The termination of the spinothalamic tract in the cat an experimental study with silver impregnation methods

Summary The termination of the spinothalamic tract (STT) in the cat has been studied light microscopically in Fink-Heimer and Nauta impregnated sections. Following lesions of the STT at various rostrocaudal levels of the spinal cord the degenerating fibres in the thalamus and subthalamus were mapped, mainly in transverse sections. The cervicothalamic tract was not injured by the lesions.The spinothalamic fibres enter the diencephalon through the mesencephalic reticular formation and terminate in the following regions: the medial portion of the magnocellular part of the medial geniculate body (MGmc), the ventrolateral portion of the medial part of the posterior nuclear complex (PO_m), the caudolateral and medial parts of the zona incerta (ZI), the nucleus centralis medialis (CeM), the nucleus parafascicularis (Pf), the lateral part of the nucleus centralis lateralis (CL), the medial and rostrolateral parts of the nucleus ventralis lateralis (VL). To reach these regions the fibres pass through the nucleus centrum medianum (CM), the nucleus subparafascicularis (SPf) and the nucleus paracentralis (Pc). The fibres that terminate in the VL pass through Forel's field H₁ and the external medullated lamina (EML). Conclusive results were not obtained concerning a termination in the CM. The spinothalamic fibres do not pass through nor terminate in the nucleus ventralis posterolateralis (VPL) and the nucleus reticularis (R). The VPL, defined as that portion of the ventral thalamus that receives terminal fibres from the dorsal column nuclei, has been found to extend rostrally only as far as Horsley-Clarke level anterior 10.5. The results strongly support the view that all the spinothalamic fibres terminate ipsilateral to their course in the ventral quadrant of the spinal cord. No signs of a somatotopical organization of the termination of the STT were found.List of Abbreviations Cd nucleus caudatus - CeM nucleus centralis medialis - CG circumaqueductal gray substance - CL nucleus centralis lateralis thalami - CM nucleus centrum medianum thalami - CP commissura posterior - CTT cervicothalamic tract - EML external medullated lamina - H₁ Forel's field H₁ - HP tractus habenulopeduncularis - LCN nucleus cervicalis lateralis - LG corpus geniculatum laterale - LP nucleus lateralis posterior thalami - MD nucleus medialis dorsalis thalami - MG corpus geniculatum mediale - MGmc corpus geniculatum mediale, pars magnocellularis - MGp corpus geniculatum mediale, pars principalis - ML medial lemniscus - MRF mesencephalic reticular formation - OT optic tract - Pc nucleus paracentralis thalami - Pf nucleus parafascicularis thalami - PO posterior group of thalamic nuclei - PO₁ lateral part of PO - PO_m medial part of PO - R nucleus reticularis thalami - SG nucleus suprageniculatus - STT spinothalamic tract - VA nucleus ventralis anterior - VL nucleus ventralis lateralis thalami - VM nucleus ventralis medialis thalami - VPI nucleus ventralis posterior inferior - VPL nucleus ventralis posterior lateralis thalami (VPL₁ + VPL_m) - VPL₁ lateral part of VPL - VPL_m medial part of VPL - VPM nucleus ventralis posterior medialis thalami - VPMpc parvocellular part of VPM - ZI zona incerta     

Somatotopic termination of the spino-olivary fibers in the cat,studied with the wheat germ agglutinin-horseradish peroxidase technique

Summary Terminal sites of the spino-olivary fibers (SOFs) were examined by the anterograde transport of wheat germ agglutinin-horseradish peroxidase in the cat. The tracer was injected at various spinal cord levels from the first cervical to the caudal segments. The SOFs derived from the C1-T1 segments terminated medially in the caudal half (levels II–VIII of Brodal) of the medial accessory olive (MAO), which projects to the A zone of the cerebellar cortex, whereas the SOFs derived from the L6-S1 segments terminated laterally in the caudal half (levels I–VIII) of the MAO. No projections were found from the T2-L5 segments to the MAO. In the dorsal accessory olive (DAO), the SOFs terminated at levels III–XIV; the DAO projects to the B zone and the C1 and C3 zones of the cerebellar cortex. The SOFs derived from the C1-C4 segments terminated in the most medial part of the DAO (levels III–XIV), followed laterally by those from the C5-T1 segments. Further laterally, the SOFs derived from the T2-L5 and the L6-S1 segments terminated in the mediolateral order at levels V–XIV. The SOFs from the L6-S1 segments occupied the most lateral part of the DAO. The present study demonstrates that there is a distinct somatotopic termination of the SOFs in the mediolateral order in the caudal MAO and the DAO.     

