Anna N Ahn

Walking and Running in the Red-Legged Running Frog, Kassina Maculata

SUMMARY Although most frog species are specialized for jumping or swimming, Kassina maculata (red-legged running frog) primarily uses a third type of locomotion during which the hindlimbs alternate. In the present study, we examined Kassinas distinct locomotory mode to determine whether these frogs walk or run and how their gait may change with speed. We used multiple methods to distinguish between terrestrial gaits: the existence or absence of an aerial phase, duty factor, relative footfall patterns and the mechanics of the animals center of mass (COM). To measure kinematic and kinetic variables, we recorded digital video as the animals moved over a miniature force platform (N=12 individuals). With respect to footfall patterns, the frogs used a single gait and walked at all speeds examined. Duty factor always exceeded 0.59. Based on COM mechanics, however, the frogs used both walking and running gaits. At slower speeds, the fluctuations in the horizontal kinetic energy (Ek) and gravitational potential energy (Ep) of the COM were largely out of phase, indicating a vaulting or walking gait. In most of the trials, Kassina used a combined gait at intermediate speeds, unlike cursorial animals with distinct gait transitions. This combined gait, much like a mammalian gallop, exhibited the mechanics of both vaulting and bouncing gaits. At faster speeds, the Ek and Ep of Kassinas COM were more in phase, indicating the use of a bouncing or running gait. Depending on the definition used to distinguish between walking and running, Kassina either only used a walking gait at all speeds or used a walking gait at slower speeds but then switched to a running gait as speed increased.

In Vivo and In Vitro Heterogeneity of Segment Length Changes in the Semimembranosus Muscle of the Toad

Many studies examine sarcomere dynamics in single fibres or length–tension dynamics in whole muscles in vivo or in vitro, but few studies link the various levels of organisation. To relate data addressing in vitro muscle segment behaviour with in vivo whole muscle behaviour during locomotion, we measured in vivo strain patterns of muscle segments using three sonomicrometry crystals implanted along a fascicle of the semimembranosus muscle in the American toad (Bufo americanus; n= 6) during hopping. The centre crystal emitted an ultrasonic signal, while the outer crystals received the signal allowing the instantaneous measurement of lengths from two adjacent muscle segments. On the first day, we recorded from the central and distal segments. On the second day of recordings, the most distal crystal was moved to a proximal position to record from a proximal segment and the same central segment. When the toads hopped a distance of two body lengths, the proximal and central segments strained −15.1 ± 6.1 and −14.0 ± 4.9 % (i.e. shortening), respectively. Strain of the distal segment, however, was significantly lower and more variable in pattern, often lengthening before shortening during a hop. From rest length, the distal segment initially lengthened by 2.6 ± 2.0 % before shortening by 6.5 ± 3.2 % at the same hop distance. Under in vitro conditions, the central segment always shortened more than the distal segment, except when passively cycled, during which the segments strained similarly. When the whole muscle was cycled sinusoidally and stimulated phasically in vitro, the two adjacent segments strained in opposite directions over much (up to 34 %) of the cycle. These differences in strain amplitude and direction imply that two adjacent segments can not only produce and/or absorb varying amounts of mechanical energy, but can also operate on different regions of their force–length and force–velocity relationships when activated by the same neural signal. Understanding regional differences in contractile dynamics within muscles is therefore important to linking our understanding of sarcomere behaviour with whole muscle behaviour during locomotion.

In situ muscle power differs without varying in vitro mechanical properties in two insect leg muscles innervated by the same motor neuron.

SUMMARY The mechanical behavior of muscle during locomotion is often predicted by its anatomy, kinematics, activation pattern and contractile properties. The neuromuscular design of the cockroach leg provides a model system to examine these assumptions, because a single motor neuron innervates two extensor muscles operating at a single joint. Comparisons of the in situ measurements under in vivo running conditions of muscle 178 to a previously examined muscle (179) demonstrate that the same inputs (e.g. neural signal and kinematics) can result in different mechanical outputs. The same neural signal and kinematics, as determined during running, can result in different mechanical functions, even when the two anatomically similar muscles possess the same contraction kinetics, force-velocity properties and tetanic force-length properties. Although active shortening greatly depressed force under in vivo-like strain and stimulation conditions, force depression was similarly proportional to strain, similarly inversely proportional to stimulation level, and similarly independent of initial length and shortening velocity between the two muscles. Lastly, passive pre-stretch enhanced force similarly between the two muscles. The forces generated by the two muscles when stimulated with their in vivo pattern at lengths equal to or shorter than rest length differed, however. Overall, differences between the two muscles in their submaximal force-length relationships can account for up to 75% of the difference between the two muscles in peak force generated at short lengths observed during oscillatory contractions. Despite the fact that these muscles act at the same joint, are stimulated by the same motor neuron with an identical pattern, and possess many of the same in vitro mechanical properties, the mechanical outputs of two leg extensor muscles can be vastly different.

