Dark-reared Chicks Research Articles

Functional Validation of the Gallus Chick's Response to the Maternal Food Call1

The term functional validation has been used to describe situations in which fully developed behaviours require appropriate environmental support during an early sensitive period if the initial level of responsiveness is to be maintained. The primary objective of the present study was to determine if an age-related decline in responsiveness to the maternal feeding call occurred in Gallus chicks and, if so, whether or not brief exposure to the call during an early responsive period would maintain responsiveness beyond that period. A decline in responsiveness to the call was observed beyond three days of age in dark-reared chicks. Exposure to the call for 7.5 min before three days of age was adequate to maintain responsiveness at five days of age.

Imprinting modulates processing of visual information in the visual wulst of chicks

BackgroundImprinting behavior is one form of learning and memory in precocial birds. With the aim of elucidating of the neural basis for visual imprinting, we focused on visual information processing.ResultsA lesion in the visual wulst, which is similar functionally to the mammalian visual cortex, caused anterograde amnesia in visual imprinting behavior. Since the color of an object was one of the important cues for imprinting, we investigated color information processing in the visual wulst. Intrinsic optical signals from the visual wulst were detected in the early posthatch period and the peak regions of responses to red, green, and blue were spatially organized from the caudal to the nasal regions in dark-reared chicks. This spatial representation of color recognition showed plastic changes, and the response pattern along the antero-posterior axis of the visual wulst altered according to the color the chick was imprinted to.ConclusionThese results indicate that the thalamofugal pathway is critical for learning the imprinting stimulus and that the visual wulst shows learning-related plasticity and may relay processed visual information to indicate the color of the imprint stimulus to the memory storage region, e.g., the intermediate medial mesopallium.

Early visual experience affects postnatal auditory responsiveness in bobwhite quail (Colinus virginianus).

Results from 6 experiments suggest perinatal visual experience interferes with postnatal auditory responsiveness in bobwhite quail (Colinus virginianus). Light- or dark-reared control chicks responded similarly to bobwhite maternal Calls A or B following hatching. Light-reared chicks that experienced 10 min/hr of Call A or B from hatching until testing preferred the familiar call at Days 2-4, but dark-reared chicks preferred the familiar call at all ages. Increased amounts of exposure to maternal calls during postnatal Days 1-2 led to auditory responsiveness deficits in light-reared chicks. Similarly, embryos exposed to 10 min/hr of prenatal visual cues required 48 hr of postnatal Call A or B exposure to demonstrate auditory discrimination. These findings highlight the linkages between developing auditory and visual systems during infancy.

NMDA receptor antagonists extend the sensitive period for imprinting

Filial imprinting in the domestic chick occurs during a sensitive period of development. The exact timing of this period can vary according to the methods used to measure imprinting. Using our imprinting paradigm, we have shown that normal, dark-reared chicks lose the ability to imprint after the second day post-hatching. Further, we reported that chicks treated 10 h after hatching with a mixture of the noncompetitive NMDA receptor antagonist ketamine (55 mg/kg) and the α 2-adrenergic receptor agonist xylazine (6 mg/kg) were able to imprint on day 8 after hatching, whereas controls treated with saline did not imprint. We now show that the effect of the ketamine–xylazine mixture can be mimicked by treating chicks with ketamine alone or with another noncompetitive NMDA receptor antagonist, MK-801 (5 mg/kg). Treating chicks with a single dose of ketamine (55 mg/kg) or with a single dose of xylazine (6 mg/kg) failed to produce the effect on the sensitive period. However, prolonging the action of ketamine by treating chicks with two doses of ketamine (at 10 and 12 h after hatching) did allow imprinting on day 8. In contrast, prolonging the action of xylazine had no effect on the sensitive period for imprinting. Chicks treated with MK-801 were also able to imprint on day 8. Thus, we have evidence that the NMDA receptor system is involved in the mechanisms that control the sensitive period for imprinting.

