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The Wulst of birds, which is generally considered homologous with the isocortex of mammals, comprises an elevation on the dorsum of the telencephalon that is particularly prominent in predatory species, especially those with large, frontally placed eyes, such as owls. The Wulst, therefore, is largely visual, but a relatively small rostral portion is somatosensory in nature. In barn owls this rostral somatosensory part of the Wulst forms a unique physical protuberance dedicated to the representation of the contralateral claw. Here we investigate whether the input to this ‘claw area’ arises from dorsal thalamic neurons that, in turn, receive their somatosensory input from the gracile nucleus. Following injections of biotinylated dextran amine into the gracile nucleus and cholera toxin B-chain into the claw area, terminations from the former and retrogradely labeled neurons from the latter overlapped substantially in the thalamic nucleus dorsalis intermedius ventralis anterior. These results indicate the existence in this species of a ‘classical’ trisynaptic somatosensory pathway from the body periphery to the telencephalic Wulst, via the dorsal thalamus, one that is likely involved in the barn owl’s predatory behavior. The results are discussed in the context of somatosensory projections, primarily in this and other avian species.

Recordings of extracellular unit activity in the ventrolateral medulla and of electyromyographic activity in either the M. scalenus, a principal inspiratory muscle, or the abdominal expiratory muscles, were used to identify inspiratory related (IR) neurons. IR neurons extended from levels caudal to the obex through the caudal level of the descending vestibular nucleus. This distribution was found to correspond to that of a subset of cells retrogradely labeled from injections of neuronal tracers into the upper thoracic spinal cord, where motoneurons innervating the M. scalenus were located by retrograde transport. Injections of biotinylated dextran amine at the recording sites resulted in projections to the spinal cord and brainstem. Bulbospinal axons traveled in the lateral funiculus, predominantly contralaterally, and terminated in relation to the dendrites and cell bodies of motoneurons innervating the M. scalenus. Brainstem nuclei receiving projections from injections at IR loci included the retroambigualis, tracheosyringeal motor nucleus, ventrolateral nucleus of the rostral medulla, infra-olivaris superior, ventrolateral parabrachial nucleus, and the dorsomedial nucleus of the intercollicular complex. In the finches, there were also bilateral projections to nucleus uvaeformis of the posterior thalamus. The spinal and brainstem projections are similar to those found in pigeon (Reinke and Wild, [1997] J. Comp. Neurol. 379:347–362), and probably mediate the intricate coordination of the vocal (syringeal) and respiratory systems for the control of vocalization. The distribution of IR neurons in birds is similar to that of the rostral ventral respiratory group (rVRG) in mammals. J. Comp. Neurol. 391:147–163, 1998. © 1998 Wiley-Liss, Inc.

We have recorded extracellular, inspiratory-related (IR) unit activity in the medulla at locations corresponding to those of neurons retrogradely labeled by injections of retrograde tracers in the lower brachial and upper thoracic spinal cord, injections that covered cell bodies and dendrites of motoneurons innervating inspiratory muscles. Bulbospinal neurons were distributed throughout the dorsomedial and ventrolateral medulla, from the spinomedullary junction through about 0.8 mm rostral to the obex. Almost all of the 104 IR units recorded were located in corresponding parts of the ventrolateral medulla, rostral to nucleus retroambigualis, where expiratory related units are found. Injections of biotinylated dextran amine at the recording sites labeled projections both to the spinal cord and to the brainstem. In the lower brachial and upper thoracic spinal cord, bulbospinal axons traveled predominantly in the contralateral dorsolateral funiculus and terminated in close relation to the dendrites of inspiratory motoneurons retrogradely labeled with cholera toxin B-chain. In the brainstem, there were predominantly ipsilateral projections to the nucleus retroambigualis, tracheosyringeal motor nucleus (XIIts), ventrolateral nucleus of the rostral medulla, infraolivary superior nucleus, ventrolateral parabrachial nucleus, and dorsomedial nucleus of the intercollicular complex. In all these nuclei, except XIIts, retrogradely labeled neurons were also found, indicating reciprocity of the connections. These results suggest the possibility of monosynaptic connections between inspiratory premotor neurons and inspiratory motoneurons, which, together with connections of IR neurons with other brainstem respiratory-vocal nuclei, seem likely to mediate the close coordination that exists in birds between the vocal and respiratory systems. The distribution of IR neurons in birds is similar to that of the rostral ventral respiratory group (rVRG) in mammals. J. Comp. Neurol. 379:347–362, 1997. © 1997 Wiley-Liss, Inc.

