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Biomedical subjects

Ole K Andersen

Publications and source records attributed to Ole K Andersen.

At least 19 recordsLinked to original sources

Psychophysical 'perceptual maps' of heat and pain sensations by direct localization of CO2 laser stimuli on the skin.

Brain activation patterns derived from neurofunctional methods are often implicitly regarded as being directly related to subjective perceptual experience in an iso- or at least homomorph manner, neglecting the operational differences between these two dimensions. This paper (a) introduces a method for assessing 'perceptual maps' of stimulation patterns presented to the body surface, providing a means to parametrically relate neural representation and subjective percept, and (b) applies this method to demonstrate the existence of 'somatotopic maps' of hot and painful stimulus patterns independent from mechanoceptive co-activation. Brief (90 ms) CO2 laser pulses were presented in an array of multiple stimulation sites on the dorsal forearms (N. radialis area, C7 dermatome) of healthy subjects. Perceived locations were indicated with a 3D tracker without touching the skin, and (mis-)localizations in distal-proximal direction were analyzed. Stimuli were localized with overall mean errors of 22 mm (SD: 16 mm) toward the wrist and 24 mm (SD: 18 mm) toward the elbow. Somatotopic representation of thermal-nociceptive stimuli could be demonstrated in all subjects, independent from mechanoceptive co-activation. The perceptual maps revealed striking individual (mis-)localization patterns, many subjects exhibiting 'stretched', some 'condensed' somatotopic representations. In estimating the mapping parameters from physical to perceptual space linear regressions generally provided a good fit (adj. R2>0.80 in 10 out of 12 subjects). Nonlinear models were advantageous in some subjects only. Our method can be useful in assessing inter-individual differences or experimentally induced shifts in somatotopic processing.

Adult↗

Modulation of heat evoked nociceptive withdrawal reflexes by painful intramuscular conditioning stimulation.

Convergence between cutaneous heat nociceptors and muscles afferents was investigated by applying a phasic, conditioning electrical stimulus to the tibialis anterior muscle (a train of five 1 ms pulses over 21 ms) at varying time intervals relative to a thermal test stimulus used for evoking the withdrawal reflex in humans. The 200 ms thermal stimulus was applied on the dorsum of the foot at an intensity of two times the pain threshold. The conditioning electrical stimulus was applied at an intensity of two times the pain threshold via a set of intramuscular needle electrodes. The conditioning-test interval was varied between -400 ms and 8,000 ms at 17 different intervals. The mean reflex onset latency of reflexes evoked by thermal stimuli alone was 354 +/- 9 ms. A facilitation of the reflex was seen when the conditioning stimulus was applied 275 ms (174 +/- 30% compared to control) and 300 ms (162 +/- 32% compared to control) after the test stimulus onset indicating sensory convergence between muscle afferents (group I-III) and cutaneous Adelta heat nociceptors arriving simultaneously at the spinal cord.

Adult↗

Secondary heat hyperalgesia induced by melittin in humans.

Melittin, which is a principal protein of honeybee venom, can induce mechanical hyperalgesia in humans. The characteristics of the melittin induced mechanical hyperalgesia are quantitatively and qualitatively different from those evoked by capsaicin. The aim of the present study was to investigate in detail secondary heat hyperalgesia induced by melittin in humans. In six healthy volunteers, 10 microg of melittin was injected intradermally on the volar forearm, and VAS score to radiant heat stimuli (focused light from a xenon lamp) was assessed around the injection site 5, 30, and 60 min after injection. For normalization purposes, a pain rating index was calculated as the individual heat evoked VAS scores obtained after melittin divided by the individual baseline VAS scores. A two-way ANOVA revealed a significant increase of the pain rating index over time (F=3.6; P=0.03). The pain rating index at 60 min was significantly larger than at 5 min (P=0.04) and at 30 min (P=0.03). These results demonstrated slowly developing secondary heat hyperalgesia after injection of melittin. A possible contribution of peripheral inflammatory responses to the manifestation of secondary heat hyperalgesia is suggested, which in reality render the distinction between the primary and secondary area of heat hyperalgesia unnecessary.

Adult↗

Quantitative assessment of nociceptive processes in conscious dogs by use of the nociceptive withdrawal reflex.

