A simplified computational model of possible hydrodynamic interactions between respiratory and swimming-related water flows in labriform-swimming fishes.

Hydrodynamic interactions in bony fishes between respiratory fluid flows leaving the opercular openings and simultaneous flows generated by movements of downstream pectoral fins are both poorly understood and likely to be complex. Labriform-swimming fishes that swim primarily by moving only their pectoral fins are good subjects for these studies. We performed a computational fluid dynamics investigation of a simplified 2D model of these interactions based on previously published experimental observations of both respiratory and pectoral fin movements under both resting and slow, steady swimming conditions in two similar labriform swimmers: the bluegill sunfish (L. macrochirus) and the largemouth bass (M. salmoides). We carried out a parametric study investigating the effects that swimming speed, strength of opercular flow and phase difference between the pectoral fin motion and the opercular opening and closing have on the thrust and sideslip forces generated by the pectoral fins during both the abduction and adduction portions of the fin movement cycle. We analyzed pressure distributions on the fin surface to determine physical differences in flows with and without opercular jets. The modeling indicates that complex flow structures emerge from the coupling between the opercular jets and vortex shedding from pectoral fins. The jets from the opercular openings appear to exert significant influence on the forces generated by the fins; they are potentially significant in the maneuverability of at least some labriform swimmers. The numerical simulations and the analysis establish a framework for the study of these interactions in various labriform swimmers in a variety of flow regimes. Similar situations in groups of fishes using other swimming modes should also be investigated.

A Comparison of Isogenic Homozygous Clone and Wildtype Zebrafish (Danio rerio): Survival and Developmental Responses to Low pH Conditions.

The value of bioassays as analytical methods for assessing the potency of particular stressors on live animal models depends on the precision of their results, which are greatly influenced by the choice of test subjects. The genetic makeup of experimental subjects varies, and, as such, so will their responses to the test environment. Genetic diversity of test populations may contribute to statistical variability; therefore, the use of genetically similar subjects may enhance the utility of bioassays. This study addresses the efficacy of using isogenic homozygous zebrafish (Danio rerio) as subjects for bioassays. Stress responses (acidic conditions) were compared during early development for gynogenetically produced isogenic homozygous line of zebrafish (C32) and wildtype (WT) zebrafish. Experiments evaluated early life stage milestones after exposure to low pH in water of a different electrolyte composition. Because the isogenic homozygous clonal (IHC) fish possessed far less genetic variability than the WT fish tested, it was predicted that the IHC fish would exhibit less variability in their response to stress. Although we found no significant differences in the variability between the responses of the IHC and WT fish, pH and water hardness level had a differential effect on the two groups. Simple strain differences may be the probable cause of the response differences to environmental stress. Factors that may affect stress response, such as heterogeneity, co-adapted gene complexes, and domestication, are discussed. Our findings and review of recent zebrafish literature stress the need for researchers to carefully consider breeding histories and trait characteristics for each potential test subject to maximize the sensitivity of the assay.

Aracaniform Swimming: A Proposed New Category of Swimming Mode in Bony Fishes (Teleostei: Tetraodontiformes: Aracanidae).

The deepwater boxfishes of the family Aracanidae are the phylogenetic sister group of the shallow-water, generally more tropical boxfishes of the family Ostraciidae. Both families are among the most derived groups of teleosts. All members of both families have armored bodies, the forward 70% of which are enclosed in rigid bony boxes (carapaces). There is substantial intragroup variation in both groups in body shapes, sizes, and ornamentation of the carapaces. Swimming-related morphology, swimming mode, biomechanics, kinematics, and hydrodynamics have been studied in detail in multiple species of the ostraciids. Ostraciids are all relatively high-performance median and paired fin swimmers. They are highly maneuverable. They swim rectilinearly with substantial dynamic stability and efficiency. Aracanids have not been previously studied in these respects. This article describes swimming-related aspects of morphology, swimming modes, biomechanics, and kinematics in two south Australian species (striped cowfish and ornate cowfish) that are possibly representative of the entire group. These species differ morphologically in many respects, both from each other and from ostraciids. There are differences in numbers, sizes, and placements of keels on carapaces. The most important differences from ostraciids are openings in the posterior edges of the carapaces behind the dorsal and anal fins. The bases of those fins in ostraciids are enclosed in bone. The openings in aracanids free the fins and tail to move. As a result, aracanids are body and caudal fin swimmers. Their overall swimming performances are less stable, efficient, and effective. We propose establishing a new category of swimming mode for bony fishes called "aracaniform swimming."