Injections of neuronal tracers were made into the dorsomedial nucleus of the intercollicular complex (DM) of pigeons and zebra finches in order to investigate the projections of this nucleus which has long been implicated in respiratory-vocal control. Despite the fact that pigeons are nonsongbirds and zebra finches are songbirds, the projections were very similar in both species. Most descended throughout the brainstem, taking ventral and dorsal trajectories, which merged in the medulla. Those descending ventrally terminated upon the ventrolateral parabrachial nucleus (PBvl), the nucleus infraolivaris superior, a nucleus of the rostral ventrolateral medulla (RVL), and the nucleus retroambigualis (RAm). Those taking a dorsal trajectory via the occipitomesencephalic tract terminated in the tracheosyringeal part of the hypoglossal nucleus (XIIts), the suprahypoglossal region, and nucleus retroambigualis. There were also substantial projections throughout an arc extending, between XIIts and RVL rostrally, and XIIts and RAm caudally. Neurons throughout this arc, which include inspiratory premotor neurons at levels straddling the obex and expiratory premotor neurons more caudally (in RAm), were retrogradely labeled from spinal injections. The DM projections were predominantly ipsilateral, but there were distinct contralateral projections to all the homologous nuclei in both species. All but the projections to PBvl and XIIts were reciprocal. In summary, the projections of DM suggest that it is able to influence all the key motor and premotor nuclei involved in patterned respiratory-vocal activity. J. Comp. Neurol. 377:392–413, 1997. © 1997 Wiley-Liss, Inc.

Injections of cholera toxin B-chain conjugated to horseradish peroxidase into individual peripheral branches of the trigeminal nerve or into the trigeminal ganglion showed that an ascending trigeminal tract (TTA) terminated in distinct ventral and dorsal divisions of the principal sensory nucleus (PrVv and PrVd, respectively), and a descending tract (TTD) terminated within pars oralis, pars interpolaris, and pars caudalis divisions of the nucleus of TTD (nTTD) and within the dorsal horn of the first six cervical spinal segments. In PrVD, mandibular, ophthalmic, and maxillary projections were predominantly located dorsally, ventrally, and medially, respectively. In nTTD, mandibular projections lay dorsomedially, ophthalmic projections lay ventrolaterally, and maxillary projections lay in between. At caudal medullary and spinal levels, mandibular projections were situated medially, ophthalmic projections were situated laterally, and maxillary projections were situated centrally. The terminations within the dorsal horn were most dense in laminae III and IV and were least dense in lamina II, with laminae III-IV also receiving topographically organised contralateral projections. Extratrigeminal projections were mainly to the external cuneate nucleus by way of a lateral descending trigeminal tract (ITTD; Dubbeldam and Karten [1978] J. Comp. Neurol. 180:661–678) and to the region of the tract of Lissauer and lamina I of the dorsal horn. Other projections were to a region medial to the apex of pars interpolaris, to the nuclei ventrolateralis anterior (Vla) and presulcalis anterior (Pas) of the solitary complex, and sparsely to the lateral reticular formation (plexus of Horsley) ventral to TTD. No projections were seen to the trigeminal motor nuclei or to the cerebellum. © 1996 Wiley-Liss, Inc.

Projections of dorsal column, spinal, and cochlear nuclei upon the central nucleus of the torus semicircularis (otherwise known as nucleus mesencephalicus lateralis, pars dorsalis, or MLd) and upon other toral nuclei were investigated in pigeon by anterograde and retrograde tracing and electrophysiological methods. The anatomical results showed that caudal regions of the dorsal column nuclei and medial lamina V of the upper four cervical spinal segments have extensive projections upon the contralateral central auditory nucleus and upon other nuclei of the torus, in particular the core portion of the preisthmic superficial area of Puelles et al. (L. Puelles, C. Rrobles, M. Martiez-de-la-Torre, and S. Martinez, 1994, J. Comp. Neurol. 340:98–125). The projections of nucleus angularis were found to terminate throughout most of the contralateral central nucleus except the dorsomedial portion at rostral levels, where the majority of the projections of nucleus laminaris were concentrated. Nucleus angularis (and to a lesser extent nucleus laminaris) was also found to have substantial projections to certain noncentral toral nuclei, in particular to the caudomedial shell nucleus of Puelles et al. (1994). As shown positively with both Nissl and cytochrome oxidase staining and negatively with substance P labeling, this nucleus is a medial extension of more caudal regions of the central nucleus, and it is suggested that it should be included as part of the auditory midbrain. The electrophysiological results confirmed the anatomical findings by showing that evoked potentials and multiunit activity can be recorded throughout the central and noncentral toral nuclei by using electrical stimulation of the radial nerve and auditory click stimuli. The core portion of the preisthmic superficial area, however, can be regarded as a distinct somatosensory nucleus of the midbrain. It is concluded that there is substantial convergence of somatosensory and auditory inputs within both central auditory and noncentral nuclei of the torus semicircularis in pigeon. © 1995 Wiley-Liss, Inc.