OBJECTIVE: To investigate the feasibility of evoking the nociceptive withdrawal reflex (NWR) from fore and hind limbs in conscious dogs, score stimulus-associated behavioral responses, and assess the canine NWR response to suprathreshold stimulations. ANIMALS: 8 adult Beagles. PROCEDURE: Surface electromyograms evoked by transcutaneous electrical stimulation of ulnaris and digital plantar nerves were recorded from the deltoideus, cleidobrachialis, biceps femoris, and tibialis cranialis muscles. Train-of-five pulses (stimulus(train)) were used; reflex threshold (I(t train)) was determined, and recruitment curves were obtained at 1.2, 1.5, and 2 x I(t train). Additionally, a single pulse (stimulus(single)) was given at 1, 1.2, 1.5, 2, and 3 x I(t train). Latency and amplitude of NWRs were analyzed. Severity of behavioral reactions was subjectively scored. RESULTS: Fore- and hind limb I(t train) values (median; 25% to 75% interquartile range) were 2.5 mA (2.0 to 3.6 mA) and 2.1 mA (1.7 to 2.9 mA), respectively. At I(t train), NWR latencies in the deltoideus, cleidobrachialis, biceps femoris, and cranial tibialis muscles were not significantly different (19.6 milliseconds [17.1 to 20.5 milliseconds], 19.5 milliseconds [18.1 to 20.7 milliseconds], 20.5 milliseconds [14.7 to 26.4 milliseconds], and 24.4 milliseconds [17.1 to 40.5 milliseconds], respectively). Latencies obtained with stimulus(train) and stimulus(single) were similar. With increasing stimulation intensities, NWR amplitude increased and correlated positively with behavioral scores. CONCLUSIONS AND CLINICAL RELEVANCE: In dogs, the NWR can be evoked from limbs and correlates with behavioral reactions. Results suggest that NWR evaluation may enable quantification of nociceptive system excitability and efficacy of analgesics in individual dogs.

Animals↗

Gradual enlargement of human withdrawal reflex receptive fields following repetitive painful stimulation.

Dynamic changes in the topography of the human withdrawal reflex receptive fields (RRF) were assessed by repetitive painful stimuli in 15 healthy subjects. A train of five electrical stimuli was delivered at a frequency of 3 Hz (total train duration 1.33 s). The train was delivered in random order to 10 electrode sites on the sole of the foot. Reflexes were recorded from tibialis anterior, soleus, vastus lateralis, biceps femoris, and iliopsoas (IL). The RRF changes during the stimulus train were assessed during standing with even support on both legs and while seated. The degree of temporal summation was depending on stimulation site. At the most sensitive part of the RRF, a statistically significant increase in reflex size was seen after two stimuli while four stimuli were needed to observe reflex facilitation at less sensitive electrode sites. Hence, the region from which reflexes could be evoked using the same stimulus intensity became larger through the train, that is, the RRF was gradually expanding. Reflexes evoked by stimuli four and five were of the same size. No reflex facilitation was seen at other stimulus sites outside the RRF. In all muscles except in IL, the largest reflexes were evoked when the subjects were standing. In the ankle joint, the main withdrawal pattern consisted of plantar flexion and inversion when the subjects were standing while dorsi-flexion was prevalent in the sitting position. Up to 35 degrees of knee and hip flexion were evoked often leading to a lift of the foot from the floor during standing. In conclusion, a gradual expansion of the RRF was seen in all muscles during the stimulus train. Furthermore, the motor programme task controls the reflex sensitivity within the reflex receptive field and, hence, the sensitivity of the temporal summation mechanism.

Analysis of Variance↗

Breakdown of accommodation in nerve: a possible role for persistent sodium current.

BACKGROUND: Accommodation and breakdown of accommodation are important elements of information processing in nerve fibers, as they determine how nerve fibers react to natural slowly changing stimuli or electrical stimulation. The aim of the present study was to elucidate the biophysical mechanism of breakdown of accommodation, which at present is unknown. RESULTS: A model of a space-clamped motor nerve fiber was developed. It was found that this new model could reproduce breakdown of accommodation when it included a low-threshold, rapidly activating, persistent sodium current. However, the phenomenon was not reproduced when the persistent sodium current did not have fast activation kinetics or a low activation threshold. CONCLUSION: The present modeling study suggests that persistent, low-threshold, rapidly activating sodium currents have a key role in breakdown of accommodation, and that breakdown of accommodation can be used as a tool for studying persistent sodium current under normal and pathological conditions.

