The cutaneous contribution to the hamstring flexor reflex in the rat: an electrophysiological and anatomical study

The location and properties of the cutaneous receptive fields responsible for detecting the flexor withdrawal reflex in the posterior head of biceps femoris (pBF) and semitendinosus (ST) components of the hamstring muscle have been examined in unanaesthetized decerebrate rats, spinalized at T10-T11. Single alpha-motoneurone efferents were recorded from the nerve to pBF and the principal head of ST and their responses to ipsi- and contralateral hindlimb skin stimulation investigated. The efferents to both muscles characteristically had a low or absent background discharge and they all had mechanoreceptive fields on the ipsilateral foot. The mechanical threshold of these fields was high with no response to light touch or brush. Fifty-four percent of these units also had a smaller and weaker contralateral mechanoreceptive field. The only apparent difference between ST and pBF efferents was that more ST efferents had contralateral fields than pBF units. Noxious, hot and cold thermal stimuli applied to the ipsilateral foot activated 56% of the efferents. Mustard oil, a chemical irritant, produced a long-lasting flexor response when applied to the ipsilateral foot. The responses of these efferents to stimulation of A beta, A delta and C cutaneous afferents in the sural nerve were also studied. Short latency reflexes were elicited in all efferents by A beta inputs, longer latency reflexes were elicited in 64% by A delta inputs and very long latency responses with long afterdischarges were found in 73% of the units to C inputs. Retrograde labelling of the hamstring motoneurones with WGA-HRP indicated that they lay in ventrolateral lamina IX extending from the caudal portion of the third lumbar segment to the junction of the 5th and 6th lumbar segments. Transganglionic labelling of small diameter primary afferent terminals in the dorsal horn of cutaneous nerves innervating the foot revealed that the longitudinal distribution corresponded closely with that of the hamstring motor nucleus. The flex-or reflex in the spinal rat provides a useful model therefore, for studying how the input in nociceptive afferents is processed and transformed within the spinal cord, to produce appropriate outputs.     

Contralateral and long range ipsilateral receptive fields of fibres in the central ends of divided dorsal roots

Action potentials were recorded in the central ends of transected feline lumbosacral dorsal rootlets after all the ipsilateral lumbar and sacral and ventral roots had been divided.

A few fibres had receptive fields on the contralateral hindlimb. A smaller number had receptive fields on the ipsilateral forelimb.

Percutaneous electrical stimulation was able to drive some fibres at such high frequencies that the fibres were considered to be branches of the distant afferent. Other fibres were probably synaptically coupled to the afferent.

These results are discussed in the context of an hypothesis that primary afferents form many extraneous or aberrant branches in the spinal cord.     

The somatotopic organization of primary afferent terminals in the superficial laminae of the dorsal horn of the rat spinal cord

Transganglionic transport of wheatgerm agglutinin conjugated horse-radish peroxidase (WGA-HRP) was used to reveal the central distribution of terminals of primary afferent fibers from peripheral nerves innervating the hind leg of the rat. In separate experiments the sizes and locations of cutaneous peripheral receptive fields were determined by electrophysiological recording techniques for each of the nerves that had been labeled with WGA-HRP. By using digital image analysis, the sizes and positions of the peripheral receptive fields were correlated with the areas of superficial dorsal horn occupied by terminals of primary afferents from each of these receptive fields. Data were obtained from the posterior cutaneous nerve of the thigh, lateral sural, sural, saphenous, superficial peroneal, and tibial nerves. The subdivisions of the sciatic nerve, the sural, lateral sural, superficial peroneal, and tibial nerves each projected to a separate and distinct region of the superficial dorsal horn and collectively formed a "U"-shaped zone of terminal labeling extending from lumbar spinal segments L2 to the caudal portions of L5. The gap in the "U" extended from L2 to the L3-4 boundary and was occupied by terminals from the saphenous nerve. Collectively, all primary afferents supplying the hindlimb occupied the medial 3/4 of the superficial dorsal horn with terminals from the tibial nerve lying most medially and occupying the largest of all the terminal fields. Afferents from the superficial peroneal lay in a zone between the medially situated tibial zone and the more laterally placed sural zone. Afferents from the posterior cutaneous nerve were located most caudally and laterally. Terminal fields from the posterior cutaneous and saphenous nerves differed from the others in having split representations caused presumably by their proximity to the mid-axial line of the limb. Comparisons between the peripheral and the central representations of each nerve revealed that 1 mm2 of surface area of the superficial dorsal horn serves approximately 600-900 mm2 of hairy skin and roughly 300 mm2 of glabrous skin. The vast majority of terminal labeling observed in the dorsal horn was found in the marginal layer and substantia gelatinosa, suggesting that small diameter afferents have an orderly somatotopic arrangement in which each portion of the skin surface is innervated by afferent fibers that terminate in preferred localities within the dorsal horn.     

