Partial coherence enhances parallelized photonic computing.

Advancements in optical coherence control1-5 have unlocked many cutting-edge applications, including long-haul communication, light detection and ranging (LiDAR) and optical coherence tomography6-8. Prevailing wisdom suggests that using more coherent light sources leads to enhanced system performance and device functionalities9-11. Our study introduces a photonic convolutional processing system that takes advantage of partially coherent light to boost computing parallelism without substantially sacrificing accuracy, potentially enabling larger-size photonic tensor cores. The reduction of the degree of coherence optimizes bandwidth use in the photonic convolutional processing system. This breakthrough challenges the traditional belief that coherence is essential or even advantageous in integrated photonic accelerators, thereby enabling the use of light sources with less rigorous feedback control and thermal-management requirements for high-throughput photonic computing. Here we demonstrate such a system in two photonic platforms for computing applications: a photonic tensor core using phase-change-material photonic memories that delivers parallel convolution operations to classify the gaits of ten patients with Parkinson's disease with 92.2% accuracy (92.7% theoretically) and a silicon photonic tensor core with embedded electro-absorption modulators (EAMs) to facilitate 0.108 tera operations per second (TOPS) convolutional processing for classifying the Modified National Institute of Standards and Technology (MNIST) handwritten digits dataset with 92.4% accuracy (95.0% theoretically).

Postcranial evidence of late Miocene hominin bipedalism in Chad.

Bipedal locomotion is one of the key adaptations that define the hominin clade. Evidence of bipedalism is known from postcranial remains of late Miocene hominins as early as 6 million years ago (Ma) in eastern Africa1-4. Bipedality of Sahelanthropus tchadensis was hitherto inferred about 7 Ma in central Africa (Chad) based on cranial evidence5-7. Here we present postcranial evidence of the locomotor behaviour of S. tchadensis, with new insights into bipedalism at the early stage of hominin evolutionary history. The original material was discovered at locality TM 266 of the Toros-Ménalla fossiliferous area and consists of one left femur and two, right and left, ulnae. The morphology of the femur is most parsimonious with habitual bipedality, and the ulnae preserve evidence of substantial arboreal behaviour. Taken together, these findings suggest that hominins were already bipeds at around 7 Ma but also suggest that arboreal clambering was probably a significant part of their locomotor repertoire.

Footprint evidence of early hominin locomotor diversity at Laetoli, Tanzania.

Bipedal trackways discovered in 1978 at Laetoli site G, Tanzania and dated to 3.66 million years ago are widely accepted as the oldest unequivocal evidence of obligate bipedalism in the human lineage1-3. Another trackway discovered two years earlier at nearby site A was partially excavated and attributed to a hominin, but curious affinities with bears (ursids) marginalized its importance to the paleoanthropological community, and the location of these footprints fell into obscurity3-5. In 2019, we located, excavated and cleaned the site A trackway, producing a digital archive using 3D photogrammetry and laser scanning. Here we compare the footprints at this site with those of American black bears, chimpanzees and humans, and we show that they resemble those of hominins more than ursids. In fact, the narrow step width corroborates the original interpretation of a small, cross-stepping bipedal hominin. However, the inferred foot proportions, gait parameters and 3D morphologies of footprints at site A are readily distinguished from those at site G, indicating that a minimum of two hominin taxa with different feet and gaits coexisted at Laetoli.

Giant lungfish genome elucidates the conquest of land by vertebrates.

