Domestic Chicks Research Articles

Blockade of muscarinic acetylcholine receptor by scopolamine impairs the memory formation of filial imprinting in domestic chicks (Gallus Gallus domesticus)

Filial imprinting in precocial birds is a useful model for studying memory formation in early learning. The intermediate medial mesopallium (IMM) in the dorsal telencephalon is one of the critical brain regions where the releases of several neurotransmitters increase after the start of imprinting training. Among the increased neurotransmitters, the role of acetylcholine in imprinting has remained unclear. Acetylcholine in the mammalian brain plays an important role in encoding new memories. The muscarinic acetylcholine receptor subtype 1 (M1 receptor) and subtype 3 (M3 receptor) in the hippocampus and cortex of mammalian brain have been shown to be necessary for memory encoding. In this study, we examined whether the imprinting acquisition in chick can be impaired by injecting muscarinic acetylcholine receptor (mAChR) antagonist scopolamine into the bilateral IMM. We show that the injection of scopolamine decreased the preference for the imprinting object in the test, but did not affect the number of approaches to the imprinting object during training. Immunoblotting and immunohistochemistry revealed that M3 receptors were expressed in the IMM. Our data suggest that acetylcholine is involved in the memory formation of imprinting through M3 receptors in the IMM. The scopolamine-injected chicks may be useful as an animal model for dementia such as Alzheimer’s disease.

A combination insecticide at sub-lethal dose debilitated the expression pattern of crucial signalling molecules that facilitate craniofacial patterning in domestic chick Gallus domesticus

Pesticides despite being agents that protect the plants and humans from noxious pests, are infamous for their potential to cause detrimental health issues in nontargeted species. In order to ascertain the latter, a set of experiments were conducted by exposing early chick embryos to a widely used combination insecticide (Ci, 50% chlorpyrifos and 5% cypermethrin). The results revealed a myriad of congenital defects pertaining to craniofacial development such as anophthalmia, microphthalmia, exencephaly as well as deformed beak and cranial structures. These teratological manifestations could be attributed to the Ci induced alteration in the titre of major regulators of neurulation and ossification. Therefore, the mRNA and/or the protein level expression pattern of genes which are reported to be involved in the craniofacial development were studied at selected time points of embryonic development. The analysis of the result showed that there have been significant alternations in the expression patterns of the signalling molecules such as SHH, WNTs, CDH1, CDH2, L1CAM, PAX6, HOX, PCNA, GLI3, BMP7, FGF8, GLIs, SOX9, RUNX2, DLX5, COL10A1, CASPASE3 etc. on embryonic days 2, 4 and/or 10. Concurrently, on day 10, whole-mount skeletal staining and biochemical estimation of hydroxyproline were carried out in the cranial tissues of the embryos. The overall result of the current study indicates that exposure to Ci during early development impede the crucial regulatory signals that orchestrate the morphogenesis of cranial neural crest cells thereby hindering the normal progression of neural tube and endochondral ossification which collectively lead to craniofacial dysmorphism in domestic chicks.

Unlearned visual preferences for the head region in domestic chicks.

Unlearned tendencies to approach animate creatures are of great adaptive value, especially for nidifugous social birds that need to react to the presence of potential social companions shortly after hatching. Domestic chicks’ preferences for taxidermized hens provided the first evidence of social predispositions. However, the nature of the stimuli eliciting this predisposition is not completely understood. Here we explore the unlearned preferences of visually naïve domestic chicks for taxidermized animals. Visually naive chicks were tested for their approach preferences between a target stimulus (an intact stuffed animal whose head region was clearly visible) and a control stimulus. After confirming the predisposition for the intact stuffed fowl hen (Exp. 1), we found an analogous preference for a taxidermized, young domestic chick over a severely scrambled version of the same stimulus, whose body structure was completely disrupted, extending to same-age individuals the results that had been obtained with taxidermized hens (Exp. 2). We also directly tested preferences for specimens whose head region is visible compared to ones whose head region was occluded. To clarify whether chicks are sensitive to species-specific information, we employed specimens of female mallard ducks and of a mammalian predator, the polecat. Chicks showed a preference for the duck stimulus whose wings have been covered over a similar stimulus whose head region has been covered, providing direct evidence that the visibility of the head region of taxidermized models drive chicks’ behaviour in this test, and that the attraction for the head region indeed extends to females of other bird species (Exp. 3). However, no similar preference was obtained with the polecat stimuli (Exp. 4). We thus confirmed the presence of unlearned visual preferences for the head region in newly-hatched chicks, though other factors can limit the species-generality of the phenomenon.

