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Behavioral Ecology Vol. 12 No. 3: 318-324
© 2001 International Society for Behavioral Ecology

Background context and decision making in hoarding gray jays

Thomas A. Waite

Department of Evolution, Ecology, and Organismal Biology, and Department of Anthropology, Ohio State University, Columbus, OH 43210-1293, USA

Address correspondence to T.A. Waite. E-mail: waite.1{at}osu.edu .

Received 2 October 1999; revised 22 July 2000; accepted 12 September 2000.


    ABSTRACT
 TOP
 ABSTRACT
 INTRODUCTION
 METHODS
 RESULTS
 DISCUSSION
 REFERENCES
 
If decision makers assign stable fitness-related values to options, preference for the most valuable of simultaneously encountered options should be independent of background context (i.e., prior options). The tendency to choose optionx versus y should be unaffected by whether the decision maker has already been given a choice betweenx' and y' or between x'' and y''. Here, food-hoarding gray jays (Perisoreus canadensis) were given an initial choice between x' (one raisin, 0.5 m into a tube) and y' (three raisins, 0.5 m) or between x'' and y'' (both identical to x'). All subjects were then given a choice between x' (one raisin, 0.3 m) and y' (three raisins, 0.7 m). In violation of the principle of irrelevant alternatives, the "market share" ofx depended on prior options. Subjects initially exposed to context {x', y'} showed a stronger preference for x than did subjects initially exposed to {x'', y'}, which implies that the jays did not assign a fixed value to each option. Subjects that initially could obtain a large reward (y') for about the same "price" (perceived danger) as a small reward (x') apparently devalued the large reward (y) in the subsequent choice. This effect may be the joint byproduct of cognitive constraints and an adaptive tendency to use information provided by the context.

Key words: choice, cognitive constraints, context, decision making, hoarding, information processing.


    INTRODUCTION
 TOP
 ABSTRACT
 INTRODUCTION
 METHODS
 RESULTS
 DISCUSSION
 REFERENCES
 
Natural selection is often assumed to favor choice behavior that maximizes fitness. This conventional view suggests, misleadingly (Houston, 1997Go; Ydenberg, 1998Go), that decision makers assign a fixed fitness-related value to each option. By extension, an animal choosing among simultaneously encountered options is expected to choose the most valuable option. Assuming natural selection favors minimization of costly errors, partial rather than absolute preference is expected (McNamara and Houston, 1987Go; Waite and Field, 2000Go), but the strength of this preference should be independent of the background context (i.e., options encountered in the past). The tendency to prefer option x over y in simultaneous encounters should be unaffected by whether the decision maker has already experienced encounters with options x' and y' or options x'' and y''. This prediction is consistent with the principle of irrelevant alternatives from choice theory (Tversky and Simonson, 1993Go) and with models of simultaneous choice from foraging theory (Stephens and Krebs, 1986Go). Thus, according to standard theories of rational choice and optimal foraging, the "attractiveness" of an option should be determined by its real (fitness-related) value, which should be independent of prior options.

Despite the parsimony of this framework, decision-making processes are known to be subject to background context effects, particularly in human consumers. For example, Simonson and Tversky (1992Go) studied choice of products (automobile tires) varying on two attributes (warranty and price). They found that subjects were more likely to choose a low-quality cheap option over a high-quality expensive option if they had already been given a choice in which a comparable difference in quality was associated with a smaller difference in price. Subjects were given an initial choice between options x' (55,000 miles, $85) and y' (75,000 miles, $91) or between x'' (30,000 miles, $25) and y'' (35,000 miles, $49). All subjects were then given a choice between x (40,000 miles, $60) and y (50,000 miles, $75). Subjects exposed to background B' = {x', y'} showed a stronger preference for x (57%) than did subjects exposed to B'' = {x'', y''} (33%). In violation of the principle of irrelevant alternatives (i.e., contrary to the normative economic expectation of rational choice theory), the subjective valuations of options x and y apparently depended on options available in the past. This seemingly maladaptive effect was interpreted as the byproduct of a cognitive illusion or bias (Simonson and Tversky, 1992Go; Tversky and Simonson, 1993Go), a systematic and directional error of judgment in violation of economic value maximization (for detailed definitions and extensive reviews of so-called cognitive illusions [including the conjunction fallacy, base-rate error, Concorde fallacy, and single-event probability problem] see Anderson, 1998Go; Fantino, 1998Go; Griffin and Buehler, 1999Go; Kahneman and Tversky, 1996Go; Nichols, 1999Go; Piattelli-Palmarini, 1994Go). Alternatively, from an evolutionary perspective, such effects are interpretable as an artifact of imposing unnecessarily narrow norms of rational choice and presenting subjects with contrived choice tasks (Gigerenzer, 2000Go). From this perspective, background context effects are even potentially representative of rational (Wernerfelt, 1995Go) or adaptive (Houston, 1997Go) use of information provided by the context (reviewed by Anderson, 1998Go; Giraldeau, 1997Go).

