Locomotor-respiratory Coupling Research Articles

Dynamics of the locomotor-respiratory coupling at different frequencies.

The locomotor-respiratory coupling (LRC) is a universal phenomenon reported for various forms of rhythmic exercise. In this study, we investigated the effect of movement and respiratory frequencies on LRC. Participants were instructed to cycle or breath in synchrony with a periodic auditory stimulation at preferred and non-preferred frequencies. LRC stability was assessed by frequency and phase coupling indexes using the theory of nonlinear coupled oscillators through the sine circle map model, and the Farey tree. Results showed a stabilizing effect of sound on LRC for all frequencies and for the two systems paced. The sound-induced effect was more prominent when the rhythm of the stimulation corresponded to the preferred frequencies. The adoption of cycling or respiratory frequencies far off preferential ones led to a loss of stability in LRC. Contrary to previous findings, our results suggest that LRC is not unidirectional-from locomotion onto respiration-but bidirectional between the two systems. They also suggest that auditory information plays an important role in the modulation of LRC.

Sound-induced stabilization of breathing and moving.

In humans and other animals, the locomotor and respiratory systems are coupled together through mechanical, neurophysiological, and informational interactions. At a macroscopic observer-environment level, these three types of interactions produce locomotor-respiratory coupling (LRC), whose dynamics are evaluated in this paper. A formal analysis of LRC is presented, exploiting tools from synchronization theories and nonlinear dynamics. The results of two recent studies, in which participants were instructed to cycle or exhale at a natural frequency or in synchrony with an external rhythmic sound, are discussed. The metronome was either absent or present (study 1) and close to or far from the natural frequency of the cycling and breathing systems (study 2). The results evidenced a stabilization of cycling, breathing, and LRC when sound was present compared to when it was absent. A decrease in oxygen consumption was also observed, accompanying the increase in sound-induced LRC stabilization. These results obtained with a simple rhythmic metronome beat have consequences for exercising while listening to music; the consequences are further explored here.

Ventilation and Locomotion in Humans: Mechanisms, Implications, and Perturbations to the Coupling of These Two Rhythms

To best sustain endurance activity, two systems must be effectively coordinated: ventilation and locomotion. Evidence has long suggested that these two mammalian rhythms are linked, yet determinants and implications of locomotor–respiratory coupling (LRC) continue to be investigated. Two general areas explaining the potential mechanisms underlying LRC are (1) neural interactions between central and peripheral controllers of locomotion and respiration, and (2) mechanical interactions between locomotor dynamics and respiratory mechanics. Additional suggested explanations for/consequence of the existence of LRC in mammals include an improved energetic cost of locomotion and a reduced sensation of breathlessness. As such, any perturbation to LRC, via alterations in breathing or kinematic patterns, could have negative performance implications to both athlete and patient populations.

The Prevalence of Abnormal Breathing Patterns During Exercise and Associations with Dynamic Upper Respiratory Tract Obstructions

IntroductionDuring canter and gallop most horses maintain a locomotor‐respiratory coupling (LRC) ratio of one breath every stride. Occasional big respiratory cycles (BRC) are described when a single breath is taken over two strides. This is considered a transient but normal feature of ventilation. However, a change to a more persistent 2:1 LRC ratio is considered to be ‘abnormal’. It has been suggested that upper respiratory tract (URT) disorders could result in a horse adopting an abnormal breathing pattern, although current information is limited.MethodsSimultaneous audio and URT endoscopy recordings from 365 horses referred for treadmill evaluation were analysed. Breathing pattern at canter and gallop was categorised at each one minute speed step: Normal: 1:1 with <2 BRC Normal: 1:1 with 2 BRC Abnormal: breathing pattern disrupted by repetitive and frequent BRC's Abnormal: alternating between periods of 2:1 and 1:1 Abnormal: 2:1 throughout Data analysis was performed using PASW 18.0. A chi‐square test was used to identify associations between breathing pattern and presence of URT obstructions.ResultsDuring the entire exercise test 30% of horses had an abnormal breathing pattern at canter or gallop, of which 23% had a 2:1 breathing pattern. The prevalence of abnormal breathing patterns was higher at low canter speeds than during strenuous exercise. There was a significant association between breathing pattern during strenuous exercise and the presence of an URT obstruction (P = 0.008).ConclusionsLoss of a 1:1 LRC ratio during canter and gallop is more common than previously realised and may be associated with dynamic URT obstructions.Ethical Animal ResearchRetrospective analysis of case records. Sources of funding: none declared. Competing interests: none.

