Aposematic colouration enhances memory formation in domestic chicks trained in a weak passive avoidance learning paradigm

The one-trial passive avoidance learning task is commonly used in avian research to explore anatomical, cellular and molecular parameters of learning and memory. Many factors are known to influence the effectiveness and/or duration of such learning events. Combinations of novel odours, such as pyrazine, and aposematic colours, such as brig ht yellow or red, have been shown to induce a long-lasting aversion to food crumbs in ‘visual’ predators, including birds such as the domestic chick (1). The aim of this study was to (a) examine whether visual complexity played a role in the generation of an aversive response to a novel visual stimulus and (b) to establish whether the duration of memory of an aversive experience could be modified by altering the visual properties of the stimulus. In the first experiment, naïve domestic chicks were trained on a weakly aversive one-trial passive avoidance bead task, in which chicks were allowed to peck at a bead coated with a 10% solution of the bitter-tasting and odorous substance methylanthranilate (MeA). The chicks were trained with (allowed to peck) one of four differently coloured beads dipped in 10% MeA. Chrome, black, yellow or black-and-yellow striped beads were used. ‘Recall’ of the aversive bead was examined by presenting the (clean) training bead 24 h after training and monitoring avoidance to it compared to a ‘neutral’ white bead. A high proportion (63%) of chicks trained with the black and yellow striped bead avoided it 24 h after training, whereas little or no avoidance was seen in response to chrome, yellow or black beads. In a second experiment naïve domestic chicks were all trained once only with a black and yellow striped bead coated in a 10% MeA solution, but this time, were tested 24 h later, once only, with either a black, a yellow or a black and yellow striped bead. Nearly 60% of chicks tested with a black and yellow striped bead showed avoidance of the bead, whereas only 23% of those tested with a black bead and 14% tested with a yellow bead showed avoidance. These results confirm the importance of complex warning colouration, when paired with a novel olfactory cue and a bitter taste, in avoidance learning. We conclude that the chicks’ response to monochromatic colours (e.g. yellow or black) is not affected by their previous experience with a conspicuously patterned stimulus (yellow and black stripes). Moreover, it suggests a predisposition for chicks to attend to aversive cues associated with ‘naturalistic’ high contrast colour cue combinations such as black and yellow.     

Piracetam facilitates long‐term memory for a passive avoidance task in chicks through a mechanism that requires a brain corticosteroid action

We investigated the effects of piracetam, a nootropic, on learning and memory formation for a passive avoidance task in day-old chicks. To test for the possible cognitive-enhancing properties of piracetam, a weak learning version of this task – whereby chicks maintain a memory to avoid pecking at a bead coated in a diluted aversant for up to 10 h – was used. Post-training (5, 30 or 60 min), but not pretraining, injections of piracetam (10 or 50 mg/kg, i.p.) increased recall for the task when the chicks were tested 24 h later. Because previous studies showed that long-term memory for the passive avoidance task is dependent upon a brain corticosteroid action, and because the efficacy of piracetam-like compounds is also modulated by corticosteroids, we tested whether the facilitating effect of piracetam was dependent upon a corticosteroid action through specific brain receptors (mineralocorticoid receptor and glucocorticoid receptor). First, increased plasma levels of corticosterone were found 5 min after piracetam injection. In addition, intracerebral administration of antagonists for each receptor type (RU28318, for mineralocorticoid receptors, and RU38486 for glucocorticoid receptors; i.c.) given before the nootropic inhibited the facilitative effect of piracetam on memory consolidation. These results give further support to a modulatory action of piracetam on the mechanisms involved in long-term memory formation through a neural action that, in this learning model, requires the activation of the two types of intracellular corticosteroid receptors.     

The corticosteroid synthesis inhibitors metyrapone and aminoglutethimide impair long-term memory for a passive avoidance task in day-old chicks

Long-term memory for a passive avoidance task in day-old chicks has proved to depend upon an action of the adrenal steroid corticosterone through specific receptors in a brain region, the intermediate medial hyperstriatum ventrale (IMHV), involved in learning the task. In this study, we questioned whether pretraining peripheral administration of drugs described to inhibit either basal levels of corticosterone – aminoglutethimide – or treatment-induced stimulated corticosterone secretion – metyrapone – might interfere with retention for the task at 24 h post-training. The results showed a dose-dependent effect of the inhibitors, with the highest doses tested for both drugs (10 and 50 mg/kg for metyrapone, and 50 mg/kg for aminoglutethimide) being amnestic for the task. Additional experiments, in which we studied possible effects of the inhibitors on concomitant aspects of learning (i.e., reactivity to novelty, and pecking pattern), show that the drugs did not affect general behavioural reactivity. The present results thus support the idea that training-induced corticosterone release plays a key role in the neurobiological processes that determine the establishment of a persistent memory for the aversively motivated avoidance response. In addition, they point to corticosteroid inhibitor drugs as potential tools for the study of the interactions between steroid hormones and cognition-enhancing compounds in this learning task.     