Thermosensory defects after cervical spinal cord lesions in the cat

Summary The behavioural thermosensitivity of cat paws was examined before and/or after restricted uni- and/or bilateral lesions had been made in the spinal cord between the first and fifth cervical segments. Unilateral lesions of the lateral funiculus, which involved at least its whole width at the level of the central canal, reproducibly were found to interfere with the contralateral sensitivity for temperature increases and/or decreases. No corresponding thermosensory deficiencies were found after unilateral lesions involving the ventral spinal quadrant or the dorsal funiculus. Various bilateral and combined lesions were made, but no cat ever developed thermoanaesthesia. The bilateral lesions included bilateral transections of: the middle parts of the lateral funiculi, the dorsal halves of the lateral funiculi, the dorsal funiculi, and the ventral spinal half.Most of our knowledge about peripheral behavioural thermosensitivity after spinal cord injury is based on observations of human patients, especially after anterolateral chordotomies. The present finding of contralateral thermosensory deficiencies after lesions of the middle part of the lateral funiculus fits with some of the clinical reports. The present failure to cause thermoanaesthesia, on the other hand, is inconsistent with the theory of a single ascending spinal pathway for behavioural thermo-sensitivity, which has emanated mainly from the clinical observations.     

Spinal projections to the cerebellar nuclei in the cat

Summary Projections from the spinal gray matter to the cerebellar nuclei in the cat have been studied using Nauta's silver technique. Following unilateral section of the ventrolateral cord at the cervical level, heavy degeneration is seen in the nucleus medialis on both sides. Scanty degeneration is present bilaterally in the nucleus interpositus. The degeneration is most intense on the contralateral side. Scanty degeneration is also present bilaterally in subnucleus medialis parvicellularis (SMP) (Flood and Jansen, 1961). No degeneration is seen in nucleus lateralis. Following unilateral section of the dorsolateral cord at the cervical level, scanty degeneration is present bilaterally in nucleus medialis and nucleus interpositus anterior. The degeneration is more pronounced ipsilaterally and is also seen in SMP on both sides. No degeneration is present in nucleus lateralis. Fibers from the ventral and dorsal spinocerebellar tracts (VSCT and DSCT) terminate bilaterally in nuclei medialis and interpositus, with the VSCT as the most important connection.     

The distribution of infrahyoid motoneurons in the cat: a retrograde horseradish peroxidase study

Summary The distribution of cell bodies and the peripheral course of axons of infrahyoid motoneurons were examined in the cat by the retrograde horseradish peroxidase method after application of the enzyme to the peripheral nerve branches supplying the infrahyoid muscles. Infrahyoid motoneurons were observed to constitute a slender cell column, which extended from a level of the caudal part of the hypoglossal nucleus usually to the most caudal level of the C1 cord segment, or occasionally to the lower levels of the C2 cord segment. The cell column was located immediately lateral to that of motoneurons of the spinal accessory nerve. In the cell column, thyrohyoid motoneurons were distributed in the medulla oblongata; sternohyoid motoneurons were located somewhat more cranially than sternothyroid motoneurons in the medulla oblongata and cervical cord. However, the level of craniocaudal distribution of thyrohyoid, sternohyoid or sternothyroid motoneurons highly overlapped. The experiments involving severance of the hypoglossal and/or cervical nerves indicated that axons of thyrohyoid and sternohyoid motoneurons passed via the roots of both hypoglossal and C1 nerves, that axons of sternohyoid motoneurons passed via the C1 nerve roots, and that axons of infrahyoid motoneurons innervating the conjugated part of the sternohyoid and sternothyroid muscles passed usually via the C1 nerve roots, or occasionally via the roots of both C1 and C2 nerves.     

