Biomedical subjects

Robert E Shadwick

Publications and source records attributed to Robert E Shadwick.

12 recordsLinked to original sources

PubMed2007

Reversibly labile, sclerotization-induced elastic properties in a keratin analog from marine snails: whelk egg capsule biopolymer (WECB).

Egg capsules from two caenogastropod whelks, Busycon canaliculatum and Kelletia kelletii, were studied to investigate the genesis of mechanical properties of nascent capsules and to formulate a biomechanical model of this material. Scanning electron microscopy revealed that the capsules possess fibrous hierarchical arrangements at all stages during processing while the mechanical integrity is developing. This suggests that an as yet uncharacterized sclerotization mechanism occurring in the ventral pedal gland primarily binds these fibrous components together. Decomposing the mechanical behavior of WECB through various physical and chemical treatments led us to develop a model for the structure and mechanical properties of this material that supports its designation as a keratin analog. Keratin mechanical models were applied to WECB in its representation as an intermediate state between matrix-free intermediate filament (IF)-type proteins and the more complex composite materials incorporating IFs such as keratin.

Animals↗

PubMed2006

Evolution of high-performance swimming in sharks: transformations of the musculotendinous system from subcarangiform to thunniform swimmers.

In contrast to all other sharks, lamnid sharks perform a specialized fast and continuous "thunniform" type of locomotion, more similar to that of tunas than to any other known shark or bony fish. Within sharks, it has evolved from a subcarangiform mode. Experimental data show that the two swimming modes in sharks differ remarkably in kinematic patterns as well as in muscle activation patterns, but the morphology of the underlying musculotendinous system (red muscles and myosepta) that drives continuous locomotion remains largely unknown. The goal of this study was to identify differences in the musculotendinous system of the two swimming types and to evaluate these differences in an evolutionary context. Three subcarangiform sharks (the velvet belly lantern shark, Etmopterus spinax, the smallspotted catshark, Scyliorhinus canicula, and the blackmouth catshark, Galeus melanostomus) from the two major clades (two galeans, one squalean) and one lamnid shark, the shortfin mako, Isurus oxyrhinchus, were compared with respect to 1) the 3D shape of myomeres and myosepta of different body positions; 2) the tendinous architecture (collagenous fiber pathways) of myosepta from different body positions; and 3) the association of red muscles with myoseptal tendons. Results show that the three subcarangiform sharks are morphologically similar but differ remarkably from the lamnid condition. Moreover, the "subcarangiform" morphology is similar to the condition known from teleostomes. Thus, major features of the "subcarangiform" condition in sharks have evolved early in gnathostome history: Myosepta have one main anterior-pointing cone and two posterior-pointing cones that project into the musculature. Within a single myoseptum cones are connected by longitudinally oriented tendons (the hypaxial and epaxial lateral and myorhabdoid tendons). Mediolaterally oriented tendons (epineural and epipleural tendons; mediolateral fibers) connect vertebral axis and skin. An individual lateral tendon spans only a short distance along the body (a fraction between 0.05 and 0.075 of total length, L, of the shark). This span is similar in all tendons along the body. Red muscles insert into the midregion of the lateral tendons. The shortfin mako differs substantially from this condition in several respects: Red muscles are internalized and separated from white muscles by a sheath of lubricative connective tissue. They insert into the anterior part of the hypaxial lateral tendon. Rostrocaudally, this tendon becomes very distinct and its span increases threefold (0.06L anteriorly to 0.19L posteriorly). Mediolateral fibers do not form distinct epineural/epipleural tendons in the mako. Since our morphological findings are in good accordance with experimental data it seems likely that the thunniform swimming mode has evolved along with the described morphological specializations.

Animals↗

PubMed2006

Kinematics of foraging dives and lunge-feeding in fin whales.

Fin whales are among the largest predators on earth, yet little is known about their foraging behavior at depth. These whales obtain their prey by lunge-feeding, an extraordinary biomechanical event where large amounts of water and prey are engulfed and filtered. This process entails a high energetic cost that effectively decreases dive duration and increases post-dive recovery time. To examine the body mechanics of fin whales during foraging dives we attached high-resolution digital tags, equipped with a hydrophone, a depth gauge and a dual-axis accelerometer, to the backs of surfacing fin whales in the Southern California Bight. Body pitch and roll were estimated by changes in static gravitational acceleration detected by orthogonal axes of the accelerometer, while higher frequency, smaller amplitude oscillations in the accelerometer signals were interpreted as bouts of active fluking. Instantaneous velocity of the whale was determined from the magnitude of turbulent flow noise measured by the hydrophone and confirmed by kinematic analysis. Fin whales employed gliding gaits during descent, executed a series of lunges at depth and ascended to the surface by steady fluking. Our examination of body kinematics at depth reveals variable lunge-feeding behavior in the context of distinct kinematic modes, which exhibit temporal coordination of rotational torques with translational accelerations. Maximum swimming speeds during lunges match previous estimates of the flow-induced pressure needed to completely expand the buccal cavity during feeding.

