How we perceive the width of grasped objects: Insights into the central processes that govern proprioceptive judgements.

Low-level proprioceptive judgements involve a single frame of reference, whereas high-level proprioceptive judgements are made across different frames of reference. The present study systematically compared low-level (grasp → $\rightarrow$ grasp) and high-level (vision → $\rightarrow$ grasp, grasp → $\rightarrow$ vision) proprioceptive tasks, and quantified the consistency of grasp → $\rightarrow$ vision and possible reciprocal nature of related high-level proprioceptive tasks. Experiment 1 (n = 30) compared performance across vision → $\rightarrow$ grasp, a grasp → $\rightarrow$ vision and a grasp → $\rightarrow$ grasp tasks. Experiment 2 (n = 30) compared performance on the grasp → $\rightarrow$ vision task between hands and over time. Participants were accurate (mean absolute error 0.27 cm [0.20 to 0.34]; mean [95% CI]) and precise ( R 2 $R^2$ = 0.95 [0.93 to 0.96]) for grasp → $\rightarrow$ grasp judgements, with a strong correlation between outcomes (r = -0.85 [-0.93 to -0.70]). Accuracy and precision decreased in the two high-level tasks ( R 2 $R^2$ = 0.86 and 0.89; mean absolute error = 1.34 and 1.41 cm), with most participants overestimating perceived width for the vision → $\rightarrow$ grasp task and underestimating it for grasp → $\rightarrow$ vision task. There was minimal correlation between accuracy and precision for these two tasks. Converging evidence indicated performance was largely reciprocal (inverse) between the vision → $\rightarrow$ grasp and grasp → $\rightarrow$ vision tasks. Performance on the grasp → $\rightarrow$ vision task was consistent between dominant and non-dominant hands, and across repeated sessions a day or week apart. Overall, there are fundamental differences between low- and high-level proprioceptive judgements that reflect fundamental differences in the cortical processes that underpin these perceptions. Moreover, the central transformations that govern high-level proprioceptive judgements of grasp are personalised, stable and reciprocal for reciprocal tasks. KEY POINTS: Low-level proprioceptive judgements involve a single frame of reference (e.g. indicating the width of a grasped object by selecting from a series of objects of different width), whereas high-level proprioceptive judgements are made across different frames of reference (e.g. indicating the width of a grasped object by selecting from a series of visible lines of different length). We highlight fundamental differences in the precision and accuracy of low- and high-level proprioceptive judgements. We provide converging evidence that the neural transformations between frames of reference that govern high-level proprioceptive judgements of grasp are personalised, stable and reciprocal for reciprocal tasks. This stability is likely key to precise judgements and accurate predictions in high-level proprioception.

Transcutaneous spinal stimulation in people with and without spinal cord injury: Effect of electrode placement and trains of stimulation on threshold intensity.

Transcutaneous spinal cord stimulation (TSS) is purported to improve motor function in people after spinal cord injury (SCI). However, several methodology aspects are yet to be explored. We investigated whether stimulation configuration affected the intensity needed to elicit spinally evoked motor responses (sEMR) in four lower limb muscles bilaterally. Also, since stimulation intensity for therapeutic TSS (i.e., trains of stimulation, typically delivered at 15-50 Hz) is sometimes based on the single-pulse threshold intensity, we compared these two stimulation types. In non-SCI participants (n = 9) and participants with a SCI (n = 9), three different electrode configurations (cathode-anode); L1-midline (below the umbilicus), T11-midline and L1-ASIS (anterior superior iliac spine; non-SCI only) were compared for the sEMR threshold intensity using single pulses or trains of stimulation which were recorded in the vastus medialis, medial hamstring, tibialis anterior, medial gastrocnemius muscles. In non-SCI participants, the L1-midline configuration showed lower sEMR thresholds compared to T11-midline (p = 0.002) and L1-ASIS (p < 0.001). There was no difference between T11-midline and L1-midline for participants with SCI (p = 0.245). Spinally evoked motor response thresholds were ~13% lower during trains of stimulation compared to single pulses in non-SCI participants (p < 0.001), but not in participants with SCI (p = 0.101). With trains of stimulation, threshold intensities were slightly lower and the incidence of sEMR was considerably lower. Overall, stimulation threshold intensities were generally lower with the L1-midline electrode configuration and is therefore preferred. While single-pulse threshold intensities may overestimate threshold intensities for therapeutic TSS, tolerance to trains of stimulation will be the limiting factor in most cases.

