The ovarian innervation in the dog: a preliminary study for the base for electro-acupuncture.

The origin of the canine ovarian sensory and sympathetic nerves was studied by applying horseradish peroxidase (HRP) or wheat germ agglutinin conjugated to HRP (WGA-HRP) to the ovarian stroma and into the ovarian bursa. HRP/WGA-HRP positive neurons were found bilaterally in the dorsal root ganglia of T10 to L4 segment with the majority located in T13 to L2. In sympathetic paravertebral ganglia, labeled neurons were distributed bilaterally in ganglia from T11 to L4 with the majorities located in segments T13 to L2. Both distributions show ipsilateral predominance. Labeled prevertebral neurons were mainly located in the aorticorenal ganglion, ovarian ganglia and caudal mesenteric ganglion. No labeled neurons were found in the dorsal motor nucleus of vagus, nodose ganglia or sacral segment from S1 to S3. This study provides the possible morphological basis of electro-acupuncture concerning the somato-visceral reflex of the ovary.     

Distribution of splenic, mesenteric and renal neurons in sympathetic ganglia in rats

The distribution of postganglionic neurons innervating the spleen, intestine and kidney in paravertebral and prevertebral sympathetic ganglia was studied in rats using retrograde transport of fluorescent dyes. Labelled cells were counted in the thoracolumbar chain ganglia T6-L4, splanchnic ganglia and the solar plexus (fusion of left and right coeliac ganglia and superior mesenteric ganglion). Most splenic neurons were located in the splanchnic ganglion (64%), mesenteric neurons in the solar plexus (96%) and renal neurons in the sympathetic chain ganglia (80%). These three groups of neurons were distributed in overlapping ganglia within the paravertebral chain. Innervation of the spleen and intestine from the chain ganglia was bilateral, whereas innervation of the kidney was almost entirely ipsilateral. In conclusion, the sympathetic postganglionic neurons controlling the spleen, intestine and kidney have their cell bodies in different ganglia. These three groups of neurons are candidates for innervation by different subgroups of preganglionic neurons.     

Segmental distribution and central projectionsof renal afferent fibers in the cat studied by transganglionic transport of horseradish peroxidase

The segmental and central distributions of renal nerve afferents in adult cats and kittens were studied by using retrograde and transganglionic transport of horseradish peroxidase (HRP). Transport of HRP from the central cut ends of the left renal nerves labeled afferent axons in the ipsilateral minor splanchnic nerves and sensory perikarya in the dorsal root ganglia from T12 to L4. The majority of labeled cells (85%) were located between L1 and L3. A few neurons in the contralateral dorsal root ganglia were also labeled. Labeled cells were not confined to any particular region within a dorsal root ganglion. Some examples of bifurcation of the peripheral and central processes within the ganglion were noted. A small number of preganglionic neurons, concentrated in the intermediolateral nucleus, were also identified in some experiments. In addition, many sympathetic postganglionic neurons were labeled in the renal nerve ganglia, the superior mesenteric ganglion, and the ipsilateral paravertebral ganglia from T12 to L3 Transganglionic transport of HRP labeled renal afferent projections to the spinal cord of kittens from T1 1 to L6, with the greatest concentrations between Ll and L3. These afferents extended rostrocaudally in Lissauer's tract and sent collaterals into lamina I. In the transverse plane, a major lateral projection and a minor medial projection were observed along the outer and inner margins of the dorsal horn, respectively. From the lateral projection many fibers extended medially in laminae V and VI forming dorsal and ventral bundles around Clarke's nucleus. The dorsal bundle was joined by collaterals from the medial afferent projection and crossed to the contralateral side. The ventral bundle extended into lamina VII along the lateroventral border of Clarke's nucleus. Some afferents in the lateral projection could be followed ventrally into the dorsolateral portion of lamina VII in the vicinity of the intermediolateral nucleus. In the contralateral spinal cord, labeled afferent fibers were mainly seen in laminae V and VI These results provide the first anatomical evidence for sites of central termination of renal afferent axons. Renal inputs to regions (laminae I, V, and VI) containing spinoreticular and spinothajamic tract neurons may be important in the mediation of supraspinal cardiovascular reflexes as well as in the transmission of activity from nociceptors in the kidney. In addition, the identification of a bilateral renal afferent projection in close proximity to the thoracolumbar autonomic nuclei is consistent with the demonstration in physiological experiments of a spinal pathway for the renorenal sympathetic reflexes.     