Central projections of primary sensory afferents to the spinal dorsal horn in the long-tailed stingray, Himantura fai

The central projections of primary sensory afferents innervating the caudal region of the pectoral fin of the long-tailed stingray (Himantura fai) were labeled by applying the lipophilic carbocyanine dye DiI to the dorsal roots in fixed tissue. These observations were complemented by examination of hemotoxylin and eosin-stained paraffin sections of the dorsal root entry zone, and transmission electron microscopy of the dorsal horn. Transverse sections of the sensory nerve and dorsal root revealed two distinct myelinated axon sizes in the sensory nerve. Although the thick and thin axons do not appear to group together in the sensory nerves and dorsal root, they segregate into a dorsally directed bundle of thin fibers and a more horizontally directed bundle of thick fibers soon after entering the spinal cord. In DiI-labeled horizontal sections, fibers were observed to enter the spinal cord and diverge into rostrally and caudally directed trajectories. Branching varicose axons could be traced in the dorsal horn gray matter in the segment of entry and about half of the adjacent rostral and caudal segments. In transverse and sagittal sections, DiI-labeled afferents were seen to innervate the superficial and, to a lesser extent, deeper laminae of the dorsal horn, but not the ventral horn. Electron microscopy of unlabeled dorsal horn sections revealed a variety of synaptic morphologies including large presynaptic elements (some containing dense-core vesicles) making synaptic contacts with multiple processes in a glomerular arrangement; in this respect, the synaptic ultrastructure is broadly similar to that seen in the dorsal horn of rodents and other mammals.     

Calcitonin gene-related peptide immunoreactivity in the cat lumbosacral spinal cord and the effects of multiple dorsal rhizotomies

This study determined the extent of the rostral projection of calcitonin gene-related peptide-like immunoreactive (CGRP-IR) primary afferents in the cat lumbosacral spinal cord. To do this we examined the distribution of CGRP-like immunoreactivity (CGRP-LI) contralateral and ipsilateral to multiple dorsal rhizotomies. In the contralateral dorsal spinal cord, CGRP-IR fibers were mostly observed in Lissauer's tract, the dorsal columns, and laminae I, II, and V. Fewer CGRP-IR fibers were observed in laminae III, IV, and VI and the area around the central canal. The location of the CGRP-LI suggests that the afferents arose from nociceptors. Unilateral dorsal rhizotomies of five consecutive segments in the lumbar enlargement caused a substantial although incomplete loss of CGRP-LI in the rhizotomized dorsal spinal cord ipsilateral to the lesions. The majority of the remaining CGRP-IR fibers were located in Lissauer's tract, the dorsal columns, and the lateral part of laminae I and V. Ventral rhizotomies or an ipsilateral hemisection in the most rostral rhizotomized segment, in addition to the dorsal rhizotomies, had no noticeable effect upon the density or location of the remaining CGRP-LI. These results suggest that the majority of the CGRP-LI within the rhizotomized region of spinal cord was contained within branches of small-diameter primary afferents that entered the spinal cord through intact dorsal roots located caudal to the rhizotomized segments of spinal cord. It is concluded that CGRP-IR small-diameter primary afferents are capable of projecting at least five segments beyond their segment of entry and supplying collaterals to the superficial and deeper layers of the dorsal horn involved in the processing of nociceptive information.     

