Succinic dehydrogenase histochemistry reveals the location of the putative primary visual and auditory areas within the dorsal ventricular ridge of Sphenodon punctatus.

In turtles, crocodilians, lizards and snakes, the dorsal ventricular ridge (DVR) is a nuclear cell mass that contains distinct visual and auditory thalamorecipient cell groups. In the tuatara (Sphenodon punctatus), the DVR is not organized into diverse cell groups but instead possesses a trilaminar cytoarchitecture resembling that characteristic of the telencephalic cortex in reptiles. To determine if visual and auditory fields might also be present in the DVR of Sphenodon punctatus, we used succinic dehydrogenase (SDH) histochemistry, which has been shown to delineate the visual and auditory fields of the DVR in turtles, crocodilians and lizards. We also used acetylcholinesterase (AChE) histochemistry to determine the boundary between the DVR and the basal ganglia in Sphenodon. We found an SDH-rich region in the neuropil ventral to the cell plate of the rostrolateral DVR and a slightly less intense SDH-rich zone in the neuropil deep to the cell plate of the ventromedial DVR. These SDH-rich zones appear to be located at the apical dendrites of the neurons of the adjacent cell plate. These SDH-rich zones were clearly located within the DVR and were distinct from the AChE-rich striatal part of the basal ganglia, which occupied the ventrolateral wall of the telencephalon. Based on findings in other reptiles, it seems likely that the SDH-rich zone in rostrolateral DVR represents the zone of termination of nucleus rotundus visual input to the DVR, whereas the zone in ventromedial DVR represents the zone of termination of nucleus reuniens auditory input. Because a trilaminar DVR such as that in Sphenodon might be the primitive DVR condition for reptiles, our results suggest that the cytoarchitecture of the DVR and the synaptic organization of its thalamic sensory input in the common ancestor of living reptiles might have been much like of the dorsal cortex.

Telencephalic connections in the Pacific hagfish (Eptatretus stouti), with special reference to the thalamopallial system.

The pallium of hagfishes (myxinoids) is unique: It consists of a superficial "cortical" mantle of gray matter which is subdivided into several layers and fields, but it is not clear whether or how these subdivisions can be compared to those of other craniates, i.e., lampreys and gnathostomes. The pallium of hagfishes receives extensive secondary olfactory projections (Wicht and Northcutt [1993] J. Comp. Neurol. 337:529-542), but there are no experimental data on its nonolfactory connections. We therefore investigated the pallial and dorsal thalamic connections of the Pacific hagfish. Injections of tracers into the pallium labeled many cells bilaterally in the olfactory bulbs. Other pallial afferents arise from the contralateral pallium, the dorsal thalamic nuclei, the preoptic region, and the posterior tubercular nuclei. Descending pallial efferents reach the preoptic region, the dorsal thalamus, and the mesencephalic tectum but not the motor or premotor centers of the brainstem. Injections of tracers into the dorsal thalamus confirmed the presence of reciprocal thalamopallial connections. In addition, these injections revealed that there is no "preferred" pallial target for the ascending thalamic fibers; instead, ascending thalamic and secondary olfactory projections overlap throughout the pallium. The mesencephalic tectum and tegmentum, which receive afferents from a variety of sensory sources, are interconnected with the dorsal thalamus; thus, ascending nonolfactory sensory information may reach myxinoid pallia via a tectal-thalamic-telencephalic route. A comparative analysis of pallial organization reveals that the subdivisions of the pallium in gnathostomes (i.e., medial, dorsal, and lateral pallia) cannot be recognized with certainty in hagfishes.

Silver lampreys (Ichthyomyzon unicuspis) lack a gonadotropin-releasing hormone- and FMRFamide-immunoreactive terminal nerve.

