Gray Projection Research Articles

Nocistatin and nociceptin exert opposite effects on the excitability of central amygdala nucleus-periaqueductal gray projection neurons

Central amygdala nucleus (CeA)-periaqueductal gray (PAG) pathway is the component of descending antinociceptive circuitry. Nociceptin/orphanin FQ (N/OFQ) and nocistatin (NST) produce supraspinal pronociceptive and antinociceptive effects, respectively. We hypothesized that opposite effects of N/OFQ and NST on supraspinal pain modulation result from their opposing effects on the excitability of CeA-PAG projection neurons. This hypothesis was tested by investigating electrophysiological effects of N/OFQ and NST on medial CeA neurons that project to PAG (CeA M-PAG). N/OFQ hyperpolarized CeA M-PAG projection neurons by enhancing inwardly rectifying potassium conductance. In contrast, NST depolarized CeA M-PAG neurons by causing the opening of TRPC cation channels via G αq/11-PLC-PKC pathway. CeA M-PAG neurons hyperpolarized by N/OFQ express CRF or neurotensin mRNA. NST-responsive CeA M-PAG neurons contain CRF or substance P mRNA. Our study provides the evidence that the molecular and cellular basis for opposite effects of N/OFQ and NST on supraspinal pain regulation is their opposing effects on the excitability of peptidergic CeA M-PAG neurons.

Spinal dis-inhibition in inflammatory pain

Inflammatory diseases and neuropathic insults trigger signaling cascades, which frequently lead to intense and long-lasting pain syndromes in affected patients. Such pain syndromes are characterized not only by an increased sensitivity to painful stimuli (hyperalgesia), but also by a qualitative change in the sensory perception of other, tactile stimuli (allodynia) and the occurrence of spontaneous pain in the absence of any sensory input. Long-term potentiation (LTP)-like changes in synaptic transmission between nociceptive C-fibers and spino-periaqueductal grey projection neurons as well as a loss of inhibitory control by GABAergic and glycinergic spinal dorsal horn neurons have repeatedly been proposed as underlying principles. While considerable evidence supports a significant contribution of C-fiber LTP to hyperalgesia, such monosynaptic plasticity cannot explain the occurrence of allodynia and spontaneous pain. In this review, we focus on mechanisms of synaptic dis-inhibition in inflammatory pain and propose that pathologically heightened pain sensitivity can be reversed by restoring synaptic inhibition with drugs that target specific spinal GABAA receptor subtypes.

11. Who is in charge? Introducing criteria to evaluate health human resource (HHR) policy documents

The maldistribution of and lack of access to health professionals continues to be a major issue for policymakers at all levels of government. Additionally, the basis by which Health Human Resource (HHR) policy is determined is unclear. Publications found in independent reports, peer-reviewed journals and most importantly, grey literature, can significantly influence or inform major policy decisions for “hot button” HHR issues (1) . We propose a framework that can be used to classify, rank and evaluate HHR policy/planning documents.
 Our framework creates six major criteria that are used to evaluate policy documents. These criteria are: 1) literature review, 2) source of primary information, 3) nature of recommendations, 4) implementation strategies, 5) credibility of authors and 6) credibility of publisher. Within each category, a score from zero to three (for criteria 1-4) or zero to two (criteria 5-6) is assigned, depending on the caliber of the document. Summing the scores from each section yields a document’s overall score.
 The intent of this measure is two-fold. Firstly, we want to create a tool that can be widely utilized by policymakers to help inform their decisions. Secondly, it can be used as a springboard to stimulate discussion and debate around HHR planning and policy formulation.
 National Information Center on Health Services Research and Health Care Technology. (NICHSR) Health Services Research and Health Policy Grey Literature Project: Summary Report. 2006. http://www.nlm.nih.gov/ nichsr/greylitreport_06.html. Accessed February 20, 2007.

