Coding of vibration by neurones of the dorsal column nuclei in the pigeon

Whereas the function of mechanoreceptors is well known in birds there are no detailed investigations of central processing. Therefore the characteristics of neurones in a first relay, the dorsal column nuclei (nuclei gracilis et cuneatus and the nucleus cuneatus externus), were studied electrophysiologically by recording responses from individual neurones to mechanical stimulation of skin and deep tissue.Of 140 units 84 were cutaneous neurones. The remainder were tap- and deep-neurones. Tap-neurones were activated by sharp brief taps. Activity in deep-neurones was comparable to joint and tendon receptors but not to muscle spindles. Thirty-four cutaneous neurones were vibration sensitive neurones, 36 were slowly adapting neurones and 14 neurones were rapidly adapting and fast habituating cells. Vibration sensitive neurones were most sensitive to 200 to 500 Hz. Frequencies above 100 Hz elicited phase coupled responses.Within the nn. gracilis et cuneatus 61 were cutaneous neurones, 13 were deep-neurones and 7 were tap-cells. Within n. cuneatus externus 17 were cutaneous units, 21 were deep-neurones and 11 were tap-neurones. No vibration sensitive neurones were found within n. cuneatus externus. This means that coding for vibration is represented in the nn. gracilis et cuneatus.Abbreviations CE n. cuneatus externus - DCN dorsal column nuclei - GC n. gracilis et cuneatus - HC Herbst corpuscle - PH phasic and habituating - Imps./s Impulses per second - INTH interval histogram - PSTH peristimulus time histogram - PHASE Phase histogram - RF receptive field - SA slowly adapting - Vec. vector strength     

Response characteristics of Herbst corpuscles in the interosseous region of the pigeon's hind limb

The Herbst corpuscle, found only in birds, is one of the morphologically distinct types of lamellated sensory nerve endings. Its response properties were studied electrophysiologically in anesthetized pigeons by recording from: (A) afferents emerging from single Herbst corpuscles located in the interosseous region of the shank, (B) from vibration-sensitive cells found in the dorsal root ganglia near the lumbosacral enlargement of the spinal cord, and (C) from nerve fibers dissected out of the sciatic nerve. Vibration-sensitive cells in the dorsal root ganglia and in sciatic nerve fibers exhibited properties that were very similar to those found at the Herbst corpuscle itself. All three recording approaches indicated that Herbst corpuscles are vibration-sensitive mechano-receptors with broad bandpass tuning curves. With all approaches, the lowest threshold always fell in the frequency range between 400 and 800 Hz. The value of the threshold, however, varied with the method, being as low as 0.08 m with method C and as high as 1 m with method A. Regardless of method, these neurons exhibited no spontaneous activity, and their firing displayed a special sensory coding pattern at high stimulus amplitude: the nerve impulses were phase locked to the stimulus cycle, exhibiting a 1:1 relationship with it up to a frequency of 500 Hz. Comparison of these data with prior behavioral data suggests that the Herbst corpuscle peripheral mechanoreceptors are part of a vibratory sensory system which acts as a warning device, given attentive behavior.Abbreviations AP action potential - INTH interspike interval histogram     

家鸽Herbst小体对振动的反应特征

鸟类的Herbst小体是一种形态特殊的感觉性神经末梢器官.本文利用电生理学方法,研究了家鸽腿部胫骨-腓骨之间的Herbst小体对振动刺激的反应特征.这种小体对振动刺激非常敏感,当振动频率在600—800赫时,它们有反应的最低阈值约为0.3微米.不同的Herbst小体的反应阈值与频率的关系曲线表明:这种小体具有明显的带通滤波的特征,对振动反应的最佳频率范围为400—1000赫.在适宜频率、超阈值强度的振动刺激下,Herbst小体能以1:1的方式作出反应,即相对于每次正弦波振动刺激都有一个锁相的神经脉冲产生.在背根脊神经节内的细胞外记录表明:对振动敏感的神经节细胞具有和Herbst小体完全相似的反应特征.     