Impact Loading and Locomotor-Respiratory Coordination Significantly Influence Breathing Dynamics in Running Humans

Locomotor-respiratory coupling (LRC), phase-locking between breathing and stepping rhythms, occurs in many vertebrates. When quadrupedal mammals gallop, 1∶1 stride per breath coupling is necessitated by pronounced mechanical interactions between locomotion and ventilation. Humans show more flexibility in breathing patterns during locomotion, using LRC ratios of 2∶1, 2.5∶1, 3∶1, or 4∶1 and sometimes no coupling. Previous studies provide conflicting evidence on the mechanical significance of LRC in running humans. Some studies suggest LRC improves breathing efficiency, but others suggest LRC is mechanically insignificant because ‘step-driven flows’ (ventilatory flows attributable to step-induced forces) contribute a negligible fraction of tidal volume. Yet, although step-driven flows are brief, they cause large fluctuations in ventilatory flow. Here we test the hypothesis that running humans use LRC to minimize antagonistic effects of step-driven flows on breathing. We measured locomotor-ventilatory dynamics in 14 subjects running at a self-selected speed (2.6±0.1 ms−1) and compared breathing dynamics in their naturally ‘preferred’ and ‘avoided’ entrainment patterns. Step-driven flows occurred at 1-2X step frequency with peak magnitudes of 0.97±0.45 Ls−1 (mean ±S.D). Step-driven flows varied depending on ventilatory state (high versus low lung volume), suggesting state-dependent changes in compliance and damping of thoraco-abdominal tissues. Subjects naturally preferred LRC patterns that minimized antagonistic interactions and aligned ventilatory transitions with assistive phases of the step. Ventilatory transitions initiated in ‘preferred’ phases within the step cycle occurred 2x faster than those in ‘avoided’ phases. We hypothesize that humans coordinate breathing and locomotion to minimize antagonistic loading of respiratory muscles, reduce work of breathing and minimize rate of fatigue. Future work could address the potential consequences of locomotor-ventilatory interactions for elite endurance athletes and individuals who are overweight or obese, populations in which respiratory muscle fatigue can be limiting.

Sound Stabilizes Locomotor-Respiratory Coupling and Reduces Energy Cost

A natural synchronization between locomotor and respiratory systems is known to exist for various species and various forms of locomotion. This Locomotor-Respiratory Coupling (LRC) is fundamental for the energy transfer between the two subsystems during long duration exercise and originates from mechanical and neurological interactions. Different methodologies have been used to compute LRC, giving rise to various and often diverging results in terms of synchronization, (de-)stabilization via information, and associated energy cost. In this article, the theory of nonlinear-coupled oscillators was adopted to characterize LRC, through the model of the sine circle map, and tested it in the context of cycling. Our specific focus was the sound-induced stabilization of LRC and its associated change in energy consumption. In our experimental study, participants were instructed during a cycling exercise to synchronize either their respiration or their pedaling rate with an external auditory stimulus whose rhythm corresponded to their individual preferential breathing or cycling frequencies. Results showed a significant reduction in energy expenditure with auditory stimulation, accompanied by a stabilization of LRC. The sound-induced effect was asymmetrical, with a better stabilizing influence of the metronome on the locomotor system than on the respiratory system. A modification of the respiratory frequency was indeed observed when participants cycled in synchrony with the tone, leading to a transition toward more stable frequency ratios as predicted by the sine circle map. In addition to the classical mechanical and neurological origins of LRC, here we demonstrated using the sine circle map model that information plays an important modulatory role of the synchronization, and has global energetic consequences.

Spinal and Pontine Relay Pathways Mediating Respiratory Rhythm Entrainment by Limb Proprioceptive Inputs in the Neonatal Rat

