How do animals 'do' geometry?
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Biomedical subjects
Publications and source records attributed to R Biegler.
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A characteristic feature of associative conditioning is that learning a predictive relationship between two events can block later learning about an added event. It is not yet well established whether blocking occurs in the spatial domain or the circumstances in which it does. We now report, using rats trained to search for hidden food near landmarks in an open field arena, that blocking can occur in spatial learning. The animals noticed the added landmark at the start of the blocking phase and explored it, but either failed to incorporate it into their spatial map or developed a representation in which only some landmarks actually control behavior. Additionally, performance at asymptote was controlled by the shape of the landmark array rather than the individual landmarks comprising it, indicating that blocking in the spatial domain may represent a failure to alter the encoded geometry of a learned array.
Two experiments were conducted to investigate the possible role of landmark stability in spatial learning. Rats were trained to search in a large arena for food hidden at a consistent distance and direction from either a single radially symmetric landmark or an array of two landmarks. We varied the relative degree to which the landmark array and/or the cues of the training context predicted the location of food, without varying the conditional probability of food being available given either cue. Experiment 1 used vestibular disorientation to ensure control of search location by experimenter-controlled cues. The results showed that making either a single landmark or a cluster of two adjacent landmarks the sole spatial predictor of reward location reduced the accuracy of search compared to a condition where both the landmark array and context cues were reliable spatial predictors. Varying global landmark stability had no effect when training was conducted using an array of two landmarks located some distance from each other. Context cues, when tested alone, triggered very little searching in appropriate locations, and the absolute magnitude of control over search was insufficient to account for the superiority of stable landmarks. The better learning with a stable landmark, and the dependence of this effect on the geometrical arrangement of landmarks, points to the conditions of spatial learning involving additional principles to those of simple associative conditioning. Experiment 2 examined landmark stability using a single landmark and fixed directional cues in the absence of vestibular disorientation. This also revealed a relative advantage of landmark stability, but animals with a landmark that moved from trial to trial did show some evidence of learning. Context cues when tested alone had minimal influence. Parametric manipulation of landmark stability offers a novel way of influencing spatial learning and thus understanding better the process through which egocentric representations of perceived space are transformed into allocentric representations of the real world.
To investigate whether spatial learning complies with associative learning theories or with theories of cognitive mapping, rats were trained in three experiments exploring the effect of variations in spatial predictive relationships. In experiment 1, it was found that making one of two landmarks the sole spatial predictor of reward, by varying the spatial relationship between reward and other cues, reduced the control over search exerted by that landmark compared with that observed when the landmark and context cues were both reliable predictors of reward location. This requirement for landmark stability rather than predictive power appears to contradict results obtained in conventional conditioning paradigms. Discrimination learning was unaffected, suggesting a dissociation between discrimination and spatial learning with respect to the influence of geometric stability. Further experiments used arrays of both single and multiple landmarks. Experiment 2 revealed that the stability of a single landmark improved accuracy of search, but also showed that local stability between a pair of landmarks that moved around the arena together was sufficient to support spatial learning. Experiment 3 examined landmark stability using fixed directional cues in the absence of vestibular disorientation. This also revealed a relative advantage of stable landmarks, but animals presented with a landmark that moved from trial to trial did show some evidence of learning. Parametric manipulation of landmark stability offers an intriguing way of influencing the process of spatial representation and thus understanding better the processes through which egocentric representations of perceived space are transformed into allocentric representations of the real world.
Neurons sensitive to both place and direction from distinct regions of the hippocampal formation, allometric relationships between spatial learning and hippocampal structure and pronounced impairments in spatial learning after lesions in this area, indicate that the hippocampal formation subserves allocentric spatial learning. To learn more about the process of spatial representation, we have developed a task that provides independent control of both landmark and directional cues. On the basis of physiological and behavioural work, this task also makes it possible to investigate the relevance of associative learning principles, such as predictability, to the spatial domain. We report here that although rats learn to discriminate between landmarks on the basis of their proximity to a reliably predicted food reward, they will only learn to use them to represent its location if they maintain stable locations within a geometric frame of reference.