The role of neuropeptides in the sacral autonomic reflex pathways of the cat

Immunohistochemical and pharmacological studies were conducted to examine the origin and function of peptidergic nerves in the sacral autonomic system of the cat. Leucine-enkephalin (L-Enk) immunoreactivity was identified in nerve terminals in peripheral ganglia on the surface of the urinary bladder and in the parasympathetic nucleus in the sacral spinal cord. In colchicine-treated animals L-Enk was also detected in sacral preganglionic neurons (sPGN) identified by retrograde transport of a fluorescent dye. L-Enk terminals in bladder ganglia are believed to arise from sPGN since the terminals were eliminated by transection of the sacral ventral roots. Pharmacological studies indicated that exogenous as well as endogenously released enkephalins have an inhibitory action at both ganglionic and spinal sites in the sacral outflow to the urinary bladder. Peptides were also associated with afferents nerves in the sacral autonomic system. The distribution of substance P, VIP and cholecystokinin in the sacral dorsal horn paralleled the distribution of visceral afferent projections as demonstrated with HRP techniques. Dye labeling combined with immunohistochemistry revealed that some dorsal root ganglion cells projecting to the pelvic viscera contain substance P or VIP.     

The effects of 4-aminopyridine on the cat spinal cord: rhythmic antidromic discharges recorded from the dorsal roots

In a previous paper, we have reported that 4-aminopyridine (4-AP, i.v., 10 mg/kg) induces in decerebrate spinal and paralyzed cats, a sustained rhythmic activity (2.5-8.5 Hz) in various muscle nerves. We describe here that similar discharges are recorded from the proximal stump of cut cutaneous nerves. The latter rhythmic activity arises from intense antidromic discharges in the dorsal roots. The rhythmic discharges are recorded from dorsal roots of both spinal cord enlargements as well as from thoracic roots. The rhythmic activity is highly synchronous among adjacent dorsal roots. Bilateral activity is also highly cross-correlated, but may be dissociated by unilateral stimulation of one dorsal root. It is not yet possible to determine the precise site where the antidromic discharges recorded from the dorsal roots are generated. 4-AP could act directly at the terminal level of the primary afferents or could activate interneurons impinging upon the terminals.     

The effects of bicuculline on the isolated spinal cord of the frog

An investigation has been made of the effects of topically applied bicuculline, a reported gamma-aminobutyric acid (GABA) antagonist, on the isolated, hemisected frog spinal cord by recording ventral and dorsal root potentials and reflexes evoked by volleys to various spinal cord inputs. Bicuculline had potent excitatory effects causing depolarization, spontaneous potentials in ventral and dorsal roots, and an increased polysynaptic ventral root reflex. More importantly, the alkaloid blocked presynaptic inhibition of orthodromic reflex activity produced by preceding ventral root stimulation and primary afferent depolarization. These effects were attributed to a demonstrated antagonism of the direct depolarizing effects of GABA on dorsal root terminals by the alkaloid. These actions of bicuculline suggest that GABA may be the transmitter responsible for primary afferent depolarization and presynaptic inhibition in the amphibian.     