Lungfishes belong to lobe-fined fish (Sarcopterygii) that, in the Devonian period, 'conquered' the land and ultimately gave rise to all land vertebrates, including humans1-3. Here we determine the chromosome-quality genome of the Australian lungfish (Neoceratodus forsteri), which is known to have the largest genome of any animal. The vast size of this genome, which is about 14× larger than that of humans, is attributable mostly to huge intergenic regions and introns with high repeat content (around 90%), the components of which resemble those of tetrapods (comprising mainly long interspersed nuclear elements) more than they do those of ray-finned fish. The lungfish genome continues to expand independently (its transposable elements are still active), through mechanisms different to those of the enormous genomes of salamanders. The 17 fully assembled lungfish macrochromosomes maintain synteny to other vertebrate chromosomes, and all microchromosomes maintain conserved ancient homology with the ancestral vertebrate karyotype. Our phylogenomic analyses confirm previous reports that lungfish occupy a key evolutionary position as the closest living relatives to tetrapods4,5, underscoring the importance of lungfish for understanding innovations associated with terrestrialization. Lungfish preadaptations to living on land include the gain of limb-like expression in developmental genes such as hoxc13 and sall1 in their lobed fins. Increased rates of evolution and the duplication of genes associated with obligate air-breathing, such as lung surfactants and the expansion of odorant receptor gene families (which encode proteins involved in detecting airborne odours), contribute to the tetrapod-like biology of lungfishes. These findings advance our understanding of this major transition during vertebrate evolution.

Turing, von Neumann, and the computational architecture of biological machines.

In the mid-1930s, the English mathematician and logician Alan Turing invented an imaginary machine which could emulate the process of manipulating finite symbolic configurations by human computers. His machine launched the field of computer science and provided a foundation for the modern-day programmable computer. A decade later, building on Turing's machine, the American-Hungarian mathematician John von Neumann invented an imaginary self-reproducing machine capable of open-ended evolution. Through his machine, von Neumann answered one of the deepest questions in Biology: Why is it that all living organisms carry a self-description in the form of DNA? The story behind how two pioneers of computer science stumbled on the secret of life many years before the discovery of the DNA double helix is not well known, not even to biologists, and you will not find it in biology textbooks. Yet, the story is just as relevant today as it was eighty years ago: Turing and von Neumann left a blueprint for studying biological systems as if they were computing machines. This approach may hold the key to answering many remaining questions in Biology and could even lead to advances in computer science.

Soft robotics informs how an early echinoderm moved.

The transition from sessile suspension to active mobile detritus feeding in early echinoderms (c.a. 500 Mya) required sophisticated locomotion strategies. However, understanding locomotion adopted by extinct animals in the absence of trace fossils and modern analogues is extremely challenging. Here, we develop a biomimetic soft robot testbed with accompanying computational simulation to understand fundamental principles of locomotion in one of the most enigmatic mobile groups of early stalked echinoderms-pleurocystitids. We show that these Paleozoic echinoderms were likely able to move over the sea bottom by means of a muscular stem that pushed the animal forward (anteriorly). We also demonstrate that wide, sweeping gaits could have been the most effective for these echinoderms and that increasing stem length might have significantly increased velocity with minimal additional energy cost. The overall approach followed here, which we call "Paleobionics," is a nascent but rapidly developing research agenda in which robots are designed based on extinct organisms to generate insights in engineering and evolution.

Humans plan for the near future to walk economically on uneven terrain.

Humans experience small fluctuations in their gait when walking on uneven terrain. The fluctuations deviate from the steady, energy-minimizing pattern for level walking and have no obvious organization. But humans often look ahead when they walk, and could potentially plan anticipatory fluctuations for the terrain. Such planning is only sensible if it serves some an objective purpose, such as maintaining constant speed or reducing energy expenditure, that is also attainable within finite planning capacity. Here, we show that humans do plan and perform optimal control strategies on uneven terrain. Rather than maintaining constant speed, they make purposeful, anticipatory speed adjustments that are consistent with minimizing energy expenditure. A simple optimal control model predicts economical speed fluctuations that agree well with experiments with humans (N = 12) walking on seven different terrain profiles (correlated with model [Formula: see text] , [Formula: see text] all terrains). Participants made repeatable speed fluctuations starting about six to eight steps ahead of each terrain feature (up to ±7.5 cm height difference each step, up to 16 consecutive features). Nearer features matter more, because energy is dissipated with each succeeding step's collision with ground, preventing momentum from persisting indefinitely. A finite horizon of continuous look-ahead and motor working space thus suffice to practically optimize for any length of terrain. Humans reason about walking in the near future to plan complex optimal control sequences.

NeuroMechFly, a neuromechanical model of adult Drosophila melanogaster.