Acceleration↗

PubMed2005

Mammal-like muscles power swimming in a cold-water shark.

Effects of temperature on muscle contraction and powering movement are profound, outwardly obvious, and of great consequence to survival. To cope with the effects of environmental temperature fluctuations, endothermic birds and mammals maintain a relatively warm and constant body temperature, whereas most fishes and other vertebrates are ectothermic and conform to their thermal niche, compromising performance at colder temperatures. However, within the fishes the tunas and lamnid sharks deviate from the ectothermic strategy, maintaining elevated core body temperatures that presumably confer physiological advantages for their roles as fast and continuously swimming pelagic predators. Here we show that the salmon shark, a lamnid inhabiting cold, north Pacific waters, has become so specialized for endothermy that its red, aerobic, locomotor muscles, which power continuous swimming, seem mammal-like, functioning only within a markedly elevated temperature range (20-30 degrees C). These muscles are ineffectual if exposed to the cool water temperatures, and when warmed even 10 degrees C above ambient they still produce only 25-50% of the power produced at 26 degrees C. In contrast, the white muscles, powering burst swimming, do not show such a marked thermal dependence and work well across a wide range of temperatures.

Animals↗

PubMed2005

Fast-start muscle dynamics in the rainbow trout Oncorhynchus mykiss: phase relationship of white muscle shortening and body curvature.

Muscle length changes of the lateral myotomal fast fibers of rainbow trout (Oncorhynchus mykiss) were measured using sonomicrometry during induced fast-starts. Simultaneous high-speed videography allowed for the analysis of midline kinematics to estimate the degree of muscle strain that occurs during body deformation. Comparison of these data was used to examine the phase relationship between local muscle shortening and local body bending during unsteady, large amplitude maneuvers. Our analysis finds that muscle shortening is temporally decoupled from body bending, probably due to the influence of hydrodynamic forces. The phase shift was such that midline curvature lagged behind muscle shortening at both the anterior (0.4 L, where L is fork length) and posterior (0.7 L) axial positions. Stronger escape responses were correlated with high peak strains and rapid strain-wave velocities, but not faster curvature-wave velocities. Under these conditions of high strain, the phase shift at the posterior position is significantly increased, whereas the anterior position fails to be affected. Curvature lag was still observed at both axial locations under conditions of low strain, suggesting that hydrodynamic forces are still significant during weaker escape responses. These data support a previous model that suggests fast-start body bending is determined by the interaction between muscle torque and hydrodynamic resistance along the body.

Animals↗

PubMed2005

Patterns of red muscle strain/activation and body kinematics during steady swimming in a lamnid shark, the shortfin mako (Isurus oxyrinchus).

The dynamics of steady swimming were examined in the shortfin mako (Isurus oxyrinchus), a member of the cartilaginous fish family Lamnidae, a family known for their morphological adaptations for high-performance locomotion and their similarity in hydromechanical design to tunas. Patterns of red muscle (RM) strain (i.e. relative length change) and activation were quantified at two axial positions ( approximately 0.4 and 0.6L, where L is total body length), using sonomicrometry and electromyography (EMG), and correlated with simultaneous measurements of dorsal midline kinematics during steady swimming ( approximately 0.5-1 L s(-1)). RM strain varied longitudinally with strain amplitudes ranging from 5.5+/-1.1% (s.e.m.) in the anterior to 8.7+/-0.9% in the posterior. We found no significant longitudinal variation in patterns of RM activation, with mean onset of activation occurring at 83-84 degrees (90 degrees is peak length) and offset at 200-210 degrees at both body positions. Likewise, duty cycles were similar: 35.5+/-1.0% in the anterior and 32.2+/-1.6% in the posterior. Comparison of the timing of waves of dorsal midline curvature and predicted strain relative to measured RM strain revealed a phase shift between RM shortening and local body bending. Furthermore, when the body is bent passively, RM shortens synchronously with the surrounding white muscle (WM) and skin, as expected. During active swimming, peaks in RM strain were delayed relative to peaks in WM strain by a mean of approximately 10% of the tailbeat cycle, with one individual as high as approximately 17% in the anterior and nearly 50% in the posterior. The longitudinal consistency in the EMG/strain phase relationship in the mako is similar to that in the leopard shark, suggesting a consistent trend among sharks using different locomotor modes. However, unlike in the leopard shark, RM shortening in the mako is physically uncoupled from deformation of the surrounding body during steady swimming, a characteristic shared between the mako and tunas.

Animals↗

PubMed2005

Cuvier's beaked whale (Ziphius cavirostris) head tissues: physical properties and CT imaging.