The effects of baclofen on the stretch reflex parameters of the cat

Experiments were done in cats decerebrated at the precollicular postmammillary level to determine how a tonic increase of presynaptic inhibition of the intraspinal terminals of muscle spindle afferents changes the mechanical properties of the soleus stretch reflex (s.r.). Baclofen, a specific GABA_B receptor agonist, was injected i.v. (1–2 mg/kg) so as to induce a tonic increase in presynaptic inhibition. The effects of baclofen on the stiffness and threshold of the s.r. were determined, respectively, from plots of stiffness vs background force and force vs length (length-tension plot). Baclofen, at these doses, had no effect on the excitation-contraction coupling properties of muscle or on the intrinsic stiffness-force relation. Changes of the soleus background force, required to obtain the stiffness vs force plots, were produced by stimulation of the contralateral common peroneal nerve or the posterior tibial nerve and occasionally by electrical stimulation in the area of the red nucleus. The stiffness of the s.r. as a function of the background force level was determined by stretching the muscle with a square pulse of 1–2 mm amplitude and 200–300 ms duration. The stiffness at each force level was calculated by dividing the change in force by the change in length, at a point where the force trace had stabilized. The length-tension relation of the s.r. was determined by stretching the muscle 12–17 mm at a constant rate of 1–2 mm/s. At all force levels, baclofen produced a significant decrease (40% or more) in the s.r. stiffness, within 10–15 min of injection as determined from the stiffness-force plots. The length-tension plots revealed that the decrease of s.r. stiffness was always accompanied by an increase in the s.r. threshold (typically 2–3 mm). It is suggested, therefore, that the s.r. threshold is not an independent variable, depending on the membrane potential of the - motoneurons, and additionally on the level of presynaptic inhibition of the muscle spindle afferent terminals. The present results also imply that it may be possible for the CNS to adaptively modify the s.r. stiffness via presynaptic inhibition of the intraspinal terminals of muscle afferents. However, any such change of s.r. stiffness will be accompanied by a change in the s.r. threshold.     

Mapping of monoamine neurones and fibres in the cat lower brainstem and spinal cord

Summary The localization of monoaminergic neurones in the medulla oblongata and the pons, and the distribution of catecholaminergic fibres in the spinal cord of the cat were investigated by means of formaldehyde-induced (FIF) or glyoxylic-acid-induced (GIF) fluorescence. Four groups of catecholamine (CA)-containing neurones were found in the following regions: (1) in the ventrolateral medulla oblongata within and adjacent to the lateral reticular nucleus, beginning slightly rostral to the medullo-spinal junction and extending rostrally to the cranial third of the inferior olive; (2) in the commissural, medial and lateral nucleus of the solitary tract; (3) cranial to the first group, closely adjacent to the facial nucleus and the superior olive; and (4) in the dorsolateral pons distributed to different nuclei, namely the nucleus coeruleus and subcoeruleus, the Koelliker-Fuse nucleus, and the medial and lateral parabrachial nuclei. The indoleamine (IA)-containing cell bodies were in general confined to the raphe nuclei, namely the nucleus raphe pallidus, nucleus raphe obscurus, nucleus raphe magnus, nucleus raphe pontis, nucleus raphe dorsalis and the central superior nucleus. A few IA-neurones were located more laterally, especially dorsal and lateral of the cranial half of the inferior olive, around the root of the hypoglossal nerve, in the lateral tegmental field and the pontine central gray. In the spinal cord most CA-fibres were found in the intermediolateral cell column. Another dense accumulation of CA-fibres was located dorsally and laterally of the central canal. The ventral and dorsal horns also contained CA-nervefibres which were slightly more numerous in the sacral spinal cord than in the more rostral parts of the spinal cord.     

Role of upper cervical inspiratory neurons studied by cross-correlation in the cat

Summary Axonal projections and synaptic connectivity of upper cervical inspiratory neurons (UCINs) were investigated in anaesthetised cats to clarify their role as propriospinal respiratory interneurons. Antidromic mapping showed axonal collaterals near phrenic and intercostal motonuclei. Of the UCINs tested, 37% had collaterals at T3-4; 55% had ipsilateral projections and 45% had contralateral projections. Ipsilateral or contralateral cross-correlations of the activity of pairs of UCINs (one on each side of the spinal cord) with the discharge of internal intercostal, external intercostal (T3-4) or phrenic nerves revealed similar features. Those with the internal intercostal and phrenic nerves were interpreted as evidence for shared or oligosynaptic excitation, those with the external intercostal nerve as shared excitation and inhibition. No evidence for monosynaptic connections was found. Monosynaptic connections could also not be demonstrated between inspiratory intercostal neurons located near (< 0.5 mm) the UCINs collateral arborizations in T3-4, examined by cross-correlation. Afferent feedback from internal intercostal nerves (T3-4) was investigated by cross-correlating nerve stimulation with UCINs activity. Ipsilateral and contralateral cross-correlograms had similar features, providing evidence for excitation in some cases and inhibition in others. Finally, cross-correlations between ipsilateral UCINs and cervical sympathetic nerves were featureless. The results suggest that the role of UCINs as part of a respiratory propriospinal control system analagous to forelimb motor control is untenable, although they may be part of an intercostal afferent feedback loop.     

