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 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.

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.

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.

The hidden hand is perceived closer to midline.

Whether visible or not, knowing the location of our hands is fundamental to how we perceive ourselves and interact with our environment. The present study investigated perceived hand location in the absence of vision in 30 participants. Their right index finger was placed 10, 20 or 30 cm away on either side of the body midline, with and without their left index finger placed 10 cm to the left of the right index. On average, at each position, participants perceived their right hand closer to the body midline than it actually was. This underestimation increased linearly with increased distance of the hand from body midline [slope 0.77 (0.74 to 0.81), mean (95% CI)]. Participants made smaller errors in perceived hand location when the right hand was in the contralateral workspace [mean difference 2.13 cm (1.57 to 2.69)]. Presence of the left hand on the support surface had little or no effect on perceived location of the right hand [mean difference [Formula: see text] cm ([Formula: see text] to 0.02)]. Overall, participants made systematic perceptual errors immediately after hand placement. The magnitude of these errors grew linearly as the hand got further away from the body midline. Because of their magnitude, these errors may contribute to errors in motor planning when visual feedback is not available. Also, these errors are important for studies in which perceived hand location is assessed after some time, for example, when studying illusions of body ownership and proprioceptive drift.

Involuntary hamstring muscle activity reduces passive hip range of motion during the straight leg raise test: a stimulation study in healthy people.

BACKGROUND: Involuntary hamstring muscle activity is present in some people during the straight leg raise test, but it is not known to what extent involuntary muscle activity limits passive joint range of motion. This study aimed to determine whether small amounts of involuntary hamstring activity limit passive hip range of motion during the straight leg raise test in healthy people. METHODS: Thirty healthy subjects were recruited from The University of Sydney. As the hamstring muscles were continuously stimulated to generate 0, 2.5, 5, 7.5 and 10% of knee flexion maximal voluntary contraction force, an investigator blinded to the amount of stimulation performed a straight leg raise test by passively raising the tested leg while keeping the knee extended. The test was stopped when the knee started to flex, at which point hip range of motion was recorded. RESULTS: On average, passive hip range of motion decreased by 0.6° for every 1% increase in knee flexion force caused by muscle activation (95% CI 0.3 to 0.9°, p = 0.0012). Subjects were instructed to fully relax when the straight leg raise test was performed, but a small amount of involuntary muscle activity (median 2.4% of maximal activation) was present during the trial without stimulation. CONCLUSIONS: Small amounts of involuntary hamstring muscles activity reduce passive hip range of motion during the straight leg raise test in healthy people. TRIAL REGISTRATION: The protocol for this study was registered with the Open Science Framework, reference: https://osf.io/fejpf/ . Registered 9 March 2017.

Time, touch and temperature affect perceived finger position and ownership in the grasp illusion.

KEY POINTS: The brain's internal model of the body and the sense of body ownership are fundamental to interaction with the world. It is thought that temporally congruent, repetitive multisensory stimuli are required to elicit a sense of body ownership. Here we investigate the ability of static cutaneous stimuli - passively grasping an artificial finger - to induce body ownership and alter perceived body position; we also investigate how physical characteristics of grasped objects alter these senses. We show that static cutaneous stimuli can alter perceived body position and induce an illusion of ownership and also that signals of temperature, texture and shape of grasped finger-sized objects influence body ownership. Thus, these aspects of human proprioception can be altered by a single sustained sensory stimulus and by the physical characteristics of held objects. ABSTRACT: Perceived body position and ownership are fundamental to our ability to sense and interact with the world. Previous work indicates that temporally congruent, repetitive multisensory stimuli are needed to alter the sense of body ownership. In the present study 30 subjects passively grasped an artificial rubber finger with their left index and thumb while their right index finger, located 12 cm below, was lightly clamped. Fingers with varied physical characteristics were also passively grasped to determine how these characteristics influenced perceived body position and ownership. Subjects immediately felt their hands to be 5.3 cm [3.4-7.3] (mean [95%CI]) closer, a feeling that remained after 3 min (6.0 cm [4.5-7.5]). By the end of the trial, perceived ownership increased by 1.2 [0.6-1.9] points on a 7-point Likert scale, with the group average moving from 'neither agree or disagree' at the start to 'somewhat agree' at the end. Compared to grasping a control rubber finger, grasping a cold, rough, oddly shaped or rectangular shaped finger-like object reduced perceived ownership. These results provide new insights into the role of cutaneous sensory receptors in defining these aspects of proprioception, and the speed with which these effects occur. Static touch rapidly induces large, sustained changes in perceived body position and prolonged exposure to these cutaneous inputs, alone, can induce a sense of body ownership. Also, certain physical characteristics of grasped objects influence the sense of body ownership.

