Intermediate And Medial Hyperstriatum Ventrale Research Articles

Early stages of memory formation in filial imprinting: Fos-like immunoreactivity and behavior in the domestic chick

Early stages of memory formation in filial imprinting were studied in domestic chicks. Chicks trained for 15 min showed strong imprinting, demonstrated by a strong preference for their training stimulus, and the time course of this preference over 2 days after training was similar to that of chicks trained for 60 min. The chicks therefore learned characteristics of the training stimulus very early during training. The intermediate and medial hyperstriatum ventrale (IMHV) is a part of the chick forebrain that is crucial for imprinting. Previous experiments have shown a learning-specific increase in Fos-like immunoreactivity, used as a marker of neuronal activity, in the IMHV after training for 60 min. The time course of Fos expression in the IMHV was measured after training for 15 min and 60 min. The same pattern of expression was found for both training times, showing a peak 120 min after the start of training. The time course of expression was stimulus-dependent. Fos expression in the IMHV, but not the hippocampus, was significantly correlated with strength of imprinting. It is concluded that the learning-specific change in Fos expression in the IMHV is associated with very early components of memory formation.

GABA, taurine and learning: release of amino acids from slices of chick brain following filial imprinting

The intermediate and medial hyperstriatum ventrale (IMHV) is a forebrain region in the domestic chick that is a site of information storage for the learning process of imprinting. We enquired whether imprinting is associated with learning-related increases in calcium-dependent, potassium-stimulated release of neurotransmitter amino acids from the IMHV. Chicks were hatched and reared in darkness until 15–30 h after hatching. They then either remained in darkness or were trained for 2 h by exposure to an imprinting stimulus. One hour later, the chicks were given a preference test and a preference score was calculated from the results of this test, as a measure of imprinting. Chicks were killed 2 h after training. Slices from the left and right IMHV of trained and untrained chicks were superfused with Krebs’ solution either with or without calcium and the superfusate assayed for arginine, aspartate, citrulline, GABA, glutamate, glycine and taurine using high-performance liquid chromatography. For calcium-containing superfusates from the left IMHV, preference score was significantly correlated with potassium-stimulated release of (i) GABA ( r=0.51, 23 d.f., P=0.008) and (ii) taurine ( r=0.77, 23 d.f., P<0.0001). There was no significant difference between the mean values of trained and untrained chicks for either compound. However, examination of the variance of the data indicated that release of both GABA and taurine increased as a result of learning. No significant correlation between preference score and release was found for any of the amino acids from the right IMHV, nor for control tissue from the left IMHV superfused with calcium-free solution. These results demonstrate that the learning process of imprinting is associated with increases in releasable pools of GABA and taurine and/or membrane excitability in the left IMHV.

Neural cell adhesion molecules, CaM kinase II and long-term memory in the chick.

The intermediate and medial hyperstriatum ventrale (IMHV) of the chick brain is a site of recognition memory for filial imprinting. Previous results have demonstrated learning-related changes in the amounts of the three major isoforms of neural cell adhesion molecule (NCAM) in the left IMHV. The increases were present 24 h after training. The present study enquired whether the increases persisted and were present 48 h after training. The brain regions analysed were the left and right IMHV and the left and right hyperstriatum accessorium (HA), a visual projection area. The alpha-subunit of calcium/calmodulin protein kinase II (CaMKIIalpha) was also assayed. There were significant correlations between a measure of the strength of learning and the amount of NCAM 180 in the right IMHV (r = +0.65; p = 0.012) but not in the left, and in the left HA (r = -0.61; p = 0.02), but not in the right. There were no learning-related changes for CaMKIIalpha. We conclude that in IMHV the effects of imprinting on NCAM 180 are expressed mainly in the left IMHV 24 h after training, but 48 h after training are expressed mainly in the right IMHV.

Neural cell adhesion molecules, learning, and memory in the domestic chick.

The intermediate and medial hyperstriatum ventrale (IMHV) of the chick forebrain is a site of recognition memory for the learning process of imprinting. The results reported here demonstrate that neural cell adhesion molecules (NCAMs) play a time-dependent role in this recognition memory. Dark-reared chicks were trained, tested, and assigned a preference score as a measure of learning. Chicks with high preference scores were designated good learners and those with lower preference scores, poor learners. Controls were untrained. Tissue was removed, 9.5 hr or 24 hr after training, from the left and right IMHV, hyperstriatum accessorium, and posterior neostriatum. Three major NCAM isoforms (180, 140, and 120 kDa) were assayed. At 24 hr only, there was in left IMHV significantly more NCAM (for each isoform) in good learners than in the other 2 groups, and also a significant correlation between the amounts of NCAM and preference scores for all isoforms; the amount predicted by each regression line at preference score 50 (no learning) did not differ significantly from the mean value for untrained controls. There were no learning-related effects in either the hyperstriatum accessorium or the posterior neostriatum.