Animal behavior emerges from an interaction between neural network dynamics, musculoskeletal properties and the physical environment. Accessing and understanding the interplay between these elements requires the development of integrative and morphologically realistic neuromechanical simulations. Here we present NeuroMechFly, a data-driven model of the widely studied organism, Drosophila melanogaster. NeuroMechFly combines four independent computational modules: a physics-based simulation environment, a biomechanical exoskeleton, muscle models and neural network controllers. To enable use cases, we first define the minimum degrees of freedom of the leg from real three-dimensional kinematic measurements during walking and grooming. Then, we show how, by replaying these behaviors in the simulator, one can predict otherwise unmeasured torques and contact forces. Finally, we leverage NeuroMechFly's full neuromechanical capacity to discover neural networks and muscle parameters that drive locomotor gaits optimized for speed and stability. Thus, NeuroMechFly can increase our understanding of how behaviors emerge from interactions between complex neuromechanical systems and their physical surroundings.

Estimation of skeletal kinematics in freely moving rodents.

Forming a complete picture of the relationship between neural activity and skeletal kinematics requires quantification of skeletal joint biomechanics during free behavior; however, without detailed knowledge of the underlying skeletal motion, inferring limb kinematics using surface-tracking approaches is difficult, especially for animals where the relationship between the surface and underlying skeleton changes during motion. Here we developed a videography-based method enabling detailed three-dimensional kinematic quantification of an anatomically defined skeleton in untethered freely behaving rats and mice. This skeleton-based model was constrained using anatomical principles and joint motion limits and provided skeletal pose estimates for a range of body sizes, even when limbs were occluded. Model-inferred limb positions and joint kinematics during gait and gap-crossing behaviors were verified by direct measurement of either limb placement or limb kinematics using inertial measurement units. Together we show that complex decision-making behaviors can be accurately reconstructed at the level of skeletal kinematics using our anatomically constrained model.

The condition for dynamic stability in humans walking with feedback control.

The walking human body is mechanically unstable. Loss of stability and falling is more likely in certain groups of people, such as older adults or people with neuromotor impairments, as well as in certain situations, such as when experiencing conflicting or distracting sensory inputs. Stability during walking is often characterized biomechanically, by measures based on body dynamics and the base of support. Neural control of upright stability, on the other hand, does not factor into commonly used stability measures. Here we analyze stability of human walking accounting for both biomechanics and neural control, using a modeling approach. We define a walking system as a combination of biomechanics, using the well known inverted pendulum model, and neural control, using a proportional-derivative controller for foot placement based on the state of the center of mass at midstance. We analyze this system formally and show that for any choice of system parameters there is always one periodic orbit. We then determine when this periodic orbit is stable, i.e. how the neural control gain values have to be chosen for stable walking. Following the formal analysis, we use this model to make predictions about neural control gains and compare these predictions with the literature and existing experimental data. The model predicts that control gains should increase with decreasing cadence. This finding appears in agreement with literature showing stronger effects of visual or vestibular manipulations at different walking speeds.

Identifying essential factors for energy-efficient walking control across a wide range of velocities in reflex-based musculoskeletal systems.

Humans can generate and sustain a wide range of walking velocities while optimizing their energy efficiency. Understanding the intricate mechanisms governing human walking will contribute to the engineering applications such as energy-efficient biped robots and walking assistive devices. Reflex-based control mechanisms, which generate motor patterns in response to sensory feedback, have shown promise in generating human-like walking in musculoskeletal models. However, the precise regulation of velocity remains a major challenge. This limitation makes it difficult to identify the essential reflex circuits for energy-efficient walking. To explore the reflex control mechanism and gain a better understanding of its energy-efficient maintenance mechanism, we extend the reflex-based control system to enable controlled walking velocities based on target speeds. We developed a novel performance-weighted least squares (PWLS) method to design a parameter modulator that optimizes walking efficiency while maintaining target velocity for the reflex-based bipedal system. We have successfully generated walking gaits from 0.7 to 1.6 m/s in a two-dimensional musculoskeletal model based on an input target velocity in the simulation environment. Our detailed analysis of the parameter modulator in a reflex-based system revealed two key reflex circuits that have a significant impact on energy efficiency. Furthermore, this finding was confirmed to be not influenced by setting parameters, i.e., leg length, sensory time delay, and weight coefficients in the objective cost function. These findings provide a powerful tool for exploring the neural bases of locomotion control while shedding light on the intricate mechanisms underlying human walking and hold significant potential for practical engineering applications.

