Rorqual Lunge-Feeding Energetics Near and Away from the Kinematic Threshold of Optimal Efficiency.

Humpback and blue whales are large baleen-bearing cetaceans, which use a unique prey-acquisition strategy-lunge feeding-to engulf entire patches of large plankton or schools of forage fish and the water in which they are embedded. Dynamically, and while foraging on krill, lunge-feeding incurs metabolic expenditures estimated at up to 20.0 MJ. Because of prey abundance and its capture in bulk, lunge feeding is carried out at high acquired-to-expended energy ratios of up to 30 at the largest body sizes (∼27 m). We use bio-logging tag data and the work-energy theorem to show that when krill-feeding at depth while using a wide range of prey approach swimming speeds (2-5 m/s), rorquals generate significant and widely varying metabolic power output during engulfment, typically ranging from 10 to 50 times the basal metabolic rate of land mammals. At equal prey field density, such output variations lower their feeding efficiency two- to three-fold at high foraging speeds, thereby allowing slow and smaller rorquals to feed more efficiently than fast and larger rorquals. The analysis also shows how the slowest speeds of harvest so far measured may be connected to the biomechanics of the buccal cavity and the prey's ability to collectively avoid engulfment. Such minimal speeds are important as they generate the most efficient lunges. Sommaire Les rorquals à bosse et rorquals bleus sont des baleines à fanons qui utilisent une technique d'alimentation unique impliquant une approche avec élan pour engouffrer de larges quantités de plancton et bancs de petits poissons, ainsi que la masse d'eau dans laquelle ces proies sont situés. Du point de vue de la dynamique, et durant l'approche et engouffrement de krill, leurs dépenses énergétiques sont estimées jusqu'à 20.0 MJ. À cause de l'abondance de leurs proies et capture en masse, cette technique d'alimentation est effectuée à des rapports d'efficacité énergétique (acquise -versus- dépensée) estimés aux environs de 30 dans le cas des plus grandes baleines (27 m). Nous utilisons les données recueillies par des capteurs de bio-enregistrement ainsi que le théorème reliant l'énergie à l'effort pour démontrer comment les rorquals s'alimentant sur le krill à grandes profondeurs, et à des vitesses variant entre 2 et 5 m/s, maintiennent des taux de dépenses énergétiques entre 10 et 50 fois le taux métabolique basal des mammifères terrestres. À densités de proies égales, ces variations d'énergie utilisée peuvent réduire le rapport d'efficacité énergétique par des facteurs entre 2x et 3x, donc permettant aux petits et plus lents rorquals de chasser avec une efficacité comparable à celle des rorquals les plus grands et rapides. Notre analyse démontre aussi comment des vitesses d'approche plus lentes peuvent être reliées à la biomécanique de leur poche ventrale extensible, et à l'habilitée des proies à éviter d'être engouffrer. Ces minimums de vitesses sont importants car ils permettent une alimentation plus efficace énergétiquement.

Why whales are big but not bigger: Physiological drivers and ecological limits in the age of ocean giants.

The largest animals are marine filter feeders, but the underlying mechanism of their large size remains unexplained. We measured feeding performance and prey quality to demonstrate how whale gigantism is driven by the interplay of prey abundance and harvesting mechanisms that increase prey capture rates and energy intake. The foraging efficiency of toothed whales that feed on single prey is constrained by the abundance of large prey, whereas filter-feeding baleen whales seasonally exploit vast swarms of small prey at high efficiencies. Given temporally and spatially aggregated prey, filter feeding provides an evolutionary pathway to extremes in body size that are not available to lineages that must feed on one prey at a time. Maximum size in filter feeders is likely constrained by prey availability across space and time.

Body Flexibility Enhances Maneuverability in the World's Largest Predator.

