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A R Light

Publications and source records attributed to A R Light.

51 records · Page 3Linked to original sources

The ultrastructure of group Ia afferent fiber synapses in the lumbosacral spinal cord of the cat.

Ia synapses in laminae VI and IX of the cat's spinal cord were examined in the electron microscope following iontophoretic injection of horseradish peroxidase (HRP) into single, identified, Ia afferent fibers from gastrocnemius muscles. Ia boutons contacting motoneuron dendrites in lamina IX contained spherical synaptic vesicles and generally contacted only one postsynaptic profile. The Ia boutons were often postsynaptic to smaller P-type axonal terminals. Consequently Ia boutons may be classified as S-boutons with axo-axonic contacts.

Animals↗

Morphology and synaptic relationships of physiologically identified low-threshold dorsal root axons stained with intra-axonal horseradish peroxidase in the cat and monkey.

The arborizations and synaptic relationships of intra-axonally stained horseradish peroxidase- (HRP) labeled primary afferent fibers to the dorsal horn of the cat and monkey spinal cord have been studied by light and electron microscopic methods. The light microscopic arborizations of the afferent fiber types (hair follicle afferents, pacinian corpuscle afferents, type I and type II slowly adapting afferents) are similar to those described by Brown and his colleagues (1) in the cat. The synaptic profiles formed by labeled afferents contain rounded synaptic vesicles. In serial thin sections, it was found that single dorsal root axons may make hundreds or thousands of synapses with neuronal structures of the dorsal horn. The vast majority of synaptic contacts are on the dendritic trees of dorsal horn neurons. The synapses made by these low-threshold afferent axons are almost all in the deeper laminae (III-VI) of the dorsal horn. The hair follicle afferent axons and the pacinian corpuscle afferents have numerous vesicle-containing structures that synapse on them to form either axoaxonal synapses or dendroaxonal synapses. The slowly adapting afferent axons are less often found to be postsynaptic to axons or dendrites. It is concluded that different physiological classes of primary afferent axons have different morphological characteristics, both at the light and electron microscopic level.

Afferent Pathways↗

Neurons with unusual response and receptive-field properties in upper laminae of cat SI cortex.

High-impedance micropipettes are used to record (both extra- and intracellularly) the electrical activity of neural elements located 550 micron or less from the pial surface of cerebral cortical areas 3a, 3b, 1, and 2 in unanesthetized cats. These elements are designated as "upper-layer SI units" and most frequently are sampled within the arm and forelimb digit sectors of areas 3b and 1. Mechanical stimulation of the skin is employed to determine the receptive field (RF) and response properties of the upper-layer units sampled. Single-shock electrical stimulation of the skin is used to obtain estimates of the minimal latency for eliciting spike discharge. Intracellular iontophoretic injection of horseradish peroxidase (HRP) is used to determine the laminar locations of the somata of the neural elements from which recordings are obtained. The receptive field (RF) and response properties of most upper-layer units sampled in areas 3b and 1 differ substantially from those of units recorded at depths greater than 550 micron from the pial surface in the same cortical fields. The members of one group of upper-layer area 3b and 1 units (U units) respond best to infrequently repeated (typically less than 0.5/s), slowly moving (1-5 cm/s) tactile stimuli. For the same units, repetitive application of slow-moving tactile stimuli to the RF typically leads to an enhancement of responsiveness accompanied by an elevation of spontaneous activity. In contrast, repetitive stimuli delivered at high velocity and at short interstimulus intervals lead to a decrease in unit responsiveness and to an absence of spontaneous activity. The members of a second group of upper-layer units (R units) respond best to moving stimuli delivered at higher velocities (5-20 cm/s), respond reliably at stimulus repetition rates well in excess of 0.5/s, and do not exhibit pronounced changes in responsiveness to repeated stimulation. The RFs of most upper-layer units (both R and U units) involve restricted regions on the contralateral upper limb, but the RFs of U units have poorly defined borders. In addition, the distribution of sensitivity within the RF of at least some U units is nonuniform and, frequently, discontinuous. Contralateral as well as ipsilateral body regions are included within the RFs for 12% of the upper-layer neurons sampled; the remainder (8%) have RFs restricted to the contralateral body.(ABSTRACT TRUNCATED AT 400 WORDS)

Animals↗

Features of laminar and somatotopic organization of lumbar spinal cord units receiving cutaneous inputs from hindlimb receptive fields.