Tissue physical properties from a Cuvier's beaked whale (Ziphius cavirostris) neonate head are reported and compared with computed tomography (CT) X-ray imaging. Physical properties measured include longitudinal sound velocity, density, elastic modulus and hysteresis. Tissues were classified by type as follows: mandibular acoustic fat, mandibular blubber, forehead acoustic fat (melon), forehead blubber, muscle and connective tissue. Results show that each class of tissues has unique, co-varying physical properties. The mandibular acoustic fats had minimal values for sound speed (1350+/-10.6 m s(-1)) and mass density (890+/-23 kg m(-3)). These values increased through mandibular blubber (1376+/-13 m s(-1), 919+/-13 kg m(-3)), melon (1382+/-23 m s(-1), 937+/-17 kg m(-3)), forehead blubber (1401+/-7.8 m s(-1), 935+/-25 kg m(-3)) and muscle (1517+/-46.8 m s(-1), 993+/-58 kg m(-3)). Connective tissue had the greatest mean sound speed and density (1628+/-48.7 m s(-1), 1087+/-41 kg m(-3)). The melon formed a low-density, low-sound-speed core, supporting its function as a sound focusing organ. Hounsfield unit (HU) values from CT X-ray imaging are correlated with density and sound speed values, allowing HU values to be used to predict these physical properties. Blubber and connective tissues have a higher elastic modulus than acoustic fats and melon, suggesting more collagen structure in blubber and connective tissues. Blubber tissue elastic modulus is nonlinear with varying stress, becoming more incompressible as stress is increased. These data provide important physical properties required to construct models of the sound generation and reception mechanisms in Ziphius cavirostris heads, as well as models of their interaction with anthropogenic sound.

Adipose Tissue↗

PubMed2004

Convergent evolution in mechanical design of lamnid sharks and tunas.

The evolution of 'thunniform' body shapes in several different groups of vertebrates, including whales, ichthyosaurs and several species of large pelagic fishes supports the view that physical and hydromechanical demands provided important selection pressures to optimize body design for locomotion during vertebrate evolution. Recognition of morphological similarities between lamnid sharks (the most well known being the great white and the mako) and tunas has led to a general expectation that they also have converged in their functional design; however, no quantitative data exist on the mechanical performance of the locomotor system in lamnid sharks. Here we examine the swimming kinematics, in vivo muscle dynamics and functional morphology of the force-transmission system in a lamnid shark, and show that the evolutionary convergence in body shape and mechanical design between the distantly related lamnids and tunas is much more than skin deep; it extends to the depths of the myotendinous architecture and the mechanical basis for propulsive movements. We demonstrate that not only have lamnids and tunas converged to a much greater extent than previously known, but they have also developed morphological and functional adaptations in their locomotor systems that are unlike virtually all other fishes.

Animals↗

PubMed2003

Steady swimming muscle dynamics in the leopard shark Triakis semifasciata.

Patterns of red muscle strain and activation were examined at three positions along the body (0.42, 0.61 and 0.72 L, where L is total body length) and correlated with simultaneous measurements of midline kinematics during steady swimming (approx. 1.0 L s(-1)) in the leopard shark Triakis semifasciata. Analysis of lateral displacement along the body indicates that the leopard shark is a subcarangiform swimmer. Longitudinal variation in red muscle strain was observed with strain amplitudes ranging from +/-3.9% in the anterior, +/-6.6% in the mid, to +/-4.8% in the posterior body position. Strain was in-phase with local midline curvature. In addition, strain amplitude calculated from a bending beam model closely matched strain measured using sonomicrometry at all three body positions. There is a high degree of similarity in red muscle activation patterns along the body between the leopard shark and many fish species, in that the onset of activation occurs during muscle lengthening while offset occurs during muscle shortening. However, we found no significant longitudinal variation in the EMG/strain phase relationship and duty cycles, with onset of muscle activation occurring at 51.4-61.8 degrees and offset at 159.7-165.2 degrees (90 degrees is peak length). This consistent pattern of activation suggests that red muscle along the entire length of the body contributes to positive power production. Thus, sharks such as Triakis may have no regional specialization in red muscle function like that seen in many teleosts, which may indicate that the evolution of differential muscle function along the body occurred after the divergence of cartilaginous and bony fishes.

Animals↗

PubMed2002

Structure and function of tuna tail tendons.

