Experimental Hydrodynamics and Evolution: Function of Median Fins in Ray-finned Fishes

The median fins of fishes consist of the dorsal, anal, and caudal fins and have long been thought to play an important role in generating locomotor force during both steady swimming and maneuvering. But the orientations and magnitudes of these forces, the mechanisms by which they are generated, and how fish modulate median fin forces have remained largely unknown until the recent advent of Digital Particle Image Velocimetry (DPIV) which allows empirical analysis of force magnitude and direction. Experimental hydrodynamic studies of median fin function in fishes are of special utility when conducted in a comparative phylogenetic context, and we have examined fin function in four ray-finned fish clades (sturgeon, trout, sunfish, and mackerel) with the goal of testing classical hypotheses of fin function and evolution. In this paper we summarize two recent technical developments in DPIV methodology, and discuss key recent findings relevant to median fin function. High-resolution DPIV using a recursive local-correlation algorithm allows quantification of small vortices, while stereo-DPIV permits simultaneous measurement of x, y, and z flow velocity components within a single planar light sheet. Analyses of median fin wakes reveal that lateral forces are high relative to thrust force, and that mechanical performance of median fins (i.e., thrust as a proportion of total force) averages 0.35, a surprisingly low value. Large lateral forces which could arise as an unavoidable consequence of thrust generation using an undulatory propulsor may also enhance stability and maneuverability. Analysis of hydrodynamic function of the soft dorsal fin in bluegill sunfish shows that a thrust wake is generated that accounts for 12% of total thrust and that the thrust generation by the caudal fin may be enhanced by interception of the dorsal fin wake. Integration of experimental studies of fin wakes, computational approaches, and mechanical models of fin function promise understanding of instantaneous forces on fish fins during the propulsive cycle as well as exploration of a broader locomotor design space and its hydrodynamic consequences.     

Experimental hydrodynamics of fish locomotion: functional insights from wake visualization

Despite enormous progress during the last twenty years in understandingthe mechanistic basis of aquatic animal propulsion—a taskinvolving the construction of a substantial data base on patternsof fin and body kinematics and locomotor muscle function—thereremains a key area in which biologists have little information:the relationship between propulsor activity and water movementin the wake. How is internal muscular force translated intoexternal force exerted on the water? What is the pattern offluid force production by different fish fins (e.g., pectoral,caudal, dorsal) and how does swimming force vary with speedand among species? These types of questions have received considerableattention in analyses of terrestrial locomotion where forceoutput by limbs can be measured directly with force plates.But how can forces exerted by animals moving through fluid bemeasured? The advent of digital particle image velocimetry (DPIV)has provided an experimental hydrodynamic approach for quantifyingthe locomotor forces of freely moving animals in fluids, andhas resulted in significant new insights into the mechanismsof fish propulsion. In this paper we present ten "lessons learned"from the application of DPIV to problems of fish locomotionover the last five years. (1) Three-dimensional DPIV analysisis critical for reconstructing wake geometry. (2) DPIV analysisreveals the orientation of locomotor reaction forces. (3) DPIVanalysis allows calculation of the magnitude of locomotor forces.(4) Swimming speed can have a major impact on wake structure.(5) DPIV can reveal interspecific differences in vortex wakemorphology. (6) DPIV analysis can provide new insights intothe limits to locomotor performance. (7) DPIV demonstrates thefunctional versatility of fish fins. (8) DPIV reveals hydrodynamicforce partitioning among fins. (9) DPIV shows that wake interactionamong fins may enhance thrust production. (10) Experimentalhydrodynamic analysis can provide insight into the functionalsignificance of evolutionary variation in fin design.     