Regenerative effect of human recombinant NGF on capsaicin-lesioned sensory neurons in the adult rat

Nerve growth factor (NGF) has the ability to increase the content of peptide transmitter in intact primary sensory afferents of the adult rat. We have previously shown that NGF can also induce a refill of peptide transmitters in capsaicin-depleted peptidergic nerve terminals of the rat paw skin upon intraplantar injection. The present study was aimed at investigating the neurochemical, immunohistochemical and functional recovery of peripheral and central terminals of capsaicin-lesioned afferents following administration of recombinant human NGF-β (rhNGF-β). The systemic capsaicin treatment in adult rats by 50 mg/kg s.c. (day 0) was followed by intraplantar rhNGF-β injections (4 μg each) into one hind paw on days 1, 2, 3, 5, 6 and by the analysis on day 8. The content of the marker peptide calcitonin gene-related peptide (CGRP) showed a 100% NGF-induced recovery in the peripheral (sciatic nerve) and central axons (lumbar dorsal roots) on the side of the NGF treatment and also in the contralateral sciatic nerve and lumbar dorsal roots. In the terminals of the hind paw skin, the recovery of the CGRP content, as measured by radioimmunoassay, was 100% in the plantar and 80% in the dorsal skin ipsilaterally, and 55% in the dorsal and plantar hind paw skin contralaterally. In the lumbar dorsal spinal cord, CGRP content recovered by 85% bilaterally. The morphological appearance of the sensory nerve terminals was visualized by CGRP-immunohistochemistry. In the paw skin, the CGRP-immunoreactive (CGRP-IR) nerve endings were restricted to a fragmentary subepidermal plexus after the capsaicin treatment, whereas the subsequent NGF treatment caused a bilateral recovery of the subepidermal plexus and an intact reinnervation of the epidermis and blood vessels with free nerve terminals. The capsaicin-induced fragmentation of the CGRP terminal plexus in laminae I and II of the lumbar spinal dorsal horn was also markedly repaired on both sides by the intraplantar NGF injections. The NGF treatment caused the CGRP nerve terminals in the spinal cord to regain their ability of releasing transmitter upon capsaicin stimulation as shown in tissue slice superfusion experiments. These results show that within one week, rhNGF-β can induce a complete reinnervation of skin and spinal cord with intact CGRP-IR nerve terminals after an acute capsaicin lesion.     

Do central terminals of intact myelinated primary afferents sprout into the superficial dorsal horn of rat spinal cord after injury to a neighboring peripheral nerve?

In order to investigate whether normal myelinated primary afferent axons sprout into the territories of adjacent injured peripheral nerve fibers in the superficial dorsal horn of the spinal cord, adult rats underwent either sectioning of the saphenous or femoral nerves on one side, or else unilateral denervation of the skin of the posterior thigh. Two weeks later cholera toxin B subunit (CTb), which is normally transported selectively by myelinated somatic primary afferents, was injected into the ipsilateral (intact) sciatic nerve. The relationship between CTb, vasoactive intestinal peptide (VIP), and binding of Bandeiraea simplicifolia isolectin B4 (IB4) was then examined in the ipsilateral dorsal horn of the second to fifth lumbar spinal segments (L2-L5). Sectioning of the femoral or saphenous nerves resulted in a reduction of IB4 binding in laminae I-II in the medial third of the dorsal horn of L2, L3, and the upper part of L4. VIP-immunoreactivity was upregulated in exactly the same regions in which IB4-binding was reduced. These correspond to the areas that were previously innervated by unmyelinated afferents in the sectioned nerves. CTb-labeling was detected in regions known to receive input from myelinated sciatic afferents: lamina I and a band extending from the inner part of lamina II (IIi) to lamina V in the L3-5 segments, and the deepest part of the dorsal horn in L2. Importantly, no CTb-labeling was detected in the outer part of lamina II (IIo) in the denervated areas. Sectioning of branches of the posterior cutaneous nerve of the thigh resulted in a reduction of IB4-binding and upregulation of VIP-immunoreactivity in the lateral part of the superficial dorsal horn of caudal L4 and L5. Again, CTb-immunoreactivity showed the normal sciatic pattern in L4-L5, with no labeling detected in lamina IIo in the denervated region. These results do not support the suggestion that the central terminals of intact myelinated afferents sprout into regions of lamina II occupied by adjacent nerves that have been axotomized peripherally.     