A dynamic foot model for predictive simulations of human gait reveals causal relations between foot structure and whole-body mechanics.

The unique structure of the human foot is seen as a crucial adaptation for bipedalism. The foot's arched shape enables stiffening the foot to withstand high loads when pushing off, without compromising foot flexibility. Experimental studies demonstrated that manipulating foot stiffness has considerable effects on gait. In clinical practice, altered foot structure is associated with pathological gait. Yet, experimentally manipulating individual foot properties (e.g. arch height or tendon and ligament stiffness) is hard and therefore our understanding of how foot structure influences gait mechanics is still limited. Predictive simulations are a powerful tool to explore causal relationships between musculoskeletal properties and whole-body gait. However, musculoskeletal models used in three-dimensional predictive simulations assume a rigid foot arch, limiting their use for studying how foot structure influences three-dimensional gait mechanics. Here, we developed a four-segment foot model with a longitudinal arch for use in predictive simulations. We identified three properties of the ankle-foot complex that are important to capture ankle and knee kinematics, soleus activation, and ankle power of healthy adults: (1) compliant Achilles tendon, (2) stiff heel pad, (3) the ability to stiffen the foot. The latter requires sufficient arch height and contributions of plantar fascia, and intrinsic and extrinsic foot muscles. A reduced ability to stiffen the foot results in walking patterns with reduced push-off power. Simulations based on our model also captured the effects of walking with anaesthetised intrinsic foot muscles or an insole limiting arch compression. The ability to reproduce these different experiments indicates that our foot model captures the main mechanical properties of the foot. The presented four-segment foot model is a potentially powerful tool to study the relationship between foot properties and gait mechanics and energetics in health and disease.

Dimensionality of locomotor behaviors in developing C. elegans.

Adult animals display robust locomotion, yet the timeline and mechanisms of how juvenile animals acquire coordinated movements and how these movements evolve during development are not well understood. Recent advances in quantitative behavioral analyses have paved the way for investigating complex natural behaviors like locomotion. In this study, we tracked the swimming and crawling behaviors of the nematode Caenorhabditis elegans from postembryonic development through to adulthood. Our principal component analyses revealed that adult C. elegans swimming is low dimensional, suggesting that a small number of distinct postures, or eigenworms, account for most of the variance in the body shapes that constitute swimming behavior. Additionally, we found that crawling behavior in adult C. elegans is similarly low dimensional, corroborating previous studies. Further, our analysis revealed that swimming and crawling are distinguishable within the eigenworm space. Remarkably, young L1 larvae are capable of producing the postural shapes for swimming and crawling seen in adults, despite frequent instances of uncoordinated body movements. In contrast, late L1 larvae exhibit robust coordination of locomotion, while many neurons crucial for adult locomotion are still under development. In conclusion, this study establishes a comprehensive quantitative behavioral framework for understanding the neural basis of locomotor development, including distinct gaits such as swimming and crawling in C. elegans.

Cholinergic innervation topography in GBA-associated de novo Parkinson's disease patients.