Here, I describe an analogous experiment designed to determine whether a background context effect could be induced in a non-human decision maker, the gray jay (Perisoreus canadensis). Jays collecting food for storage were given an initial choice between options x' (small reward) and y' (large reward) or between x'' and y'' (both identical to x'). All of these initial options could be obtained for an intermediate "price." All subjects were then given a choice between a small cheap reward (x) and a large expensive reward (y). According to classical theories of rational choice (Bell et al., 1988Go) and optimal foraging (Stephens and Krebs, 1986Go), the tendency to choose x versus y should be unaffected by any such manipulation of background context. Evidence to the contrary would challenge the notion that non-human animals are virtually immune to cognitive illusions because their decision making processes are shaped by natural selection (see also Arkes and Ayton, 1999Go).


    METHODS
 TOP
 ABSTRACT
 INTRODUCTION
 METHODS
 RESULTS
 DISCUSSION
 REFERENCES
 
Subjects and study area
The gray jay (Perisoreus canadensis), a year-round resident of boreal forests of North America, lives in social groups on all-purpose territories (see Strickland and Ouellet, 1993Go for a detailed account; see also Waite and Strickland, 1997Go). During summer and autumn, jays routinely make hundreds of food caches per day, placing each saliva-coated bolus in a separate arboreal site (Waite and Reeve, 1993Go, 1994Go). They rely on these caches throughout the winter and even use them to provision young.

Twenty semi-tame, free-ranging gray jays, identifiable by unique color-band combinations, were used as subjects. These subjects comprised seven adult females and eight adult males, two philopatric juvenile females, two philopatric juvenile males, and one immigrant (unrelated) juvenile female.

The experiment was conducted in Algonquin Provincial Park, Ontario, Canada (45°33' N, 78°38' W; see Strickland, 1991Go, for detailed description) between 22 and 26 October 1998. All tests were conducted between 0750 and 1757. Air temperature varied between -2 and 15°C. No precipitation occurred during the tests.

Experimental procedure
The experiment was designed to measure the effect of manipulated background context on binary choice. Options varied along two attributes, quality (number of raisins) and price (distance into a tube). Presumably, the jays perceived greater danger with greater distance into the tube, as suggested by their hesitant sidling into the tubes, especially during performance tests (see below). By entering farther into a tube, a jay's ability to escape the surprise attack of a predator would be compromised. Instead of instantly flying to cover upon detecting the oncoming predator, the jay would have to hop to the opening of the tube and then flee. The choice task thus measured the jays' tendency to choose a low-quality, safe option versus a high-quality, dangerous option as influenced by background context. Defined by previously encountered options, the background context was manipulated as described below.

The experimental set-up consisted of two tubes, both containing food (Figure 1). Subjects were required to hop into either tube during each visit to obtain the food reward. The semi-cylindrical (radius = 25 cm) tubes were 1.2-m long, made of 1-cm welded wire mesh (hardware cloth), and closed at one end. The open ends were situated equidistant from a standard 25-cm-high perch. From this perch, subjects could inspect the contents of the two tubes simultaneously. Food was placed on white plastic discs (diameter = 17.8 cm), positioned at specified distances into the tubes. Immediately upon each arrival, the subject could choose between the two options. As soon as the jay collected the food (one or three raisins) and flew to hoard it in (one or three) nearby arboreal sites, the food was replenished (as described below) before the jay returned to the set-up, usually within a minute. Detailed descriptions of hoarding behavior in this species are published elsewhere (Waite and Ydenberg, 1996Go, references therein).