The terminal nerve is a ganglionated cranial nerve with peripheral processes that enter the nasal cavity and centrally directed processes that enter the forebrain. Members of all classes of gnathostomes have been found to possess a terminal nerve, some components of which demonstrate immunoreactivity to the peptides Phe-Met-Arg-Phe-NH2 (FMRFamide) and gonadotropin-releasing hormone (GnRH). To explore the possibility that lampreys possess a terminal nerve, we examined the distribution of these peptides in the silver lamprey, Ichthyomyzon unicuspis, by using antisera to FMRFamide and to four forms of GnRH. We found cells with FMRFamide-like immunoreactivity in the preoptic area and the isthmal gray region of the mesencephalon, and found labeled fibers throughout the preoptic-infundibular region. Occasional labeled fibers were scattered through many regions of the brain, including the optic nerve and olfactory bulb; however, unlike species that possess a terminal nerve, lampreys have no immunoreactive cells or fibers in the olfactory nerve or nasal epithelia. In addition, we observed GnRH-immunoreactive cell bodies in the preoptic area of all animals and in the ventral hypothalamus of one individual. Most of the labeled fibers extended ventrally to the hypothalamus, with other fibers extending throughout the striatum and hypothalamic-neurohypophyseal region. A few fibers in other regions, including the optic nerve, were also labeled; we detected no immunoreactivity in the olfactory bulb, olfactory nerve, or nasal epithelia. The use of different GnRH antisera resulted in remarkably similar patterns of labeling of both cells and fibers. In summary, we did not observe either GnRH or FMRFamide-like immunoreactivity in the olfactory regions that represent the typical path of terminal nerve fibers, nor were we able to locate a terminal nerve ganglion. We conclude that lampreys may lack a terminal nerve, and that the previously described fiber bundle extending from the nasal sac to the ventral forebrain may constitute an extra-bulbar olfactory pathway.

Embryonic taste buds develop in the absence of innervation.

It has been hypothesized that taste buds are induced by contact with developing cranial nerve fibers late in embryonic development, since descriptive studies indicate that during embryonic development taste cell differentiation occurs concomitantly with or slightly following the advent of innervation. However, experimental evidence delineating the role of innervation in taste bud development is sparse and equivocal. Using two complementary experimental approaches, we demonstrate that taste cells differentiate fully in the complete absence of innervation. When the presumptive oropharyngeal region was taken from a donor axolotl embryo, prior to its innervation and development of taste buds, and grafted ectopically on to the trunk of a host embryo, the graft developed well-differentiated taste buds. Although grafts were invaded by branches of local spinal nerves, these neurites were rarely found near ectopic taste cells. When the oropharyngeal region was raised in culture, numerous taste buds were generated in the complete absence of neural elements. Taste buds in grafts and in explants were identical to those found in situ both in terms of their morphology and their expression of calretinin and serotonin immunoreactivity. Our findings indicate that innervation is not necessary for complete differentiation of taste receptor cells. We propose that taste buds are either induced in response to signals from other tissues, such as the neural crest, or arise independently through intrinsic patterning of the local epithelium.

The forebrain of gnathostomes: in search of a morphotype.

A morphotype of the forebrain of gnathostomes, i.e. those characters that must have been present in the forebrain of ancestral gnathostomes, was generated by using out-group analysis to identify the shared primitive characters present in the forebrains of extant gnathostomes. The nature of morphotypes and the steps in generating a morphotype are described. Because hypotheses of phylogenetic relationships profoundly affect the resulting morphotype, current hypotheses of gnathostome interrelationships are reviewed, and particular attention is paid to the problematic relationships of lobe-finned fishes. Ontogenetic studies provide the most common basis for how neural characters are grouped, and a review of the developmental literature indicates that gnathostome forebrains are segmented, with the diencephalon arising from a rostral parencephalic neuromere, which subsequently forms anterior and posterior divisions, and a more caudal synencephalic neuromere. Unfortunately, there is no agreement concerning the number of segments that form the secondary prosencephalon (telencephalon and hypothalamus). For this reason, the characters of the secondary prosencephalon must be analyzed in a topological manner. An out-group analysis of the characters of the diencephalon of extant gnathostomes reveals that the diencephalon of ancestral gnathostomes must have arisen from three segments: an anterior parencephalic segment, which gave rise to intermediate, ventrolateral and ventromedial thalamic nuclei; a posterior parencephalic segment, which gave rise to dorsal and ventral habenular nuclei, anterior, dorsal posterior, dorsal central, and, possibly, lateral posterior thalamic nuclei, and posterior tubercular nuclei; a synencephalic segment, which gave rise to pretectal nuclei, accessory optic nuclei and the nucleus of the medial longitudinal fascicle. The pretectal and posterior tubercular regions of ray-finned fishes appear to be highly derived, due to extensive cellular proliferations that give rise to numerous nuclei. The secondary prosencephalon of ancestral gnathostomes was probably divided rostrally into inverted and evaginated cerebral hemispheres, with paired olfactory bulbs arising directly from the hemispheres, and caudally into preoptic and hypothalamic areas. The cerebral hemispheres likely comprised a dorsally situated pallium divided into medial, dorsal and lateral pallial formations, as well as an intercalated pallial nucleus situated ventrolateral to the lateral pallium, and a ventrally situated subpallium divided medially into septal nuclei and a medial amygdalar nucleus and laterally into a corpus striatum. Both pallial and subpallial centers of ancestral gnathostomes probably received ascending thalamic and posterior tubercular inputs, with telencephalic efferent pathways terminating primarily in the hypothalamus, posterior tubercle and midbrain tegmentum. An out-group analysis further indicates that some taxa in each gnathostome radiation exhibit highly derived telencephalic characters due to the independent expansion of one or more pallial formations.