Cutaneous sensory afferents recorded from the nervus intramandibularis ofGallus gallus vardomesticus

Summary The responses of single sensory afferent nerve fibres were recorded from small nerve bundles of the intramandibular nerve of the chicken following thermal and mechanical stimulation of the beak. Thermoreceptors, nociceptors and mechanoreceptors were identified and their responses characterized.Of the thermoreceptors identified 11 units were classified as cold receptors, which responded to cooling the receptive field by increasing the discharge rate and had conduction velocities in the range 0.83 to 4.4 m/s. Only one warm unit was identified.Two classes of nociceptors were identified: mechano-thermal (polymodal) nociceptors and high threshold mechanical nociceptors. The discharge characteristics and stimulus-response curves of both types were described. While the mechanothermal nociceptors were exclusively C-fibres (c.v. 0.4 to 1.86 m/s), the high threshold mechanoreceptors contained both C and A delta fibres (c.v. 1 to 5.5 m/s). Thermal response thresholds for the mechano-thermal units ranged from 41 to 50 °C with mechanical thresholds of 2 to over 50 g. Mechanical thresholds for the high threshold units ranged from 5 to over 50 g.The mechanoreceptors were either slowly or rapidly adapting. The pattern of response together with stimulus-response curves were presented for the slowly adapting units. Conduction velocities of the slowly adapting units varied from 0.7 to 20 m/s and mechanical threshold from 0.1 to 2 g. On the basis of their response to a vibrating, and a ramp-and-hold mechanical stimulus, the rapidly adapting units were divided into Herbst and Grandry units with only the Herbst units responding accurately to the vibrating stimulus. Both units had fibres conducting in the 50 m/s range with thresholds in the 0.1 to 10 g range.The results are discussed in relation to the receptors found in other avian species and mammalian peripheral sensory afferents.Abbreviations c.v. conduction velocity - RA rapidly adapting (receptors) - SA slowly adapting (receptors)     

Sensory Innervation of the Raccoon Forepaw: 1. Receptor Types in Glabrous and Hairy Skin and Deep Tissue

Electrophysiological recordings were made from the median, ulnar, radial, and dorsal ulnar nerves to determine the types of mechanosensory receptors serving glabrous and hairy skin surfaces of the raccoon forepaw. In addition to the cutaneous mechanoreceptors, fibers innervating deep tissues were also recorded from each of these nerves. These included sensory fibers innervating muscles, joints, claws, and the subcutaneous pulp.

The array of receptors serving raccoon glabrous skin was the same as found in monkeys and humans: Rapidly adapting (RA), slowly adapting (SA), and Pacinian (Pc) fibers were characterized. Pacinian fibers have been rarely described in previous physiological studies of the raccoon peripheral nerves, but in the present study they composed between 14% and 18% of the glabrous skin mechanoreceptors recorded. A distal-proximal gradient in the density of skin innervation was evident for all three types of receptors.

Receptors characterized in the hairy skin of the dorsal paw were similar to those described in other mammals, and included both down and guard hair afferents, non-hair-associated RA fibers, and SA I and SA II fibers. The relative proportions of these fibers differed from those generally reported for the hairy skin of other mammals. SA hair-associated afferent fibers, which have been reported previously only in primate hairy skin, were also found in large numbers in the raccoon. Similarities and differences in the frequency and types of receptors innervating the raccoon forepaw, the forepaws of other mammals, and the hands of primates (including humans) are discussed.     

Vagal afferent and reflex responses to changes in surface osmolarity in lower airways of dogs.

In anesthetized dogs we examined the sensitivity of afferent vagal endings in the lungs to changes in airway fluid osmolarity. Injection of 0.25-0.5 ml/kg water or hyperosmotic sodium chloride solutions (1,200-2,400 mmol/l) into a lobar bronchus caused bradycardia, arterial hypotension, apnea followed by rapid shallow breathing, and contraction of tracheal smooth muscle. All effects were abolished by vagotomy. We examined the sensory mechanisms initiating these effects by recording afferent vagal impulses arising from the lung lobe into which the liquids were injected. Water stimulated pulmonary and bronchial C-fibers and rapidly adapting receptors; isosmotic saline and glucose solutions were ineffective. Hyperosmotic saline (1,200-9,600 mmol/l, 0.25-1 ml/kg) stimulated these afferents in a concentration-dependent manner. Stimulation began 1-10 s after the injection and sometimes continued for several minutes. Responses of slowly adapting stretch receptors varied. Our results suggest that non-isosmotic fluid in the lower airways initiates defense reflexes by stimulating pulmonary and bronchial C-fibers and rapidly adapting receptors. Conceivably, stimulation of these afferents as a result of evaporative water loss from airway surface liquid could contribute to exercise-induced asthma.     