Encouraging responsible reporting practices in the Instructions to Authors of neuroscience and physiology journals: There is room to improve.

Journals can substantially influence the quality of research reports by including responsible reporting practices in their Instructions to Authors. We assessed the extent to which 100 journals in neuroscience and physiology required authors to report methods and results in a rigorous and transparent way. For each journal, Instructions to Authors and any referenced reporting guideline or checklist were downloaded from journal websites. Twenty-two questions were developed to assess how journal Instructions to Authors address fundamental aspects of rigor and transparency in five key reporting areas. Journal Instructions to Authors and all referenced external guidelines and checklists were audited against these 22 questions. Of the full sample of 100 Instructions to Authors, 34 did not reference any external reporting guideline or checklist. Reporting whether clinical trial protocols were pre-registered was required by 49 journals and encouraged by 7 others. Making data publicly available was encouraged by 64 journals; making (processing or statistical) code publicly available was encouraged by ∼30 of the journals. Other responsible reporting practices were mentioned by less than 20 of the journals. Journals can improve the quality of research reports by mandating, or at least encouraging, the responsible reporting practices highlighted here.

Upper Limb Function but Not Proprioception is Impaired in Essential Tremor: A Between-Groups Study and Causal Mediation Analysis.

Background: Essential tremor (ET) is characterized by abnormal oscillatory muscle activity and cerebellar involvement, factors that can lead to proprioceptive deficits, especially in active tasks. The present study aimed to quantify the severity of proprioceptive deficits in people with ET and estimate how these contribute to functional impairments. Methods: Upper limb sensory, proprioceptive and motor function was assessed inindividuals with ET (n = 20) and healthy individuals (n = 22). To measure proprioceptive ability, participants discriminated the width of grasped objects and the weight of objects liftedwith the wrist extensors. Causal mediation analysis was used to estimate the extentthat impairments in upper limb function in ET was mediated by proprioceptive ability. Results: Participants with ET had impaired upper limb function in all outcomes, and had greater postural and kinetic tremor. There were no differences between groups in proprioceptive discrimination of width (between-group mean difference [95% CI]: 0.32 mm [-0.23 to 0.87 mm]) or weight (-1.12 g [-7.31 to 5.07 g]). Causal mediation analysis showed the effect of ET on upper limb function was not mediated by proprioceptive ability. Conclusions: Upper limb function but not proprioception was impaired in ET. The effect of ET on motor function was not mediated by proprioception. These results indicate that the central nervous system of people with ET is able to accommodate mild to moderate tremor in active proprioceptive tasks that rely primarily on afferent signals from muscle spindles.

Muscle electromyographic activity normalized to maximal muscle activity, not to Mmax, better represents voluntary activation.