The diencephalon of the Pacific herring, Clupea harengus: cytoarchitectonic analysis.

The cytoarchitecture of nuclei in the preoptic area, ventral thalamus, dorsal thalamus, epithalamus, hypothalamus, posterior tuberculum, synencephalon, and pretectum and the accessory optic nuclei was analyzed in the clupeomorph teleost, Clupea harengus. Plesiomorphic (evolutionarily primitive) and apomorphic (evolutionarily derived) features of nuclei were identified by cladistic analysis. Plesiomorphic features include the cytoarchitectonic organization of most of the preoptic nuclei, the somewhat scattered cells of nucleus ventrolateralis, the compact, oval shape of nucleus intermedius, the presence of dorsoventrally oriented laminae in the central posterior nucleus, and most features of the hypothalamic nuclei. Also plesiomorphic are the presence of a thick, prominent paraventricular organ, a nucleus of the paraventricular organ, a nucleus tuberis posterior, and a preglomerular complex in which the boundaries between multiple nuclei are relatively difficult to distinguish. Additionally, the cytoarchitecture of the three synencephalic nuclei present in Clupea, the presence of small cells in nucleus pretectalis superficialis pars parvicellularis and of larger, scattered cells in nucleus pretectalis superficialis pars magnocellularis, the presence of large cells in the dorsal accessory optic nucleus that form a rostrocaudally oriented column, and the feature of a small, cell-sparse ventral accessory optic nucleus are plesiomorphic. Apomorphic features include the presence of a single, large, circular lamina that surrounds a central neuropil in all but the most caudal part of nucleus anterior, a lack of bilateral asymmetry in the habenular nuclei, the relatively small size of the periventricular nucleus of the posterior tuberculum, the presence of two, distinguishable caudomedial nuclei in the posterior tuberculum, elongation and folding of the neuropil of nucleus pretectalis superficialis pars parvicellularis, and the relatively large size of nucleus pretectalis superficialis pars magnocellularis and the posterior pretectal nucleus.

The diencephalon and optic tectum of the longnose gar, Lepisosteus osseus (L.): cytoarchitectonics and distribution of acetylcholinesterase.

The cytoarchitecture of nuclei in the diencephalon and the distribution of acetylcholinesterase (AChE) in the diencephalon and optic tectum were analyzed in the longnose gar, Lepisosteus osseus, a non-teleost actinopterygian fish. Nuclei were identified in the preoptic area, thalamus, posterior tubercle, hypothalamus, synencephalon, and pretectum which are homologous to like-named nuclei in teleosts and other non-teleost actinopterygians. Of particular note, a nucleus in the rostral diencephalon, nucleus rostrolateralis, which has previously been identified only in the osteoglossomorph Pantodon, is present in the long-nose gar. The posterior pretectal nucleus, previously identified in teleosts and in the bowfin Amia, is also present in gars. The small size of the posterior pretectal nucleus in gars supports the hypothesis that this nucleus was small plesiomorphically. The distribution of AChE in the diencephalon and optic tectum corresponds in most respects to that found in teleosts. The superficial pretectal nuclei, including the posterior pretectal nucleus, are strongly positive for AChE. In contrast, most of the nuclei within the preglomerular complex are negative for AChE. Acetylcholinesterase is present in some of the fibers in the optic tracts and in most retinorecipient nuclei, as well as in some other nuclei and tracts.