Early learning and the development of filial preferences in the chick

Newly hatched domestic chicks ( Gallus gallus domesticus) rapidly form a social preference for a conspicuous stimulus to which they are exposed. The learning process involved is known as filial imprinting. When chicks are exposed to an audio-visual compound stimulus, both auditory and visual learning are enhanced. The enhancement of visual imprinting is virtually abolished when chicks are exposed separately to the auditory element, either before or after training with the audio-visual compound. Simultaneous exposure to the two elements of the compound is superior to sequential exposure in achieving the enhancement of visual learning. These results are unlike Pavlovian conditioning, but are consistent with an interpretation of imprinting as a form of within-event learning, where links are formed between the representations of the elements of the compound, that can be weakened by separate exposure to an element. Apart from imprinting, chicks may show a developing predisposition to approach stimuli resembling conspecifics. The predisposition emerges in dark-reared chicks given some non-specific experience during a sensitive period, and is expressed as a relatively general preference for stimuli with a head and neck region. In the natural situation, the animal’s response may be biased by the predisposition, and through imprinting it then learns the characteristics of individuals.

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.

Pharmacological Extension of the Sensitive Period for Imprinting in Gallus domesticus

Parsons, C. H. and L. J. Rogers. Pharmacological extension of the sensitive period for imprinting inGallus domesticus. Physiol Behav 62(6) 1303–1310, 1997.—Precocial animals, such as the chick, exhibit a form of learning termed filial imprinting. The chick’s sensitive period for filial imprinting is restricted to the first few days after hatching. The neural mechanism that terminates the sensitive period is not fully understood. It is thought to be an experience-dependent event because once a chick has imprinted, it will not readily imprint on another stimulus. However, even dark-reared chicks eventually lose the ability to imprint, which suggests that the ending of the sensitive period may not be entirely experience-dependent. The present study investigates factors that may contribute to the ending of the sensitive period. In our experiments, dark-reared chicks were unable to imprint after Day 2 posthatching, but chicks treated 10 h after hatching with an intramuscular injection of the noncompetitive N-methyl- d-aspartate (NMDA) receptor antagonist ketamine (55 mg/kg) and the α 2-adrenoceptor agonist xylazine (6 mg/kg) (KX) imprinted on a stuffed hen 8 days after hatching. Similarly treated chicks did not imprint on a red and black box, although the box was an effective imprinting stimulus for Day 2 chicks. Chicks treated with KX at 20 or 40 h posthatching or on Day 4 or 7 as well as controls treated with pyrogen-free saline were unable to imprint on Day 8.

Delayed Induction of a Filial Predisposition in the Chick after Anaesthesia

BOLHUIS, J. J. AND G. HORN. Delayed induction of a filial predisposition in the chick after anaesthesia. PHYSIOL BEHAV 62(6) 1235–1239, 1997.—In domestic chicks, Gallus gallus domesticus, early filial preferences are formed as the result of at least two processes: learning by exposure to conspicuous objects (filial imprinting) and a developing predisposition to approach some stimuli rather than others. The predisposition may be measured in visually naive chicks, e.g. as a preference for a rotating stuffed jungle fowl hen rather than for a rotating red box. Non-specific experience, such as handling or being placed in a running wheel in darkness, for a short time, is sufficient to induce the predisposition. In Experiment 1, the existence of a sensitive period for the induction of the predisposition, between approximately 14 and 42 h after hatching, was confirmed. The putative effect of anaesthesia on the induction of the predisposition was investigated in Experiment 2. Soon after hatching dark-reared chicks received injections of the anaesthetic equithesin (0.12 ml, IP), or saline, or they did not receive injections (controls). The chicks were subsequently placed in running wheels for a total of 2 h at either 24 h or 48 h after hatching. The following day the chicks’ preferences were tested. In the 24 h-chicks, the saline and control groups showed a significant mean preference for the stuffed fowl, whilst the equithesin group did not. In contrast, in the 48 h-groups of chicks, only the equithesin group showed a significant mean preference for the stuffed fowl. These results suggest that the anaesthetic equithesin affects a developing filial predisposition in the domestic chick, and does so by delaying the onset of a sensitive period for the induction of the predisposition.