The coordination of locomotion and respiration is widespread among mammals, although the underlying neural mechanisms are still only partially understood. It was previously found in neonatal rat that cyclic electrical stimulation of spinal cervical and lumbar dorsal roots (DRs) can fully entrain (1:1 coupling) spontaneous respiratory activity expressed by the isolated brainstem/spinal cord. Here, we used a variety of preparations to determine the type of spinal sensory inputs responsible for this respiratory rhythm entrainment, and to establish the extent to which limb movement-activated feedback influences the medullary respiratory networks via direct or relayed ascending pathways. During in vivo overground locomotion, respiratory rhythm slowed and became coupled 1:1 with locomotion. In hindlimb-attached semi-isolated preparations, passive flexion-extension movements applied to a single hindlimb led to entrainment of fictive respiratory rhythmicity recorded in phrenic motoneurons, indicating that the recruitment of limb proprioceptive afferents could participate in the locomotor-respiratory coupling. Furthermore, in correspondence with the regionalization of spinal locomotor rhythm-generating circuitry, the stimulation of DRs at different segmental levels in isolated preparations revealed that cervical and lumbosacral proprioceptive inputs are more effective in this entraining influence than thoracic afferent pathways. Finally, blocking spinal synaptic transmission and using a combination of electrophysiology, calcium imaging and specific brainstem lesioning indicated that the ascending entraining signals from the cervical or lumbar limb afferents are transmitted across first-order synapses, probably monosynaptic, in the spinal cord. They are then conveyed to the brainstem respiratory centers via a brainstem pontine relay located in the parabrachial/Kölliker-Fuse nuclear complex.

Locomotor-respiratory coupling patterns and oxygen consumption during walking above and below preferred stride frequency

Locomotor respiratory coupling patterns in humans have been assessed on the basis of the interaction between different physiological and motor subsystems; these interactions have implications for movement economy. A complex and dynamical systems framework may provide more insight than entrainment into the variability and adaptability of these rhythms and their coupling. The purpose of this study was to investigate the relationship between steady state locomotor-respiratory coordination dynamics and oxygen consumption [Formula: see text] of the movement by varying walking stride frequency from preferred. Twelve male participants walked on a treadmill at a self-selected speed. Stride frequency was varied from -20 to +20% of preferred stride frequency (PSF) while respiratory airflow, gas exchange variables, and stride kinematics were recorded. Discrete relative phase and return map techniques were used to evaluate the strength, stability, and variability of both frequency and phase couplings. Analysis of [Formula: see text] during steady-state walking showed a U-shaped response (P = 0.002) with a minimum at PSF and PSF - 10%. Locomotor-respiratory frequency coupling strength was not greater (P = 0.375) at PSF than any other stride frequency condition. The dominant coupling across all conditions was 2:1 with greater occurrences at the lower stride frequencies. Variability in coupling was the greatest during PSF, indicating an exploration of coupling strategies to search for the coupling frequency strategy with the least oxygen consumption. Contrary to the belief that increased strength of frequency coupling would decrease oxygen consumption; these results conclude that it is the increased variability of frequency coupling that results in lower oxygen consumption.

Stabilizing the Locomotor-Respiratory Coupling Using a Metronome to Save Energy

The Locomotor-Respiratory Coupling (LRC) is often evidenced by phase- or frequency-locking patterns. The model of the sine circle map is used here to characterize LRC. Several studies have suggested that a sound emitted by an external metronome can stabilize the LRC. Participants in our task were asked during a cycling exercise to synchronize either their respiration or their pedaling rate with an external auditory stimulus corresponding to their preferred respiratory and pedaling frequencies respectively. Our results showed a significant reduction in energy expenditure when participants breathed in sync with the auditory stimulation, but not accompanied by a change in the stabilization of LRC. A large within- as well as between-participants LRC variability, together with the spontaneous adoption of the most stable pace, contributes to explain this result.

A model of locomotor-respiratory coupling in quadrupeds

Locomotion and respiration are not independent phenomena in running mammals because locomotion and respiration both rely on cyclic movements of the ribs, sternum, and associated musculature. Thus, constraints are imposed on locomotor and respiratory function by virtue of their linkage. Specifically, locomotion imposes mechanical constraints on breathing that require the respiratory cycle to be synchronized with gait. Thus, many mammals, including humans, synchronize respiration with the movement of the limbs during locomotion. For example, quadrupeds synchronize locomotor and respiratory cycles at a 1:1 ratio (stride/breath) over a wide range of speeds. Interestingly, quadrupeds maintain an almost constant stride frequency (and therefore respiratory frequency) at different speeds. To increase speed, quadrupeds lengthen their stride. Accordingly, to increase minute ventilation, quadrupeds must increase tidal volume since respiratory rate is coupled with stride frequency. We developed a simple, inexpensive, and easy to build model to demonstrate this concept. A model was chosen because models significantly enhance student understanding. Students are drawn into discussion by the power of learning that is associated with manipulating and thinking about objects. Building and using this model strengthen the concept that locomotor-respiratory coupling provides a basis for the appropriate matching of lung ventilation to running speed and metabolic power.