Valproate Exposure in ovo Attenuates the Acquisition of Social Preferences of Young Post-hatch Domestic Chicks.

Embryonic exposure to valproic acid (VPA) is known to produce sociability deficits, resembling human autistic phenotypes, in several vertebrate species. Animals living in groups prefer the proximity of peers and have the ability to perceive and to respond to social signals for modifying behavior. Chicks of Galliform birds, known to display early preference behaviors, have been used extensively for adaptive learning studies. Young precocial birds seem to be useful models also for studying the effect of embryonic VPA treatment. Here, domestic chicken eggs were injected with sodium valproate (200 μl of 35 μmol/L solution) or with vehicle (distilled water) on the 14th day of incubation. After hatching, the chicks were tested for one-trial passive avoidance learning at day 1, vocalization due to isolation as a measure of stress level (day 2), approach preference to large versus small groups of age-matched conspecifics (day 5), and to those with normal versus blurred head features (day 7). In addition, we tested the preference of birds to conspecifics reared in group versus those reared in isolation (day 9), as well as the preference of chicks to familiar versus non-familiar conspecifics (day 21). Our findings confirm previous reports concerning an adverse effect of VPA on embryonic development, including a tendency for aborted or delayed hatching and, occasionally, for locomotor disorders in a small percentage of birds (eliminated from later studies). Otherwise, VPA treatment did not impair motor activity or distress level. Memory formation for the aversive stimulus and discrimination of colors were not impaired by VPA treatment either. Innate social predispositions manifested in approach preferences for the larger target group or for the birds with natural facial features remained unaffected by VPA exposure. The most prominent finding was attenuation of social exploration in VPA-exposed birds (expressed as the frequency of positional switches between two stimulus chicks after the first choice), followed by a deficit in the recognition of familiar conspecifics, unfolding at the end of the third week. Social exploration and recognition of familiar individuals are the key elements impaired at this stage. The results underline the importance of early social exploration in ASD.

Selective response of the nucleus taeniae of the amygdala to a naturalistic social stimulus in visually naive domestic chicks

The detection of animate beings at the onset of life is important for phylogenetically distant species, such as birds and primates. Naïve chicks preferentially approach a stimulus resembling a conspecific (a stuffed fowl) over a less naturalistic one (a scrambled version of the stuffed fowl, presenting the same low-level visual features as the fowl in an unnatural configuration). The neuronal mechanisms underlying this behavior are mostly unknown. However, it has been hypothesized that innate social predispositions may involve subpallial brain areas including the amygdala. Here we asked whether a stuffed hen would activate areas of the arcopallium/amygdala complex, in particular the nucleus taeniae of the amygdala (TnA) or septum. We measured brain activity by visualizing the immediate early gene product c-Fos. After exposure to the hen, TnA showed higher density of c-Fos expressing neurons, compared to chicks that were exposed to the scrambled stimulus. A similar trend was present in the lower portion of the arcopallium, but not in the upper portion of the arcopallium or in the septum. This demonstrates that at birth the TnA is already engaged in responses to social visual stimuli, suggesting an important role for this nucleus in the early ontogenetic development of social behavior.

