The recollection of past experiences allows us to recall what a particular event was, and where and when it occurred, a form of memory that is thought to be unique to humans. It is known, however, that food-storing birds remember the spatial location and contents of their caches. Furthermore, food-storing animals adapt their caching and recovery strategies to the perishability of food stores, which suggests that they are sensitive to temporal factors. Here we show that scrub jays (Aphelocoma coerulescens) remember 'when' food items are stored by allowing them to recover perishable 'wax worms' (wax-moth larvae) and non-perishable peanuts which they had previously cached in visuospatially distinct sites. Jays searched preferentially for fresh wax worms, their favoured food, when allowed to recover them shortly after caching. However, they rapidly learned to avoid searching for worms after a longer interval during which the worms had decayed. The recovery preference of jays demonstrates memory of where and when particular food items were cached, thereby fulfilling the behavioural criteria for episodic-like memory in non-human animals.

Studies of psychiatric disorders have traditionally focused on emotional symptoms such as depression, anxiety and hallucinations. However, poorly controlled cognitive deficits are equally prominent and severely compromise quality of life, including social and professional integration. Consequently, intensive efforts are being made to characterize the cellular and cerebral circuits underpinning cognitive function, define the nature and causes of cognitive impairment in psychiatric disorders and identify more effective treatments. Successful development will depend on rigorous validation in animal models as well as in patients, including measures of real-world cognitive functioning. This article critically discusses these issues, highlighting the challenges and opportunities for improving cognition in individuals suffering from psychiatric disorders.

Cognition presents evolutionary research with one of its greatest challenges. Cognitive evolution has been explained at the proximate level by shifts in absolute and relative brain volume and at the ultimate level by differences in social and dietary complexity. However, no study has integrated the experimental and phylogenetic approach at the scale required to rigorously test these explanations. Instead, previous research has largely relied on various measures of brain size as proxies for cognitive abilities. We experimentally evaluated these major evolutionary explanations by quantitatively comparing the cognitive performance of 567 individuals representing 36 species on two problem-solving tasks measuring self-control. Phylogenetic analysis revealed that absolute brain volume best predicted performance across species and accounted for considerably more variance than brain volume controlling for body mass. This result corroborates recent advances in evolutionary neurobiology and illustrates the cognitive consequences of cortical reorganization through increases in brain volume. Within primates, dietary breadth but not social group size was a strong predictor of species differences in self-control. Our results implicate robust evolutionary relationships between dietary breadth, absolute brain volume, and self-control. These findings provide a significant first step toward quantifying the primate cognitive phenome and explaining the process of cognitive evolution.psychology | behavior | comparative methods | inhibitory control | executive function S ince Darwin, understanding the evolution of cognition has been widely regarded as one of the greatest challenges for evolutionary research (1). Although researchers have identified surprising cognitive flexibility in a range of species (2-40) and potentially derived features of human psychology (41-61), we know much less about the major forces shaping cognitive evolution (62-71). With the notable exception of Bitterman's landmark studies conducted several decades ago (63, 72-74), most research comparing cognition across species has been limited to small taxonomic samples (70, 75). With limited comparable experimental data on how cognition varies across species, previous research has largely relied on proxies for cognition (e.g., brain size) or metaanalyses when testing hypotheses about cognitive evolution (76-92). The lack of cognitive data collected with similar methods across large samples of species precludes meaningful species comparisons that can reveal the major forces shaping cognitive evolution across species, including humans (48,70,89,(93)(94)(95)(96)(97)(98). SignificanceAlthough scientists have identified surprising cognitive flexibility in animals and potentially unique features of human psychology, we know less about the selective forces that favor cognitive evolution, or the proximate biological mechanisms underlying this process. We tested 36 species in two problemsolving tasks measuring self-control and evaluated the leading hypotheses regarding how ...