Animals↗

Orderly activation of human motor neurons using electrical ramp prepulses.

OBJECTIVE: Conventional electrical stimulation (rectangular pulses) recruits large before small diameter motor neurons and motor neurons close to the electrode before more distant motor neurons. The present study investigated the possibility for changing the recruitment order of electrical stimuli with sub-threshold ramp prepulses. METHODS: The median nerve was stimulated using surface electrodes at the wrist and elbow. Compound motor action potentials were recorded from abductor pollicis brevis and flexor carpi radialis. Stimulus-response curves, nerve conduction velocity and excitation thresholds of abductor pollicis brevis and flexor carpi radials, with and without ramp prepulses, were recorded in order to study the effect of ramp prepulses on axonal excitability. RESULTS: The conduction velocity of the initial response (10% of the maximal response) was decreased by 4.3+/-0.83m/s with ramp prepulses (500ms, 80% of the excitation threshold). The ramp prepulses also had a differential effect on the excitation thresholds of abductor pollicis brevis and flexor carpi radialis. In addition, ramp prepulses increased the threshold of 10% of the maximal response more than the threshold of 90% of the maximal response. CONCLUSIONS: These results demonstrate that large diameter and motor neurons close to the electrode accommodate more to ramp prepulses than small diameter and distant motor neurons, which suggests that ramp prepulses may be used to change the recruitment order of rectangular pulses. SIGNIFICANCE: This technique of ramp prepulses allows stimulation of alternate subsets of motor nerves.

Adult↗

Facilitation and inhibition of withdrawal reflexes following repetitive stimulation: electro- and psychophysiological evidence for activation of noxious inhibitory controls in humans.

A systematic evaluation of nociceptive withdrawal reflexes and pain rating was undertaken in order to explore the mechanisms underlying temporal summation of repetitive electrocutaneous stimulation in healthy individuals (n=12; age=27.5+/-1.5 years). Five-second subreflex threshold (RT) electrocutaneous stimulation at different frequencies (single stimulus, 5, 10, and 20 Hz) and intensities (0.6RT and 0.8RT) was applied on the dorsum of the foot, and the withdrawal reflex from the ipsilateral biceps femoris muscle was measured. The subjects scored the pain intensity on a visual analogue scale (0-100 mm) for the beginning, the middle and the end phase of the 5 s series of stimulation, and the respective averaged reflex size was calculated. The reflex size increased at stimulus frequencies 10 Hzx0.8RT and 20 Hzx0.8RT as compared with 5 Hzx0.8RT (SNK, P<0.05), and by an increase in current intensity from 0.6RT to 0.8RT (SNK, P<0.05). Pain intensity increased with the increase in the current intensity from 0.6RT to 0.8RT (SNK, P<0.05). Profound activation of inhibition following electrocutaneous pain stimuli was demonstrated by reduction in pain intensity and reflex size during the last second as compared with the first second at 0.6RT current intensity (SNK, P<0.05). The pain intensity peaked between 5 and 10 Hz (P<0.05) and was reduced at 20 Hz for current intensities at 0.8RT (P<0.05). This study provides evidence for both frequency dependent central integration of the repetitive electrocutaneous stimuli and activation of a pain inhibitory system by psychophysical and electrophysiological means, demonstrating the delicate balance between neuronal facilitation and inhibition in the human pain system.

Adult↗

Secondary heat hyperalgesia detected by radiant heat stimuli in humans: evaluation of stimulus intensity and duration.

Diverging observations on secondary hyperalgesia to heat stimuli have been reported in the literature. No studies have investigated the importance of heat stimulus intensity and duration for the assessment of secondary heat hyperalgesia. The present study was designed to investigate systematically (1) if pain sensitivity to radiant heat stimuli (focused Xenon light) is altered in the area of secondary punctuate hyperalgesia induced by intradermal injection of capsaicin and (2) if heat stimulus duration and intensity had an influence on the ability to detect secondary heat hyperalgesia. Pain ratings to radiant heat stimuli from a focused xenon lamp were assessed within the area of secondary punctuate hyperalgesia in fifteen volunteers before and after intradermal injection of capsaicin. The stimulus conditions were systematically varied between three intensity levels (0.8, 1.0 and 1.2 x heat pain threshold (PT)) and four duration steps (200, 350, 500 and 750 ms). The present study shows that long duration (350-750 ms) and low intensity (0.8 and 1.0 x PT) radiant heat stimuli were adequate to detect secondary heat hyperalgesia.

