Emersion and Terrestrial Locomotion of the Northern Snakehead (Channa argus) on Multiple Substrates.

Most fishes known for terrestrial locomotion are small and/or elongate. Northern snakeheads (Channa argus) are large, air-breathing piscivores anecdotally known for terrestrial behaviors. Our goals were to determine their environmental motivations for emersion, describe their terrestrial kinematics for fish 3.0-70.0 cm and compare kinematics among four substrates. For emersion experiments, C. argus was individually placed into aquatic containers with ramps extending through the surface of the water, and exposed to 15 ecologically-relevant environmental conditions. For kinematic experiments, fish were filmed moving on moist bench liner, grass, artificial turf, and a flat or tilted rubber boat deck. Videos were digitized for analysis in MATLAB and electromyography was used to measure muscular activity. Only the low pH (4.8), high salinity (30 ppt), and high dCO2 (10% seltzer solution) treatments elicited emersion responses. While extreme, these conditions do occur in some of their native Asian swamps. Northern snakeheads >4.5 cm used a unique form of axial-appendage-based terrestrial locomotion involving cyclic oscillations of the axial body, paired with near-simultaneous movements of both pectoral fins. Individuals ≤3.5 cm used tail-flip jumps to travel on land. Northern snakeheads also moved more quickly on complex, three-dimensional substrates (e.g., grass) than on smooth substrates (e.g., bench liner), and when moving downslope. Release of snakeheads onto land by humans or accidentally by predators may be more common than voluntary emersion, but because northern snakeheads can respire air, it may be necessary to factor in the ability to spread overland into the management of this invasive species.

Emersión y locomoción terrestre de la cabeza de serpiente del norte (Channa argus) en múltiples sustratos (Emersion and terrestrial locomotion of the northern snakehead (Channa argus) on multiple substrates) La mayoría de los peces conocidos por locomoción terrestre son pequeños y/o alargados. Las cabezas de serpiente del norte (Channa argus) son grandes pesces piscívoros que respiran aire, anecdóticamente conocidos por sus comportamientos terrestres. Nuestros objetivos fueron determinar sus motivaciones ambientales para la emersión, describir su cinemática terrestre para peces de 3, 0 a 70, 0 cm y comparar la cinemática entre cuatro sustratos. Para los experimentos de emersión, C. argus se colocó individualmente en contenedores acuáticos con rampas que se extienden a través de la superficie del agua y fueron expuesto a quince condiciones ambientales ecológicamente pertinentes. Para los experimentos cinemáticos, los peces se filmaron moviéndose sobre un revestimiento de banco húmedo, césped, césped artificial y una cubierta de bote de goma plana o inclinada. Los videos se digitalizaron para su análisis en MATLAB y se usó electromiografía para medir la actividad muscular. Solo los tratamientos de bajo pH (4, 8), alta salinidad (30 partes por mil) y alto dCO2 (solución de agua de Seltz 10%) provocaron respuestas de emersión. Aunque son extremas, estas condiciones si ocurren en algunos de sus pantanos asiáticos nativos. Las cabezas de serpiente del norte >4, 5 cm usaron una forma única de locomoción terrestre basada en movimientos apéndiculares-axiales que involucra oscilaciones cíclicas del cuerpo axial, junto con movimientos casi simultáneos de ambas aletas pectorales. Los individuos de ≤3, 5 cm usaron saltos de cola para moverse en tierra. Las cabezas de serpiente del norte también se movían más rápidamente en sustratos tridimensionales complejos (ej., césped) que en sustratos lisos (ej., revestimiento de banco), y al moverse cuesta abajo. La liberación de cabezas de serpiente en la tierra por humanos o accidentalmente por depredadores puede ser más común que la emersión voluntaria, pero debido a que las cabezas de serpiente del norte pueden respirar aire, puede ser necesario tener en cuenta la capacidad de propagarse por tierra en el manejo de esta especie invasora. Translated to Spanish by YE Jimenez (yordano_jimenez@brown.edu).

The effect of fiber-type heterogeneity on optimized work and power output of hindlimb muscles of the salamander Ambystoma tigrinum.