Origins of the renal innervation in the primate, Macaca fascicularis

The origins of the renal efferent and afferent nerves in 5 cynomolgus monkeys (Macaca fascicularis) were studied by using the retrograde transport of horseradish peroxidase (HRP) and horseradish peroxidase-wheat germ agglutinin (HRP-WGA). The cut ends of the right renal nerves were soaked for 30-45 min in solutions consisting of 15% HRP and 1% HRP-WGA. Three or four days later the animals were killed and the tissues examined for the presence of retrogradely labeled neurons, HRP-filled cells were observed, with rare exceptions, only in ganglia ipsilateral to the side of tracer application. Renal efferent neurons (4648-14565 cells per animal) were found in relatively equal numbers in prevertebral and paravertebral (sympathetic chain) ganglia. Labeled prevertebral cells were distributed among the renal (52%), aorticorenal (32%) and superior mesenteric (16%) ganglia, whereas labeled paravertebral neurons were mainly located in chain ganglia T11-L3, with 94% of these located in L1-3. Labeled renal sensory neurons (31-543 per animal) constituted less than 5% of all labeled cells and were found in ipsilateral dorsal root ganglia T10-L3, with (80%) in T12 and L1. The labeled sensory neurons ranged from 18-64 microns in diameter (X = 32.4 microns). With the exception of a single cell in one animal, no labeled neurons were observed in the nodose ganglia. Many parallels were observed between the organization of the renal plexuses of macaques and humans, suggesting the utility of the non-human primate as an experimental model for functional studies of renal innervation in humans.     

Sympathetic and sensory neurons projecting into the cervical sympathetic trunk in the chicken.

The cell bodies of the sensory and sympathetic pre- and postganglionic neurons projecting into the cervical sympathetic trunk were retrogradely labeled with horseradish peroxidase in the chicken. Preganglionic neurons were located in the spinal segments T1-T6 (maximum T2), postganglionic neurons in the paravertebral ganglia T1-T3 (maximum T1) and sensory neurons in the dorsal root ganglia T1-T4 (maximum T1). Labeled preganglionic neurons were widely distributed across the intermediate gray matter and lateral funiculus, but the majority of them were located in the intermediomedial area dorsolateral to the central canal. The short and long axis diameters of labeled preganglionic neurons in this area decreased caudally. From the data of the present study, it is estimated that about 4190 preganglionic, about 450 postganglionic and about 390 sensory neurons project into the cervical sympathetic trunk cranial to the paravertebral ganglion T1 in the chicken.     

Muscle sensory neurons in the spinal ganglia in the rat determined by the retrograde transport of horseradish peroxidase

The method of retrograde transport of horseradish peroxidase (HRP) was used to identify muscle sensory neurons in the spinal ganglia in the rat. Experiments were conducted on 25 albino rats. Injections of 0.06 to 0.08 ml 2 to 20% Sigma type VIHRP were made unilaterally into anterior tibial muscle. Cells of origin of muscle receptors and motor endings in the same area where HRP was administered were demonstrated. The labeled cells, medium to large, were found in fourth and fifth lumbar ganglia ipsilateral to the site of injection. Simultaneously, labeled neurons were also found in the ipsilateral ventral horn of the same cord segments as the labeled sensory ganglia.     

Origin of sympathetic and sensory innervation of the elbow joint in the rat: A retrograde axonal tracing study with wheat germ agglutinin conjugated horseradish peroxidase

After injection of wheat germ agglutinin conjugated horseradish peroxidase (WGA-HRP) into the elbow joint of adult rats, labeled neurons were found in the stellate and the T2-T4 ganglia of the ipsilateral sympathetic trunk, and also in dorsal root ganglia at the C4–T4 levels. Most labeled sympathetic cells, 90% or more, were located in the stellate ganglion. The sensory innervation to the joint originated mainly from the dorsal root ganglia at the levels of C7–T1.     