Clathrin proteins and recognition memory

Strong converging evidence indicates that the intermediate and medial part of the hyperstriatum ventrale (IMHV) of the chick forebrain is a site of recognition memory for the learning process of imprinting. Clathrin proteins have been implicated in synaptic plasticity. In the present study we demonstrate for the first time that they are involved in vertebrate learning. Chicks were trained by exposure to a conspicuous object and their preference for it versus a novel object subsequently measured as a preference score (an index of learning). Trained chicks with low preference scores were classed as “poor learners” and those with high preference scores as “good learners”. An additional group of chicks was untrained (“dark-reared”). Tissue was removed from the left and right IMHV, hyperstriatum accessorium and posterior neostriatum 9.5 h or 24 h after training. Clathrin heavy chain and clathrin light chains a and b were assayed using sodium dodecyl sulphate–polyacrylamide gel electrophoresis and immunoblotting. In the IMHV, and only for clathrin heavy chain, was there a significant effect of training. The effect occurred 24 h but not 9.5 h after training, and was significant only in the left IMHV. In this region at 24 h, there was (i) significantly more clathrin heavy chain in good learners than in dark-reared chicks, and (ii) a significant positive correlation between the amount of clathrin heavy chain and preference score; the amount of protein present in the dark-reared chicks did not differ significantly from the amount predicted from the regression line for trained chicks performing at chance (preference score 50). These findings imply that for the left IMHV, visual experience per se, locomotor activity and other side effects of training did not affect the amount of clathrin heavy chain. Rather, the increase observed was a function of the amounts chick learned and, because it was delayed, is likely to be involved in long-term memory. The results for clathrin heavy chain taken together suggest that enhanced presynaptic events in the IMHV, possibly associated with an increase in synaptic vesicle release/uptake, are important in the recognition memory underlying imprinting.

2240 Cells containing c-fos-like protein in the rat brain after stimulation on the external membrane of the uterus

The concentrations of adrenaline, dopamine and noradrenaline were measured in 3 regions of the domestic chick telencephalon: (a) the Wulst; (b) a medial forebrain sample comprising mainly the intermediate part of the medial hyperstriatum ventrale (IMHV); and (c) a basal forebrain sample comprising mainly paleostriatum augmentatum. There was no significant left/right hemispheric asymmetry in the concentration of any of these catecholamines in any region studied. Adrenaline was undetectable in the Wulst and medial forebrain samples and only trace amounts were found in the basal forebrain samples of 1-day-old, light-reared chicks. Dopamine concentrations of 9.13 ± 1.13 (S.E.M.) ng/g were present in the Wulst, 16.66 ± 2.56 ng/g in the medial forebrain and 121.19 ± 33.06 ng/g in the basal forebrain samples at hatching. These levels did not alter with age or with visual experience of an imprinting stimulus during the first 50 h post-hatch. At hatching, noradrenaline concentrations of 35.83 ± 8.61 ng/g were present in the Wulst, 26.09 ± 3.75 ng/g in the medial forebrain and 53.13 ± 7.85 in the basal forebrain samples. The noradrenaline concentrations in the Wulst and medial forebrain samples increased significantly over the first 50 h post-hatch in dark-reared chicks. Visual experience increased noradrenaline levels in all 3 regions of the telencephalon studied.