Single-unit recordings from 312 units of lamina I-VII of the lumbar spinal cord of unanesthetized, decerebrate, T8 spinal cats were used to determine the somatotopic and laminar organization of spinal neurons responding to cutaneous stimulation of the hindlimb. Properties of cells confined to different Rexed laminae (I-VII) were shown to differ in several respects, including responses to variations in stimulus intensity, receptive-field areas, spontaneous frequencies, and central delays. Spinal cord neurons with similarly localized cutaneous receptive fields were found to be organized in sagittally oriented rectangular columns. These columns were 7 to at least 20 mm long (rostral-caudal axis), 0.5-1.0 mm wide, and could encompass laminae I-VII in depth. Touch, pressure, and pinch were effective excitatory inputs into each column subserving a given receptive-field location. A map of the somatotopic organization of units in the horizontal plane is presented, which in general confirms previous reports and in particular deals with the organization of units with receptive fields on the plantar cushion and individual toes.

Afferent Pathways↗

The ultrastructure and synaptic connections of serotonin-immunoreactive terminals in spinal laminae I and II.

In order to study the synaptic relationships of serotonin (5-HT)-containing axons, boutons in laminae I and II of the cat spinal cord were labeled for serotonin with peroxidase-antiperoxidase immunocytochemistry. Labeled boutons were examined with the light microscope and recut into serial ultrathin sections for examination with the electron microscope. Labeled axons exhibiting boutons were sagittally oriented, and were most numerous in lamina I and outer lamina II (IIo) and least numerous in inner lamina II (IIi). Two types of labeled boutons were observed ultrastructurally. A relatively rare, large, scalloped or egg-shaped bouton, which contained many mitochondria and dense core vesicles, was found in laminae I and IIo. A smaller dome-shaped bouton, which contained fewer dense core vesicles and round or pleomorphic, clear vesicles, was found throughout laminae I and II. Both types commonly established symmetrical synaptic contacts with the distal portion of a dendritic tree, rarely with proximal portions or cell somas, and never with axon terminals. The results suggest that there are heterogeneous serotonergic systems that may selectively modify different inputs postsynaptically to functionally different types of neurons in the superficial dorsal horn of the spinal cord.

Animals↗

Synaptic complexes formed by functionally defined primary afferent units with fine myelinated fibers.

The individual fine myelinated fibers of cutaneous mechanical nociceptors and "D-hair" receptors were identified by electrophysiological recording with micropipette electrodes in cats and monkeys. Their intraspinal terminations were labeled by iontophoresing horseradish peroxidase intracellularly and subsequent diaminobenzidine histochemistry. These terminations were examined with light and electron microscopy to determine the nature and organization of their synaptic contacts. Myelinated fibers of the mechanical nociceptors became unmyelinated before exhibiting many enlargements that made multiple synaptic contacts in the marginal zone (lamina I) and lamina V. Pre- or postsynaptic contacts were found only on enlargements. In the marginal zone of the cat, enlargements made simple axodendritic contacts or were scalloped, central terminals in glomeruli. In glomeruli, myelinated mechanical nociceptor enlargements were presynaptic to several dendritic appendages and postsynaptic to two different types of profiles. One type was interpreted as a presynaptic axon terminal, the other as a presynaptic, vesicle-containing, dendritic appendage. In lamina V of the cat the nociceptor synaptic complexes were similar, but simpler, and only axonal profiles were found to be presynaptic to them. In the monkey marginal zone and deep nucleus proprius, myelinated nociceptor terminations formed the central element of glomeruli, which consisted of postsynaptic dendritic appendages and presynaptic axon terminals. D-hair axons terminated in large numbers of enlargements in the nucleus proprius (laminae III and IV) and inner substantia gelatinosa (lamina IIi). Their large rounded enlargements formed the central terminals in glomeruli and were presynaptic to both ordinary and vesicle-containing dendritic appendages; the presynaptic dendritic profiles also often contacted each other. Profiles interpreted as axonal in origin were the only terminals presynaptic to the primary ending within the D-hair glomeruli. The results suggest that transfer of primary afferent information occurs only at enlargements of the primary fiber and that each primary fiber enters into more than one kind of synaptic arrangement. They also point out that synaptic glomeruli are common to functionally different types of primary afferent fibers and that the internal organization of glomeruli varies with the kind of primary fiber and the locus of the complex.

