Differences in delay, not ratios, control choice in concurrent chains

In two experiments, pigeons were trained with concurrent-chains schedules, wherein responding to equal initial links measured preference between variable-interval terminal links. Absolute terminal-link duration was varied by keeping constant the difference between the terminal-link delays and forcing their ratio to change. Delay-reduction theory scales value relative to a common temporal context and requires that delay differences control choice. Thus, preference should remain invariant. Most competing accounts, including the matching law and a strong form of Weber's law, require that preference vary with the delay ratio. Experiment 1 employed standard concurrent chains, in which terminal-link position and color were confounded. Although average preference remained constant, individual preferences were highly variable and inconsistent, possibly due to carryover of position biases across conditions. In an attempt to reduce variability, Experiment 2 used a modified concurrent-chains procedure. Preference at different terminal-link durations was assessed simultaneously to prevent order effects, and terminal-link position was alternated randomly across trials to minimize the impact of position biases. In Experiment 2, both individual and mean preferences showed the constant-difference invariance. Overall, choice was controlled by terminal-link differences, not ratios.     

Assessment and choice: an operant simulation of foraging in patches

Pigeons were presented with an operant simulation of two prey patches using concurrent random-ratio schedules of reinforcement. An unstable patch offered a higher initial reinforcement probability, which then declined unpredictably to a zero reinforcement probability in each session. A stable patch offered a low but unvarying reinforcement probability. When the reinforcement probability declined to zero in a single step, the birds displayed shorter giving-up times in the unstable patch when the ratio between the initial reinforcement probabilities in the unstable and stable patches was greater and when the combined magnitude of the reinforcement probabilities in the two patches was greater. When the unstable patch declined in two steps, the birds behaved as if their giving-up times were influenced heavily by events encountered during the most recent step of the double-step change. This effect was observed, however, only when the reinforcement probability in that step was .04, not when it was .06. All of these data agree with the predictions of a capture-probability model based on a comparison of the estimated probability of receiving a reinforcer in the current patch with that in alternative patches.     

Delay reduction and optimal foraging: variable-ratio search in a foraging analogue.

The present study investigated conditions under which the conditioned reinforcement principles of delay-reduction theory and views based on simple maximization of reinforcement rate make ordinally opposing predictions with respect to foraging-related choice behavior. The use of variable-ratio schedules in the choice phase also represents an extension of delay-reduction theory to schedules that may better mimic the effort involved in searching. Pigeons responded on modified concurrent-chains schedules in which equal variable-ratio schedules led to unequal variable-interval outcomes and unequal reinforcer amounts. All 4 subjects completed a minimum of two replications of conditions for which the predictions of delay-reduction theory and a simple rate-maximizing theory were opposed. Results were consistent with delay reduction's ordinal predictions in 11 of 11 replications of the divergent predictions favoring the smaller, more immediate alternative. The predictions of rate maximization were upheld only when they were consistent with those of delay reduction. Results are discussed in terms of conditioned reinforcement, sensitivity to reductions in delay to food, and possible rules of thumb that may be useful in characterizing foraging.     

Zebrafish choice behavior is sensitive to reinforcer rate,immediacy, and magnitude ratios

Behavioral flexibility has, in part, been defined by choice behavior changing as a function of changes in reinforcer payoffs. We examined whether the generalized matching law quantitatively described changes in choice behavior in zebrafish when relative reinforcer rates, delays/immediacy, and magnitudes changed between two alternatives across conditions. Choice was sensitive to each of the three reinforcer properties. Sensitivity estimates to changes in relative reinforcer rates were greater when 2 variable-interval schedules were arranged independently between alternatives (Experiment 1a) than when a single schedule pseudorandomly arranged reinforcers between alternatives (Experiment 1b). Sensitivity estimates for changes in relative reinforcer immediacy (Experiment 2) and magnitude (Experiment 3) were similar but lower than estimates for reinforcer rates. These differences in sensitivity estimates are consistent with studies examining other species, suggesting flexibility in zebrafish choice behavior in the face of changes in payoff as described by the generalized matching law.     