In human applied physiology studies, the amplitude of recorded muscle electromyographic activity (EMG) is often normalized to maximal EMG recorded during a maximal voluntary contraction. When maximal contractions cannot be reliably obtained (e.g. in people with muscle paralysis, anterior cruciate ligament injury, or arthritis), EMG is sometimes normalized to the maximal compound muscle action potiential evoked by stimulation, the Mmax. However, it is not known how these two methods of normalization affect the conclusions and comparability of studies. To address this limitation, we investigated the relationship between voluntary muscle activation and EMG normalized either to maximal EMG or to Mmax. Twenty-five able-bodied adults performed voluntary isometric ankle plantarflexion contractions to a range of percentages of maximal voluntary torque. Ankle torque, plantarflexor muscle EMG, and voluntary muscle activation measured by twitch interpolation were recorded. EMG recorded at each contraction intensity was normalized to maximal EMG or to Mmax for each plantarflexor muscle, and the relationship between the two normalization approaches quantified. A slope >1 indicated EMG amplitude normalized to maximal EMG (vertical axis) was greater than EMG normalized to Mmax (horizontal axis). Mean estimates of the slopes were large and had moderate precision: soleus 8.7 (95% CI 6.9 to 11.0), medial gastrocnemius 13.4 (10.5 to 17.0), lateral gastrocnemius 11.4 (9.4 to 14.0). This indicates EMG normalized to Mmax is approximately eleven times smaller than EMG normalized to maximal EMG. Normalization to maximal EMG gave closer approximations to the level of voluntary muscle activation assessed by twitch interpolation.

Quality Output Checklist and Content Assessment (QuOCCA): a new tool for assessing research quality and reproducibility.

Research must be well designed, properly conducted and clearly and transparently reported. Our independent medical research institute wanted a simple, generic tool to assess the quality of the research conducted by its researchers, with the goal of identifying areas that could be improved through targeted educational activities. Unfortunately, none was available, thus we devised our own. Here, we report development of the Quality Output Checklist and Content Assessment (QuOCCA), and its application to publications from our institute's scientists. Following consensus meetings and external review by statistical and methodological experts, 11 items were selected for the final version of the QuOCCA: research transparency (items 1-3), research design and analysis (items 4-6) and research reporting practices (items 7-11). Five pairs of raters assessed all 231 articles published in 2017 and 221 in 2018 by researchers at our institute. Overall, the results were similar between years and revealed limited engagement with several recommended practices highlighted in the QuOCCA. These results will be useful to guide educational initiatives and their effectiveness. The QuOCCA is brief and focuses on broadly applicable and relevant concepts to open, high-quality, reproducible and well-reported science. Thus, the QuOCCA could be used by other biomedical institutions and individual researchers to evaluate research publications, assess changes in research practice over time and guide the discussion about high-quality, open science. Given its generic nature, the QuOCCA may also be useful in other research disciplines.

Proprioception: a new look at an old concept.

Proprioception, which can be defined as the awareness of the mechanical and spatial state of the body and its musculoskeletal parts, is critical to motor actions and contributes to our sense of body ownership. To date, clinical proprioceptive tests have focused on a person's ability to detect, discriminate, or match limb positions or movements, and reveal that the strength of the relationship between deficits in proprioception and physical function varies widely. Unfortunately, these tests fail to assess higher-level proprioceptive abilities. In this Perspective, we propose that to understand fully the link between proprioception and function, we need to look beyond traditional clinical tests of proprioception. Specifically, we present a novel framework for human proprioception assessment that is divided into two categories: low-level and high-level proprioceptive judgments. Low-level judgments are those made in a single frame of reference and are the types of judgments made in traditional proprioceptive tests (i.e., detect, discriminate or match). High-level proprioceptive abilities involve proprioceptive judgments made in a different frame of reference. For example, when a person indicates where their hand is located in space. This framework acknowledges that proprioception is complex and multifaceted and that tests of proprioception should not be viewed as interchangeable, but rather as complimentary. Crucially, it provides structure to the way researchers and clinicians can approach proprioception and its assessment. We hope this Perspective serves as the catalyst for discussion and new lines of investigation.

Transcutaneous spinal cord stimulation combined with locomotor training to improve walking ability in people with chronic spinal cord injury: study protocol for an international multi-centred double-blinded randomised sham-controlled trial (eWALK).

