Changes in stretch reflexes and muscle stiffness with age in prepubescent children.

Musculo-articular stiffness of the triceps surae (TS) increases with age in prepubescent children, under both passive and active conditions. This study investigates whether these changes in muscle stiffness influence the amplitude of the reflex response to muscle stretch. TS stiffness and reflex activities were measured in 46 children (7-11 yr old) and in 9 adults. The TS Hoffmann reflex (H reflex) and T reflex (tendon jerk) in response to taping the Achilles tendon were evaluated at rest and normalized to the maximal motor response (Mmax). Sinusoidal perturbations of passive or activated muscles were used to evoke stretch reflexes and to measure passive and active musculoarticular stiffness. The children's Hmax-to-Mmax ratio did not change with age and did not differ from adult values. The T-to-Mmax ratio increased with age but remained significantly lower than in adults. Passive stiffness also increased with age and was correlated with the T-to-Mmax ratio. Similarly, the children's stretch reflex and active musculoarticular stiffness were significantly correlated and increased with age. We conclude that prepubescent children have smaller T reflexes and stretch reflexes than adults, and the lower musculoarticular stiffness is mainly responsible for these smaller reflexes, as indicated by the parallel increases in reflex and stiffness.     

Mechanisms of reflexes induced by esophageal distension

We investigated the mechanisms of esophageal distension-induced reflexes in decerebrate cats. Slow air esophageal distension activated esophago-upper esophageal sphincter (UES) contractile reflex (EUCR) and secondary peristalsis (2P). Rapid air distension activated esophago-UES relaxation reflex (EURR), esophago-glottal closure reflex (EGCR), esophago-hyoid distraction reflex (EHDR), and esophago-esophagus contraction reflex (EECR). Longitudinal esophageal stretch did not activate these reflexes. Magnitude and timing of EUCR were related to 2P but not injected air volume. Cervical esophagus transection did not affect the threshold of any reflex. Bolus diversion prevented swallow-related esophageal peristalsis. Lidocaine or capsaicin esophageal perfusion, esophageal mucosal layer removal, or intravenous baclofen blocked or inhibited EURR, EGCR, EHDR, and EECR but not EUCR or 2P. Thoracic vagotomy blocked all reflexes. These six reflexes can be activated by esophageal distension, and they occur in two sets depending on inflation rate rather than volume. EUCR was independent of 2P, but 2P activated EUCR; therefore, EUCR may help prevent reflux during peristalsis. All esophageal peristalsis may be secondary to esophageal stimulation in the cat. EURR, EHDR, EGCR, and EECR may contribute to belching and are probably mediated by capsaicin-sensitive, rapidly adapting mucosal mechanoreceptors. GABA-B receptors also inhibit these reflexes. EUCR and 2P are probably mediated by slowly adapting muscular mechanoreceptors. All six reflexes are mediated by vagal afferent fibers.     

Influence of long-term spaceflight on neuromechanical properties of muscles in humans.

Reflex and elastic properties of the triceps surae (TS) were measured on 12 male cosmonauts 28-40 days before a 3- to 6-mo spaceflight, 2 or 3 days after return (R+2/+3) and a few days later (R+5/+6). H reflexes to electrical stimulations and T reflexes to tendon taps gave the reflex excitability at rest. Under voluntary contractions, reflex excitability was assessed by the stretch reflex, elicited by sinusoidal length perturbations. Stiffness measurements concerned the musculoarticular system in passive conditions and the musculotendinous complex in active conditions. Results indicated 1) no changes (P > 0.05) in H reflexes, whatever the day of test, and 2) increase in T reflexes (P < 0.05) by 57%, despite a decrease (P < 0.05) in musculoarticular stiffness (11%) on R+2/+3. T reflexes decreased (P < 0.05) between R+2/+3 and R+5/+6 (-21%); 3) increase in stretch reflexes (P < 0.05) on R+2/+3 by 31%, whereas it decreased (P < 0.05) between R+2/+3 and R+5/+6 (-29%). Musculotendinous stiffness was increased (P < 0.05) whatever the day of test (25%). Links between changes in reflex and stiffness were also studied by considering individual data. At R+2/+3, correlated changes between T reflexes and musculoarticular stiffness suggested that, besides central adaptive phenomena, musculoarticular structures took part in the reflex adaptation. This mechanical contribution was confirmed when data collected at R+2/+3 and R+5/+6 were used because correlations between changes in stretch reflexes and musculotendinous stiffness were improved. In conclusion, the present study shows that peripheral influences take part in reflex changes in gravitational unloaded muscles, but can only be revealed when central influences are reduced.     