Moderate Dehydration Decreases Locomotor Performance of the Ghost Crab, Ocypode quadrata

The effect of dehydration on the aerobic metabolism and endurance of sustained, terrestrial locomotion was determined for the ghost crab, Ocypode quadrata. The rate of evaporative water loss, measured as the percentage of decrease in body mass per hour, was influenced by ambient temperature (Ta). Increasing Ta from 24°C to 30°C (40%-50% relative humidity) increased the rate of water loss from 2.3% h⁻¹ ± 0.2% h⁻¹ to 3.6% h⁻¹ ± 0.6% h⁻¹. Crabs were divided into three treatment groups to determine the effect of dehydration on aerobic metabolism: hydrated control crabs, slowly dehydrated crabs, and rapidly dehydrated crabs. Hydrated control crabs lost only 1.2% of their initial body mass. Slowly dehydrated crabs were dehydrated by 3.6% of their initial body mass at a rate of 2.3% h⁻¹. Finally, rapidly dehydrated crabs were dehydrated by 3.6% of their initial body mass at a rate of 3.6% h⁻¹. The maximal rate of oxygen consumption (V̇o2max) determined during treadmill exercise was decreased by 30% for slowly dehydrated crabs and by 70% for rapidly dehydrated crabs, as compared to hydrated controls. The minimum cost of locomotion was independent of the dehydration state for hydrated and slowly dehydrated crabs but was 62% lower for rapidly dehydrated crabs. Endurance was correlated with the speed at which V̇o2max was attained (the maximum aerobic speed [MAS]). The MAS was highest for hydrated control crabs and was decreased by 32% for slowly dehydrated crabs and by 68% for rapidly dehydrated crabs. We conclude that moderate dehydration can substantially decrease the ghost crab s capacity for sustained, terrestrial locomotion.

Kinematic Evidence for Superfast Locomotory Muscle in Two Species of Teneriffiid Mites

SUMMARY Locomotory muscles typically operate over a narrow range of contraction frequencies, characterized by the predominant fiber types and functional roles. The highest documented frequencies in the synchronous sound-producing muscles of insects (550 Hz) and toadfish (200 Hz) far exceed the contraction frequencies observed in weight-bearing locomotory muscles, which have maximum documented frequencies below 15–30 Hz. Laws of scaling, however, predict that smaller arthropods may employ stride frequencies exceeding this range. In this study we measured running speed and stride frequency in two undescribed species of teneriffiid mites from the coastal sage scrub of southern California. Relative speeds of both species [129–133 body lengths (BL)s–1] are among the fastest documented for any animal. Measured stride frequencies for both species far exceed those documented for any weight-bearing locomotory muscle, with measured values for one species ranging from 93 Hz at 25°C to 111 Hz at 45°C. Stride frequencies either closely approximate or, for one species, exceed predicted values based on an interspecific scaling of frequency and animal mass. Consequently, while the ultra-high frequencies of these muscles must depend on appropriately scaled kinetics of the calcium transient and contraction–relaxation cycle, these do not appear to limit the operating frequencies during running. The predicted low muscle forces operating at these very high frequencies evidently suffice for locomotion, probably because of the larger relative muscle force generated by smaller animals.

HOW MUSCLES FUNCTION – THE WORK LOOP TECHNIQUE

![Figure][1] Anna Ahn discusses Bob Josephson’s 1985 paper entitled: ‘Mechanical power output from striated muscle during cyclic contraction’. A copy of the paper can be obtained from The primary function of muscle is to generate

Effect of temperature on leg kinematics in sprinting tarantulas (Aphonopelma hentzi): high speed may limit hydraulic joint actuation.

ABSTRACT Tarantulas extend the femur–patella (proximal) and tibia–metatarsal (distal) joints of their legs hydraulically. Because these two hydraulically actuated joints are positioned in series, hemolymph flow within each leg is expected to mechanically couple the movement of the joints. In the current study, we tested two hypotheses: (1) at lower temperatures, movement of the two in-series hydraulic joints within a leg will be less coupled because of increased hemolymph viscosity slowing hemolymph flow; and (2) at higher temperatures, movement of the two in-series hydraulic joints will be less coupled because the higher stride frequencies limit the time available for hemolymph flow. We elicited maximal running speeds at four ecologically relevant temperatures (15, 24, 31 and 40°C) in Texas Brown tarantulas (Aphonopelma hentzi). The spiders increased sprint speed 2.5-fold over the temperature range by changing their stride frequency but not stride length. The coefficient of determination for linear regression (R2) of the proximal and distal joint angles was used as the measure of the degree of coupling between the two joints. This coupling coefficient between the proximal and distal joint angles, for both forelegs and hindlegs, was significantly lowest at the highest temperature at which the animals ran the fastest with the highest stride frequencies. The coordination of multiple, in-series hydraulically actuated joints may be limited by operating speed. Highlighted Article: Tarantulas extend their leg joints hydraulically. The coupling between in-series, hydraulically actuated joints was lowest at the highest temperature when the animals ran the fastest with the highest stride frequencies. The coordination of multiple, in-series hydraulically actuated joints may be limited by operating speed.