STUDY DESIGN: An international multi-centred, double-blinded, randomised sham-controlled trial (eWALK). OBJECTIVE: To determine the effect of 12 weeks of transcutaneous spinal stimulation (TSS) combined with locomotor training on walking ability in people with spinal cord injury (SCI). SETTING: Dedicated SCI research centres in Australia, Spain, USA and Scotland. METHODS: Fifty community-dwelling individuals with chronic SCI will be recruited. Participants will be eligible if they have bilateral motor levels between T1 and T11, a reproducible lower limb muscle contraction in at least one muscle group, and a Walking Index for SCI II (WISCI II) between 1 and 6. Eligible participants will be randomised to one of two groups, either the active stimulation group or the sham stimulation group. Participants allocated to the stimulation group will receive TSS combined with locomotor training for three 30-min sessions a week for 12 weeks. The locomotor sessions will include walking on a treadmill and overground. Participants allocated to the sham stimulation group will receive the same locomotor training combined with sham stimulation. The primary outcome will be walking ability with stimulation using the WISCI II. Secondary outcomes will record sensation, strength, spasticity, bowel function and quality of life. TRIAL REGISTRATION: ANZCTR.org.au identifier ACTRN12620001241921.

Do interoception and attending to the upper limbs affect body ownership and body representation in the grasp illusion?

Passively grasping an unseen artificial finger induces ownership over this finger and an illusory coming together of one's index fingers: a grasp illusion. Here we determine how interoceptive ability and attending to the upper limbs influence this illusion. Participants passively grasped an unseen artificial finger with their left index finger and thumb for 3 min while their right index finger, located 12 cm below, was lightly clamped. Experiment 1 (n = 30) investigated whether the strength of the grasp illusion (perceived index finger spacing and perceived ownership) is related to a person's level of interoceptive accuracy (modified heartbeat counting task) and sensibility (Noticing subscale of the Multidimensional Assessment of Interoceptive Awareness). Experiment 2 (n = 30) investigated the effect of providing verbal or tactile cues to guide participants' attention to their upper limbs. On their own, neither interoceptive accuracy and sensibility or verbal and tactile cueing had an effect on the grasp illusion. However, verbal cueing increased the strength of the grasp illusion in individuals with lower interoceptive ability. Across the observed range of interoceptive accuracy and sensibility, verbal cueing decreased perceived index spacing by 5.6 cm [1.91 to 9.38] (mean [95%CI]), and perceived ownership by ∼3 points on a 7-point Likert scale (slope -0.93 [-1.72 to -0.15]). Thus, attending to the upper limbs via verbal cues increases the strength of the grasp illusion in a way that is inversely proportional to a person's level of interoceptive accuracy and sensibility.

Negligible epimuscular myofascial force transmission between the human rectus femoris and vastus lateralis muscles in passive conditions.

PURPOSE: There have been contradictory reports of the effects of epimuscular myofascial force transmission in humans. This study investigated the transmission of myofascial force to the human vastus lateralis muscle by determining whether vastus lateralis slack angle changed with hip angle. Since the distance between the origin and insertion of the vastus lateralis muscle does not change when hip angle changes, any change in vastus lateralis slack angle with hip position can be attributed to epimuscular myofascial force transmission. METHODS: Nineteen young adults were tested in hip flexed ([Formula: see text]) and neutral ([Formula: see text]) positions. Ultrasound images of the vastus lateralis muscle were obtained as the knee was passively flexed at [Formula: see text]/s. The knee angle at which vastus lateralis muscle fascicles began to lengthen was used to identify muscle slack angle. RESULTS: Overall, there was a negligible effect of hip position on vastus lateralis slack angle ([Formula: see text] [[Formula: see text] to 1.9]; mean [95% confidence interval]). However, a small and variable effect was noted in 3/19 participants. CONCLUSION: This result indicates that, over the range of joint angles tested here, there is little or no epimuscular myofascial force transmission between the vastus lateralis muscle and neighbouring bi-articular structures under passive conditions. More broadly, this result provides additional evidence that epimuscular myofascial force transmission tends to be small and variable under passive conditions in healthy human muscle.