Blue whales are often characterized as highly stable, open-ocean swimmers who sacrifice maneuverability for long-distance cruising performance. However, recent studies have revealed that blue whales actually exhibit surprisingly complex underwater behaviors, yet little is known about the performance and control of these maneuvers. Here, we use multi-sensor biologgers equipped with cameras to quantify the locomotor dynamics and the movement of the control surfaces used by foraging blue whales. Our results revealed that simple maneuvers (rolls, turns, and pitch changes) are performed using distinct combinations of control and power provided by the flippers, the flukes, and bending of the body, while complex trajectories are structured by combining sequences of simple maneuvers. Furthermore, blue whales improve their turning performance by using complex banked turns to take advantage of their substantial dorso-ventral flexibility. These results illustrate the important role body flexibility plays in enhancing control and performance of maneuvers, even in the largest of animals. The use of the body to supplement the performance of the hydrodynamically active surfaces may represent a new mechanism in the control of aquatic locomotion.

How Baleen Whales Feed: The Biomechanics of Engulfment and Filtration.

Baleen whales are gigantic obligate filter feeders that exploit aggregations of small-bodied prey in littoral, epipelagic, and mesopelagic ecosystems. At the extreme of maximum body size observed among mammals, baleen whales exhibit a unique combination of high overall energetic demands and low mass-specific metabolic rates. As a result, most baleen whale species have evolved filter-feeding mechanisms and foraging strategies that take advantage of seasonally abundant yet patchily and ephemerally distributed prey resources. New methodologies consisting of multi-sensor tags, active acoustic prey mapping, and hydrodynamic modeling have revolutionized our ability to study the physiology and ecology of baleen whale feeding mechanisms. Here, we review the current state of the field by exploring several hypotheses that aim to explain how baleen whales feed. Despite significant advances, major questions remain about the processes that underlie these extreme feeding mechanisms, which enabled the evolution of the largest animals of all time.

Photograph-based ergonomic evaluations using the Rapid Office Strain Assessment (ROSA).

The Rapid Office Strain Assessment (ROSA) was developed to assess musculoskeletal disorder (MSD) risk factors for computer workstations. This study examined the validity and reliability of remotely conducted, photo-based assessments using ROSA. Twenty-three office workstations were assessed on-site by an ergonomist, and 5 photos were obtained. Photo-based assessments were conducted by three ergonomists. The sensitivity and specificity of the photo-based assessors' ability to correctly classify workstations was 79% and 55%, respectively. The moderate specificity associated with false positive errors committed by the assessors could lead to unnecessary costs to the employer. Error between on-site and photo-based final scores was a considerable ∼2 points on the 10-point ROSA scale (RMSE = 2.3), with a moderate relationship (ρ = 0.33). Interrater reliability ranged from fairly good to excellent (ICC = 0.667-0.856) and was comparable to previous results. Sources of error include the parallax effect, poor estimations of small joint (e.g. hand/wrist) angles, and boundary errors in postural binning. While this method demonstrated potential validity, further improvements should be made with respect to photo-collection and other protocols for remotely-based ROSA assessments.

Middle ear capillary haemangioma causing vestibulocochlear symptoms: a case report.

PROBLEM: A 58-year-old man presented with transient vertigo and pulsatile tinnitus. METHODS: High-resolution computed tomography, magnetic resonance imaging, excision, and subsequent immunohistochemical assays were performed. RESULTS: Imaging showed a soft tissue mass in the epitympanum and mastoid with bone erosion of the tegmen tympani and a dural tail sign, suggesting meningioma. Subsequently, because of signs of clinical progression, a canal-wall-up attico-antromastoidectomy was performed, with near-complete removal of a granulomatous, ossifying, haemorrhagic mass. CONCLUSIONS: Radiological imaging was critical in determining the extent of the mass and excluding other pathologies. Due to the atypical clinical and radiological signs, the final diagnosis of capillary haemangioma of the middle ear and temporal bone was made only after surgical resection and histopathological examination with immunohistochemistry, which excluded meningioma. The contiguous occurrence of cutaneous capillary haemangioma of the lateral face and neck was an important clue to the diagnosis.

Validation of the Dutch version of the Spatial Hearing Questionnaire.

