Wild hummingbirds can use the geometry of a flower array

This research was supported in part by ASAB to SDH, and a Natural Sciences and Engineering Research Council of Canada Discovery Grant to TAH.Animals use cues from their environment to orient in space and to navigate their surroundings. Geometry is a cue whose informational content may originate from the metric properties of a given environment, and its use has been demonstrated in the laboratory in nearly every species of animal tested. However, it is not clear whether geometric information, used by animals typically tested in small, rectangular boxes, is directly relevant to animals in their natural environment. Here we present the first data that confirm the use of geometric cues by a free-living animal in the wild. We trained rufous hummingbirds to visit a rectangular array of four artificial flowers, one of which was rewarded. In some trials a conspicuous landmark cued the reward. Following array translocation and rotation, we presented hummingbirds with three tests. When trained and tested with the landmark, or when trained and tested without it, hummingbirds failed to show geometric learning. However, when trained with a landmark but tested without it, hummingbirds produced the classic geometric response, showing that they had learned the geometric relationships (distance and direction) of several non-reward visual elements of the environment. While it remains that the use of geometry to relocate a reward may be an experimental artefact, it is one cue that is not confined to the laboratory.PostprintPeer reviewe

Metabolic state alters economic decision making under risk in humans

Background: Animals' attitudes to risk are profoundly influenced by metabolic state (hunger and baseline energy stores). Specifically, animals often express a preference for risky (more variable) food sources when below a metabolic reference point (hungry), and safe (less variable) food sources when sated. Circulating hormones report the status of energy reserves and acute nutrient intake to widespread targets in the central nervous system that regulate feeding behaviour, including brain regions strongly implicated in risk and reward based decision-making in humans. Despite this, physiological influences per se have not been considered previously to influence economic decisions in humans. We hypothesised that baseline metabolic reserves and alterations in metabolic state would systematically modulate decision-making and financial risk-taking in humans. Methodology/Principal Findings: We used a controlled feeding manipulation and assayed decision-making preferences across different metabolic states following a meal. To elicit risk-preference, we presented a sequence of 200 paired lotteries, subjects' task being to select their preferred option from each pair. We also measured prandial suppression of circulating acyl-ghrelin (a centrally-acting orexigenic hormone signalling acute nutrient intake), and circulating leptin levels (providing an assay of energy reserves). We show both immediate and delayed effects on risky decision-making following a meal, and that these changes correlate with an individual's baseline leptin and changes in acyl-ghrelin levels respectively. Conclusions/Significance: We show that human risk preferences are exquisitely sensitive to current metabolic state, in a direction consistent with ecological models of feeding behaviour but not predicted by normative economic theory. These substantive effects of state changes on economic decisions perhaps reflect shared evolutionarily conserved neurobiological mechanisms. We suggest that this sensitivity in human risk-preference to current metabolic state has significant implications for both real-world economic transactions and for aberrant decision-making in eating disorders and obesity

Corvid Re-Caching without ‘Theory of Mind’: A Model

Scrub jays are thought to use many tactics to protect their caches. For instance, they predominantly bury food far away from conspecifics, and if they must cache while being watched, they often re-cache their worms later, once they are in private. Two explanations have been offered for such observations, and they are intensely debated. First, the birds may reason about their competitors' mental states, with a ‘theory of mind’; alternatively, they may apply behavioral rules learned in daily life. Although this second hypothesis is cognitively simpler, it does seem to require a different, ad-hoc behavioral rule for every caching and re-caching pattern exhibited by the birds. Our new theory avoids this drawback by explaining a large variety of patterns as side-effects of stress and the resulting memory errors. Inspired by experimental data, we assume that re-caching is not motivated by a deliberate effort to safeguard specific caches from theft, but by a general desire to cache more. This desire is brought on by stress, which is determined by the presence and dominance of onlookers, and by unsuccessful recovery attempts. We study this theory in two experiments similar to those done with real birds with a kind of ‘virtual bird’, whose behavior depends on a set of basic assumptions about corvid cognition, and a well-established model of human memory. Our results show that the ‘virtual bird’ acts as the real birds did; its re-caching reflects whether it has been watched, how dominant its onlooker was, and how close to that onlooker it has cached. This happens even though it cannot attribute mental states, and it has only a single behavioral rule assumed to be previously learned. Thus, our simulations indicate that corvid re-caching can be explained without sophisticated social cognition. Given our specific predictions, our theory can easily be tested empirically