Afferent and efferent connections of the thalamic eminence in the axolotl, Ambystoma mexicanum.

Afferent and efferent connections of the thalamic eminence of the axolotl were determined using the fluorescent compound DiI as a tracer. The thalamic eminence is connected reciprocally with a number of telencephalic and diencephalic areas, particularly with the medial pallium, the amygdala and the preoptic region. Efferent connections are widespread throughout the ipsilateral diencephalon. These findings are discussed in relation to the homology of this nucleus, especially its homologue in agnathan brains.

Immunocytochemical localization of luteinizing hormone-releasing hormone (LHRH) in the nervus terminalis and brain of the big brown bat, Eptesicus fuscus.

Little is known about the immunohistochemistry of the nervous system in bats. This is particularly true of the nervus terminalis, which exerts strong influence on the reproductive system during ontogeny and in the adult. Luteinizing hormone-releasing hormone (LHRH) was visualized immunocytochemically in the nervus terminalis and brain of juvenile and adult big brown bats (Eptesicus fuscus). The peripheral LHRH-immunoreactive (ir) cells and fibers (nervus terminalis) are dispersed along the basal surface of the forebrain from the olfactory bulbs to the prepiriform cortex and the interpeduncular fossa. A concentration of peripheral LHRH-ir perikarya and fibers was found at the caudalmost part of the olfactory bulbs, near the medioventral forebrain sulcus; obviously these cells mediate between the bulbs and the remaining forebrain. Within the central nervous system (CNS), LHRH-ir perikarya and fibers were distributed throughout the olfactory tubercle, diagonal band, preoptic area, suprachiasmatic and supraoptic nuclei, the bed nuclei of stria terminalis and stria medullaris, the anterior lateral and posterior hypothalamus, and the tuber cinereum. The highest concentration of cells was found within the arcuate nucleus. Fibers were most concentrated within the median eminence, infundibular stalk, and the medial habenula. The data obtained suggest that this distribution of LHRH immunoreactivity may be characteristic for microchiropteran (insectivorous) bats. The strong projections of LHRH-containing nuclei in the basal forebrain (including the arcuate nucleus) to the habenula, may indicate close functional contact between these brain areas via feedback loops, which could be important for the processing of thermal and other environmental stimuli correlated with hibernation.

The visually related posterior pretectal nucleus in the non-percomorph teleost Osteoglossum bicirrhosum projects to the hypothalamus: a DiI study.

This study was done to elucidate the ancestral (plesiomorphic) condition for visual pathways to the hypothalamus in teleost fishes. Three patterns of pretectal organization can be discerned morphologically and histochemically in teleosts. Their taxonomic distribution suggests that the intermediately complex pattern (seen in most teleost groups) is ancestral to both the elaborate pattern (seen in percomorphs) and the simple pattern (seen in cyprinids). The pretectal nuclei involved can be demonstrated with acetylcholinesterase histochemistry selectively and reliably in different species of teleosts, suggesting that the same-named nuclei are homologous in representatives of the three different patterns. Whereas there are visual pathways to the hypothalamus in both the elaborate (percomorph) and the simple (cyprinid) patterns, different pretectal and hypothalamic nuclei are involved. Thus visual hypothalamic pathways in these two patterns would not appear to be homologous. In this study, circuitry within the third, i.e., the intermediately complex, pattern is investigated. It is demonstrated that visual pathways project via the pretectum to the hypothalamus in Osteoglossum bicirrhosum and that they are very similar to the visual pathways in the elaborate pattern. This suggests that the circuitry in the intermediately complex pattern, as represented by Osteoglossum, is plesiomorphic (evolutionarily primitive) and the circuitry in both the simple pattern (seen in cyprinids) and the elaborate pattern (seen in percomorphs) is apomorphic (evolutionarily derived) for teleosts.