A NEW SPINICAUDATAN GENUS (CRUSTACEA: ‘CONCHOSTRACA’) FROM THE LATE CRETACEOUS OF MADAGASCAR

Abstract: A new spinicaudatan genus and species, Ethmosestheria mahajangaensis gen. et sp. nov., is described from the Anembalemba Member (Upper Cretaceous, Maastrichtian) of the Maevarano Formation, Mahajanga Basin, Madagascar. This is the first spinicaudatan reported from the post‐Triassic Mesozoic of Madagascar. The new species is assigned to the family Antronestheriidae based on the cavernous or sievelike ornamentation on the carapace. Of well‐documented Mesozoic spinicaudatan genera, Ethmosestheria mahajangaensis is most closely related to Antronestheria Chen and Hudson from the Great Estuarine Group (Jurassic) of Scotland. However, relatively poor documentation of the ornamentation of most Gondwanan Mesozoic spinicaudatan species precludes detailed comparison among taxa. Ethmosestheria mahajangaensis exhibits ontogenetic trends in carapace growth: a change in carapace outline from subcircular/subelliptical to elliptical, and from very wide juvenile growth bands to narrow adult growth bands. Ornamentation style, however, does not vary with ontogeny. Ethmosestheria mahajangaensis individuals lived in temporary pools in a broad channel‐belt system within a semiarid environment; preserved desiccation structures on carapaces indicate seasonal drying out of pools within the river system. Specimens of Ethmosestheria mahajangaensis are preserved with exquisite detail in debris flow deposits; these are the first spinicaudatans reported from debris flow deposits. These deposits also contain a varied vertebrate fauna, including dinosaurs, crocodyliforms, turtles, and frogs. Rapid entombment of the spinicaudatan carapaces likely promoted early fossil diagenesis leading to highly detailed preservation.     

Repeatability of Habitat-Associated Divergence in Shell Shape of Turtles

Repeated patterns of phenotypic divergence between environments across disparate taxa provide strong evidence for the generation of adaptive phenotypes. Flow velocity is an important selective force in aquatic habitats; however, among vertebrates, the study of its effects on morphology has been limited almost exclusively to fully-aquatic bony fishes. We tested whether three confamilial species of semi-aquatic freshwater turtle (family Emydidae: Graptemys pseudogeographica, Graptemys nigrinoda, and Pseudemys concinna) displayed similar patterns of phenotypic divergence in carapace shape between fast- and slow-flowing aquatic environments. We used (1) geometric morphometrics to quantify shell shape, (2) multivariate analysis of variance to test the effects of species, sex, and flow, and (3) phenotypic trajectory analysis to compare patterns of divergence for six species-sex groups. We found significant effects on shell shape for all factors. In general, ecomorphs from fast-flowing habitats had flatter shells than those from slow-flowing habitats. Furthermore, results of trajectory analysis indicate that the degree to which, as well as the way in which, ecomorphs differed were concordant across all species. Our findings demonstrate that the effects of flow are not limited to fully-aquatic vertebrates, and provide evidence of the ability of flow to drive repeatable phenotypic divergence in tetrapods.     

Measurement of distribution of maximum index-fingertip force in all directions at fingertip in flexion/extension plane

The purpose of the present study is to investigate whether distribution patterns of the maximum fingertip force in all directions from 0 deg to 360 deg around the index fingertip are the same among subjects. Distributions of the maximum index fingertip force were measured at four finger postures for five subjects (healthy males, ages 21 to 25). It became apparent that distribution patterns of the fingertip force were very similar among subjects, through the application of an analysis of variance (level of significance: 5%) to the measurement results. In the production of the maximum fingertip force, Valero-Cuevas et al. [1998, 2000] reported that patterns of control signals for driving muscle forces were common among subjects. The results of the present study indicate that patterns of maximum fingertip force are also similar among subjects. Therefore, the possibility is high that the static transfer characteristic for index fingers from input (i.e., control signals to muscles) to output (ie., the maximum fingertip force) is also common among subjects.     

Aquatic prey capture in ray-finned fishes: a century of progress and new directions

