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Layer 7 is one of the retinorecipient layers of the avian optic tectum. However, little information is available about the neuronal organization of this layer and its implications for visual function. Golgi impregnation was used to investigate the retinal input to and the neuronal architecture of layer 7 of the chick optic tectum, which forms a narrow band between the two cell-dense layers 6 and 8. Anterograde tracers were also used to investigate the afferent and efferent connections of layer 7, in both the light and the electron microscope, together with GABA immunogold labelling. Three types of radial neuron were defined according to the origin and course of their axons. The perikarya of these neurons were situated in tectal layers 10-11. The principal dendrites of these radial neurons ascended to the tectal surface and gave rise to dendritic side-branches in layer 7. These dendritic side-branches received asymmetric synapses from the terminations of retinal fibre arborisations. Type 2 radial neurons, whose axons arose from the deep pole of the perikaryon or occasionally from a basal dendrite, were shown to project to the nucleus isthmi pars magnocellularis, which has previously been demonstrated to be GABAergic and to project to glomerulus-like complexes in tectal layers 4-5. In these layers, the dendritic branches of layer 13 neurons that project to the nucleus rotundus have previously been shown to receive retinal fibre input. Therefore, the retinal input to layer 7 may be able to modulate the transmission of information to the visual thalamus, by way of a feed-back loop to layers 4-5 of the tectum involving the nucleus isthmi pars magnocellularis.

Calcitonin gene-related peptide (CGRP)-containing neurones have been implicated in the transmission of visceral sensory information to the cortex and in the control of arterial blood pressure in mammals. However, little is known about its function in other vertebrates. As a first step toward investigating the function of CGRP in birds, its distribution was studied in the domestic chick and quail brain by means of immunocytochemistry, by using antibodies against rat CGRP. The distribution of CGRP immunoreactivity in the chick and quail central nervous system was found to be similar. CGRP-immunoreactive (CGRPi) perikarya were not present in the telencephalon. In the diencephalon, CGRPi perikarya were present mainly in the shell of the thalamic nucleus ovoidalis, the nucleus semilunaris paraovoidalis, the nucleus dorsolateralis posterior thalami, and in the hypothalamic nucleus of the ansa lenticularis. In the brainstem, CGRPi perikarya were present in the nucleus mesencephalicus nervi trigemini, the nucleus tegmenti ventralis, the locus coeruleus, the nucleus linearis caudalis and in the parabrachial region. In addition CGRPi perikarya were found in the motor nuclei of the III, IV, V, VI, VII, IX, X, and XII cranial nerves. The telencephalon contained CGRPi fibres within the paleostriatal complex (mainly in the ventral paleostriatum), parts of the neostriatum and ventral hyperstriatum, parts of the archistriatum, and the septum. In the diencephalon, the densest plexus of CGRPi fibres was observed in the dorsal reticular thalamus. A less dense CGRPi innervation was present in some dorsal thalamic nuclei and in the medial and periventricular hypothalamus. The pretectum and midbrain tegmentum also contained CGRPi fibres, whereas the optic tectum was virtually devoid of immunolabelling. Scattered CGRPi fibres were observed in the central grey and neighbouring pontine areas. Some of the sensory fibres of the trigeminal, vagal, glossopharyngeal, and spinal nerves were also CGRPi. The results of comparative studies indicate that the presence of CGRP in some thalamo-telencephalic projections is a primitive feature of the forebrain of amniotes. Therefore, the brain areas giving rise to and receiving such a projection in different vertebrates, are likely to be homologous.

Neurons and fibres in the chick and homing pigeon hippocampus were described following Golgi impregnation. Two principal classes of neurons were distinguished: projection neurons with distant projecting axons and spiny dendrites, and local circuit neurons. In the homing pigeon and chicken hippocampus there are three types of projection neurons: pyramidal, pyramidal-like and multipolar. The pyramidal and pyramidal-like neurons are only found in the central 'pyramidal' layer of the hippocampus whereas multipolar neurons are present in the suprapyramidal, pyramidal and infrapyramidal layers. The axon of projection neurons typically emits several varicose collaterals from the initial section. Most of these collaterals extend along the infrapyramidal layer of the hippocampus, while others ascend to the pyramidal and suprapyramidal layers where they branch. The number of impregnated axon collaterals was higher in the homing pigeon than in the chick hippocampus. A variety of multi-angular/ovoid local circuit neurons ranging from small to large size are found in the homing pigeon and chick hippocampus. Their axons develop local arborization of varicose branches, the extent of which varies with the type of local circuit neurons. The density of GABA immunopositive local circuit neurons was found to be greater in the homing pigeon than in the chick. The profuse arborization of projection neuron axon collaterals and the higher density of GABA-immunopositive local circuit neurons in the homing pigeon hippocampus may underlie the differences in hippocampal function between the homing pigeon and chick, and this complex local connectivity may contribute to the ability of spatial orientation and memory.

