Stepping outside the box in considering the C/T ratio

Rate expectancy theory (RET) predicts that in Pavlovian procedures conditioned responding will be directly related to the ratio of time spent in the experimental context (C) relative to the trial time (T) or duration of the conditioned stimulus (CS). This prediction was discussed in the context of three experiments. The first and third experiments involved sexual conditioning in quail [Learn Motiv 31 (2000) 211; J Exp Psychol Anim Behav Processes 27 (2001) 269]; the second experiment involved conditioning rats with food as the unconditioned stimulus (US) [Learn Motiv 28(1997) 465]. In each experiment, one type of conditioned response was directly related to the C/T ratio (as predicted by RET) but another conditioned response was inversely related. In addition, the conditioned behavior that occurred with low C/T ratios was controlled by contextual cues rather than the putative CS. The implications of these findings for possible boundary conditions of RET were discussed. The discussion revealed serious shortcomings in the characterization of context conditioning by RET.     

Modeling the effects of enclosure size on geometry learning

Several recent studies have shown that chickens, fish, and humans trained to find a reward in a corner of a rectangular enclosure with distinctive features rely more on the geometry of the enclosure in small enclosures and rely more on the features in large enclosures. Here, these results are modeled using a recent associative model of geometry learning [Miller, N.Y., Shettleworth, S.J., 2007. Learning about environmental geometry: an associative model. J. Exp. Psychol. Anim. B 33, 191–212]. By adjusting the salience of either geometric or featural information or both the model is capable of reproducing much of the data on the effects of enclosure size on geometry learning.     

Rats have trouble associating all three parts of the time-place-event memory code

The ability of animals to associate an event with predictable time and place information confers a major biological advantage. The current research uses a variety of procedures and paradigms (e.g. place preference, radial arm maze, Morris water maze, T-maze, go no-go) to show that rats, unlike pigeons [e.g. Anim Learn Behav 22 (1994) 143] do not readily make an event-time-place association. They do make associations between event-time and event-place information, however. These findings are in disagreement with Gallistel's (The Organization of Learning, MIT Press, Cambridge, MA ) theory that claims that animals automatically store a memory code that has these three pieces of information. The present research is in line with the work of others who also find that rats do not readily make daily time-place associations [Behav Processes 23 (1997) 232; Behav Processes 52 (2000) 11; Behav Processes 49 (2000) 21; Anim Learn Behav 28 (2000) 298]. An interesting finding that did emerge from the present research was that at least some rats can use a circadian timer to solve a time-of-day discrimination if the task is a go no-go discrimination.     

Effects of post-treatment retention interval and context on neophobia and conditioned taste aversion

We have repeatedly observed that a delay between acquisition and test, and the nature of the context in which the delay is spent, modulates latent inhibition (LI) of conditioned taste aversion (CTA; e.g. [Anim. Learn. Behav. 28 (2000) 389; Anim. Learn. Behav. 30 (2002) 112]). The present paper analysed the effects of delayed testing and treatment context after flavor exposure on the recovery of neophobia (Experiment 1) and on extinction after simple conditioning (Experiment 2). Two experiments were conducted with the same factorial design (2x2: 1 day versus 21 days of delay between first and second stage, and home versus experimental cages as place of experimental treatment). There were independent effects of both variables on habituation of neophobia and conditioning strength as measured on extinction trials. The long delay produced a reduction of neophobia (Experiment 1) and an increase in conditioning (Experiment 2). In addition, more of the flavored solution was consumed when the experimental treatment was conducted in the home cage than in the experimental cage (Experiment 1), and there was stronger conditioning when the delay period took place in the experimental cages than in the home cages (Experiment 2). The implications of these results for LI, as well as their relevance for experiments that use the CTA paradigm, are discussed.     

