Context-dependent foraging decisions in rufous hummingbirds

A core assumption implicit in economic models of animal choice is that subjects assign absolute utilities to options that are independent of the type and number of alternatives available. Humans sometimes appear to violate this assumption and employ relative, as opposed to absolute, currencies when making choices. Recent evidence suggests that animals too might sometimes employ relative choice mechanisms. We tested this idea by measuring the foraging preferences of rufous hummingbirds (Selasphorus rufus) faced with choices analogous to those in which human use of relative currencies is evident. The birds experienced three treatments: a binary choice between two artificial flower types designated concentration (20 microl, 40% sucrose solution) and volume (40 microl, 20%), and two trinary treatments in which a third decoy option (either concentration decoy: 10 microl, 30% or volume decoy: 30 microl, 10%) was added to the set. The birds' preferences differed significantly across the three treatments. In the trinary treatments, the effect of the decoy options was to increase the preference for the option that dominated the decoy. These results are similar to those reported in the human choice literature, and are compatible with the hummingbirds using a relative evaluation mechanism in decision making.     

Starlings uphold principles of economic rationality for delay and probability of reward

Rationality principles are the bedrock of normative theories of decision-making in biology and microeconomics, but whereas in microeconomics, consistent choice underlies the notion of utility; in biology, the assumption of consistent selective pressures justifies modelling decision mechanisms as if they were designed to maximize fitness. In either case, violations of consistency contradict expectations and attract theoretical interest. Reported violations of rationality in non-humans include intransitivity (i.e. circular preferences) and lack of independence of irrelevant alternatives (changes in relative preference between options when embedded in different choice sets), but the extent to which these observations truly represent breaches of rationality is debatable. We tested both principles with starlings (Sturnus vulgaris), training subjects either with five options differing in food delay (exp. 1) or with six options differing in reward probability (exp. 2), before letting them choose repeatedly one option out of several binary and trinary sets of options. The starlings conformed to economic rationality on both tests, showing strong stochastic transitivity and no violation of the independence principle. These results endorse the rational choice and optimality approaches used in behavioural ecology, and highlight the need for functional and mechanistic enquiring when apparent violations of such principles are observed.     

Context‐dependent female preference for multiple ornaments in the bearded reedling

While it is well established that females prefer to mate with well‐ornamented males, the influence of perceptive and cognitive processes on the expression of female mate choice is still poorly known. It has been suggested that the female perception of a male's attractiveness is not absolute, but depends on the other males with which he is compared that have been previously encountered (comparative evaluation). We investigated whether mate preference in bearded reedlings (Panurus biarmicus) is dependent on or independent of social context in relation to two different traits: beard and tail lengths. Each female had a choice between two to three males with different modifications of beard and tail. For each female, three different experiments were conducted (one binary and two trinary tests). We found that when females are presented with options that vary antagonistically with respect to two ornaments (binary test), some individuals prefer one trait while others the other trait. This indicates that in our bearded reedlings population exists a mate preference polymorphisms. Moreover, we found that the presence of a third stimulus, irrespective of the initial preference, reduced the strength of the initial preference – what we can call a “preference dilution effect.” Our results suggest that the female's choice may be constrained by her cognitive abilities when she is simultaneously presented with several options varying for two uncorrelated traits.     

Natural selection and context-dependent values

A simple model in which an animal makes a choice between two simultaneously available foraging options is used to show that we cannot expect natural selection to assign an absolute value (based on fitness) to each option. The example shows that the value of an option depends on its context; in particular, it depends on the option with which it is paired. This dependence results in a pattern of choice that violates a stochastic form of transitivity.     

Rationality in risk-sensitive foraging choices by starlings

Normative models of choice usually predict preferences between alternatives by computing their value according to some criterion, then identifying the alternative with greatest value. An important consequence of this procedure is captured in the economic concept of rationality, defined through a number of principles that are necessary for the existence of an ordinal scale of value upon which organisms base their choices. Violations of these principles, such as some recently reported breaches of transitivity and regularity in birds and honeybees, have strong implications for the understanding of decision mechanisms in humans and nonhumans alike. We investigated rationality in risky choice using European starlings, Sturnus vulgaris. Birds had to choose between two or three food sources, each associated with a different variance in delay to reward. In three experiments, starlings were strongly risk prone, showing regular and consistent preferences in binary and trinary choices. Preferences also satisfied weak and strong stochastic transitivity. Our results extend the generality of previous research in risk-sensitive foraging to situations where more than two alternatives are present and suggest that violations of rationality in risk-sensitive choices may be expressed only under restricted sets of conditions. Copyright 2002 The Association for the Study of Animal Behaviour. Published by Elsevier Science Ltd. All rights reserved.      