The three-dimensional microanatomy of Meissner corpuscles in monkey palmar skin

The Meissner corpuscle is a rapidly-adapting mechanoreceptor in the dermal papillae of digital skin. For an analysis of how the sensory endings detect tissue deformations, an examination of their fine structure and relationships with dermal collagen was carried out in the Japanese monkey, Macaca fuscata, using a combination of three methods: SEM of cell architecture denuded by 6N sodium hydroxide maceration, SEM of collagen networks exposed by a mild alkaline corrosion, and TEM according to a conventional procedure. Observations showed the sensory corpuscles to be represented by a stack of discoid components consisting of flattened axon terminals sandwiched between Schwann cell lamellae, as reported previously. Each corpuscle was entirely covered by a connective tissue capsule, which was linked with the basal aspect of the epidermis by dermal collagen fibers. Margins of the discoid components of the corpuscles were serrated with numerous fine projections of lamellar Schwann cells, which tightly held collagen trabeculae on the inner aspect of the pericorpuscular capsule. Central portions of the discoids, on the other hand, displayed extremely smooth surfaces, which were covered by a thick layer of basal lamina-like matrix. The former portions of the discoids appear susceptible to mechanical deformations of surrounding tissues, while the latter may follow the tissue movements rather slowly because of their indirect linkage with the dermal collagen network. The resulting distortions of the axon endings during dynamic phases of the tissue deformations will be in favor of the generation of rapidly adapting receptor potentials in the sensory corpuscle.     

Immunohistochemical study of cutaneous nerves in the emu

The distribution and chemical content of cutaneous nerves in 3- to 13-day-old emu chicks (Dromaius novaehollandiae) were examined by using double-labelling immunohistochemistry. Seven different subpopulations of cutaneous nerves were identified based on their neurochemistry. No intraepidermal nerve fibres were found. However, axons were located within the dermis and were often associated with blood vessels, pennamotor muscles and feather follicles or innervated Herbst corpuscles. Both similarities and differences exist between subpopulations of cutaneous nerves in the emu and volant birds. As in volant birds, a subpopulation of cutaneous axons innervates the superficial skin layers and contains immunoreactivity to both substance P and calcitonin gene-related peptide (CGRP). This suggests that the neuropeptide content of these presumptive free nerve endings is conserved throughout the evolution of birds. In contrast, Herbst corpuscles in the emu are innervated by axons that contain immunoreactivity for CGRP or neuropeptide Y (NPY) but that lack the calbindin D-28k immunoreactivity found in fibres innervating Herbst corpuscles of volant birds. Herbst corpuscles therefore may have a different chemical content in a flightless species from that in volant birds.This work was supported by an Australian Postgraduate Award (Industry; CO9942100) and the Flinders Institute for Health and Medical Research.     

Two types of P neurons in the medullary dorsal respiratory cellular group in cats

Electrophysiological properties of P neurons localized in the medullary dorsal respiratory cellular group and of vagal afferent fibers innervating these neurons were studied in acute experiments on nembutal-anesthetized cats with preserved spontaneous respiration. P neurons were shown to form a non-homogeneous cellular population. They generated phasic discharges during the whole inspiration period, but differed in their responses to lung inflation. These findings allowed us to classify P neurons as slowly adapting and rapidly adapting units, probably activated by slowly and rapidly adapting pulmonary receptors, respectively. Sensitivity of the slowly adapting P neurons to activation by the corresponding receptors and the mechanisms underlying the participation of the two types of P neurons in the reflex feedback between the respiratory center and lungs are discussed.Neirofiziologiya/Neurophysiology, Vol. 26, No. 3, pp. 211–217, May–June, 1994.     