Distinct mechanisms for expression of Fos-like immunoreactivity and synaptic potentiation in telencephalic hyperstriatum of the quail chick

In the intermediate and medial hyperstriatum ventrale (IMHV), a telencephalic region essentially involved in the initial processes of early learning tasks in poultry chicks, induction of an immediate early gene c- fos correlates significantly with the degree of learning (K.V. Anokhin, R. Mileusnic, I.Y. Shamakina, S.P.R. Rose, Effects of early experience on c- fos gene expression in the chick forebrain, Brain Res. 544 (1991) 101–107; B.J. McCabe, G. Horn, Learning-related changes in Fos-like immunoreactivity in the chick forebrain after imprinting, Proc. Natl. Acad. Sci. USA 91 (1994) 11417–11421). In slices of IMHV in vitro, on the other hand, tetanic stimulation at a low frequency induces a potentiation of synaptic responses (P.M. Bradley, B.D. Burns, A.C. Webb, Potentiation of synaptic responses in slices from the chick forebrain, Proc. R. Soc. Lond. B. 243 (1991) 19–24; T. Matsushima, K. Aoki, Potentiation and depotentiation of DNQX-sensitive fast excitatory synaptic transmission in telencephalon of the quail chick, Neurosci. Lett. 185 (1995) 179–182). In this study, we have examined a possible causal link between these two forms of activity-dependent processes, c- fos expression and synaptic potentiation. C- fos was visualized immunohistochemically using antibody raised against the Fos-protein, and potentiation was evaluated on the basis of field potential responses to local electrical stimulation. Tetanic stimulation (5 Hz×300 pulses) was required for potentiation, but not for c- fos expression. Conversely, a negative correlation appeared between them, and slices with relatively high density of Fos-like immunoreactive cells around the stimulation site failed to show potentiation. Furthermore, drugs similarly effective in blocking potentiation (such as AP5 (NMDA receptor antagonist) and bicuculline (GABA A receptor antagonist)) had different effects on the c- fos induction. While AP5 had minor, if any, effects on c- fos expression, bicuculline enhanced it selectively around the site of stimulation. Our results suggest that these two processes are basically distinct, and could represent different aspects in the formation of memory traces in IMHV.

Differential distribution of protein kinase C (PKCαβ and PKCγ) isoenzyme immunoreactivity in the chick brain

Protein kinase C (PKC) is involved in neural plasticity. The phosphorylation of the myristoylated alanine-rich protein kinase C substrate (MARCKS) in the left intermediate and medial hyperstriatum ventrale (IMHV) of the chick brain has been shown previously to correlate significantly with the strength of learning in filial imprinting. The distribution of PKCα,βI,βII and PKCγ in the brain of 1-day-old dark-reared chicks was determined immunocytochemically, using the monoclonal antibodies MC5 and 36G9, raised against purified PKCαβ and PKCγ, respectively. PKCγ-stained cells were distributed widely in the telencephalon, including all hyperstriatal structures (including the IMHV), the hippocampus, neostriatum, ectostriatum and archistriatum. There were fewer stained cells in the septum and the least cellular staining was in the paleostriatum primitivum. Fluorescent double-labelling with neuron-specific enolase (NSE) and with the glial calcium-binding protein S100 suggested that PKCγ immunoreactivity was present in neurones but not in glia. The distribution of PKCαβ-stained cells was more limited, with staining in the archistriatum, hippocampus and septum but not in the hyperstriatum. However, there was PKCαβ-staining of some fibres in the IMHV (but little elsewhere in the hyperstriatum ventrale), in the neostriatum, paleostriatal complex and the lobus parolfactorius. Double-labelling with NSE and S100 revealed PKCαβ/S100-positive glial cells present in the paleostriatal region only. There was some PKCαβ-staining of putative neurones in the hippocampus, septum and archistriatum. The differential distribution of PKC isoenzymes suggests that in the IMHV some axonal inputs contain PKCαβ whereas some postsynaptic cells contain the γ form of PKC.