Topographical organization of group II afferent input in the rat spinal cord

The organization of neurons in the lumbar enlargement of the rat spinal cord processing information conveyed by group II afferents of hind-limb muscle nerves has been investigated by using cord dorsum and intraspinal field potential recording. Group II afferents of different muscle nerves were found to evoke their strongest synaptic actions in specific segments of the lumbar cord. Group II afferents of quadriceps and deep peroneal nerves evoked potentials mainly at the rostral end of the lumbar enlargement (L1-rostral L3), whereas group II afferents of gastrocnemius-soleus and hamstring nerves evoked their main synaptic actions at the caudal end of the lumbar enlargement (L5). In the central lumbar segments (caudal L3–L4), the largest group II potentials were produced by afferents of tibialis posterior and, to a lesser degree, flexor digitorum longus. Field potentials evoked by group II afferents of quadriceps, tibialis posterior, and flexor digitorum longus were largest in the dorsal horn (up to 600 μV), but also occurred in the ventral horn where they were sometimes preceded by group I field potentials. In contrast, field potentials evoked by group II afferents of gastrocnemius-soleus and hamstring nerves were restricted to the dorsal horn. These results indicate that neurons in different segments of the rat lumbar spinal cord process information from group II afferents of different hind-limb muscles. Furthermore, the topographical organization of group II neuronal systems in the rat is similar in several respects to that in the cat and may therefore represent a general organizational feature of the mammalian spinal cord. J. Comp. Neurol. 394:357–373, 1998. © 1998 Wiley-Liss, Inc.     

Immunoreactive dynorphin B in sacral primary afferent fibers of the cat

Immunocytochemical analysis of the distribution of dynorphin B terminals in the sacral spinal cord of the cat revealed a pattern of staining very similar to that produced with antisera directed against the primary afferent derived, putative neurotransmitter, vasoactive intestinal polypeptide. Labeled axons and terminals were concentrated in lamina I and V and there was dense fiber staining in the tract of Lissauer. Of particular interest was the presence of immunoreactive axons in attached dorsal rootlets. To specifically focus on the possibility that some of the sacral primary afferent fibers are dynorphin-immunoreactive, we first tried to increase perikaryal labeling in the sacral dorsal root ganglia by topical treatment with colchicine. This did not produce immunoreactive labeling of cell bodies in the ganglia. Unilateral multiple dorsal rhizotomy (L5 to coccygeal 1), however, significantly decreased the staining of dynorphin-immunoreactive axons and terminals in the tract of Lissauer and in the dorsal horn of sacral segments ipsilateral to the deafferentation. No changes were detected in the lumbar cord. Finally, radioimmunoassay of caudal lumbar and sacral dorsal root ganglia was performed. Measurable immunoreactivity was found in all ganglia assayed, but, consistent with the histochemical analysis, sacral ganglia contained the highest concentration of immunoreactive dynorphin B. These data indicate that a significant component of the sacral spinal cord dynorphin terminal immunoreactivity derives from primary afferent fibers.(ABSTRACT TRUNCATED AT 250 WORDS)     

Alterations in the distribution of stimulus-evoked c-fos in the spinal cord after neonatal peripheral nerve injury in the rat