Histological and electrophysiological investigations on the vibration-sensitive receptors (Herbst corpuscles) in the wing of the pigeon (Columba livia)

Summary The Herbst corpuscles (HCs) of the pigeon's wing were investigated both histologically and electrophysiologically. All HCs found in the wing were lamellated, basic type corpuscles without any specialized structures. Their lengths ranged from 67 to 853 m (mean = 310 m). Unexpected findings were their large number (about 1000 in the manual part of the wing), their irregular distribution and their preferred orientation (approximately parallel or at right angles to the primary feather follicles). The HCs were highly sensitive to vibrational stimuli applied to wing feathers. Their electrophysiological behaviour has the following characteristics: no spontaneous activity, phase-locked nerve impulses, a 11 stimulus-response relation up to at least 660 Hz at sufficiently high stimulus amplitudes, and a sensitivity to stimulus frequencies up to 1800 Hz. The best frequencies of 52 receptive units for which complete threshold curves were obtained lay between 100 and 900 Hz, 67% of the best frequencies were between 200 and 400 Hz. The threshold amplitudes at best frequency ranged from 0.5 to 150 m. Two virtually non-overlapping mechanosensitive areas on the wing were identified. One is a very narrow band along the frontal edge and the other covers a large area of the remaining wing. They correspond with the two branches of the radial nerve. The histological and electrophysiological findings suggest that the HCs are part of a vibrational sensory system that is principally involved in flight control. The actual aerodynamic state during flight could be detected by the frontal receptive area, and flight behaviour could be adjusted accordingly. The effectiveness of these corrective reactions would then be assessed on the basis of air current changes along the caudal wing edge by the caudal receptive area. Despite some physiological differences considerable similarities between HCs and Pacinian corpuscles support the hypothesis that they evolved from a unique ancestral lamellar receptor.Abbreviations HC(s) Herbst corpuscle(s) - PC(s) Pacinian corpuscle(s) All experiments were carried out at the Ruhr-Universität Bochum, Lehrstuhl für Allgemeine Zoologie     

Afferent neural pathways in cough and reflex bronchoconstriction

Cough and bronchoconstriction are airway reflexes that protect the lung from inspired noxious agents. These two reflexes can be evoked both from the larynx and tracheobronchial tree and also from some extrarespiratory sites. Within the airways, certain sites are particularly sensitive to stimulation of cough (larynx and points of proximal airway branching), whereas bronchoconstriction can be triggered from the whole of the tracheobronchial tree. In the larynx, "irritant" receptors with myelinated afferents mediate cough and bronchoconstriction. Little seems to be known about laryngeal nonmyelinated afferents and their reflexes. In the tracheobronchial tree and lung, slowly adapting stretch receptors (SARs) and rapidly adapting stretch receptors (RARs) have opposing effects on airway tone, the former mediating bronchodilation and the latter bronchoconstriction. In cough, on the other hand, they operate concurrently, a mediatory role for RARs and a facilitatory role for SARs. C-fiber endings (bronchial and pulmonary) mediate bronchoconstriction. Inhalation of so-called "selective" C-fiber stimulants induces cough, but excitation of RARs has not been eliminated, and the possibility also exists that the cough is secondary to other lung actions mediated by these nerve endings. Although cough and bronchoconstriction may be mediated by the same type of receptor, they seem to have separate afferent neural pathways.     

Stretch receptors of the lungs in the frog Rana temporaria

In experiments on isolated lungs, studies have been made on the activity of single afferent fibers during simulated expiration and inspiration. Both slowly adapting and rapidly adapting voluminous stretch receptors were found in the frog lungs. The latter type exhibits the activity also during fast simulated expiration. Using acetylcholine and histamine, it was shown that the level of excitability of the receptors investigated depends on functional condition of the plain muscles in the pulmonary wall.     

The Effects of Aging on Information-Processing Channels in the Sense of Touch: I. Absolute Sensitivity

Thresholds for detecting vibrotactile signals of variable frequency applied to the thenar eminence of the hand by small and large contactors were measured in subjects ranging in age from 10 to 89 years. Thresholds were found to increase as a function of age, but the rate of increase was greater after than before the age of 65 years. The rate of loss of vibrotactile sensitivity was substantially greater in the P channel (mediated by Pacinian corpuscles) than in the NP I channel (mediated by rapidly adapting fibers), the NP II channel (mediated by slowly adapting type II fibers), or the NP HI channel (mediated by slowly adapting type I fibers). Women were frequently found to have greater sensitivity than men.     