History-dependence of muscle slack length following contraction and stretch in the human vastus lateralis.

KEY POINTS: In reduced muscle preparations, the slack length and passive stiffness of muscle fibres have been shown to be influenced by previous muscle contraction or stretch. In human muscles, such behaviours have been inferred from measures of muscle force, joint stiffness and reflex magnitudes and latencies. Using ultrasound imaging, we directly observed that isometric contraction of the vastus lateralis muscle at short lengths reduces the slack lengths of the muscle-tendon unit and muscle fascicles. The effect is apparent 60 s after the contraction. These observations imply that muscle contraction at short lengths causes the formation of bonds which reduce the effective length of structures that generate passive tension in muscles. ABSTRACT: In reduced muscle preparations, stretch and muscle contraction change the properties of relaxed muscle fibres. In humans, effects of stretch and contraction on properties of relaxed muscles have been inferred from measurements of time taken to develop force, joint stiffness and reflex latencies. The current study used ultrasound imaging to directly observe the effects of stretch and contraction on muscle-tendon slack length and fascicle slack length of the human vastus lateralis muscle in vivo. The muscle was conditioned by (a) strong isometric contractions at long muscle-tendon lengths, (b) strong isometric contractions at short muscle-tendon lengths, (c) weak isometric contractions at long muscle-tendon lengths and (d) slow stretches. One minute after conditioning, ultrasound images were acquired from the relaxed muscle as it was slowly lengthened through its physiological range. The ultrasound image sequences were used to identify muscle-tendon slack angles and fascicle slack lengths. Contraction at short muscle-tendon lengths caused a mean 13.5 degree (95% CI 11.8-15.0 degree) shift in the muscle-tendon slack angle towards shorter muscle-tendon lengths, and a mean 5 mm (95% CI 2-8 mm) reduction in fascicle slack length, compared to the other conditions. A supplementary experiment showed the effect could be demonstrated if the muscle was conditioned by contraction at short lengths but not if the relaxed muscle was held at short lengths, confirming the role of muscle contraction. These observations imply that muscle contraction at short lengths causes the formation of bonds which reduce the effective length of structures that generate passive tension in muscles.

Human motoneurone excitability is depressed by activation of serotonin 1A receptors with buspirone.