Morphological changes associated with stages of memory formation in the chick following passive avoidance training

Memory formation following learning is presumed to result from modification in the efficacy of neural circuitry, either through strengthening of pre-existing synapses, or formation of new contacts. An ideal paradigm to investigate memory formation is one-trial passive avoidance training of day-old chicks, in which the birds learn to avoid pecking a bead coated with an aversive substance, methylanthranilate. Following training, a sequence of biochemical, electro-physiological, pharmacological and morphological events takes place within two loci in the forebrain, the intermediate and medial hyperstriatum ventrale (IMHV), and part of the paleostriatal complex, the lobus parolfactorius (LPO). Our data reviewed here suggest that the initial acquisition of memory involves population changes in the fine spatial organization of synaptic vesicles and active zones in synapses in the IMHV whereas longer-term changes are more prominent in the LPO and involve, primarily, a bilateral increase in the density of synapses and dendritic spines. The short-term synaptic changes are as dynamic as the molecular changes which have hitherto been considered the preserve of short-term correlates of memory formation.

Training in chicks alters PSA-N-CAM distribution in forebrain cell membranes.

The intermediate and medial hyperstriatum ventrale (IMHV) in the chick is involved in memory formation following one-trial passive avoidance training. Possible links between neural cell adhesion molecule (N-CAM) distribution and memory consolidation were examined in an immunoelectron microscope study of IMHV 6 h after training. An antibody against alpha 2,8 polysialic acid (PSA), characteristic for embryonic N-CAM, and one against the protein backbones of N-CAM were labelled (post-embedding), separately, with 15 nm immunogold, and binding to their epitopes was analysed using the statistics of point processes. No difference in labelling levels between control and trained chicks, for either antibodies, was found, both groups of birds showing that cell membrane regions are 10-30 times more enriched in N-CAM isoforms, and have a two-fold greater proportion of PSA-N-CAM, than tissue as a whole. However, 375-400 nm regular arrays of labelled PSA-N-CAM were revealed statistically in cell membranes of control, but not trained, chicks, which may be related to the possible involvement of such membrane PSA domains in memory-related neuroplasticity.

Learning-related alterations in the visual responsiveness of neurons in a memory system of the chick brain.

The intermediate and medial hyperstriatum ventrale (IMHV) of the chick brain is known to be essential for the learning process of imprinting. The activity of neurons was recorded from the left IMHV of 2-day-old unanaesthetized chicks while the chicks were free to move in a running wheel. The chicks were either raised in complete darkness or visually trained (imprinted) with a set duration of exposure to a visual image. The first group of these birds was trained by exposure for 100 min to a rotating red box and the second was trained by similar exposure to a rotating blue cylinder. A third group was left untrained. Training more than doubled the proportion of sites that responded to the stimulus used to train the bird, relative to the proportion of sites responsive to the other stimulus and to the proportion of sites responsive in the untrained birds; the learning-related increase was selective and highly significant. Behavioural monitoring indicated that the enhanced responsiveness could not be explained by overt differences in the alertness, attentiveness or movements of the birds. No significant effect of training was found on the proportion of sites responsive to a rotating stuffed jungle fowl or to the sound of a maternal call. The response at certain sites selectively signalled the presence of the training stimulus, while at others the response showed generalization across stimulus shape or colour. There was a non-specific effect of training upon the pattern of spontaneous discharges of the neurons: the numbers of spikes occurring in clusters (bursts) was significantly reduced in trained birds compared with the dark reared controls.

Object Recognition and Sensitive Periods: A Computational Analysis of Visual Imprinting

Using neural and behavioral constraints from a relatively simple biological visual system, we evaluate the mechanism and behavioral implications of a model of invariant object recognition. Evidence from a variety of methods suggests that a localized portion of the domestic chick brain, the intermediate and medial hyperstriatum ventrale (IMHV), is critical for object recognition. We have developed a neural network model of translation-invariant object recognition that incorporates features of the neural circuitry of IMHV, and exhibits behavior qualitatively similar to a range of findings in the filial imprinting paradigm. We derive several counter-intuitive behavioral predictions that depend critically upon the biologically derived features of the model. In particular, we propose that the recurrent excitatory and lateral inhibitory circuitry in the model, and observed in IMHV, produces hysteresis on the activation state of the units in the model and the principal excitatory neurons in IMHV. Hysteresis, when combined with a simple Hebbian covariance learning mechanism, has been shown in this and earlier work (Földiák 1991; O'Reilly and McClelland 1992) to produce translation-invariant visual representations. The hysteresis and learning rule are responsible for a sensitive period phenomenon in the network, and for a series of novel temporal blending phenomena. These effects are empirically testable. Further, physiological and anatomical features of mammalian visual cortex support a hysteresis-based mechanism, arguing for the generality of the algorithm.