Irrational choices in hummingbird foraging behaviour

It is conventionally assumed that, when animals evaluate alternative options, the value assigned to an option is absolute and independent of the other options available. It follows that animal choices should exhibit the rational property of regularity whereby the proportion of choices for an option cannot be increased by the addition of further options to the choice set. However, violations of regularity occur in human decision making, suggesting that humans may use comparative evaluation mechanisms whereby the value of an option is computed relative to the other options available. For example, in the asymmetrically dominated decoy effect the preference for a target option over a competitor is altered by the addition of a decoy option that is inferior to the target and competitor on one attribute, but lies between them on a second. We tested whether foraging wild rufous hummingbirds, Selasphorus rufus, would demonstrate violations of regularity in response to an asymmetrically dominated decoy. Sixteen birds chose between three artificial flower types (Target: 15 μl, 40% sucrose; Competitor: 45 μl, 30%; Decoy: 10 μl, 35%) in Binary (Target versus Competitor) and Trinary (Target versus Competitor versus Decoy) treatments. We predicted higher preference for the Target in the Trinary treatment. The birds ranked the three options in the same order in the Binary and Trinary treatments (Competitor>Target>Decoy). Seven birds showed violations of regularity, six increasing their absolute preference for the Competitor in the Trinary treatment. Overall, relative preference for the Competitor over the Target was higher in the Trinary than in the Binary treatment. These changes in preference are incompatible with an absolute evaluation mechanism.     

Pigs’ preferences for rooting materials measured in a three-choice maze-test

The aim of this experiment was to investigate pigs’ preferences for rooting materials. Eighteen materials were allocated to six categories each of which consisted of three similar materials based on characteristics such as structure, size of particles, complexity, destructibility and digestibility. Twelve pairs of pigs chose among the three materials of each of the six categories in a balanced design. Within each category each pair was given four instantaneous choices among the three materials in a three-armed maze. ‘No choice’ was scored if the pigs did not enter one of the maze-arms within 90 s. Thus there were four options in each choice situation. The results were analysed using a random utility model incorporating random intercepts to account for the repeated testing of the same animals. The pigs expressed clear preferences within the category EARTH, where compost and peat were preferred to wood-shavings and no choice. In the category CHIP the most probable rank-order was spruce chip, willow chip, fir chip and no choice, while in the category ROUGH the most probable rank-order was maize-silage, grass silage, sugar beets and no choice. However, in these two categories none of the probabilities were sufficiently large to signify a preference for any of the three materials although the probabilities of the ‘no choice’ option were low. The pigs expressed no preferences among any of the four options including ‘no choice’ in the categories TOY (sisal robe, Bite-Rite, wooden beam), HAY (alfalfa hay mixed with straw, seed grass hay, barley straw with under-seed), and STRAW (long straw, chopped straw and straw pellets).     

I Choose,Therefore I Like: Preference for Faces Induced by Arbitrary Choice

Behavioral choice alters one’s preference rather than simply reflecting it. This effect to fit preferences with past choice, is known as “choice-induced preference change.” After making a choice between two equally attractive options, one tends to rate the chosen option better than they initially did and/or the unchosen option worse. The present study examined how behavioral choice changes subsequent preference, using facial images for the choice options as well as blind choice techniques. Participants rated their facial preference for each face, and chose between two equally preferred faces and subsequently rated their facial preference. Results from four experiments demonstrated that randomly chosen faces were more preferred only after participants were required to choose “a preferred face,” (in Experiment 1) but not “an unpreferred face,” (in Experiment 2) or “a rounder face” (in Experiment 3). Further, preference change was still observed after participants were informed that choices were actually random (in Experiment 4). Our findings provide new and important implications characterizing the conditions under which random choice changes preference, and show that people are tempted to make a biased evaluation even after they know that they did not make the choice for themselves.     