OBJECTIVES: This study aimed to validate the Dutch version of the Spatial Hearing Questionnaire (SHQ). The original English SHQ by Tyler et al. is a useful and reliable self-reported assessment of spatial hearing. It has excellent construct validity and significantly correlates with other measures of auditory function. Our goal is to have similarly high internal consistency reliability and construct validity for the Dutch version of the SHQ. METHODS: We obtained the Dutch version of the original American English SHQ by the process of translation and back-translation. The SHQ was assessed in 71 patients at the Ear, Nose and Throat (ENT) department of Antwerp University Hospital. Internal consistency reliability was evaluated using Cronbach's alpha value. Factor analysis was performed and compared to results from previous psychometric analysis of the original SHQ. Construct validity was checked by comparing scores obtained in patients with asymmetric and symmetric hearing loss. RESULTS: The internal consistency of the Dutch version of the SHQ was very high with a Cronbach's alpha value of 0.98. Comparison of a group with asymmetric and symmetric impaired patients revealed significant differences in the SHQ total score and most subscale scores, emphasizing good construct validity. CONCLUSIONS: The SHQ was successfully translated to Dutch. The psychometric characters of the Dutch questionnaire are similar to the original SHQ in English, with high internal consistency reliability. The Dutch version of the SHQ, Vragenlijst voor Ruimtelijk Horen, is a valid and reliable instrument for measuring spatial hearing abilities in both clinic and research settings.

Third mobile window associated with suspected otosclerotic foci in two patients with an air-bone gap.

OBJECTIVE: To demonstrate the need for computed tomography imaging of the temporal bone in patients clinically suspected of otosclerosis who present with atypical symptoms or audiological findings. CASE REPORTS: We present two patients with bilateral conductive hearing loss and suspected otosclerosis in whom third mobile window lesions were revealed. The first patient had bilateral large vestibular aqueducts and bilateral fenestral otosclerotic foci. Computed tomography imaging of the second case revealed bilateral superior semicircular canal dehiscence and bilateral cochlear clefts, mimicking an otosclerotic focus in the fissula ante fenestram. CONCLUSION: Differentiating third mobile window lesions from otosclerosis as the cause of a conductive hearing loss is essential before considering stapes surgery, as such treatment would be unnecessary and potentially harmful.

Mechanics, hydrodynamics and energetics of blue whale lunge feeding: efficiency dependence on krill density.

Lunge feeding by rorqual whales (Balaenopteridae) is associated with a high energetic cost that decreases diving capacity, thereby limiting access to dense prey patches at depth. Despite this cost, rorquals exhibit high rates of lipid deposition and extremely large maximum body size. To address this paradox, we integrated kinematic data from digital tags with unsteady hydrodynamic models to estimate the energy budget for lunges and foraging dives of blue whales (Balaenoptera musculus), the largest rorqual and living mammal. Our analysis suggests that, despite the large amount of mechanical work required to lunge feed, a large amount of prey and, therefore, energy is obtained during engulfment. Furthermore, we suggest that foraging efficiency for blue whales is significantly higher than for other marine mammals by nearly an order of magnitude, but only if lunges target extremely high densities of krill. The high predicted efficiency is attributed to the enhanced engulfment capacity, rapid filter rate and low mass-specific metabolic rate associated with large body size in blue whales. These results highlight the importance of high prey density, regardless of prey patch depth, for efficient bulk filter feeding in baleen whales and may explain some diel changes in foraging behavior in rorqual whales.

Scaling of lunge feeding in rorqual whales: an integrated model of engulfment duration.