Electronic image stabilization algorithm based on fixed-lag smooth filtering

An algorithm using fixed-lag smooth filtering to improve the precision of EIS algorithm is presented in this paper. Firstly, gray projection algorithm (GPA) is used to calculate the interframe global motion vectors. Secondly, the fixed-lag smooth filter is used to smoothen the motion track of the original video sequence, according to different motion models. At last, the original sequence is compensated by using the compensation vectors calculated from filtering. This algorithm has been proved superior in filtering precision by experiments of video sequences in different motion scenes. This algorithm also satisfies the real-time request.

Ultrastructural analysis of ventrolateral periaqueductal gray projections to the A7 catecholamine cell group.

Stimulation of neurons in the ventrolateral periaqueductal gray produces antinociception that is mediated in part by pontine noradrenergic neurons. Previous light microscopic analysis provided suggestive evidence for a direct projection from neurons in the ventrolateral periaqueductal gray to noradrenergic neurons in the A7 cell group that innervate the spinal cord dorsal horn. Therefore, the present ultrastructural study used anterograde tracing combined with tyrosine hydroxylase immunoreactivity to provide definitive evidence that neurons in the ventrolateral periaqueductal gray form synapses with the somata and dendrites of noradrenergic neurons of the A7 cell group. Injections of the anterograde tracers biotinylated dextran amine or Phaseolus vulgaris leucoagglutinin into the ventrolateral periaqueductal gray of Sasco Sprague-Dawley rats yielded a dense innervation in the region of the lateral pons containing the A7 cell group. Electron microscopic analysis of anterogradely labeled terminals (n=401) in the region of the A7 cell group indicated that approximately 10% of these formed plasmalemmal appositions to tyrosine hydroxylase-immunoreactive dendrites with no intervening astrocytic processes. About 23% of these were asymmetric synapses, 10% were symmetric synapses, and 67% did not exhibit clearly differentiated synaptic specializations. The majority of anterogradely labeled terminals (60%) formed plasmalemmal appositions with dendrites and somata that lacked detectable tyrosine hydroxylase immunoreactivity. About 35% of these were symmetric synapses, 9% were asymmetric synapses and 56% did not form synaptic specializations. Approximately 30% of all anterogradely labeled terminals displayed features characteristic of axo-axonic synapses.The present results provide direct ultrastructural evidence to support the hypothesis that the analgesia produced by stimulation of neurons in the ventrolateral periaqueductal gray is mediated, in part, by activation of spinally projecting noradrenergic neurons in the A7 catecholamine cell group.

Lamina I-periaqueductal gray (PAG) projections represent only a limited part of the total spinal and caudal medullary input to the PAG in the cat

The periaqueductal gray is well known for its involvement in nociception control, but it also plays an important role in the emotional motor system. To accomplish these functions the periaqueductal gray receives input from the limbic system and from the caudal brainstem and spinal cord. Earlier studies gave the impression that the majority of the periaqueductal gray projecting cells in caudal brainstem and spinal cord are located in the contralateral lamina I, which is involved in nociception. The present study in the cat, however, demonstrates that of all periaqueductal gray projecting neurons in the contralateral caudal medulla less than 7% was located in lamina I. Of the spinal periaqueductal gray projecting neurons less than 29% was located in lamina I. However, within the spinal cord large segmental differences exist: in few segments of the enlargements the lamina I-periaqueductal gray projecting neurons represent a majority. In conclusion, although the lamina I-periaqueductal gray projection is a very important nociceptive pathway, it constitutes only a limited part of the total projection from the caudal medulla and spinal cord to the periaqueductal gray. These results suggest that a large portion of the medullo- and spino-periaqueductal gray pathways conveys information other than nociception.