Residence time and choice in concurrent foraging schedules

Five pigeons were trained on a concurrent-schedule analogue of the “some patches are empty” procedure. Two concurrently available alternatives were arranged on a single response key and were signaled by red and green keylights. A subject could travel between these alternatives by responding on a second yellow “switching” key. Following a changeover to a patch, there was a probability (p) that a single reinforcer would be available on that alternative for a response after a time determined by the value of λ, a probability of reinforcement per second. The overall scheduling of reinforcers on the two alternatives was arranged nonindependently, and the available alternative was switched after each reinforcer. In Part 1 of the experiment, the probabilities of reinforcement, ρ_red and ρ_green, were equal on the two alternatives, and the arranged arrival rates of reinforcers, λ_red and λ_green, were varied across conditions. In Part 2, the reinforcer arrival times were arranged to be equal, and the reinforcer probabilities were varied across conditions. In Part 3, both parameters were varied. The results replicated those seen in studies that have investigated time allocation in a single patch: Both response and time allocation to an alternative increased with decreasing values of λ and with increasing values of ρ, and residence times were consistently greater than those that would maximize obtained reinforcer rates. Furthermore, both response- and time-allocation ratios undermatched mean reinforcer-arrival time and reinforcer-frequency ratios.     

Vantage perspective in analogue trauma memories: an experimental study

Vantage perspective during recall is thought to affect the emotionality and accessibility of distressing memories. This study aimed to test the effects of vantage perspective during recall on memory associated distress and intrusion development. An adapted version of the trauma film paradigm was used in an experimental design with three conditions. Participants were asked to listen to eyewitness reports of car accidents (e.g. Trauma Analogue Induction) and imagine the scenes vividly using mental imagery. Afterwards, they were asked to recall the most distressing scene from field perspective, observer perspective, or to recall a neutral image from observer perspective (control condition) (e.g. Trauma Analogue Recall). Recall from field perspective resulted in higher negative mood, state-anxiety, and a higher number of short-term intrusions compared to the observer perspective condition and control condition. Negative mood and state-anxiety were mediators in the relationship between vantage perspective and intrusions. In comparison to observer perspective, field perspective increased the amount of short-term intrusions as a result of higher levels of negative mood and state-anxiety after memory retrieval. Future research on the interaction between vantage perspective at recall and negative mood and anxiety effects is warranted.     

Variation in inhibitory control does not influence social rank,foraging efficiency,or risk taking,in red junglefowl females

Animal Cognition - Individual variation in cognition, seen in many taxa, is not well understood, despite its potential evolutionary consequences. Inhibitory control is an aspect of cognition which...     

Locus of control,choice, and satisfaction with an assigned task

P. E. Spector's (1982, Psychological Bulletin, 91, 482–497) hypothesis that under conditions of freedom of choice persons characterized by an internal locus of control will experience greater satisfaction with an undesirable task than persons characterized by an external locus of control is considered in light of distinctions between perceived control and perceived choice. Perceived choice is proposed as an intervening variable implicit in Spector's argument. This reasoning was tested with 46 male and female college student participants in a 2 (Choice) × 2 (Locus of Control) factorial design experiment. An interaction of Choice × Locus of Control on task satisfaction was obtained such that internals tended to be more satisfied when given a choice while externals experienced greater satisfaction when not given a choice. The implications of this result for Spector' hypothesis and for the manner in which undesirable work is assigned are discussed.     

