Andrew R. Lammers

Ontogenetic allometry in the locomotor skeleton of specialized half-bounding mammals

Specialization for a locomotor behaviour may affect limb bone morphology throughout ontogeny. Ontogenetic development of the limb skeletons of two mammalian species, which are behaviourally specialized for the half-bounding gait (Chinchilla lanigera and Oryctolagus cuniculus), were compared to two similarly-sized species which are not specialized for half-bounding (Rattus norvegicus and Monodelphis domestica). Limb bone lengths and anteroposterior diameters (mediolateral diameters for the radius and metacarpal) were measured from radiographs taken throughout the ontogeny of each species. Body mass was also measured repeatedly during growth. Bone measurements were regressed against body mass, as well as forelimb bone length vs serially homologous hindlimb bone length, bone length vs total limb length and bone length vs width. Similar comparisons were made among adults of each species using ratios. Although there were many significant differences among species, overall there were few consistent differences in adult scaling ratios or ontogenetic allometry slopes between specialized and generalized groups. Adult specialized half-bounders had significantly narrower tibiae and metatarsals than the gait-generalized runners. Specialized half-bounders usually had similar slopes for hindlimb length vs width ontogenetic comparisons, but the non-specialized species did not group together. However, there were two patterns that occurred among all four species: (1) hindlimb bone lengths nearly always grew faster than the serially homologous forelimb bone lengths in all species; (2) proximal elements usually increased in length proportionally faster than distal elements. In conclusion, small mammals may share strong developmental constraints that govern their relative growth rates. It is also likely that there are different selective pressures on juveniles and adults, but that these selective pressures may not be different between specialized and unspecialized runners during ontogeny.

Different parts of erector spinae muscle fatigability in subjects with and without low back pain

BACKGROUND CONTEXT There is conflicting evidence regarding erector spinae muscle fatigability because previous studies have not considered the thoracic and lumbar components separately. These muscles have very different mechanical responses and, therefore, would be recruited differentially for the chosen task. PURPOSE The present study was conducted to compare whether fatigability differences exist in the thoracic and lumbar parts of the erector spinae muscles in subjects with and without low back pain (LBP). STUDY DESIGN This cross-sectional study was conducted in the Motion Analysis Lab at Cleveland State University. PATIENT SAMPLE The study sample included 40 subjects with LBP and 40 subjects without LBP. OUTCOME MEASURES The fatigability of the erector spinae muscles was compared based on median frequency of electromyography (EMG) versus time. The level of pain of each subject was also compared using the Oswestry Disability Index. METHODS Fatigue measurements were evaluated between groups based on the assessed sides as well as the thoracic and lumbar parts of the erector spinae muscles using surface EMG. A modified version of the isometric fatigue test as introduced by Sorensen was used to test the endurance of the erector spinae muscles. RESULTS There were significant median EMG frequency (F((1, 78))=28.82, p=.001) differences in the thoracic and lumbar parts of the erector spinae muscles between subjects with and without LBP. The thoracic part had a significantly lower median EMG frequency than the lumbar part in subjects with LBP. The thoracic and lumbar parts of the erector spinae muscles had interactions with group (F((1, 78))=47.88, p=.01] and age (F((1, 78))=16.51, p=.01). CONCLUSIONS The results of this study suggested that subjects with LBP demonstrated higher fatigability of the erector spinae muscles at the thoracic part than at the lumbar part. The increased fatigability of the thoracic part needs to be emphasized in rehabilitation strategies for subjects with LBP. In addition, as age increased, the median frequency of the lumbar part of the erector spinae muscles significantly decreased. Understanding the anatomical and biomechanical characteristics of the erector spinae muscle may enhance clinical outcomes and rehabilitation strategies for subjects with LBP.

ONTOGENY OF SEXUAL DIMORPHISM IN CHINCHILLA LANIGERA (RODENTIA: CHINCHILLIDAE)

Abstract Growth patterns that lead to sexual dimorphism in adults are not well quantified. We measured 49 skeletal dimensions in male and female Chinchilla lanigera from radiographs of growing individuals taken during 320 days. Measurements for each individual were fit with a nonlinear Gompertz equation to quantify growth patterns. Differences in Gompertz parameters between sexes were compared with a t-test. Most significant differences between sexes in growth and final size were in the pelvic girdle (which formed the birth canal) and viscerocranium. Sexual dimorphism in the viscerocranium may support the hypothesis that differences in use of ecological niche often causes sexual dimorphism where females are larger than males.