Adult↗

Repetitive painful stimulation produces an expansion of withdrawal reflex receptive fields in humans.

The aims of the present study were to investigate whether temporal summation of the nociceptive withdrawal reflex depends on the stimulation site on the sole of the human foot, and to characterize the reflex receptive fields (RRF) of lower limb muscles to repetitive stimulation. The cutaneous RRFs were assessed in 15 subjects in sitting position by recording the EMG from five lower leg muscles and the kinematic responses (ankle, knee, and hip joints) to repetitive painful electrical stimulation. The stimulus consisted of a series of five stimuli (frequency: 3 Hz) delivered randomly at 10 different sites on the sole of the foot. The size of the reflexes increased generally between the first and the second stimulus, however, the increment depended on the stimulation site. In tibialis anterior, the RRF covered the distal sole of the foot and gradually expanded during the stimulus train. No expansion toward the heel area was detected. In soleus, the reflexes were facilitated after the second stimulus at all sites and remained in this state until the last stimulus. In vastus lateralis, biceps femoris, and iliopsoas a gradual expansion of the RRF was seen, resulting in RRFs covering the lateral, distal foot, and part of the proximal foot (iliopsoas). Knee and hip flexion were evoked at all sites. Ankle dorsiflexion was evoked at the distal foot, while ankle plantarflexion was evoked at the heel. The enlargement of the RRF reflects spinal temporal summation leading to gradually stronger reflex responses. The degree of temporal summation was dependent on stimulation site. The facilitation of the withdrawal reflex responses due to repetitive stimulation might have potential applications in the rehabilitation engineering field, where these reflexes could be used to assist gait of patients with central nervous system injuries.

Adult↗

Effect of romifidine on the nociceptive withdrawal reflex and temporal summation in conscious horses.

OBJECTIVE: To investigate the action of a single IV administration of romifidine on the thresholds of the nociceptive withdrawal reflex (NWR) and temporal summation in conscious horses. ANIMALS: 10 adult horses. PROCEDURE: Single electrical stimulations were applied on the digital nerves to evoke NWR from the left forelimb and hind limb. Repeated electrical stimulations (10 stimuli, 5 Hz) were given to obtain temporal summation. Surface electromyographic reflex activity was recorded from the common digital extensor and cranial tibial muscles. After baseline assessment of NWR and temporal summation thresholds, romifidine (80 microg x kg(-1), IV) was administered. Successive determinations of NWR and temporal summation thresholds were performed 5, 25, and 55 minutes after administration. RESULTS: Romifidine significantly increased the current intensities necessary to evoke NWR and temporal summation in forelimbs and hind limbs of horses. Values were significantly higher than baseline values 55 minutes after romifidine administration. After administration of romifidine, a facilitation of reflex components of tactile origin was observed when repeated stimulations were applied. CONCLUSIONS AND CLINICAL RELEVANCE: Results confirm antinociceptive activity of romifidine and may represent an objective demonstration of the well-known hypersensitivity to tactile stimuli observed in horses receiving alpha2-adrenoreceptor agonists in clinical practice. Romifidine can be included in analgesic and anesthetic protocols to provide additional analgesia in horses.

Adrenergic alpha-Agonists↗

Expansion of nociceptive withdrawal reflex receptive fields in spinal cord injured humans.