Movement of the ribs in supine humans for small and large changes in lung volume.

An object-tracking algorithm was used on computed tomography (CT) images of the thorax from six healthy participants and nine participants with chronic obstructive pulmonary disease (COPD) to describe the movement of the ribs between the static lung volumes of functional residual capacity (FRC) and total lung capacity (TLC). The continuous motion of the ribs during tidal breathing was also described using four-dimensional CT datasets from seven participants with thoracic esophageal malignancies. Rib motion was defined relative to a local joint coordinate system where rotations about the axes that predominantly affected the anteroposterior and transverse diameters of the rib cage were referred to as pump-handle and bucket-handle movements, respectively. Between TLC and FRC, pump-handle movements were 1.8 times larger in healthy participants than in participants with COPD, in line with their 1.6 times larger inspiratory capacities. However, when rib motion was normalized to the change in lung volume, pump-handle movements were similar for healthy participants and participants with COPD. We found no differences in bucket-handle movements between participant groups before and after normalization. Pump-handle movement was the dominant rib motion between FRC and TLC, on average four times greater than bucket-handle movement in healthy participants. For expiratory tidal volume, pump-handle movements were 20% smaller than bucket-handle movements. When normalized to tidal volume and compared with inspiratory capacity, pump-handle movements were smaller and bucket-handle movements were larger during tidal breathing. The findings suggest that the pump-handle and bucket-handle components of rib motion vary for small and large changes in lung volume.NEW & NOTEWORTHY Rib movements over inspiratory capacity are comparable for healthy participants and participants with chronic obstructive pulmonary disease when normalized to the change in lung volume. The kinematics of the ribs during tidal breathing were described from four-dimensional computed tomography images. For large changes in lung volume with inspiratory capacity, pump-handle movements of the ribs are four times greater than bucket-handle movements, whereas at tidal volume, pump-handle movements are 20% smaller than bucket-handle movements.

Estimation of maximal muscle electromyographic activity from the relationship between muscle activity and voluntary activation.

Maximal muscle activity recorded with surface electromyography (EMG) is an important neurophysiological measure. It is frequently used to normalize EMG activity recorded during passive or active movement. However, the true maximal muscle activity cannot be determined in people with impaired capacity to voluntarily activate their muscles. Here, we determined whether maximal muscle activity can be estimated from muscle activity produced during submaximal voluntary activation. Twenty-five able-bodied adults (18 males, mean age 29 yr, range 19-64 yr) participated in the study. Participants were seated with the knee flexed 90° and the ankle in 5° of dorsiflexion from neutral. Participants performed isometric voluntary ankle plantarflexion contractions at target torques, in random order: 1, 5, 10, 15, 25, 50, 75, 90, 95, and 100% of maximal voluntary torque. Ankle torque, muscle activity in soleus, medial and lateral gastrocnemius muscles, and voluntary muscle activation determined using twitch interpolation were recorded. There was a strong loge-linear relationship between measures of muscle activation and muscle activity in all three muscles tested. Linear mixed models were fitted to muscle activation and loge-transformed EMG data. Each 1% increase in muscle activation increased muscle activity by a mean of 0.027 ln(mV) [95% confidence interval (CI) 0.025 to 0.029 ln(mV)] in soleus, 0.025 ln(mV) [0.022 to 0.028 ln(mV)] in medial gastrocnemius, and 0.028 ln(mV) [0.026 to 0.030 ln(mV)] in lateral gastrocnemius. The relationship between voluntary muscle activation and muscle activity can be described with simple mathematical functions. In future, it should be possible to normalize recorded muscle activity using these types of functions.NEW & NOTEWORTHY Muscle activity is often normalized to maximal muscle activity; however, it is difficult to obtain accurate measures of maximal muscle activity in people with impaired voluntary neural drive. We determined the relationship between voluntary muscle activation and plantarflexor muscle activity across a broad range of muscle activation values in able-bodied people. The relationship between voluntary muscle activation and muscle activity can be described with simple mathematical functions capable of estimating maximal muscle activity.