Short-term changes in the numerical density of synapses in the intermediate and medial hyperstriatum ventrale following one-trial passive avoidance training in the chick

Previous ultrastructural studies using stereological counting techniques, based on assumptions regarding shape, size, and orientation of synapses, have suggested synaptic remodeling occurred at least 24 hr after one-trial passive avoidance training in day-old chicks. The present study estimates the mean synaptic density (Nv syn) in a region of the chick forebrain known to be involved in memory formation, the intermediate and medial hyperstriatum ventrale (IMHV), 1 and 24 hr following one-trial passive avoidance training. A stereological technique, the "disector," that makes no assumptions about size, shape, and orientation of synapses was used in the synaptic analyses. The density of axospinous synapses increased by approximately 77% at 1 hr posttraining in the right IMHV of chicks (M-trained) that learned to avoid a bitter-tasting bead, compared to those (W-trained controls) that peck a water-coated bead. A measure of the postsynaptic density size, the mean projected height of synapses (H), was 57% smaller 1 hr posttraining in the right IMHV of M-trained chicks. These differences were not found at 24 hr posttraining. We suggest that structural modification of synapses may be a key part of the processes involved in short-term memory formation.

Spatial re-arrangement of the vesicle apparatus in forebrain synapses of chicks 30 min after passive avoidance training

A quantitative ultrastructural study of synapses was carried out in the forebrain IMHV (intermediate and medial hyperstriatum ventrale) of 1-day-old chicks 30 min after training to avoid pecking at a bead coated with methyl anthranilate. In 10 birds (5 control and 5 trained), the length, curvature and the number of the synaptic active zone profiles were measured, and the active zone profile length was observed to increase in trained chicks. The spatial arrangement of synaptic vesicles with respect to the active zone was examined using a statistical stereological approach. This showed that vesicles are not located uniformly but accumulate in two spatial pools which appear to rearrange following training, with a greater number of vesicles near the active zone. These data may reflect subtle changes in the functional efficacy of synapses in the IMHV in the initial phases of memory formation.

Co-expression of Fos immunoreactivity in protein kinase (PKCγ)-positive neurones: quantitative analysis of a brain region involved in learning

The expression of the gamma protein kinase C isoenzyme (PKCγ) and of the c- fos immediate early gene protein product Fos in the intermediate and medial hyperstriatum ventrale (IMHV) of day-old chicks was determined immunocytochemically. Previous research has shown that (a) there is a learning-related increase in the expression of Fos in the IMHV of the chick after imprinting; (b) PKCγ is expressed in neurones in most regions of the chick forebrain, including the IMHV. In the present study it was found that in imprinted chicks, 96.5% of neurones in the IMHV that expressed Fos also stained positively for PKCγ. These results raise the possibility of a functional connection between PKCγ activation and c- fos expression in neurones in general, and in particular in neurones in the IMHV that are involved in learning.

Quantitative Autoradiographic Demonstration of Changes in Binding to NMDA-sensitive [3H]Glutamate and [3H]MK801, but not [3H]AMPA Receptors in Chick Forebrain 30 min After Passive Avoidance Training.

