Is tilting behaviour at low swimming speeds unique to negatively buoyant fish? Observations on steelhead trout,Oncorhynchus mykiss,and bluegill,Lepomis macrochirus

When swimming at low speeds, steelhead trout and bluegill sunfish tilted the body at an angle to the mean swimming direction. Trout swam using continuous body/caudal fin undulation, with a positive (head-up) tilt angle ( 0 , degrees) that decreased with swimming speed ( u , cm s⁻¹) according to: 0 =(164±96).u^{(−1.14±0.41)} (regression coefficients; mean±2 s.e. ). Bluegill swimming gaits were more diverse and negative (head down) tilt angles were usual. Tilt angle was −3·0 ± 0.9° in pectoral fin swimming at speeds of approximately 0.2–1.7 body length s⁻¹ (Ls⁻¹; 3–24 cm s⁻¹), −4.5 ±2.6° during pectoral fin plus body/caudal fin swimming at 1·2–1·7 L s⁻¹ (17–24cm s⁻¹), and −5.0± 1.0° during continuous body/caudal fin swimming at 1.6 and 2.5 L s⁻¹ (22 and 35cm s⁻¹). At higher speeds, bluegill used burst-and-coast swimming for which the tilt angle was 0.1±0.6°. These observations suggest that tilting is a general phenomenon of low speed swimming at which stabilizers lose their effectiveness. Tilting is interpreted as an active compensatory mechanism associated with increased drag and concomitant increased propulsor velocities to provide better stabilizing forces. Increased drag associated with trimming also explains the well-known observation that the relationship between tail-beat frequency and swimming speed does not pass through the origin. Energy dissipated because of the drag increases at low swimming speeds is presumably smaller than that which would occur with unstable swimming.     

Functional morphology of the fin rays of teleost fishes

Ray‐finned fishes are notable for having flexible fins that allow for the control of fluid forces. A number of studies have addressed the muscular control, kinematics, and hydrodynamics of flexible fins, but little work has investigated just how flexible ray‐finned fish fin rays are, and how flexibility affects their response to environmental perturbations. Analysis of pectoral fin rays of bluegill sunfish showed that the more proximal portion of the fin ray is unsegmented while the distal 60% of the fin ray is segmented. We examined the range of motion and curvatures of the pectoral fin rays of bluegill sunfish during steady swimming, turning maneuvers, and hovering behaviors and during a vortex perturbation impacting the fin during the fin beat. Under normal swimming conditions, curvatures did not exceed 0.029 mm^?1 in the proximal, unsegmented portion of the fin ray and 0.065 mm^?1 in the distal, segmented portion of the fin ray. When perturbed by a vortex jet traveling at approximately 1 ms^?1 (67 ± 2.3 mN s.e. of force at impact), the fin ray underwent a maximum curvature of 9.38 mm^?1. Buckling of the fin ray was constrained to the area of impact and did not disrupt the motion of the pectoral fin during swimming. Flexural stiffness of the fin ray was calculated to be 565 × 10^?6 Nm². In computational fluid dynamic simulations of the fin‐vortex interaction, very flexible fin rays showed a combination of attraction and repulsion to impacting vortex dipoles. Due to their small bending rigidity (or flexural stiffness), impacting vortices transferred little force to the fin ray. Conversely, stiffer fin rays experienced rapid small‐amplitude oscillations from vortex impacts, with large impact forces all along the length of the fin ray. Segmentation is a key design feature of ray‐finned fish fin rays, and may serve as a means of making a flexible fin ray out of a rigid material (bone). This flexibility may offer intrinsic damping of environmental fluid perturbations encountered by swimming fish. J. Morphol. 274:1044–1059, 2013. © 2013 Wiley Periodicals, Inc.     

Opercular differential pressure as a predictor of metabolic oxygen demand in the starry flounder

The feasibility of using a differential pressure sensor connected to an acoustic telemetry device to monitor opercular activity as a correlate of oxygen consumption was investigated. Four starry flounders Platichthys stellatus were fitted with a miniature differential pressure sensor mounted close to the operculum. A cannula was connected to the sensor and inserted under the operculum, inside the branchial cavity. Measurements of oxygen consumption and opercular activity were carried out over a broad range of metabolic activity, from the post‐surgery stress (high metabolic rate) to routine metabolic rate the following day. Relationships between differential pressure changes (rate and amplitude) were highly correlated with oxygen consumption ( r ² = 0·74 and 0·60 respectively). The results indicate that monitoring opercular activity offers an alternative method for measuring aerobic metabolism in free‐swimming fishes in nature.     