Stimulus control and response bias in an analogue prey-detection procedure

The present study compared the performance of 6 pigeons trained to detect luminance differences in two different signal-detection procedures. Exposed to a three-key array, the pigeons were trained to peck the left key when the brighter of two light intensities had been presented on the center key and to peck the right key when the dimmer of two light intensities had been presented on the center key. Procedure A was a standard signal-detection procedure in which left/bright and right/dim responses produced food reinforcement and left/dim and right/bright responses produced periods of timeout. Procedure B was designed to simulate some of the contingencies operating in a prey-detection situation. Left-key responses produced reinforcement following the brighter center-key stimulus and a period of timeout following the dimmer center-key stimulus. Right-key responses always produced a short period of timeout irrespective of the stimulus. Within each procedure, the duration of timeout arranged for false alarms (left/dim responses) was varied between 3 s and 120 s. Measures of accuracy and response bias were compared between the two procedures. The timeout manipulation produced systematic, but relatively small, changes in these measures when right/dim responses (i.e., correct rejections) produced reinforcement (Procedure A). Arranging timeout for right/dim responses in Procedure B produced greater variability in accuracy and response bias than did arranging reinforcement, but this variability was not related to timeout duration. Overall, discrimination accuracy was considerably higher when right/dim responses produced timeout than when they resulted in reinforcement, and accuracy was accompanied by a large bias toward the response associated with reinforcement. These results are consistent with a recently proposed model of signal detection.     

Relative reinforcer rates and magnitudes do not control concurrent choice independently

One assumption of the matching approach to choice is that different independent variables control choice independently of each other. We tested this assumption for reinforcer rate and magnitude in an extensive parametric experiment. Five pigeons responded for food reinforcement on switching-key concurrent variable-interval variable-interval schedules. Across conditions, the ratios of reinforcer rates and of reinforcer magnitudes on the two alternatives were both manipulated. Control by each independent variable, as measured by generalized-matching sensitivity, changed significantly with the ratio of the other independent variable. Analyses taking the model-comparison approach, which weighs improvement in goodness-of-fit against increasing number of free parameters, were inconclusive. These analyses compared a model assuming constant sensitivity to magnitude across all reinforcer-rate ratios with two alternative models. One of those alternatives allowed sensitivity to magnitude to vary freely across reinforcer-rate ratios, and was less efficient than the common-sensitivity model for all pigeons, according to the Schwarz-Bayes information criterion. The second alternative model constrained sensitivity to magnitude to be equal for pairs of reinforcer-rate ratios that deviated from unity by proportionately equal amounts but in opposite directions. This model was more efficient than the common-magnitude-sensitivity model for 2 of the pigeons, but not for the other 3. An analysis of variance, carried out independently of the generalized-matching analysis, also showed a significant interaction between the effects of reinforcer rate and reinforcer magnitude on choice. On balance, these results suggest that the assumption of independence inherent in the matching approach cannot be maintained. Relative reinforcer rates and magnitudes do not control choice independently.     

Digit ratios,but not facial width-to-height ratios,are associated with the priority placed on attending to faces versus bodies

Testosterone has been linked to a sexual preference for bodies rather than faces. Low digit ratio (2D:4D) and high facial width-to-height ratio (WHR) are associated with high prenatal testosterone. We tested whether low 2D:4D and high facial WHR were correlated with a preference for bodies over faces. Our sample consisted of 109 college students (64 males). A two-way analysis of variance demonstrated a significant main effect of digit ratio on the priority placed on paying attention to faces or bodies such that low 2D:4D was linked to a preference for bodies, but the effect of sex was not significant. There were no significant interaction effects. Another two-way analysis of variance revealed no significant effects for facial WHR and attentional priority but significant associations between sex and attentional priority. There were no significant interaction effects. Our findings indicate that individuals with low digit ratios tended to pay more attention to bodies than to faces compared with individuals with higher digit ratios, independently of sex. We also found that males tended to pay more attention to bodies than to faces compared with females, independently of facial WHR.     