Mechanics of torque generation during quadrupedal arboreal locomotion

Quadrupedal animals moving on arboreal substrates face unique challenges to maintain stability. The torque generated by the limbs around the long axis of a branch during locomotion may clarify how the animals remain stable on arboreal supports. We sought to determine what strategy gray short-tailed opossums (Monodelphis domestica) use to exert torque and avoid toppling. The opossums moved across a branch trackway about half the diameter of their bodies. Part of the trackway was instrumented to measure substrate reaction forces and torque around the long axis of the branch. Kinematic analysis was used to estimate the center of pressure of the manus and pes; from center of pressure and vertical and mediolateral forces, the torque generated by substrate reaction forces versus muscular effort could be determined. Forelimbs generated significantly greater torque than hindlimbs, which is probably explained by the greater weight-bearing role of the forelimbs. Fore- and hindlimbs generated torque in opposite directions because contralateral fore- and hindlimbs typically contacted the branch. Torque generated by muscular effort, however, was often in the same direction in both fore- and hindlimbs. The muscle-generated torque is likely the result of mediolateral movement of the center of mass caused by mediolateral undulation of the torso. These results bear an important implication for the study of arboreal locomotion: center of mass dynamics are at least as important as static positions. M. domestica is a good representative for a primitive mammal, and comparisons with arboreal specialists will shed light on how proficient arboreal locomotion evolved.

Torque around the center of mass: dynamic stability during quadrupedal arboreal locomotion in the Siberian chipmunk (Tamias sibiricus)

When animals travel on tree branches, avoiding falls is of paramount importance. Animals swiftly running on a narrow branch must rely on movement to create stability rather than on static methods. We examined how Siberian chipmunks (Tamias sibiricus) remain stable while running on a narrow tree branch trackway. We examined the pitch, yaw, and rolling torques around the center of mass, and hypothesized that within a stride, any angular impulse (torque during step time) acting on the center of mass would be canceled out by an equal and opposite angular impulse. Three chipmunks were videotaped while running on a 2cm diameter branch trackway. We digitized the videos to estimate center of mass and center of pressure positions throughout the stride. A short region of the trackway was instrumented to measure components of the substrate reaction force. We found that positive and negative pitch angular impulse was by far the greatest in magnitude. The anterior body was pushed dorsally (upward) when the forelimbs landed simultaneously, and then the body pitched in the opposite direction as both hindlimbs simultaneously made contact. There was no considerable difference between yaw and rolling angular impulses, both of which were small and equal between fore- and hindlimbs. Net angular impulses around all three axes were usually greater than or less than zero (not balanced). We conclude that the chipmunks may balance out the torques acting on the center of mass over the course of two or more strides, rather than one stride as we hypothesized.

Mechanics of generating friction during locomotion on rough and smooth arboreal trackways

SUMMARY Traveling on arboreal substrates is common among most small mammals living anywhere vegetation grows. Because arboreal supports vary considerably in surface texture, animals must be able to adjust their locomotor biomechanics to remain stable on such supports. I examined how gray short-tailed opossums (Monodelphis domestica), which are generalized marsupials living on or near the ground, adjust to travel on rough and smooth 2 cm-diameter arboreal trackways. Limb contact position was determined via high-speed videography, and substrate reaction force was measured by an instrumented section of each branch trackway. Normal and shear forces were calculated from substrate reaction force and limb contact position around the branch trackways. Normal force is greater in forelimbs, probably because of the forelimbs greater weight support role. Shear force was identical between limb pairs, most likely because of interactions between vertical force, limb placement, mediolateral force, and torque. The opossums adjusted to the smooth trackway mainly by reducing speed, changing footfall patterns and increasing normal force. I predict that arboreal specialists will show less change in performance between rough and smooth arboreal trackways because of their greater ability to grasp or maintain contact with arboreal substrates.

Central nervous system integration of sensorimotor signals in oral and pharyngeal structures: oropharyngeal kinematics response to recurrent laryngeal nerve lesion

Safe, efficient liquid feeding in infant mammals requires the central coordination of oropharyngeal structures innervated by multiple cranial and spinal nerves. The importance of laryngeal sensation and central sensorimotor integration in this system is poorly understood. Recurrent laryngeal nerve lesion (RLN) results in increased aspiration, though the mechanism for this is unclear. This study aimed to determine the effect of unilateral RLN lesion on the motor coordination of infant liquid feeding. We hypothesized that 1) RLN lesion results in modified swallow kinematics, 2) postlesion oropharyngeal kinematics of unsafe swallows differ from those of safe swallows, and 3) nonswallowing phases of the feeding cycle show changed kinematics postlesion. We implanted radio opaque markers in infant pigs and filmed them pre- and postlesion with high-speed videofluoroscopy. Markers locations were digitized, and swallows were assessed for airway protection. RLN lesion resulted in modified kinematics of the tongue relative to the epiglottis in safe swallows. In lesioned animals, safe swallow kinematics differed from unsafe swallows. Unsafe swallow postlesion kinematics resembled prelesion safe swallows. The movement of the tongue was reduced in oral transport postlesion. Between different regions of the tongue, response to lesion was similar, and relative timing within the tongue was unchanged. RLN lesion has a pervasive effect on infant feeding kinematics, related to the efficiency of airway protection. The timing of tongue and hyolaryngeal kinematics in swallows is a crucial locus for swallow disruption. Laryngeal sensation is essential for the central coordination in feeding of oropharyngeal structures receiving motor inputs from different cranial nerves.