KEY POINTS: In the adult turtle spinal cord, action potential generation in motoneurones is inhibited by spillover of serotonin to extrasynaptic serotonin 1A (5-HT1A ) receptors at the axon initial segment. We explored whether ingestion of the 5-HT1A receptor partial agonist, buspirone, decreases motoneurone excitability in humans. Following ingestion of buspirone, two tests of motoneurone excitability showed decreases. F-wave areas and persistence in an intrinsic muscle of the hand were reduced, as was the area of cervicomedullary motor evoked potentials in biceps brachii. Our findings suggest that activation of 5-HT1A receptors depresses human motoneurone excitability. Such a depression could contribute to decreased motoneurone output during fatiguing exercise if there is high serotonergic drive to the motoneurones. ABSTRACT: Intense serotonergic drive in the turtle spinal cord results in serotonin spillover to the axon initial segment of the motoneurones where it activates serotonin 1A (5-HT1A ) receptors and inhibits generation of action potentials. We examined whether activation of 5-HT1A receptors decreases motoneurone excitability in humans by determining the effects of a 5-HT1A receptor partial agonist, buspirone, on F waves and cervicomedullary motor evoked potentials (CMEPs). In a placebo-controlled double-blind study, 10 participants were tested on two occasions where either placebo or 20 mg of buspirone was administered orally. The ulnar nerve was stimulated supramaximally to evoke F waves in abductor digiti minimi (ADM). CMEPs and the maximal M wave were elicited in biceps brachii by cervicomedullary stimulation and brachial plexus stimulation, respectively. Following buspirone intake, F-wave area and persistence, as well as CMEP area, were significantly decreased. The mean post-pill difference in normalized F-wave areas and persistence between buspirone and placebo days was -27% (-42, -12; 95% confidence interval) and -9% (-16, -2), respectively. The mean post-pill difference in normalized CMEP area between buspirone and placebo days showed greater variation and was -31% (-60, -2). In conclusion, buspirone reduces motoneurone excitability in humans probably via activation of 5-HT1A receptors at the axon initial segment. This has implications for motor output during high drive to the motoneurones when serotonin may spill over to these inhibitory receptors and consequently inhibit motoneurone output. Such a mechanism could potentially contribute to fatigue with exercise.

Time course of human motoneuron recovery after sustained low-level voluntary activity.

Motoneurons often fire repetitively and for long periods. In sustained voluntary contractions the excitability of motoneurons declines. We provide the first detailed description of the time course of human motoneuron recovery after sustained activity at a constant discharge rate. We recorded the discharge of single motor units (MUs, n = 30) with intramuscular wire electrodes inserted in triceps brachii during weak isometric contractions. Subjects (n = 15) discharged single MUs at a constant frequency (∼10 Hz) with visual feedback for prolonged durations (3-7 min) until rectified surface electromyogram (sEMG) of triceps brachii increased by ∼100%. After a rest of 1-2, 15, 30, 60, 120, or 240 s, subjects briefly resumed the contraction with the target MU at the same discharge rate. Each MU was tested with three to four rest periods. The magnitude of sEMG was increased when contractions were resumed, and the target motoneuron discharged at the test frequency following rest intervals of 2-60 s (P = 0.001-0.038). The increased sEMG indicates that greater excitatory drive was needed to discharge the motoneuron at the test rate. The increase in EMG recovered exponentially with a time constant of 28 s but did not return to baseline even after a rest period of ∼240 s. Thus the decline in motoneuron excitability from a weak contraction takes several minutes to recover fully.

Vestibular contribution to balance control in the medial gastrocnemius and soleus.

The soleus (Sol) and medial gastrocnemius (mGas) muscles have different patterns of activity during standing balance and may have distinct functional roles. Using surface electromyography we previously observed larger responses to galvanic vestibular stimulation (GVS) in the mGas compared with the Sol muscle. However, it is unclear whether this difference is an artifact that reflects limitations associated with surface electromyography recordings or whether a compensatory balance response to a vestibular error signal activates the mGas to a greater extent than the Sol. In the present study, we compared the effect of GVS on the discharge behavior of 9 Sol and 21 mGas motor units from freely standing subjects. In both Sol and mGas motor units, vestibular stimulation induced biphasic responses in measures of discharge timing [11 ± 5.0 (mGas) and 5.6 ± 3.8 (Sol) counts relative to the sham (mean ± SD)], and frequency [0.86 ± 0.6 Hz (mGas), 0.34 ± 0.2 Hz (Sol) change relative to the sham]. Peak-to-trough response amplitudes were significantly larger in the mGas (62% in the probability-based measure and 160% in the frequency-based measure) compared with the Sol (multiple P < 0.05). Our results provide direct evidence that vestibular signals have a larger influence on the discharge activity of motor units in the mGas compared with the Sol. More tentatively, these results indicate the mGas plays a greater role in vestibular-driven balance corrections during standing balance.