The effects of alcohol consumption and expectancies in an experimental date rape analogue

This study examined the impact of perceived token resistance and the psychological and pharmacological effects of alcohol consumption on men's discrimination of when a female wants her partner to stop his sexual advances. In a 2 (alcohol vs. no alcohol) × 2 (expectancy vs. no expectancy) × 2 (perceived token resistance vs. no resistance) randomized factorial design, male college students were exposed to an audiotape of a date rape. Before listening, participants were told that on the previous date the woman did not resist sexual contact or that she initially objected to the contact but the man was able to obtain the level of intimacy he desired. Relative to participants assigned to the no-alcohol expectancy or no-alcohol consumption group, participants in the alcohol expectancy and alcohol consumption groups took significantly longer to determine that the man should refrain from attempting further sexual contact. The implications of the findings are discussed.This investigation was carried out by the first author under the supervision of the second author in partial fulfillment of the requirements for the degree of Doctor of Philosophy.     

Information, choice, and reactions to stress: a field experiment in a blood bank with laboratory analogue.

Two experiments examined the effects of various operations of personal control on reactions to stress. The first study incorporated two features into the blood drawing procedure at a blood bank: providing donors with accurate information and allowing donors to choose the arm to be used. Measurement of nurses' actions to prevent donors from fainting and self-reports of discomfort revealed that the combination of choice and information was somewhat effective in reducing distress. However, providing either information or choice alone was more effective. In a second laboratory study using a cold pressor stimulus as stressor, subjects given a choice (the option to terminate the aversive stimulus and choice of hand used) showed a reduction of aftereffects on a measure of attention to detail. Subjects given information but not choice also showed this reduction. Combining information and choice was no different from either treatment alone. Taken together, the results of both studies indicate that moderate levels of choice and information are optimal for coping with stress. An explanation was suggested based on a contextually determined relationship amomg choice, information, and perceived control.     