The archistriatum of the domestic chick has been implicated in both fear behaviour and learning. However, relatively little is known about its organisation. The efferent connections of discrete anatomical regions of the chick archistriatum were therefore investigated by iontophoresis of the anterograde tracer Phaseolus vulgaris leucoagglutinin into its anterior, dorsal intermediate, ventral intermediate, medial, and posterior parts. The results of this study suggest that the chick archistriatum can be divided into two basic divisions according to whether they project to the following limbic structures: the hippocampal formation, septal areas, lobus parolfactorius, nucleus accumbens, ventral paleostriatum, and dorsomedial thalamus. The limbic archistriatum includes the posterior archistriatum and extends rostrally through the ventral intermediate archistriatum into the anterior archistriatum. The non-limbic archistriatum comprises the dorsal intermediate and medial archistriatum and largely gives rise to specific sensory, somatosensory, and motor telencephalofugal efferents. There may not be distinct borders between these two divisions of the chick archistriatum.

The chick archistriatum has been implicated in avoidance learning and filial imprinting. However, its role in these learning paradigms may be due to the inhibition of normal avoidance responses, since the avian archistriatum has been shown to play a role in fear/avoidance behaviour. The involvement of the archistriatum in the expression of unlearned fear/avoidance behaviour was therefore investigated in two day-old chicks. Chicks were exposed individually to a novel 'open field' for 5 min. Behaviour was recorded on videotape for analysis. In a separate but concurrent experiment, bilateral archistriatal lesions, sham archistriatal lesions or lateral cerebral lesions were made in day-old chicks which were then exposed to the 'open field' arena. At the end of each exposure a novel object was dropped near the chick. Chicks with archistriatal lesions generally displayed greater movement, more pecking behaviour and spent more time near the centre of the 'open field' than other chicks. There were no differences between the treatment groups in latencies to move or begin peeping. The behaviour of untreated chicks in the 'open field' was similar to that of sham-lesioned chicks and there was no effect of hatch on behaviour. Upon exposure to a novel object, indices of fear and avoidance were not changed in lesioned chicks. These results demonstrate that in the young chick, the archistriatum may be involved in the response to mild or intermediate levels of environment or isolation-related stress, but does not appear to be important for overt fear responses or avoidance of novel objects. Taken together with the results of previous work, the data suggests that the archistriatum may be directly involved in avoidance learning and imprinting.

The chick archistriatum receives afferents from the intermediate part of the medial hyperstriatum ventrale (IMHV) and projects to the lobus parolfactorius (LPO). There is functional evidence to suggest that the IMHV and the LPO are connected, but there is no anatomical evidence for a direct connection between the two structures. The aim of the current study was to characterize the termination pattern of medial hyperstriatal afferents within the archistriatum to determine whether the archistriatum may act as a relay between the IMHV and LPO. Following iontophoresis of Phaseolus vulgaris leucoagglutinin into the medial hyperstriatum ventrale (including the IMHV) of 1-week-old domestic chicks, anterogradely labelled fibers were observed to descend through the medial neostriatum and paleostriatum to enter the archistriatum. These medial hyperstriatum ventrale afferents arborised profusely to give varicose axon branches within all except the anterior part of the archistriatum. However, the greatest density was present in the ventral part of the intermediate archistriatum. Electron microscope examination of Phaseolus lectin immunocytochemistry and Golgi impregnation revealed that medial hyperstriatum ventrale axons formed multiple asymmetric synapses with dendritic spines (head and neck regions) on the terminal and preterminal dendritic segments of densely spiny archistriatal projection neurons. Medial hyperstriatum ventrale afferents were not observed to contact calbindin immunoreactive, presumptive "local circuit" neurons, within the archistriatum, despite a spatial overlap in their distribution. These results suggest that the archistriatum may be capable of mediating the transfer of information from the IMHV to the LPO.

The avian archistriatum has been demonstrated to play a role in agonistic behaviours and avoidance learning. However, the extent of its role in learning is unknown. The involvement of the archistriatum in the learning process of filial imprinting was therefore investigated in day-old chicks. Bilateral archistriatal lesions, lateral cerebral area lesions or sham archistriatal penetrations were made in dark-reared, day-old chicks, which were subsequently exposed to either a rotating red box or blue cylinder for 2 x 1 h training sessions. Three hours later, the approach of chicks to their training object and to the other, novel object was measured. Chicks with archistriatal lesions ran a similar distance towards each stimulus and therefore failed to display a preference for their training object. However, chicks with sham archistriatal penetrations or lateral cerebral area lesions exhibited a significant preference for the object they had been trained upon. These results demonstrate that the archistriatum is essential for the expression of an imprinted preference. All chicks approached their training object significantly more on their second compared to their first training exposure, suggesting that some aspects of imprinting behaviour remain intact in chicks with archistriatal lesions. Taken together with the results of previous work, the current data suggest that the archistriatum may be involved in retention of significant aspects of the imprinting experience, or in motivation to approach imprinting objects.