Transmission in a locomotor-related group Ib pathway from hindlimb extensor muscles in the cat

It has been previously shown that phasic stimulation of group I afferents from ankle and knee extensor muscles may entrain and/or reset the intrinsic locomotor rhythm; these afferents are thus acting on motoneurones through the spinal rhythm generators. It was also concluded that the major part of these effects originates from Golgi tendon organ Ib afferents. Transmission in this pathway to lumbar motoneurones has now been investigated during fictive locomotion in spinal cats injected with nialamide and l-DOPA, and in decerebrate cats with stimulation of the mesencephalic locomotor region. In spinal cats injected with nialamide and l-DOPA, it was possible to evoke long-latency, long-lasting reflexes upon stimulation of high threshold afferents before spontaneous fictive locomotion commenced. During that period, stimulation of ankle and knee extensor group I afferents evoked oligosynaptic excitation of extensor motoneurones, rather than the classical Ib inhibition. Furthermore, a premotoneuronal convergence (spatial facilitation) between this group I excitation and the crossed extensor reflex was established. During fictive locomotion, in both preparations, the transmission in these group I pathways was phasically modulated within the step cycle. During the flexor phase, the group I input cut the depolarised (active) phase in flexor motoneurones and evoked EPSPs in extensor motoneurones; during the extensor phase, the group I input evoked smaller EPSPs in extensor motoneurones and had virtually no effect on flexor motoneurones. The above results suggest that the group I input from extensor muscles is transmitted through the spinal rhythm generator and more particularly, through the extensor half-centre. The locomotor-related group I excitation had a central latency of 3.5–4.0 ms. The excitation from ankle extensors to ankle extensors remained after a spinal transection at the caudal part of L6 segment; the interneurones must therefore be located in the L7 and S1 spinal segments. Candidate interneurones for mediating these actions were recorded extracellularly in lamina VII of the 7th lumbar segment. Responses to different peripheral nerve stimulation (high threshold afferents and group I afferents bilaterally) were in concordance with the convergence studies in motoneurones. The interneurones were rhythmically active in the appropriate phases of the fictive locomotor cycle, as predicted by their response patterns. The synaptic input to, and the projection of these candidate interneurones must be fully identified before their possible role as components of the spinal locomotor network can be evaluated.     

Behavioural thermosensitivity after bilateral lesions of the lateral funiculi in the cervical spinal cord of the cat

Summary The behavioural thermosensitivity of six cats was measured before and after single stage, symmetrical, bilateral, surgical lesions of the cervical spinal cord. The lesions were aimed at an area in the most ventral parts of the dorsal halves of the lateral funiculi. Unilateral lesions of that area have previously been found to cause reproducible, although subtotal, contralateral thermosensory defects, which were attributed to interruption of the thermosensory spinothalamic pathway. The lesions of three of the present cats were found to be incomplete, and those animals showed no postoperative thermosensory deficiency. Two of the cats with complete lesions showed marked post-operative defects, especially immediately after the operations, but the third cat with a complete lesion showed no postoperative thermosensory defect at all. The differences between the last three animals have been compared to the irregularity of previous reports about thermosensitivity after spinal cord lesions in man and animals, and may depend on the testing technique, rather than differences of thermosensitivity per se.     

Changes in synaptology of adult cat spinal α-motoneurons after axotomy

The aim of this electron-microscopic study was to analyze the distribution of synaptic contacts on the cell bodies and dendrites of permanently axotomized adult cat spinal α-motoneurons. Following transection and ligation of the medial gastrocnemius nerve, the synaptic covering of the cell bodies and three different dendritic compartments of homonymous α-motoneurons was analyzed quantitatively at 3, 6, and 12 weeks postoperatively. The synaptic boutons were classified according to their size and the shape of their synaptic vesicles. On the soma, a transient increase in the number of boutons was noted at 3 weeks and 6 weeks postoperatively, while after 12 weeks the bouton number had decreased to half of its normal value. The transient increase was mainly due to an increase in the number of F-type boutons. At 12 weeks postoperatively, the synaptic covering was reduced by 83% on the soma and by 57% on the proximal dendrites. In the distal dendritic regions, the values for synaptic covering remained largely unchanged. In summary, axotomized motoneurons exhibit a reduction in synaptic covering which is maximal on the cell body and becomes less pronounced centrifugally along the dendrites. However, if also taking into account the loss of distal dendritic branches that occurs in axotomized motoneurons, the total loss of boutons is several times larger in the dendrites than on the soma. Received: 18 October 1996 / Accepted: 13 June 1997     