Functional consequences of structural differences in stingray sensory systems. Part I: mechanosensory lateral line canals.

Short range hydrodynamic and electrosensory signals are important during final stages of prey capture in elasmobranchs (sharks, skates and rays), and may be particularly useful for dorso-ventrally flattened batoids with mouths hidden from their eyes. In stingrays, both the lateral line canal and electrosensory systems are highly modified and complex with significant differences on ventral surfaces that relate to feeding ecology. This study tests functional hypotheses based on quantified differences in sensory system morphology of three stingray species, Urobatis halleri, Myliobatis californica and Pteroplatytrygon violacea. Part I investigates the mechanosensory lateral line canal system whereas part II focuses on the electrosensory system. Stingray lateral line canals include both pored and non-pored sections and differ in branching complexity and distribution. A greater proportion of pored canals and high pore numbers were predicted to correspond to increased response to water flow. Behavioral experiments were performed to compare responses of stingrays to weak water jets mimicking signals produced by potential prey at velocities of 10-20 cm s(-1). Bat rays, M. californica, have the most complex and broadly distributed pored canal network and demonstrated both the highest response rate and greater response intensity to water jet signals. Results suggest that U. halleri and P. violacea may rely on additional sensory input, including tactile and visual cues, respectively, to initiate stronger feeding responses. These results suggest that stingray lateral line canal morphology can indicate detection capabilities through responsiveness to weak water jets.

Functional consequences of structural differences in stingray sensory systems. Part II: electrosensory system.

Elasmobranch fishes (sharks, skates and rays) possess highly sensitive electrosensory systems, which enable them to detect weak electric fields such as those produced by potential prey organisms. Different species have unique electrosensory pore numbers, densities and distributions. Functional differences in detection capabilities resulting from these structural differences are largely unknown. Stingrays and other batoid fishes have eyes positioned on the opposite side of the body from the mouth. Furthermore, they often feed on buried prey, which can be located non-visually using the electrosensory system. In the present study we test functional predictions based on structural differences in three stingray species (Urobatis halleri, Pteroplatytrygon violacea and Myliobatis californica) with differing electrosensory system morphology. We compare detection capabilities based upon behavioral responses to dipole electric signals (5.3-9.6 microA). Species with greater ventral pore numbers and densities were predicted to demonstrate enhanced electrosensory capabilities. Electric field intensities at orientation were similar among these species, although they differed in response type and orientation pathway. Minimum voltage gradients eliciting feeding responses were well below 1 nVcm(-1) for all species regardless of pore number and density.

Can systems biology help to separate evolutionary analogies (convergent homoplasies) from homologies?

Convergent evolutionary analogies (homoplasies) of many kinds occur in diverse phylogenetic clades/lineages on both the animal and plant branches of the Tree of Life. Living organisms whose last common ancestors lived millions to hundreds of millions of years ago have later converged morphologically, behaviorally or at other levels of functionality (from molecular genetics through biochemistry, physiology and other organismic processes) as a result of long term strong natural selection that has constrained and channeled evolutionary processes. This happens most often when organisms belonging to different clades occupy ecological niches, habitats or environments sharing major characteristics that select for a relatively narrow range of organismic properties. Systems biology, broadly defined, provides theoretical and methodological approaches that are beginning to make it possible to answer a perennial evolutionary biological question relating to convergent homoplasies: Are at least some of the apparent analogies actually unrecognized homologies? This review provides an overview of the current state of knowledge of important aspects of this topic area. It also provides a resource describing many homoplasies that may be fruitful subjects for systems biological research.