Locomotor–respiratory synchronization after body weight supported treadmill training in incomplete tetraplegia: a case report

Case report. To report on the respiratory and kinematic changes associated with body weight supported treadmill training (BWSTT) in an individual with an incomplete cervical spinal cord injury (SCI). Human Locomotion Research Laboratory, Vancouver, BC, Canada. A 38-year-old man with an incomplete SCI at the C1/C2 level, graded on the American Spinal Injury Association Impairment Scale (AIS) as AIS D, participated in this case study. He performed three BWSTT sessions (45 min) per week for 12 weeks. Before and after training, respiration was measured at the mouth using a pneumotachograph while treadmill walking at 1.6 km h(-1). Completion of 12 weeks of BWSTT resulted in a lowering of minute ventilation during treadmill exercise. A coupling of locomotion and respiration was observed during treadmill walking after the training program, whereas the relationship was initially absent. This case illustrates that BWSTT can result in a reduced ventilatory demand and promote locomotor-respiratory coupling during walking in an individual with an incomplete cervical SCI.

The gross morphology and histochemistry of respiratory muscles in bottlenose dolphins, Tursiops truncatus

Most mammals possess stamina because their locomotor and respiratory (i.e., ventilatory) systems are mechanically coupled. These systems are decoupled, however, in bottlenose dolphins (Tursiops truncatus) as they swim on a breath hold. Locomotion and ventilation are coupled only during their brief surfacing event, when they respire explosively (up to 90% of total lung volume in approximately 0.3 s) (Ridgway et al. 1969 Science 166:1651-1654). The predominantly slow-twitch fiber profile of their diaphragm (Dearolf 2003 J Morphol 256:79-88) suggests that this muscle does not likely power their rapid ventilatory event. Based on Bramble's (1989 Amer Zool 29:171-186) biomechanical model of locomotor-respiratory coupling in galloping mammals, it was hypothesized that locomotor muscles function to power ventilation in bottlenose dolphins. It was further hypothesized that these muscles would be composed predominantly of fast-twitch fibers to facilitate the bottlenose dolphin's rapid ventilation. The gross morphology of craniocervical (scalenus, sternocephalicus, sternohyoid), thoracic (intercostals, transverse thoracis), and lumbopelvic (hypaxialis, rectus abdominis, abdominal obliques) muscles (n = 7) and the fiber-type profiles (n = 6) of selected muscles (scalenus, sternocephalicus, sternohyoid, rectus abdominis) of bottlenose dolphins were investigated. Physical manipulations of excised thoracic units were carried out to investigate potential actions of these muscles. Results suggest that the craniocervical muscles act to draw the sternum and associated ribs craniodorsally, which flares the ribs laterally, and increases the thoracic cavity volume required for inspiration. The lumbopelvic muscles act to draw the sternum and caudal ribs caudally, which decreases the volumes of the thoracic and abdominal cavities required for expiration. All muscles investigated were composed predominantly of fast-twitch fibers (range 61-88% by area) and appear histochemically poised for rapid contraction. These combined results suggest that dolphins utilize muscles, similar to those used by galloping mammals, to power their explosive ventilation.

An Improved Model for Simulating Obstructive Lung Disease

Pulmonary function testing with the use of spirometers is useful for enhancing students' understanding of normal lung volumes, capacities, and flow rates. Furthermore, the spirogram can be an excellent tool for understanding how lung diseases alter pulmonary function parameters. For example,

Intercostal and Abdominal Respiratory Motoneurons in the Neonatal Rat Spinal Cord: Spatiotemporal Organization and Responses to Limb Afferent Stimulation

Respiration requires the coordinated rhythmic contractions of diverse muscles to produce ventilatory movements adapted to organismal requirements. During fast locomotion, locomotory and respiratory movements are coordinated to reduce mechanical conflict between these functions. Using semi-isolated and isolated in vitro brain stem-spinal cord preparations from neonatal rats, we have characterized for the first time the respiratory patterns of all spinal intercostal and abdominal motoneurons and explored their functional relationship with limb sensory inputs. Neuroanatomical and electrophysiological procedures were initially used to locate intercostal and abdominal motoneurons in the cord. Intercostal motoneuron somata are distributed rostrocaudally from C(7)-T(13) segments. Abdominal motoneuron somata lie between T(8) and L(2). In accordance with their soma distributions, inspiratory intercostal motoneurons are recruited in a rostrocaudal sequence during each respiratory cycle. Abdominal motoneurons express expiratory-related discharge that alternates with inspiration. Lesioning experiments confirmed the pontine origin of this expiratory activity, which was abolished by a brain stem transection at the rostral boundary of the VII nucleus, a critical area for respiratory rhythmogenesis. Entrainment of fictive respiratory rhythmicity in intercostal and abdominal motoneurons was elicited by periodic low-threshold dorsal root stimulation at lumbar (L(2)) or cervical (C(7)) levels. These effects are mediated by direct ascending fibers to the respiratory centers and a combination of long-projection and polysynaptic descending pathways. Therefore the isolated brain stem-spinal cord in vitro generates a complex pattern of respiratory activity in which alternating inspiratory and expiratory discharge occurs in functionally identified spinal motoneuron pools that are in turn targeted by both forelimb and hindlimb somatic afferents to promote locomotor-respiratory coupling.

