Abstract: It is often assumed that there is more than one kind of learning--or more than one memory system--each of which is specialized for a different function. Yet, the criteria by which the varieties of learning and memory should be distinguished are seldom clear. Learning and memory phenomena can differ from one another across species or situations (and thus be specialized) in a number of different ways. What is needed is a consistent theoretical approach to the whole range of learning phenomena, and one is explored here. Parallels and contrasts in the study of sensory systems illustrate one way to integrate the study of general mechanisms with an appreciation of species-specific adaptations.

Abstract: Marsh tits (Parus palustris) store single food items in scattered locations and recover them hours or days later. Some properties of the spatial memory involved were analyzed in two laboratory experiments. In the first, marsh tits were offered 97 sites for storing 12 seeds. They recovered a median of 65% of them 2-3 hr later, making only two errors per seed while doing so. Over trials, they used some sites more often than others, but during recovery they were more likely to visit a site of any preference value if they had stored a seed there that day than if they had not. Recovery performance was much worse if the experimenters moved the seeds between storage and recovery. A fixed search strategy that had some of the same average properties as the tits' search behavior also did worse than the real birds. In Experiment 2, any tendency to visit the same sites on successive daily tests in the aviary was placed in opposition to memory for storage sites by allowing the tits to store more seeds 2 hr after storing a first batch. They tended to avoid individual storage sites holding seeds from the first batch. When the tits searched for all the seeds 2 hr later, they tended to recover more seeds from the second batch than from the first, i.e., there was a recency effect.

Abstract: Animals frequently use signals that travel further than the spacing between individuals. For every intended recipient of a given signal there are likely to be many other individuals that receive information. Eavesdropping on signalling interactions between other individuals provides a relatively cost-free method of assessing future opponents or mates. Male great tits (Parus major) extract relative information from such interactions between individuals unknown to them. Here, we show that male great tits can take information gathering a stage further and obtain more information about a previously unencountered intruder, by the hitherto unknown capability of combining information gathered by eavesdropping with that derived from their own direct interaction with an individual. Prior experience with an intruder (A) was achieved by subjecting a focal male to different levels of intrusion simulated using interactive playback. This intruder (A) then took part in a simulated interaction with an unknown male (B) outside the territorial boundary of the focal males. In response to subsequent intrusion by the second male (B), focal males showed low song output in response to males that had lost to a male that the subject was able to beat. Males of known high quality, or those about which information was ambiguous, elicited a high level of song output by focal males. We discuss the implications of this finding for the evolution of communication and social behaviour.

Abstract: Theoretical models have demonstrated the possibility of stable cost-free signalling of need between relatives. The stability of these cost-free equilibria depends on the indirect fitness cost of cheating and deceiving a donor into giving away resources. We show that this stability is highly sensitive to the distribution of need among signallers and receivers. In particular, cost-free signalling is likely to prove stable only if there is very large variation in need (such that the least-needy individuals stand to gain much less than the most-needy individuals from additional resources). We discuss whether these conditions are likely to be found in altricial avian breeding systems--the most intensively studied instance of signalling of need between relatives. We suggest that cost-free signalling is more likely to prove stable and will provide parents with more information during the earlier phases of chick growth, when parents can more easily meet the demands of a brood (and chicks are more likely to reach satiation). Later, informative yet cost-free signalling is unlikely to persist.

Abstract: Animal communication generally occurs in the environment of a network of several potential signallers and receivers. Within a network environment, it is possible to gain relative information about conspecifics by eavesdropping on signalling interactions. We presented male great tits with the opportunity to gain such information by simulating singing interactions using two loudspeakers. Interactions were presented so that relevant information was not available in the absolute singing behaviour of either individual, only in the relative timing of their songs in the interaction as a whole. We then assayed the information extracted by focal males by subsequently introducing one of the 'interactants' (i.e. loudspeakers) into the territory of the focal male. Focal males responded with a reduced song output to males that had just 'lost' an interaction. Focal males did not respond significantly differently to 'winners' as compared with intruders recently involved in an interaction that contained no consistent information. Focal males also responded by switching song types more often when encountering males that had recently been involved in a low-intensity interaction. These results provide the clearest evidence yet that male songbirds extract information from signal interactions between conspecifics in the field.