The greatest step in vertebrate history: a paleobiological review of the fish-tetrapod transition.

Recent discoveries of previously unknown fossil forms have dramatically transformed understanding of many aspects of the fish-tetrapod transition. Newer paleobiological approaches have also contributed to changed views of which animals were involved and when, where, and how the transition occurred. This review summarizes major advances made and reevaluates alternative interpretations of important parts of the evidence. We begin with general issues and concepts, including limitations of the Paleozoic fossil record. We summarize important features of paleoclimates, paleoenvironments, paleobiogeography, and taphonomy. We then review the history of Devonian tetrapods and their closest stem group ancestors within the sarcopterygian fishes. It is now widely accepted that the first tetrapods arose from advanced tetrapodomorph stock (the elpistostegalids) in the Late Devonian, probably in Euramerica. However, truly terrestrial forms did not emerge until much later, in geographically far-flung regions, in the Lower Carboniferous. The complete transition occurred over about 25 million years; definitive emergences onto land took place during the most recent 5 million years. The sequence of character acquisition during the transition can be seen as a five-step process involving: (1) higher osteichthyan (tetrapodomorph) diversification in the Middle Devonian (beginning about 380 million years ago [mya]), (2) the emergence of "prototetrapods" (e.g., Elginerpeton) in the Frasnian stage (about 372 mya), (3) the appearance of aquatic tetrapods (e.g., Acanthostega) sometime in the early to mid-Famennian (about 360 mya), (4) the appearance of "eutetrapods" (e.g., Tulerpeton) at the very end of the Devonian period (about 358 mya), and (5) the first truly terrestrial tetrapods (e.g., Pederpes) in the Lower Carboniferous (about 340 mya). We discuss each of these steps with respect to inferred functional utility of acquired character sets. Dissociated heterochrony is seen as the most likely process for the evolutionarily rapid morphological transformations required. Developmental biological processes, including paedomorphosis, played important roles. We conclude with a discussion of phylogenetic interpretations of the evidence.

Body-induced vortical flows: a common mechanism for self-corrective trimming control in boxfishes.

Boxfishes (Teleostei: Ostraciidae) are marine fishes having rigid carapaces that vary significantly among taxa in their shapes and structural ornamentation. We showed previously that the keels of the carapace of one species of tropical boxfish, the smooth trunkfish, produce leading edge vortices (LEVs) capable of generating self-correcting trimming forces during swimming. In this paper we show that other tropical boxfishes with different carapace shapes have similar capabilities. We conducted a quantitative study of flows around the carapaces of three morphologically distinct boxfishes (spotted boxfish, scrawled cowfish and buffalo trunkfish) using stereolithographic models and three separate but interrelated analytical approaches: digital particle image velocimetry (DPIV), pressure distribution measurements, and force balance measurements. The ventral keels of all three forms produced LEVs that grew in circulation along the bodies, resembling the LEVs produced around delta-winged aircraft. These spiral vortices formed above the keels and increased in circulation as pitch angle became more positive, and formed below the keels and increased in circulation as pitch angle became more negative. Vortices also formed along the eye ridges of all boxfishes. In the spotted boxfish, which is largely trapezoidal in cross section, consistent dorsal vortex growth posterior to the eye ridge was also present. When all three boxfishes were positioned at various yaw angles, regions of strongest concentrated vorticity formed in far-field locations of the carapace compared with near-field areas, and vortex circulation was greatest posterior to the center of mass. In general, regions of localized low pressure correlated well with regions of attached, concentrated vorticity, especially around the ventral keels. Although other features of the carapace also affect flow patterns and pressure distributions in different ways, the integrated effects of the flows were consistent for all forms: they produce trimming self-correcting forces, which we measured directly using the force balance. These data together with previous work on smooth trunkfish indicate that body-induced vortical flows are a common mechanism that is probably significant for trim control in all species of tropical boxfishes.