Neurokinin-1 Receptors Modulate the Excitability of Expiratory Neurons in the Ventral Respiratory Group

We studied the role of neurokinin-1 receptors (NK1-R) on the excitability of expiratory (E) neurons (tonic discharge, E(TONIC); augmenting, E(AUG); decrementing, E(DEC)) throughout the ventral respiratory group, including Bötzinger Complex (BötC) using extracellular single-unit recording combined with pressurized picoejection in decerebrate, arterially perfused juvenile rats. Responses evoked by picoejection of the NK1-R agonist, [Sar9-Met(O2)11]-substance P (SSP) were determined before and after the selective NK1-R antagonist, CP99,994. SSP excited 20 of 35 expiratory neurons by increasing the number of action potentials per burst (+33.7 +/- 6.5% of control), burst duration (+20.6 +/- 7.9% of control), and peak firing frequency (+16.2 +/- 4.8% of control; means +/- SE). Pretreatment with CP99,994 completely blocked SSP-evoked excitation in a subset of neurons tested, supporting the notion that SSP excitation was mediated through NK1-R activation. Because we had previously shown that E(AUG) neurons were crucial to locomotor-respiratory coupling (LRC), we reasoned that blockade of NK1-R would alter LRC by preventing somatic-evoked excitation of E(AUG) neurons. Blockade of NK1-Rs by CP99,994 in the BötC severely disrupted LRC and prevented somatic-evoked excitation of E(AUG) neurons. These findings demonstrate that LRC is dependent on endogenous SP release acting via NK1-Rs on E(AUG) neurons of the BötC. Taken together with our earlier finding that inspiratory off-switching by the Hering-Breuer Reflex requires endogenous activation of NK1-Rs through activation of NK1-Rs on E(DEC) neurons, we suggest that endogenous release of substance P in the BötC provides a reflex pathway-dependent mechanism to selectively modulate respiratory rhythm.

Factors influencing variation in locomotor‐respiratory coupling in Standardbred Trotters in the field

A close relationship between limb and respiratory rhythms has been shown in clinically healthy galloping horses due to mechanical constraints in the thoracic region. This synchronisation leads to a 1/1 ratio between stride frequency (SF) and respiratory frequency (RF) during galloping. Very little is known about locomotor-respiratory coupling (LRC) during fast trot. To investigate stride and respiratory rates during a standardised exercise test on the track in Standardbred Trotters. Forty-four French trotters age 2-10 years performed a standardised exercise test consisting of three 3 min steps at constant speeds. Speed (V) of exercise varied between 500 and 750 m/min. Variables measured during exercise: SF, heart rate, RF using a microphone between the nostrils, blood lactate concentration. Physiological variables V4 and V200 were calculated and race performance index recorded. There was no age-related difference in RF or in LRC. Two LRC strategies were found: around 1/1 ratio and 3/2 ratio between SF and RF, respectively. A relationship between SF/RF ratio and physiological parameters showed a higher LRC in good performers because of a lower RF during submaximal and maximal exercise. RF is significantly lower and LRC significantly higher in good compared to poor performers. Investigation is required to ensure that the breathing techniques of trotters may be used in the selection process of racehorses.