Convergent inputs from articular,cutaneous and muscle receptors onto ascending tract cells in the cat spinal cord

Summary 1.Responses were recorded from 160 ascending tract cells in segments L4 to L6 of the spinal cord in chloralose anaesthetized, spinalized cats. The tract cells were identified by antidromic activation following stimulation of pathways in the lateral and ventral funiculi at the level of the spinal cord transection at the thoracolumbar junction. Axonal conduction velocities ranged from 9 to 114 m/s. 2. A sample of 152 of the neurones examined could be subdivided according to the distribution of their receptive fields into 49 cells activated just from receptors located in skin (s cells), 17 neurones excited by receptors in deep tissues (d cells), 15 units with a convergent input from receptors in skin and deep tissues (sd cells), and 25 neurones with a convergent input from the knee joint and either skin (sj cells), deep tissues (dj cells) or both (sdj cells). No receptive fields could be demonstrated for the remaining 46 neurones. 3. S and sj cells were found almost exclusively in the dorsal horn, whereas many d, sd, sdj and dj units were in the ventral horn. Almost all of the cells that lacked receptive fields were in the ventral horn or intermediate grey. 4. Ninety-one of 158 cells (56%) demonstrated no background activity. Of these, 43 cells (27%) lacked receptive fields. Many of the silent neurones were in the ventral horn, but some were in the dorsal horn. Of 25 cells having knee joint input, 18 (72%) had background activity. 5. All of the neurones that had a receptive field in the knee joint also had a convergent input from receptors in other tissues. In 3 cases, there was a receptive field in the skin over the foot (sj cells). For 16 cells, receptive fields included not only the knee joint but also skin and deep tissue (sdj cells). Usually, the cutaneous receptive field was near the knee joint, but sometimes it was remote, such as on the foot. The deep receptive fields were chiefly in the muscles of the thigh and/or leg. For 6 dj cells, the receptive fields included not only the knee joint but also deep fields like those of sdj cells. 6. Cutaneous receptive fields were classified as low threshold (cells excited best by innocuous intensities of mechanical stimulation), wide dynamic range (cells activated by weak mechanical stimuli, but the best responses were to noxious stimuli) or high threshold (innocuous stimuli had little effect, but noxious mechanical stimuli produced a vigorous discharge). Similarly, stimulation of the knee joint with weak mechanical stimuli could excite some neurones, while others could be activated by weak or strong articular stimuli but were excited best by noxious stimuli, and still other neurones were activated by knee joint stimuli only if the intensity was noxious. 7. In several instances, contralateral receptive fields were noted. These were generally in deep tissue or in the knee joint. 8. It was concluded that many of the responses to articular stimulation of the spinal cord ascending tract cells examined in this study could have been mediated by the fine afferent fibres that supply the knee joint. Although further work will be required to determine which particular ascending tracts transmit nociceptive information concerning the knee joint, it can be proposed that many of the responses demonstrated here were likely to play a role in either joint pain of in triggering responses associated with joint pain.     

Mesencephalic projections to the first cervical segment in the cat

Mesencephalic neurons projecting to the upper cervical spinal cord were examined by mapping the distributions of labeled cells after injecting fluorescent tracers or wheat-germ agglutinin conjugated to horseradish peroxidase (WGA-HRP) into the C1 segment. Injections into the central or deep regions of the ventral horn produced retrograde labeling in cells of several mesencephalic regions. The majority of cells were found contralaterally in the superior colliculus and red nucleus, and ipsilaterally in and around the interstitial nucleus of Cajal (INC), in the cuneiform region, and in the fields of Forel. Smaller numbers of cells were located in the periaqueductal gray matter, nucleus annularis, and magnocellular nucleus of the posterior commissure. Dorsomedial injections in the ventral horn near the ventral commissure labeled only a subset of these projections, including cells in the mesencephalic reticular formation adjacent to the INC and in the nucleus annularis. Dorsolateral injections labeled some cells in the superior colliculus and were particularly effective at labeling cells in the red nucleus. These results suggest that at least ten different cell groups project to the ventral horn of the first cervical segment. Most, but not all, groups originate from regions implicated previously in the control of eye or head movements.     