Female mate choice in convict cichlids is transitive and consistent with a self-referent directional preference

Introduction

One of the most important decisions that an animal has to make in its life is choosing a mate. Although most studies in sexual selection assume that mate choice is rational, this assumption has not been tested seriously. A crucial component of rationality is that animals exhibit transitive choices: if an individual prefers option A over B, and B over C, then it also prefers A over C.

Results

We assessed transitivity in mate choice: 40 female convict cichlids had to make a series of binary choices between males of varying size. Ninety percent of females showed transitive choices. The mean preference index was significantly higher when a female chose between their most preferred and least preferred male (male 1 vs. male 3) compared to when they chose between males of adjacent ranks (1 vs. 2 or 2 vs. 3). The results are consistent with a simple underlying preference function leading to transitive choice: females preferred males about one third larger than themselves. This rule of thumb correctly predicted which male was preferred in 67% of the cases and the ordering in binary choices in 78% of cases.

Conclusions

This study provides the first evidence for strong stochastic transitivity in a context of mate choice. The females exhibited ordinal preferences and the direction and magnitude of these preferences could be predicted from a simple rule. The females do not necessarily compare two males to choose the best; it is sufficient to use a self-referent evaluation. Such a simple decision rule has important implications for the evolution of the mating strategies and it is consistent with patterns of assortative mating repeatedly observed at population level.

     

Choice in multi-alternative environments: A trial-by-trial implementation of the Sequential Choice Model

The sequential choice model (SCM) proposes that latencies to accept options presented alone can be used to predict preferences between these options when they are presented simultaneously. SCM has been proposed and tested in experiments where only two alternatives were present. To further challenge the model, we trained and tested European starlings (Sturnus vulgaris) in an environment with a background of four alternatives differing in delay to reinforcement. Unexpected binary choices between the six possible pairs of alternatives were used to assess preference. The model's predictions of the strength of preference roughly corresponded to the bird's choices for each of the six choice situations. More importantly, a trial-by-trial test of the model correctly predicted 84% of all individual choice trials.     

Cognitive mechanisms of risky choice: is there an evaluation cost?

We contrast two classes of choice processes, those assuming time-consuming comparisons and those where stimuli for each option act independently, competing for expression by cross censorship. The Sequential Choice Model (SCM) belongs in the latter category, and has received empirical support in several procedures involving deterministic alternatives. Here we test this model in risky choices. In two treatments, each with five conditions, European starlings (Sturnus vulgaris) faced choices between options with unpredictable outcomes and risk-free alternatives. In the delay treatment the five conditions involved choices between a variable option offering two equiprobable delays to reward and a fixed option with delay differing between conditions. The amount treatment was structurally similar, but amount of reward rather than delay was manipulated. As assumed (and required) by the SCM, latency to respond in no-choice trials reflected each option's richness with respect to the background alternatives, and, crucially, preferences in simultaneous choices were predictable from latencies to each option in forced trials. However, we did not detect reliable differences in response times between forced and choice trials, neither the lengthening expected from evaluation models nor the shortening expected from the SCM.     

Genetic dissimilarity,genetic diversity,and mate preferences in humans

It is clear that genes at the major histocompatibility complex (MHC) are involved in mate preferences in a range of species, including humans. However, many questions remain regarding the MHC's exact influence on mate preference in humans. Some research suggests that genetic dissimilarity and individual genetic diversity (heterozygosity) at the MHC influence mate preferences, but the evidence is often inconsistent across studies. In addition, it is not known whether apparent preferences for MHC dissimilarity are specific to the MHC or reflect a more general preference for genome-wide dissimilarity, and whether MHC-related preferences are dependent on the context of mate choice (e.g., when choosing a short-term and long-term partner). Here, we investigated whether preferences for genetic dissimilarity are specific to the MHC and also whether preferences for genetic dissimilarity and diversity are context dependent. Genetic dissimilarity (number of alleles shared) influenced male, but not female, partner preferences, with males showing a preference for the faces of MHC-dissimilar females in both mating contexts. Genetic diversity [heterozygosity (H) and standardized mean (d²)] influenced both male and female preferences, regardless of mating context. Females preferred males with greater diversity at MHC loci (H) and males preferred females with greater diversity at non-MHC loci (d²) in both contexts. Importantly, these findings provide further support for a special role of the MHC in human sexual selection and suggest that male and female mate preferences may work together to potentially enhance both male and female reproductive success by increasing genetic diversity in offspring.     