Neonatal peripheral nerve injury results in a significant rearrangement of the central terminals of surviving axotomized and adjacent intact primary afferents in the dorsal horn of the spinal cord. This study investigates the ability of these afferents to make functional contacts with dorsal horn cells, using c-fos expression as a marker of synaptic activation. Graded electrical stimulation at A- or C-fiber strength of either the neonatally axotomized sciatic nerve or the adjacent uninjured saphenous nerve was performed in adult rats. Stimulation of the contralateral uninjured nerve served as a control. Quantitative examination of the number and distribution of c-fos-labeled cells in the spinal cord laminae was performed. Electrical stimulation of the previously axotomized sciatic nerve at A-fiber intensity resulted in many labeled profiles in laminae I-V of the lumbar spinal cord on the experimental as compared to the contralateral side. Electrical stimulation of uninjured saphenous nerve or saphenous-nerve-innervated skin (using pin electrodes) at A-fiber intensity did not evoke c-fos. Stimulation of the saphenous nerve at C-fiber intensity, however, resulted in a significant increase in the number and distribution of c-fos-labeled profiles in laminae I-V on the experimental side as compared to the contralateral control side. The results show that the distribution of c-fos-expressing cells after neonatal nerve injury is compatible with the previously demonstrated distribution of sprouting of primary afferents belonging to an uninjured nerve adjacent to an injured nerve, and that the surviving axotomized afferents are capable of transmitting signals to postsynaptic cells. These findings indicate that Abeta afferent stimulation of injured but not uninjured afferents elicits c-fos expression in postsynaptic cells. This may reflect an injury-induced maintenance of a normal developmental process whereby Abeta stimulation elicits c-fos in dorsal horn neurons.     

Primary afferent depolarization evoked by electroacupuncture in the lumbar cord of the cat

In precollicular decerebrate cats, experiments were performed to ascertain the presence of primary afferent depolarization at the slowly conducting fiber terminals of the sural nerve, in an attempt to substantiate our previous postulation of a possible presynaptic mechanism underlying acupuncture analgesia in the spinal cord. A well correlated temporal course has been observed to exist between the negativity of dorsal root potential, suppression of sural polysynaptic reflexes, and increased excitability of sural primary afferent terminals, under the influence of the same electroacupuncture to the left tsusanli point in the hindlimb. Furthermore, by a collision test and conduction velocity measurement, the acupuncture-evoked primary afferent depolarization thus indicated was found to occur solely at the terminals of the slowly conducting fibers of the sural nerve, fibers believed to transmit “pain” impulses. As primary afferent depolarization has powerful inhibitory actions and its existence is well demonstrated in the spinal cord and trigeminal system, we suggest that acupuncture can also utilize this well established mechanism in modulating “pain” information at the primary afferent level in the spinal cord. This spinal presynaptic inhibitory mechanism, however, is thought to be only a part of an overall process underlying the production of acupuncture analgesia.     

The development and postnatal organization of primary afferent projections to the rat thoracic spinal cord

Primary afferent projections to the thoracic spinal cord in fetal and postnatal rats were labelled by applying horseradish peroxidase (HRP) to the central stumps of cut peripheral nerves. Diaminobenzidine (DAB) and tetramethyl benzidine (TMB) histochemical processing procedures were used to reveal the HRP reaction product. In postnatal rats, individual muscle nerves were labelled to reveal the organization of muscle afferent projections to the motor nuclei. The terminals of muscle afferents were distributed widely across the dendritic arbors of motoneurons supplying the same muscles. No spatial segregation of the terminations of different populations of muscle afferents was discernable. Afferents supplying different regions of the skin were labelled by applying HRP to the dorsal and ventral primary rami of the spinal nerves. Afferents in the dorsal rami projected to lateral portions of both the ipsilateral and contralateral dorsal horns while afferents in the ventral rami projected to the medial portions of both dorsal horns. The projections of the dorsal rami were shifted caudally relative to those of the ventral rami. This relationship reflects the fact that the regions of skin innervated by the dorsal rami are displaced caudally relative to those innervated by the corresponding ventral rami. In fetuses, dorsal rami were labelled alone or in combination with ventral rami. These experiments disclosed the time course of development of the projections to different laminae of the spinal gray matter and revealed that afferents in the two primary rami project to appropriate regions in the ipsilateral and contralateral dorsal horns from the very outset.     