Stress impairs optimal behavior in a water foraging choice task in rats

Stress is a biologically significant social–environmental factor that plays a pervasive role in influencing human and animal behaviors. While stress effects on various types of memory are well characterized, its effects on other cognitive functions are relatively unknown. Here, we investigated the effects of acute, uncontrollable stress on subsequent decision-making performance in rats, using a computer vision-based water foraging choice task. Experiencing stress significantly impaired the animals'' ability to progressively bias (but not maintain) their responses toward the larger reward when transitioning from equal to unequal reward quantities. Temporary inactivation of the amygdala during stress, however, blocked impairing effects on decision making.It is now well documented that exposure to uncontrollable stress can produce alterations in multiple brain–memory systems in humans and animals (McEwen and Sapolsky 1995; Kim and Diamond 2002; Joels et al. 2006; Shors 2006; Luethi et al. 2008). In humans, impairments in long-term, but not short-term, verbal recall tasks have been observed in people with post-traumatic stress disorder (PTSD) (Bremner et al. 1995) or Cushing''s disease (a hypercortisolemia condition) (Starkman et al. 1992) and in healthy individuals subjected to stress (Lupien et al. 1997) or exposed to stress levels of cortisol (Newcomer et al. 1994). In rodents, stress and corticosterone administration interfere with spatial and working memory (Diamond and Rose 1994; de Quervain et al. 1998; Kim et al. 2001) and potentiate aversive conditioning (Shors et al. 1992; Maier et al. 1995). Further, a number of stress-associated neurobiological changes have been identified (e.g., in hippocampus, medial prefrontal cortex, and amygdala) subserving different memory functions (Arnsten and Goldman-Rakic 1998; Kim and Yoon 1998; Vyas et al. 2003; Holmes and Wellman 2009).Although stress effects on memory have been well studied, far less is known about whether (and in what manner) stress influences other higher cognitive functions. The present study investigated the effects of acute, uncontrollable stress (60-min restraint + 60 intermittent tailshocks) on subsequent decision-making performance in rats. The stress procedure, in which animals learn that they can neither escape nor predict an aversive experience, was adapted from earlier studies (e.g., Maier and Seligman 1976; Kim et al. 1996). Decision making was assessed using an automated Figure-8-shaped maze on which rats were motivated to forage for water rewards in two different locations under equal and unequal quantity conditions (Fig. 1). In addition to behavioral stress, we examined the effects of corticosterone administration and inactivation of the amygdala during stress on decision making. Both corticosterone (a glucocorticoid hormone released in response to stress) and the amygdala (a structure crucial in defensive behavior) have been implicated in mediating neurocognitive effects of stress (McEwen and Sapolsky 1995; Kim and Diamond 2002).Open in a separate windowFigure 1.Decision-making task. Thirst-motivated rats were trained to forage for water on an automated Figure-8-shaped maze. A computer algorithm controlled the raising and lowering of four gates (represented by rectangles), the delivery of water (blue circles), and tracking of the animal''s location on the maze. During the baseline days, both left and right sides of the maze presented 0.04 mL of water at 80% probability (equal reward trials). Following each trial (a left or right loop), the animal returned to the center bridge to start the next trial (40 trials per day). During the bias test days, one side of the maze (counterbalanced) offered 0.12 mL of water at 80% probability, while the other side continued to present 0.04 mL of water at 80% probability (unequal reward trials). Animals received either stress, CORT injections, or neither (for group details, see text).Experimentally naïve male Charles River Sprague–Dawley rats (initially weighing 275–300 gm) were singly housed and maintained on a reverse 12-h light-dark cycle (lights on at 19:00 h). After 7 d of acclimation and for the duration of the experiment, daily water access was restricted to maintain approximately 85% of the animal''s normal body weight. Food was available ad libitum throughout the experiment. All experiments were conducted during the dark phase of the cycle and in strict compliance with the University of Washington Institutional Animal Care and Use Committee guidelines. Under anesthesia (30 mg/kg ketamine and 2.5 mg/kg xylazine, i.p.) amygdala (AMYG) animals were chronically implanted with 26-gauge guide cannulae (Plastics One) bilaterally into the amygdala (from bregma: anteroposterior, −2.3 mm; mediolateral, ±5 mm; dorsoventral, −7.7 to 8.0 mm). During 10–15 d of postoperative recovery, each dummy cannula was removed and replaced with a clean one.Muscimol free base (Sigma-Aldrich), dissolved in artificial cerebrospinal fluid (10 mM at pH ∼7.4), was microinfused into the amygdala (bilaterally) via 33-gauge infusion cannulae that protruded 1 mm beyond the guide cannulae (cf. Kim et al. 2005). An infusion volume of 0.3 μL (per side) was delivered using a Harvard PHD2000 syringe pump (Harvard Apparatus) over the course of 3 min. Animals were returned to their home cages for 30 min before undergoing the stress procedure. We based the timing of inactivation on previous findings that pre-stress but not immediate post-stress inactivation of the amygdala interferes with stress effects on hippocampal long-term potentiation (LTP) and spatial memory (Kim et al. 2005). Based on studies that examined ³H-muscimol spreading (Krupa et al. 1993; Arikan et al. 2002) in the cerebellum, in which 1 μL diffused a radius of 1.6–2.0 mm, it was estimated that 0.3 μL of muscimol would spread within a radius of approximately 0.5–0.7 mm from the infusion needle tip. Hence, it is likely that infused muscimol would have diffused to the central, lateral, and basal nuclei of the amygdala and possibly to portions of adjacent neighboring structures. The BODIPY TMR-X muscimol