Irrational decision-making in an amoeboid organism: transitivity and context-dependent preferences

Most models of animal foraging and consumer choice assume that individuals make choices based on the absolute value of items and are therefore ‘economically rational’. However, frequent violations of rationality by animals, including humans, suggest that animals use comparative valuation rules. Are comparative valuation strategies a consequence of the way brains process information, or are they an intrinsic feature of biological decision-making? Here, we examine the principles of rationality in an organism with radically different information-processing mechanisms: the brainless, unicellular, slime mould Physarum polycephalum. We offered P. polycephalum amoebas a choice between food options that varied in food quality and light exposure (P. polycephalum is photophobic). The use of an absolute valuation rule will lead to two properties: transitivity and independence of irrelevant alternatives (IIA). Transitivity is satisfied if preferences have a consistent, linear ordering, while IIA states that a decision maker''s preference for an item should not change if the choice set is expanded. A violation of either of these principles suggests the use of comparative rather than absolute valuation rules. Physarum polycephalum satisfied transitivity by having linear preference rankings. However, P. polycephalum''s preference for a focal alternative increased when a third, inferior quality option was added to the choice set, thus violating IIA and suggesting the use of a comparative valuation process. The discovery of comparative valuation rules in a unicellular organism suggests that comparative valuation rules are ubiquitous, if not universal, among biological decision makers.     

Context-dependent decisions among options varying in a single dimension

Contrary to theories of rational choice, adding alternatives to a choice set can change the choices made by both humans and animals. This is usually done by adding an inferior decoy to a choice set of two favoured options that are characterized on two distinct dimensions. We presented wild, free-living rufous hummingbirds (Selasphorus rufus) with choices between two or three options that varied in a single dimension only. The options varied in concentration, in volume or in corolla length. When the options varied in concentration, the addition of a medium option to a choice set of a low and a high concentration caused birds to increase their preference for the high option. However, they decreased their preference for the high concentration option when a low option was added to a choice set of high and medium concentrations. When the options varied only in volume, the addition of a high volume option to a choice set of low and medium options decreased the birds’ preference for the medium option. We saw no effects of adding a third option when the options varied in corolla length alone. Hummingbirds, then, make context-dependent decisions even when the options vary in only a single dimension although which effect occurs seems to depend on the dimension being manipulated. None of the current theories alone adequately explain these results.     

Fixation patterns in simple choice reflect optimal information sampling

Simple choices (e.g., eating an apple vs. an orange) are made by integrating noisy evidence that is sampled over time and influenced by visual attention; as a result, fluctuations in visual attention can affect choices. But what determines what is fixated and when? To address this question, we model the decision process for simple choice as an information sampling problem, and approximate the optimal sampling policy. We find that it is optimal to sample from options whose value estimates are both high and uncertain. Furthermore, the optimal policy provides a reasonable account of fixations and choices in binary and trinary simple choice, as well as the differences between the two cases. Overall, the results show that the fixation process during simple choice is influenced dynamically by the value estimates computed during the decision process, in a manner consistent with optimal information sampling.     

A fruit in the hand or two in the bush? Divergent risk preferences in chimpanzees and bonobos

Human and non-human animals tend to avoid risky prospects. If such patterns of economic choice are adaptive, risk preferences should reflect the typical decision-making environments faced by organisms. However, this approach has not been widely used to examine the risk sensitivity in closely related species with different ecologies. Here, we experimentally examined risk-sensitive behaviour in chimpanzees (Pan troglodytes) and bonobos (Pan paniscus), closely related species whose distinct ecologies are thought to be the major selective force shaping their unique behavioural repertoires. Because chimpanzees exploit riskier food sources in the wild, we predicted that they would exhibit greater tolerance for risk in choices about food. Results confirmed this prediction: chimpanzees significantly preferred the risky option, whereas bonobos preferred the fixed option. These results provide a relatively rare example of risk-prone behaviour in the context of gains and show how ecological pressures can sculpt economic decision making.     