The caudal tendons in tunas and other scombrid fish link myotomal muscle directly to the caudal fin rays, and thus serve to transfer muscle power to the hydrofoil-like tail during swimming. These robust collagenous tendons have structural and mechanical similarity to tendons found in other vertebrates, notably the leg tendons of terrestrial mammals. Biochemical studies indicate that tuna tendon collagen is composed of the (alpha1)(2),alpha2 heterotrimer that is typical of vertebrate Type I collagen, while tuna skin collagen has the unusual alpha1,alpha2,alpha3 trimer previously described in the skin of some other teleost species. Tuna collagen, like that of other fish, has high solubility due to the presence of an acid-labile intermolecular cross-link. Unlike collagen in mammalian tendons, no differences related to cross-link maturation were detected among tendons in tuna ranging from 0.05 to 72 kg (approx. 0.25-6 years). Tendons excised post-mortem were subjected to load cycling to determine the modulus of elasticity and resilience (mean of 1.3 GPa and 90%, respectively). These material properties compare closely to those of leg tendons from adult mammals that can function as effective biological springs in terrestrial locomotion, but the breaking strength is substantially lower. Peak tendon forces recorded during steady swimming appear to impose strains of much less than 1% of tendon length, and no more than 1.5% during bursts. Thus, the caudal tendons in tunas do not appear to function as elastic storage elements, even at maximal swimming effort.

Aging↗

PubMed2002

Effects of longitudinal body position and swimming speed on mechanical power of deep red muscle from skipjack tuna (Katsuwonus pelamis).

The mechanical power output of deep, red muscle from skipjack tuna (Katsuwonus pelamis) was studied to investigate (i) whether this muscle generates maximum power during cruise swimming, (ii) how the differences in strain experienced by red muscle at different axial body locations affect its performance and (iii) how swimming speed affects muscle work and power output. Red muscle was isolated from approximately mid-way through the deep wedge that lies next to the backbone; anterior (0.44 fork lengths, ANT) and posterior (0.70 fork lengths, POST) samples were studied. Work and power were measured at 25 degrees C using the work loop technique. Stimulus phases and durations and muscle strains (+/- 5.5 % in ANT and +/- 8 % in POST locations) experienced during cruise swimming at different speeds were obtained from previous studies and used during work loop recordings. In addition, stimulus conditions that maximized work were determined. The stimulus durations and phases yielding maximum work decreased with increasing cycle frequency (analogous to tail-beat frequency), were the same at both axial locations and were almost identical to those used by the fish during swimming, indicating that the muscle produces near-maximal work under most conditions in swimming fish. While muscle in the posterior region undergoes larger strain and thus produces more mass-specific power than muscle in the anterior region, when the longitudinal distribution of red muscle mass is considered, the anterior muscles appear to contribute approximately 40% more total power. Mechanical work per length cycle was maximal at a cycle frequency of 2-3 Hz, dropping to near zero at 15 Hz and by 20-50% at 1 Hz. Mechanical power was maximal at a cycle frequency of 5 Hz, dropping to near zero at 15 Hz. These fish typically cruise with tail-beat frequencies of 2.8-5.2 Hz, frequencies at which power from cyclic contractions of deep red muscles was 75-100% maximal. At any given frequency over this range, power using stimulation conditions recorded from swimming fish averaged 93.4 +/- 1.65% at ANT locations and 88.6 +/- 2.08% at POST locations (means +/- S.E.M., N=3-6) of the maximum using optimized conditions. When cycle frequency was held constant (4 Hz) and strain amplitude was increased, work and power increased similarly in muscles from both sample sites; work and power increased 2.5-fold when strain was elevated from +/- 2 to +/- 5.5%, but increased by only approximately 12% when strain was raised further from +/- 5.5 to +/- 8%. Taken together, these data suggest that red muscle fibres along the entire body are used in a similar fashion to produce near-maximal mechanical power for propulsion during normal cruise swimming. Modelling suggests that the tail-beat frequency at which power is maximal (5 Hz) is very close to that used at the predicted maximum aerobic swimming speed (5.8 Hz) in these fish.

Animals↗

PubMedDate not supplied

Mechanical characterization of an unusual elastic biomaterial from the egg capsules of marine snails (Busycon spp.).

Egg capsule material serves as a putative protection mechanism for developing snail embryos facing the perils of the marine environment. We conducted the first quantitative study of this acellular structural protein with the goals of characterizing its chemical and mechanical properties and the relationship of these properties to its biological protective function. We have found that this protein polymer exhibits long-range elasticity with an interesting recoverable yield evidenced by an order of magnitude decrease in elastic modulus (apparent failure) that begins at 3%-5% strain. This material differs significantly from other common structural proteins such as collagen and elastin in mechanical response to strain. Qualitative similarities in stress/strain behavior to keratin, another common structural protein, are more than coincidental when composition and detailed mechanical quantification are considered. This suggests the possibility of alpha-helical structure and matrix organization that might be similar in these two proteins. Indeed, the egg capsule protein may be closely related to vertebrate keratins such as intermediate filaments. We conclude that while this material's bimodal tensile properties may serve as useful protection against the impact loading egg capsules encounter in the intertidal zone, the full biological importance of these capsules is not known.

Amino Acids↗