Discussions of the evolution of intelligence have focused on monkeys and apes because of their close evolutionary relationship to humans. Other large-brained social animals, such as corvids, also understand their physical and social worlds. Here we review recent studies of tool manufacture, mental time travel, and social cognition in corvids, and suggest that complex cognition depends on a ''tool kit'' consisting of causal reasoning, flexibility, imagination, and prospection. Because corvids and apes share these cognitive tools, we argue that complex cognitive abilities evolved multiple times in distantly related species with vastly different brain structures in order to solve similar socioecological problems.

Knowledge of and planning for the future is a complex skill that is considered by many to be uniquely human. We are not born with it; children develop a sense of the future at around the age of two and some planning ability by only the age of four to five. According to the Bischof-Köhler hypothesis, only humans can dissociate themselves from their current motivation and take action for future needs: other animals are incapable of anticipating future needs, and any future-oriented behaviours they exhibit are either fixed action patterns or cued by their current motivational state. The experiments described here test whether a member of the corvid family, the western scrub-jay (Aphelocoma californica), plans for the future. We show that the jays make provision for a future need, both by preferentially caching food in a place in which they have learned that they will be hungry the following morning and by differentially storing a particular food in a place in which that type of food will not be available the next morning. Previous studies have shown that, in accord with the Bischof-Köhler hypothesis, rats and pigeons may solve tasks by encoding the future but only over very short time scales. Although some primates and corvids take actions now that are based on their future consequences, these have not been shown to be selected with reference to future motivational states, or without extensive reinforcement of the anticipatory act. The results described here suggest that the jays can spontaneously plan for tomorrow without reference to their current motivational state, thereby challenging the idea that this is a uniquely human ability.

According to the 'mental time travel hypothesis' animals, unlike humans, cannot mentally travel backwards in time to recollect specific past events (episodic memory) or forwards to anticipate future needs (future planning). Until recently, there was little evidence in animals for either ability. Experiments on memory in food-caching birds, however, question this assumption by showing that western scrub-jays form integrated, flexible, trial-unique memories of what they hid, where and when. Moreover, these birds can adjust their caching behaviour in anticipation of future needs. We suggest that some animals have elements of both episodic-like memory and future planning.

Social life has costs associated with competition for resources such as food. Food storing may reduce this competition as the food can be collected quickly and hidden elsewhere; however, it is a risky strategy because caches can be pilfered by others. Scrub jays (Aphelocoma coerulescens) remember 'what', 'where' and 'when' they cached. Like other corvids, they remember where conspecifics have cached, pilfering them when given the opportunity, but may also adjust their own caching strategies to minimize potential pilfering. To test this, jays were allowed to cache either in private (when the other bird's view was obscured) or while a conspecific was watching, and then recover their caches in private. Here we show that jays with prior experience of pilfering another bird's caches subsequently re-cached food in new cache sites during recovery trials, but only when they had been observed caching. Jays without pilfering experience did not, even though they had observed other jays caching. Our results suggest that jays relate information about their previous experience as a pilferer to the possibility of future stealing by another bird, and modify their caching strategy accordingly.

The ‘social intelligence hypothesis’ was originally conceived to explain how primates may have evolved their superior intellect and large brains when compared with other animals. Although some birds such as corvids may be intellectually comparable to apes, the same relationship between sociality and brain size seen in primates has not been found for birds, possibly suggesting a role for other non-social factors. But bird sociality is different from primate sociality. Most monkeys and apes form stable groups, whereas most birds are monogamous, and only form large flocks outside of the breeding season. Some birds form lifelong pair bonds and these species tend to have the largest brains relative to body size. Some of these species are known for their intellectual abilities (e.g. corvids and parrots), while others are not (e.g. geese and albatrosses). Although socio-ecological factors may explain some of the differences in brain size and intelligence between corvids/parrots and geese/albatrosses, we predict that the type and quality of the bonded relationship is also critical. Indeed, we present empirical evidence that rook and jackdaw partnerships resemble primate and dolphin alliances. Although social interactions within a pair may seem simple on the surface, we argue that cognition may play an important role in the maintenance of long-term relationships, something we name as ‘relationship intelligence’.