Comparative cytoarchitectonic analysis of some visual pretectal nuclei in teleosts.

The posterior pretectal nucleus, which in Osteoglossum receives second order visual input and projects to the inferior lobe of the hypothalamus, was identified and characterized in species from all major groups of non-neoteleost teleosts. The hypothesis that the posterior pretectal nucleus in these species is homologous to both the pars intermedius of the superficial pretectal nucleus and nucleus glomerulosus in acanthopterygians is supported by multiple similarities in relative position and cytoarchitecture. Nucleus corticalis, which receives retinal input and projects to the posterior pretectal nucleus (or to nucleus glomerulosus), was identified in species belonging to three of the four major teleost radiations. Both the posterior pretectal nucleus and nucleus corticalis are plesiomorphic for teleosts. The presence of glomeruli in the posterior pretectal nucleus and nucleus glomerulosus in esocids and acanthopterygians, respectively, and the presence of two nuclei, the pars intermedius and nucleus glomerulosus, in acanthopterygians, as opposed to one nucleus, the posterior pretectal nucleus, are apomorphies.

Visual pathways in elasmobranchs: organization and phylogenetic implications.

Although earlier experimental studies of the visual system in elasmobranch fishes suggested that these fishes possess fewer primary retino-recipient nuclei than other gnathostome vertebrates, recent studies utilizing more sensitive tracing methods indicate that most elasmobranch species possess ten primary retinofugal targets in addition to the optic tectum. Furthermore, many species appear to exhibit bilateral retinal projections to these nuclei. Similarly, initial claims that the organization of the visual thalamus of elasmobranchs is more primitive than that of most other gnathostomes--in that elasmobranchs possess only a single thalamic nucleus that receives both retinal and tectal inputs and that only a single thalamo-telencephalic projection exists to the telencephalon--have been refuted. Many, if not all, elasmobranchs possess a rostrally located dorsal thalamic nucleus (anterior thalamic nucleus), that receives retinal and tectal inputs and projects bilaterally to the dorsal and medial pallium, and a more caudally and dorsally located thalamic nucleus, the dorsal posterior thalamic nucleus, that receives bilateral tectal input and projects to the ventrolateral periventricular area and/or dorsal pallium of the telencephalon. Thus the thalamic organization of elasmobranch fishes is similar to that of other gnathostomes.

Nuclear organization of the bullfrog diencephalon.

A cytoarchitectonic analysis was performed on the diencephalic nuclei of the bullfrog, Rana catesbeiana. The epithalamus contains two widely recognized habenular nuclei. The thalamus has three subdivisions: dorsal and ventral thalamus, and posterior tuberculum. The dorsal thalamus may be further parcelled into anterior, middle, and posterior zones. Connectional data from other studies support this zonation. The anterior zone projects to the telencephalic pallium. The middle zone nuclei receive a strong input from the midbrain roof and project to the telencephalic striatal complex. The posterior zone nuclei do not appear to project to the telencephalon; they may eventually be placed in the pretectum, a transitional area between the diencephalon and mesencephalon. Two of the ventral thalamic populations have been frequently placed in the dorsal thalamus and called the nucleus rotundus and the lateral geniculate nucleus. These terms imply homology with sauropsid dorsal thalamic nuclei, but our analysis and current connectional information do not support such homologies. We have given these populations more neutral names. The hypothalamus is divisible into a preoptic and infundibular hypothalamus, and the preoptic area can be further separated into anterior and posterior preoptic areas. The posterior area contains the magnocellular preoptic nucleus and a dorsal arm of this nucleus, often placed in the ventral thalamus, was recognized. We have tentatively placed the posterior entopeduncular nucleus in the hypothalamus.

Localization of neurons afferent to the optic tectum in longnose gars.