Most vertebrate muscles are composed of a mixture of fiber types. However, studies of muscle mechanics have concentrated on homogeneous bundles of fibers. Hindlimb muscles of the tiger salamander, Ambystoma tigrinum, present an excellent system to explore the consequences of fiber heterogeneity. Isometric twitches and work loops were obtained in vitro from two muscles, the m. iliotibialis pars posterior (heterogeneous, containing types I, IIa and IIb fibers) and the m. iliofibularis (nearly homogeneous for type IIa fibers). Maximal isometric twitch and tetanic stresses in m. iliotibialis posterior were significantly greater than in iliofibularis. Work loops were obtained over a range of frequencies (0.5-3.0 Hz) and strains (2-6% muscle length) that encompassed the observed ranges in vivo. Work per cycle from the homogeneous iliofibularis declined from 1.5-3.0 Hz, while that from the heterogeneous m. iliotibialis posterior increased from 0.5 Hz to 2.5 Hz and declined at 3.0 Hz. Power output from the iliofibularis rose with frequency to at least 3 Hz; power from the iliotibialis posterior rose with frequency to 2.5 Hz and declined thereafter. Mass-specific work per cycle and power output were higher in iliofibularis than iliotibialis posterior over most frequencies and strains tested.

Fetlock joint kinematics differ with age in Thoroughbred [was thoroughbred] racehorses.

Fetlock joint kinematics during galloping in 2-, 3-, 4-, and 5-year-old Thoroughbreds in race training were quantified to determine if differences due to age could account for the observation that 2-year old Thoroughbred racehorses incur a high number of injuries to the bones and soft tissues in the distal forelimbs during training and at the outset of racing. Twelve Thoroughbred racehorses were videotaped in the sagittal plane at 250 frames/s during their daily galloping workout on a 7/8 mile sand-surface training track. Four galloping strides were recorded for each horse and subsequently digitized to determine fetlock joint angles of the leading forelimb during the limb support period of a stride. Four kinematic variables were measured from each stride's angular profile: angle of fetlock joint dorsi-flexion at mid-stance, negative angular velocity, positive angular velocity and time from hoof impact to mid-stance phase of limb support. The 2-year old Thoroughbreds had significantly quicker rates of dorsi-flexion of their fetlock joints than 3- (p=0.01), 4- (p=0.01), and 5-year old (p<0.01) Thoroughbreds following impact of the leading forelimb during moderate galloping (avg. 14 m/s). Higher rates of dorsi-flexion in young Thoroughbreds may reflect immaturity (lack of stiffness) of the suspensory apparatus tissues.

Trading force for speed: why superfast crossbridge kinetics leads to superlow forces.

Superfast muscles power high-frequency motions such as sound production and visual tracking. As a class, these muscles also generate low forces. Using the toadfish swimbladder muscle, the fastest known vertebrate muscle, we examined the crossbridge kinetic rates responsible for high contraction rates and how these might affect force generation. Swimbladder fibers have evolved a 10-fold faster crossbridge detachment rate than fast-twitch locomotory fibers, but surprisingly the crossbridge attachment rate has remained unchanged. These kinetics result in very few crossbridges being attached during contraction of superfast fibers (only approximately 1/6 of that in locomotory fibers) and thus low force. This imbalance between attachment and detachment rates is likely to be a general mechanism that imposes a tradeoff of force for speed in all superfast fibers.

Motor patterns and kinematics during backward walking in the pacific giant salamander: evidence for novel motor output.