Cells containing C-fos-like protein in the rat brain after stimulation on the external membrane of the uterus

The concentrations of adrenaline, dopamine and noradrenaline were measured in 3 regions of the domestic chick telencephalon: (a) the Wulst; (b) a medial forebrain sample comprising mainly the intermediate part of the medial hyperstriatum ventrale (IMHV); and (c) a basal forebrain sample comprising mainly paleostriatum augmentatum. There was no significant left/right hemispheric asymmetry in the concentration of any of these catecholamines in any region studied. Adrenaline was undetectable in the Wulst and medial forebrain samples and only trace amounts were found in the basal forebrain samples of 1-day-old, light-reared chicks. Dopamine concentrations of 9.13 ± 1.13 (S.E.M.) ng/g were present in the Wulst, 16.66 ± 2.56 ng/g in the medial forebrain and 121.19 ± 33.06 ng/g in the basal forebrain samples at hatching. These levels did not alter with age or with visual experience of an imprinting stimulus during the first 50 h post-hatch. At hatching, noradrenaline concentrations of 35.83 ± 8.61 ng/g were present in the Wulst, 26.09 ± 3.75 ng/g in the medial forebrain and 53.13 ± 7.85 in the basal forebrain samples. The noradrenaline concentrations in the Wulst and medial forebrain samples increased significantly over the first 50 h post-hatch in dark-reared chicks. Visual experience increased noradrenaline levels in all 3 regions of the telencephalon studied.

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.

Induction and Development of a Filial Predisposition in the Chick

Abstract Filial preferences in young domestic chicks, Gallus gallus domesticus, are influenced by at least two systems: one involved in learning about stimuli to which the animals are exposed, and the other a predisposition to approach stimuli resembling conspecifics. The predisposition is manifest in dark-reared chicks as an emerging preference for a rotating, stuffed jungle fowl over a rotating red box. In previous studies, 24 h-old visually naive chicks were placed in running wheels for 2 h. Throughout this time the chicks were in darkness. A significant preference for the stuffed fowl was found 24 h, but not 2 h later. In the present study it was shown that the predisposition becomes manifest at 10 h (Experiment la) and 5 h (Experiment 1 b) after either placement in running wheels, or after the chicks had been trained by exposure to a rotating red box. In Experiment 2 it was shown that placement in running wheels was not necessary for the predisposition to emerge. However, handling the birds was sufficient for the induction of a significant preference, expressed 5 h later. Exposure to a maternal call for 2 h (Experiment 3) was also sufficient to induce the development of the visual predisposition 24 h later. These results show that a visual predisposition to approach certain stimuli becomes manifest in dark-reared chicks between 2 and 5 h after the experience of handling, and that exposure to a maternal call is also sufficient to induce this predisposition.

3H]MK-801 binding asymmetry in the IMHV region of dark-reared chicks is reversed by imprinting

Glutamate NMDA-type receptor binding in the intermediate medial hyperstriatum ventrale (IMHV) of dark-hatched chicks is lateralized. This lateralization was found to be markedly influenced by imprinting. In dark-reared chicks the binding of the selective ligand [ 3H]MK-801 was two-fold higher in the right IMHV than in the left IMHV. In contrast, imprinted chicks have significantly higher levels of [ 3H]MK-801 binding in the left IMHV region than in the right IMHV. Imprinting results in 2a learning-related increase in NMDA-type receptor binding levels in the left IMHV, whereas [ 3H]MK-801 binding levels in the right IMHV remain unchanged by imprinting. Thus, the plasticity present in the NMDA-type receptor system and associated with imprinting appears to occur in the left hemisphere only.

Learning-related changes in Fos-like immunoreactivity in the chick forebrain after imprinting.