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.     

The role of the pectoral fins in body trim of sharks

In a large aquarium the leopard shark Triakis semifasciata , sand tiger shark Odontaspis taurus , sandbar shark Carcharhinus plumbeus , and spiny dogfish Squalus acanthias cruised steadily at 0·1-0·7 body lengths s^-1. Relative to the trajectory of the shark, the pectoral fins were maintained at a positive angle of ttack regardless of vertical direction. For level swimming the mean angle of attack for the pectoral fin was 11±1·7, 10·1±1·3°, 9·3±1·3°, and 15·0±0·0 for T. semifasciata , C. plumbeus , O. taurus , and S. acanthias , respectively. The long axis of the body was canted at an angle of attack for T. semifasciata and S. acanthias , but trim was maintained during level swimming for C. plumbeus and O. taurus . Hydrodynamic analysis of the body and fin design of T. semifasciata indicated that the pectoral fins could develop suffcient pitching moment to maintain depth and keep the body in trim. Demonstration of positive angles of attack support the hypothesis that lift is generated in the anterior body to counterbalance the lift produced by the heterocercal tail.     

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.     

Loaches of the genus Lepidocephalichthys, from Manipur, with description of a new species

A new species Lepidocephalichthys manipurensis , is described, confined to the Chandel district of Manipur near the adjoining borderland areas of Manipur, India and Burma (Myanmar). It is distinguished by the following combination of characters: a sharp small black dot just above the upper base of the caudal fin; 8–9 dorsal short dark ashy brown bars from occiput to the base of the caudal fin; absence of scales on the vertex of head; caudal fin fork, pectoral, ventral and anal fins are non-immaculate; least depth of caudal peduncle 50·00–60·24% (mean 54·09) of its length and 7·69–8·11% L , S (mean 8·22); caudal fin with 4–5 W-shaped bands: a single dark black stripe from tip of snout to eye; 8–11 spots on the mid lateral line; and 30–31 vertebrae. A key to the identification of lepidocephalid loaches of Manipur is provided.     

A new species of the genusamblyeleotris (pisces: Gobiidae) from Japan

A new shrimp-associated goby,Amblyeleotris melanocephala, is described on the basis of specimens from Okinoshima Island. Kochi Prefecture, and Okinawa Island, Okinawa Prefecture, Japan. The species is distinguished from other members of the genus by the following combination of characters: head dark brown, a few yellow spots on pectoral fin base and opercular margin, 13 second dorsal and 13 anal fin soft rays, 20 pectoral fin rays, longitudinal scales 92–101, proportional length of interpelvic connecting membrane relative to longest pelvic fin ray (CM-value) 0.46–0.55, presence of a ventral frenum, midline of nape naked, sides scaled above midpoint between preopercle and opercle.     

The locomotory function of the fins in the squid Loligo pealei

Kinematic data of high spatial and temporal resolution, acquired from image sequences of adult long-finned squid, Loligo pealei, during steady swimming in a flume, were used to examine the role of fins and the coordination between fin and jet propulsion in squid locomotion. Fin shape and body outlines were digitized and used to calculate fin wave speed, amplitude, frequency, angle of attack, body deformation, speed, and acceleration. L. pealei were observed to have two fin gait patterns with a transition at 1.4-1.8 mantle lengths per second (Lm s-1) marked by alternation between the two patterns. Fin motion in L. pealei exhibited characteristics of both traveling waves and flapping wings. At low speeds, fin motion was more wave-like; at high speeds, fin motion was more flap-like and was marked by regular periods during which the fins were wrapped tightly against the mantle. Fin cycle frequencies were dependent on swimming speed and gait, and obvious coordination between the fins and jet were observed. Fin wave speed, angle of attack, and body acceleration confirmed the role of fins in thrust production and revealed a role of fins at all swimming speeds by a transition from drag-based to lift-based thrust when fin wave speed dropped below swimming speed. Estimates of peak fin thrust were as high as 0.44-0.96 times peak jet thrust in steady swimming over the range of swimming speeds observed. Fin downstrokes generally contributed more to thrust than did upstrokes, especially at high speeds.     