The Physiologic Impact of Unilateral Recurrent Laryngeal Nerve (RLN) Lesion on Infant Oropharyngeal and Esophageal Performance

Recurrent laryngeal nerve (RLN) injury in neonates, a complication of patent ductus arteriosus corrective surgery, leads to aspiration and swallowing complications. Severity of symptoms and prognosis for recovery are variable. We transected the RLN unilaterally in an infant mammalian animal model to characterize the degree and variability of dysphagia in a controlled experimental setting. We tested the hypotheses that (1) both airway protection and esophageal function would be compromised by lesion, (2) given our design, variability between multiple post-lesion trials would be minimal, and (3) variability among individuals would be minimal. Individuals’ swallowing performance was assessed pre- and post-lesion using high speed VFSS. Aspiration was assessed using the Infant Mammalian Penetration-Aspiration Scale (IMPAS). Esophageal function was assessed using two measures devised for this study. Our results indicate that RLN lesion leads to increased frequency of aspiration, and increased esophageal dysfunction, with significant variation in these basic patterns at all levels. On average, aspiration worsened with time post-lesion. Within a single feeding sequence, the distribution of unsafe swallows varied. Individuals changed post-lesion either by increasing average IMPAS score, or by increasing variation in IMPAS score. Unilateral RLN transection resulted in dysphagia with both compromised airway protection and esophageal function. Despite consistent, experimentally controlled injury, significant variation in response to lesion remained. Aspiration following RLN lesion was due to more than unilateral vocal fold paralysis. We suggest that neurological variation underlies this pattern.

Stability During Arboreal Locomotion

Arboreal locomotion – traveling on the branches, twigs, and trunks of trees and woody shrubs – is very common among mammals. Most primates, many rodents, marsupials, carnivores, and even an occasional artiodactyl travel on arboreal substrates to forage, escape predators, and acquire shelter. Arboreal supports are usually far enough from the ground that a slip or fall could cause serious injury or death, or deprive the animal of a mate, food, or energy. Thus, stability is of great importance for an animal traveling on arboreal supports. The considerable variation among arboreal supports makes stability during locomotion a mechanical challenge. Supports vary in diameter, slope, compliance, texture, direction (that is, bends or curves in a branch), and number and distribution. Furthermore there may be interaction among these variables; for example, compliance varies with diameter – thinner branches are more compliant than thick branches. Also, the thin branches frequently have leaves that act like sails in the wind, causing even more movement in the substrate. Substrate texture often varies with diameter, where narrow twigs have smoother bark than large branches or trunks. Therefore one might expect a considerable number of morphological, behavioral, and biomechanical mechanisms to enhance stability on arboreal supports. Stability can be divided into two categories: static and dynamic. Static stability is the process by which objects at rest remain stable, i.e., neither move (translation) nor rotate about a point or axis. For example, a table is statically stable because the forces and moments (torques) produced by gravity (weight) are balanced by ground reaction forces and the moments generated by them. One way an animal might remain stable is by not moving and adhering to or gripping the support; this definition is the ultimate example of static stability in an animal. Although this strategy allows no movement, it is nevertheless a valid locomotor strategy for an animal attempting to travel on an arboreal support subjected to a sudden gust of wind or other disturbance (Stevens, 2003). This analysis also applies when the animal walks very slowly, but fails when it walks or runs at considerable speed. Because the distribution of the mass is changing from one instant to the next, the forces and torques necessary to maintain static stability would also change with time. That is, it requires an active control by the nervous system. Because stability is critical, it is very likely that the animal employs both active and passive control (Full et al., 2002). Passive control can be due to dynamic processes of the animal’s body, and is referred to as dynamic stability. For example, a hiker might cross a stream or river by running across a fallen log; the rotation of the limbs around the hips and shoulder

The effects of substrate texture on the mechanics of quadrupedal arboreal locomotion in the gray short-tailed opossum (Monodelphis domestica)

Among small mammals, the ability to move on tree trunks, branches, and twigs is nearly ubiquitous. Performance and locomotor mechanics on arboreal substrates may be influenced by variation in the coefficient of friction between the hands/feet of the animal and the surface of the arboreal substrate. To test this, I examined speed, substrate reaction forces, and torque around the long axis of two cylindrical trackways with rough and smooth surfaces in gray short-tailed opossums (Monodelphis domestica). Speed was determined with videography, and forces and torques were measured by an instrumented section of the trackway. The opossums traveled more slowly on the smooth arboreal trackway. There was also significant interaction between limb (forelimbs, hindlimbs) and substrate texture (rough, smooth) in braking, propulsive, and laterally directed impulses. Running on the smooth trackway had the effect of reducing some between-limb (forelimb vs. hindlimb) differences. Stability on the rough trackway was probably maintained by relatively high momentum, but on the smooth trackway, the opossums used static methods (many limbs contacting the substrate, greater muscular effort, lower momentum) to remain stable and avoid toppling. Clearly, momentum and dynamics are often important biomechanical considerations for this generalized mammal. Highly arboreal animals can remain dynamically stable on a wider variety of substrate textures.