Afferent Pathways↗

A high voltage electrometer for recording and iontophoresis with fine-tipped, high resistance microelectrodes.

A high voltage electrometer is described which incorporates a controlled current source, direct current monitoring, balance bridge, electrode resistance, and capacitance compensation test circuits. This device is suitable for making biophysical measurements and iontophoresing dyes or enzymes through extremely fine micropipettes with impedances an order of magnitude higher than conventional micropipettes. Such electrodes are useful for recording from the small neurons of the central nervous system in vivo.

Animals↗

Reexamination of the dorsal root projection to the spinal dorsal horn including observations on the differential termination of coarse and fine fibers.

Primary afferent fibers in the lumbar, sacral, and caudal spinal segments of several mammals (rat, cat, monkey) were stained by applying horseradish peroxidase to the proximal part of cut dorsal rootlets and reacting the tissue histochemically after several hours of survival. The stained fibers' pattern of termination in the dorsal horn was similar in all three species, with many bouton-like enlargements in the ipsilateral marginal zone, substantia gelatinosa, and nucleus proprius, as well as a few projections at each level to the dorsal commissure and contralaterally to the ventral border of the nucleus proprius. Partial lesions of dorsal rootlets in monkey revealed that the thin fibers comprising the lateral division end principally in the marginal zone and substantial gelatinosa, while the thick fibers of the medial division terminate in the nucleus proprius and deeper regions, contributing little to the substantia gelatinosa and marginal zone. On the basis of the termination patterns observed for whole and partly sectioned rootlets, the superficial dorsal horn can be divided into at least four regions. (1) The marginal zone (lamina I of cat) appears to receive terminations from intermediate (smaller myelinated) fibers; (2) the outer substantia gelatinosa (outer lamina II) receives many terminations from the very finest afferent fibers; (3) the inner substantia gelatinosa (inner lamina II) receives endings from some of the finest fibers and also from intermediate (smaller myelinated) fibers; and (4) the superficial part of the nucleus proprius (lamina III) receives endings from intermediate and large diameter dorsal root fibers.

Animals↗

Spinal termination of functionally identified primary afferent neurons with slowly conducting myelinated fibers.

Single primary afferent myelinated fibers from cutaneous receptors of cat and monkey were functionally identified by recording from the spinal cord with micropipettes filled with horseradish peroxidase (HRP). Relatively slowly conducting fibers (less than 40 m/sec) from high threshold mechanoreceptors (mechanical nociceptors) and two types of low threshold mechanoreceptor (D-hair and field) were selected for staining. Iontophoresis of the HRP and subsequent histochemical reaction stained the axons recorded from and their collaterals, including terminations, for several millimeters. The termination patterns in the two species proved essentially identical. Ipsilaterally, the mechanical nociceptor fibers terminated principally in the dorsal horn's marginal zone and in the ventral parts of the nucleus proprius (lamina V in the cat). Some of these nociceptors also had terminals in the midline just dorsal to the central canal, contralaterally in the marginal zone, and at the base of the opposite nucleus proprius. In contrast, the D-hair primary afferent axons terminated in the dorsal part of the nucleus proprius overlapping into the innermost portion of the substantia gelatinosa. The field receptor fibers terminated predominantly in the middle part of the nucleus proprius. These results suggest that there is a highly specialized central projection of primary afferent endings which is related to sensory function and not to fiber diameter. The marginal zone and most dorsal parts of the substantia gelatinosa receive direct projections from cutaneous nociceptors but do not have direct input from cutaneous receptors transmitting activity initiated by innocuous stimulation.

Animals↗

Morphological features of functionally defined neurons in the marginal zone and substantia gelatinosa of the spinal dorsal horn.

