pterogobius virgo foraged mostly on polychaetes by volume from among several available prey items by digging in the sandy bottom. The digging behavior comprised swing of only pectoral fins or of both pectoral fins and body. Pectoral fin swing exposed the cryptic prey within the bottom, and fins and body swing exposed the prey and washed the sediment away. The swings were repeatedly and continuously conducted at a site during the daytime, making a pit several centimeters deep in which the fish was located. After the prey was exposed, the fish immediately and rapidly picked up the prey. Polychaetes were abundant prey in the sediment, occurring in the layer 3–5 cm deep from the bottom surface in the study area. In this goby, spot-fixed fin digging, the first documentation of feeding habits in gobies, may be effective for feeding on the most valuable prey, i.e., polychaetes, which may be otherwise unavailable for this fish. Received: April 24, 2001 / Revised: April 26, 2002 / Accepted: May 7, 2002     

Fibin, a novel secreted lateral plate mesoderm signal, is essential for pectoral fin bud initiation in zebrafish

We identified a novel secreted protein, fibin, in zebrafish, mice and humans. We inhibited its function in zebrafish embryos by injecting antisense fibin morpholino oligonucleotides. A knockdown of fibin function in zebrafish resulted in no pectoral fin bud initiation and abolished the expression of tbx5, which is involved in the specification of pectoral fin identification. The lack of pectoral fins in fibin-knockdown embryos was partially rescued by injection of fibin RNA. fibin was expressed in the lateral plate mesoderm of the presumptive pectoral fin bud regions. Its expression region was adjacent to that of tbx5. fibin expression temporally preceded tbx5 expression in presumptive pectoral fin bud regions, and not abolished in tbx5-knockdown presumptive fin bud regions. In contrast, fibin expression was abolished in retinoic acid signaling-inhibited or wnt2b-knockdown presumptive fin bud regions. These results indicate that fibin is a secreted signal essential for pectoral fin bud initiation in that it potentially acts downstream of retinoic acid and wnt signaling and is essential for tbx5 expression. The present findings have revealed a novel secreted lateral plate mesoderm signal essential for fin initiation in the lateral plate mesoderm.     

Outgrowth by fin motor axons in wildtype and a finless mutant of the Japanese medaka fish

The outgrowth of motor axons to the developing pectoral fin of the Japanese medaka fish (Oryzias latipes) was investigated both in wildtype embryos and in the pectoral finless (pl) mutants in which adults are missing pectoral fins. Late in embryogenesis the pectoral fin is a simple limb which contains two antagonist muscles which are innervated by presumptive motor neurons from the first four spinal segments (S1-4). The pectoral fin develops from a fin bud located in S1 and S2 centered on the border between S1 and S2 and, as with other limbs, one of the earliest signs of differentiation is the apical ectodermal ridge (AER). By the time the AER is well formed the growth cones of the presumptive motor neurons have reached the base of the fin bud and formed a plexus by extending toward the fin bud upon emergence from the spinal cord. This is especially evident on the ventral surface of the metamerically arranged axial muscles. For example, growth cones from S2 extend in a diagonal direction (both anterior and lateral) towards the fin bud. One hypothesis which can account for the pattern of motor outgrowth is that growth cones are attracted to the base of the fin bud, perhaps via a long distance cue. This hypothesis was tested by examining outgrowth of segmental nerves in pl embryos in which the fin buds arrest early in development following the initial appearance of the AER. In pl, nerves from S1-4 converged to form a plexus at the base of the abnormal fin bud, but the pattern of outgrowth varied from wildtype in a way consistent with a diminished capacity of the fin bud to attract segmental nerves to it.     