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Figure 1 Top view of experimental set-up showing simultaneously available but mutually exclusive options under background set B', background set B'', and the target set. Rectangles represent 1.2-m-long tubes made of welded-wire mesh, open circles within the tubes represent white plastic discs, small filled circles represent the food reward (raisins), and filled circles at the open ends of the tubes represent a perch from which the subject could simultaneously inspect the contents of the two tubes. The positioning of the two options was switched after each visit by the subject. See Methods for additional details.

 

To test a particular individual, I positioned myself near the known core of its territory and whistled to attract it. I then conducted a performance test in which the prospective subject was given a choice between identical options (one raisin, same distance into tube). During this 15-visit session, raisins were positioned 0.3 m into the tubes during the first five visits, 0.5 m during the next five visits, and 0.7 m during the final five visits. To train the subjects to treat the task as an exclusive binary choice, I flushed any jay that attempted to enter both tubes during a given visit. No individual attempted to do so more than twice. All individuals passed the performance test, entering one of the tubes during each visit, collecting the raisin and transporting it to a nearby arboreal site for storage. Throughout this and subsequent tests, raisins were offered to all accompanying jays to minimize the subject's tendency to adjust its choice behavior in response to interference competition (Waite and Ydenberg, 1996Go). Following this test, the set-up was moved ~60 m to minimize the subject's tendency to adjust load size in response to density-dependent risk of cache theft (Waite and Ydenberg, 1996Go).

Following a 10-min hiatus, the subject was exposed to background context B' or B'' (Figure 1) during each of 25 consecutive visits. Half the subjects were exposed to B' and the other half were exposed to B''. To avoid an order effect, background context was randomly assigned for the first subject and then alternated across successive subjects. Under B', subjects were given a choice between option x' (one raisin, 0.5 m into tube) and y' (three raisins, 0.5 m). Under B'', subjects were given a choice between x'' and y'' (both identical to x'). To avoid a side bias, the positioning of the rewards was randomized initially and then switched after each visit (even for B''). After the last visit, the set-up was moved (~60 m).

The target-set test (Figure 1) was then conducted, following a 10-min intertrial interval. During this test, all subjects were given a choice between x (one raisin, 0.3 m) and y (three raisins, 0.7 m) during each of 25 consecutive visits. The positioning of x and y was randomized initially and then switched after each visit. Each subject was tested once, following the initial 25-visit exposure to either B' or B''. (Beginning with the onset of the background test, B' subjects completed the experiment [50 hoarding trips] in 44 min [SD = 6.2], on average, while B'' subjects did so in 56 min [SD = 34.3].) I thus used a between-subjects design in which 10 subjects were tested following exposure to context B' and 10 other subjects were tested following exposure to B''.

Data analysis
Analyses used SigmaStat (1997Go) routines. The Mann-Whitney U test was used to test whether the tendency to choose x rather than y differed between subjects previously exposed to background B'' versus those previously exposed to B''. Wilcoxon signed-ranks tests were used to test whether the tendency to choose x' or x'' during the background-set test differed from the tendency to choose x in the target-set test. Wilcoxon tests were also used to evaluate any preferences for x' versus y', x'' versus y'', and x versus y. These tests compared the observed proportion of x', x'', or x choices with random expectation (0.5). Finally, Spearman's rank correlation was used to evaluate the degree of autocorrelation (lag = 1) in each subject's 25 consecutive choices during the target-set test. For each series of choices (where x = 0 and y = 1), C1, 2,..., 25, the correlation between Ct and Ct-1 was calculated. For any subject whose choices were autocorrelated (at the 0.05{alpha}-level), logistic regression was used to evaluate whether the subject's tendency to choose x versus y increased or decreased across the 25 consecutive choices. All tests were two-tailed.