Pituitary adenylate cyclase activating polypeptide-immunoreactive sensory neurons innervate rat adrenal medulla

Rat adrenal chromaffin cells were invested by a dense network of nerve fibers immunoreactive to pituitary adenylate cyclase activating polypeptide-38 (PACAP-IR). Immunohistochemical studies demonstrated the presence of PACAP-IR in nodose and dorsal root ganglion cells, but not in neurons of the intermediolateral cell column and other autonomic nuclei of the thoracic and upper lumbar spinal cord. Somata of the T7 to T12 paravertebral ganglia were PACAP-negative. A few lightly labeled neurons were occasionally noted in the dorsal motor nucleus of the vagus. Injection of the retrograde tracer Fluorogold into the left adrenal medulla 3 days prior to sacrifice resulted in the labeling of a population of neurons in the ipsilateral spinal cord intermediolateral cell column (T1 to L1), ipsilateral and contralateral nodose ganglia and ipsilateral dorsal root ganglia from T7 to T10 inclusive. A small number of lightly labeled somata was occasionally noted in the dorsal motor nucleus of the vagus. Combined retrograde tracing and PACAP immunohistochemistry showed that a population of Fluorogold-containing nodose and dorsal root ganglion cells were also PACAP-positive. Pre-treatment of the rats with capsaicin caused a marked reduction of the PACAP-IR in the adrenal gland as well as in the superficial layers of the dorsal horn and caudal spinal trigeminal nucleus. These findings, in conjunction with the apparent absence of PACAP-IR in spinal sympathetic preganglionic neurons, sympathetic postganglionic neurons, and dorsal motor nucleus of the vagus, raise the possibility that PACAP-IR fibers observed in the adrenal medulla are primarily sensory in origin. As a corollary, catecholamine secretion from chromaffin cells may be modulated by the peptidergic sensory afferents in addition to the cholinergic sympathetic preganglionic nerve fibers.     

Increased renal interstitial hydrostatic pressure causes c-fos expression in the rat''s spinal cord dorsal horn

To describe a sympathetic afferent circuit, interstitial hydrostatic pressure in the left kidney was increased in anesthetized rats for 1.5 h to activate renal mechanoreceptor afferents. Following renal afferent stimulation, the number of immunocytochemically stained cells for the immediate early gene c-fos was increased within the dorsal horn of the spinal cord. Relative to the surgical control procedure, increasing renal interstitial hydrostatic pressure produced more immunocytochemically stained cells per tissue section in laminae I and II of the dorsal horn both ipsilateral and contralateral to the stimulated kidney in the three most caudal thoracic spinal segments. Further, the number of c-fos immunocytochemically stained cells per section in the dorsal horn ipsilateral to the stimulated kidney was 28% greater than the number of stained cells contralateral to it. The staining patterns in the dorsal horns of stimulated and control animals were similar with most labeled cells in laminae I and II. These results indicate that (1) c-fos immunocytochemical staining may be useful for tracing specific sympathetic afferent pathways, (2) sensory pathways affected by increased renal interstitial hydrostatic pressure include spinal neurons located at lower thoracic levels, and (3) some of this sympathetic afferent pathway is located contralateral to the stimulated kidney. Neurons in the contralateral dorsal horn activated by renal stimulation may mediate renorenal reflexes.     

Sexual dimorphism in sympathetic preganglionic neurons of the rat hypogastric nerve

Horseradish peroxidase (HRP) applied to one hypogastric nerve labelled sensory neurons in T11-L3 dorsal root ganglia (DRG) bilaterally and preganglionic neurons (PGN) in the spinal cord segments T13-L3. An average of 130 small DRG neurons were labelled per animal (male or female). These were concentrated in the L1 + L2 DRGs (92%). About 75% were located ipsilateral to the site of HRP application. Central projections from DRG neurons were noted throughout Lissauer's tract and in the marginal zones (medial and lateral) near the borders of Lissauer's tract. A short projection was also seen extending to the dorsolateral funiculus. More than 90% of the preganglionic neurons were located in segments L1 + L2. Most of these were found in the dorsal commissural nucleus (75%) and most of the remainder were located bilaterally in the intermediolateral columns. Somewhat more intermediolateral neurons were labelled on the ipsilateral side than on the contralateral side. There were a few intercalating neurons and a very few funicular cells. An average of 415 PGNs were labelled in the male animals and 110 in the females, demonstrating a strong sexual dimorphism. No dimorphism was found in the sensory components.     