Immunocytochemical localization of amino acid neurotransmitter candidates in the ventral horn of the cat spinal cord: a light microscopic study

The distribution of immunoreactivities to six amino acids, possibly related to synaptic function, was investigated in the motor nucleus of the cat L7 spinal cord (laminae VII and IX) using a postembedding peroxidase-antiperoxidase technique. Consecutive 0.5 m transverse sections of plastic-embedded tissue were incubated with antisera raised against protein-glutaraldehyde conjugates of -aminobutyric acid (GABA), glycine, aspartate, glutamate, homocysteate, and taurine. This method allowed localization of the different immunoreactivities in individual cell profiles. The results showed that all these amino acids, except homocysteate, could be clearly detected in either neuronal or glial elements in the ventral horn. In cell bodies of neurons in lamina VII, immunoreactivity was observed for aspartate, glutamate, GABA, and glycine. Adjacent section analysis revealed that combinations of immunoreactivity for glycine/glutamate/aspartate, GABA/glycine/glutamate/aspartate and glutamate/aspartate, respectively, may occur in one and the same cell. In the motor nuclei (lamina IX), immunoreactivity to amino acids was observed in two types of neuron. Large cells, probably representing -motoneurons, were harboring immunoreactivity to both glutamate and aspartate, while a few small neurons in this area displayed a colocalization of glycine, glutamate, and aspartate. Dendrites and axons in the motor nuclei cocontained glycine/glutamate/aspartate, GABA/glycine/glutamate/aspartate, and glutamate/aspartate immunoreactivities. In both laminae VII and IX, taurine-like immunoreactivity was absent in neuronal cell bodies, but highly concentrated in perivascular cells and small cells with a morphology resembling that of glial cells. A punctate immunolabeling, in all probability representing labeling of nerve terminals, could be demonstrated in the ventral horn for GABA, glycine, and glutamate, but not with certainty for aspartate or taurine. A quantitative estimate of the covering of cell bodies of -motoneuron size by immunoreactive puncta revealed that glycine immunoreactive terminal-like structures were most abundant (covering 26–42% of the somatic membrane), while glutamate immunoreactive terminals were seen least frequently (5–9% covering). GABA-immunoreactive terminals covered from 10 to 24% of the soma surface. A colocalization of GABA and glycine immunoreactivities in putative nerve terminals could be shown both in the neuropil and in close relation to cell bodies of motoneurons. These results suggest that among the studied amino acids probably only three, namely GABA, glycine, and glutamate, can be considered to be neurotransmitter candidates in the ventral horn of the cat spinal cord.     

Alterations in motoneuron properties induced by acute dorsal spinal hemisection in the decerebrate cat

Summary Using intracellular recording techniques, we studied the response characteristics of two separate populations of triceps surae motoneurons in unanesthetized decerebrate cats, recorded before and after low thoracic hemisection of the spinal cord. In each preparation, we studied the response properties of one group of motoneurons and the protocol was then repeated for a separate group, immediately following the dorsal hemisection. In each group, we examined both the minimum firing rates of motoneurons during intracellular current injection and a range of cellular properties, including input resistance, rheobase current and afterhyperpolarization time course and magnitude. Although earlier studies from this laboratory have shown substantial reductions in minimum firing rate in reflexively active motoneurons in the hemisected decerebrated preparation, the response of motoneurons to intracellular current injection in the current preparation proved to be quite different. Minimum firing rates were either normal or even somewhat higher in the post-lesion group, while the time course of the afterhyperpolarization was shortened. Moreover, these effects were not evenly distributed across the motoneuron pool. The rate effect was most evident in motoneurons with higher conduction velocity, while the afterhyperpolarization effect occurred predominantly in motoneurons with lower conduction velocity. Neither of these effects could be accounted for by lesion-induced changes in other cellular properties. We conclude that tonically active neurons with descending axons traversing dorsolateral white matter may influence both the discharge characteristics and membrane properties of spinal motoneurons in novel ways, presumably by modifying voltage or calcium activated motoneuronal conductances. The previously described reactions in the the firing rate of motoneurons after such lesions appear to be mediated by different means, perhaps by alterations in synaptic input from segmental interneurons.     