The head of ray-finned fishes is structurally complex and is composed of numerous bony, muscular, and ligamentous elements capable of intricate movement. Nearly two centuries of research have been devoted to understanding the function of this cranial musculoskeletal system during prey capture in the dense and viscous aquatic medium. Most fishes generate some amount of inertial suction to capture prey in water. In this overview we trace the history of functional morphological analyses of suction feeding in ray-finned fishes, with a particular focus on the mechanisms by which suction is generated, and present new data using a novel flow imaging technique that enables quantification of the water flow field into the mouth. We begin with a brief overview of studies of cranial anatomy and then summarize progress on understanding function as new information was brought to light by the application of various forms of technology, including high-speed cinematography and video, pressure, impedance, and bone strain measurement. We also provide data from a new technique, digital particle image velocimetry (DPIV) that allows us to quantify patterns of flow into the mouth. We believe that there are three general areas in which future progress needs to occur. First, quantitative three-dimensional studies of buccal and opercular cavity dimensions during prey capture are needed; sonomicrometry and endoscopy are techniques likely to yield these data. Second, a thorough quantitative analysis of the flow field into the mouth during prey capture is necessary to understand the effect of head movement on water in the vicinity of the prey; three-dimensional DPIV analyses will help to provide these data. Third, a more precise understanding of the fitness effects of structural and functional variables in the head coupled with rigorous statistical analyses will allow us to better understand the evolutionary consequences of intra- and interspecific variation in cranial morphology and function.     

Tail blow energy and carapace fractures in a large glyptodont (Mammalia, Xenarthra)

Glyptodonts had muscular tails terminating, in many cases, with a rigid bony sheath which, in Doedicurus and Panochthus, was a formidable club. Some carapaces show fractures which have been interpreted as resulting from intraspecific fights. We estimate the energy that the tail muscles could have supplied for a blow and the energy required to fracture the carapace, and obtain results of the same order of magnitude. Thus it is not unreasonable to interpret die observed damage as resulting from fights. A space between the thoracic and lumbar vertebral column and the carapace may have been occupied by a fatty pad which would have served as a protective cushion, reducing the force of impacts.     

Linear intermolt growth in Americamysis bahia (Molenock 1969) (Mysida,Mysidae)

Two phases of allometry, prepubertal, and pubertal, are evident in A. bahia, equally dividing the first six instars. Coefficients of variation of the sizes of individuals’ abdomens, carapaces, and standard lengths significantly decrease in intermolt four, suggesting a synchronization of the size at which individuals enter the pubertal phase. Growth in standard length is best expressed over the first six post-larval instars by a quadratic equation, but within instars 85%, or more, of the expansion occurs linearly during the intermolt period, with only about 15% occurring during ecdysis. Estimations of carapace:abdomen allometry indicate an average ratio of 1:1.97 during the intermolt and 1:1.68 during ecdysis. The average molt:intermolt ratio in the carapace expansion is 1:3.16, and in the abdomen, 1:3.71, indicating that proportional growth between the carapace and the abdomen is continuous during both the intermolt and molt phases, but occur at different rates and proportions during the two periods.     

Musculoskeletal determinants of pelvic sucker function in hawaiian stream gobiid fishes: Interspecific comparisons and allometric scaling

Gobiid fishes possess a distinctive ventral sucker, formed from fusion of the pelvic fins. This sucker is used to adhere to a wide range of substrates including, in some species, the vertical cliffs of waterfalls that are climbed during upstream migrations. Previous studies of waterfall‐climbing goby species have found that pressure differentials and adhesive forces generated by the sucker increase with positive allometry as fish grow in size, despite isometry or negative allometry of sucker area. To produce such scaling patterns for pressure differential and adhesive force, waterfall‐climbing gobies might exhibit allometry for other muscular or skeletal components of the pelvic sucker that contribute to its adhesive function. In this study, we used anatomical dissections and modeling to evaluate the potential for allometric growth in the cross‐sectional area, effective mechanical advantage (EMA), and force generating capacity of major protractor and retractor muscles of the pelvic sucker (m. protractor ischii and m. retractor ischii) that help to expand the sealed volume of the sucker to produce pressure differentials and adhesive force. We compared patterns for three Hawaiian gobiid species: a nonclimber (Stenogobius hawaiiensis), an ontogenetically limited climber (Awaous guamensis), and a proficient climber (Sicyopterus stimpsoni). Scaling patterns were relatively similar for all three species, typically exhibiting isometric or negatively allometric scaling for the muscles and lever systems examined. Although these scaling patterns do not help to explain the positive allometry of pressure differentials and adhesive force as climbing gobies grow, the best climber among the species we compared, S. stimpsoni, does exhibit the highest calculated estimates of EMA, muscular input force, and output force for pelvic sucker retraction at any body size, potentially facilitating its adhesive ability. J. Morphol. 2013. © 2013 Wiley Periodicals, Inc.     