    RESULTS
 TOP
 ABSTRACT
 INTRODUCTION
 METHODS
 RESULTS
 DISCUSSION
 REFERENCES
 
The tendency to choose option x over y was stronger among subjects initially exposed to background B' than among those initially exposed to B'' (U = 153.5, p <.001; Figure 2). In fact, nine of the 10 subjects exposed to B' showed a stronger preference for x than did all 10 subjects exposed to B''. Subjects exposed to B' showed a preference for y' over x' during the background-set test (median proportion y' = 0.74; Z = 2.706, p =.007; Table 1), but showed a preference for x over y (median proportion x = 0.92; Z = 2.812, p =.005; Figure 2) during the subsequent target-set test. By contrast, subjects exposed to B'' showed no significant preference during either the background-set test (median proportion x'' = 0.5; Z = -0.36, p =.72; Table 1) or target-set test (median proportion x = 0.62; Z = 1.381, p =.17; Figure 2). Among subjects exposed to B', the tendency to choose x (median = 0.92; Figure 2) differed significantly from the prior tendency to choose x' (median = 0.26; Z = 2.807, p =.005; Table 1). However, among subjects exposed to B'', the tendency to choose x (median = 0.62; Figure 2) did not differ significantly from the prior tendency to choose x'' (median = 0.5; Z = 1.227, p =.22; Table 1).



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Figure 2 Proportion of (25) visits during the target-set tests in which subjects chose option x rather than option y. Ten subjects had been exposed to background context B' and 10 others to B''. See Figure 1 and Methods for additional details. Horizontal lines represent the median, boxes encompass the 25th through 75th percentiles, and circles indicate the minimum and maximum observed proportions.

 

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Table 1 Proportion choice x (median, 25th-75th percentiles) during background-set test
 

No significant autocorrelation in the tendency to choose x versus y was detected for any of the subjects previously exposed to background B'' (all p's >.15), and only one significant autocorrelation was found among the subjects previously exposed to background B'' (rs = -.45, p =.026; all other p's >.07). Logistic regression was used to detect any (increasing or decreasing) trend in this subject's tendency to choose x versus y across the 25-visit target-set test. No strong trend was suggested by the odds ratio (0.59), an estimate of the increase in odds of choosing y in the next visit. The 60% confidence interval (0.47, 0.75) of the odds ratio encompassed 0.5, which implies that the subject's tendency to choose x versus y did not change significantly across the 25 visits in the target-set test.


    DISCUSSION
 TOP
 ABSTRACT
 INTRODUCTION
 METHODS
 RESULTS
 DISCUSSION
 REFERENCES
 
Contrary to classical theories of rational choice, the jays' choice behavior was influenced by background context. According to the principle of irrelevant alternatives, the market share of an option should be independent of other options, including those encountered in the past (Tversky and Simonson, 1993Go). While minimizing costly errors (Houston, 1997Go; Waite and Field, 2000Go), subjects should prefer the most valuable of simultaneously encountered options, and the strength of this preference should be independent of past alternatives. In violation of this principle, jays differed in their tendency to choose option x versus y depending on whether they had been exposed to background B' or B'' (Figure 2). This finding is surprising given the body of work showing that gray jays often make decisions that closely match predictions based on the rate-maximization currency (Waite and Ydenberg, 1994aGo, bGo, 1996Go, and references therein). The observed effect of background context implies that the jays' subjective valuation of options was not absolute or fixed. Instead, their perception of an option's value apparently depended on theoretically irrelevant options encountered in the past. The influence of recent experience on foraging decisions is well known (reviewed by Giraldeau, 1997Go), but unlike the present study, these studies have focused on the role of information acquisition via statistical sampling in stochastic environments (see further discussion below).

The ideas that rewards have relative as well as absolute effects on choice behavior and that these effects depend on an animal's recent experience have been long recognized, though largely overlooked, in the psychological literature (see Flaherty, 1996Go for a fascinating historical account). For example, in one experiment in Crespi's (1942Go) classic study, rats were trained to obtain a food reward at the end of a runway. Three groups of rats were differentially rewarded during the first day of the experiment. Subjects in one group received a 256-unit reward (5.12 g of dog food), those in another group received a 64-unit reward, and those in a control group received a 16-unit reward. Mean performance (running speed) was highest in subjects receiving the 256-unit reward, intermediate in those receiving the 64-unit reward, and lowest in the control subjects. Beginning on the second day of the experiment, all subjects received a 16-unit reward. The running speed of the subjects that had been shifted downward from 256 or 64 units was compared with that of the control subjects. Paradoxically, the shifted subjects abruptly reduced their running speed below that of the control subjects, even though all subjects now received identical 16-unit rewards. This so-called negative contrast effect persisted throughout the experiment (i.e., over the subsequent 7 days). Although the experimental procedures and conceptual framework of this study differed from those of the present study, the findings are nonetheless reminiscent of the observed effect of prior options on the jays' behavior and are likewise difficult to explain from a functional perspective.