Variability of Neural Activation During Walking in Humans: Short Heels and Big Calves

People come in different shapes and sizes. In particular, calf muscle size in humans varies considerably. One possible cause for the different shapes of calf muscles is the inherent difference in neural signals sent to these muscles during walking. In sedentary adults, the variability in neural control of the calf muscles was examined with muscle size, walking kinematics and limb morphometrics. Half the subjects walked while activating their medial gastrocnemius (MG) muscles more strongly than their lateral gastrocnemius (LG) muscles during most walking speeds (‘MG-biased’). The other subjects walked while activating their MG and LG muscles nearly equally (‘unbiased’). Those who walked with an MG-biased recruitment pattern also had thicker MG muscles and shorter heel lengths, or MG muscle moment arms, than unbiased walkers, but were similar in height, weight, lower limb length, foot length, and exhibited similar walking kinematics. The relatively less plastic skeletal system may drive calf muscle size and motor recruitment patterns of walking in humans.

Exceptional running and turning performance in a mite

ABSTRACT The Southern California endemic mite Paratarsotomus macropalpis was filmed in the field on a concrete substrate and in the lab to analyze stride frequency, gait and running speed under different temperature conditions and during turning. At ground temperatures ranging from 45 to 60°C, mites ran at a mean relative speed of 192.4±2.1 body lengths (BL) s−1, exceeding the highest previously documented value for a land animal by 12.5%. Stride frequencies were also exceptionally high (up to 135 Hz), and increased with substrate temperature. Juveniles exhibited higher relative speeds than adults and possess proportionally longer legs, which allow for greater relative stride lengths. Although mites accelerated and decelerated rapidly during straight running (7.2±1.2 and −10.1±2.1 m s−2, respectively), the forces involved were comparable to those found in other animals. Paratarsotomus macropalpis employs an alternating tetrapod gait during steady running. Shallow turns were accomplished by a simple asymmetry in stride length. During tight turns, mites pivoted around the tarsus of the inside third leg (L3), which thus behaved like a grappling hook. Pivot turns were characterized by a 42% decrease in turning radius and a 40% increase in angular velocity compared with non-pivot turns. The joint angle amplitudes of the inner L2 and L3 were negligible during a pivot turn. While exceptional, running speeds in P. macropalpis approximate values predicted from inter-specific scaling relationships. Highlighted Article: The mite Paratarsotomus macropalpis attains the highest relative speed and stride frequency documented for any animal.

Different Segments within Vertebrate Muscles Can Operate on Different Regions of Their Force–Length Relationships

To relate in vivo behavior of fascicle segments within a muscle to their in vitro force-length relationships, we examined the strain behavior of paired segments within each of three vertebrate muscles. After determining in vivo muscle activity patterns and length changes of in-series segments within the semimembranosus muscle (SM) in the American Toad (Bufo americanus) during hopping and within the sternohyoid (SH) muscle in the rat (Rattus rattus) during swallowing, and of spatially separated fascicles within the medial gastrocnemius (MG) muscle in the rat during trotting, we measured their corresponding in vitro (toad) or in situ (rat) force-length relationships (FLRs). For all three muscles, in vivo strain heterogeneity lasted for about 36-57% of the behavior cycle, during which one segment or fascicle shortened while the other segment or fascicle simultaneously lengthened. In the toad SM, the proximal segment shortened from the descending limb across the plateau of its FLR from 1.12 to 0.91 of its optimal length (Lo), while the distal segment lengthened (by 0.04 ± 0.04 Lo) before shortening down the ascending limb from 0.94 to 0.83 Lo. In the rat SH muscle, the proximal segment tended to shorten on its ascending limb from 0.90 to 0.85 Lo while the distal segment tended to lengthen across Lo (0.96-1.12 Lo). In the rat MG muscle, in vivo strains of proximal fascicles ranged from 0.72 to 1.02 Lo, while the distal fascicles ranged from 0.88 to 1.11 Lo. Even though the timing of muscle activation patterns were similar between segments, the heterogeneous strain patterns of fascicle segments measured in vivo coincided with different operating ranges across their FLRs simultaneously, implying differences in force-velocity behavior as well. The three vertebrate skeletal muscles represent a diversity of fiber architectures and functions and suggest that patterns of in vivo contractile strain and the operating range over the FLR in one muscle region does not necessarily represent other regions within the same muscle.