In birds the red nucleus is the most rostral cell group in the brain having projections to all levels of the spinal cord (Cabot et al., Prog. Brain Res., 57:79–108, 1982), but its sources of afferents are incompletely known. In order to determine these, a series retrograde and anterograde tracing experiments was carried out, largely with cholera toxin B-chain conjugated to horseradish peroxidase. The results show that a sparse and diffuse projection to the red nucleus arises from deep regions of the hyperstriatum accessorium (HA) of the anterior Wulst, and that a much more dense projection arises from the caudal part of the nucleus principalis precommissuralis and the medial part of the medial spiriform nucleus (SpMm). These last two sources were themselves shown to receive a substantial projection from HA of the anterior Wulst. The red nucleus was also shown to project upon the cerebellar cortex of lobule VI, and SpM upon the cerebellar cortex of lobules VI through IX (Karten and Finger, Brain Res., 102:335–338, 1976; Clarke, J. Comp. Neurol., 174:535–552, 1977). Double retrograde labelling experiments with fluorescein and rhodamine labelled latex microspheres injected into the cerebellar cortex and spinal cord showed that the rubrocerebellar cortical neurons are a different population from, although intermixed with, the rubrospinal neurons. © 1992 Wiley-Liss, Inc.

The ascending and descending projections of the parabrachial nuclear complex in the pigeon have been charted with autoradiographic and histochemical (WGA-HRP) techniques. The ascending projections originate from a group of subnuclei surrounding various components of the brachium conjunctivum, namely, the superficial lateral, dorsolateral, dorsomedial, and ventromedial subnuclei. The projections are predominantly ipsilateral and travel in the quintofrontal tract. They are primarily to the medial and lateral hypothalamus (including the periventricular nucleus and the strata cellulare internum and externum), certain dorsal thalamic nuclei, the nucleus of the pallial commissure, the bed nucleus of the stria terminalis, the ventral paleostriatum, the olfactory tubercle, the nucleus accumbens, and a dorsolateral nucleus of the posterior archistriatum. There are weaker or more diffuse projections to the rostral locus coeruleus (cell group A8), the compact portion of the pedunculopontine tegmental nucleus, the central grey and intercollicular region, the ventral area of Tsai, the medial spiriform nucleus, the nucleus subrotundus, the anterior preoptic area, and the diagonal band of Broca. The parabrachial subnuclei have partially differential projections to these targets, some of which also receive projections from the nucleus of the solitary tract (Arends, Wild, and Zeigler: J. Comp. Neurol. 278:405-429, '88). Most of the targets, particularly those in the basal forebrain (viz., the periventricular nucleus and the strata cellulare internum and externum of the hypothalamus, the bed nucleus of the stria terminalis, and its lateral extension into the ventral paleostriatum, which may be comparable with the substantia innominata), have reciprocal connections with the parabrachial and solitary tract subnuclei and therefore may be said to compose parts of a “visceral forebrain system” analogous to that described in the rat (Van der Kooy et al: J. Comp. Neurol. 224:1-24, '84). The descending projections to the lower brainstem arise in large part from a ventrolateral subnucleus that may be comparable with the Kolliker Fuse nucleus of mammals. They are mainly to the ventrolateral medulla, nucleus ambiguus, and massively to the hypoglossal nucleus, particularly its tracheosyringeal portion. These projections are therefore likely to be importantly involved in the control of vocalization and respiration (Wild and Arends: Brain Res. 407:191-194, '87). Some of these results have been presented in abstract form (Wild, Arends, and Zeigler: Soc. Neurosci. Abst. 13:308, '87).