More reminiscent of the present results, and perhaps more paradoxical, are the results of recent work on choice behavior in pigeons (reviewed by Shettleworth, 1998Go). In one experiment (Belke, 1992Go), pigeons were trained on two pairs of concurrent variable interval (VI) schedules. (On a VI schedule, intervals between successive reinforcements vary randomly around a specified [mean] VI value.) In alternating periods, subjects preferred option VI:20 s (white key) over VI:40 (red) and option VI:40 (green) over VI:80 (yellow), where the numerals represent the mean interval between successive rewards. Not surprisingly, in both pairs of schedules, subjects preferred the option with the shorter mean delay to reward (i.e., VI:20-white over VI:40-red, and VI:40-green over VI:80-yellow). However, when offered the choice between red and green, the subjects strongly preferred green, even though both colors signaled VI:40 s and hence the principle of value (or rate) maximization would predict no preference.

Even more paradoxical are the results of a follow-up experiment (Gibbon, 1995Go) in which pigeons, beyond showing a preference where none was expected, showed a preference opposite that predicted by value maximization. Specifically, the pigeons strongly preferred the equivalent of VI:40-green over VI:20-white. Thus, the VI:40 option from the VI:40,VI:80 background context was more "attractive" than the VI:20 option from the VI:20,VI:40 background context, even though the delay to reward was twice as long for the preferred option. While it is conceivable that the jays' choice behavior is representative of the usually adaptive use of information provided by recent experience (see below), the pigeons' choice behavior in Gibbon's experiment seems to have no reasonable functional explanation. Because the pigeons' preference directly contradicts value maximization, only a blithe adaptationist would be quick to interpret it as being somehow adaptive. After all, how could it be adaptive to respond to recent experience by strongly and persistently preferring an option that is only half as profitable as an alternative? Instead, this finding suggests that background context may produce a cognitive illusion or bias, a directional error of judgment in violation of value maximization (for reviews see Anderson, 1998Go; Fantino, 1998Go; Griffin and Buehler, 1999Go; Kahneman and Tversky, 1996Go; Nichols, 1999Go; Piattelli-Palmarini, 1994Go; but see Gigerenzer, 1996).

The effect of background context observed in the gray jays, though less dramatic, is likewise inconsistent with value maximization. It is consistent, though, with a descriptive (nonfunctional) hypothesis of choice behavior in human consumers. The tradeoff contrast hypothesis (Tversky and Simonson, 1993Go) predicts an increased tendency to choose a low-quality cheap option over a high-quality expensive option if the decision maker has already been given a choice in which a comparable difference in quality was associated with a smaller difference in price. In agreement with this prediction, gray jays showed a strong preference for a low-quality cheap option (Figure 2) if they had already been given a choice in which the same difference in quality was associated with no difference in price (Figure 1). Subjects initially offered a three-for-the-price-of-one deal showed a strong subsequent preference for the small reward. Thus, the large but now expensive reward was less "attractive" on this background. A higher-than-background difference between the price of two options, as in the present experiment, may prompt the decision maker to undervalue the higher-quality option. If fixed values were assigned to each option, decision makers would be immune to such effects, which occur apparently because the background influences the subjective valuation of options.

To explore more formally the potential influence of background context on subjective valuation, I describe here a model developed to explain context effects in human consumers (Tversky and Simonson, 1993Go). The purpose is not to test the model, but to explore whether conditions like those in the present experiment could lead to paradoxical preferences and to generate predictions to be tested in future experiments. This modeling exercise reinforces the notion that models of decision making should recognize that an option's fitness value is context dependent rather than fixed (Houston, 1997Go) and that cognitive biases can lead to paradoxical departures from value maximization. Whatever the underlying mechanisms, the model shows how comparative evaluation (Shafir, 1994Go) of options varying along multiple dimensions can lead to paradoxical effects of background context on choice. By contrast, standard theories of choice cannot account for such effects.