Distinct effect of early and late embryonic light-stimulation on chicks' lateralization

Embryonic light exposure affects similarly functional lateralization in fish and birds. While the light acts on an asymmetric habenular system during the first post fertilization hours in zebrafish, in the domestic chicks it shapes the thalamofugal visual pathway affecting the right retinal photoreceptors in the last stages before hatching. However, recent evidence has shown that also in chicks a precocial embryonic time window seems open to light action. Here we addressed the issue of whether the light effect is comparable in the early and late sensitive periods of chicks' embryonic development by testing birds coming from early (EL) and late (LL) light stimulated eggs compared to dark maintained eggs (DK) under different conditions of vision in a gravel-grain task. The perseveration of pecks directed to irrelevant elements revealed that in all chicks the right hemisphere was heavily attracted by the novel elements when tested with the left eye. When using the right eye, instead, only DK chicks attended repeatedly to distractors whereas LL and EL chicks showed a left hemisphere advantage for fine discrimination and sustained attention; conversely, when tested binocularly, LL chicks perseverated significantly more than both DK and EL chicks, likely compensating the distraction with the analysis carried out by both hemispheres. For the first time, we unveiled a fine graded difference between the light modulation exerted during the two time windows, adding evidence to the idea that genes and environmental factors interplay in several separate routes to the modulation of the neurodevelopment of cerebral lateralization in vertebrates.

Context learning before birth: evidence from the chick embryo.

Learning contextual information to form associative memories with stimuli of interest is an important brain function in both human and non-human animals. Intuitively, one would expect that such a sophisticated cognitive skill develops postnatally, as the organism starts exploring the surrounding environment to search for significant contingencies among stimuli. Here we show, instead, that even before hatching, domestic chicks are capable of forming associative memories between discrete alerting sounds and the surrounding context, as attested by the fact that habituation of the freezing response to the sounds is affected by the context of stimulation. This finding indicates that, while in the egg, chicks recognize and learn the context in which they are stimulated. Hence, context learning in chicks is an innate brain function already active before birth, which can provide an immediate survival advantage to the newborns of this precocial avian species.

Similar predator aversion for natural prey with diverse toxicity levels

Müllerian mimicry between chemically defended species arises from selection exerted by predators in which individuals benefit from higher survival when they share the same warning signal. However, despite sharing warning signals, co-mimetic species harbour a diversity of toxins at a range of different concentrations. This variation may affect the rate of predator avoidance learning and therefore the dynamics of mimicry. Here, to understand the nature of mimetic relationships in natural communities of butterflies and moths, we compared protection against predators induced by chemical defences of 13 lepidopteran species belonging to six mimicry complexes. Protection was estimated by quantifying the extent of avoidance learning, using domestic chicks, Gallus gallus domesticus, as model predators. We showed that most co-mimics were avoided at similarly high rates, with the exception of two species eliciting markedly slower rates of avoidance. Assuming our model and natural predators behave similarly and cannot distinguish co-mimics visually, the similar avoidance learning they induce supports the contention that mutualistic relationships among these co-mimetic species might be predominant in natural communities, despite large variation in toxin concentrations. Indeed, by comparing our estimated avoidance learning rate to mean toxin concentration, we found that prey with a two- to three-fold difference in toxin content generated similar avoidance learning indices. This lack of a direct relationship between prey defence level and predator avoidance learning points to alternative evolutionary mechanisms promoting the evolution of high levels of toxins.

Response of passerine birds and chicks to larvae and pupae of ladybirds

1. Few, if any, experimental tests have demonstrated the anti‐predator protection of the developmental stages of ladybirds (Coleoptera: Coccinellidae) against vertebrates, despite the fact that both the visual appearance of ladybirds and the content of defensive compounds fulfil the definition of an aposematic prey.2. In this study, avian predators of three species were confronted with fourth‐instar larvae and pupae of the harlequin ladybird (Harmonia axyridis) – a large, conspicuous, and toxic ladybird species.3. The selected predators differed in their individual experience and attitude to ladybirds. Wild‐caught great tits (Parus major) strongly avoided attacking ladybirds in general, whereas wild‐caught tree sparrows (Passer montanus) were willing to include ladybirds in their diet. Domestic chicks (Gallus gallus domesticus) have never been confronted with ladybirds but usually show avoidance of aposematic signals. In this study, great tits and domestic chicks avoided both developmental stages, but in the case of chicks the avoidance had to be learned over the course of repeated encounters. Sparrow avoidance was significantly lower, with more than one‐third of the prey being attacked and eaten.4. The protection of both developmental stages of ladybirds is similar to adults, despite substantially different visual appearance.