Sympathetic and afferent somata projecting in hindlimb nerves and the anatomical organization of the lumbar sympathetic nervous system of the rat

The anatomy of the sympathetic pathways from the spinal cord to the lumbar sympathetic trunk and the inferior mesenteric ganglion was studied systematically in the rat. Details of the arrangements of white and gray rami communicantes, sympathetic trunk ganglia, the intermesenteric nerve, and the lumbar splanchnic nerves are summarized. A modified nomenclature for the segmental ganglia of the paravertebral sympathetic chain is proposed. Cell bodies of sensory and sympathetic axons projecting to the skin and skeletal muscle of the rat hindlimb were labeled retrogradely with horseradish peroxidase (HRP) in order to study numbers, segmental distribution, and location of the somata of these neurons quantitatively. HRP was applied to the nerves supplying skeletal muscle (gastrocnemius-soleus, GS), hairy skin (sural, SU; saphenous, SA) and to a mixed nerve (tibial, TI). All sensory somata and 96.4% of the sympathetic cell bodies were located ipsilaterally. Sensory somata were commonly restricted to two adjacent dorsal root ganglia (usually L3-4 for SA; L4-5 for GS, TI; L5-6 for SU). Although the sympathetic somata were more widely distributed rostrocaudally (four to six segments), their maximum was always located one or two segments more cranially than the sensory outflow, i.e., corresponding to the rami communicantes grisei. From the data, it is estimated that 420 sympathetic and 530 afferent neurons project into GS, 590 and 3,610 into SU, 920 and 3,750 into SA, and 1,070 and 5,760 into TI. These absolute neuron numbers are compared with electron microscopic fiber counts from the literature.     

Afferent innervation of the lower oesophageal sphincter of the cat. An HRP study

Labeling of afferent neurons by the retrograde axonal transport of horseradish peroxidase (HRP) was performed on anaesthetized cats in order to examine the afferent innervation of the lower oesophageal sphincter (LOS), involving both the vagal and the sympathetic nerves. The labeled cells, whose fibres follow the sympathetic pathways were found in dorsal root ganglia from T1 to L2. Nerve section experiments indicated that the main pathways involved were the splanchnic nerves, as expected from classical data. Additional pathways passing through the sympathetic cardiac branch emerging from the stellate ganglion and the thoracic sympathetic branches were also evidenced. This work corroborated the electrophysiological data showing the richness of the LOS sensory vagal innervation. Nevertheless, in this case the difficulties related to the HRP technique are particularly enhanced since the abdominal sensory vagal fibres can be affected by HRP injections.     

Horseradish peroxidase localization of the sympathetic postganglionic neurons innervating the cat heart

The localization of the sympathetic postganglionic neurons innervating the cat heart has been investigated by using retrograde axonal transport of horseradish peroxidase (HRP). HRP was injected into the subepicardial layers of 4 different cardiac regions. The animals were sacrificed 72-96 h later and fixed by perfusion via the left ventricle. The paravertebral sympathetic ganglia from the superior cervical, middle cervical and stellate ganglia to T10 ganglia were removed and processed for HRP identification. Following injections of HRP into the apex of the heart, the sinoatrial (SA) nodal region and the ventral wall of the right ventricle, we observed that HRP-labeled sympathetic neurons were localized predominantly in the right stellate ganglia, and to a lesser extent, in the right superior and middle cervical ganglia, and left stellate ganglia. Fewer labeled cells were found in the right T4-T6. T8 and T9. After HRP injection into the dorsal wall of the left ventricle, HRP-labeled cells were present mainly in the left stellate ganglia.     

An HRP study of the central projections of primary trigeminal neurons which innervate tooth pulps in the cat

Trigeminal ganglia and brain stem of adult cats were studied following HRP injections into tooth pulps or after exposure of the cut end of the inferior alveolar nerve to HRP. Ipsilateral ganglion cells within a wide range of sizes were labeled in both experimental situations, whereas no labeled cells were observed in the contralateral ganglion in any animal. Labeled central branches of tooth pulp and inferior alveolar neurons were observed in all subdivisions of the ipsilateral trigeminal sensory complex. Terminal labeling in the tooth pulp experiments was confined to the dorsomedial parts of the main sensory nucleus and subnuclei oralis and interpolaris. Caudal to the obex terminal labeling was restricted to the medial halves of laminae I, IIa and V of the medullary dorsal horn. In the inferior alveolar nerve experiments dense terminal labeling was observed in the dorsal parts of the main sensory nucleus and subnuclei oralis and interpolaris. Caudal to the obex terminal labeling was located throughout laminae I to V in contrast to the tooth pulp experiments. Neither of the two experimental situations offers any evidence for a bilateral or contralateral brain stem projection of primary trigeminal neurons.     