Rorqual whales (Balaenopteridae) obtain their food by lunge feeding, a dynamic process that involves the intermittent engulfment and filtering of large amounts of water and prey. During a lunge, whales accelerate to high speed and open their mouth wide, thereby exposing a highly distensible buccal cavity to the flow and facilitating its inflation. Unsteady hydrodynamic models suggest that the muscles associated with the ventral groove blubber undergo eccentric contraction in order to stiffen and control the inflation of the buccal cavity; in doing so the engulfed water mass is accelerated forward as the whale's body slows down. Although the basic mechanics of lunge feeding are relatively well known, the scaling of this process remains poorly understood, particularly with regards to its duration (from mouth opening to closure). Here we formulate a new theory of engulfment time which integrates prey escape behavior with the mechanics of the whale's body, including lunge speed and acceleration, gape angle dynamics, and the controlled inflation of the buccal cavity. Given that the complex interaction between these factors must be highly coordinated in order to maximize engulfment volume, the proposed formulation rests on the scenario of Synchronized Engulfment, whereby the filling of the cavity (posterior to the temporomandibular joint) coincides with the moment of maximum gape. When formulated specifically for large rorquals feeding on krill, our analysis predicts that engulfment time increases with body size, but in amounts dictated by the specifics of krill escape and avoidance kinematics. The predictions generated by the model are corroborated by limited empirical data on a species-specific basis, particularly for humpback and blue whales chasing krill. A sensitivity analysis applied to all possible sized fin whales also suggests that engulfment duration and lunge speed will increase intra-specifically with body size under a wide range of predator-prey scenarios. This study provides the theoretical framework required to estimate the scaling of the mass-specific drag being generated during engulfment, as well as the energy expenditures incurred.

Passive versus active engulfment: verdict from trajectory simulations of lunge-feeding fin whales Balaenoptera physalus.

Lunge-feeding in rorqual whales represents the largest biomechanical event on Earth and one of the most extreme feeding methods among aquatic vertebrates. By accelerating to high speeds and by opening their mouth to large gape angles, these whales generate the water pressure required to expand their mouth around a large volume of prey-laden water. Such large influx is facilitated by highly extensible ventral groove blubber (VGB) associated with the walls of the throat (buccal cavity). Based on the mechanical properties of this tissue, previous studies have assumed lunge-feeding to be an entirely passive process, where the flow-induced pressure driving the expansion of the VGB is met with little resistance. Such compliant engulfment would be facilitated by the compliant properties of the VGB that have been measured on dead specimens. However, adjoining the ventral blubber are several layers of well-developed muscle embedded with mechanoreceptors, thereby suggesting a capability to gauge the magnitude of engulfed water and use eccentric muscle action to control the flux of water into the mouth. An unsteady hydrodynamic model of fin whale lunge-feeding is presented here to test whether engulfment is exclusively passive and compliant or involves muscle action. The model is based on the explicit simulation of the engulfed water as it interacts with the buccal cavity walls of the whale, under different heuristically motivated cavity forces. Our results, together with their comparison with velocity data collected in the field, suggest that adult rorquals actively push engulfed water forward from the very onset of mouth opening in order to successfully complete a lunge. Interestingly, such an action involves a reflux of the engulfed mass rather than the oft-assumed rebound, which would occur mainly at the very end of a lunge sequence dominated by compliant engulfment. Given the great mass of the engulfed water, reflux creation adds a significant source of hydrodynamic drag to the lunge process, but with the benefit of helping to circumvent the problem of removing prey from baleen by enhancing the efficiency of cross-flow filtration after mouth closing. Reflux management for a successful lunge will therefore demand well-coordinated muscular actions of the tail, mouth and ventral cavity.

Less is more: high pass filtering, to remove up to 99% of the surface EMG signal power, improves EMG-based biceps brachii muscle force estimates.

It is generally assumed that raw surface EMG (sEMG) should be high pass filtered with cutoffs of 10-30 Hz to remove motion artifact before subsequent processing to estimate muscle force. The purpose of the current study was to explore the benefits of filtering out much of the raw sEMG signal when attempting to estimate accurate muscle forces. Twenty-five subjects were studied as they performed rapid static, anisotonic contractions of the biceps brachii. Biceps force was estimated (as a percentage of maximum) based on forces recorded at the wrist. An iterative approach was used to process the sEMG from the biceps brachii, using progressively greater high pass cutoff frequencies (20-440 Hz in steps of 30 Hz) with first and sixth order filters, as well as signal whitening, to determine the effects on the accuracy of EMG-based biceps force estimates. The results indicate that removing up to 99% of the raw sEMG signal power resulted in significant and substantial improvements in biceps force estimates. These findings challenge previous assumptions that the raw sEMG signal power between about 20 and 500 Hz should used when estimating muscle force. For the purposes of force prediction, it appears that a much smaller, high band of sEMG frequencies may be associated with force and the remainder of the spectrum has little relevance for force estimation.