No role for direct touch using the pectoral fins,as an information gathering strategy in a blind fish

Blind Mexican cave fish (Astyanax fasciatus) lack a functional visual system and have been shown to sense their environment using a technique called hydrodynamic imaging, whereby nearby objects are detected by sensing distortions in the flow field of water around the body using the mechanosensory lateral line. This species has also been noted to touch obstacles, mainly with the pectoral fins, apparently using this tactile information alongside hydrodynamic imaging to sense their surroundings. This study aimed to determine the relative contributions of hydrodynamic and tactile information during wall following behaviour in blind Mexican cave fish. A wall was custom built with a ‘netted’ region in its centre, which provided very similar tactile information to a solid tank wall, but was undetectable using hydrodynamic imaging. The fish swam significantly closer to and collided more frequently with the netted region of this wall than the solid regions, indicating that the fish did not perceive the netted region as a solid obstacle despite being able to feel it as such with their pectoral fins. We conclude that the touching of objects with the pectoral fins may be an artefact of the intrinsic link between pectoral fin extensions and tail beating whilst swimming, and does not function to gather information. During wall following, hydrodynamic information appears to be used strongly in preference to tactile information in this non-visual system.     

The morphology of the cephalic lobes and anterior pectoral fins in six species of batoids

Many benthic batoids utilize their pectoral fins for both undulatory locomotion and feeding. Certain derived, pelagic species of batoids possess cephalic lobes, which evolved from the anterior pectoral fins. These species utilize the pectoral fins for oscillatory locomotion while the cephalic lobes are used for feeding. The goal of this article was to compare the morphology of the cephalic lobes and anterior pectoral fins in species that possess and lack cephalic lobes. The skeletal elements (radials) of the cephalic lobes more closely resembled the radials in the pectoral fin of undulatory species. Second moment of area (I), calculated from cephalic lobe radial cross sections, and the number of joints revealed greater flexibility and resistance to bending in multiple directions as compared to pectoral fin radials of oscillatory species. The cephalic lobe musculature was more complex than the anterior pectoral fin musculature, with an additional muscle on the dorsal side, with fiber angles running obliquely to the radials. In Rhinoptera bonasus, a muscle presumably used to help elevate the cephalic lobes is described. Electrosensory pores were found on the cephalic lobes (except Mobula japonica) and anterior pectoral fins of undulatory swimmers, but absent from the anterior pectoral fins of oscillatory swimmers. Pore distributions were fairly uniform except in R. bonasus, which had higher pore numbers at the edges of the cephalic lobes. Overall, the cephalic lobes are unique in their anatomy but are more similar to the anterior pectoral fins of undulatory swimmers, having more flexibility and maneuverability compared to pectoral fins of oscillatory swimmers. The maneuverable cephalic lobes taking on the role of feeding may have allowed the switch to oscillatory locomotion and hence, a more pelagic lifestyle. J. Morphol. 274:1070–1083, 2013. © 2013 Wiley Periodicals, Inc.     

Lycodes nishimurai, a new deep-sea eelpout (Teleostei: Zoarcidae) from the Sea of Japan

A new zoarcid fish, Lycodes nishimuari, is described on the basis of four specimens from the Yamato Bank, the Sea of Japan, at depths of 642–669 m. Although this species resembles Lycodes diapterus, L. hubbsi, L. pectoralis, L. macrochir, and L. nakamurae in having a deep notch in the pectoral fin and a single lateral line on the body, it is distinguished from them by the following combination of characters: no scales on pectoral fin base; scales on dorsum before dorsal fin origin; suborbital, preopercular, and last postorbital pores with risen rim; whitish lips; no whitish transverse bars on body; pectoral fin membrane and anterior part of dorsal fin membrane without black spots; basal color of vertical fins black. Supplementary material to this paper is available in electronic format at .     

Methoprene Photolytic Compounds Disrupt Zebrafish Development,Producing Phenocopies of Mutants in the Sonic Hedgehog Signaling Pathway

Environmental chemicals have been proposed to impact endocrine or retinoid pathways, causing developmental abnormalities in humans and other vertebrates. Presented evidence shows that exposure of zebrafish embryos to sunlight-induced photolytic products of the pesticide methoprene results in developmental defects in the head, heart, pectoral fins, and somites, and in spinal motor and optic nerve axons. Exposed embryos are phenocopies of zebrafish you-type mutants and, as in the mutant sonic-you, show underexpression of the signaling protein sonic hedgehog. Reduced expression of sonic hedgehog is also displayed in embryos treated with the retinoic acid synthesis inhibitor citral. This study identifies citral-related compounds as embryonic signaling disruptors of potential environmental concern.     