Visual Imprinting in Birds: Behavior, Models, and Neural Mechanisms.

Filial imprinting is a process, readily observed in precocial birds, whereby a social attachment is established between a young animal and an object that is typically (although not necessarily) a parent. During a perinatal sensitive period, the young animal learns characteristics of the object (the imprinting stimulus) simply by being exposed to it and will subsequently recognize and selectively approach this stimulus. Imprinting can thus establish a filial bond with an individual adult: a form of social cohesion that may be crucial for survival. Behavioral predispositions can act together with the learning process of imprinting in the formation, maintenance, and modification of the filial bond. Memory of the imprinting stimulus, as well as being necessary for social recognition, is also used adaptively in perceptual classification of sensory signals. features of an imprinting stimulus, such as similarity or difference between stimulus components, can also be recognized. Studies of domestic chicks have elucidated the neural basis of much of the above behavior. This article discusses (1) principal behavioral characteristics of filial imprinting and related predispositions, (2) theoretical models that have been developed to account for this behavior, and (3) physiological results elucidating the underlying neural mechanisms. Interactions between these different levels of analysis have resulted in advancement of all of them. Taken together, the different approaches have helped define strategies for investigating mechanisms of learning, memory, and perception.

Embryonic Exposure to Valproic Acid Affects Social Predispositions for Dynamic Cues of Animate Motion in Newly-Hatched Chicks.

Early predispositions to preferentially orient toward cues associated with social partners have been documented in several vertebrate species including human neonates and domestic chicks. Human newborns at high familiar risk of Autism Spectrum Disorder (ASD) show differences in their attention toward these predisposed stimuli, suggesting potential impairments in the social-orienting mechanisms in ASD. Using embryonic exposure to valproic acid (VPA) we modeled ASD behavioral deficits in domestic chicks. To investigate social predispositions toward animate motion in domestic chicks, we focused on self-propulsion, using two video-animations representing a simple red circle moving at constant speed (speed-constant) or one that was changing its speed (accelerating and decelerating; speed-change). Using a spontaneous choice test for the two stimuli, we compared spontaneous preferences for stimuli that autonomously change speed between VPA- and vehicle-injected chicks. We found that the preference for speed changes was abolished in VPA-injected chicks compared to vehicle-injected controls. These results add to previous findings indicating similar impairments for static social stimuli and suggest a specific effect of VPA on the development of mechanisms that enhance orienting toward animate stimuli. These findings strengthen the hypothesis of an early impairment of predispositions in the early development of ASD. Hence, early predispositions are a potentially useful tool to detect early ASD symptoms in human neonates and to investigate the molecular and neurobiological mechanisms underlying the onset of this neurodevelopmental disorder.

Spontaneous and light-induced lateralization of immediate early genes expression in domestic chicks

Exposure of domestic chicks’ eggs to light during embryo incubation stimulates asymmetrically the two eye-systems, reaching selectively the right eye (left hemisphere) and inducing asymmetries at the behavioral and neural level. Surprisingly, though, some types of lateralization have been observed also in dark incubated chicks, especially at the behavioral level. Here we investigate the mechanisms subtending the development of lateralization, in the presence and in the absence of embryonic light exposure. We measured the baseline level of expression for the immediate early gene product c-Fos, used as an indicator of the spontaneous level of neural activity and plasticity in four areas of the two hemispheres (preoptic area, septum, hippocampus and intermediate medial mesopallium). Additional DAPI staining measured overall cell density (regardless of c-Fos expression), ruling out any confound due to underlying asymmetries in cell density between the hemispheres. In different brain areas, c-Fos expression was lateralized either in light- (septum) or in dark-incubated chicks (preoptic area). Light exposure increased c-Fos expression in the left hemisphere, suggesting that c-Fos expression could participate to the known effects of light stimulation on brain asymmetries. Interestingly, this effect was visible few days after the end of the light exposure, revealing a delayed effect of light exposure on c-Fos baseline expression in brain areas outside the visual pathways. In the preoptic area of dark incubated chicks, we found a rightward bias for c-Fos expression, revealing that lateralization of the baseline level of activity and plasticity is present in the developing brain also in the absence of light exposure.