Golgi-like immunoperoxidase staining of dopamine neurons in the reticular formation of the rat brainstem using antibody to tyrosine-hydroxylase

Cell bodies of sympathetic and sensory axons projecting via the superficial peroneal (SP) nerve supplying hairy skin of the distal hindlimb have been labeled retrogradely with horseradish peroxidase (HRP) on both sides of three cats in which the left SP nerve had been cut and ligated about 5 months previously. Three SP nerves from unoperated cats have also been studied. The location, size, and numbers of labeled somata have been determined from serial sections of lumbosacral dorsal root and sympathetic chain ganglia after standard histochemical processing. The numbers of myelinated fibers in each nerve have also been counted. The segmental distributions of both sympathetic and sensory cell bodies were identical bilaterally in each operated animal, but the number of labeled neurons was reduced on the lesioned side. There were only about 31% of sympathetic and about 51% of sensory somata relative to the numbers on the contralateral side. The average total number of neurons labeled from SP nerves in unoperated animals was about 8% higher than on the control side of operated animals. The average number of myelinated fibers in the neuromatized nerves was not reduced with respect to that in the contralateral nerve and both of these were not significantly different from the number in unoperated animals. The dimensions of samples of labeled sympathetic and sensory somata were reduced on the side with the neuroma, both in comparison with the contralateral side and with unlabeled neurons at the same levels. The mean cross-sectional area of profiles of sympathetic ganglion cells was 76% of the control; of sensory ganglion cells, 65% of the control. Assuming that HRP labeling was not impaired, we conclude that large numbers of neurons with unmyelinated axons had degenerated in the neuromatized cutaneous nerves.     

The afferent and sympathetic components of the lumbar spinal outflow to the colon and pelvic organs in the cat. I. The hypogastric nerve

The cell bodies of the lumbar sensory and sympathetic pre- and postganglionic neurons that project to the pelvic organs in the hypogastric nerve of the cat have been labeled retrogradely with horseradish peroxidase applied to the central end of their cut axons. The numbers, segmental distribution, location, and size of these labeled somata have been determined quantitatively. Afferent and preganglionic cell bodies were located bilaterally in dorsal root ganglia and spinal cord segments L3-L5, with the maximum numbers in L4. Very few cells lay rostral to L3. Afferent cell bodies were generally very small in cross-sectional area relative to the entire population in the dorsal root ganglia. Most of the preganglionic cell bodies lay clustered just medial to the region of the intermediolateral column and extended caudally well beyond its usual limit in the upper part of L4. These neurons were, on the average, larger than the cells of the intermediolateral column itself, with the largest cells lying in the most medial positions. Most of the post-ganglionic somata were in the ipsilateral distal lobe of the inferior mesenteric ganglion, while some (usually less than 10%) lay in accessory ganglia along the lumbar splanchnic nerves and in paravertebral ganglia L3-L5. Postganglionic somata in the inferior mesenteric ganglion were larger than both labeled and unlabeled ganglion cells in the paravertebral ganglia. From the data, it is estimated that about 1,300 afferent neurons, about 1,700 preganglionic neurons, and about 17,000 postganglionic neurons project in each hypogastric nerve in the cat.     

The renal afferent pathways in the rat: a pseudorabies virus study

Retrograde tract tracing studies have indicated that dorsal root ganglion cells from T₈ to L₂ innervate the rat's left kidney. Electrophysiology studies have indicated that putative second-order sympathetic afferents are found in the dorsal horn at spinal segments T₁₀ to L₁ in laminae V–VII. Here, the spread of pseudorabies virus through renal sensory pathways was examined following 2–5 days post-infection (PI) and the virus was located immunocytochemically using a rabbit polyclonal antibody. Two days PI, dorsal root ganglion neurons (first-order sympathetic afferents) were infected with PRV. An average of 1.2, 0.8, 2.1 and 4.4% of the infected dorsal root ganglion neurons were contralateral to the injected kidney at spinal segments T₁₀, T₁₁, T₁₂ and T₁₃, respectively. Four days PI, infected neurons were detected within laminae I and II of the dorsal horn of the caudal thoracic and upper lumbar spinal cord segments. The labeling patterns in the spinal cord are consistent with previous work indicating the location of renal sympathetic sensory pathways. The nodose ganglia were labeled starting 4 days PI, suggesting the involvement of parasympathetic sensory pathways. Five days PI, infected neurons were found in the nucleus tractus solitarius. In the present study, it was unclear whether the infected neurons in the nucleus tractus solitarius are part of sympathetic or parasympathetic afferent pathways or represent a convergence of sensory information. Renal denervation prevented the spread of the virus into the dorsal root ganglia and spinal cord. Sectioning the dorsal roots from T₁₀–L₃ blocked viral spread into the spinal cord dorsal horn, but did not prevent infection of neurons in dorsal root ganglion nor did it prevent infection of putative preganglionic neurons in the intermediolateral cell column. The present results indicated that renal afferent pathways can be identified after pseudorabies virus infection of the kidney. Our results suggest that renal afferents travel in sympathetic and parasympathetic nerves and that this information may converge at the NTS.     