Birds have well-developed basal ganglia within the telencephalon, including a striatum consisting of the medially located lobus parolfactorius (LPO) and the laterally located paleostriatum augmentatum (PA), Relatively little is known, however, about the extent and organization of the telencephalic “cortical” input to the avian basal ganglia (i. e., the avian “corticostriatal” projection system). Using retrograde and anterograde neuroanatomical pathway tracers to address this issue, we found that a large continuous expanse of the outer pallium projects to the striatum of the basal ganglia in pigeons. This expanse includes the Wulst and archistriatum as well as the entire outer rind of the pallium intervening between Wulst and archistriatum, termed by us the pallium externum (PE). In addition, the caudolateral neostriatum (NCL), pyriform cortex, and hippocampal complex also give rise to striatal projections in pigeon. A restricted number of these pallial regions (such as the “limbic” NCL, pyriform cortex, and ventral/caudal parts of the archistriatum) project to such ventral striatal structures as the olfactory tubercle (TO), nucleus accumbens (Ac), and bed nucleus of the stria terminalis (BNST). Such “limbic” pallial areas also project to medialmost LPO and lateralmost PA, while the hyperstriatum accessorium portion of the Wulst, the PE, and the dorsal parts of the archistriatum were found to project primarily to the remainder of LPO (the lateral two-thirds) and PA (the medial four-fifths). The available evidence indicates that the diverse pallial regions projecting to the striatum in birds, as in mammals, are parts of higher order sensory or motor systems. The extensive corticostriatal system in both birds and mammals appears to include two types of pallial neurons: (1) those that project to both striatum and brainstem (i. e., those in the Wulst and the archistriatum) and (2) those that project to striatum but not to brainstem (i. e., those in the PE). The lack of extensive corticostriatal projections from either type of neuron in anamniotes suggests that the anamniote-amniote evolutionary transition was marked by the emergence of the corticostriatal projection system as a prominent source of sensory and motor information for the striatum, possibly facilitating the role of the basal ganglia in movement control. © 1995 Wiley-Liss, Inc.

Nucleus uvaeformis (Uva), previously identified as a component of song control circuitry in songbirds, and nucleus dorsolateralis posterior thalami, pars caudalis (DLPc) in pigeon, were compared with respect to their relative positions in the dorsolateral part of the posterior thalamus, their cell types, and their afferent and efferent projections. Both nuclei are closely related to the habenulointerpeduncular tract, have similar cell types, and receive a dense projection from deep layers of the optic tectum, predominantly ipsilaterally, and a distinct projection from the dorsal column and external cuneate nuclei, predominantly contralaterally. Recordings of multiple unit activity evoked by visual and somatosensory stimuli were used to guide injections of tracer into either DLPc or Uva, and the projections to the telencephalon were charted. Both nuclei were found to have a major terminal field in the medial part of the ipsilateral neostriatum intermedium (NI), known as nucleus interfacialis (NIf) in songbirds, and a minor terminal field in the roof of the neostriatum caudale (NC). In pigeon, the DLPc terminations in NC were within a region known as neostriatum dorsale (Nd), and, in male songbirds, the Uva terminations were in the high vocal center (HVC). Recordings of visual and somatosensory evoked activity were then used to guide injections of tracer into NI, and the afferent and efferent projections were again compared in pigeon and songbirds. The projections from either DLPc or Uva were confirmed, and terminal fields were observed either in Nd in pigeon, the dorsolateral part of NC in female songbirds, or HVC in male songbirds. Injections of tracer into either Nd or HVC confirmed their sources of afferents in DLPc or Uva, respectively, and in NI, but there was incomplete overlap of the distribution of retrogradely labelled cells in NI and the terminal fields of DLPc or Uva. It is concluded that DLPc and Uva are comparable nuclei having similar afferent and efferent projections relaying visual and somatosensory information to the telencephalon. The possible role of this information in vocal control is discussed.

The descending, efferent projections of nucleus robustus archistriatalis were investigated in male zebra finches and greenfinches with injections of either biotinylated dextran amine or cholera toxin B-chain conjugated to horseradish peroxidase. The results show that in addition to the well-known projections to the tracheosyringeal motor nucleus and the dorsomedial nucleus of the intercollicular complex, there are other projections of comparable density to the ipsilateral nucleus ambiguus and nucleus retroambigualis. Within nucleus ambiguus, robustus axons terminate in close proximity to laryngeal motoneurons which were retrogradely labelled in the same bird by injections of cholera B-chain into the laryngeal muscles; and within nucleus retroambigualis robustus axons terminate in relation to bulbospinal neurons previously shown to project to regions of spinal cord containing motoneurons innervating abdominal expiratory muscles (J.M. Wild, Brain Res. 606:119–124, 1993). These projections of nucleus robustus thus seem well placed to coordinate syringeal, laryngeal, and expiratory muscle activity during vocalization. Other relatively sparse, but distinct, projections of nucleus robustus were found to nucleus dorsolateralis anterior thalami, pars medialis, to a narrow region between the superior olivary nucleus and the spinal lemniscus, and to the rostral ventrolateral medulla. Neurons in these last two locations were retrogradely labelled bilaterally following injections of cholera B-chain into nucleus retroambigualis of one side. Together with sparse contralateral projections of nucleus robustus to all brainstem targets receiving ipsilateral projections, potential pathways are thus identified by which the respiratory-vocal activity controlled by one side of the lower medulla can be influenced by the nucleus robustus of either side, thereby possibly bringing about bilateral coordination of respiratory-vocal output. © 1993 Wiley-Liss, Inc.