The responses of leg and trunk muscles to sudden unloading of the hands: implications for balance and spine stability.

OBJECTIVE: To examine the anticipatory and responsive actions of leg and trunk muscles, and their role in whole-body and spine control in situations of sudden unloading of the hands in the sagittal plane. DESIGN: EMG and force plate measures were used to determine the baseline, anticipatory responses and post unloading responses of selected trunk and leg muscles under different conditions of unload timing knowledge. BACKGROUND: Postural muscles have been observed to increase activation in anticipation of a known loading situation to decrease the overall effect of an impulsive load delivered to the spine. It is thought that this increased activation places the spine in a more stable state, thereby reducing the likelihood of injury. Comparisons have not been made previously of the responses of postural muscles to unloading conditions where the certainty of unload timing is varied. METHODS: Eleven male subjects, holding a 6.8 kg load in the hands, were subjected to three different unloading conditions: (1) voluntary load drop; (2) known timing of load release; (3) unknown timing of load release. Anterior-posterior center of pressure data, as well as EMG activity on 8 right side muscles, were collected for 10 trials in each condition. RESULTS: Anterior-posterior center of pressure responses were significantly different (P<0.05) between each of the three conditions. Lumbar erector spinae and thoracic erector spinae significantly decreased anticipatory activity as knowledge of the unload timing increased. Five of the eight monitored muscles demonstrated significantly decreased response levels as knowledge of the timing of unloading increased. CONCLUSIONS: When an unload is self-triggered, preparatory adjustments can be made which reduce the overall postural perturbation to the body, and the spine in particular, while minimizing the responsive activity of trunk muscles. RELEVANCE: Spinal instability has been identified as a risk factor for low back injury during trunk loading. This study demonstrates that, in situations of sudden unloading, knowledge of the timing of the unloading may lead to anticipatory actions of postural muscles which actually decrease spinal stability, thereby increasing the risk of injury were an unexpected perturbation to occur.

The influence of horizontal velocity on interlimb symmetry in normal walking.

Changes in horizontal velocity (HV) are known to influence many biomechanical characteristics of human locomotion. The purpose of the present study was to investigate this phenomenon with respect to the interlimb symmetry of walking in a normal population. Peak and temporal ground reaction force data from both feet of 20 able-bodied males were collected at each of three relative velocity conditions (slow, normal and fast). These data were analyzed using of a series of 2 x 3 repeated measures ANOVAs, which revealed a high degree of interlimb (bilateral) symmetry across HV conditions despite significant intralimb (unilateral) changes. In contrast to this primary finding were two significant interaction effects for the stance time and peak vertical force at push-off measures respectively. These interactions indicated greater asymmetries at the slow HV condition with a trend toward improved symmetry at higher velocities. Although these results may provide some theoretical insight into the underlying nature of symmetry in gait, their overall magnitude does not seem to invalidate the current widespread use of symmetry assumptions in clinical and research settings today.

Quantification of subcellular glycogen in resting human muscle: granule size, number, and location.

A few qualitative investigations suggested that location of muscle glycogen (G) granules in specific sites may be associated with distinct metabolic roles. Similarly, it has been suggested that the acid-soluble and -insoluble G fractions (macro- and proglycogen, respectively) are different metabolic pools and also could exist as separate entities. We employed a transmission electron microscopic technique to quantify subcellular G particle size, number, and location in human vastus lateralis biopsies of 11 resting men. The intra- and interobserver variability for the various measures was generally <4%. Granule size and number were quantified in subcellular compartments (subsarcolemmal, intra- and intermyofibrillar). Subcellular location was critical: G was more densely concentrated in the subsarcolemmal than in the myofibrillar space, whereas the single-particle volume was greater in the latter. Single-particle diameter ranged from 10 to 44 etam and followed a continuous, normal distribution. This implies that proglycogen is not a distinct entity, but rather that pro- and macroglycogen are divisions of smaller and larger molecules. These results demonstrate a compartmentalized pattern of subcellular G deposition in human skeletal muscle for both the size and density of granules.