Day-old domestic chicks (Gallus domesticus) were trained on a one-trial passive avoidance task in which the aversive stimulus was an unpleasant tasting substance, methyl anthranilate. Thirty minutes later, localization of N-methyl-d-aspartic acid (NMDA)-sensitive [3H]glutamate receptor binding sites, [3H]MK801 and [3H]AMPA binding sites in 17 regions of the forebrain of methylanthranilate-trained and control (water-trained) chicks was determined using quantitative receptor autoradiography. Significant differences in binding to both MK801- and NMDA-sensitive glutamate receptors, but not alpha-amino-3-hydroxy-5-methyl-4-isoxazolepropionic acid (AMPA) receptors, were found in three regions of the forebrain of trained compared to control chicks; two of these regions have been implicated from previous lesion, biochemical and morphological studies as playing a key role in the process of memory formation and storage following passive avoidance training. For NMDA-sensitive [3H]glutamate receptors, significant elevations in binding were observed in two regions, the left intermediate and medial hyperstriatum ventrale (IMHV) (39%) and the lobus parolfactorius (LPO) (34%), at 30 min post-training, but a decrease (44%) occurred in binding to the lateral neostriatum. Significant increases in binding to MK801 receptors were observed in the left IMHV (19%) and right IMHV (13%), and left LPO (22%) at 30 min post-training, though there was a decrease in the right LPO (15%). These findings, coupled with those described in a previous paper from our group (Burchuladze and Rose, Eur. J. Neurosci., 4, 533 - 538, 1992), demonstrate that a glutamate receptor subtype is involved in learning and memory formation in the chick.

Quantitation of synaptic, neuronal and glial development in the intermediate and medial hyperstriatum ventrale (IMHV) of the chick Gallus domesticus, pre- and post-hatch

A quantitative analysis was made of the development of synapses, neurons and glia in both left and right intermediate and medial hyperstriatum ventrale (IMVH) of the forebrain of the domestic chick, Gallus domesticus from 16 days in ovo to 9 days post-hatch. There was a marked increase in total synapse numerical density ( N νsyn , from 10 synapses per 100 μm 3 at 16 days in ovo to 50 synapses per 100 μm 3 at 9 days post-hatch; no significant left/right hemispheric differences were evident but there were differences between the development profiles for synapses with asymmetric as compared to symmetrical synaptic junctions. In contrast to synaptic development, the number of neurons per unit volume, N νneu , decreased by approximately 50% from the value at 16 days in ovo to that at day 0 (hatching). The neuronal population density remained unchanged to 3 days post-hatch and then, in 9-day-old birds, declined to almost a quarter of the original population size; no significant left/right hemisphere differences were evident. Glial cells declined in number from 16 days in ovo to 19 days in ovo, and then gradually increased to 9 days post-hatch. When the synapse to neuron ratio was examined, a trend was observed of a gradual increase with age, but no hemispheric differences were present. It is concluded that these changes are major events which must be considered in experiments aimed at determining the effects of behavioural, and/or environmental manipulations, on the morphology of the IMHV because they may mask other, more subtle, structural changes that occur as a result of behavioural experiences.

Classical conditioning as an animal model for analyzing characteristics of information processing - an evoked potential study

Early stages of memory formation in filial imprinting were studied in domestic chicks. Chicks trained for 15 min showed strong imprinting, demonstrated by a strong preference for their training stimulus, and the time course of this preference over 2 days after training was similar to that of chicks trained for 60 min. The chicks therefore learned characteristics of the training stimulus very early during training. The intermediate and medial hyperstriatum ventrale (IMHV) is a part of the chick forebrain that is crucial for imprinting. Previous experiments have shown a learning-specific increase in Fos-like immunoreactivity, used as a marker of neuronal activity, in the IMHV after training for 60 min. The time course of Fos expression in the IMHV was measured after training for 15 min and 60 min. The same pattern of expression was found for both training times, showing a peak 120 min after the start of training. The time course of expression was stimulus-dependent. Fos expression in the IMHV, but not the hippocampus, was significantly correlated with strength of imprinting. It is concluded that the learning-specific change in Fos expression in the IMHV is associated with very early components of memory formation.

Mechanisms of information storage after imprinting in the domestic chick

Early stages of memory formation in filial imprinting were studied in domestic chicks. Chicks trained for 15 min showed strong imprinting, demonstrated by a strong preference for their training stimulus, and the time course of this preference over 2 days after training was similar to that of chicks trained for 60 min. The chicks therefore learned characteristics of the training stimulus very early during training. The intermediate and medial hyperstriatum ventrale (IMHV) is a part of the chick forebrain that is crucial for imprinting. Previous experiments have shown a learning-specific increase in Fos-like immunoreactivity, used as a marker of neuronal activity, in the IMHV after training for 60 min. The time course of Fos expression in the IMHV was measured after training for 15 min and 60 min. The same pattern of expression was found for both training times, showing a peak 120 min after the start of training. The time course of expression was stimulus-dependent. Fos expression in the IMHV, but not the hippocampus, was significantly correlated with strength of imprinting. It is concluded that the learning-specific change in Fos expression in the IMHV is associated with very early components of memory formation.