Direct comparison of measured and calculated total knee replacement force envelopes during walking in the presence of normal and abnormal gait patterns

Knee joint forces measured from instrumented implants provide important information for testing the validity of computational models that predict knee joint forces. The purpose of this study was to validate a parametric numerical model for predicting knee joint contact forces against measurements from four subjects with instrumented TKRs during the stance phase of gait. Model sensitivity to abnormal gait patterns was also investigated. The results demonstrated good agreement for three subjects with relatively normal gait patterns, where the difference between the mean measured and calculated forces ranged from 0.05 to 0.45 body weights, and the envelopes of measured and calculated forces (from three walking trials) overlapped. The fourth subject, who had a "quadriceps avoidance" external moment pattern, initially had little overlap between the measured and calculated force envelopes. When additional constraints were added, tailored to the subject's gait pattern, the model predictions improved to complete force envelope overlap. Coefficient of multiple determination analysis indicated that the shape of the measured and calculated force waveforms were similar for all subjects (adjusted coefficient of multiple correlation values between 0.88 and 0.92). The parametric model was accurate in predicting both the magnitude and waveform of the contact force, and the accuracy of model predictions was affected by deviations from normal gait patterns. Equally important, the envelope of forces generated by the range of solutions substantially overlapped with the corresponding measured envelope from multiple gait trials for a given subject, suggesting that the variable strategic processes of in vivo force generation are covered by the solution range of this parametric model.     

A simple, inexpensive system for digital particle image velocimetry (DPIV) in biomechanics

Functional morphology and biomechanics seek to reveal the mechanistic bases of organismal functions and the physical principles involved at the phenotype-environment interface. Characterization of fluid flow (air or water) within and around organismal structures is an example of this approach. Digital particle imaging velocimetry (DPIV) has been exploited in a variety of biological systems to visualize fluid flow associated with animal movement. DPIV employs particles suspended in air or water that are illuminated by a laser light sheet and recorded with a high-speed video camera. Software tracks particle movement across a specified number of video frames, generating vector diagrams showing patterns of fluid flow through time. As powerful as DPIV methods are, they are limited in application by the high cost and complexity of the equipment required. In this article, we describe a simple DPIV system that substitutes widely available, inexpensive consumer components for scientific-grade equipment to achieve low cost (<$1,000 total) and high accuracy (total error calculated to be approx. 6%, as compared with 5% in professional systems). We have employed this system successfully in our studies on the fluid dynamics of chemosensory tongue-flicking in snakes. This system can be used for research and teaching in labs that typically cannot afford the expense or commitment of a traditional DPIV apparatus and is particularly suited for obtaining preliminary data required to justify further grant and institutional support.     

Force measurements on single molecular contacts through evanescent wave microscopy

We introduce a new method to apply controlled forces on single molecules. The motion of a micron-sized bead attached to a solid surface through a single molecular contact is tracked by evanescent wave microscopy as a force is exerted through a flow. We report measurements of the streptavidin-biotin bond rupture force obtained with this technique. We also obtain detailed measurements of the balance of forces involved in detaching an adhering bead with a flow. A small lateral force translates into a much bigger normal force on the attachment point. This effect is relevant for the interpretation of common cell adhesion assays.     

Feeding biomechanics and theoretical calculations of bite force in bull sharks (Carcharhinus leucas) during ontogeny

Evaluations of bite force, either measured directly or calculated theoretically, have been used to investigate the maximum feeding performance of a wide variety of vertebrates. However, bite force studies of fishes have focused primarily on small species due to the intractable nature of large apex predators. More massive muscles can generate higher forces and many of these fishes attain immense sizes; it is unclear how much of their biting performance is driven purely by dramatic ontogenetic increases in body size versus size-specific selection for enhanced feeding performance. In this study, we investigated biting performance and feeding biomechanics of immature and mature individuals from an ontogenetic series of an apex predator, the bull shark, Carcharhinus leucas (73–285 cm total length). Theoretical bite force ranged from 36 to 2128 N at the most anterior bite point, and 170 to 5914 N at the most posterior bite point over the ontogenetic series. Scaling patterns differed among the two age groups investigated; immature bull shark bite force scaled with positive allometry, whereas adult bite force scaled isometrically. When the bite force of C. leucas was compared to those of 12 other cartilaginous fishes, bull sharks presented the highest mass-specific bite force, greater than that of the white shark or the great hammerhead shark. A phylogenetic independent contrast analysis of anatomical and dietary variables as determinants of bite force in these 13 species indicated that the evolution of large adult bite forces in cartilaginous fishes is linked predominantly to the evolution of large body size. Multiple regressions based on mass-specific standardized contrasts suggest that the evolution of high bite forces in Chondrichthyes is further correlated with hypertrophication of the jaw adductors, increased leverage for anterior biting, and widening of the head. Lastly, we discuss the ecological significance of positive allometry in bite force as a possible “performance gain” early in the life history of C. leucas.     