Judgements of hand location and hand spacing show minimal proprioceptive drift.

With a visual memory of where our hands are, their perceived location drifts. We investigated whether the perceived location of one hand or the spacing between two hands drifts in the absence of visual memories or cues. In 30 participants (17 females, mean age 27 years, range 20-45 years), perceived location of the right index finger was assessed when it was 10 cm to the right or left of the midline. Perceived spacing between the index fingers was assessed when they were spaced 20 cm apart, centred on the midline. Testing included two conditions, one with ten measures at 30 s intervals and another where a 3 min delay was introduced after the fifth measure. Participants responded by selecting a point on a ruler or a line from a series of lines of different lengths. Overall, participants mislocalised their hands closer to the midline. However, there was little to no drift in perceived index finger location when measures were taken at regular intervals (ipsilateral slope: 0.073 cm/measure [[Formula: see text] to 0.160], mean [99% CI]; contralateral slope: 0.045 cm/measure [[Formula: see text] to 0.120]), or across a 3 min delay (ipsilateral: ([Formula: see text] cm [[Formula: see text] to 0.17]; contralateral: [Formula: see text] cm [[Formula: see text] to 0.24]). There was a slight drift in perceived spacing when measures were taken at regular intervals (slope: [Formula: see text] cm/measure [[Formula: see text] to [Formula: see text]]), but none across a 3 min delay (0.08 cm [[Formula: see text] to 1.24]). Thus, proprioceptive-based perceptions of where our hands are located or how they are spaced drift minimally or not at all, indicating these perceptions are stable.

Stability of perception of the hand's aperture in a grasp.

KEY POINTS: How we judge the location of our body parts can be affected by a range of factors that change how our brain interprets proprioceptive signals. We examined the effect of several such factors on how we perceive an object's width and the spacing between our thumb and fingers when grasping. Grasp-related perceptions were slightly wider when using all digits, in line with our tendency to grasp larger objects with the entire hand. Surprisingly, these perceptions were not affected by the frames of reference for judgements (object width versus grasp aperture), whether the object was grasped actively or passively, or the strength of the grasp. These results show that the brain maintains a largely stable representation of the hand when grasping stationary objects. This stability may underpin our dexterity when grasping a vast array of objects. ABSTRACT: Various factors can alter how the brain interprets proprioceptive signals, leading to errors in how we perceive our body and execute motor tasks. This study determined the effect of critical factors on hand-based perceptions. In Experiment 1, 20 participants grasped without lifting an unseen 6.5 cm-wide object with two grasp configurations: thumb and all fingers, and thumb and index finger. Participants reported perceived grasp aperture (body reference frame) or perceived object width (external reference frame) using visual charts. In Experiment 2, 20 participants grasped the object with three grasp intensities (1, 5 and 15% maximal grasp force) actively or passively and reported perceived grasp aperture. A follow-up experiment addressed whether results from Experiment 2 were influenced by the external force applied during passive grasp. Overall, there was a mean difference of 0.38 cm (95% confidence interval (CI), 0.12 to 0.63) between the two grasp configurations (all digits compared to thumb and index finger). Perceived object width compared to perceived grasp aperture differed by only -0.04 cm (95% CI, -0.30 to 0.21). There was no real effect of grasp intensity on perceived grasp aperture (-0.01 cm; 95% CI, -0.03 to 0.01) or grasp type (active versus passive; 0.18 cm; 95% CI, -0.19 to 0.55). Overall, grasp-related perceptions are slightly wider when using all digits, in line with our tendency to grasp larger objects with the entire hand. The other factors - frame of reference, grasp intensity and grasp type - had no meaningful effect on these perceptions. These results provide evidence that the brain maintains a largely stable representation of the hand.