Retinoic Acid Is Involved in the Metamorphosis of the Anal Fin into an Intromittent Organ,the Gonopodium,in the Green Swordtail (Xiphophorus hellerii)

In poeciliid fish the male anal fin has been transformed into a gonopodium, an intromittent organ required for internal fertilization. Elevated testosterone levels induce metamorphosis of a subset of anal fin rays to grow and form the specialized terminal structures of the gonopodium. The molecular mechanisms underlying these processes are largely unknown. Here, we investigated whether retinoic acid (RA) signaling is involved in gonopodium development in the swordtail Xiphophorus hellerii. We showed that aldh1a2, a RA synthesizing enzyme, and the RA receptors, rar-ga and rar-gb, are expressed in anal fins during metamorphosis. aldh1a2 expression is regulated by testosterone in a concentration-dependent manner and is up-regulated in both hormone-induced and naturally developing gonopodia. Androgen receptor (ar), a putative regulator of gonopodial development, is co-expressed with aldh1a2 and the RA receptors in gonopodial rays. Importantly, experimental increase of RA signaling promoted growth of the gonopodium and increased the number of new segments. Based on gene expression analyses and pharmacological manipulation of gonopodium development, we show that the RA signaling pathway is activated in response to androgen signaling and promotes fin ray growth and development during the metamorphosis of the anal fin into the gonopodium.     

Tri-phasic expression of posterior Hox genes during development of pectoral fins in zebrafish: implications for the evolution of vertebrate paired appendages

During development of the limbs, Hox genes belonging to the paralogous groups 9-13 are expressed in three distinct phases, which play key roles in the segmental patterning of limb skeletons. In teleost fishes, which have a very different organization in their fin skeletons, it is not clear whether a similar patterning mechanism is at work. To determine whether Hox genes are also expressed in several distinct phases during teleost paired fin development, we re-analyzed the expression patterns of hox9-13 genes during development of pectoral fins in zebrafish. We found that, similar to tetrapod Hox genes, expression of hoxa/d genes in zebrafish pectoral fins occurs in three distinct phases, in which the most distal/third phase is correlated with the development of the most distal structure of the fin, the fin blade. Like in tetrapods, hox gene expression in zebrafish pectoral fins during the distal/third phase is dependent upon sonic hedgehog signaling (hoxa and hoxd genes) and the presence of a long-range enhancer (hoxa genes), which indicates that the regulatory mechanisms underlying tri-phasic expression of Hox genes have remained relatively unchanged during evolution. Our results suggest that, although simpler in organization, teleost fins do have a distal structure that might be considered comparable to the autopod region of limbs.     

Functional morphology of the pectoral fins in bamboo sharks, Chiloscyllium plagiosum: benthic vs. pelagic station-holding

Bamboo sharks (Chiloscyllium plagiosum) are primarily benthic and use their relatively flexible pectoral and pelvic fins to rest on and move about the substrate. We examined the morphology of the pectoral fins and investigated their locomotory function to determine if pectoral fin function during both benthic station-holding and pelagic swimming differs from fin function described previously in leopard sharks, Triakis semifasciata. We used three-dimensional kinematics and digital particle image velocimetry (DPIV) to quantify pectoral fin function in five white-spotted bamboo sharks, C. plagiosum, during four behaviors: holding station on the substrate, steady horizontal swimming, and rising and sinking during swimming. During benthic station-holding in current flow, bamboo sharks decrease body angle and adjust pectoral fin angle to shed a clockwise fluid vortex. This vortex generates negative lift more than eight times that produced during open water vertical maneuvering and also results in an upstream flow that pushes against the posterior surface of the pectoral fin to oppose drag. In contrast, there is no evidence of significant lift force in the wake of the pectoral fin during steady horizontal swimming. The pectoral fin is held concave downward and at a negative dihedral angle during steady horizontal swimming, promoting maneuverability rather than stability, although this negative dihedral angle is much less than that observed previously in sturgeon and leopard sharks. During sinking, the pectoral fins are held concave upward and shed a clockwise vortex with a negative lift force, while in rising the pectoral fin is held concave downward and sheds a counterclockwise vortex with a positive lift force. Bamboo sharks appear to sacrifice maneuverability for stability when locomoting in the water column and use their relatively flexible fins to generate strong negative lift forces when holding position on the substrate and to enhance stability when swimming in the water column.