Spatial coordination of compensatory eye movements in vertebrates: form and function

The semicircular canals of the labyrinth of vertebrates provide one way of motion detection in three-dimensional space. The fully developed form of the vertebrate labyrinth consists of six semicircular canals, three on each side of the head, whose spatial arrangement (vertical canals are placed diagonally in the head, horizontal canals are oriented earth horizontally) follows three interconnected principles: 1) bilateral symmetry, 2) push-pull operational mode, and 3) mutual orthogonality. Other sensory and motor systems related to vestibular reflexes, such as the extraocular muscles or the "optokinetic" coordinate axes encoded in the activity of the visually driven cells of the accessory optic system, share the same geometrical framework. This framework is also reflected in the anatomical networks mediating compensatory eye movements, linking each of the semicircular canals to a particular set of extraocular muscles (so-called principal vestibuloocular reflex connections to yoke muscles). These classical vestibulo-oculomotor relationships have been verified at many levels of the vertebrate hierarchy, including lateral- and frontal-eyed animals. The particular spatial orientation of the semicircular canals requires further comment and phylogenetic evaluation. The spatial arrangement of the vertical canals is already present in fossil ostracoderms, and is also exemplified in lampreys, the modern forms of once abundant agnathan species that populated the Silurian and Devonian oceans. The lampreys and ostracoderms lack horizontal canals, which appear later in all descendent vertebrates. The fully developed vertebrate labyrinth with its six semicircular canals displays distinct differences that are obvious when comparing distant taxa (e.g. elasmobranchs versus other vertebrates). Whereas the common crus of the semicircular canals in teleosts through mammals is formed between the anterior and the posterior semicircular canal, it occurs between the anterior and the horizontal canal in elasmobranchs. However, despite this morphological difference, these two vertebrate labyrinth prototypes constitute a functionally identical solution. A similar analysis holds for certain invertebrate species (crab, octopus, squid), which display an even wider variety in the physical expressions of movement detection systems when compared to vertebrates. Although the physical expressions of motion detection systems differ in the animal kingdom, the functional solutions (providing the best signal-to-noise ratio) with adherence to bilateral symmetry, push-pull operational mode, and mutual orthogonality are identical.(ABSTRACT TRUNCATED AT 400 WORDS)     

Modeling movement disorders--CRPS-related dystonia explained by abnormal proprioceptive reflexes

Humans control their movements using adaptive proprioceptive feedback from muscle afferents. The interaction between proprioceptive reflexes and biomechanical properties of the limb is essential in understanding the etiology of movement disorders. A non-linear neuromuscular model of the wrist incorporating muscle dynamics and neural control was developed to test hypotheses on fixed dystonia. Dystonia entails sustained muscle contractions resulting in abnormal postures. Lack of inhibition is often hypothesized to result in hyperreflexia (exaggerated reflexes), which may cause fixed dystonia. In this study the model-simulated behavior in case of several abnormal reflex settings was compared to the clinical features of dystonia: abnormal posture, sustained muscle contraction, increased stiffness, diminished voluntary control and activity-aggravation. The simulation results were rated to criteria based on characteristic features of dystonia. Three abnormal reflex scenarios were tested: (1) increased reflex sensitivity-increased sensitivity of both the agonistic and antagonistic reflex pathways; (2) imbalanced reflex offset-a static offset to the reflex pathways on the agonistic side only; and (3) imbalanced reflex sensitivity-increased sensitivity of only the agonistic reflex pathways. Increased reflex sensitivity did not fully account for the features of dystonia, despite distinct motor dysfunction, since no abnormal postures occurred. Although imbalanced reflex offset did result in an abnormal posture, it could not satisfy other criteria. Nevertheless, imbalanced reflex sensitivity with unstable force feedback in one of the antagonists closely resembled all features of dystonia. The developed neuromuscular model is an effective tool to test hypotheses on the underlying pathophysiology of movement disorders.     