Spinal origins of preganglionic B and C neurons that innervate paravertebral sympathetic ganglia nine and ten of the bullfrog

These experiments were designed to characterize the distribution, morphology, and number of spinal preganglionic neurons that selectively innervate the B- and C-type sympathetic neurons in paravertebral ganglia 9 and 10 of the bullfrog. For this purpose, horseradish peroxidase (HRP) was applied to the anterior end of the sectioned sympathetic chain between ganglia 8 and 9. Subsequent retrograde axonal transport of the HRP labeled ipsilateral spinal neurons whose cell bodies form a column having rostral and caudal boundaries that are, respectively, just caudal to the level of spinal nerve 4 and midway between the entry zones of spinal nerves 7 and 8. In all segments, the labeled preganglionic somata were found in the lateral half of the spinal gray and slightly dorsal to the central canal; a position analogous to that of the intermediolateral cell column in mammals. Most labeled preganglionic neurons were multipolar in shape, and the cell bodies lying between spinal nerves 4 and 5 were, on average, larger than those found between spinal nerves 7 and 8. In transverse sections that were cut near spinal nerve 5, the axons of preganglionic neurons could be seen to exit the cord through ventral roots. Counts of labeled preganglionic neurons indicate that an average +/- S.D. of 338 +/- 89 cells innervate ganglia 9 and 10. Selective labeling of preganglionic B neurons, by cutting spinal nerves 7 and 8 central to their rami communicantes at the time of HRP application, revealed an average +/- S.D. of 137 +/- 31 cells that lie exclusively between spinal nerves 4 and 6. By contrast, selective labeling of preganglionic C neurons, by cutting the sympathetic chain rostral to ganglion 7 at the time of HRP application, revealed an average +/- S.D. of 187 +/- 77 cells in an adjacent portion of the preganglionic column that is bounded by spinal nerve 6 and by a point midway between spinal nerves 7 and 8. These results thus demonstrate a clear segmental segregation between the preganglionic B and C neurons that innervate ganglia 9 and 10.     

Retrograde tracing shows that CGRP-immunoreactive nerves of rat trachea and lung originate from vagal and dorsal root ganglia

The origins of sensory innervation of the lower respiratory tract are thought to be principally the nodose and jugular ganglia of the vagus nerve. It has been suggested and partially demonstrated that there is also a component arising from dorsal root ganglia, but the segmental levels involved are not known precisely. We have therefore investigated the origins of sensory nerves within the rat respiratory tract, particularly those containing calcitonin gene-related peptide (CGRP), using the technique of retrograde axonal tracing combined with immunohistochemistry. Injections of True blue were made into extra-thoracic trachea (n = 4 rats) and percutaneously into the right and left lung (n = 4 each). Retrogradely labelled neuronal perikarya were detected in vagal and dorsal root ganglia, and sympathetic chain ganglia. CGRP-immunoreactive cells were seen only in vagal and dorsal root ganglia. Tracheal innervation arose bilaterally in the vagal sensory ganglia but those on the right side represented the principal source; the majority of CGRP-containing neurons occurred in the jugular ganglion. A very small component of labelling occurred in spinal ganglia at levels C2-C6. The sensory innervation of the lungs was seen to arise predominantly from the ipsilateral dorsal root ganglia (45% of cells CGRP-immunoreactive) at levels T1-T6. In contrast to the trachea, the contribution of vagal sensory neurones to the lungs appeared to be less than that of the spinal ganglia. These results show that the sensory innervation of the rat lungs has a major origin in the dorsal root ganglia, in which almost half of the involved neurons contain CGRP, and confirm that most CGRP-immunoreactive nerves in the trachea arise in the right jugular ganglion.