A Principle of Neuromechanical Matching for Motor Unit Recruitment in Human Movement

What determines which motor units are active in a motor task? In the respiratory muscles, motor units are recruited according to their mechanical advantages. We describe a principle of motor unit recruitment by neuromechanical matching due to mechanisms in the spinal cord that sculpt descending drive to motoneurons. This principle may be applicable to movements in nonrespiratory muscles

Critical considerations of the contribution of the corticomotoneuronal pathway to central fatigue

Neural drive originating in higher brain areas reaches exercising limb muscles through the corticospinal-motoneuronal pathway, which links the motor cortex and spinal motoneurones. The properties of this pathway have frequently been observed to change during fatiguing exercise in ways that could influence the development of central fatigue (i.e. the progressive reduction in voluntary muscle activation). However, based on differences in motor cortical and motoneuronal excitability between exercise modalities (e.g. single-joint vs. locomotor exercise), there is no characteristic response that allows for a categorical conclusion about the effect of these changes on functional impairments and performance limitations. Despite the lack of uniformity in findings during fatigue, there is strong evidence for marked ‘inhibition’ of motoneurones as a direct result of voluntary drive. Endogenous forms of neuromodulation, such as via serotonin released from neurones, can directly affect motoneuronal output and central fatigue. Exogenous forms of neuromodulation, such as brain stimulation, may achieve a similar effect, although the evidence is weak. Non-invasive transcranial direct current stimulation can cause transient or long-lasting changes in cortical excitability; however, variable results across studies cast doubt on its claimed capacity to enhance performance. Furthermore, with these studies, it is difficult to establish a cause-and-effect relationship between brain responsiveness and exercise performance. This review briefly summarizes changes in the corticomotoneuronal pathway during various types of exercise, and considers the relevance of these changes for the development of central fatigue, as well as the potential of non-invasive brain stimulation to enhance motor cortical excitability, motoneuronal output and, ultimately, exercise performance. (Figure presented.)

Respiratory muscle activity in voluntary breathing tracking tasks: Implications for the assessment of respiratory motor control

How the involuntary (bulbospinal) and voluntary (corticospinal) pathways interact in respiratory muscle control is not established. To determine the role of excitatory corticobulbar pathways in humans, studies typically compare electromyographic activity (EMG) or evoked responses in respiratory muscles during hypercapnic and voluntary tasks. Although ventilation is matched between tasks by having participants track signals of ventilation, these tasks may not result in matched respiratory muscle activity. The aim of this study was to describe respiratory muscle activity and ribcage and abdominal excursions during two different voluntary conditions, compared to hypercapnic hyperventilation. Ventilation was matched in the voluntary conditions via (i) a simple target of lung volume (‘volume tracking’) or (ii) targets of both ribcage and abdominal excursions, adjusted to end-expiratory lung volume in hypercapnic hyperventilation (‘bands tracking’). Compared to hypercapnic hyperventilation, respiratory parameters such as tidal volume were similar, but the ratio of ribcage to abdominal excursion was higher for both voluntary tasks. Inspiratory scalene and parasternal intercostal muscle activity was higher in volume tracking, but diaphragm and abdominal muscle activity showed little to no change. There were no differences in muscle activity in bands tracking for any muscle, compared to hypercapnic hyperventilation. An elevated ratio of ribcage to abdominal excursion in the bands tracking task indicates that participants could not accurately match the targets in this condition. Inspiratory muscle activity is altered in some muscles in some voluntary tasks, compared to hypercapnia. Therefore, differences in muscle activity should be considered in interpretation of studies that use these protocols to investigate respiratory muscle control

Use of a physiological profile to document upper limb motor impairment in ageing and in neurological conditions

Profiling performance in the physiological domains underpinning upper limb function (such as strength, sensation, coordination) provides insight into an individual's specific impairments. This compliments the traditional medical ‘diagnosis’ model that is currently used in contemporary medicine. From an initial battery of 13 tests in which data were collected across the adult lifespan (n = 367, 20–95 years) and in those with neurological conditions (specifically, multiple sclerosis (n = 40), Parkinson's disease (n = 34), and stroke (n = 50)), six tests were selected to comprise a core upper limb physiological profile assessment (PPA). This comprised measures of handgrip strength, simple reaction time, finger dexterity, tactile sensation, bimanual coordination, and a functional task. Individual performance in each of these tests can be compared to a reference population score (devised from our database of healthy individuals aged under 60 years), informing the researcher or clinician how to best direct an intervention or treatment for the individual based on their specific impairment(s). Lastly, a composite score calculated from the average performance across the six tests provides a broad overview of an individual's overall upper limb function. Collectively, the upper limb PPA highlights specific impairments that are prevalent within distinct pathologies and reveals the magnitude of upper limb motor impairment specific to each condition. (Figure presented.)