conjugate (Invitrogen) was infused in the same manner as muscimol free base (cf. Allen et al. 2008) to image the spread of reversible amygdalar inactivation.Corticosterone (CORT) animals received three daily subcutaneous injections of 3 mg/kg corticosterone (suspended in sesame oil; Sigma-Aldrich) 30 min prior to bias testing.Rats undergoing stress were restrained in Plexiglas tubes and presented with 60 tailshocks (1-mA intensity, 1-sec duration, 5- to 115-sec variable intershock interval) for 60 min (Kim et al. 2005). Animals were divided into four groups: control, stress, amygdalar inactivation plus stress (AMYG), and daily corticosterone (CORT).Following 2 d of habituation (to transport, maze, and room ambiance), all animals underwent successive shaping and testing phases. The dimensions and automatic features of the Figure-8 maze have been described previously (Pedigo et al. 2006; Yoon et al. 2008); for details, see Supplemental material. During shaping, each rat is placed into the center runway with all four gates in the up position (Supplemental Fig. S1). After 3 sec, the front and one of the side gates drop, until the rat on its own volition moves out of the center and onto the open side runway. At this point, the lowered front and side gates rise (to prevent the rat from going backward), water is delivered to both the open arm and the center spout, and the back gate drops. The rat consumes water from the open arm, and a new trial begins when he returns to and consumes water from the center arm. There was a 3-sec delay between each trial. During shaping, left and right choices were thus forced choices and were presented in a pseudorandom pattern, such that there was an equal number of both in a complete session (40 laps). Rats underwent shaping once daily until they met predetermined criteria: completion of 40 laps in less than 30 min, and less than five back edge errors (i.e., after making a choice, running up the opposite arm instead of going back to the center arm). The automated program controlled the gates and water delivery, according to the rat''s position on the maze.During baseline testing (Supplemental Fig. S2), the rewards dispensed on left and right arms were equal in both volume (0.04 mL) and probability (0.8). Each rat remained on baseline testing until he demonstrated a stable left/right choice pattern across three consecutive days. If a slight preference to one side was present, the opposite arm would be the increased reward side when bias testing commenced.Stress and AMYG rats were exposed to restraint + tailshock stress 1 d before their first bias test, and CORT rats were given corticosterone injections 30 min prior to each bias test. During bias testing (Supplemental Fig. S3), the reward value on one side was tripled in volume (0.12 mL) while the value for the other side remained constant (0.04 mL). Control, AMYG, and CORT animals were given three consecutive days of bias tests, while stress animals underwent six consecutive days of bias tests.At the completion of behavioral testing, animals were overdosed with urethane and perfused intracardially with 0.9% saline followed by 10% buffered formalin. The brains were removed and stored in 10% formalin overnight and then kept in 30% sucrose solution until they sank. Transverse sections (50 μm) were taken through the extent of the cannulae tract, mounted on gelatin-coated slides, and stained with cresyl violet to verify cannulae placements.The visit number to the left (or right) side of the maze for baseline and bias days were normalized to the mean left (or right) visits across the three baseline days. Statistical comparisons between groups were examined using ANOVA. For a significant difference (P < 0.05), post-hoc comparisons were performed using Tukey''s honestly significant difference test.Thirst-motivated rats readily learned to forage for water on the maze, and when left and right sides of the maze provided the same quantity (0.04 mL) and probability (80%) of water, the animals made comparable numbers of left and right visits (during 40 laps daily) that were stable across three baseline days (Fig. 2A). The 80% probability (a partial reinforcement schedule) was used so that animals frequently explored both sides of the maze. After animals demonstrated stable baseline choices, the volume of water on one side was tripled (0.12 mL at 80%), while the other side remained constant (0.04 mL at 80%). Overall, bias performance (choice of the larger reward) increased across the first three bias test days (repeated-measures ANOVA; main effect of day, F_(2,54) = 78.16, P < 0.001). However, the four groups differentially increased their bias across days (group × day interaction, F_(6,54) = 4.14, P < 0.01). Specifically, stressed rats displayed a significantly slower rate of bias toward the larger reward than did the controls (P < 0.01, Tukey). The stressed rats did ultimately develop a bias compared with their baseline choices (F_(6,62) = 8.44, P < 0.001). This bias was first significant on the fourth day (P < 0.05, Dunnett t-test). However, even after 6 d of bias testing, their bias (127 ± 5.7%, mean ± SEM) did not reach the level of controls'' third day bias (182 ± 12.2%). Unlike the behavioral stress group, however, animals that received three daily corticosterone injections (3 mg/kg, subcutaneously) prior to testing chose the larger reward side more frequently (173 ± 6.7%, bias day 3) and did not differ from the control group (P > 0.7, Tukey). When the amygdalae were inactivated during stress (Fig. 3), these animals behaved like controls (P > 0.7, Tukey) and increased their visit frequency toward the larger reward side of the maze (174 ± 13.0%, bias day 3) (Fig. 2A). Although control, CORT, and AMYG animals developed strong bias responses, they did not exclusively visit the larger reward side of the maze because on 20% of trials they did not receive a reward.Open in a separate windowFigure 2.Stress effects on decision making. (A) All groups of animals showed comparable visits to left and right sides of the maze during the three baseline days. When transitioning from equal to unequal reward trials, stressed rats (n = 7) displayed an impaired ability to bias their responses toward the larger reward side compared with control (n = 10), AMYG (n = 7), and CORT (n = 7) rats (P = 0.002, group × bias day interaction). (B) Example visit maps of a control rat during baseline and bias days (40 laps each).Open in a separate windowFigure 3.