The most common genetic risk factors for Parkinson's disease are GBA1 mutations, encoding the lysosomal enzyme glucocerebrosidase. Patients with GBA1 mutations (GBA-PD) exhibit earlier age of onset and faster disease progression with more severe cognitive impairments, postural instability and gait problems. These GBA-PD features suggest more severe cholinergic system pathologies. PET imaging with the vesicular acetylcholine transporter ligand 18F-F-fluoroethoxybenzovesamicol (18F-FEOBV PET) provides the opportunity to investigate cholinergic changes and their relationship to clinical features in GBA-PD. The study investigated 123 newly diagnosed, treatment-naïve Parkinson's disease subjects-with confirmed presynaptic dopaminergic deficits on PET imaging. Whole-gene GBA1 sequencing of saliva samples was performed to evaluate GBA1 variants. Patients underwent extensive neuropsychological assessment of all cognitive domains, motor evaluation with the Unified Parkinson's Disease Rating Scale, brain MRI, dopaminergic PET to measure striatal-to-occipital ratios of the putamen and 18F-FEOBV PET. We investigated differences in regional cholinergic innervation between GBA-PD carriers and non-GBA1 mutation carriers (non-GBA-PD), using voxel-wise and volume of interest-based approaches. The degree of overlap between t-maps from two-sample t-test models was quantified using the Dice similarity coefficient. Seventeen (13.8%) subjects had a GBA1 mutation. No significant differences were found in clinical features and dopaminergic ratios between GBA-PD and non-GBA-PD at diagnosis. Lower 18F-FEOBV binding was found in both the GBA-PD and non-GBA-PD groups compared to controls. Dice (P < 0.05, cluster size 100) showed good overlap (0.7326) between the GBA-PD and non-GBA-PD maps. GBA-PD patients showed more widespread reduction in 18F-FEOBV binding than non-GBA-PD when compared to controls in occipital, parietal, temporal and frontal cortices (P < 0.05, FDR-corrected). In volume of interest analyses (Bonferroni corrected), the left parahippocampal gyrus was more affected in GBA-PD. De novo GBA-PD show a distinct topography of regional cholinergic terminal ligand binding. Although the Parkinson's disease groups were not distinguishable clinically, in comparison to healthy controls, GBA-PD showed more extensive cholinergic denervation compared to non-GBA-PD. A larger group is needed to validate these findings. Our results suggest that de novo GBA-PD and non-GBA-PD show differential patterns of cholinergic system changes before clinical phenotypic differences between carriers versus non-carrier groups are observable.

Designing minimal and scalable insect-inspired multi-locomotion millirobots.

In ant colonies, collectivity enables division of labour and resources1-3 with great scalability. Beyond their intricate social behaviours, individuals of the genus Odontomachus4, also known as trap-jaw ants, have developed remarkable multi-locomotion mechanisms to 'escape-jump' upwards when threatened, using the sudden snapping of their mandibles5, and to negotiate obstacles by leaping forwards using their legs6. Emulating such diverse insect biomechanics and studying collective behaviours in a variety of environments may lead to the development of multi-locomotion robotic collectives deployable in situations such as emergency relief, exploration and monitoring7; however, reproducing these abilities in small-scale robotic systems with simple design and scalability remains a key challenge. Existing robotic collectives8-12 are confined to two-dimensional surfaces owing to limited locomotion, and individual multi-locomotion robots13-17 are difficult to scale up to large groups owing to the increased complexity, size and cost of hardware designs, which hinder mass production. Here we demonstrate an autonomous multi-locomotion insect-scale robot (millirobot) inspired by trap-jaw ants that addresses the design and scalability challenges of small-scale terrestrial robots. The robot's compact locomotion mechanism is constructed with minimal components and assembly steps, has tunable power requirements, and realizes five distinct gaits: vertical jumping for height, horizontal jumping for distance, somersault jumping to clear obstacles, walking on textured terrain and crawling on flat surfaces. The untethered, battery-powered millirobot can selectively switch gaits to traverse diverse terrain types, and groups of millirobots can operate collectively to manipulate objects and overcome obstacles. We constructed the ten-gram palm-sized prototype-the smallest and lightest self-contained multi-locomotion robot reported so far-by folding a quasi-two-dimensional metamaterial18 sandwich formed of easily integrated mechanical, material and electronic layers, which will enable assembly-free mass-manufacturing of robots with high task efficiency, flexibility and disposability.

A soft microrobot with highly deformable 3D actuators for climbing and transitioning complex surfaces.