Injections of cholera toxin B subunit conjugated to horseradish peroxidase (CTB-HRP) were made into the lingual branch of the hypoglossal nerve in four species of finch in order to identify the innervation of the mechanoreceptors of the dermal papillae of the tongue, and simultaneously to determine the pattern of central projections of lingual hypoglossal afferents. The results showed that hypoglossal fibers innervate all the Herbst corpuscles and terminal cell receptors of the elaborately organized papillae of the dorsum of the tongue, of the shorter papillae in the ventral tongue, and the loose collection of Herbst corpuscles in the subpapillary region. Labelled fibers were also observed in the intralingual glands, in the intrinsic tongue muscles, and in the posterodorsal epithelium where they formed budlike structures. Retrogradely labelled cell bodies were located in the jugular ganglion and their central processes ascended and descended throughout the brainstem within the descending trigeminal tract (TTD). Terminal fields were observed within the dorsolateral part of the nucleus caudalis of TTD, predominantly ipsilaterally, and within the medial part of the dorsal horn of the first 4–6 cervical segments bilaterally. There were dense patches of termination over a dorsolateral subnucleus of the interpolated nucleus of TTD, and within two regions of the principal sensory trigeminal nucleus: a large one laterally and a small one medially. Terminal fields were also observed within the nucleus ventralis lateralis anterior of the rostral solitary complex, and within adjacent nuclei, which are probably equivalent to the dorsal sensory nuclei of the facial and glossopharyngeal nerves of other avian species. The results are interpreted in the light of the role of the tongue in species-specific patterns of feeding in finches, and the possible requirement for the central integration of touch and taste.

The ascending projections of the dorsal column and external cuneate nuclei (DCN/CuE) in the pigeon were investigated in anterograde tracing experiments by using autoradiography or wheat germ agglutinin conjugated to horseradish peroxidase (WGA-HRP). The results show that the majority of ascending projections decussate via internal arcuate fibers to form a contralateral medial lemniscus which ascends in a ventral position. In the brainstem, terminal fields were observed in the ventral lamella of the inferior olive (OI), the parabrachial nuclei (PB) of the dorsolateral pons, the intercollicular nucleus (ICo) of the midbrain, and the nucleus pretectalis diffusus (PD). In the diencephalon there were terminal fields in the strata cellulare externum and internum (SCE and SCI) of the caudal hypothalamus; in the intercalated (ICT), ventrolateral (VLT), and reticular nuclei of the ventral thalamus; in the nuclei principalis precommissuralis (PPC), spiriform medialis (SpM), and dorsolateralis posterior, pars caudalis (cDLP) of the caudal thalamus; and in the nuclei dorsalis intermedius ventralis anterior (DIVA), dorsolateralis posterior, pars rostralis (rDLP), dorsolateralis anterior (DLA), and dorsolateralis anterior, pars medialis (DLM) of the rostrodorsal thalamus. The origins of these projections within the DCN/CuE complex were verified in retrograde tracing experiments with WGA-HRP and were found to be partly differentiable with respect to their targets. The projections to DIVA, rDLP, DLA, DLM, cDLP, and SpM arise from all rostrocaudal levels of the DCN/CuE complex; those to ICo arise from caudomedial nuclear regions, while those to the hypothalamus and ventral thalamus arise from rostrolateral nuclear regions. Projections to PB arise from lamina I neurons of the dorsal horn of upper cervical spinal cord segments and from CuE. No evidence was found of a projection to the cerebellum. The distribution of the cells of origin of the medial lemniscus (ML) within the DCN/CuE complex was found to be largely coextensive with the areas of termination of primary spinal (Wild: J. Comp. Neurol. 240:377–395, '85) and some trigeminal (Dubbledam and Karten: J. Comp. Neurol. 180:661–678, '78) afferents. Furthermore, the areas of termination of the ML within the rostrodorsal and caudal thalamus are also either coextensive or closely associated with nuclei which provide a somatosensory projection to separate regions of the telencephalon (Wild: Brain Res. 412:205–223, '87). There are thus clear similarities in the overall pattern of somatosensory projections in the pigeon and in many mammalian species.