Naïve 3-day-old domestic chicks (Gallus gallus) are attracted to discrete acoustic patterns characterizing natural vocalizations.

Nonsongbirds can produce rhythmical sounds that, at times, have been shown to be meaningful in their communication. This raises the possibility that rhythm is a separate ability that might have evolved earlier than song. We asked whether nearly completely naïve domestic chicks perceive rhythm and respond in specific ways to different rhythmic patterns. To do so, specific constituent parameters of rhythmicity were used based on the sound of a natural mother hen's cluck. The sound samples created ranged from a continuous sound to articulated rhythmic patterns of alternating strong and weak events. Chicks' reactivity to the patterns was tested over a series of sound exposure experiments by their propensity to operate a running wheel toward the acoustic source, a paradigm simulating chicks' natural affiliative response to the hen's call. Results showed that motor activity increased markedly when acoustic events were discrete (compared with continuous), and significantly when accent structure was faster (compared with slower rates). Similar to human infants, chicks showed a significant preference for pulsed over continuous patterns. Chicks also ran harder toward calls with fast strong pulsating events, suggesting that different arrangements of events in time can be differently arousing, but independently of whether the events were presented in a regular or nonregular fashion. (PsycINFO Database Record (c) 2019 APA, all rights reserved).

Environmental complexity: A buffer against stress in the domestic chick.

Birds kept in commercial production systems can be exposed to multiple stressors from early life and this alters the development of different morphological, immunological and behavioural indicators. We explore the hypothesis that provision of a complex environment during early life, better prepares birds to cope with stressful events as well as buffers them against future unpredictable stressful episodes. In this study, 96 one day old pullets were randomly distributed in eight pens (12 birds/pen). Half of the chicks (N = 48) were assigned to a Complex Environment (CENV: with perches, a dark brooder etc.) the others to a Simple Environment (SENV: without enrichment features). Half of the birds from each of these treatments were assigned to a No Stress (NSTR, 33°C) or to an acute Cold Stress (CSTR, 18–20°C) treatment during six hours on their second day of life. At four weeks of age, chicks with these four different backgrounds were exposed to an Intermittent Stressful Challenges Protocol (ISCP). In an immunological test indicative of pro-inflammatory status Phytohemagglutinin-P (PHA-P), the response of CSTR birds was ameliorated by rearing chicks in a CENV as they had a similar response to NSTR chicks and a significantly better pro-inflammatory response than those CSTR birds reared in a SENV (five days after the CSTR treatment was applied). A similar better response when coping with new challenges (the ISCP) was observed in birds reared in a CENV compared to those from a SENV. Birds reared in the CENV had a lower heterophil/lymphocyte ratio after the ISCP than birds reared in SENV, independently of whether or not they had been exposed to CSTR early in life. No effects of stress on general behaviour were detected, however, the provision of a CENV increased resting behaviour, which may have favoured stress recover. Additionally, we found that exposure to cold stress at an early age might have rendered birds more vulnerable to future stressful events. CSTR birds had lower humoral immune responses (sheep red blood cells induced antibodies) after the ISCP and started using elevated structures in the CENV later compared to their NSTR conspecifics. Our study reflects the importance of the early provision of a CENV in commercial conditions to reduce negative stress-related effects. Within the context of the theory of adaptive plasticity, our results suggest that the early experience of the birds had long lasting effects on the modulation of their phenotypes.