The projection from the chick optic tectum to the nucleus rotundus was investigated by iontophoresis of the anterograde tracer Phaseolus vulgaris lectin (PHA-L) into the optic tectum and by Golgi impregnation. PHA-L labelled fibres were observed within the ipsilateral nucleus rotundus and to course through the supraoptic decussation to the contralateral nucleus rotundus. They appeared similar to fibres evident in Golgi impregnated material. Before reaching the nucleus rotundus, PHA-L labelled fibres gave off short terminal branches in the caudoventral thalamic nucleus. Within the ipsilateral nucleus rotundus, the labelled fibres formed an extensive fibre net with a dense core and a less dense periphery. PHA-L labelled fibres in the contralateral nucleus rotundus exhibited a qualitatively similar but sparser distribution. There appeared to be a topographic representation of the optic tectum within the nucleus rotundus. Electron microscopy revealed that PHA-L labelled tectal fibres established asymmetric synapses with principal neurons in the nucleus rotundus.

Investigation of the chick archistriatum revealed that it was not possible to distinguish distinct sub-regions on the basis of their Golgi architecture, despite the fact that the pigeon archistriatum has been suggested to comprise "amygdaloid" and "sensorimotor" divisions. Nine sub-types of projection neuron and four of local circuit neuron were identified in the chick archistriatum. Projection neurons were generally larger with thicker dendrites than those in other regions of the chick telencephalon. A greater proportion of these projection neurons also had highly densely spiny dendrites. These findings suggest that archistriatal neurons receive a large, possibly complex afferentation and project considerable distances. Large and medium-sized local circuit neurons with varicose axon branches and relatively large neurogliaform cells were characteristic of the archistriatum. All types of archistriatal neuron were identified in both 28 and one day old chicks. However, presumptive highly densely spiny projection neurons appeared relatively immature in the one day old chick archistriatum.

The effects of the noradrenergic neurotoxin DSP4 on imprinting in domestic chicks were investigated. Chicks received either DSP4 or distilled water and were trained 60 hr after hatching by exposing them to either a rotating red box or a stuffed jungle fowl. Noradrenaline concentration was determined in Wulst and medial and basal forebrain samples. The medial forebrain sample comprised mainly the intermediate part of the medial hyperstriatum ventrale, a region previously shown to be intimately involved in imprinting. The DSP4 treatment reduced forebrain noradrenaline levels by about 65%. DSP4 profoundly impaired imprinting in box-trained chicks but not in fowl-trained chicks. In box-trained chicks, the strength of imprinting was positively correlated with noradrenaline concentration in both medial and basal forebrain samples. It is suggested on the basis of these and previous studies that some of the neural structures and mechanisms involved in imprinting on the fowl differ from those involved in imprinting on the box.

The intermediate part of the medial component of the hyperstriatum ventrale (IMHV) of the chick telencephalon is implicated in the learning process of imprinting. The afferent connections of this region were investigated using the retrograde neuronal tracers horseradish peroxidase (HRP), nuclear yellow and fast blue. Labelled cells were found (i) ipsilaterally in the visual Wulst, neostriatum pars intermedia and caudalis, the posterior part of the hyperstriatum ventrale, the dorsal part of the archistriatum, paleostriatum augmentatum, dorsomedial part of the thalamus, and (ii) bilaterally in the hippocampus, area septalis, ventromedial part of the thalamus and the stratum album centrale of the tectum opticum. The efferent projections of the IMHV were studied using the anterograde tracer, [3H]leucine. Labelling was seen ipsilaterally over the dorsolateral part of the hemispherium telencephali, the paleostriatum augmentatum and the posterior and dorsal parts of the archistriatum. Double labelling with nuclear yellow and fast blue revealed (i) neurons in the IMHV which project to both the paleostriatum augmentatum and the archistriatum, and (ii) neurons in the hippocampus which project bilaterally to the IMHV.

The retrograde fluorescent tracers Fast Blue and Nuclear Yellow were injected into the intermediate part of the medial hyperstriatum ventrale (IMHV) of the chick telencephalon. After the appropriate survival times, the brains were processed for acetylcholinesterase (AChE) histochemistry using the thiocholine technique. In some telencephalic regions, fluorescent retrograde labelling and brown AChE reaction product were observed simultaneously in the same neurones by using a combination of ultraviolet and bright-field illumination. These AChE-positive afferents of IMHV arose from the hippocampus, septum, septal nuclei, hyperstriatum ventrale and neostriatum.