多値パターン投影による3次元形状測定 (第3報)

多値パターン投影による3次元形状測定 (第3報)

The ventrolateral periaqueductal gray projects to caudal brainstem depressor regions: a functional-anatomical and physiological study

The reaction of shock, a precipitous, life-threatening fall in arterial pressure and heart rate, is evoked often by the combination of deep pain and blood loss following traumatic injury. A similar “shock-like” pattern of response can be evoked by excitation of the ventrolateral midbrain periaqueductal gray. Further, ventrolateral periaqueductal gray neurons are selectively activated by deep somatic or visceral pain and haemorrhage. The pathways mediating ventrolateral periaqueductal gray evoked hypotension and bradycardia are not known. In this study, the projections from the ventrolateral periaqueductal gray to “cardiovascular” regions in the caudal medulla of the rat were examined. Injections of the anterograde tracer, biotinylated dextran amine at physiologically-defined, ventrolateral periaqueductal gray depressor sites, revealed strong projections to the caudal midline medulla and to the depressor region of the caudal ventrolateral medulla. Injections of excitatory amino acids established that substantial falls in arterial pressure could be evoked from the ventrolateral periaqueductal gray-recipient parts of the caudal midline medulla. Injections of the retrograde tracer, cholera toxin subunit B at physiologically-defined, depressor sites in the caudal midline medulla and the caudal ventrolateral medulla confirmed the existence of substantial projections from the ventrolateral periaqueductal gray. Although previous studies have emphasized the importance of projections from the ventrolateral periaqueductal gray to the pressor region of the rostral ventrolateral medulla, this study has revealed the existence of strong ventrolateral periaqueductal gray projections to depressor regions within the caudal medulla (caudal midline medulla and caudal ventrolateral medulla) which likely contribute to ventrolateral periaqueductal gray-mediated hypotension and bradycardia.

Discrete subregions of the rat midbrain periaqueductal gray project to nucleus ambiguus and the periambigual region

We investigated the organization of projections from the rat midbrain periaqueductal gray to nucleus ambiguus and the periambigual region using retrograde and anterograde tract tracing techniques. Retrograde tracing results revealed that neurons that project to nucleus ambiguus arise from three discrete, longitudinally organized columns of neurons located in the supraoculomotor central gray, lateral and ventrolateral periaqueductal gray. Anterograde tracing studies demonstrated that projections from these three columns of periaqueductal gray neurons terminate with topographic specificity in nucleus ambiguus and the periambigual region. Double-labelling studies demonstrated that periaqueductal gray neurons terminate in close contiguity to cholinergic neurons in the compact, semicompact, loose and external formations of nucleus ambiguus. The present results suggest that projections from periaqueductal gray to nucleus ambiguus may mediate, in part, certain cardiovascular adjustments and vocalizations produced by stimulation of periaqueductal gray.

Extensive projections from the midbrain periaqueductal gray to the caudal ventrolateral medulla: a retrograde and anterograde tracing study in the rat.

We investigated the innervation of the caudal ventrolateral medulla by the midbrain periaqueductal gray in the rat using retrograde and anterograde tract-tracing. Iontophoretic injection of Fluoro-Gold or cholera toxin B subunit into the caudal ventrolateral medulla resulted in retrogradely labeled neurons in discrete regions of the periaqueductal gray. These labeled cells were observed throughout the rostrocaudal extent of the periaqueductal gray and were distributed (as percentage of total labeled cells) in its lateral (53-67%), ventrolateral (14-28%), ventromedial (7-16%) and dorsomedial aspects (7-10%). About 70-72% of labeled cells were found in the caudal half of the periaqueductal gray and 28-30% in the rostral half. In the ventromedial periaqueductal gray, more labeled cells were seen in the contralateral side (5-13%) than the ipsilateral side (2-3%), whereas for other periaqueductal gray areas labeling was preferentially ipsilateral. Phaseolus vulgaris leucoagglutinin anterograde tracing was used to confirm the retrograde labeling results. Following iontophoretic injection into the periaqueductal gray, labeled fibers and terminals were observed throughout the rostrocaudal extent of the caudal ventrolateral medulla. Injections in the lateral and/or ventrolateral aspect of the periaqueductal gray yielded more anterograde labeling in the ipsilateral than the contralateral caudal ventrolateral medulla, while injections in the ventromedial aspect of the periaqueductal gray produced labeling preferentially in the contralateral caudal ventrolateral medulla. The present study indicates that specific regions of the periaqueductal gray project to the caudal ventrolateral medulla and may regulate cardiovascular and respiratory functions through these connections.