OBJECTIVE: In spinal cord injured (SCI) subjects, exaggerated withdrawal reflexes associated with a dominant flexor pattern irrespective of stimulation site have been reported. In the present study, withdrawal reflex receptive field (RRF) was determined in complete SCI subjects (N=9). METHODS: Distributed electrical stimulation was applied to the sole of the foot, and reflexes in tibialis anterior, soleus, biceps femoris, and vastus lateralis muscles were recorded together with knee and ankle movement trajectories. A group of spinally intact subjects (N=10) were included as controls. With the subjects in supine position, stimulation was applied to 10 different sites on the foot sole. Based on the tibialis anterior reflex threshold for stimulation on the mid foot sole, two stimulus intensities (1.1 times the reflex threshold and 1.4 times the reflex threshold) were used for all 10 sites. RESULTS: In SCI subjects, dorsi-flexion dominated independent of stimulus site and the tibialis anterior RRF covered the entire foot sole in contrast to a well-defined tibialis anterior receptive field at the medial, distal foot sole in the spinally intact subjects. Further, the soleus RRF also covered the entire sole in the SCI subjects. The reflexes in biceps femoris and vastus lateralis muscles were small and associated with weak knee flexion at all 10 sites in the SCI subjects and in the controls. CONCLUSIONS: The RRF of the ankle flexor and the ankle extensor muscles both covered the entire sole of the foot indicating an expansion of the RRFs following spinal cord injury. The expansion is most likely due to lack of descending inhibitory control and/or increased sensitivity of the spinal reflex loop in the SCI subjects. SIGNIFICANCE: The study improves the understanding of spinal reflex control in spinal intact and spinal cord injured subjects.

Adult↗

Evidence for spinal cord hypersensitivity in chronic pain after whiplash injury and in fibromyalgia.

Patients with chronic pain after whiplash injury and fibromyalgia patients display exaggerated pain after sensory stimulation. Because evident tissue damage is usually lacking, this exaggerated pain perception could be explained by hyperexcitability of the central nervous system. The nociceptive withdrawal reflex (a spinal reflex) may be used to study the excitability state of spinal cord neurons. We tested the hypothesis that patients with chronic whiplash pain and fibromyalgia display facilitated withdrawal reflex and therefore spinal cord hypersensitivity. Three groups were studied: whiplash (n=27), fibromyalgia (n=22) and healthy controls (n=29). Two types of transcutaneous electrical stimulation of the sural nerve were applied: single stimulus and five repeated stimuli at 2 Hz. Electromyography was recorded from the biceps femoris muscle. The main outcome measurement was the minimum current intensity eliciting a spinal reflex (reflex threshold). Reflex thresholds were significantly lower in the whiplash compared with the control group, after both single (P=0.024) and repeated (P=0.035) stimulation. The same was observed for the fibromyalgia group, after both stimulation modalities (P=0.001 and 0.046, respectively). We provide evidence for spinal cord hyperexcitability in patients with chronic pain after whiplash injury and in fibromyalgia patients. This can cause exaggerated pain following low intensity nociceptive or innocuous peripheral stimulation. Spinal hypersensitivity may explain, at least in part, pain in the absence of detectable tissue damage.

Adult↗

Comparison of hyperalgesia induced by capsaicin injection and controlled heat injury: effect on temporal summation.

The relationship between induction of central sensitization and facilitation of temporal summation to repetitive stimulation is still unclear. The aim of this study was to investigate temporal summation before and after the induction of secondary hyperalgesia by two different experimental methods: capsaicin injection and controlled heat injury. The effect of each injury model was assessed on a separate day with an interval of at least 5 days. Twelve healthy volunteers participated. Each experiment was performed using electrical, radiant heat, mechanical impact, and punctuate stimuli consecutively. The pain threshold (PT) to a single stimulus and the summation threshold to five repetitive stimuli for electrical (2 Hz) and radiant heat (0.83 Hz) were assessed within the secondary hyperalgesic area. The degree of temporal summation for stimulus intensities of 0.8, 1.0, and 1.2 times the baseline pain thresholds were evaluated by the increase in visual analogue scale (VAS) scores from the first to the fifth stimulus of the train. Further, the degrees of temporal summation were assessed for mechanical impact and punctuate stimuli within the primary and secondary hyperalgesic areas. The contra-lateral forearm served as control (no injury). The pain threshold and the summation threshold to electrical and heat stimuli decreased significantly within the secondary hyperalgesic area after the injury induced by both heat injury or capsaicin injection. However, there was no temporal summation for heat and electrical stimuli in either model. In contrast, for the mechanical impact and punctuate mechanical stimuli the degree of temporal summation was significantly facilitated in the secondary hyperalgesic areas compared with the baseline and the control arm in both models. In the primary hyperalgesic area, the degree of temporal summation was facilitated to mechanical impact and punctuate stimuli but only following the capsaicin injection. In conclusion, the temporal summation mechanism for mechanical stimuli was facilitated in the secondary hyperalgesic area.