Mating context and menstrual phase affect women's preferences for male voice pitch

Fundamental frequency (F₀) is the vocal acoustic parameter closest to what we perceive as pitch. Men speak at a lower F₀ than do women, even controlling for body size. Although the developmental and anatomical reasons for this sex difference are known, the evolutionary reasons are not. By examining fertility-related variation in women's preferences for men's voices, the present study tests the hypothesis that female choice for good genes influenced the evolution of male voice pitch (VP). Unlike previous correlational studies that did not consider the effects of menstrual phase and mating context on women's preferences for male VP, the present study includes these variables and utilizes experimental pitch (P) manipulations. Results indicate that low VP is preferred mainly in short-term mating contexts rather than in long-term, committed ones, and this mating context effect is greatest when women are in the fertile phase of their ovulatory cycles. Moreover, lower male F₀ correlated with higher self-reported mating success. These findings are consistent with the hypothesis that an association between low male VP and heritable fitness led to the evolution of the observed patterns in women's P preferences and men's mating success and that these patterns influenced the evolution of low VP in men. However, alternative explanations are considered.     

Too much of a good thing? Variety is confusing in mate choice

Choice variety is supposed to increase the likelihood that a chooser's preferences are satisfied. To assess the effects of variety on real-world mate choice, we analysed human dating decisions across 84 speed-dating events (events in which people go on a series of sequential 'mini-dates'). Results showed that choosers made fewer proposals (positive dating decisions) at events in which the available dates showed greater variety across such attributes as age, height, occupation and education, and this effect was particularly strong when choosers were confronted with a larger number of opposite-sex speed daters. Additionally, participants attending events in which the available options showed greater variety across these attributes were less likely to choose the consensually preferred mate option and more likely to choose no one at all. In contexts in which time is a limited resource, choice variety-rather than facilitating choice quality or increasing choosiness-is confusing and potentially detrimental to choice quality.     

Low-quality females prefer low-quality males when choosing a mate

Mate choice studies routinely assume female preferences for indicators of high quality in males but rarely consider developmental causes of within-population variation in mating preferences. By contrast, recent mate choice models assume that costs and benefits of searching or competing for high-quality males depend on females'' phenotypic quality. A prediction following from these models is that manipulation of female quality should alter her choosiness or even the direction of her mating preferences. We here provide (to our knowledge) the first example where an experimental manipulation of female quality induced a mating preference for low-quality males. Zebra finches (Taeniopygia guttata) reared in small or large experimental broods became high- or low-quality adults, respectively. Only high-quality females preferred high-quality males'' mate-advertising songs, while all low-quality females preferred low-quality males'' song. Subsequent breeding trials confirmed this pattern: latency until egg laying was shortest in quality-matched pairs, indicating that quality-matched birds were accepted faster as partners. Females produced larger eggs when mated with high-quality males, regardless of their own quality, indicating consensus regarding male quality despite the expression of different choices. Our results demonstrate the importance of considering the development of mating preferences to understand their within-population variation and environmentally induced change.     

Rational mate choice decisions vary with female age and multidimensional male signals in swordtails

Biologists have long been interested in intransitive preferences: circular preferences in which options cannot be ranked and no single option dominates, similar to a game of rock‐paper‐scissors. Intransitive preferences violate rational decision‐making, an assumption made by models of evolution by mate choice. Despite its potential importance in the study of sexual selection, few studies have tested for intransitive preferences. Even fewer have asked whether females differ in whether they choose mates transitively or intransitively and what factors might predict (in)transitive choice. Though intransitive choice is thought to be more common as options become more complex, this prediction is untested in animals. To fill this gap, we tested whether female Xiphophorus nigrensis swordtails can rank digitally animated males differing in size, courtship intensity, or both size and courtship intensity, and whether female responses were predicted by a female's age. Females choosing among males that varied only in size showed higher than expected levels of intransitivity, whereas females choosing among males that varied in their courtship or both properties did not. Older females were more likely to be irrational than younger females when evaluating male size, suggesting that experience modifies transitive decision‐making processes. These results show that mate choice irrationality may vary by a female's experience and the signal characteristics during decision‐making.