Stabilization of the gaze in chameleons: visual and vestibular reflexes

Some visual, vestibular and proprioceptive reflexes which contribute to gaze (head + eye) stabilization were quantified in the chameleon. All the reflexes were analysed in the horizontal plane, and the visual reflexes were also studied in the vertical plane. In restrained-head animals, both the optokinetic nystagmus (OKN) and the vestibulo-ocular reflex (VOR) had low gains. In free-head animals, the head (opto-collic or vestibulo-collic reflex) and eye (OKN or VOR) responses added their effects, thus improving gaze stabilization, especially during vestibular stimulation. Cervical stimulation provoked both a cervico-ocular reflex (COR) in the compensatory direction and a large number of saccades. The saccadic response was especially marked in the presence of patterned visual surroundings.     

Activity of the laryngeal abductor and adductor muscles during cough, expiration and aspiration reflexes in cats

We studied the temporal relationships and the patterns of electromyographic activities of the posterior cricoarytenoid and thyreoarytenoid muscles (laryngeal abductor and adductor), the diaphragm and abdominal muscles in anesthetized cats during mechanically induced tracheobronchial and laryngopharyngeal coughs, expiration and aspiration reflexes. The posterior cricoarytenoid muscle activity reached the maxima just before the peak of diaphragmatic activity in both types of cough and aspiration reflexes and slightly before the top of abdominal muscle activity in coughs and the expiration reflex. Thus, this muscle contributes to the inspiratory phase of coughs and aspiration reflex and also to the expulsive phase of coughs and the expiration reflex. The thyreoarytenoid muscle presented strong discharges in the compressive phase of coughs and expiration reflex (during the rising part of the abdominal muscle activity) and in the subsequent laryngoconstriction (following the diaphragmal and/or abdominal muscle activity) in all four reflexes. This muscle was also slightly activated at the beginning of the aspiration reflex. The existence of four phases of the cough reflex is also discussed.     

Reflex regulation of the circulation after stimulation of cardiac receptors by prostaglandins

Prostaglandins (PGs) are potent vasoactive substances that may participate in the control of coronary blood flow, platelet aggregation, and inflammation. An important action of PGs may be the stimulation of c fibers in general and vagal cardiac c fibers in particular. The Bezold-Jarisch reflex after intracoronary injection of Veratrum alkaloids is very similar to the vagal bradycardia elicited by stimulation of cardiac PG synthesis or injection of prostacyclin (PGI2). The characteristic features of this reflex are 1) stimulation of c fibers, 2) inferoposterior wall location of receptors, 3) vagal afferents, 4) vagal efferents to the heart, 5) sympathetic efferents to peripheral blood vessels, and 6) interaction with other reflexes. Vagal cardiac c fibers are activated by intracoronary injections of PGI2 or arachidonic acid, resulting in a vagal reflex bradycardia and hypotension due to withdrawal of peripheral alpha-adrenergic tone to resistance vessels. The cardiac receptors are located predominantly in the inferoposterior wall of the left ventricle. When stimulated by PGs, cardiac receptors may also modify the regulation of arterial pressure by the baroreflexes, altering the inverse relationship between systemic arterial pressure and heart rate. Thus, there is a striking parallelism between the veratridine-induced Bezold-Jarisch reflex and PG-induced cardiac reflexes, although the physiological and clinical significance of these reflexes remains to be determined.     