In a series of retrograde tracing studies involving the injection of WGA-HRP into the thalamus of the pigeon, labeled neurons were consistently observed in anterior regions of the vestibular nuclei. Following small dorsal thalamic injections, labeled neurons were located predominantly in rostroventrolateral regions of the superior vestibular nucleus, less numerously within the ventral part of the lateral vestibular nucleus, and least numerously within the medial vestibular nucleus. Following large dorsal thalamic injections, many more vestibular neurons were labeled, and these were distributed more extensively throughout anterior parts of the superior, lateral, and medial nuclei. No labeled neurons were found in the descending nucleus. Injections of tritiated amino acids into vestibular nuclei revealed a terminal field within the dorsal thalamic nucleus: dorsolateralis posterior, pars rostralis. The location of this field between auditory, somatosensory, and paleostriatally and neostriatally projecting nuclei suggests a general similarity to the organization of vestibulothalamic projections in mammals.

With the aid of autoradiographic and histochemical (WGA-HRP) tracing techniques, the projections of the nucleus of the nucleus of the tractus solitarius (nTS) in the pigeon have been delineated and related to the viscerotopic organization of the nucleus. As in mammals, nTS projects to both brainstem and forebrain structures. At medullary levels, projections were seen to nTS itself, to the dorsal motor nucleus of the vagus and to the subjacent and more ventral reticular formation. There is a substantial projection to the parabrachial nuclear complex with terminations in all its subnuclei and minor projections to locus coeruleus and several mesencephalic areas, including the ventral area of Tsai, the nucleus of the ascending brachium conjunctivum, and the compact portion of the tegmental pedunculopontine nucleus. At diencephalic levels, projections to the hypothalamus (magnocellular periventricular nucleus, stratum cellulare internum and externum) and dorsal thalamus were seen. Terminal fields within the basal telencephalon included the nucleus of the pallial commissure, the bed nucleus of the stria terminalis, and the nucleus accumbens. The organization of nTS projections in pigeons is correlated with the pattern of inputs to specific nTS subnuclei. Lateral tier subnuclei receiving cardiovascular and pulmonary inputs project upon the ventrolateral reticular formation and the ventrolateral parabrachial complex. Medial tier subnuclei receiving gustatory and gastrointestinal inputs project upon dorsal and medial parabrachial nuclei. Transparabrachial projections arise from nTS subnuclei receiving little or no primary input from the viscera.

The process of transganglionic transport was used to determine the pattern of primary afferent projections to the spinal cord and brainstem in the pigeon by (1) applying horseradish peroxidase (HRP) to various peripheral nerves in the leg or wing, (2) by injecting HRP-lectin into feather follicles of the wing or tail, and (3) by injecting HRP-lectin into various muscles of the leg or wing. In the spinal cord major peripheral nerves were represented heavily throughout the dorsal horn laminae but sparsely in more ventral laminae. The representations of these different nerves tended to be located in different mediolateral regions of the dorsal horn. Cutaneous nerves and feather follicles were represented predominantly in laminae I and II, and different sets of follicles were represented in different mediolateral regions of these laminae. Afferent labelling from muscles of the leg and wing was located in the lateral portion of the dorsal horn, predominantly in laminae I, II, and IV. In the caudal medulla the representation of the leg within the gracile nucleus was medial to and separate from that of the wing within the cuneate nucleus (Cu). The wing representation, however, extended laterally throughout the external cuneate nucleus (CuE) and lateral regions of the descending trigeminal tract. There was less evidence of separation of the limb representations at more rostral medullary levels where they both occupied predominantly CuE. Afferent labelling from cutaneous nerves and feather follicles was distributed lightly throughout Cu and CuE, and from muscles of both limbs primarily throughout CuE. There was also a small but specific projection from the limbs to the nucleus of the solitary tract, and from the wing to the principal sensory trigeminal nucleus. These results are discussed within a comparative context with a view to highlighting the similarities and differences in the pattern of primary afferent central projections in different vertebrates.