Intrapulmonary receptors in the bullfrog: Sensitivity to CO2

Summary Single-unit vagal afferent recordings were made on 55 intrapulmonary receptors in 15 anesthetized or decerebrate bullfrogs. Intrapulmonary CO₂ concentration and intrapulmonary pressure were controlled independently by unidirectionally ventilating the lungs. No CO₂ receptors (insensitive to stretch of the lung) of the kind reported in birds and reptiles were found; all 55 receptors were mechano-sensitive. Of these mechanoreceptors, 39 adapted slowly to inflation of the lung and 16 adapted rapidly. Thirtythree of the slowly-adapting receptors and 15 of the rapidly adapting receptors decreased their discharge frequency as intrapulmonary CO₂ concentration was increased. Inflating the lung enhanced CO₂ sensitivity. The results indicate that the frog possesses CO₂-sensitive pulmonary mechanoreceptors similar to those of mammals and reptiles.Abbreviations P _ip intrapulmonary pressure - fractional inspired concentration of CO₂ The authors wish to thank Dalyn Wilson for his help in gathering the experimental animals and for his technical assistance. The study was supported in part by a grant-in-aid from the American Heart Association, Kansas Affiliate, Inc. Contribution No. 78-185-J Department of Anatomy and Physiology, KAES, Kansas State University, Manhattan, Kansas 66506, USA.     

Two-dimensional receptive fields of midbrain lateral line units in the goldfish, <Emphasis Type="Italic">Carassius auratus</Emphasis>

We determined the receptive fields of midbrain lateral line units in goldfish, Carassius auratus, with a 50 Hz vibrating sphere placed at various azimuths and elevations alongside the fish and studied how responses were affected by different directions of sphere vibration. The receptive fields of toral lateral line units, in contrast to those of primary afferent nerve fibers, did not represent the pressure gradient pattern generated by a vibrating sphere. Thus, unlike primary afferents, single toral lateral line units did not code for source location in their spatial discharge patterns. The two-dimensional receptive fields were round, horizontally or vertically stretched, or complex. While some toral lateral line units were sensitive to the direction of sphere vibration others were not.     

Hairy Skin: Psychophysical Channels and Their Physiological Substrates

Experiments were conducted in which threshold-frequency characteristics were measured on the hairy skin of the forearm of human observers. Thresholds were measured with two stimulus probe areas (2.9 and 0.008 cm²) at three skin-surface temperatures (15d`, 30d`, and 40d`C). The results suggest that whereas glabrous skin uses four distinct channels of information, only three channels may be involved in mediating the sense of touch for hairy skin. The three channels are defined as P_h (Pacinian, hairy skin), NP_{h low} (non-Pacinian, hairy skin, low frequencies) and NP_{h mid} (non-Pacinian, hairy skin, middle frequencies). In addition, it is proposed that the neural substrates for the three psychophysically characterized channels are, respectively, the Pacinian corpuscle (PC) nerve fibers, the slowly adapting type II (SAII) fibers, and the rapidly adapting (RA) fibers.     

Activité cholinestérasique des corpuscules sensoriels cutanés de Herbst et de Grandry

Résumé L'activité cholinestérasique (ChE) a été étudiée dans les corpuscules de Herbst et de Grandry du bec de canard par la méthode de Karnovsky. Le BW 284 C 51, l'iso-OMPA et l'ésérine ont été utilisés comme inhibiteurs. Une activité cholinestérasique non spécifique (nsChE) a été identifiée dans les corpuscules. Elle est présente dès le début de leur différenciation.Corpuscule de Herbst: l'activité nsChE est localisée dans les cellules du bulbe interne (citernes périnucléaires, reticulum endoplasmique granulaire, vésicules associées à l'apppareil de Golgi) et à la surface de la membrane plaamique des cellules du bulbe interne et de la terminaison nerveuse.Corpuscule de Grandry: l'activité nsChE est localisée dans les cellules satellites (citernes périnucléaires, reticulum endoplasmique granulaire) et à la surface de la membrane plasmique des cellules satellites, des cellules de Grandry et de la terminaison nerveuse.Une forte activité nsChE a été mise en évidence dans certaines zones de contact entre les cellules dites sensorielles (cellules du bulbe interne et de Grandry) et les terminaisons nerveuses. Une réaction positive a été observée au niveau des mitochondries des cellules satellites et de la terminaison nerveuse du corpuscule de Grandry, et dans des vésicules du réticulum endoplasmique lisse des terminaisons nerveuses.Aucune activité ChE n'a été détectée dans les cellules capsulaires, les cellules de l'espace interne, les cellules de Grandry, et dans les vésicules synaptiques.Ces résultats ont été discutés en relation avec ce que l'on sait sur l'activité ChE des autres types de corpuseules sensoriels. La signification fonctionnelle de cette activité ChE et le problème de l'origine des différentes catégorics cellulaires qui constituent les corpuscules de Herbst et de Grandry ont été envisagés.