Influence of CPAP on reflex responses to tracheal irritation in anesthetized humans

We investigated the effects of lung inflation during continuous positive airway pressure breathing (CPAP) on airway defensive reflexes in 10 enflurane-anesthetized spontaneously breathing humans. The airway defensive reflexes were induced by instillation into the trachea of 0.5 ml of distilled water at two different levels of end-expiratory pressure (0 and 10 cmH2O CPAP). The tracheal irritation at an end-expiratory pressure of 0 cmH2O caused a variety of reflex responses including apnea, spasmodic panting, expiration reflex, cough reflex, an increase in heart rate, and an increase in blood pressure. Lung inflation during CPAP of 10 cmH2O did not exert any influence on these reflex responses in terms of the types, latencies, and durations of reflex responses although the intensity of the expiration reflex and cough reflex was augmented by lung inflation. Our results suggest that the pulmonary stretch receptors do not play an important role in the mechanisms of airway defensive reflexes in humans.     

Functional characteristics and topography of spinal pathways conducting high- and low-threshold startle reflexes

Electrophysiological and topographical properties of the spinal tract systems involved in two functional types of startle reflexes were studied in chloralose anesthetized cats: a high-threshold reflex produced by intense peripheral nerve stimulation (spinobulbo-spinal, SBS, reflex) and a low-threshold reflex evoked by tactile (T-reflex) and acoustic (A-reflex) stimulation. Maximum conduction velocity of descending transmission of the high-threshold reflex, at 30 m/sec, was perceptibly lower than that of low-threshold reflexes, at 85 m/sec for T-reflex and 100 m/sec for A-reflex. Mean conduction velocity for SBS and T-reflexes were 40.2 and 70.8 m/sec respectively. Perceptible differences were also found in the topography of spinal and especially ascending pathways of these reflexes. It was established by partial spinal cord destruction that accomplishment of T-reflex depended on the integrity of ascending pathways of the dorsal and dorsolateral funiculi and the SBS reflex on preservation of the dorsolateral, ventrolateral and (partially) of ventral funiculi. Descending pathways of the reflexes under study were revealed mainly in the ventrolateral and ventral funiculi and those of the SBS reflex mainly in the first of these. Findings also show the noticeable similarity between the organization of both T- and A-reflex descending pathways. The functional organization of the spinal pathways of a variety of startle reflexes is discussed.A. A. Bogomolets Institute of Physiology, Academy of Sciences of the Ukrainian SSR, Kiev. Translated from Neirofiziologiya, Vol. 18, No. 4, pp. 486–496, July–August, 1986.     

Effects of stimulating the periaqueductal gray matter on high- and low-threshold startle reflexes

The effects of stimulating the periaqueductal gray matter (PAG) on two types of startle reflex (spino-bulbo-spinal reflex produced by intensive stimulation of the peripheral nerves and low-threshold tactile spino-reticulo-spinal reflex) as well as high-threshold jaw-opening reflex arising in response to tooth pulp stimulation were investigated in cats anesthetized with chloralose. Simulating most PAG test sites led to pronounced inhibition of jaw-opening reflex, profound depression of spino-bulbo-spinal reflex, and moderate inhibition of tactile reflexes. The facilitatory effect of stimulating a number of PAG sites on the latter reflexes was demonstrated. Effects of PAG stimulation fell into two classes: brief, measurable in hundreds of msec and more prolonged, measured in minutes and seconds. Findings would indicate certain differences between the effects of PAG stimulation low-threshold (non-nociceptive) and high-threshold (nociceptive) startle reflexes, of which the possible mechanisms are discussed.A. A. Bogomolets Institute of Physiology, Academy of Sciences of the Ukrainian SSR, Kiev. Translated from Neirofiziologiya, Vol. 21, No. 1, pp. 71–78, January–February, 1989.     

Supraspinal influences on the penile reflexes of the male rat: a comparison of the effects of copulation, spinal transection, and cortical spreading depression.

