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In the Mid-Session Reversal task (MSR), an animal chooses between two options, S1 and S2. Rewards follow S1 but not S2 from trials 1-40, and S2 but not S1 from trials 41-80. With pigeons, the psychometric function relating S1 choice proportion to trial number starts close to 1 and ends close to 0, with indifference (PSE) close to trial 40. Surprisingly, pigeons make anticipatory errors, choosing S2 before trial 41, and perseverative errors, choosing S1 after trial 40. These errors suggest that they use time into the session as the preference reversal cue. We tested this timing hypothesis with 10 Spotless starlings. After learning the MSR task with a T-s Inter-Trial Interval (ITI), they were exposed to either 2 T or T/2 ITIs during testing. Doubling the ITI should shift the psychometric function to the left and halve its PSE, whereas halving the ITI should shift the function to the right and double its PSE. When the starlings received one pellet per reward, the ITI manipulation was effective: The psychometric functions shifted in the direction and by the amount predicted by the timing hypothesis. However, non-temporal cues also influenced choice. published

Researchers have unraveled multiple cases in which behavior deviates from rationality principles. We propose that such deviations are valuable tools to understand the adaptive significance of the underpinning mechanisms. To illustrate, we discuss in detail an experimental protocol in which animals systematically incur substantial foraging losses by preferring a lean but informative option over a rich but non-informative one. To understand how adaptive mechanisms may fail to maximize food intake, we review a model inspired by optimal foraging principles that reconciles sub-optimal choice with the view that current behavioral mechanisms were pruned by the optimizing action of natural selection. To move beyond retrospective speculation, we then review critical tests of the model, regarding both its assumptions and its (sometimes counterintuitive) predictions, all of which have been upheld. The overall contention is that (a) known mechanisms can be used to develop better ultimate accounts and that (b) to understand why mechanisms that generate suboptimal behavior evolved, we need to consider their adaptive value in the animal’s characteristic ecology.

To contrast the classic version of the Scalar Expectancy Theory (SET) with the Behavioral Economic Model (BEM), we examined the effects of trial frequency on human temporal judgments. Mathematical analysis showed that, in a temporal bisection task, SET predicts that participants should show almost exclusive preference for the response associated with the most frequent duration, whereas BEM predicts that, even though participants will be biased, they will still display temporal control. Participants learned to emit one response (R[S]) after a 1.0-s stimulus and another (R[L]) after a 1.5-s stimulus. Then the effects of varying the frequencies of the 1.0-s and 1.5-s stimuli were assessed. Results were more consistent with BEM than with SET. Overall, this research illustrates how the impact of non-temporal factors on temporal discrimination may help us to contrast associative models such as BEM with cognitive models such as SET. Deciding between these two classes of models has important implications regarding the relations between associative learning and timing. This article is part of a Special Issue entitled: Associative and Temporal Learning., This research was supported by NIH grant MH033881. Jeremie Jozefowiez and Armando Machado acknowledge support from the Fundacao para a Ciencia e a Tecnologia as well as from the European project COST ISCH Action TD0904 "Time in Mental activity" (www.timely-cost.eu). We would like to thank Sean Gannon and Sarah Sterling for help running parts of the experiments, Mario Laborda, Bridget McConnell, Gonzalo Miguez, and James Witnauer for comments on an earlier version of this manuscript. Correspondence concerning this article should be addressed to Jeremie Jozefowiez, laboratoire URECA, Universite Lille Nord de France, Campus de Lille3, Domaine Universitaire du Pont de Bois, BP 60149, 58653 Villeneuve d'Ascq Cedex, France. jeremie.jozefowiez@univ-lille3fr.