Flow Patterns Around the Carapaces of Rigid-bodied, Multi-propulsor Boxfishes (Teleostei: Ostraciidae).

Boxfishes (Teleostei: Ostraciidae) are rigid-body, multi-propulsor swimmers that exhibit unusually small amplitude recoil movements during rectilinear locomotion. Mechanisms producing the smooth swimming trajectories of these fishes are unknown, however. Therefore, we have studied the roles the bony carapaces of these fishes play in generating this dynamic stability. Features of the carapaces of four morphologically distinct species of boxfishes were measured, and anatomically-exact stereolithographic models of the boxfishes were constructed. Flow patterns around each model were investigated using three methods: 1) digital particle image velocimetry (DPIV), 2) pressure distribution measurements, and 3) force balance measurements. Significant differences in both cross-sectional and longitudinal carapace morphology were detected among the four species. However, results from the three interrelated approaches indicate that flow patterns around the various carapaces are remarkably similar. DPIV results revealed that the keels of all boxfishes generate strong longitudinal vortices that vary in strength and position with angle of attack. In areas where attached, concentrated vorticity was detected using DPIV, low pressure also was detected at the carapace surface using pressure sensors. Predictions of the effects of both observed vortical flow patterns and pressure distributions on the carapace were consistent with actual forces and moments measured using the force balance. Most notably, the three complementary experimental approaches consistently indicate that the ventral keels of all boxfishes, and in some species the dorsal keels as well, effectively generate self-correcting forces for pitching motions-a characteristic that is advantageous for the highly variable velocity fields in which these fishes reside.

Hydrodynamic stability of swimming in ostraciid fishes: role of the carapace in the smooth trunkfish Lactophrys triqueter (Teleostei: Ostraciidae).

The hydrodynamic bases for the stability of locomotory motions in fishes are poorly understood, even for those fishes, such as the rigid-bodied smooth trunkfish Lactophrys triqueter, that exhibit unusually small amplitude recoil movements during rectilinear swimming. We have studied the role played by the bony carapace of the smooth trunkfish in generating trimming forces that self-correct for instabilities. The flow patterns, forces and moments on and around anatomically exact, smooth trunkfish models positioned at both pitching and yawing angles of attack were investigated using three methods: digital particle image velocimetry (DPIV), pressure distribution measurements, and force balance measurements. Models positioned at various pitching angles of attack within a flow tunnel produced well-developed counter-rotating vortices along the ventro-lateral keels. The vortices developed first at the anterior edges of the ventro-lateral keels, grew posteriorly along the carapace, and reached maximum circulation at the posterior edge of the carapace. The vortical flow increased in strength as pitching angles of attack deviated from 0 degrees, and was located above the keels at positive angles of attack and below them at negative angles of attack. Variation of yawing angles of attack resulted in prominent dorsal and ventral vortices developing at far-field locations of the carapace; far-field vortices intensified posteriorly and as angles of attack deviated from 0 degrees. Pressure distribution results were consistent with the DPIV findings, with areas of low pressure correlating well with regions of attached, concentrated vorticity. Lift coefficients of boxfish models were similar to lift coefficients of delta wings, devices that also generate lift through vortex generation. Furthermore, nose-down and nose-up pitching moments about the center of mass were detected at positive and negative pitching angles of attack, respectively. The three complementary experimental approaches all indicate that the carapace of the smooth trunkfish effectively generates self-correcting forces for pitching and yawing motions--a characteristic that is advantageous for the highly variable velocity fields experienced by trunkfish in their complex aquatic environment. All important morphological features of the carapace contribute to producing the hydrodynamic stability of swimming trajectories in this species.