For a given background B, the decision rule is to chose x from option set S = {x, y} if and only if the subjective value VB(x, S) exceeds VB(y, S) (see Table 2 for definitions of symbols). As done elsewhere (Waite and Field, 2000Go), it would be straightforward to incorporate the notion that decision-making processes are inevitably subject to error (McNamara and Houston, 1987Go), particularly when the fitness consequences of erroneous choice are trivial (Houston, 1997Go). Future work, both theoretical and experimental, should explore the possibility that paradoxical effects of context are more common when the fitness consequences of such erroneous choice are smaller. Animals may be especially vulnerable to background effects when the fitness consequences are trivial. However, to simplify the present analysis, the decision maker is expected to choose—repeatedly and consistently—the option with higher subjective value:

(1)
where v(x) is the true (fitness-related, context-free) value of x, the second term describes the effect of background context, and the third term describes the effect of local context (i.e., the choice set S currently available). Elsewhere (Shafir S, Waite TA, Smith B, unpublished data), gray jays and honey bees were shown to violate the principle of irrelevant alternatives in response to a change in local context (i.e., the addition of an option to the choice set) (see also Hurly and Oseen, 1999Go). With no apparent adaptive basis (and in violation of the condition termed regularity; Huber et al., 1982Go; Tversky and Simonson, 1993Go), the jays responded to the introduction of an "unattractive" option by increasing their absolute preference for the more similar (less preferred) of the two original alternatives. This local context effect strengthens the inference from the present findings that the jays do not assign fixed values to options. Instead, they apparently use a comparative method of evaluation, assigning flexible values to options based on comparisons with current (Hurly and Oseen, 1999Go; Shafir, 1994Go; Shafir et al., 1989Go; Tversky, 1969Go; Waite, 2001Go) and past (this study) alternatives.


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Table 2 Symbols used in model of context-dependent choice
 

Here, I explore the flexible valuation of options as influenced by background context alone. To isolate the effect of background context, I eliminate the local-context component of the model:

(2)
This simplification is consistent with the design of the present experiment, where the target set (Figure 1) comprised a single local context {x, y}. Assuming the attributes (quality and price) of each option contribute additively to the option's real value and to the effect of background, the subjective value of an option is described by:

(3)
where vi is the real value along attribute q (quality) or p (price), bi is the relative weight of the background effect along q or p (satisfying bq + bp = 1), and ß controls the overall weight of the background. Here, I use this expression to explore the influence of these parameters on the subjective values of options in a binary choice set.

Table 3 summarizes two numerical examples. In the first example, the real values of option x (= vq(xq) + vp(xp) = 1 + 3) and y (= vq(yq) + vp(yp) = 3 + 1) are identical. For ß = 0, the subjective value of an option equals its real value (Equation 3 reduces to value maximization). Therefore, the subjective value of the options is identical in the two contexts and no preference is predicted. However, for ß = 10, the subjective value of x exceeds that of y on background B', reflecting the greater weighting of price (bp = 0.75) under that context. The model predicts a preference for x under B', but no preference under B''. In the second example, the real value of option y (= 3 + 1) exceeds that of x (= 1 + 2). Thus, for ß = 0, the subjective value of y exceeds x (by the same degree in both contexts). However, for ß = 10, the model predicts a preference for x under B' and for y under B''. This preference reversal reflects the interaction between the asymmetry in true value and the greater relative weighting of price under B'. Figure 3 illustrates the effects of variation in ß and bp where the true values of options x and y are identical. As ß approaches zero (Figure 3a), or bp approaches 0.5 (Figure 3b), the difference in the subjective values of x and y declines linearly to zero. Thus, the model generates predictions (e.g., Figure 3b) that are qualitatively consistent with the measured effect of background context in our experiment (Figure 2). For animals faced with the choice between options varying along two dimensions (e.g., quality and price), as in our experiment, background context has the potential of inducing dramatic shifts and even reversals in preference.