Wild hummingbirds rely on landmarks not geometry when learning an array of flowers

Funding: Natural Sciences and Engineering Research Council of Canada (TAH, TAOF, DMZ) and the Association for the Study of Animal Behaviour (SDH).Rats, birds or fish trained to find a reward in one corner of a small enclosure tend to learn the location of the reward using both nearby visual features and the geometric relationships of corners and walls. Because these studies are conducted under laboratory and thereby unnatural conditions, we sought to determine whether wild, free-living rufous hummingbirds (Selasphorus rufus) learning a single reward location within a rectangular array of flowers would similarly employ both nearby visual landmarks and the geometric relationships of the array. Once subjects had learned the location of the reward, we used test probes in which one or two experimental landmarks were moved or removed in order to reveal how the birds remembered the reward location. The hummingbirds showed no evidence that they used the geometry of the rectangular array of flowers to remember the reward. Rather, they used our experimental landmarks, and possibly nearby, natural landmarks, to orient and navigate to the reward. We believe this to be the first test of the use of rectangular geometry by wild animals, and we recommend further studies be conducted in ecologically relevant conditions in order to help determine how and when animals form complex geometric representations of their local environments.PostprintPeer reviewe

Wild, free-living hummingbirds can learn what happened, where and in which context

Studies in the laboratory have shown that animals can combine multiple kinds of information to form integrated memories for rules and events. Less is known about how animals make use of these integrated memories in the wild. Here we tested whether wild, free-living, rufous hummingbirds, Selasphorus rufus, could learn to identify rewarded flowers in a naturalistic foraging situation, by remembering, over multiple exposures, what flower was rewarded, where and in which context. Birds were presented with boards on which four artificial flowers were mounted, one containing a food reward, the others containing water. Which flower (its colour and location) contained a reward was indicated in one condition by the presence of visually distinctive background boards and in a second condition by the sequential order in which the boards were presented. In both conditions, birds combined these pieces of information and learned to use the context to determine which of the four flowers was rewarded. Although they were not required to do so here, it is possible that these birds might be able to combine pieces of information to form integrated memories for single events. (C) 2014 The Association for the Study of Animal Behaviour. Published by Elsevier Ltd. All rights reserved.</p

Three-dimensional space:locomotory style explains memory differences in rats and hummingbirds

While most animals live in a three-dimensional world, they move through it to different extents depending on their mode of locomotion: terrestrial animals move vertically less than do swimming and flying animals. As nearly everything we know about how animals learn and remember locations in space comes from two-dimensional experiments in the horizontal plane, here we determined whether the use of three-dimensional space by a terrestrial and a flying animal was correlated with memory for a rewarded location. In the cubic mazes in which we trained and tested rats and hummingbirds, rats moved more vertically than horizontally, whereas hummingbirds moved equally in the three dimensions. Consistent with their movement preferences, rats were more accurate in relocating the horizontal component of a rewarded location than they were in the vertical component. Hummingbirds, however, were more accurate in the vertical dimension than they were in the horizontal, a result that cannot be explained by their use of space. Either as a result of evolution or ontogeny, it appears that birds and rats prioritize horizontal versus vertical components differently when they remember three-dimensional space

Wild, free-living rufous hummingbirds do not use geometric cues in a spatial task

In the laboratory, many species orient themselves using the geometric properties of an enclosure or array and geometric information is often preferred over visual cues. Whether animals use geometric cues when relocating rewarded locations in the wild, however, has rarely been investigated. We presented free-living rufous hummingbirds with a rectangular array of four artificial flowers to investigate learning of rewarded locations using geometric cues. In one treatment, we rewarded two of four flowers at diagonally opposite corners. In a second treatment, we provided a visual cue to the rewarded flower by connecting the flowers with “walls” consisting of four dowels (three white, one blue) laid on the ground connecting each of the flowers. Neither treatment elicited classical geometry results; instead, hummingbirds typically chose one particular flower over all others. When we exchanged that flower with another, hummingbirds tended to visit the original flower. These results suggest that (1) hummingbirds did not use geometric cues, but instead may have used a visually derived cue on the flowers themselves, and (2) using geometric cues may have been more difficult than using visual characteristics. Although hummingbirds typically prefer spatial over visual information, we hypothesize that they will not use geometric cues over stable visual features but that they make use of small, flower-specific visual cues. Such cues may play a more important role in foraging decisions than previously thought