Discharge properties of human diaphragm motor units with ageing

Key points: Ageing is associated with changes in the respiratory system including in the lungs, rib cage and muscles. Neural drive to the diaphragm, the principal inspiratory muscle, has been reported to increase during quiet breathing with ageing. We demonstrated that low-threshold motor units of the human diaphragm recruited during quiet breathing have similar discharge frequencies across age groups and shorter discharge times in older age. With ageing, motor unit action potential area increased. We propose that there are minimal functionally significant changes in the discharge properties of diaphragm motor units with ageing despite remodelling of the motor unit in the periphery. Abstract: There are changes in the skeletal, pulmonary and respiratory neuromuscular systems with healthy ageing. During eupnoea, one study has shown relatively higher crural diaphragm electromyographic activity (EMG) in healthy older adults (>51 years) than in younger adults, but these measures may be affected by the normalisation process used. A more direct method to assess neural drive involves the measurement of discharge properties of motor units. Here, to assess age-related changes in neural drive to the diaphragm during eupnoea, EMG was recorded from the costal diaphragm using a monopolar needle electrode in participants from three age groups (n ≥ 7 each): older (65–80 years); middle-aged (43–55 years) and young (23–26 years). In each group, 154, 174 and 110 single motor units were discriminated, respectively. A mixed-effects linear model showed no significant differences between age groups for onset (group mean range 9.5–10.2 Hz), peak (14.1–15.0 Hz) or offset (7.8–8.5 Hz) discharge frequencies during eupnoea. The motor unit recruitment was delayed in the older group (by ∼15% of inspiratory time; p = 0.02 cf. middle-aged group) and had an earlier offset time (by ∼15% of inspiratory time; p = 0.04 cf. young group). However, the onset of multiunit activity was similar across groups, consistent with no global increase in neural drive to the diaphragm with ageing. The area of diaphragm motor unit potentials was ∼40% larger in the middle-aged and older groups (P < 0.02), which indicates axonal sprouting and re-innervation of muscle fibres associated with ageing, even in middle-aged participants

Reflex response to airway occlusion in human inspiratory muscles when recruited for breathing and posture

Briefly occluding the airway during inspiration produces a short-latency reflex inhibition in human inspiratory muscles. This occlusion reflex seems specific to respiratory muscles; however, it is not known whether the reflex inhibition has a uniform effect across a motoneuron pool when a muscle is recruited concurrently for breathing and posture. In this study, participants were seated and breathed through a mouthpiece that occluded inspiratory airflow for 250 ms at a volume threshold of 0.2 liters. The reflex response was measured in the scalene and sternocleidomastoid muscles during 1) a control condition with the head supported in space and the muscles recruited for breathing only, 2) a postural condition with the head unsupported and the neck flexors recruited for both breathing and to maintain head posture, and 3) a large-breath condition with the head supported and the volume threshold raised to between 0.8 and 1.0 liters to increase inspiratory muscle activity. When normalized to its preocclusion mean, the reflex response in the scalene muscles was not significantly different between the large-breath and control conditions, whereas concomitant recruitment of these muscles for posture control reduced the reflex response by half compared with the control condition. A reflex response occurred in sternocleidomastoid when it contracted phasically as an accessory muscle for inspiration during the large-breath condition. These results indicate that the occlusion reflex does not produce a uniform effect across the motoneuron pool and that afferent inputs for this reflex most likely act via intersegmental networks of premotoneurons rather than at a motoneuronal level

Changes in pharyngeal collapsibility and genioglossus reflex responses to negative pressure during the respiratory cycle in obstructive sleep apnoea