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Table 3 Numerical examples of predicted effect of background context on subjective values of options (Equation 3)
 


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Figure 3 Numerical example of the subjective value, V(·), of options x and y, and the subjective advantage of option x over y, A(x, y) = V(x) — V(y), as a function of: (a) the induced weight of the background context, ß, and (b) the weight of the price, bp. Lines generated by solving Equation 3. Assumed parameter values: both (a) and (b), bq = 0.25, vq(x) = 1, vq(y) = 3, vd(x) = 3, and vd(y) = 1; (a) only, bp = 0.75; and (b) only, ß = 5.

 

Although it may be tempting to view such paradoxical effects strictly as byproducts of intrinsic cognitive biases (Tversky and Simonson, 1993Go), these apparent deficits could have an adaptive basis (Houston, 1997Go), especially when decisions are made under uncertainty (e.g., Anderson, 1998Go; Bateson and Kacelnik, 1996Go; Dall et al., 1999Go; Kamil et al., 1993Go; Sih, 1992Go; Stephens, 1989Go; reviewed by Dukas, 1998Go; Giraldeau, 1997Go). Animals may routinely cope with uncertainty about tradeoffs between attributes by using information provided by the background. If so, background effects may be especially dramatic during early stages of exposure to novel tasks, as in the present experiment. If an animal in a stochastic environment uses information to update its valuation of options (Giraldeau, 1997Go), the influence of background may become less dramatic as the animal gains experience with an array of option sets (equivalently, ß may become smaller). (From this perspective, because the jays were exposed to a single background context and so were not able to acquire information from a stochastic array of option sets, it is not surprising that the jays' preferences were steady throughout the target-set test.) Similarly, as an animal gains experience with a choice involving a quality-price tradeoff, the relative weighting of price (danger) may decline (bp may become smaller) as the animal gains experience, especially if the animal habituates to the danger. (The jays' perception of danger may have stabilized during the performance and background-set tests.) If so, an animal may become increasingly willing to pay a high price for a given valuable reward (Figure 3) (cf. Lima and Bednekoff, 1999Go). Thus, background context effects may be common if animals making tradeoffs between attributes initially tend to overweight the background (see above) or overestimate the predation risk (as theorized by Abrams, 1995Go; Bouskila et al., 1995Go, and references therein).

This speculation prompts a testable prediction: if animals faced with simultaneous choice under uncertainty and predation risk behave as if they weight background or price less heavily as they acquire information, the effect of background should decline with experience. The background effect should be weakened by increased exposure to an array of background option sets, and choice behavior should converge on value maximization as assigned values and hence preferences become more stable. Moreover, closer convergence on value maximization should be expected in situations where the fitness costs of erroneous choice are higher (see Houston, 1997Go; Waite and Field, 2000Go). Consistent with the theme of the emerging field of cognitive ecology (Dukas, 1998Go; Giraldeau, 1997Go; Pepperberg et al., 1998Go; Real, 1991Go, 1993Go; Shettleworth, 1998Go), experimental tests of these predictions could help resolve the conflict between the view that context effects are cognitive illusions (Tversky and Simonson, 1993Go) versus the view that such effects are artifactual (Gigerenzer, 2000Go; see also Anderson, 1998Go) or may even represent rational (Wernerfelt, 1995Go) or adaptive (Houston, 1997Go) use of information provided by the context.


    ACKNOWLEDGEMENTS
 
This article is dedicated to Dan Strickland—naturalist and corvidophile extraodinaire—on the occasion of his (nominal) retirement. I thank Dan for three decades' worth of background information, Sharoni Shafir for background context (inspiration and prior collaboration), John Vucetich for discussion, and two anonymous reviewers for valuable comments on the manuscript.


    REFERENCES
 TOP
 ABSTRACT
 INTRODUCTION
 METHODS
 RESULTS
 DISCUSSION
 REFERENCES
 
Abrams PA, 1995. Overestimation versus underestimation of predation risk: a reply to Bouskila et al. Am Nat 145: 1020-1024.

Anderson JL, 1998. Embracing uncertainty: the interface between Bayesian statistics and cognitive psychology. Cons Ecol [online] 2: 2. URL: http://www.consecol.org/vol2/iss1/art2.

Arkes HR, Ayton P, 1999. The sunk cost and Concorde effects: are humans less rational than lower animals? Psych Bull 125: 591-600.

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