An imprinting object rapidly acquires high attractiveness when associated with food delivery

Abstract Young precocial birds develop a preference for an imprinting object by mere visual exposure to it in the absence of conventional physiological reinforcement. The lack of the necessity of conventional reinforcement for imprinting, however, does not mean that such reinforcement is unimportant. The evidence presented here shows that an imprinting object rapidly acquires high attractiveness to young chicks when it is associated with food provisioning. Domestic chicks, Gallus gallus domesticus, were first exposed to two different imprinting objects in the absence of any reinforcement. Subsequently, two groups of chicks received a single feeding session wherein they were provided with prey from one of the imprinting objects. A third group served as a control in which the chicks were exposed to one of the imprinting objects and prey delivery in an unpaired way. Finally, all chicks received two choice tests to assess their preferences for the two imprinting objects. The chicks that received food from an imprinting object strongly preferred that object to the alternative familiar-only object, and preferred the familiar-only object to a novel object. The control group did not show any preference between the two imprinting objects, but preferred the unpaired imprinting object to a novel object. These results suggest that primary-need reinforcers such as food contribute to increasing the attractiveness of an imprinting object by promoting rapid associative learning.

GABA-A and GABA-B Receptors in Filial Imprinting Linked With Opening and Closing of the Sensitive Period in Domestic Chicks (Gallus gallus domesticus).

Filial imprinting of domestic chicks has a well-defined sensitive (critical) period lasting in the laboratory from hatching to day 3. It is a typical model to investigate the molecular mechanisms underlying memory formation in early learning. We recently found that thyroid hormone 3,5,3′-triiodothyronine (T3) is a determinant of the sensitive period. Rapid increases in cerebral T3 levels are induced by imprinting training, rendering chicks imprintable. Furthermore, the administration of exogenous T3 makes chicks imprintable on days 4 or 6 even after the sensitive period has ended. However, how T3 affects neural transmission to enable imprinting remains mostly unknown. In this study, we demonstrate opposing roles for gamma-aminobutyric acid (GABA)-A and GABA-B receptors in imprinting downstream of T3. Quantitative reverse transcription polymerase chain reaction and immunoblotting showed that the GABA-A receptor expression increases gradually from days 1 to 5, whereas the GABA-B receptor expression gradually decreases. We examined whether neurons in the intermediate medial mesopallium (IMM), the brain region responsible for imprinting, express both types of GABA receptors. Immunostaining showed that morphologically identified putative projection neurons express both GABA-A and GABA-B receptors, suggesting that those GABA receptors interact with each other in these cells to modulate the IMM outputs. The roles of GABA-A and GABA-B receptors were investigated using various agonists and antagonists. Our results show that GABA-B receptor antagonists suppressed imprinting on day 1, while its agonists made day 4 chicks imprintable without administration of exogenous T3. By contrast, GABA-A receptor agonists suppressed imprinting on day 1, while its antagonists induced imprintability on day 4 without exogenous T3. Furthermore, both GABA-A receptor agonists and GABA-B receptor antagonists suppressed T3-induced imprintability on day 4 after the sensitive period has ended. Our data from these pharmacological experiments indicate that GABA-B receptors facilitate imprinting downstream of T3 by initiating the sensitive period, while the GABA-A receptor contributes to the termination of the sensitive period. In conclusion, we propose that opposing roles of GABA-A and GABA-B receptors in the brain during development determine the induction and termination of the sensitive period.

Thyroid Hormone Sensitizes the Imprinting-Associated Induction of Biological Motion Preference in Domestic Chicks.

Filial imprinting is associated with induction of predisposed preference to animations that bear visual features of Johansson's biological motion (BM), and the induction is limited to a few days after hatching. As thyroid hormone (3,5,3′-triiodothyronine, T3) plays a critical role in determining the sensitive period of imprinting, we examined if exogenously applied T3 (or iopanoic acid, IOP; a selective inhibitor for converting enzymes) could also sensitize (or desensitize) the BM induction. Chicks were trained by using a non-BM stimulus (rotating red toy) according to a conventional imprinting procedure. Trained chicks were tested for preference to a point-light BM animation (walking chick) over a non-BM animation (linear motion), and for the preference for the familiarized stimulus (red toy) over an unfamiliar one (yellow toy). In 1-day chicks, those injected with IOP showed significantly lower scores than controls on both BM and imprinting tests. In 4-days chicks, those injected with T3 showed higher scores than control, but the difference in BM score was not significant. Imprinting and the accompanying T3 surge may be necessary for the predisposed BM preference to appear in 1-day chicks. Even after the conventional sensitive period is over, exogenous T3 can partly re-sensitize the BM preference as it does imprinting.