The locomotory function of the fins in the squid Loligo pealei

Kinematic data of high spatial and temporal resolution, acquired from image sequences of adult long-finned squid, Loligo pealei, during steady swimming in a flume, were used to examine the role of fins and the coordination between fin and jet propulsion in squid locomotion. Fin shape and body outlines were digitized and used to calculate fin wave speed, amplitude, frequency, angle of attack, body deformation, speed, and acceleration. L. pealei were observed to have two fin gait patterns with a transition at 1.4-1.8 mantle lengths per second (L_m s^-1) marked by alternation between the two patterns. Fin motion in L. pealei exhibited characteristics of both traveling waves and flapping wings. At low speeds, fin motion was more wave-like; at high speeds, fin motion was more flap-like and was marked by regular periods during which the fins were wrapped tightly against the mantle. Fin cycle frequencies were dependent on swimming speed and gait, and obvious coordination between the fins and jet were observed. Fin wave speed, angle of attack, and body acceleration confirmed the role of fins in thrust production and revealed a role of fins at all swimming speeds by a transition from drag-based to lift-based thrust when fin wave speed dropped below swimming speed. Estimates of peak fin thrust were as high as 0.44-0.96 times peak jet thrust in steady swimming over the range of swimming speeds observed. Fin downstrokes generally contributed more to thrust than did upstrokes, especially at high speeds.     

Pectoral fin beat frequency predicts oxygen consumption during spontaneous activity in a labriform swimming fish (Embiotoca lateralis)

The objective of this study was to identify kinematic variables correlated with oxygen consumption during spontaneous labriform swimming. Kinematic variables (swimming speed, change of speed, turning angle, turning rate, turning radius and pectoral fin beat frequency) and oxygen consumption (MO₂) of spontaneous swimming in Embiotoca lateralis were measured in a circular arena using video tracking and respirometry, respectively. The main variable influencing MO₂ was pectoral fin beat frequency (r ² = 0.71). No significant relationship was found between swimming speed and pectoral fin beat frequency. Complementary to other methods within biotelemetry such as EMG it is suggested that such correlations of pectoral fin beat frequency may be used to measure the energy requirements of labriform swimming fish such as E. lateralis in the field, but need to be taken with great caution since movement and oxygen consumption patterns are likely to be quite different in field situation compared to a small lab tank. In addition, our methods could be useful to measure metabolic costs of growth and development, or bioassays for possible toxicological effects on fish.     

Pectoral Fin of the Megamouth Shark: Skeletal and Muscular Systems,Skin Histology,and Functional Morphology

This is the first known report on the skeletal and muscular systems, and the skin histology, of the pectoral fin of the rare planktivorous megamouth shark Megachasma pelagios. The pectoral fin is characterized by three features: 1) a large number of segments in the radial cartilages; 2) highly elastic pectoral fin skin; and 3) a vertically-rotated hinge joint at the pectoral fin base. These features suggest that the pectoral fin of the megamouth shark is remarkably flexible and mobile, and that this flexibility and mobility enhance dynamic lift control, thus allowing for stable swimming at slow speeds. The flexibility and mobility of the megamouth shark pectoral fin contrasts with that of fast-swimming sharks, such as Isurus oxyrhinchus and Lamna ditropis, in which the pectoral fin is stiff and relatively immobile.     

The evolution of underwater flight: The redistribution of pectoral fin rays,in manta rays and their relatives (Myliobatidae)