Afferent pathways to the optic tectum in the longnose gar were determined by unilateral tectal injections of HRP. Retrogradely labeled cells were observed in the ipsilateral caudal portion of the rostral entopeduncular nucleus and bilaterally in the rostral half of the lateral zone of area dorsalis of the telencephalon. The following diencephalic cell groups were also labeled following tectal injections: the ipsilateral anterior, ventrolateral, and ventromedial thalamic nuclei, the periventricular pretectal nucleus, and the central pretectal nucleus (bilaterally); the ventromedial thalamic and central pretectal nuclei revealed the largest number of labeled cells. At midbrain levels, retrogradely labeled cells were seen in the ipsilateral torus longitudinalis, nucleus isthmi, and accessory optic nucleus; cells were labeled bilaterally in the torus semicircularis and a rostral tegmental nucleus. Only a few cells were labeled in the contralateral optic tectum, suggesting that few of the fibers of the intertectal commissure are actually commissural to the tectum. At hindbrain levels, retrogradely labeled cells were seen bilaterally in the locus coeruleus, the superior, medial, and inferior reticular formations, the eurydendroid cells of the cerebellum, and the nucleus of the descending trigeminal tract; the contralateral dorsal funicular nucleus also exhibited labeling. Clearly, the tectum in gars receives a substantial number of nonvisual afferents from all major brain areas, most of which have been reported in other vertebrates. The functional significance of these afferent sources and their probable homologues in other vertebrate groups are discussed.

Primary retinal targets in the Atlantic loggerhead sea turtle, Caretta caretta.

Autoradiographic analysis distinguished twelve primary retinal targets in the diencephalon and the mesencephalon of the Atlantic loggerhead sea turtle, Caretta caretta. While the majority of fibers terminate contralaterally, sparse labelling is seen over ipsilateral thalamic nuclei. The dorsal optic nucleus is the most expansive retinal target in the dorsal thalamus. Four nuclei ventral and one dorsal, to the dorsal optic nucleus, receive retinal input. Before terminating in the optic tectum, labelled fibers pass through the pretectum terminating in four nuclei. Within the superficial zone of the optic tectum, three terminal zones are recognized. A distinct accessory tegmental tract separates from the main optic tract terminating in the basal optic nucleus. While such a multiplicity of retinal targets occurs among other reptiles, birds and mammals, it is presently impossible to accurately recognize visual homologies among amniotic vertebrates.

Retinal projections in the Australian lungfish.

Autoradiographic analysis of the primary retinofugal projections in the Australian lungfish reveals contralateral retinal projections to a ventral portion of the periventricular preoptic nucleus, throughout its rostrocaudal extent, and to 4 distinct terminal fields in the thalamus. Only one of these thalamic fields (t4) likely receives dendrites solely from dorsal thalamic neurons. Thalamic terminal field 1 probably receives dendrites from both dorsal and ventral thalamic neurons, and fields 2 and 3 from only ventral thalamic neurons. Contralateral retinofugal fibers terminate in the pretectum and in the superficial and central tectal zones. The central tectal terminal field is restricted to the medial one-third of the tectum. At pretectal levels a contralateral basal optic tract arises from the marginal optic tract and terminates along the lateral edge of the tegmentum, as a series of glomerular puffs, and in the rostral pole of a superficial isthmal nucleus. The Australian lungfish, unlike the African and South American lungfish, possesses ipsilateral retinal projections to all of the nuclei that receive contralateral retinal input.

The organization of monoamine-containing neurons in the brain of the sunfish (Lepomis gibbosus) as revealed by fluorescence microscopy.