Convergence in the lumbar spinal cord of pathways activated by splanchnic nerve and hind limb cutaneous nerve stimulation

Interneurons in the L6–L7 spinal segments were examined for convergence of input froh hind limb cutaneous nerves and the ipsilateral greater splanchnic nerve in anesthetized cats. The locations of all encountered units were mapped in relation to Rexed's laminae. Cells exhibiting viscerosomatic convergence were found throughout the gray matter with the majority of units being located in the intermediate and ventral gray. A comparison of the patterns of neuronal activity and the time course of the initial positive deflection of the cord dorsum potential elicited by splanchnic nerve volleys indicates a causal relationship. The effects observed in the cord after splanchnic nerve stimulation were dependent on excitation of Aγδ fibers. Neurons exhibiting viscerosomatic convergence received input from low-threshold cutaneous afferents.     

Organization of the primary projections of the lateral line nerves in the lamprey Lampetra japonica

The lateral line sensory system of Lampetra japonica is innervated by the anterior and posterior lateral line nerves. The anterior lateral line nerve innervates all electroreceptors throughout the body and mechanoreceptors of the head. The posterior lateral line nerve innervates trunk mechanoreceptors. The anterior lateral line nerve consists of two ganglia (anterior lateral line and intracapsular) and four major peripheral branches (superficial ophthalmic, buccal, hyomandibular, and recurrent nerves). The posterior lateral line nerve has one posterior lateral line ganglion and one peripheral branch. The location and central projection patterns of the primary sensory neurons of these branches of the lateral line nerves were studied with the aid of horseradish peroxidase labeling. The ganglion cells of the buccal nerve were found in the rostral half, and those of the hyomandibular nerve were found in the caudal half of the medial part of the anterior lateral line ganglion. The lateral part of the anterior lateral line ganglion contains ganglion cells of the recurrent nerve and the superficial ophthalmic nerve. The rostral half of the intracapsular ganglion contains ganglion cells of the recurrent, hyomandibular, and buccal nerves. The ganglion cells of the posterior lateral line nerve were found in the posterior lateral line ganglion. The buccal nerve afferents terminated mainly in the lateral part of the ipsilateral mechanoreceptive medial nucleus. The peripheral part of the electroreceptive dorsal nucleus also received several afferents. The hyomandibular afferents terminated ipsilaterally in the central part of the medial nucleus and in the dorsolateral part of the dorsal nucleus. Some afferents of the hyomandibular nerve ascended and descended in the descending nucleus of the trigeminal nerve near its dorsal margin. The ventral nucleus, the primary nucleus of the VIIIth nerve, received a few fibers of the buccal and hyomandibular nerves. In the recurrent nerve, the fibers of the lateral part of the anterior lateral line ganglion terminated throughout the entire dorsal nucleus, and the fibers of the intracapsular ganglion projected to the dorsolateral part of the nucleus. The afferents of the posterior lateral line nerve terminated in the medial part of the ipsilateral medial nucleus and in the lateral part of the contralateral medial nucleus. In the cerebellar area, afferents of the anterior lateral line nerve were located laterally to those of the posterior lateral line nerve. Several fibers terminated in some branchiomotor nuclei, the cerebellar crest, and the dorsal gray near the obex level. No efferent cell bodies were found in the place where efferent neurons of the VIIIth nerve have been previously reported.     

Presynaptic and long-lasting postsynaptic inhibition during penicillin-induced spinal seizures

In experiments on seven cats we tested the hypothesis that the epileptogenic effect of penicillin (PCN) on the spinal cord is mediated by a reduction of presynaptic inhibition. PCN-induced spinal hyperactivity was not associated with changes in either the presynaptic inhibition of extensor monosynaptic reflexes by conditioning volleys in flexor muscle nerves, or in evoked dorsal root potentials. Long-lasting inhibition of monosynaptic reflexes by repetitive cutaneous nerve volleys, shown by intracellular recording to be associated with prolonged inhibitory postsynaptic potentials (IPSPs), was also not changed by PCN. Antagonism of either pre- or postsynaptic spinal inhibition is not a necessary condition for induction of spinal seizures by PCN.     