The intermediate and medial part of the hyperstriatum ventrale (IMHV) is a part of the chick forebrain that is critical for the learning process of imprinting and may be a site of information storage. Chicks were either trained on an imprinting stimulus or dark-reared. Trained chicks were classified as good or poor learners by their preference score (a measure of the strength of imprinting). A monoclonal antibody against the immediate early gene product Fos was applied to sections through IMHV and other forebrain regions. In the IMHV, significantly more immunopositive nuclei were counted in good learners than in poor learners or dark-reared chicks. There was a positive correlation between counts of labeled nuclei and preference score that was not attributable to sensory activity per se, locomotor activity during training, or a predisposition to learn well; rather, the results indicated that the change in Fos immunoreactivity in the IMHV was related to learning. In the hyperstriatum accessorium, significantly fewer immunopositive nuclei were counted in good learners than in poor learners or in dark-reared chicks. In the dorsolateral hippocampal region, more immunopositive nuclei were counted in trained than in dark-reared chicks. No significant effects of training were found in the anterior hyperstriatum ventrale, lobus parolfactorius, neostriatum, medial hippocampal region, or ventrolateral hippocampal region, but counts in this last region were positively correlated with training approach. The results for IMHV implicate Fos or Fos-related proteins in memory processes and pave the way for the identification of the cell types that show the learning-related increase in gene expression.

Learning-dependent Changes in the Responses to Visual Stimuli of Neurons in a Recognition Memory System.

When young chicks are trained by exposing them to a conspicuous object they learn its characteristics. The learning process is known as imprinting. In the present study neuronal activity in a region crucial for imprinting was shown to be affected by training and by the object on which the chicks had been trained. The region is the intermediate and medial part of the left hyperstriatum ventrale (left IMHV). No such effects were found in a visual projection area, the left hyperstriatum accessorium. Domestic chicks were imprinted on either a rotating red box (n=7 chicks) or a rotating blue box (n=8). When the chicks were approximately 48 h old they were anaesthetized and multiple-unit activity was recorded in simultaneous, single penetrations through each of the two regions. Records were also made from eight dark-reared chicks. Whilst recording, the red or blue box, placed in front of the contralateral eye, was switched on to give a total of 20 rotations, the interval between each rotation being 10 s. The alternative stimulus was then presented 20 times. Unit activity in the 3 s before and after stimulus onset was compared and the data for each of the 20 presentations were combined. In the left IMHV 18 out of a total of 115 recording sites (16%) responded significantly to the stimuli; in the left hyperstriatum accessorium 39 out of 126 recording sites (26%) did so. Measures of unit activity at each recording site were combined for a given penetration to provide a 'mean penetration response'. The response to the red box differed from the response to the blue box in the left IMHV of dark-reared chicks. After training with the blue box the response to both boxes was similar to the response to the blue box in dark-reared birds. After training with the red box the response to both boxes was similar to the response to the red box in dark-reared birds. No significant effects were found in the left hyperstriatum accessorium. The two training boxes were virtually identical apart from the differences in colour and brightness. Training appeared to stabilize the response of the visually naive left IMHV to the training stimulus whilst changing its response to the alternative, but similar stimulus. That is, one consequence of training is that the two stimuli are placed in the same category, and this neural change may provide a basis for stimulus generalization. The underlying neural system is modelled and a mechanism that allows such stimuli to be discriminated is proposed.

The effect of the neurotoxin DSP4 on the development of a predisposition in the domestic chick.

Under certain conditions, dark-reared chicks preferentially approach objects resembling conspecifics. In the current study we investigated the effect of the catecholaminergic neurotoxin DSP4 on this predisposition. After hatching, chicks received an intraperitoneal injection of either DSP4 or vehicle (controls). The chicks were individually placed in running wheels for 2 x 1 hr, 24 hr before testing. The chicks were maintained in darkness until given a preference test, which involved the simultaneous presentation of an illuminated red box and a stuffed fowl at 36, 48, 60, 66, or 72 hr after hatching. The approach of each chick toward the two objects was recorded over 5 min and a preference score calculated. Control chicks placed in the running wheels at 24 and 36 hr posthatch and tested 24 hr later showed a significant preference for the fowl compared to the red box. In contrast, DSP4-treated chicks did not show such a preference at corresponding ages, but did express a greater preference for the fowl when placed in the running wheels at 42 and 48 hr posthatch and tested 24 hr later. These results suggest that experience of the running wheel or of associated factors, e.g., handling, must occur during a restricted period of time to influence the emergence of the predisposition to approach the stuffed fowl. DSP4 treatment delays the onset of this period.