The morphological organization of the monoamine-containing neurons in the brain of the sunfish (Lepomis gibbosus) was studied by means of the Falck-Hillarp histofluorescence method. No attempt was made to distinguish between norepinephrine and dopamine, both primary catecholamines (CA) yielding a similar yellow-green fluorescence after paraformaldehyde treatment. In the brain stem of this teleost fish, three groups of CA-containing neuronal somata have been found. First, there is a small collection of CA perikarya located just caudal to the obex of the fourth ventricle. The neurons of this medullo-sinal group give rise to numerous CA fibers many of which ascend within the central portion of the medulla. Intermingled with these CA fibers are some CA cells that constitute the central medullary group. The CA perikarya of this group are scattered between the levels of cranial nerves X and VIII. The tegmentum of the isthmus also contains a small group of very closely packed CA neurons. The large-sized CA cells of the isthmal group are located dorsolateral to the medial longitudinal fasciculus, partly within the periventricular gray. High densities of CA varicosities were also disclosed in various brain stem structures such as the optic tectum, the torus semicircularis and the cerebeller valvula. In addition, numerous serotonin (5-HT)-type neuronal somata were found in the raphe region of the brain stem, particularly at caudal mesencephalic, isthmal and rostral medullary levels. A large number of CA cell bodies were visualized in the sunfish hypothalamus. Most of them form two populations of small, round cells that are located along and partly within the ependymal walls of the posterior and lateral recesses of the third ventricle. These bipolar cells possess one short club-like process protruding into the ventricle and their thin ependymofugal processes contribute to the CA innervation of numerous hypothalamic regions. Large CA neurons apparently without direct CSF contact also occur in the area of nucleus posterior tuberis, at the level of the mesodiencephalic junction. Although the hypothalamic inferior lobes are devoid of CA cell bodies they are heavily innervated by CA axons. The sunfish telencephalon also receives a strikingly massive and complex monoaminergic innervation. Numerous CA fibers which are first observed at the level of the preoptic area, ascend through the central zone of the telencephalon and arborize profusely particularly within the medial zone of area dorsalis telencephali. Other CA fibers, as well as abundant fine 5-HT varicosities were found in the lateral zone of area dorsalis. Although the exact origin of the telencephalic CA afferents in Lepomis is not known, part of it may arise from the isthmal CA cell group which appears similar to the locus coeruleus of reptiles, birds and mammals.

Retinofugal projections in the lepidosirenid lungfishes.

Autoradiographic and silver methods indicate that the African and South American lungfishes, Protopterus and Lepidosiren, lack ipsilateral retinal projections. Contralaterally, the retina projects to the preoptic nucleus of the hypothalamus, to four discrete areas located in the lateral neuropil of the thalamus, to a superficial pretectal neuropil, to the upper half of the tectal neutropil, and to a laterally situated basal optic neuropil located in the rostral tegmentum. The overall pattern of the primary retinofugal projections is markedly similar to that of amphibians which suggests that lungfishes may be more closely related to amphibians than to actinopterygian fishes. Neotenic trends in both lepidosirenid lungfishes and urodeles may be expressions of parallelism, hence Latimeria and Neoceratodus must be examined to resolve this phylogenetic problem. A 300-fold range in the size of the eye, indicated by the number of ganglion cells present, occurs among lungfishes, salamanders and frogs. This variation may have implications for recognizing the morphological expression of selection operating on the visual systems of lepidosirenids and amphibians.

Telencephalic efferents of the tiger salamander Ambystoma tigrinum tigrinum (Green).

The efferent projections of the telencephalon in the tiger salamander were examined by the Nauta and Fink-Heimer methods following unilateral hemispherectomies, rostral hemispheric ablations and pallial lesions. The cerebral hemisphere connects with most areas of the contralateral hemisphere via the pallial, anterior and habenular commissures. The descending fibers travel in the medial and lateral forebrain bundles and in the tracts comprising the stria medullaris. Degenerating fibers and terminals were present throughout the diencephalon but were more abundant ipsilaterally. Fibers reach the pretectum and optic tectum via dorsal and ventral pathways. There is a heavy projection to the midbrain tegmentum and a sparse projection to the tectum via the ipsilateral lateral forebrain bundle. This tract continues into the medulla oblongata and the cervical spinal cord. Rostral and dorsal hemispheric ablations revealed that the majority of fibers forming the olfacto-peduncular tract originate in the ventral, rostral one-third of the hemisphere. It was also determined that the majority of the descending efferent fibers located in the lateral forebrain bundle originate from the caudal lateral hemispheric wall, and that these fibers form connections characteristic of mammalian corticofugal and striatofugal systems. The cytoarchitecture and connections of the caudal lateral hemispheric wall suggest that it is homologous to parts of motor isocortex and amygdala of amniotes.