The penile reflexes of the rat were observed on interruption of the copulatory behavior sequence after intromission and ejaculation in the initial ejaculatory series, after the penultimate series, during sexual exhaustion, and during recovery from sexual exhaustion 24 and 72 hr later. These were compared to the reflexes of the normal rat in control conditions, to those of the male rat after spinal transection, and to those of the sexually rested and sexually exhausted male rat under cortical spreading depression (CSD). It was concluded that (1) the stimuli associated with copulation evoke disinhibition of the penile reflexes, these showing the short reflex latencies observed in the spinal animal. The release of the spinal mechanisms is lost within 30 min of the last copulatory event. CSD further inhibits reflex responsivity. (2) Stimuli associated with intromission provoke acceleration of the normal rhythmic presentation of reflexes seen in the normal and spinal rat, resulting in a decrease in the duration of intervals between reflex clusters and an increase in reflex number. This excitation decays within about 15 min after intromission. (3) The increase in degree of penile extension and percentage of penile flips after spinal transection suggests tonic inhibition of reflex intensity in the normal rat. The decrease in capacity to attain full erection with the approach of sexual exhaustion suggests an increase in this inhibition. This does not recover during a rest period but instead intensifies. CSD effects did not mimic the effects of spinal transection but instead depressed reflex excitability. The relationship of these changes to the copulatory behavior pattern is discussed.     

An experimental psychophysiological approach to human bradycardiac reflexes

Bradycardic reflexes in man are both of scientific and clinical interest. Using the methods of experimental psychophysiology, control over relevant independent variables permits the study of fine-grained temporal physiologic response topographies, and of psychological factors that may modify the reflex. In addition, information can also be sought through interdisciplinary collaborations with experimental physiologists in order to shed light on the mechanism of the reflexes. These general features of the approach are illustrated by presenting data on two bradycardic reflex preparations: the laboratory dive analog, and the 90-degree negative tilt. The dive-analog studies have shown that a) the dive-reflex proper is a late-occurring bradycardia accompanied by a late-occurring vasoconstriction; and b) for the elicitation of this reflex, both breath-holding and face immersion are necessary. In addition, the physiologic manipulation of temperature affects the reflex in an inverse way over the range of 10 degrees to 40 degrees C, while the sense of control (a psychological variable) attenuates the reflex. The negative-tilt preparation produces a bradycardic response that is ideal as a Pavlovian unconditional response. Some Pavlovian conditioning arrangements, especially an "imaginational" form, do produce significant conditional bradycardic responding, and this has both potential clinical (e.g., biofeedback-related) and theoretical (e.g., S-R vs. S-S accounts of Pavlovian conditioning) applications. The paper ends with a comment on the cognitive paradigm shift in psychology. Although this shift is of importance, it is suggested that it is also important to "remember the response."     

The Validity and Reliability of Motion Analysis in Patellar Tendon Reflex Assessment

Background

The deep tendon reflex assessments that are essential to the accurate diagnosis of neurological or neuromuscular disorders are conducted subjectively in clinical neurology. Our aim was to assess deep tendon reflexes objectively with a new reflex quantification method.

Methodology/Principal Findings

The present study used a motion analysis technique to collect quantitative measurements for both the input and output of normal patellar tendon reflex. Reflex responses were measured as knee angles. The patellar tendon reflexes of 100 healthy subjects were examined using 6 levels of tendon taps, where all the assessments were captured using motion capture system. A linear relationship was found between the experimental maximum tapping velocity and tapping angle (coefficient of determination = 0.989), which was consistent with the theoretical values. Tapping velocities were predictable according to tapping angles. The findings proved the reproducibility of tapping method in producing consistent input. The reflex amplitude was consistent between two randomly assigned groups, and linearly proportionate to the tapping velocity.

Conclusions/Significance

The findings on reflex amplitude indicate that motion analysis is a valid and reliable method of assessing and measuring deep tendon reflexes.     