Anatomical evidence for inputs to ventrolateral medullary catecholaminergic neurons from the midbrain periaqueductal gray of the rat

Previous studies have shown that the midbrain periaqueductal gray (PAG) projects to the ventrolateral medulla (VLM). Here, we studied PAG projections to the area of A1/C1 neurons in the VLM in the rat using phaseolus vulgaris leucoagglutinin (PHA-L) anterograde tracing combined with immunocytochemistry for tyrosine hydroxylase (TH) or phenylethanolamine N-methyl transferase (PNMT). Following PAG injections, PHA-L labeled fibers and terminals were intermingled among TH-immunoreactive (TH-ir) neurons in the VLM. High-power light microscopic examination revealed that some of the PHA-L labeled varicose fibers and boutons were in close contiguity with TH-ir elements. Such apparent appositions appeared more frequently on TH-ir elements in the A1 area than on TH-ir or PNMT-ir neurons ni the C1 area. These results indicate that some PAG inputs to the VLM may directly innervate A1/C1 neurons

Hypoxia sensitive neurons in the caudal hypothalamus project to the periaqueductal gray

Previous studies have demonstrated that the caudal hypothalamus modulates the respiratory responses to hypoxia and hypercapnia. In addition, many of the neurons in this area have a basal discharge related to the cardiac and/or respiratory cycles and are stimulated by hypoxia or hypercapnia. The purpose of the present study was to determine if these hypothalamic neurons project to a known cardiorespiratory area, the periaqueductal gray in the rat. In a first set of experiments, rhodamine-tagged microspheres were injected into the periaqueductal gray (PAG) to determine the areas of the caudal hypothalamus that project to the PAG. These studies revealed that the caudal hypothalamus sends strong ipsilateral and weak contralateral projections to the PAG. In a second set of experiments, single unit recordings were made from neurons in the caudal hypothalamus; the basal discharge of these neurons were examined with signal averaging techniques. Each neuron ( n = 79) was tested for a response to inhalation of a hypoxic (10% O 2) and a hypercapnic (5% CO 2) gas. Antidromic activation techniques were then used to determine if neurons in the caudal hypothalamus send projections to or through the PAG. Nineteen percent ( n = 15) of the hypothalamic neurons studied could be activated from the PAG; approximately 53% ( n = 8) of these were excited by hypoxia and 27% ( n = 4) by hypercapnia. Most of these neurons tested (42 of 64 neurons) had a basal discharge related temporally to the cardiac and/or respiratory cycles. These findings suggest that a caudal hypothalamic to periaqueductal gray projection is involved in the integrated response to hypoxia.

Distinct populations of neurons in the ventromedial periaqueductal gray project to the rostral ventral medulla and abducens nucleus

Following microinjections of a colloidal gold complex into the nucleus paragigantocellularis (PGi) of the ventral medulla, and of latex microspheres into the nucleus abducens (Abd) of the same animal, retrogradely labeled neurons were identified in the region of the contralateral supraoculomotor nucleus (SOM) in the ventromedial periaqueductal gray (PAG). Neurons labeled from the Abd were found in the SOM in the ventromedial PAG throughout the midbrain, as well as scattered ventrally in the oculomotor nucleus. Neurons in the SOM area retrogradely labeled from the PGi were most numerous rostral to the dorsal raphe nucleus and extended throughout the level of the oculomotor nucleus. Direct comparison of the two labels revealed that the neurons that project to the Abd were located slightly more ventrally than PGi-projecting neurons. Almost no doubly labeled neurons were identified, although singly labeled neurons formed adjacent but separate populations. These results indicate that neurons in the SOM area projecting to the PGi are distinct from those projecting to the Abd, and that the PGi-projecting neurons are probably not pre-oculomotor neurons. Given their more dorsal location and the nature of their target neurons in the rostral ventrolateral medulla, PGi-projecting neurons may be related to visceromotor, as opposed to oculomotor functions.