Convergence of skin reflex and corticospinal effects in segmental and propriospinal pathways to forelimb motoneurones in the cat

The organization of facilitatory convergence from cutaneous afferents (Skin) and the corticospinal tract (pyramidal tract, Pyr) in pathways to forelimb motoneurones of mainly distal muscles was studied in anaesthetized cats by analysing postsynaptic potentials (PSPs), which were spatially facilitated by combinations of stimuli to the two sources at different time intervals. Conditioning Pyr volleys facilitated Skin-evoked PSPs of fixed (1.2–3.6 ms) central latencies (Skin PSPs), suggesting that disynaptic and polysynaptic skin reflex pathways are facilitated from the pyramidal tract. The shortest latencies (1.2–1.7 ms) of pyramidal facilitation suggested direct connection of pyramidal fibres with last order neurones of skin reflex pathways. Conditioning Skin volleys facilitated Pyr-evoked PSPs of fixed, mostly disynaptic latencies (1.0–2.5 ms; Pyr PSPs), suggesting that pyramido-motoneuronal pathways are facilitated from Skin at a premotoneuronal level. The shortest pathway from skin afferents to the premotor neurones appeared to be monosynaptic. Although Pyr and Skin volleys were mutually facilitating, the facilitation curve of Pyr PSPs and that of Skin PSPs were discontinuous to each other, with the peak facilitation at different Skin-Pyr volley intervals. Transection of the dorsal column (DC) at the C5/C6 border had little effect on the latencies or amplitudes evoked by maximal stimulation and the pyramidal facilitation of Skin PSPs. In contrast, the facilitation of Pyr PSPs by Skin stimulation was greatly decreased after the DC transection, and the facilitation curve of Pyr PSPs was continuous to that of Skin PSPs, with no separate peak. Latencies of Pyr PSPs ranged similarly to those in DC intact preparations. More rostral DC transection (C4/C5 border) reduced Skin-facilitated Pyr excitatory PSPs (EPSPs) less than C5/C6 lesions, suggesting that the C5 segment also contains neurones mediating Skin-facilitated Pyr EPSPs. The results show that convergence from skin afferents and the corticospinal tract occurs at premotor pathways of different cervical segments. We suggest that corticospinal facilitation of skin reflex occurs mostly in the brachial segments and Skin facilitation of cortico-motoneuronal effects takes place largely in the rostral cervical segments and partly in the brachial segments.     

Dopaminergic control of transmission from group II muscle afferents to spinal neurones in the cat and guinea-pig

The effects of dopamine and its agonists on transmission from muscle afferents to spinal neurones were investigated in the cat and guinea-pig spinal cord, by measuring the drug effects on the amplitude of monosynaptic field potentials evoked by electrical stimulation of group I and group II muscle afferents. Local iontophoretic application of dopamine, the dopamine D1/D5 agonist SKF-38393 and the D2/D3/D4 agonist quinpirole all depressed the group II field potentials evoked at the base of the dorsal horn. Group II field potentials in the intermediate zone were depressed by dopamine to a similar degree as the dorsal horn field potentials, whereas the dopamine agonists were without effect upon them. The intermediate zone field potentials evoked by group I muscle afferents were not depressed by any of the drugs. The dopamine-evoked depression of the group II-evoked field potentials in the dorsal horn in the guinea-pig spinal cord was reduced by the simultaneous application of haloperidol. The results demonstrate that dopamine receptors mediate the depression of transmission from group II muscle afferents to interneurones in the dorsal horn, but not to neurones in the intermediate zone of the spinal cord.     

Segmental and supraspinal control of synaptic effectiveness of functionally identified muscle afferents in the cat