Adenosine–amino acid interactions in the chick brain: a role in passive avoidance learning

The present work describes interactions between adenosine and the amino acids glutamate and GABA in slices of intermediate medial hyperstriatum ventrale (IMHV), an area of the chick brain known to be involved in learning and memory events associated with a one-trial passive avoidance task. In slices derived from the IMHV of untrained chicks, the A₁ receptor agonist N⁶-cyclohexyladenosine (CHA; 10 μM) specifically inhibited glutamate release. Conversely, cyclopentyltheophylline (CPT; 100 μM an A₁ antagonist) increased glutamate release from the slices and blocked the CHA-induced inhibition of glutamate. The A₂ receptor agonist 2-p-(2-carboxylethyl)-phenylamino-5′-N-ethylcarboxamido adenosine hydrochloride (CGS 21680) selectively increased glutamate release when applied at 5 μM while it selectively inhibited GABA release at a lower concentration (10 nM). The addition of NMDA to the medium, resulted in increased adenosine release equivalent to that found following stimulation with 50 mM KCl. Both the NMDA and the KCl-induced increases were eliminated by addition of -2-amino-5 phosphopentanoic acid ( -AP5), an NMDA-receptor antagonist. Slices prepared from the IMHV of chicks following successful training on the task showed enhanced adenosine release 30 min, 1, 3 and 6.5 h after training compared to chicks trained to peck a water-coated bead. The results show that changes in adenosine release from the IMHV accompany memory formation in the chick. We suggest that adenosine–amino acid transmitter interactions potentially via the activation of NMDA receptors, a necessary step in long-term memory formation for the task, may modulate the formation of memory for the one-trial passive avoidance task.     

The cannabinoid CB1 receptor antagonist rimonabant dose-dependently inhibits memory recall in the passive avoidance task in domestic chicks (Gallus domesticus)

The effect of endocannabinoids on synaptic plasticity has been demonstrated in a variety of species and brain regions. Relatively little is known about the localization and significance of cannabinoid (CB) receptors in the avian brain. The objective of the present study was to investigate the effect of a specific CB₁ receptor antagonist upon the acquisition and consolidation of memory in young domestic chicks. One-day-old domestic chicks (Gallus domesticus) were trained and tested by the passive avoidance paradigm. Systemic (i.p.) administration of the CB₁ receptor antagonist rimonabant in a dose of 1 mg/kg 30 min before the training failed to affect learning, but a similar treatment 30 min before the recall (5.5 h after training) attenuated the retention in 60% of animals. In another set of animals, a dose of 0.01 mg/kg produced no significant impairment, whereas doses 0.1 mg/kg and 1.0 mg/kg resulted in significant attenuation in passive avoidance performance when tested 30 min prior to recall. The results are discussed in terms of a putative mediating role of CB receptors in the consolidation of memory.     

Increases in NMDA receptor binding are specifically related to memory formation for a passive avoidance task in the chick: a quantitative autoradiographic study

One-day-old chicks (Gallus domesticus) were trained on a one-trial passive avoidance task in which the aversive stimulus was an unpleasant tasting substance, methylanthranilate (MeA). Control birds were presented with a water (W) coated bead. Five minutes after training a group of MeA-trained chicks were given a brief sub-convulsive trans-cranial electric shock, which rendered half amnesic whilst the remainder were able to show recall for the aversive stimulus. Thirty minutes after training birds were killed and quantitative receptor autoradiography was used to determine NMDA sensitive [³H]l-glutamate binding in specific regions of the forebrain of: (i) MeA-trained chicks; (ii) water-control chicks; (iii) MeA-trained electroshocked chicks showing recall; and (iv) MeA-electroshocked chicks amnesic for the aversive stimulus. Increases (80% in the left lobus parolfactorius and 67% in the left intermediate medial hyperstriatum ventrale) in NMDA sensitive [³H]l-glutamate binding occurred in electro-shocked chicks which showed recall of the aversive experience but were absent in MeA-trained chicks rendered amnesic by electro-shock. The increased binding in electroshocked MeA-trained birds which showed recall was similar to that observed previously in MeA-trained birds (without electroshock), compared to water control birds, whereas binding levels in MeA-trained electroshocked amnesic birds were not different from those of water control birds. These data argue strongly that alterations in binding to glutamate receptor sub-types are specific to memory formation for the passive avoidance task.