Midbrain periaqueductal gray projections to the dorsomedial medulla in the rabbit

The present study sought to determine the existence of projections from the midbrain periaqueductal gray (PAG) to the nucleus of the solitary tract (NTS) and dorsal motor nucleus of the vagus nerve (DMN) in the rabbit. Fast Blue injections into the NTS/DMN complex revealed a population of retrogradely labeled cells within the ventrolateral PAG. Deposits of wheat germ agglutinin/horseradish peroxidase (WGA/HRP) into the ventrolateral PAG revealed terminal label within the dorsomedial, lateral, ventrolateral, intermediate, and commissural subnuclei of the NTS. Label was also observed within the DMN and a heavy concentration encapsulated this nucleus. These data suggest that the projection from the PAG to the NTS/DMN complex may represent a substrate by which the PAG may influence autonomic and cardiovascular regulation, particularly during emotional arousal.

Projections from the periaqueductal gray to the rostromedial pericoerulear region and nucleus locus coeruleus: Anatomic and physiologic studies

Previous studies showed that the nucleus locus coeruleus (LC) receives two major afferent inputs from 1) nucleus paragigantocellularis and 2) nucleus prepositus hypoglossi, both in the rostral medulla. Recent reports suggested that the midbrain periaqueductal gray (PAG) projects to the rostromedial pericoerulear area and LC. Since the PAG is a major site for control of central antinociception, and since descending noradrenergic fibers have been implicated in pain modulation, we have investigated in detail the functional anatomy of projections from PAG to the dorsolateral pontine tegmentum. A combined anatomical and electrophysiological approach was used to assess the organization and synaptic influence of PAG on neurons in the rostromedial pericoerulear region and in LC proper. Injections of the tracer wheatgerm agglutinin conjugated to horseradish peroxidase encompassing LC proper and the rostromedial pericoerulear area retrogradely labeled neurons in PAG located lateral and ventrolateral to the cerebral aqueduct; injections restricted to LC proper did not consistently label PAG neurons. Deposits of the anterograde axonal tracer Phaseolus vulgaris leucoagglutinin into this same region of PAG labeled axons that robustly innervated the zone rostral and medial to LC. Only sparse fibers were observed in LC proper. Consistent with these results, focal electrical stimulation of LC antidromically activated only a few PAG neurons (6 of 100); all of these driven cells were located lateral and ventrolateral to the cerebral aqueduct. The majority of neurons in the rostromedial pericoerulear area were robustly activated by single pulse stimulation of PAG. In contrast, single pulse electrical stimulation of lateral PAG produced weak to moderate synaptic activation of some LC neurons; stimulation of ventrolateral PAG produced predominant inhibition of LC discharge, perhaps through recurrent collaterals subsequent to antidromic activation of neighboring LC cells. Taken together, these results indicate that PAG strongly innervates the region rostral and medial to LC, including Barrington's nucleus, but only weakly innervates LC proper. Although recent studies indicate that the dendrites of LC neurons ramify heavily and selectively in the rostromedial pericoerulear region, the results of the present physiological studies suggest that PAG preferentially targets rostromedial pericoerulear neurons rather than LC dendrites.

