Imprinting on time-structured acoustic stimuli in ducklings

Filial imprinting is a dedicated learning process that lacks explicit reinforcement. The phenomenon itself is narrowly heritably canalized, but its content, the representation of the parental object, reflects the circumstances of the newborn. Imprinting has recently been shown to be even more subtle and complex than previously envisaged, since ducklings and chicks are now known to select and represent for later generalization abstract conceptual properties of the objects they perceive as neonates, including movement pattern, heterogeneity and inter-component relationships of same or different. Here, we investigate day-old Mallard ( Anas platyrhynchos ) ducklings’ bias towards imprinting on acoustic stimuli made from mallards’ vocalizations as opposed to white noise, whether they imprint on the temporal structure of brief acoustic stimuli of either kind, and whether they generalize timing information across the two sounds. Our data are consistent with a strong innate preference for natural sounds, but do not reliably establish sensitivity to temporal relations. This fits with the view that imprinting includes the establishment of representations of both primary percepts and selective abstract properties of their early perceptual input, meshing together genetically transmitted prior pre-dispositions with active selection and processing of the perceptual input.

Steps towards a computational ethology: an automatized, interactive setup to investigate filial imprinting and biological predispositions

Soon after hatching, the young of precocial species, such as domestic chicks or ducklings, learn to recognize their social partner by simply being exposed to it (imprinting process). Even artificial objects or stimuli displayed on monitor screens can effectively trigger filial imprinting, though learning is canalized by spontaneous preferences for animacy signals, such as certain kinds of motion or a face-like appearance. Imprinting is used as a behavioural paradigm for studies on memory formation, early learning and predispositions, as well as number and space cognition, and brain asymmetries. Here, we present an automatized setup to expose and/or test animals for a variety of imprinting experiments. The setup consists of a cage with two high-frequency screens at the opposite ends where stimuli are shown. Provided with a camera covering the whole space of the cage, the behaviour of the animal is recorded continuously. A graphic user interface implemented in Matlab allows a custom configuration of the experimental protocol, that together with Psychtoolbox drives the presentation of images on the screens, with accurate time scheduling and a highly precise framerate. The setup can be implemented into a complete workflow to analyse behaviour in a fully automatized way by combining Matlab (and Psychtoolbox) to control the monitor screens and stimuli, DeepLabCut to track animals’ behaviour, Python (and R) to extract data and perform statistical analyses. The automated setup allows neuro-behavioural scientists to perform standardized protocols during their experiments, with faster data collection and analyses, and reproducible results.

Imprinting on time-structured acoustic stimuli in ducklings

Filial imprinting is a dedicated learning process that lacks explicit reinforcement. The phenomenon itself is narrowly heritably canalized, but its content, the representation of the parental object, reflects the circumstances of the newborn. Imprinting has recently been shown to be even more subtle and complex than previously envisaged, since ducklings and chicken are now known to select and represent for later generalization abstract conceptual properties of the objects they perceive as neonates, including movement pattern, heterogeneity, and inter-component relationships of same or different. Here we investigate whether day-old Mallard ducklings (Anas platyrhynchos) imprint on the temporal separation between duos of brief acoustic stimuli, and whether they generalize such timing information to novel sound types. Subjects did discriminate temporal structure when imprinted and tested on natural duck calls, but not when using white noise for imprinting or testing. Our data confirm that imprinting includes the establishment of neural representations of both primary percepts and abstract properties of candidate objects, meshing together genetically transmitted prior knowledge with selected perceptual input.

Resurgence of an Inborn Attraction for Animate Objects via Thyroid Hormone T3

For inexperienced brains, some stimuli are more attractive than others. Human neonates and newly hatched chicks preferentially orient towards face-like stimuli, biological motion, and objects changing speed. In chicks, this enhances exposure to social partners, and subsequent attachment trough filial imprinting. Early preferences are not steady. For instance, preference for stimuli changing speed fades away after 2 days in chicks. To understand the physiological mechanisms underlying these transient responses, we tested whether early preferences for objects changing speed can be promoted by thyroid hormone 3,5,3′-triiodothyronine (T3). This hormone determines the start of imprinting’s sensitive period. We found that the preference for objects changing speed can be re-established in female chicks treated with T3. Moreover, day-1 chicks treated with an inhibitor of endogenous T3 did not show any preference. These results suggest that the time windows of early predispositions and of sensitive period for imprinting are controlled by the same molecular mechanisms.

Stability and individual variability of social attachment in imprinting

Filial imprinting has become a model for understanding memory, learning and social behaviour in neonate animals. This mechanism allows the youngs of precocial bird species to learn the characteristics of conspicuous visual stimuli and display affiliative response to them. Although longer exposures to an object produce stronger preferences for it afterwards, this relation is not linear. Sometimes, chicks even prefer to approach novel rather than familiar objects. To date, little is known about how filial preferences develop across time. This study aimed to investigate filial preferences for familiar and novel imprinting objects over time. After hatching, chicks were individually placed in an arena where stimuli were displayed on two opposite screens. Using an automated setup, the duration of exposure and the type of stimuli were manipulated while the time spent at the imprinting stimulus was monitored across 6 days. We showed that prolonged exposure (3 days vs 1 day) to a stimulus produced robust filial imprinting preferences. Interestingly, with a shorter exposure (1 day), animals re-evaluated their filial preferences in functions of their spontaneous preferences and past experiences. Our study suggests that predispositions influence learning when the imprinting memories are not fully consolidated, driving animal preferences toward more predisposed stimuli.

Resurgence of a perinatal attraction for animate objects via thyroid hormone T3

SummaryFor inexperienced brains, some stimuli are more attractive than others. Human neonates and newly-hatched chicks preferentially orient towards face-like stimuli, biological motion, and objects changing speed. In chicks, this enhances exposure to social partners, and subsequent attachment trough filial imprinting. Early preferences are not steady. The preference for stimuli changing speed fades away after three days in chicks. To understand the physiological mechanisms underlying these transient responses, we tested whether the early preferences for objects changing speed can be promoted by thyroid hormone 3,5,3’-triiodothyronine (T3). This hormone determines the start of imprinting’s sensitive period. We found that the preference for objects changing speed can be re-established in female chicks treated with T3. Moreover, day-one chicks treated with an inhibitor of endogenous T3 did not show any preference. These results suggest that the time windows of early predispositions and of high plasticity are controlled by the same molecular mechanisms.

Research evidence from studies on filial imprinting, attachment, and early life stress: a new route for scientific integration

Attachment is a concept that was developed and researched in developmental psychology in uptake of findings on filial imprinting from ethology. In the present period, however, attachment concepts are increasingly applied to and investigated in animal research, thereby translating back criteria that were established for human infants. It herein appears that findings on filial imprinting are becoming more and more forgotten, whilst basic findings in human infants are not reflected in investigations on attachment in animals. To re-integrate both domains, the present article undertakes the effort in briefly reviewing and recapitulating basic findings in human attachment and recent research on filial imprinting. In specific, replicated were critical roles of the conversion of thyroid prohormone by 2 iodothyronine deiodinase (Dio2) into triiodothyronine (T3) in the regulation of the timing of imprinting learning. Because of the interactions of T3 with oxytocinergic and dopaminergic neurones of the hypothalamic paraventricular nucleus, these findings provide new neuroendocrinological insight for possible relations with both attachment and metabolic sequelae of early life stress. Necessary is a mutual integration of all recent advances in the yet separated fields.