A Mechanism for Vocal-Respiratory Coupling in the Mammalian Parabrachial Nucleus

Mammalian vocalizations require the precise coordination of separate laryngeal and respiratory motor pathways. Precisely how and where in the brain vocal motor patterns interact with respiratory rhythm control is unknown. The parabrachial nucleus (PB) is known to mediate key respiratory reflexes and is also considered a principle component of the mammalian vocal motor pathway, making it a likely site for vocal-respiratory interactions, yet a specific role for the PB in vocalizing has yet to be demonstrated. To investigate the role of the PB in vocal-respiratory coordination, we pharmacologically manipulated synaptic activity in the PB while spontaneously vocalizing horseshoe bats were provoked to emit either short, single syllable or long, multisyllabic vocal motor patterns. Iontophoresis of the GABAA agonist muscimol (MUS) into the lateral PB extended expiratory durations surrounding all vocalizations and increased mean call durations. Alternatively, application of the GABAA antagonist bicuculline methiodide (BIC) shortened expirations and call durations. In addition, BIC eliminated the occurrence of multisyllabic vocalizations. BIC caused a mild increase in quiet breathing rates, whereas MUS tended to slow quiet breathing. The results indicate that GABAA receptor-mediated inhibition in the lateral PB modulates the time course of respiratory phase switching during vocalizing, and is needed for proper coordination of calling and breathing in mammals. We hypothesize that vocal-respiratory rhythm entrainment is achieved at least in part via mechanisms similar to other forms of locomotor-respiratory coupling, namely somatosensory feedback influences on respiratory phase-switching in the lateral PB.

Neuro-mechanical and chemical influences on locomotor respiratory coupling in humans

Mechanisms of coordination between breathing and movement during dynamic exercise are still a matter of debate. This study aimed to examine the degree of coordination between breathing and arm propulsion patterns and to compare the relative contribution of neuro-mechanical and chemical factors. Thirteen trained cross-country skiers performed constant submaximal 6-min roller skiing exercises on a motorized driven treadmill in two different poling techniques (V2A and V2 skating techniques) at two exercise intensity levels corresponding to the first and second ventilatory thresholds. The timing of arm propulsion movements in V2A and V2 techniques was considered as a mechanical/neural influence on breathing whereas exercise intensity represented the metabolic demand to breathing. The degree of coordination, expressed as the percentage of breaths presenting a constant phase interval (time between an arbitrarily chosen point of the arm movement cycle and the onset of expiration) was significantly higher in V2A than V2 (P<0.05) while exercise intensity had no effect on the degree of coordination. We concluded that locomotor-respiratory coupling occurs in cross-country skiing as simulated by roller skiing because of strong influences from neuro-mechanical factors.

Degree of Coordination between Breathing and Rhythmic Arm Movements During Hand Rim Wheelchair Propulsion

This study aimed (i) to quantify the spontaneous coordination between breathing and hand rim wheelchair propulsion, (ii) to manipulate arm movement frequency and assess its effects on spontaneous coordination, and (iii) to investigate the hypothesis that entrainment of breathing improves economy of locomotion and leads to a lower rate of perceived exertion (RPE) compared with spontaneous breathing. Nine male, able-bodied participants completed four bouts of 6 min submaximal steady state exercise at 60 % of maximal propulsion velocity on a wheelchair ergometer, with spontaneous breathing and arm frequencies (Fspont), with 20 % higher and lower arm frequencies (F +20 and F -20, respectively) compared to Fspont accompanied with spontaneous breathing frequency, and by synchronising expiration phase with pushing time and inspiration phase with upper limb recovery time (C). Oxygen uptake and propulsion frequency were continuously recorded. The degree of coordination was expressed as a percentage of inspiration starting in the same phase of the wheelchair propulsion cycle (i.e. pushing and recovery times). No difference in degree of coordination was observed between Fspont, F -20 and F +20 conditions (49.2 +/- 12.1 %, 49.1 +/- 29.0 % and 48.2 +/- 29.4 %, respectively). Oxygen uptake increased significantly during C condition while RPE was significantly lower for C and F -20 (p < 0.05) compared to F +20 conditions. Contrary to what we expected, entrainment of breathing using a monofrequency ratio (C) induced a higher energy cost probably due to the mechanical properties of the wheelchair propulsion activity itself. In conclusion, this study showed that the same locomotor-respiratory coupling occurred during hand rim wheelchair propulsion regardless of the arm movement frequency, and that entrainment of breathing did not improve economy of locomotion.

Dynamic stability of locomotor respiratory coupling during cycling in humans

We explored the locomotor respiratory coupling (LRC) during a 50-min constant-load submaximal cycling exercise. A 4-week recombinant human erythropoietin (r-HuEPO) treatment improved participants’ aerobic capabilities, but did not elicit significant changes in LRC. The distributions of the respiratory frequency over pedalling frequency ratios were systematically bimodal, with a preferred use of 1/3 and 1/2, and a progressive shift of the higher mode from 1/3 towards 1/2 with exercise duration. These results are interpreted in the framework of the sine circle map as the result of coordination dynamics between the physiological subsystems involved in the breathing pedalling cooperation.