Relevance of crustacean carapace wettability for fouling

Carapace wettability and density of fouling organisms (bacteria, diatoms, protozoa, fungi, macro-organisms) were investigated for 45 crustacean species (Hoplocarida, Decapoda) from 15 families in the Gulf of Thailand. The results show that crustaceans can create and maintain characteristic carapace wettabilities. About 21 species (47%) possess highly wettable carapaces with contact angles below 20°. Contact angles between 20° and 40° were recorded for four species (2%), angles between 40° and 60° for eight species (4%) and from 60° to 70° for 11 (24%) species. One species, Alpheus euphrosyne (Alpheidae, Decapoda), exhibited an extremely low surface wettability (contact angle: 91°). Densities of colonisers and contact angles did not correlate. Very low wettability by water ( > 90°) may only contribute little to fouling reduction in A. euphrosyne which showed the most hydrophobic carapace surface and was colonised by the lowest numbers of bacteria among all species and no other colonisers at all. We conclude that surface wettability is of little relevance for antifouling defence in crustaceans.     

Push or Pull? The light‐weight architecture of the Daphnia pulex carapace is adapted to withstand tension,not compression

Daphnia (Crustacea, Cladocera) are well known for their ability to form morphological adaptations to defend against predators. In addition to spines and helmets, the carapace itself is a protective structure encapsulating the main body, but not the head. It is formed by a double layer of the integument interconnected by small pillars and hemolymphatic space in between. A second function of the carapace is respiration, which is performed through its proximal integument. The interconnecting pillars were previously described as providing higher mechanical stability against compressive forces. Following this hypothesis, we analyzed the carapace structure of D. pulex using histochemistry in combination with light and electron microscopy. We found the distal integument of the carapace to be significantly thicker than the proximal. The pillars appear fibrous with slim waists and broad, sometimes branched bases where they meet the integument layers. The fibrous structure and the slim‐waisted shape of the pillars indicate a high capacity for withstanding tensile rather than compressive forces. In conclusion they are more ligaments than pillars. Therefore, we measured the hemolymphatic gauge pressure in D. longicephala and indeed found the hemocoel to have a pressure above ambient. Our results offer a new mechanistic explanation of the high rigidity of the daphniid carapace, which is probably the result of a light‐weight construction consisting of two integuments bound together by ligaments and inflated by a hydrostatic hyper‐pressure in the hemocoel. J. Morphol. 277:1320–1328, 2016. © 2016 Wiley Periodicals, Inc.     

EFFECTS OF INTRINSIC AND EXTRINSIC FACTORS ON POPULATION FRAGMENTATION IN THREE SPECIES OF NORTH AMERICAN MINNOWS (TELEOSTEI: CYPRINIDAE)

Geographic patterns of genetic variation (mitochondrial DNA [mtDNA] and allozymes) were used to examine effects of intrinsic characteristics (e.g., vagility, habitat specificity, and reproductive behaviors) and extrinsic factors (e.g., climatic and geological history) on population fragmentation. The three species of cyprinid fishes examined (Tiaroga cobitis, Meda fulgida, and Agosia chrysogaster) occupied similar historical ranges within the lower Colorado River drainage, but differ in intrinsic characteristics conducive to population fragmentation. Relationships among populations were similar across species, reflecting common historical influences, but results indicate the distribution of variation among species is strongly affected by intrinsic characteristics. Variation within two species (T. cobitis and M. fulgida) is subdivided among populations, suggesting little gene flow among rivers. In contrast, similarity of A. chrysogaster populations throughout the Gila River drainage supports the hypothesis that levels of gene flow are high for this species. Levels of mtDNA divergence were much higher than expected for both T. cobitis and A. chrysogaster suggesting long-term isolation of geographic regions. These results indicate that both long-term and short-term extrinsic factors have shaped basic patterns of variation within these fishes; however, the intrinsic characteristics of each species have strongly affected the population genetic structure of these fishes.     