Medio-lateral balance adjustments preceding reflexive limb withdrawal are modified by postural demands.

We have recently observed medio-lateral balance adjustments (BA) preceding reflexive stepping elicited by noxious stimulation. While task specific modulation is evident for BA prior to voluntary leg movement, it is unclear whether rapid BA reactions (prior to 'reflexive' stepping) represent a generic response to evoked limb withdrawal or can be modified to suit task-conditions. This study was designed to establish whether the CNS is able to modify rapid onset latency BAs to match task conditions. Reflexive stepping was evoked by applying a noxious stimulus (50 ms stimulus train, 1 ms pulses, 300 Hz, 4 x perceptual threshold) to the plantar surface of the either the left or right foot. Task conditions were varied prior to stimulation by having subjects maintain one of three different static positions: (1) lean left (70% body weight (BW) on left), (2) neutral (50% BW both sides), (3) lean right (70% BW on right). BAs were denoted by centre-of-pressure (CoP) excursions towards the swing foot after the onset of noxious stimulation (average onset latency of 128 ms). There was a significant increase in frequency of occurrence and a significant increase in magnitude of CoP shift when the stimulation was applied to a loaded limb (leaning with 70% BW on the stimulated foot) as compared to an unloaded limb (30% BW). In addition, 78% of loaded trials featured steps taken with the unstimulated foot, which delayed removal of the stimulated foot. Collectively, the results indicate modifiability of the very rapid onset balance adjustments that precede the onset of limb withdrawal revealing complex control of balance exists even over very brief latencies.

The in vivo dynamic response of the human spine to rapid lateral bend perturbation: effects of preload and step input magnitude.

STUDY DESIGN: A repeated measures design was used to determine the effects that combinations of two preloads and two added loads have on spine mechanics both before and during the response to the added load. OBJECTIVE: To investigate the effects of varying initial isometric and added step input load magnitudes on mechanical and electromyographic responses of the trunk during sudden loading that causes lateral bending moments. SUMMARY OF BACKGROUND DATA: Cocontractions of the antagonistic and agonistic muscles of the trunk are required for stability during loading of the spine. In several in vivo studies, it was observed that trunk muscle cocontraction serves a functional role before the application of unexpected or sudden loads. The response of agonistic and antagonistic trunk muscles to rapid lateral bend moments would provide further insight into the dynamic stability mechanisms of the spine. METHODS: In this study, 13 men maintained an upright standing posture while resisting the application of lateral bend moments produced by four different loading conditions comprising combinations of two preloads (5% or 15% of the maximum isometric lateral bend moment) and two added loads (20% or 30%). The preloading was used to develop different initial levels of trunk stiffness before the application of the added loads. The lateral bend moment and angular rotation of the trunk were measured, as well as the surface electromyogram amplitudes of the bilateral internal oblique, external oblique, rectus abdominus, lumbar erector spinae, and thoracic erector spinae muscles. Dependent measures were recorded during the steady state preload conditions, and peak values were recorded after the load was added. RESULTS: Higher added loads resulted in higher peak lateral bend rotations, and higher preloads resulted in lower rotations. The patterns of response were similar for the peak lateral bend moments and the electromyogram amplitudes from four of the five agonistic muscles. The thoracic erector spinae excepted, each of the other four muscles demonstrated larger responses in the agonistic muscles. However, all of the antagonistic muscles showed some increase in electromyogram activity in response to the added load. The thoracic erector spinae appeared to have the role of counteracting the flexor moments created by the abdominal muscles and the maintenance of spine stability. The agonistic external obliques and lumbar erector spinae had the largest responses to the added load. A comparison of the 35% loading conditions showed an increased response of the trunk to the 5% + 30% condition (with lower initial trunk stiffness), as compared with the 15% + 20% condition. CONCLUSIONS: The findings from this study show that higher levels of preactivation can serve to increase spine compression and trunk muscle stiffness, thereby attenuating the lateral displacements caused by rapid loading. Furthermore, antagonistic muscles were observed to respond rapidly to such perturbations with large increases in activation when preactivation and spine stability were low. The trunk muscles monitored all were larger, multisegmental muscles. The results from this study lend support to previous studies suggesting that the larger multisegmental muscles make a significant contribution to spinal stability.