(Top) Histological reconstruction of injection cannulae placement tips in the amygdala. (Bottom) A photomicrograph of fluorophore-conjugated muscimol (0.3 μL over 3 min) spread in the amygdala. The red fluorescence is overlaid with a dark field image.We then examined whether stress might have produced alterations in motor, motivation, and reference memory performances that hindered the animals'' ability to bias their responses toward the larger reward. The latency to complete 40 laps of the first bias test session (Supplemental Fig. S4A) showed a trend of stress animals completing the bias test faster than the other three groups, but this group × day interaction was not significant (F_(6,54) = 1.89, P > 0.05). Stress also did not impair the animals'' reference memory of the maze (Supplemental Fig. S4B). That is, after making a left or right visit, stressed animals readily re-entered the center runway to start the next trial (one-way ANOVA; average baseline and first three bias days, F_(3,41) = 0.20, P > 0.8), whereas control, CORT, and AMYG animals displayed an increased propensity to investigate the other side before re-entering the center, particularly as bias testing progressed (repeated-measures ANOVA; main effect of day, F_(2,54) = 4.84, P < 0.05).Our results indicate that rats clearly demonstrate the capacity to change their foraging behavior to acquire a larger water reward when transitioning from equal to unequal quantities, and that such behavioral flexibility is vulnerable to acute, uncontrollable stress. Specifically, rats that experienced 1 h of restraint stress + 60 intermittent tailshocks were significantly impaired in biasing their responses toward the side of the maze with a larger quantity of water. This effect on bias was not due to any lingering post-stress motivational or motor effects, as stress did not increase the latency to complete the bias test. Daily corticosterone injections did not interfere with this task, indicating that corticosterone elevation per se cannot reproduce behavioral stress effects on behavioral flexibility. However, similar to previous stress–memory studies (Kim et al. 2001; Waddell et al. 2008), amygdalar inactivation during stress effectively blocked this effect. This suggests that the amygdala plays a crucial role in mediating stress effects across different cognitive domains.Although stress altered the rats'' behavior in our choice-based task, it remains unclear precisely which neural and cognitive systems were affected. For instance, the impairment of behavioral flexibility might be an indirect consequence of stress effects on hippocampal memory functioning. The stress paradigm used here is known to impede LTP in the CA1 hippocampus and hinder spatial memory (Foy et al. 1987; Kim et al. 2001). However, corticosterone, which also impedes LTP (McEwen and Sapolsky 1995) and spatial memory when administered 30 min before testing (de Quervain et al. 1998), did not impair behavioral flexibility. The impairment to choose the larger reward may be due to stress effects on working memory, such that the rats cannot remember (and thus learn) that one reward is larger. However, if true, the stress-induced memory impairment is unusually persistent in our task: Rats were affected through at least 6 d beyond the stress exposure. Because acquisition and retrieval of information are integral components of decision making, the contribution of stress-associated changes in learning and memory cannot be fully excluded. Another possibility is that stressed rats are more likely to use habitual rather than flexible strategies, even when a change in behavior may be optimal. Consistent with this explanation are recent findings that chronic stress exposure increases habit-based responding, with corresponding atrophy and hypertrophy of goal-directed and habit-based neural circuitry, respectively (Dias-Ferreira et al. 2009). The reliance on habit memory is also increased following anxiogenic drug infusions into the amygdala (Elliott and Packard 2008), which further implicates amygdalar modulatory activity during and after stress exposure in decreasing flexible behavior. If the stress experience did increase perseverative choice behavior, this may partially explain previously observed associations between distress and perseveration in humans (Robinson et al. 2006). Alternately, stress may have disrupted the reward circuitry and impaired the ability to discriminate between the two reward values from the two side arms, in which case stress effects on a dopamine-related reward circuit (Schultz et al. 1997) should be explored. However, this cannot be the sole explanation because post-bias stress did not affect the animals'' bias behavior toward the larger reward (Supplemental Table S1), and nonstressed rats resume equal arm visits when rewards are returned to baseline values (data not shown).The present findings reveal that a single exposure to an acutely stressful experience is enough to affect an animal''s behavior on a simple forging task for several days. There is accumulating evidence that exposure to stress increases amygdala and decreases prefrontal activity in both humans and animals (for review, see Arnsten 2009) and that even a single stress exposure alters cellular morphology in prefrontal cortex (Izquierdo et al. 2006). This shift in neural activity may promote the use of one cognitive strategy over another (e.g., habitual versus flexible). To address this, future studies need to investigate brain structures implicated in decision making, including the prefrontal and the parietal cortices (Gold and Shadlen 2007; Lee 2008), for their susceptibility to stress. Regardless, the present findings, to our knowledge, provide the first direct evidence that acute uncontrollable stress persistently impairs decision-making performance in animals and that this effect is dependent upon amygdalar activity during stress.     