Batoids are a diverse clade of flat cartilaginous fishes that occur primarily in benthic marine habitats. The skates and rays typically use their flexible pectoral fins for feeding and propulsion via undulatory swimming. However, two groups of rays have adopted a pelagic or bentho‐pelagic lifestyle and utilize oscillatory swimming—the Myliobatidae and Gymnuridae. The myliobatids have evolved cephalic lobes, anteriorly extended appendages that are optimized for feeding, while their pectoral fins exhibit several modifications that likely arose in association with functional optimization of pelagic cruising via oscillatory flight. Here, we examine variation in fin ray distribution and ontogenetic timing of fin ray development in batoid pectoral fins in an evolutionary context using the following methods: radiography, computed tomography, dissections, and cleared and stained specimens. We propose an index for characterizing variation in the distribution of pectoral fin rays. While undulatory swimmers exhibit symmetry or slight anterior bias, we found a posterior shift in the distribution of fin rays that arose in two distinct lineages in association with oscillatory swimming. Undulatory and oscillatory swimmers occupy nonoverlapping morphospace with respect to fin ray distribution illustrating significant remodeling of pectoral fins in oscillatory swimmers. Further, we describe a derived skeletal feature in anterior pectoral fins of the Myliobatidae that is likely associated with optimization of oscillatory swimming. By examining the distribution of fin rays with clearly defined articulation points, we were able to infer evolutionary trends and body plan remodeling associated with invasion of the pelagic environment. Finally, we found that the number and distribution of fin rays is set early in development in the little skate, round stingray, and cownose ray, suggesting that fin ray counts from specimens after birth or hatching are representative of adults and therefore comparable among species.     

Gait transition and oxygen consumption in swimming striped surfperch Embiotoca lateralis Agassiz

A flow-through respirometer and swim tunnel was used to estimate the gait transition speed ( U _p-c) of striped surfperch Embiotoca lateralis , a labriform swimmer, and to investigate metabolic costs associated with gait transition. The U _p-c was defined as the lowest speed at which fish decrease the use of pectoral fins significantly. While the tail was first recruited for manoeuvring at relatively low swimming speeds, the use of the tail at these low speeds [as low as 0·75 body (fork) lengths s⁻¹, L _F s⁻¹) was rare (<10% of the total time). Tail movements at these low speeds appeared to be associated with occasional slow manoeuvres rather than providing power. As speed was increased beyond U _p-c, pectoral fin (PF) frequencies kept increasing when the tail was not used, while they did not when PF locomotion was aided by the tail. At these high speeds, the tail was employed for 40–50% of the time, either in addition to pectoral fins or during burst-and-coast mode. Oxygen consumption increased exponentially with swimming speeds up to gait transition, and then levelled off. Similarly, cost of transport ( C _T) decreased with increasing speed, and then levelled off near U _p-c. When speeds ≥ U _p-c are considered, C _T is higher than the theoretical curve extrapolated for PF swimming, suggesting that PF swimming appears to be higher energetically less costly than undulatory swimming using the tail.     

大麻哈鱼卵黄囊期仔鱼异速生长及其生态学意义

运用实验生态学的方法, 对大麻哈鱼(Oncorhynchus keta Walbaum)卵黄囊期仔鱼的异速生长及器官优先发育在早期生存和环境适应上的生态学意义进行了研究。结果表明, 大麻哈鱼卵黄囊期仔鱼的感觉、摄食, 呼吸和游泳等器官快速分化, 许多关键器官均存在异速生长现象。在身体各部分中, 头部和尾部为正异速生长, 躯干部为负异速生长, 体高有先增大后减小的趋势; 在头部器官中, 眼径、口宽、吻长和眼后头长均为正异速生长; 在游泳器官中, 胸鳍、腹鳍、背鳍、臀鳍、背鳍基、臀鳍基和尾鳍均为正异速生长, 脂鳍为负异速生长, 其中, 腹鳍在全长25.31 mm、12日龄出现生长拐点, 但拐点前后均为正异速生长。大麻哈鱼卵黄囊期仔鱼感觉、摄食, 呼吸和游泳等器官的快速发育, 使出膜后的仔鱼在最短的时间内获得了与早期生存密切相关的各种能力, 对适应复杂多变的外界环境具有重要的生态学意义。       

Comparing aging precision of calcified structures in shovelnose sturgeon

Age estimates for population analysis must be precise. We assessed the usefulness of pectoral fin rays, sphenoids, opercula, and dorsal scutes of shovelnose sturgeonScaphirhynchus platorynchus (n = 30) as aging structures based on ease of collection, distinctness of annuli, and measures of precision both between and within readers. We also determined how age estimates from paired fin rays of individuals were related (n = 106). Pectoral fin rays generated the highest within‐reader precision (100% within 2 years) followed by sphenoids (58%), opercula (56%), and dorsal scutes (49%). Ages estimated by the pectoral fin ray also had higher between‐reader agreement (80% within 1 year) than did those from the operculum (60%), sphenoid (59%), or dorsal scute (56%). Likewise, age estimates from pectoral fin rays had the lowest mean coefficient of variation (8.2%) followed by sphenoids (9.9%), opercula (11.3%), and dorsal scutes (11.5%). Only the operculum produced biased estimates between readers. Ages from paired fin rays agreed poorly (36% exact, 30% within 1 year) although no aging bias occurred. The pectoral fin ray is typically used to age shovelnose sturgeon. Because uncertainty about accuracy and precision of age estimates from this structure remains, shovelnose sturgeon management objectives that result from age data should remain conservative.     