Preparatory balance adjustments precede withdrawal response to noxious stimulation in standing humans.

Self-initiated leg movement in standing humans is preceded by a medio-lateral preparatory balance adjustment (PBA); however, such preparatory balance control is often absent in reflex-like stepping responses evoked by whole-body instability. The presence or absence of the PBA may reflect a task-dependent modulation of the response serving to preserve lateral stability (PBA present) or avoid delay in the lifting of the foot (PBA absent). To examine whether such task-dependent modulation can occur during more stereotypical limb movements, we examined spinally-mediated withdrawal responses evoked by noxious stimulation of the foot. Results showed that rapid limb withdrawal was preceded by a large PBA when subjects were standing but not when they were supine. The PBA caused limb withdrawal to the noxious stimulation to be delayed. However, the onset of the PBA in the standing trials was equivalent in timing to the onset latency of the classic withdrawal responses recorded during the supine trials. Evidence of a preparatory balance adjustment evoked, in advance of a delayed withdrawal response, at very rapid latencies (underlying muscle activation at 70-120 ms) may raise new questions about the neural mechanisms underlying the co-ordination of balance and movement.

The in vivo dynamic response of the spine to perturbations causing rapid flexion: effects of pre-load and step input magnitude.

OBJECTIVE: To evaluate the impact of muscle pre-activation levels and load magnitude on the response of the trunk to loading conditions causing rapid flexion. DESIGN: Eight male subjects were asked to maintain an upright standing posture while resisting the application of forward flexion moments produced by four different loading conditions consisting of combinations of two pre-loads (4% or 16% of the maximum extensor moment) and two added loads (12% or 24%). Pre-loading was used to develop different initial levels of trunk muscle activity prior to the application of the added loads. Of special interest were the two conditions that resulted in total final loads of 28%. BACKGROUND: Cocontraction of the antagonistic and agonistic muscles of the trunk are required to provide stability during normal physiological loading conditions. In several in vivo studies, levels of trunk muscle cocontraction have been observed prior to the application of unexpected or sudden loads. Forces from the abdominal muscles have been proposed to provide stability when extensor moments are generated. The response of trunk muscles to rapid flexor moments would provide further insight into the dynamic stability mechanisms of the spine. METHODS: Measurements were made of the trunk extensor moments, angular displacement of the trunk and unilateral surface EMG amplitudes of three abdominal and three trunk extensor muscles. Values were recorded during the isometric pre-load and for the maximum magnitude of each variable in response to the added load. RESULTS: Higher pre-loads resulted in lower flexion rotations of the spine and higher added loads caused larger rotations. With increasing magnitudes of final loads, a corresponding increase in trunk extensor moments and trunk muscle cocontraction was observed. The largest activations were observed in the lumbar erector spinae and thoracic erector spinae muscles, while smaller yet substantial EMG activity was observed in the internal oblique and external oblique. A comparison of the 28% loading conditions showed an increased response of the trunk to the [4 + 24] loading condition (with lower initial trunk stiffness) when compared to the [16 + 12] loading condition. CONCLUSIONS: Pre-activation of trunk extensor muscles can serve to reduce the flexion displacements caused by rapid loading. The abdominal oblique muscles, especially external oblique, will rapidly increase their activation levels in response to rapid loading. These changes are more pronounced when pre-activation levels are low, resulting in lower initial trunk stiffness and spine compression force. It is proposed that these factors will ultimately affect spine stability and the risk of injury. RELEVANCE: The results of this study provide insight into several mechanisms involved in the dynamic stability of the spine. Injuries can be caused by unexpected and rapid loading of the spine. A study of the mechanisms available to respond to such perturbations is important to an understanding of spine mechanics and the etiology of low back injury.