Localization of cholecystokininlike immunoreactivity in the rat spinal cord,with particular reference to the autonomic innervation of the pelvic organs

The distribution of cholecystokinin in the spinal cord was investigated by immunohistochemistry. Throughout the length of the spinal cord cholecystokinin immunoreactivity was found in laminae I and II, in the spinal re-ticular nucleus, and in the surroundings of the central canal. On the basis of the cholecystokinin pattern lamina II could be divided into a dorsal and ventral part. In the lumbar and sacral spinal cord additional terminal fields of cholecystokinin immunoreactive boutons unique to these levels were found. They corresponded to the intermediolateral nucleus and to the medial lumbar sympathetic nucleus dorsal to the central canal in the first and second lumbar segment. Also the intermediolateral nucleus in L₆–S₁ received a dense cholecystokinin positive input. Moreover, the area surrounding the central canal in L₆–S₁, contained many cholecystokinin immunoreactive structures. Combined retrograde tracing and immunocytochemistry revealed that the two cholecystokinin terminal fields characteristic for L₁–L₂ and that sur-rounding the intermediolateral nucleus in L₆–S₁ were situated corresponding to preganglionic neurons innervating pelvic organs through the hypo-gastric nerve or the pelvic nerves. It thus appears that the unique lumbosacral cholecystokinin is related to nuclei influencing pelvic structures, pointing to a special need for regulation of the organs involved in evacuation and sexual functions. Moreover, it is demonstrated that the caudal part of the spinal sympathetic system differs from the more cranial part with respect to type of afferent connections. The origin of the spinal cholecystokinin was investigated and it was found that neither complete transection of the spinal cord nor ipsilateral sectioning of three or four dorsal roots induced visible changes in the cholecystokinin staining pattern. Treatment of the caudal spinal cord with colchicine revealed the presence of cholecystokinin immunoreactive neurons in the intermediate gray, at the lateral border of the dorsal horn, in the dorsal horn proper, and in the substantia gelatinosa. These findings indicate that the majority of spinal cholecystokinin has a spinal origin.     

Presynaptic modulation of synaptic effectiveness of afferent and ventrolateral tract fibers in the frog spinal cord

In the isolated frog spinal cord, antidromic stimulation of motor nerves produces intraspinal field potentials with a characteristic spatial distribution. When recording from the ventral horn, there is a short latency (1–2 msec) response corresponding to activity generated by antidromic activation of motoneuron cell bodies and proximal dendrites. In addition, in the dorsal horn, a delayed wave (12–13 msec latency) corresponding in time with the negative dorsal root potential is also recorded. This wave (VR-SFP) is positive at the dorsal surface of the cord and inverts to negativity at more ventral regions. The negative VR-SFP is maximum between 300–500 μm depth from the dorsal surface and decays with increasing depth towards the motor nucleus. Six days after chronic section of the dorsal roots L7 to L9 in one side of the spinal cord, stimulation of the motor nerves on the deafferented side produces only the early response attributable to antidromic activation of motoneurons. No distinctive VR-SFPs are recorded at any depth within the cord. These findings are consistent with the interpretation that afferent fiber terminals are the current generators of the VR-SFP. The presynaptic and postsynaptic focal potentials recorded in the motor nucleus after stimulation of the ventrolateral tract, as well as the corresponding synaptic potentials electrotonically recorded from the ventral roots, are not depressed during conditioning stimulations which produce primary afferent depolarization. This contrasts with the depression of the presynaptic and post-synaptic focal potentials and synaptic potentials produced by stimulation of sensory fibers. It is concluded that, unlike the afferent fiber terminals, the terminals of the ventrolateral tract are not subjected to a presynaptic modulation of the type involving primary afferent depolarization.