Locomotion with flexible propulsors: II. Computational modeling of pectoral fin swimming in sunfish

This paper describes a computational fluid dynamics (CFD) based investigation of the pectoral fin hydrodynamics of a bluegill sunfish. The pectoral fin of this fish undergoes significant shape-change during its abduction-adduction cycle and the effect of this deformation on the thrust performance remains far from understood. The current study is part of a combined experimental-numerical approach wherein the numerical simulations are being used to examine features and issues that are not easily amenable to the experiments. These numerical simulations are highly challenging and we briefly describe the computational methodology that has been developed to handle such flows. Finally, we describe some of the key computational results including wake vortex topologies and hydrodynamics forces.     

Locomotory capacity of Baltic Sea and freshwater populations of the threespine stickleback (Gasterosteus aculeatus)

Two freshwater populations and one marine population (Baltic Sea) of threespine stickeback (Gasterosteus aculeatus) from Northeastern Germany were studied with regard to locomotory capacity: sustained swimming performance, activities of key enzymes in axial muscle, pectoral fin muscle and heart, and morphology. We postulated that life history differences between migratory Baltic Sea and resident freshwater populations could have led to a divergence in their locomotory capacity. The activity of citrate synthase (CS) in pectoral muscle correlated with critical swimming speed. Critical swimming speed, aerobic and anaerobic capacity of the pectoral fin muscle were population-specific. The Baltic Sea sticklebacks had a higher locomotory capacity (activity of CS in pectoral muscle, critical swimming speed) than sticklebacks of one freshwater population. However, another freshwater population expressed a similar locomotory capacity as the Baltic Sea population. In addition, Baltic Sea sticklebacks had a greater mass and lower anaerobic capacity of the pectoral fin muscle than the freshwater sticklebacks. The results are interpreted as an indication of a proceeding divergence between marine and resident freshwater populations and between freshwater populations of G. aculeatus originating from marine ancestors. The migratory Baltic Sea sticklebacks had better morphological prerequisites for sustained swimming than both freshwater populations, but there was no general difference in the locomotory capacity between marine and freshwater sticklebacks. However, their morphology could favour a more effective locomotion in the Baltic Sea sticklebacks.     

Assessing the ecological relevance of swimming performance traits: a case study of bluegill sunfish (<Emphasis Type="Italic">Lepomis macrochirus</Emphasis>)

A variety of fish species show habitat-related variation in traits associated with swimming performance and foraging behavior. This commonly manifests as a distinction between open water and shallow water littoral ecotypes. In bluegill sunfish (Lepomis macrochirus), open water fish exhibit greater energy economy and speed during sustained locomotion than those from the littoral, whereas littoral fish were more maneuverable than their open water counterparts. These distinctions are associated with variation in diet and foraging behavior and may represent a resource polyphenism that enhances fitness through more effective exploitation of particular habitat types. A lack of field data means that polyphenisms have not been placed in context with swimming behavior in the field. We have used 3D videography to quantify bluegill field swimming performance in open water and littoral habitats. This revealed patterns of performance variation that parallel the trait variation previously established in the laboratory. Open water fish utilized faster average swimming speeds than inshore fish, while indicators of nonlinearity and unsteadiness were greater in the littoral fish. There are, however, differences in propulsive behavior between the field and laboratory. Pectoral-fin-powered, median-paired fin swimming is rarely employed by open water fish. Field body-caudal fin swimming involves short sequences of propulsive tail beats interspersed with gliding, rather than the repeated propulsive cycles employed under steady-state conditions. This suggests a need to re-evaluate the applicability of steady-state performance traits to behavior and fitness in the field and highlights the general importance of obtaining field performance data.