The climbing microrobots have attracted growing attention due to their promising applications in exploration and monitoring of complex, unstructured environments. Soft climbing microrobots based on muscle-like actuators could offer excellent flexibility, adaptability, and mechanical robustness. Despite the remarkable progress in this area, the development of soft microrobots capable of climbing on flat/curved surfaces and transitioning between two different surfaces remains elusive, especially in open spaces. In this study, we address these challenges by developing voltage-driven soft small-scale actuators with customized 3D configurations and active stiffness adjusting. Combination of programmed strain distributions in liquid crystal elastomers (LCEs) and buckling-driven 3D assembly, guided by mechanics modeling, allows for voltage-driven, complex 3D-to-3D shape morphing (bending angle > 200°) at millimeter scales (from 1 to 10 mm), which is unachievable previously. These soft actuators enable development of morphable electroadhesive footpads that can conform to different curved surfaces and stiffness-variable smart joints that allow different locomotion gaits in a single microrobot. By integrating such morphable footpads and smart joints with a deformable body, we report a multigait, soft microrobot (length from 6 to 90 mm, and mass from 0.2 to 3 g) capable of climbing on surfaces with diverse shapes (e.g., flat plane, cylinder, wavy surface, wedge-shaped groove, and sphere) and transitioning between two distinct surfaces. We demonstrate that the microrobot could navigate from one surface to another, recording two corresponding ceilings when carrying an integrated microcamera. The developed soft microrobot can also flip over a barrier, survive extreme compression, and climb bamboo and leaf.

Walking is like slithering: A unifying, data-driven view of locomotion.

Legged movement is ubiquitous in nature and of increasing interest for robotics. Most legged animals routinely encounter foot slipping, yet detailed modeling of multiple contacts with slipping exceeds current simulation capacity. Here we present a principle that unifies multilegged walking (including that involving slipping) with slithering and Stokesian (low Reynolds number) swimming. We generated data-driven principally kinematic models of locomotion for walking in low-slip animals (Argentine ant, 4.7% slip ratio of slipping to total motion) and for high-slip robotic systems (BigANT hexapod, slip ratio 12 to 22%; Multipod robots ranging from 6 to 12 legs, slip ratio 40 to 100%). We found that principally kinematic models could explain much of the variability in body velocity and turning rate using body shape and could predict walking behaviors outside the training data. Most remarkably, walking was principally kinematic irrespective of leg number, foot slipping, and turning rate. We find that grounded walking, with or without slipping, is governed by principally kinematic equations of motion, functionally similar to frictional swimming and slithering. Geometric mechanics thus leads to a unified model for swimming, slithering, and walking. Such commonality may shed light on the evolutionary origins of animal locomotion control and offer new approaches for robotic locomotion and motion planning.

Low effective mechanical advantage of giraffes' limbs during walking reveals trade-off between limb length and locomotor performance.

Giraffes (Giraffa camelopardalis) possess specialized locomotor morphology, namely elongate and gracile distal limbs. While this contributes to their overall height and enhances feeding behavior, we propose that the combination of long limb segments and modest muscle lever arms results in low effective mechanical advantage (EMA, the ratio of in-lever to out-lever moment arms), when compared with other cursorial mammals. To test this, we used a combination of experimentally measured kinematics and ground reaction forces (GRFs), musculoskeletal modeling, and inverse dynamics to calculate giraffe forelimb EMA during walking. Giraffes walk with an EMA of 0.34 (±0.05 SD), with no evident association with speed within their walking gait. Giraffe EMA was about four times lower than expectations extrapolated from other mammals, ranging from 0.03 to 297 kg, and this provides further evidence that EMA plateaus or even diminishes in mammals exceeding horse size. We further tested the idea that limb segment length is a factor which determines EMA, by modeling the GRF and muscle moment arms in the extinct giraffid Sivatherium giganteum and the other extant giraffid, Okapia johnstoni. Giraffa and Okapia shared similar EMA, despite a four to sixfold difference in body mass (Okapia EMA = 0.38). In contrast, Sivatherium, sharing a similar body mass with Giraffa, had greater EMA (0.59), which we propose reflects behavioral differences, such as a somewhat increased capability for athletic performance. Our modeling approach suggests that limb length is a determinant of GRF moment arm magnitude and that unless muscle moment arms scale isometrically with limb length, tall mammals are prone to low EMA.

Vertebral level specific modulation of paraspinal muscle activity based on vestibular signals during walking.