The vestibulo-ocular reflex of hypergravity rats

The vertebrate vestibular system detects linear (otolith organs) and angular (semicircular canals) acceleration. The function of the otolith system is twofold, 1: perception of linear acceleration of the head, and 2: assessment of the spatial orientation of the head relative to the vector of gravity. Because of the latter function, a change of gravity will affect the vestibular input which, in turn, may have a wide range of serious physiological effects, for instance on ocular reflexes. The function of the vestibulo-ocular reflex (VOR) is to stabilize the visual image on the retina. Measurement of this VOR provides a method to investigate the (processing within the) vestibular system. Discrimination between gravity and linear acceleration, caused by movement of the head, is not possible. Therefore, information from the otolith system must be constantly compared with additional information from other sensory systems in order to solve the inherent ambiguity between tilt and translation. In this processing, cues from the semicircular canals also play a role. During parabolic flight, experiments can be performed at altered gravity levels for brief periods of time. On earth, the only effective possibility to manipulate gravity for longer periods of time is a centrifuge. Together with experiments in weightlessness during orbital flight, these methods form useful tools to investigate the influence of gravity on physiology. In our laboratory, rats have been kept inside a centrifuge at 2.5 g during their entire life-span (i.e. including gestation).     

Effect of Inhibition of NO Synthases on Cardiogenic Depressor Reflexes in Different Animal Species

We studied the role of the nitric oxide (NO) system in the realization of cardiogenic depressor reflexes evoked by stimulation of cardiac receptors by veratrine (reproduction of the Bezold–Jarish reflex). Acute experiments were performed on anesthetized dogs and rats: we tested the effects of inhibition of dissimilar isoforms of NO synthase (NOS) and paid special attention to possible species-related differences in realization of the reflex responses. We found that systemic inhibition of NOS by L-nitro-N-arginine (L-NNA, 30 mg/kg, i.v.) significantly decreased the depressor reflex reaction in dogs. Vasomotor dilatatory reactions of the peripheral vessels underwent considerable modifications and in some cases were converted into vasoconstrictory responses. Selective inhibition of neuronal NOS (nNOS) by 7-nitroindazole (7-NI, 25 mg/kg, i.p.) exerted no effect on the development of cardiogenic depressor reflexes in dogs. At the same time, systemic inhibition of NOS in the course of reproduction of cardiogenic depressor reflexes in rats resulted in intensification of depressor responses, while inhibition of nNOS decreased these reactions. Thus, we first demonstrated the role of NO in the realization of cardiogenic depressor reflexes under in vivo conditions and described species-related peculiarities of the involvement of the NO system in the development of these reflexes. We also demonstrated the dependence of formation of cardiogenic depressor reflexes on the predominant involvement of one NOS type or another.     

Possible contributions of CPG activity to the control of rhythmic human arm movement

There is extensive modulation of cutaneous and H-reflexes during rhythmic leg movement in humans. Mechanisms controlling reflex modulation (e.g., phase- and task-dependent modulation, and reflex reversal) during leg movements have been ascribed to the activity of spinal central pattern generating (CPG) networks and peripheral feedback. Our working hypothesis has been that neural mechanisms (i.e., CPGs) controlling rhythmic movement are conserved between the human lumbar and cervical spinal cord. Thus reflex modulation during rhythmic arm movement should be similar to that for rhythmic leg movement. This hypothesis has been tested by studying the regulation of reflexes in arm muscles during rhythmic arm cycling and treadmill walking. This paper reviews recent studies that have revealed that reflexes in arm muscles show modulation within the movement cycle (e.g., phase-dependency and reflex reversal) and between static and rhythmic motor tasks (e.g., task-dependency). It is concluded that reflexes are modulated similarly during rhythmic movement of the upper and lower limbs, suggesting similar motor control mechanisms. One notable exception to this pattern is a failure of contralateral arm movement to modulate reflex amplitude, which contrasts directly with observations from the leg. Overall, the data support the hypothesis that CPG activity contributes to the neural control of rhythmic arm movement.     

Self-organization of reflexive behavior from spontaneous motor activity

In mammals, the development of reflexes is often regarded as an innate process. However, recent findings show that fetuses are endowed with favorable conditions for ontogenetic development. In this article, we hypothesize that the circuitry of at least some mammalian reflexes can be self-organized from the sensory and motor interactions brought forth in a musculoskeletal system. We focus mainly on three reflexes: the myotatic reflex, the reciprocal inhibition reflex, and the reverse myotatic reflex. To test our hypothesis, we conducted a set of experiments on a simulated musculoskeletal system using pairs of agonist and antagonist muscles. The reflex connectivity is obtained by producing spontaneous motor activity in each muscle and by correlating the resulting sensor and motor signals. Our results show that, under biologically plausible conditions, the reflex circuitry thus obtained is consistent with that identified in relation to the analogous mammalian reflexes. In addition, they show that the reflex connectivity obtained depends on the morphology of the musculoskeletal system as well as on the environment that it is embedded in.     