Connections between the central nucleus of the amygdala and the midbrain periaqueductal gray: Topography and reciprocity

Previous reports indicate that the midbrain periaqueductal gray and the central nucleus of the amygdala are interconnected but the organization of these projections has not been characterized. We have analyzed this reciprocal circuitry using anterograde and retrograde tracing methods and image analysis. Our findings reveal that innervation of periaqueductal gray from the central nucleus of the amygdala is extensive and discretely organized along the rostrocaudal axis of periaqueductal gray. In addition, the reciprocal projection from periaqueductal gray to the central nucleus of the amygdala is more extensive and more highly organized than previously suggested. Multiple or single discrete injections of wheatgerm agglutinin-horseradish peroxidase into several rostrocaudal levels of periaqueductal gray retrogradely labeled a substantial population of neurons, predominantly located in the medial division of the central nucleus of the amygdala. Tracer injections into the central nucleus revealed a high degree of spatial organization in the projection from this nucleus to periaqueductal gray. Two discrete longitudinally directed columns in dorsomedial and lateral/ventrolateral periaqueductal gray are heavily targeted by central amygdalar inputs throughout the rostral one-half to two-thirds of periaqueductal gray. Beginning at the level of dorsal raphe and continuing caudally, inputs from the central nucleus terminate more uniformly throughout the ventral half of periaqueductal gray. In addition, a substantial population of periaqueductal gray neurons were retrogradely labeled from the central nucleus of the amygdala; these were heterogeneously distributed along the rostrocaudal axis of periaqueductal gray, and included both raphe and non-raphe neurons. Thus, the present study demonstrates that periaqueductal gray receives heavy, highly organized projections from the central nucleus of the amygdala and, in turn, has reciprocal connections with the central nucleus. Previous studies have demonstrated that longitudinally organized columns of output neurons located in dorsomedial and lateral/ventrolateral periaqueductal gray project to the ventral medulla. Thus, there may be considerable overlap between the two longitudinally organized terminal input columns from the central nucleus of the amygdala and the two longitudinal columns of descending projection neurons from periaqueductal gray to the ventral medulla. The central nucleus of the amygdala has been implicated in a variety of emotional/cognitive functions ranging from fear and orienting responses, defensive and aversive reactions, associative conditioning, cardiovascular regulation, and antinociception. Many of these same functions are strongly represented in the periaqueductal gray. It is noteworthy that the present results demonstrate that lateral periaqueductal gray, a preeminent central trigger site for behavioral and autonomic components of the defense/aversion response, is heavily targeted by inputs from the central nucleus of the amygdala at all levels of periaqueductal gray.(ABSTRACT TRUNCATED AT 400 WORDS)

Autoradiographic and electrophysiological evidence for excitatory amino acid transmission in the periaqueductal gray projection to nucleus raphe magnus in the rat

Selective retrograde labelling was used as an autoradiographic method to identify possible excitatory amino acid afferents to nucleus raphe magnus (NRM). Injections of 25–50 nl 10 −2 or 10 −3 M d-[ 3H]aspartate into the NRM resulted in prominent labelling of cells in ventrolateral mesencephalic periaqueductal gray (PAG). Electrophysiologically, stimulation in ventrolateral PAG excited cells in NRM with a latency of 2–12 ms. With many cells, microelectrophoretic application of the excitatory amino acid antagonists, kynurenate and γ- d-glutamyl-glycine, resulted in a reversible reduction of the PAG-evoked response. Selective antagonists of N- methyl- d-aspartate (NMDA) were less effective. It is suggested that neurones in the ventrolateral PAG projecting to NRM utilize an excitatory amino acid or structurally related compound as a transmitter, and that this transmitter acts on receptors of the non-NMDA type.

Ascending projections to the mammillary nuclei in the rat: a study using retrograde and anterograde transport of wheat germ agglutinin conjugated to horseradish peroxidase.