Adult↗

Investigation of the facilitation of the nociceptive withdrawal reflex evoked by repeated transcutaneous electrical stimulations as a measure of temporal summation in conscious horses.

OBJECTIVE: To investigate whether facilitation of the nociceptive withdrawal reflex (NWR) can be evoked and quantified as a measure of temporal summation from the distal aspect of the left forelimb and hind limb in standing nonsedated horses via repeated stimulations of various subthreshold intensities and frequencies. ANIMALS: 10 adult horses. PROCEDURE: Surface electromyographic activity evoked by stimulation of the digital palmar and plantar nerves was recorded from the common digital extensor and cranial tibial muscles. For each horse, the NWR threshold intensity to a single stimulus was determined for the forelimb and hind limb. Repeated stimulations were performed at subthreshold intensities and at frequencies of 2, 5, and 10 Hz. The reflex amplitude was quantified, and the behavioral responses accompanying the stimulations were scored. RESULTS: Repeated stimulations at subthreshold intensities were able to summate and facilitate the NWR in conscious horses. The reflex facilitation was significantly related to the intensity of the repeated stimuli, whereas no effect of stimulation frequency was found. Reaction scores increased significantly for increasing stimulation intensities. CONCLUSIONS AND CLINICAL RELEVANCE: Temporal summation obtained by repeated stimulations of subthreshold intensity appears to represent a new tool for investigating nociceptive pathophysiologic processes in horses; this experimental model may be useful to examine the mode of action and efficacy of analgesic and anesthetic interventions and possibly to assess sensory dysfunction in clinical settings.

Analysis of Variance↗

Modulation of lower limb withdrawal reflexes during gait: a topographical study.

The aim of this study was to investigate the modulation and topography of the nociceptive withdrawal reflex elicited by painful electrical stimulation of the foot sole during gait. Fifteen healthy volunteers participated in this study. Cutaneous electrical stimulation was delivered on five locations of the foot sole after heel-contact, during foot-flat, after heel-off, and during the mid-swing phase of the gait cycle during treadmill walking. Reflexes were recorded from muscles of the ipsilateral and contralateral legs. Furthermore, the kinematic responses in the sagittal plane of the ipsilateral ankle, knee, and hip joints were recorded. Reflexes in the distal muscles showed a site-dependent modulation. The largest responses in tibialis anterior were evoked at the arch of the foot and the smallest at the heel (P < 0.05). The largest soleus responses were also elicited at the arch of the foot (P < 0.04). The EMG responses in flexors and extensors of the knee and extensors of the contralateral leg were generally not dependent on the stimulation site. The response at the three joints showed site dependency, especially during the swing phase where maximal flexion was obtained by stimulation at the arch of the foot (P < 0.05). The withdrawal reflex was modulated during the gait cycle and presented distinctive characteristics for the different muscles studied. Minimal kinematic responses were observed during stance in contrast to swing phase. Modulation of the reflex probably ensures an appropriate withdrawal but primarily secures balance and continuity of movement.

Adult↗

Foot-sole reflex receptive fields for human withdrawal reflexes in symmetrical standing position.

Human withdrawal-reflex receptive fields were assessed in 10 healthy subjects during standing with even support on both legs. Two electrical-stimulus intensities (1.2 and 2.2 times the pain threshold, PTh) were used. The painful stimuli were delivered in random order to 12 positions distributed over the foot sole. Tibialis anterior (TA), soleus (SO), vastus lateralis (VL), semitendinosus (ST), and iliopsoas (IL) reflexes were recorded. Further, the vertical force was recorded and the center or pressure (CoP) was assessed in the frontal and sagittal planes on both legs. Reflexes were observed at both intensities with the strongest reflexes at the high intensity. Around the ankle joint, SO reflexes dominated, which is in contrast to previous observations for subjects sitting. An unloading of the limb was found on the stimulated leg associated with a simultaneous loading of the contralateral leg. The shift in load was most pronounced for stimulation of the heel. The flexors ST and IL also had strong reflexes with reflex patterns correlated to the pattern of unloading. The shift in vertical force was accomplished by a move of the CoP in the anterior direction on the stimulated limb (contraction of SO), which simultaneously caused a small movement of the CoP in the lateral direction. In the present standing conditions, the ankle extensor played a dominant role in the withdrawal pattern in contrast to previous studies during sitting, relaxed conditions.

Adult↗