The postnatal development of benzodiazepine receptor sites in the chick optic lobe is modulated by environmental lighting

The present paper describes the ability of benzodiazepin receptor sites to undergo light mediated-plastic changes during the early postnatal development of the chick optic lobe. The postnatal development pattern of these receptors was studied under different levels of light stimulation, i.e. normal-, light- and dark-rearing. At hatching the specific binding of [ 3H]Flunitrazepam was 0.23 ± 0.01 pmol/mg protein. The developmental profile shows a sharp and transient peak of receptor overexpression between the 1st and the 2nd postnatal day in the three experimental groups. Between the 2nd and the 6th day significant differences were found between the three groups, being this difference maximal during the peak of over-expression. In fact, on the 2nd day the specific [ 3H]Flunitrazepam binding showed an increase of 17% ( P < 0.0005) and a decrease of 34% ( P < 0.0005) for light- and dark-reared animals as compared with normally-reared ones. The changes in receptor density were transient since from the 6th day onward they gradually disappeared, being almost identical in the three groups by the day 15. At this moment the number of benzodiazepine receptor sites stabilized at the adult level. Scatchard analysis at the 2nd postnatal day revealed that the differences observed in the high affinity benzodiazepine binding sites between the three groups were due to modifications in the total number of binding sites while the affinity remained unchanged. The maximal number of binding sites were: 2.76 ± 0.03, 3.40 ± 0.01 and 1.46 ± 0.11 pmol/mg protein in normally-, light- and dark-reared chicks, respectively; while the apparent dissociation constants were unaffected. The present work shows that variations in the light intensity during the early postnatal life affect the postnatal developmental pattern of benzodiazepine receptor sites. This effect is maximal at the time of appearance of the peak of receptor overexpression (1–2 postnatal day) suggesting that the peak corresponds to a brief period of high plasticity for the receptor development.

Glutamic acid decarboxylase in different areas of the developing chick central nervous system

The temporal course of the development of GAD activity in GABAergic neurons was studied in the chick retina, optic lobe and cerebellum. The developmental pattern of GAD activity was similar in the three areas studied, showing typical sigmoideal curves, which reached a maximal value at the 3rd post-hatching day. Kinetic studies during development revealed that Km remained unchanged while Vmax increased 3-fold in the retina (48.99 +/- 0.84 nmol/hr/mg protein), almost 4-fold in the optic lobe (162.77 +/- 4.32 nmol/hr/mg protein) and 3.5 fold in the cerebellum (69.30 +/- 1.26 nmol/hr/mg protein). The developmental pattern of GAD activity in homogenates of the three areas studied from dark-reared and light-reared chicks with respect to normal light-dark cycle animals showed no significant differences. These results indicate that the increase in GAD activity during development are not due to a change in the affinity for its substrate but rather to changes in the concentration of the enzyme. The developmental pattern of GAD activity in the chick visual system was not affected by environmental conditions suggesting that the developmental profile is light-independent.

Neural bases of recognition memory investigated through an analysis of imprinting

Through a learning process known as imprinting, the young of some animals, including the domestic chick, come to recognize an object by being exposed to it. Visually naive chicks vigorously approach a wide range of objects. After an adequate period of exposure to one object chicks selectively approach it in a recognition test. The nervous system of dark-reared chicks is not a tabula rasa, as chicks have predispositions to approach some stimuli rather than others. Nevertheless, visual imprinting leads to changes in a nervous system that may not have been 'marked' by previous visual experience, and so encourages the hope of discovering the neural bases of the learning process. The intermediate and medial part of the hyperstriatum ventrale, a sheet of cells within the cerebral hemispheres, plays a crucial role in visual imprinting, particularly in the memory process of recognition. The cellular and sub-cellular changes that take place in this part of the hyperstriatum ventrale after imprinting are described. The right and left hyperstriatum ventrale regions play different roles in the imprinting process, and evidence is given for the existence of multiple memory systems in the chick brain.