Functional characteristics of spinal neurons located in the marginal zone (lamina I) and substantia gelatinosa (lamina II) were compared to their structural features by intrcellularly staining the source of unitary potentials with horseradish peroxidase (HRP) in unanesthetized, spinal cats. The responses of postsynaptic units to graded electrical volleys in intact dorsal roots and to physiological stimulation revealed that the peripheral excitatory input to neurons of the region is dominated by slowly conducting afferent fibers; often, the input to a given element is largely from a particular class of receptors. One type commonly seen received its principal peripheral excitation from low threshold mechanoreceptors with A delta or C afferent fibers. Mechanoreceptive elements often exhibited a marked, prolonged habituation and many were not excited by afferent volleys. Other units were predominantly excited by nociceptors with myelinated or unmyelinated fibers, or by thermoreceptors with unmyelinated fibers. A few units (principally the thermoreceptive) showed substantial ongoing activity which was modulated by sensory stimulation, but most had little or none. The HRP staining revealed neuronal morphology in fine detail. No relationship between neuronal configuration and physiological response was discerned. Soma location was not always linked to afferent input, although the cell bodies of nociceptive and thermoreceptive neurons tended to be in lamina I or outer lamina II (SGo) while those of the innocuous mechanoreceptive meurons tended to be in inner lamina II (SGi). The locus of a neuron's major dendritic arborization was more closely related to the source(s) of peripheral excitation. Cells excited by nociceptors with myelinated fibers had major dendritic projections in the marginal zone. Cells excited by nociceptors or thermoreceptors with unmyelinated fibers had important dendritic branching in the SGo. Innocuous mechanoreceptive neurons had primary dendritic arborization in the SGi when the input derived from unmyelinated fibers, or in the SGi and extending into the outer nucleus proprius (lamina III) when the afferent drive came from A delta fibers. These findings support the concept that laminae I and II constitute a major termination region for thin primary afferent fibers, myelinated fibers from nociceptors ending principally in lamina I and unmyelinated fibers from nociceptors, thermoreceptors, and mechanoreceptros terminating predominantly in lamina II. Substantial integrative and distributive functions can be expected of such an afferent termination zone.

Animals↗

The morphology of the spinal cord efferent and afferent neurons contributing to the ventral roots of the cat.

Horseradish peroxidase was applied to proximal ventral roots of the coccygeal and sacral spinal cord of cats. Subsequent histochemical reaction resulted in extensive staining of spinal cord neurons that had processes in the ventral roots. This procedure was used to study four issues concerning ventral root neurons. (1) Extensive transverse dendritic arborizations were revealed for large and small neurons presumed to be alpha and gamma motoneurons respectively. Dendrites from these neurons were found to project heavily into the ipsilateral white matter, both laterally and ventrally. Dendrites also projected extensively through the anterior commissure, attaining the contralateral grey and white matter. (2) Medially-located efferent neurons were found to contribute the contralateral dendrites as well as some dorsally-directed dendrites. Laterally-located neurons projected dendrites extensively into the lateral and ventral white matter. (3) Stained neurons were found in the intermediolateral cell column, and were presumed to be preganglionic efferent neurons. Some of these neurons projected dendrites into the marginal zone of the dorsal horn, while others sent dendrites medially toward the central canal. (4) Stained fibers, presumed to be primary afferents, were found to enter from the ventral roots and course to the dorsal horn. Most of these fibers were small in diameter and distributed boutons predominantly to the substantia gelatinosa. A few large ventral root afferent fibers were observed that distributed boutons mostly to the nucleus proprius.

Animals↗

Estrous cycle modulation of nociceptive behaviors elicited by electrical stimulation and formalin.

The impact of circulating ovarian hormones on nociceptive behaviors elicited by phasic and tonic stimuli was evaluated in rats using two behavioral tests: an operant escape task and the formalin test. The operant escape task was structured to separately evaluate hindlimb flexion reflexes, the latency of escape, and the amplitude of peak vocalization to a series of phasic electrocutaneous stimuli (0.05-0.8 mA), whereas the formalin test evaluated nociceptive behaviors elicited by tonic stimulation following a subcutaneous injection of dilute formalin (1%). Hindlimb reflex amplitude, escape latency, and peak vocalization varied across the estrous cycle, such that rats were most sensitive to electrical stimuli during proestrus (reflex and escape latency) and diestrus (vocalization). Furthermore, morphine-induced (3 mg/kg sc) attenuation of hindlimb reflex amplitude was sensitive to estrous cycling. During proestrus, morphine produced less attenuation of hindlimb reflex amplitude than during nonproestrus phases. However, estrous cycling did not alter nociceptive behaviors elicited by 1% formalin. These data support the notion that circulating ovarian hormones may differentially modulate behaviors associated with phasic and tonic pain.

Analgesics, Opioid↗