Evoking muscle responses by electrical vestibular stimulation (EVS) may help to understand the contribution of the vestibular system to postural control. Although paraspinal muscles play a role in postural stability, the vestibulo-muscular coupling of these muscles during walking has rarely been studied. This study aimed to investigate how vestibular signals affect paraspinal muscle activity at different vertebral levels during walking with preferred and narrow step width. Sixteen healthy participants were recruited. Participants walked on a treadmill for 8 min at 78 steps/min and 2.8 km/h, at two different step width, either with or without EVS. Bipolar electromyography was recorded bilaterally from the paraspinal muscles at eight vertebral levels from cervical to lumbar. Coherence, gain, and delay of EVS and EMG responses were determined. Significant EVS-EMG coupling (P < 0.01) was found at ipsilateral and/or contralateral heel strikes. This coupling was mirrored between left and right relative to the midline of the trunk and between the higher and lower vertebral levels, i.e. a peak occurred at ipsilateral heel strike at lower levels, whereas it occurred at contralateral heel strike at higher levels. EVS-EMG coupling only partially coincided with peak muscle activity. EVS-EMG coherence slightly, but not significantly, increased when walking with narrow steps. No significant differences were found in gain and phase between the vertebral levels or step width conditions. In summary, vertebral level specific modulation of paraspinal muscle activity based on vestibular signals might allow a fast, synchronized, and spatially co-ordinated response along the trunk during walking. KEY POINTS: Mediolateral stabilization of gait requires an estimate of the state of the body, which is affected by vestibular afference. During gait, the heavy trunk segment is controlled by phasic paraspinal muscle activity and in rodents the medial and lateral vestibulospinal tracts activate these muscles. To gain insight in vestibulospinal connections in humans and their role in gait, we recorded paraspinal surface EMG of cervical to lumbar paraspinal muscles, and characterized coherence, gain and delay between EMG and electrical vestibular stimulation, during slow walking. Vestibular stimulation caused phasic, vertebral level specific modulation of paraspinal muscle activity at delays of around 40 ms, which was mirrored between left, lower and right, upper vertebral levels. Our results indicate that vestibular afference causes fast, synchronized, and spatially co-ordinated responses of the paraspinal muscles along the trunk, that simultaneously contribute to stabilizing the centre of mass trajectory and to keeping the head upright.

Collagen architecture and biomechanics of gracilis and adductor longus muscles from children with cerebral palsy.

Cerebral palsy (CP) describes some upper motoneuron disorders due to non-progressive disturbances occurring in the developing brain that cause progressive changes to muscle. While longer sarcomeres increase muscle stiffness in patients with CP compared to typically developing (TD) patients, changes in extracellular matrix (ECM) architecture can increase stiffness. Our goal was to investigate how changes in muscle and ECM architecture impact muscle stiffness, gait and joint function in CP. Gracilis and adductor longus biopsies were collected from children with CP undergoing tendon lengthening surgery for hamstring and hip adduction contractures, respectively. Gracilis biopsies were collected from TD patients undergoing anterior cruciate ligament reconstruction surgery with hamstring autograft. Muscle mechanical testing, two-photon imaging and hydroxyproline assay were performed on biopsies. Corresponding data were compared to radiographic hip displacement in CP adductors (CPA), gait kinematics in CP hamstrings (CPH), and joint range of motion in CPA and CPH. We found at matched sarcomere lengths muscle stiffness and collagen architecture were similar between TD and CP hamstrings. However, CPH stiffness (R2 = 0.1973), collagen content (R2 = 0.5099) and cross-linking (R2 = 0.3233) were correlated to decreased knee range of motion. Additionally, we observed collagen fibres within the muscle ECM increase alignment during muscular stretching. These data demonstrate that while ECM architecture is similar between TD and CP hamstrings, collagen fibres biomechanics are sensitive to muscle strain and may be altered at longer in vivo sarcomere lengths in CP muscle. Future studies could evaluate the impact of ECM architecture on TD and CP muscle stiffness across in vivo operating ranges. KEY POINTS: At matched sarcomere lengths, gracilis muscle mechanics and collagen architecture are similar in TD patients and patients with CP. In both TD and CP muscles, collagen fibres dynamically increase their alignment during muscle stretching. Aspects of muscle mechanics and collagen architecture are predictive of in vivo knee joint motion and radiographic hip displacement in patients with CP. Longer sarcomere lengths in CP muscle in vivo may alter collagen architecture and biomechanics to drive deficits in joint mobility and gait function.