Helmut Prior

Mirror-Induced Behavior in the Magpie (Pica pica): Evidence of Self-Recognition

Comparative studies suggest that at least some bird species have evolved mental skills similar to those found in humans and apes. This is indicated by feats such as tool use, episodic-like memory, and the ability to use ones own experience in predicting the behavior of conspecifics. It is, however, not yet clear whether these skills are accompanied by an understanding of the self. In apes, self-directed behavior in response to a mirror has been taken as evidence of self-recognition. We investigated mirror-induced behavior in the magpie, a songbird species from the crow family. As in apes, some individuals behaved in front of the mirror as if they were testing behavioral contingencies. When provided with a mark, magpies showed spontaneous mark-directed behavior. Our findings provide the first evidence of mirror self-recognition in a non-mammalian species. They suggest that essential components of human self-recognition have evolved independently in different vertebrate classes with a separate evolutionary history.

Asymmetry pays: visual lateralization improves discrimination success in pigeons

Functional cerebral asymmetries, once thought to be exclusively human, are now accepted to be a widespread principle of brain organization in vertebrates [1]. The prevalence of lateralization makes it likely that it has some major advantage. Until now, however, conclusive evidence has been lacking. To analyze the relation between the extent of cerebral asymmetry and the degree of performance in visual foraging, we studied grain-grit discrimination success in pigeons, a species with a left hemisphere dominance for visual object processing [2,3]. The birds performed the task under left-eye, right-eye or binocular seeing conditions. In most animals, right-eye seeing was superior to left-eye seeing performance, and binocular performance was higher than each monocular level. The absolute difference between left- and right-eye levels was defined as a measure for the degree of visual asymmetry. Animals with higher asymmetries were more successful in discriminating grain from grit under binocular conditions. This shows that an increase in visual asymmetry enhances success in visually guided foraging. Possibly, asymmetries of the pigeons visual system increase the computational speed of object recognition processes by concentrating them into one hemisphere while preventing the other side of the brain from initiating conflicting search sequences of its own.

Role of a neuronal small non-messenger RNA: behavioural alterations in BC1 RNA-deleted mice

BC1 RNA is a small non-messenger RNA common in dendritic microdomains of neurons in rodents. In order to investigate its possible role in learning and behaviour, we compared controls and knockout mice from three independent founder lines established from separate embryonic stem cells. Mutant mice were healthy with normal brain morphology and appeared to have no neurological deficits. A series of tests for exploration and spatial memory was carried out in three different laboratories. The tests were chosen as to ensure that different aspects of spatial memory and exploration could be separated and that possible effects of confounding variables could be minimised. Exploration was studied in a barrier test, in an open-field test, and in an elevated plus-maze test. Spatial memory was investigated in a Barnes maze and in a Morris water maze (memory for a single location), in a multiple T-maze and in a complex alley maze (route learning), and in a radial maze (working memory). In addition to these laboratory tasks, exploratory behaviour and spatial memory were assessed under semi-naturalistic conditions in a large outdoor pen. The combined results indicate that BC1 RNA-deficient animals show behavioural changes best interpreted in terms of reduced exploration and increased anxiety. In contrast, spatial memory was not affected. In the outdoor pen, the survival rates of BC1-depleted mice were lower than in controls. Thus, we conclude that the neuron-specific non-messenger BC1 RNA contributes to the aptive modulation of behaviour.

Asymmetries of representation in the visual system of pigeons

ALTHOUGH functional asymmetries in the course of visual information processing have been known for a long time in humans as well as in non-human species, the structural basis of these asymmetries is largely unknown. We now report that due to an asymmetry of commissural projections in the pigeon the left nucleus rotundus of the ascending tectofugal visual system predominantly represents inputs from both eyes while the right nucleus rotundus mainly represents the contralateral left eye. We suggest that a comparable organization exists for several asymmetries in humans. A representation of both hemifields can provide the dominant hemisphere with direct access to all stimulus features when objects cross the vertical meridian.

Left-hemispheric superiority for visuospatial orientation in homing pigeons

To test for lateralisation of visuospatial orientation during homing, pigeons who had binocularly learned the homeward route from remote release sites were tested monocularly on either their left or their right eye for homing performance. In two experiments with three different release sites, birds using their right eye showed considerably better homing performance. If sun compass information was available, there was no difference in the direction of vanishing. Without this information, a difference between pigeons using their left or right eye emerged. Results show that visuospatial orientation in birds can be lateralised in favour of the left brain hemisphere and lend further support to the view that vision is important for pigeons homing on a familiar route. Cognitive mechanisms which might account for the observed pattern of lateralisation are discussed.