Cells of origin of ascending afferents to the mammillary nuclei and the afferents' fields of termination within these nuclei were studied by using retrograde and anterograde transport of wheat germ agglutinin conjugated to horseradish peroxidase in the rat. The pars compacta of the superior central nucleus projects bilaterally to the median region of the medial mammillary nucleus. The ventral tegmental nucleus projects ipsilaterally to the medial mammillary nucleus, except for its median region, in a topographic manner such that the rostrodorsolateral part of the ventral tegmental nucleus projects to the medial quadrant of the medial mammillary nucleus; the rostroventromedial part projects to the dorsal quadrant; the caudodorsolateral part projects to the ventral quadrant; and the caudoventromedial part projects to the lateral quadrant. These projection fields extend throughout the longitudinal axis of the medial mammillary nucleus, except for its most caudal region, to which only the dorsolateral part of the ventral tegmental nucleus projects. This nucleus also projects topographically to the ipsilateral dorsal premammillary nucleus; the rostral part of the ventral tegmental nucleus projects to the dorsal part of the dorsal premammillary nucleus, whereas the caudal part projects to the ventral part. The periaqueductal gray around the dorsal tegmental nucleus projects bilaterally to the supramammillary nucleus. The pars alpha of the pontine periaqueductal gray projects bilaterally to the peripheral part of the lateral mammillary nucleus, whereas the pars ventralis of the dorsal tegmental nucleus projects ipsilaterally to the lateral mammillary nucleus. The results show that the tegmentomammillary projections are organized in a gradient fashion, with the rostral to caudal position of cells of origin within the tegmental nuclei of Gudden being reflected by the medial to lateral position of fields of termination within the mammillary nuclei.

Neurochemical identification of afferents onto spinomedullary neurons in the rat spinal cord central gray matter

The synaptic relationship between spinal cord central gray projection neurons and immunocytochemically identified afferents in the rat were examined at the light microscopic level using the combined techniques of retrogradely transported True blue and serotonin (5-HT), enkephalin (ENK), and substance P (SP) immunocytochemistry. At L 42-L 6, numerous retogradely labeled neurons could be identified around the central canal after large bulbar injections of True blue. Of these projection neurons, 75% were apposted by 5-HT varicosities, 57% by ENK varicosities and 58% by SP varicosities. Hemisection of the spinal cord produced a marked reduction in the amount of 5-HT immunoreactivity and the number of putative 5-HT contacts observed on neurons of the spinal cord central gray. A small decrease in SP immunoreactivity and putative contacts was seen after dorsal rhizotomy. Neither rhizotomy nor hemisection produced discernable changes in ENK immunofluorescence. Based on the distributions of 5-HT, ENK and SP in the spinal cord, we suggest that a more precise delineation of lamina X in the rat can be made according to immunocytochemical rather than strictly morphological criteria.

Tectoreticular pathways in the turtle, Pseudemys scripta. II. Morphology of tectoreticular cells.

The morphology of tectoreticular neurons in turtles was examined with serial section reconstructions of neurons retrogradely filled with HRP. Six classes of tectal neurons project into the three tectobulbar pathways characterized in the preceding paper (Sereno, '85). (1) Large multipolar neurons with somata in the central gray layers, and with moderately branched dendrites sometimes spanning over a millimeter, project into the dorsal tectobulbar pathway, TBd. Their dendrites are covered with fine spicules and tend to arborize in the lower third of the superficial gray layers. (2) Medium-sized neurons with multiple radial dendrites and somata in the central white and upper periventricular layers probably project into the ipsilateral intermediate tectobulbar pathway, TBi. Their dendrites also bear fine spicules and usually reach the tectal surface. (3) Small radial cells in the periventricular layers, and (4) small bitufted radial cells in the superficial gray project into the small caliber component of the ipsilateral ventral tectobulbar pathway, TBv(sm). (5) Medium-sized central gray neurons with stratified dendrites, and (6) medium-sized central gray neurons with horizontal dendrites probably project into the medium caliber component of the ventral tectobulbar pathway, TBv(med). In contrast to TBd and TBi neurons, these last four classes emit a spray of long, filamentous dendritic appendages in the central gray and have dendritic arbors near the top of the superficial gray. The morphology of the neurons described in this and the preceding paper is briefly discussed in light of current ideas about tectally mediated sensorimotor transformations.