Kinematic and motor patterns during forward and backward walking in the salamander Dicamptodon tenebrosus were compared to determine whether the differences seen in mammals also apply to a lower vertebrate with sprawling posture and to measure the flexibility of motor output by tetrapod central pattern generators. During treadmill locomotion, electromyograms (EMGs) were recorded from hindlimb muscles of Dicamptodon while simultaneous high-speed video records documented movement of the body, thigh, and crus and allowed EMGs to be synchronized to limb movements. In forward locomotion, the trunk was lifted above the treadmill surface. The pelvic girdle and trunk underwent smooth side-to-side oscillations throughout the stride. At the beginning of the stance phase, the femur was protracted and the knee joint extended. The knee joint initially flexed in early stance and then extended as the foot pushed off in late stance, reaching maximum extension just before foot lift-off. The femur retracted steadily throughout the stance. In the swing phase, the femur rapidly protracted, and the leg was brought forward in an "overhand crawl" motion. In backward walking, the body frequently remained in contact with the treadmill surface. The pelvic girdle, trunk, and femur remained relatively still during stance phase, and most motion occurred at the knee joint. The knee joint extended throughout most of stance, as the body moved back, away from the stationary foot. The knee flexed during swing. Four of five angles showed significantly smaller ranges in backward than in forward walking. EMGs of forward walking showed that ventral muscles were coactive, beginning activity just before foot touchdown and ceasing during the middle of stance phase. Dorsal muscles were active primarily during swing. Backward locomotion showed a different pattern; all muscles except one showed primary activity during the swing phase. This pattern of muscle synergy in backward walking never was seen in forward locomotion. Also, several muscles demonstrated lower burst rectified integrated areas (RIA) or durations during backward locomotion. Multivariate statistical analysis of EMG onset and RIA completely separated forward and backward walking along the first principal component, based on higher RIAs, longer durations of muscle activity, and greater synergy between ventral muscles during early stance in forward walking. Backward walking in Dicamptodon uses a novel motor pattern not seen during forward walking in salamanders or during any other locomotor activity in previously studied tetrapods. The central neuronal mechanisms mediating locomotion in this primitive tetrapod are thus capable of considerable plasticity.

HINDLIMB KINEMATICS DURING TERRESTRIAL LOCOMOTION IN A SALAMANDER (DICAMPTODON TENEBROSUS)

A quantitative study of hindlimb kinematics during terrestrial locomotion in a non-specialized salamander was undertaken to allow comparisons with limb movements in other groups of tetrapods. Five Dicamptodon tenebrosus were videotaped at 200 fields s-1 walking on a treadmill. Coordinates of marker points on the salamander's midline, pelvic girdle and left hindlimb were digitized through at least three strides at both a walk (0.77 SVL s-1, where SVL is snout­vent length) and a trot (2.90 SVL s-1). Marker coordinates were used to compute kinematic variables summarizing trunk flexion, pelvic girdle rotation, femoral protraction/retraction and knee flexion/extension. The stride is characterized by uninterrupted trunk and pelvic girdle oscillation, femoral retraction throughout stance phase, and knee flexion in early stance followed by extension. Mean angular excursions are: trunk, 66 °; pelvic girdle, 38.5 °; pelvic girdle­femur, 106 °; and knee, 65 °. The hindlimb and pelvic girdle also show a complicated pattern of lateral movement related to knee flexion/extension and periods of support by the contralateral hindlimb during the step cycle. Dicamptodon shares the following features of the hindlimb step cycle with other tetrapod taxa: rotation of the pelvic girdle through a 30­40 ° arc, femoral retraction beginning simultaneously with and persisting throughout stance phase, flexion of the knee in early stance, and extension of the knee in late stance.

METAMORPHIC AND SPEED EFFECTS ON HINDLIMB KINEMATICS DURING TERRESTRIAL LOCOMOTION IN THE SALAMANDER DICAMPTODON TENEBROSUS

The kinematics of the hindlimb during terrestrial treadmill locomotion in Dicamptodon tenebrosus were compared between larval and metamorphosed individuals at different speeds. Coordinates of marker points on the salamander's midline, pelvic girdle and left hindlimb were digitized from high-speed videos (200 fields s-1). These yielded kinematic variables describing trunk flexion, pelvic girdle rotation, femoral protraction/retraction and knee flexion/extension. A three-way analysis of variance tested for mean differences among individuals, speeds and metamorphic stages for each variable. No significant overall effects of metamorphosis were found, although several variables showed significant stage x individual effects. Multivariate analyses revealed that the variance in kinematics of the larvae was significantly greater than that of the metamorphosed salamanders. Several variables showed significant speed effects or strong trends, among them stride length (increases with speed), cycle duration (decreases), contact interval (decreases) and phase variables describing the relative timing between minimum/maximum angles and the beginning of stance/swing phase. Such changes with speed are consistent with those shown for diverse arthropods and tetrapods and suggest that changes in stride length and timing events during a stride represent a general mechanism for effecting an increase in locomotor speed.