State-dependent learning and suboptimal choice: when starlings prefer long over short delays to food

Recent studies have used labels such as 'work ethics', 'sunk costs' and 'state-dependent preferences' for apparent anomalies in animals' choices. They suggest that preference between options relates to the options' history, rather than depending exclusively on the expected payoffs. For instance, European starlings, Sturnus vulgaris, trained to obtain identical food rewards from two sources while in two levels of hunger preferred the food source previously associated with higher hunger, regardless of the birds' state at the time of testing. We extended this experimentally and theoretically by studying starlings choosing between sources that differed not only in history but also in the objective properties (delay until reward) of the payoffs they delivered. Two options (PF and H) were initially presented in single-option sessions when subjects were, respectively, prefed or hungry. While option PF offered a delay until reward of 10 s in all treatments, option H delivered delays of 10, 12.5, 15 and 17.5 s in four treatments. When training was completed, we tested preference between the options. When delays in both options were equal (10 s), the birds strongly preferred H. When delay in H was 17.5 s, the birds were indifferent, with intermediate results for intermediate treatments. Preference was not mediated by disrupted knowledge of the delays. Thus, preferences were driven by past state-dependent gains, rather than by the joint effect of the birds' state at the time of choice and knowledge of the absolute properties of each alternative, as assumed in state-dependent, path-independent models of optimal choice.