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Cholinesterase activity in the Herbst and Grandry cutaneous sensory corpusclesHistochemical study at the light and electron microscope

Summary Cholinesterase activity (ChE) was localized in the Herbst and Grandry cutaneous sensory corpuscle of the duck beak with Karnovsky's method. Controls with BW 284 C 51, iso-OMPA and eserine were carried out. A non-specific cholinesterase activity (nsChE) was identified in the corpuscles. This enzyme activity is present from the beginning of their differenciation.Herbst corpuscle: nsChE activity was localized in the inner bulb cells (perinuclear cisterna, granular endoplasmic reticulum, vesicles associated with the Golgi complex) and along the plasma membrane of inner bulb cells and nerve ending.Grandry corpuscle: nsChE activity was localized in the satellite cells (perinuclear cisterna, granular endoplasmic reticulum) and along the plasma membrane of satellite cells, Grandry cells, and nerve ending.A high nsChE activity was evident in certain zones of contact between the so-called sensory cells (inner bulb and Grandry cells) and the nerve endings. A positive reaction was seen in mitochondria of satellite cells and nerve ending for the Grandry corpuscle, and in smooth endoplasmic vesicles of the nerve endings.No ChE activity was detected in capsular cells, inner space cells, Grandry cells and in synaptic vesicles.These results are discussed in relation with what is known about ChE activity of other sensory corpuscles. The functional significance of this ChE activity and the problem of the origin of the different cells of Herbst and Grandry corpuscles have been considered.

     

Cutaneous sensitivity to touch and low-frequency vibration in selachians

In 46 isolated and 4in situ preparations of perioral skin, afferent impulse patterns, leading from cutaneous mechanoreceptors and responding to orthogonal square as well as sine-wave stimuli, were investigated in selachians (Scyliorhinus canicula).

Functional Properties of Slowly Adapting Mechanoreceptors in Cat Footpad Skin

The functional properties of slowly adapting (SA) afferent fibers innervating cat footpad skin were examined. Measurements were taken of receptive field area; spontaneous activity (< 1 impulse/sec); the slope of the stimulus-response curve for steady indentations up to 2 mm in amplitude; variability of the interimpulse intervals, as measured by the coefficient of variation of time interval histograms; decay of the response to steady indentation; and sensitivity to sinusoidal vibration (most sensitive at 5-10 Hz). Where comparable tests were performed on glabrous and hairy skin SA fibers, the functional properties of those in glabrous skin more closely resembled SAI fibers than SAII fibers. Additional results from glabrous skin SA fibers suggest that it is distortion of the nerve endings rather than steady indentation or compression that leads to a brisk response. On the measures described above, there appeared to be only one functional class of SA fiber innervating the cat footpad skin.     

Somatosensory activity relevant for motor output. A summary of presentations and discussion

Several aspects of the function of receptors which contribute to somatic sensations are reviewed. First, there is evidence for a role of large-diameter cutaneous afferents in the reflex regulation of precision movements by the hand. Second, large-diameter muscle afferents from the intrinsic muscles of the hand, probably from primary muscle spindle afferents, can evoke specific sensations of finger movement. Third, the variable relationship between discharges in human C fibers from the hand and the specific sensation of pain is investigated. Activity in large-diameter cutaneous afferents can probably modify this sensation. Finally, the properties of small-diameter afferent fibers innervating joints are shown to be consistent with a role in the reflex regulation of joint integrity.