Motor units in the human medial gastrocnemius muscle are not spatially localized or functionally grouped.

KEY POINTS: Human medial gastrocnemius (MG) motor units (MUs) are thought to occupy small muscle territories or regions, with low-threshold units preferentially located distally. We used intramuscular recordings to measure the territory of muscle fibres from MG MUs and determine whether these MUs are grouped by recruitment threshold or joint action (ankle plantar flexion and knee flexion). The territory of MUs from the MG muscle varied from somewhat localized to highly distributed, with approximately half the MUs spanning at least half the length and width of the muscle. There was also no evidence of regional muscle activity based on MU recruitment thresholds or joint action. The CNS does not have the means to selectively activate regions of the MG muscle based on task requirements. ABSTRACT: Human medial gastrocnemius (MG) motor units (MUs) are thought to occupy small muscle territories, with low-threshold units preferentially located distally. In this study, subjects (n = 8) performed ramped and sustained isometric contractions (ankle plantar flexion and knee flexion; range: ∼1-40% maximal voluntary contraction) and we measured MU territory size with spike-triggered averages from fine-wire electrodes inserted along the length (seven electrodes) or across the width (five electrodes) of the MG muscle. Of 69 MUs identified along the length of the muscle, 32 spanned at least half the muscle length (≥ 6.9 cm), 11 of which spanned all recording sites (13.6-17.9 cm). Distal fibres had smaller pennation angles (P < 0.05), which were accompanied by larger territories in MUs with fibres located distally (P < 0.05). There was no distal-to-proximal pattern of muscle activation in ramp contraction (P = 0.93). Of 36 MUs identified across the width of the muscle, 24 spanned at least half the muscle width (≥ 4.0 cm), 13 of which spanned all recording sites (8.0-10.8 cm). MUs were not localized (length or width) based on recruitment threshold or contraction type, nor was there a relationship between MU territory size and recruitment threshold (Spearman's rho = -0.20 and 0.13, P > 0.18). MUs in the human MG have larger territories than previously reported and are not localized based on recruitment threshold or joint action. This indicates that the CNS does not have the means to selectively activate regions of the MG muscle based on task requirements.

Is this my finger? Proprioceptive illusions of body ownership and representation.

Body 'ownership' defines which things belong to us and can be manipulated by signals from cutaneous or muscle receptors. Whether signals from muscle proprioceptors on their own influence perceived ownership is unknown. We used finger-joint movement to induce illusory ownership of an artificial finger without vision. We coupled the subject's index finger to an artificial finger 12 cm above it. The experimenter held the subject's other index finger and thumb on the artificial finger and passively moved them congruently or incongruently for 3 min with the index finger and the grasping index finger and thumb intact or anaesthetised. When intact, congruent movement (19 subjects) reduced perceived vertical distance between index fingers to 1.0 (0.0, 2.0) cm [median (IQR)] from 3.0 (3.0, 4.0) cm with incongruent movement (P < 0.01). Simply grasping the artificial finger reduced perceived spacing between the grasping and test index fingers from 6.0 (5.0, 9.0) cm to 3.0 (3.0, 6.0) cm (P < 0.01), a new grasp illusion. Digital anaesthesia eliminated this grasp effect, after which congruent movement still reduced the perceived spacing between the index fingers to 1.0 (0.0, 2.75) cm compared to 4.0 (3.25, 6.0) cm with incongruent movement (P < 0.001). Subjects more strongly agreed that they were holding their own finger after congruent but not incongruent movement (P < 0.01). We propose that the brain generates possible scenarios and tests them against available sensory information. This process can function without vision or motor commands, and with only one channel of somatic information.

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.