Morphological adaptation of shape to flow: Microcurrents around lotic macroinvertebrates with known Reynolds numbers at quasi-natural flow conditions

Summary Using Laser Doppler Anemometry we measured current velocities in the median plane around dead lotic macroinvertebrates in a flume which reproduced natural near bottom hydraulics. We investigated specimens of the gastropods Ancylus, Acroloxus, and Potamopyrgus, the amphipod Gammarus, and the larval caddisflies Anabolia, Micrasema, and Silo of various size, various alignment to the flow or which were otherwise manipulated in order to clarify certain questions of adaptation of shape or case building style to flow, or the effects of flow on field distribution patterns. The steepest velocity gradients close to the animals were found near areas of their bodies protruding furthest into the flow. In such regions the rates of potential diffusive exchange processes, the potential corrasion (abrasion through suspended solids), and, for larger specimens, the lift forces (directed towards the water surface) must be highest. Posterior of these areas growing boundary layers formed above those species whose upper contour was approximately parallel to the upstream-downstream direction of the flow. All specimens removed momentum from the flow and thus experience a drag force (directed downstream). From the complete data set we derived the following general conclusions about the physical effects of potential morphological adaptations, taking into consideration diffusion through boundary layers, corrasion, lift forces, friction and pressure drag forces: The physical significance of these five factors generally depends on the Reynolds number of an animal and is largely affected by flow separation, which was significantly related to the ratio of body length to height and the slope of the posterior contour. A simultaneous effective morphological adaptation to all five factors is physically impossible and, in addition, would have to change from life at low (e.g. a young, small specimen of a species) to life at high (e.g. a fully grown specimen of the same species) Reynolds number.     

The distribution and orientation of epizoic barnacles on crabs

The distribution and orientation of Balanus crenatus on Carcinus maenas is described. Elminius modestus was also found on C. maenas but it was not common. Both species of barnacle were also found on Cancer pagurus, although neither species of barnacle appeared to be as common on Cancer as on Carcinus. Possible reasons for this are discussed.
Treating the carapaces of similar sized C. maenas as sampling units it is clear that the distribution of B. crenatus over these sampling units is not random but is aggregated. This probably arises because of the gregarious nature of the cyprids when they settle out but other possibilities are considered. B. crenatus on the carapace of C. maenas is found exclusively in the grooves and depressions on the carapace and this is because the cyprids preferentially settle in concavities.
The orientation of B. crenatus also shows a consistent pattern, with the cirral nets facing predominantly backwards. This pattern could arise because the barnacle cyprids orientate to water currents, generated by the exhalant respiratory currents, flowing forwards over the carapace.     

Size,but not sex,predicts pinch force and exoskeleton mechanical properties in crayfish of the genus Faxonius

Studies of animal weaponry and defensive structures rarely take into consideration their underlying mechanical properties. We measured the compressive strength and thickness of the exoskeleton of the claw (chela) in two North American crayfish species, Faxonius virilis and F. limosus. We performed similar measures on the carapace, a body region not directly involved in agonistic contests. Males of both species generated significantly stronger maximum pinch forces than females. However, these differences can be attributed to differences in claw size between the sexes. The thickness (ultrastructure) of the claw exoskeleton was a significant predictor of its compressive strength and likely explained the difference in compressive strength we observed between the two species. Neither claw thickness nor claw compressive strength was correlated with maximum pinch force. Additionally, we found that crayfish body size was a strong predictor of carapace compressive strength and thickness, whereas sex was not. The claw had greater compressive strength and thickness than the corresponding values for the carapace. Our study shows that the mechanical properties of the crayfish exoskeleton are largely a function of size and highlights the need to integrate mechanical properties into studies of animal morphology and performance.