Prenatal Sensory Stimulation Induces BDNF Gene Expression in the Brain and Potentiates the Development of Species-Specific Predisposition in Newborn Chicks.

We studied the effects of light and non-specific sound stimulation of domestic chick embryos on their filial preference as well as on the expression of two transcriptional factors c-Fos and Egr-1 and neurotrophin BDNF in the embryo brain. Prenatal light stimulation increased preference of the "natural" object, thus producing a priming effect. In the brain of E19 embryos, c-Fos and Egr-1 were expressed at a high basal level and neither light nor sound stimulation affected the number of cells expressing these factors. BDNF mRNA was also present in a number of brain areas of non-stimulated embryos, but light and sound stimulation enhanced the expression of BDNF mRNA in brain structures associated with filial imprinting. These findings suggest that BDNF is probably involved in the effects of prenatal priming on the development of species-specific behavior.

Micro-RNAs, their target proteins, predispositions and the memory of filial imprinting

Visual imprinting is a learning process whereby young animals come to prefer a visual stimulus after exposure to it (training). The intermediate medial mesopallium (IMM) in the domestic chick forebrain is critical for visual imprinting and contributes to molecular regulation of memory formation. We investigated the role of micro-RNAs (miRNAs) in such regulation. Twenty-four hours after training, miRNA spectra in the left IMM were compared between chicks with high preference scores (strong memory for imprinting stimulus), and chicks with low preference scores (weak memory for imprinting stimulus). Using criteria of significance and expression level, we chose gga-miR-130b-3p for further study and found that down-regulation correlated with learning strength. No effect was detected in posterior nidopallium, a region not involved in imprinting. We studied two targets of gga-miR-130b-3p, cytoplasmic polyadenylation element binding proteins 1 (CPEB-1) and 3 (CPEB-3), in two subcellular fractions (P2 membrane-mitochondrial and cytoplasmic) of IMM and posterior nidopallium. Only in the left IMM was a learning-related effect observed, in membrane CPEB-3. Variances from the regression with preference score and untrained chicks suggest that, in the IMM, gga-miR-130b-3p level reflects a predisposition, i.e. capacity to learn, whereas P2 membrane-mitochondrial CPEB-3 is up-regulated in a learning-specific way.

Ecological Validity of Impulsive Choice: Consequences of Profitability-Based Short-Sighted Evaluation in the Producer-Scrounger Resource Competition

Results of intertemporal choice paradigm have been accounted for mostly by psychological terms such as temporal discounting of subjective value. Inability to wait for delayed gratification (choice impulsiveness, as opposed to self-control) is often taken to represent violated rationality. If viewed from foraging ecology, however, such impulsiveness can be accountable as adaptive adjustments to requirements in nature. First, under the circumstance where foragers stochastically encounter food items, the optimal diet-menu model suggests that each option must be evaluated by profitability (e/h), which is the ratio of energetic gain (e) per handling time (h), a short-sighted currency. As h includes the delay, profitability will be hyperbolically lower for long-delay food. Second, because of the resource competition between producing and scrounging foragers, profitability of the producer’s gain will critically depend on the scrounger’s behaviors. We first constructed an analytical model. The model predicted that the profitability of small and short-delay food option (SS) can be higher than that of the large and long-delay alternative (LL), depending on the duration in which the producer can monopolize a food patch (finder’s share). Next, we conducted numerical simulations on the assumption of variable food amount in each patch with realistic set of behavioral parameters. Although non-linearity of profitability function largely reduced profitability for variable amount of food, SS still can have a higher profitability than LL when the finder’s share is small. Because SS is consumed more quickly, it is more resistant against scrounging than LL. In good accordance, foraging domestic chicks form a synchronized flock and show socially-facilitated investment of effort. If raised in competition, chicks develop a higher degree of choice impulsiveness.