A temporal limit on the effect of future food on current performance in an analogue of foraging and welfare

Rats obtained access to food twice each 24-hour period. The first session was a work session in which food was available on a progressive-ratio schedule. During the second session, which occurred between 1 and 23 hours after the work session, food was freely available up to a fixed total intake each 24 hours. The situation resembled elements of several real world circumstances, including the choice between continuing to forage in a rapidly depleting patch and waiting for a better patch, and between working now and receiving a guaranteed income later. The purpose of the experiment was to explore the time period over which future access to reward could affect current responding. Contrary to what might be expected from recent theorizing, anticipation of future food delayed by an hour or more after the start of the work session had no effect on current performance. Food intake was high and constant during work sessions except for a prefeeding effect that occurred when the free session closely preceded the next day's work session. Also, an increase in the difficulty of the work schedule increased the amount of work and the maximum price paid for food as if the work session were the only time food was available. The results indicate the importance of considering temporal limits in theories that require animals to integrate input over time to determine the allocation of resources among alternatives.     

An experimental analysis of optimal foraging behaviour in patchy environments

A series of experiments was designed to explore the cognitive mechanisms involved in optimal foraging models by using the behavioural controls of operant methodology. Rats were trained to press one of two levers to obtain reinforcement on a progressive variable-interval schedule, which modelled food patch depletion; the schedule was reset by pressing the other lever. Thus both duration (residence time in a patch) and rate-related (interval before and after the final reward) measures were obtained. Experiment 1, which manipulated environmental stability and quality, and Experiment 2, which varied travel time between patches, found results that supported the marginal value theorem (Charnov, 1976) and suggested that rats adjust capture rate to the environment average by monitoring the length of the interval between rewards. Experiment 3 modelled the clumping of food items and found that capture rate was now adjusted by adoption of a fixed giving-up time. Finally, Experiments 4a and 4b ruled out a time expectancy hypothesis by manipulating the number of food clumps and the series of inter-reinforcement intervals. Overall the experiments demonstrate the value of modelling foraging strategies in operant apparatus, and suggest that rats adopt rate predictive strategies when deciding to switch patches.