Visual lateralization and homing in pigeons

The aim of our study was to analyse the components of visual lateralization in pigeon homing, a large-scale spatial task. In a series of 13 releases, birds were tested as binocular controls or monocularly with the right or left-eye covered. Occlusion of either eye had a significant effect on initial orientation and homing performance. Vanishing bearings were deflected to the side of the open eye, vanishing intervals were longer, and homing speed was reduced. These parameters were affected to a different degree. Initial orientation was markedly lateralized, with birds using their right-eye deviating less from the mean of control birds and showing significantly less variance. One minute after release, the deviation and variance were similarly large in both monocular groups. However, while the right-eyed birds improved their performance until leaving the release site, the left-eyed birds failed to do so. Vanishing intervals were similar in both monocular groups, but homing speed was reduced to a lesser extent in pigeons using the right-eye. The degree of lateralization varied across different releases, but superiority of the right-eye/left hemisphere prevailed. Lateralization did not depend on familiarity with the release site. This suggests that the crucial processes involved the eyes, but did not depend on visual memory of landscape features at the release site. Results reveal, for the first time, asymmetries of directional orientation as an essential component of lateralized homing performance. As likely mechanisms we suggest hemispheric differences in magnetic compass orientation and in the adjustment to optic flow.

Dissociation of spatial reference memory, spatial working memory, and hippocampal mossy fiber distribution in two rat strains differing in emotionality

Rats of the inbred strains DA/Han and BDE/Han were compared on two complex spatial learning tasks, a spatial reference memory task in a 16-unit multiple T-maze and a spatial working memory task in an eight-arm radial-maze. In addition, sizes of hippocampal mossy fiber terminal fields were measured. BDE rats showed marked superiority in multiple T-maze learning whereas DA rats outperformed BDE rats on the radial-maze task. DA rats had significantly larger intra- and infrapyramidal mossy fiber terminal fields (IIP-MF). This is consistent with findings from other studies suggesting that large IIP-MF are related to excellent spatial radial-maze learning, but it also indicates that size of IIP-MF is correlated with processing of a specific type of spatial information rather than with overall spatial abilities. BDE rats had more extended suprapyramidal mossy fiber projections (SP-MF) and a larger hilus. Rats of both strains differed in exploratory behaviour and emotionality: DA rats revealed little freezing and had a high rearing activity, whereas BDE rats showed frequent freezing and reared rarely. Results suggest that IIP-MF are involved with flexible expression of memory, updating environmental information and parallel processing whereas SP-MF might be linked to processing of familiar information. Presumably, emotional factors contribute to performance differences.

Filling-in with colour: Different modes of surface completion

We investigated the figural dynamics of filling-in processes in figures with more than one possible figure-ground organisation. Using a central disk and two concentric rings as well as similar stimuli consisting of three nested squares or parallel stripes, we tested for filling-in with different equiluminant colour combinations. We observed four modes of filling-in: First, in most of the cases, the inner ring assumed the colour of the central disk and outer ring (M1). Second, the central disk became filled-in with the colour of the inner ring, without any colour change on the outer ring (M2). Third, in a first step, the colour of the inner ring spread onto the central disk; then, in a second step, the colour of the outer ring spread over the whole stimulus (M3). This two step filling-in process has not been reported so far. Fourth, a mode (M4) was sometimes observed that was characterised by the central disk and outer ring assuming the colour of the inner ring. Thus, colour filling-in or colour spreading proceeded both in a centripetal (periphery to fovea) as well as a centrifugal direction. The colours red and yellow proved to be stronger inducers than blue and green. Conversely, the latter colours became filled-in more easily than the former. The filled-in colour was always that of the inducing stimulus, i.e., there was no colour mixture. This suggests a long-range, neural process underlying filling-in under these conditions.

Lateralization of magnetic compass orientation in pigeons

The aim of our study was to test for lateralization of magnetic compass orientation in pigeons. Having shown that pigeons are capable of learning magnetic compass directions in an operant task, we wanted to know whether the brain hemispheres contribute differently and how the lateralization pattern relates to findings in other avian species. Birds that had learnt to locate food in an operant chamber by means of magnetic directions were tested for lateralization of magnetic compass orientation by temporarily covering one eye. Successful orientation occurred under all conditions of viewing. Thus, pigeons can perceive and process magnetic compass directions with the right eye and left brain hemisphere as well as the left eye and right brain hemisphere. However, while the right brain hemisphere tended to confuse the learned direction with its opposite (axial response), the left brain hemisphere specifically preferred the correct direction. Our findings demonstrate bilateral processing of magnetic information, but also suggest qualitative differences in how the left and the right brain deal with magnetic cues.

Effects of predictable and unpredictable intermittent noise on spatial learning in rats

The effects of predictable (periodic) and unpredictable (aperiodic) intermittent noise of moderate intensity (68 dB) on the learning of a complex T-maze by genetically defined rats were investigated. In Experiment 1, three groups (n=8) of rats learned a multiple T-maze, one group under control conditions, one group with predictable intermittent noise and one group with unpredictable intermittent noise. Results showed a profound effect of noise on learning and behavioural scores. Noise-exposed animals made less errors, finished their trials sooner and explored less. There was no difference between predictable and unpredictable noise. Further tests, during which formerly noise-exposed groups learned a new route under control conditions (Experiment 2) or the former controls learned a new route with noise (Experiment 3), suggest that the effects of noise on learning were caused by an effect of noise on memory formation and/or retrieval, rather than by long-term shifts in behavioural strategies.