The present investigation documents the patterns of primary afferent depolarization (PAD) of single, functionally identified muscle afferents from the medial gastrocnemius nerve in the intact, anesthetized cat. Classification of the impaled muscle afferents as from muscle spindles or from tendon organs was made according to several criteria, which comprised measurement of conduction velocity and electrical threshold of the peripheral axons, and the maximal frequency followed by the afferent fibers during vibration, as well as the changes in discharge frequency during longitudinal stretch, the projection of the afferent fiber to the motor pool, and, in unparalyzed preparations, the changes in afferent activity during a muscle twitch. In confirmation of a previous study, we found that most muscle spindle afferents (46.1–66.6%, depending on the combination of criteria utilized for receptor classification) had a type A PAD pattern. That is, they were depolarized by stimulation of group I fibers of the posterior biceps and semitendinosus (PBSt) nerve, but not by stimulation of cutaneous nerves (sural and superficial peroneus) or the bulbar reticular formation (RF), which in many cases inhibited the PBSt-induced PAD. In addition, we found a significant fraction of muscle spindle primaries that were depolarized by stimulation of group I PBSt fibers and also by stimulation of the bulbar RF. Stimulation of cutaneous nerves produced PAD in 9.1–31.2% of these fibers (type B PAD pattern) and no PAD in 8.2–15.4% (type C PAD pattern). In contrast to muscle spindle afferents, only the 7.7–15.4% of fibers from tendon organs had a type A PAD pattern, 23–46.1% had a type B and 50–61.5% a type C PAD pattern. These observations suggest that the neuronal circuitry involved in the control of the synaptic effectiveness of muscle spindles and tendon organs is subjected to excitatory as well as to inhibitory influences from cutaneous and reticulospinal fibers. As shown in the accompanying paper, the balance between excitation and inhibition is not fixed, but can be changed by crushing the afferent axons in the peripheral nerve and allowing subsequent reconnection of these afferent fibers with muscle receptors.     

Sensitivity of monosynaptic test reflexes to facilitation and inhibition as a function of the test reflex size: a study in man and the cat

Summary In parallel experiments on humans and in the cat it was investigated how the sensitivity of monosynaptic test reflexes to facilitation and inhibition varies as a function of the size of the control test reflex itself. In man the monosynaptic reflex (the Hoffmann reflex) was evoked in either the soleus muscle (by stimulation of the tibial nerve) or the quadriceps muscle (by stimulation of the femoral nerve). In the decerebrate cat monosynaptic reflexes were recorded from the nerves to soleus and medial gastrocnemius muscles; they were evoked by stimulation of the proximal ends of the sectioned L7 and S1 dorsal roots. Various excitatory and inhibitory spinal reflex pathways were used for conditioning the test reflexes (e.g. monosynaptic Ia excitation, disynaptic reciprocal inhibition, cutaneous inhibition, recurrent inhibition, presynaptic inhibition of the Ia fibres mediating the test reflex). It was shown that the additional number of motoneurones recruited in a monosynaptic test reflex by a constant excitatory conditioning stimulus was very much dependent on the size of the test reflex itself. This dependency had the same characteristic pattern whatever the conditioning stimulus. With increasing size of the test reflex the number of additionally recruited motoneurones first increased, then reached a peak (or plateau) and finally decreased. A similar relation was also seen with inhibitory conditioning stimuli. The basic physiological factors responsible for these findings are discussed. Finally, the implications for the interpretation of experiments in man with the H-reflex technique are considered.     

Depression of transmission from group II muscle afferents by electrical stimulation of the cuneiform nucleus in the cat

The effects of short trains of electrical stimuli applied within the cuneiform nucleus and the subcuneiform region were examined on transmission from group I and group II muscle afferents to first-order spinal neurons. Variations in the effectiveness of transmission from these afferents were assessed from changes in the sizes of the monosynaptic component of extracellular field potentials evoked following stimulation of muscle nerves. Field potentials evoked from group II muscle afferents in the dorsal horn of the midlumbar and sacral segments and in the intermediate zone of the midlumbar segments were reduced when the test stimuli applied to peripheral nerves were preceded by conditioning stimulation of the cuneiform nucleus or the subcuneiform region. The depression occurred at conditioning-testing intervals of 20–400 ms, being maximal at intervals of 32–72 ms for dorsal horn potentials and 40–100 ms for intermediate zone potentials. At the shortest intervals, both group II and group I field potentials in the intermediate zone were depressed. Conditioning stimulation of the cuneiform nucleus depressed group II field potentials nearly as effectively as conditioning stimulation of the coerulear or raphe nuclei. We propose that the nonselective depression of transmission from group I and II afferents at short intervals is due to the activation of reticulospinal pathways by cells or fibers stimulated within the cuneiform area. We also propose that the selective depression of transmission from group II afferents at long intervals is mediated at least partly by monoaminergic pathways, in view of the similarity of the effects of conditioning stimulation of the cuneiform nucleus and of the brainstem monoaminergic nuclei and by directly applied monoamines (Bras et al. 1990). In addition, it might be caused by primary afferent depolarization mediated by non-monoaminergic fibers (Riddell et al. 1992).