Indicators used in livestock to assess unconsciousness after stunning: a review

Assessing unconsciousness is important to safeguard animal welfare shortly after stunning at the slaughter plant. Indicators that can be visually evaluated are most often used when assessing unconsciousness, as they can be easily applied in slaughter plants. These indicators include reflexes originating from the brain stem (e.g. eye reflexes) or from the spinal cord (e.g. pedal reflex) and behavioural indicators such as loss of posture, vocalisations and rhythmic breathing. When physically stunning an animal, for example, captive bolt, most important indicators looked at are posture, righting reflex, rhythmic breathing and the corneal or palpebral reflex that should all be absent if the animal is unconscious. Spinal reflexes are difficult as a measure of unconsciousness with this type of stunning, as they may occur more vigorous. For stunning methods that do not physically destroy the brain, for example, electrical and gas stunning, most important indicators looked at are posture, righting reflex, natural blinking response, rhythmic breathing, vocalisations and focused eye movement that should all be absent if the animal is unconscious. Brain stem reflexes such as the cornea reflex are difficult as measures of unconsciousness in electrically stunned animals, as they may reflect residual brain stem activity and not necessarily consciousness. Under commercial conditions, none of the indicators mentioned above should be used as a single indicator to determine unconsciousness after stunning. Multiple indicators should be used to determine unconsciousness and sufficient time should be left for the animal to die following exsanguination before starting invasive dressing procedures such as scalding or skinning. The recording and subsequent assessment of brain activity, as presented in an electroencephalogram (EEG), is considered the most objective way to assess unconsciousness compared with reflexes and behavioural indicators, but is only applied in experimental set-ups. Studies performed in an experimental set-up have often looked at either the EEG or reflexes and behavioural indicators and there is a scarcity of studies that correlate these different readout parameters. It is recommended to study these correlations in more detail to investigate the validity of reflexes and behavioural indicators and to accurately determine the point in time at which the animal loses consciousness.     

Postnatal Temporal,Spatial and Modality Tuning of Nociceptive Cutaneous Flexion Reflexes in Human Infants

Cutaneous flexion reflexes are amongst the first behavioural responses to develop and are essential for the protection and survival of the newborn organism. Despite this, there has been no detailed, quantitative study of their maturation in human neonates. Here we use surface electromyographic (EMG) recording of biceps femoris activity in preterm (<37 weeks gestation, GA) and term (≥37 weeks GA) human infants, less than 14 days old, in response to tactile, punctate and clinically required skin-breaking lance stimulation of the heel. We show that all infants display a robust and long duration flexion reflex (>4 seconds) to a single noxious skin lance which decreases significantly with gestational age. This reflex is not restricted to the stimulated limb: heel lance evokes equal ipsilateral and contralateral reflexes in preterm and term infants. We further show that infant flexion withdrawal reflexes are not always nociceptive specific: in 29% of preterm infants, tactile stimulation evokes EMG activity that is indistinguishable from noxious stimulation. In 40% of term infants, tactile responses are also present but significantly smaller than nociceptive reflexes. Infant flexion reflexes are also evoked by application of calibrated punctate von Frey hairs (vFh), 0.8–17.2 g, to the heel. Von Frey hair thresholds increase significantly with gestational age and the magnitude of vFh evoked reflexes are significantly greater in preterm than term infants. Furthermore flexion reflexes in both groups are sensitized by repeated vFh stimulation. Thus human infant flexion reflexes differ in temporal, modality and spatial characteristics from those in adults. Reflex magnitude and tactile sensitivity decreases and nociceptive specificity and spatial organisation increases with gestational age. Strong, relatively non-specific, reflex sensitivity in early life may be important for driving postnatal activity dependent maturation of targeted spinal cord sensory circuits.