Key points: Impaired pharyngeal anatomy and increased airway collapsibility is a major cause of obstructive sleep apnoea (OSA) and a mediator of its severity. Upper airway reflexes to changes in airway pressure provide important protection against airway closure. This study shows increased pharyngeal collapsibility and attenuated genioglossus reflex responses during expiration in people with OSA. Abstract: Upper airway collapse contributes to obstructive sleep apnoea (OSA) pathogenesis. Pharyngeal dilator muscle activity varies throughout the respiratory cycle and may contribute to dynamic changes in pharyngeal collapsibility. However, whether upper airway collapsibility and reflex responses to changes in airway pressure vary throughout the respiratory cycle in OSA is unclear. Thus, this study quantified differences in upper airway collapsibility and genioglossus electromyographic (EMG) activity and reflex responses during different phases of the respiratory cycle. Twelve middle-aged people with OSA (2 female) were fitted with standard polysomnography equipment: a nasal mask, pneumotachograph, two fine-wire intramuscular electrodes into the genioglossus, and a pressure catheter positioned at the epiglottis and a second at the choanae (the collapsible portion of the upper airway). At least 20 brief (∼250 ms) pressure pulses (∼−11 cmH2O at the mask) were delivered every 2–10 breaths during four conditions: (1) early inspiration, (2) mid-inspiration, (3) early expiration, and (4) mid-expiration. Mean baseline genioglossus EMG activity 100 ms prior to pulse delivery and genioglossus reflex responses were quantified for each condition. The upper airway collapsibility index (UACI), quantified as 100 × (nadir choanal − epiglottic pressure)/nadir choanal pressure during negative pressure pulses, varied throughout the respiratory cycle (early inspiration = 43 ± 25%, mid-inspiration = 29 ± 19%, early expiration = 83 ± 19% and mid-expiration = 95 ± 11% (mean ± SD) P < 0.01). Genioglossus EMG activity was lower during expiration (e.g. mid-expiration vs. mid-inspiration = 76 ± 23 vs. 127 ± 41% of early-inspiration, P < 0.001). Similarly, genioglossus reflex excitation was delayed (39 ± 11 vs. 23 ± 7 ms, P < 0.001) and reflex excitation amplitude attenuated during mid-expiration versus early inspiration (209 ± 36 vs. 286 ± 80%, P = 0.009). These findings may provide insight into the physiological mechanisms of pharyngeal collapse in OSA

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 29yr, range 19-64yr) 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 logetransformed 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

The detection and sensory perception of inspiratory resistive loads in people with chronic tetraplegia

This study investigated sensations of breathing following tetraplegia. Fifteen people with chronic tetraplegia and fifteen healthy able-bodied controls matched for age, sex, height, and weight participated. Sensations of breathing were quantified by determining the threshold for detecting an added resistance during inspiration. In a separate task, the perceived magnitudes of six suprathreshold resistive loads were determined with a modified Borg scale. The detection threshold of 0.34 cmH2O/L/s [standard deviation (SD) 0.14] in the tetraplegia group was higher than the 0.23 cmH2O/L/s (SD 0.10) threshold for able-bodied controls (P = 0.004). Both participant groups perceived larger loads to be more effortful, with the Borg effort rating increasing linearly with the peak inspiratory pressure generated at each load. The relationship between Borg effort rating and peak inspiratory pressure was steeper in participants with tetraplegia than in able-bodied controls (P = 0.001), but there was no difference when pressure was divided by maximal inspiratory pressure (P = 0.95). Despite a higher detection threshold, the findings suggest that the perceived magnitude of a suprathreshold inspiratory load is not impaired in chronic tetraplegia and that load magnitude perception is related to the maximal, and not absolute, inspiratory muscle force. NEW & NOTEWORTHY Sensations of breathing are thought to be impaired following chronic tetraplegia. The detection threshold for an added resistive load during inspiration was higher in people with tetraplegia than in healthy able-bodied participants. However, for inspiratory loads above the detection threshold, the perceived magnitude of a resistive load as a function of the peak inspiratory pressure was greater in tetraplegia. Load magnitude perception was comparable between participant groups when peak pressure was divided by maximal inspiratory pressure