Effect of stimulus orderability and reinforcement history on transitive responding in pigeons

Transitive responding in humans and non-human animals has attracted considerable attention because of its presumably inferential nature. In an attempt to replicate our earlier study with crows [Lazareva, O.F., Smirnova, A.A., Bagozkaja, M.S., Zorina, Z.A., Rayevsky, V.V., Wasserman, E.A., 2004. Transitive responding in hooded crows requires linearly ordered stimuli. J. Exp. Anal. Behav. 82, 1-19], we trained pigeons to discriminate overlapping pairs of colored squares (A+ B-, B+ C-, C+ D-, and D+ E-). For some birds, the colored squares, or primary stimuli, were followed by a circle of the same color (feedback stimuli) whose diameter decreased from A to E (Ordered Feedback group); these circles were made available to help order the stimuli along a physical dimension. For other birds, all of the feedback stimuli had the same diameter (Constant Feedback group). In later testing, novel choice pairs were presented, including the critical BD pair. The pigeons' reinforcement history with Stimuli B and D was controlled, so that the birds should not have chosen Stimulus B during the BD test. Unlike the crows, the pigeons selected Stimulus B over Stimulus D in both ordered and Constant Feedback groups, suggesting that the orderability of the post-choice feedback stimuli did not affect pigeons' transitive responding. Post hoc simulations showed that associative models [Wynne, C.D.L., 1995. Reinforcement accounts for transitive inference (TI) performance. Anim. Learn. Behav. 23, 207-217; Siemann, M., Delius, J.D., 1998. Algebraic learning and neural network models for transitive and non-transitive responding. Eur. J. Cogn. Psychol. 10, 307-334] failed to predict pigeons' responding in the BD test.     

Stimulus representation in SOP: I. Theoretical rationalization and some implications

THE SOP MODEL [INFORMATION PROCESSING IN ANIMALS: Memory Mechanisms, Erlbaum, Hillsdale, NJ, 1981, p. 5] is described in terms of its assumed stimulus representation, network characteristics, and rules for learning and performance. It is shown how several Pavlovian conditioning phenomena can be accounted on the basis of the model's presumed stimulus representation. Challenges to the SOP model prompted the adoption of a componential stimulus representation in: AESOP [Contemporary Learning Theories: Pavlovian Conditioning and the Status of Traditional Learning Theory, Erlbaum, Hillsdale, NJ, 1989, p. 149], this was a dual representation of the unconditioned stimulus (US), and C-SOP [Contemporary Learning: Theory and Application, Erlbaum, Mahwah, NJ, 2001, p. 23], this was a multi-component representation of the conditioned stimulus (CS). The assumption of a componential CS representation, where large numbers of elements can be separately learned about, necessitated a modification of the learning rule. The modified, "constrained" rule was found useful to explain timing characteristics of Pavlovian conditioned responses, as well as data offered by Rescorla [J. Exp. Psychol. Anim. Behav. Process. 26 (2000) 428; Q. J. Exp. Psychol. 54B (2001) 53; J. Exp. Psychol. Anim. Behav. Process. 28 (2002) 163] showing that stimuli trained in compound do not share the same quantitative fate.     

Matching-to-sample by pigeons: the dissociation of comparison choice frequency from the probability of reinforcement

It has been proposed that comparison choice in matching-to-sample should depend on two factors, the relative probability of reinforcement associated with each of the comparison stimuli and the conditional probability of each comparison stimulus being correct given presentation of one of the samples. DiGian and Zentall [DiGian, K.A., Zentall, T.R., 2007. Matching-to-sample in pigeons: in the absence of sample memory, sample frequency is a better predictor of comparison choice than the probability of reinforcement for comparison choice. Learn Behav. 35, 242-261] have shown that sample frequency together with the probability of choosing each of the comparison stimuli in training can influence comparison choice when delays are introduced, when the number of reinforcements associated with each of the comparison stimuli is equated. Furthermore, Zentall and Clement [Zentall, T.R., Clement, T.S., 2002. Memory mechanisms in pigeons: Evidence of base-rate neglect. J. Exp. Psych.: Anim. Behav. Proc. 28, 111-115] have found that sample frequency can affect comparison choice when delays are introduced independently of the number of choices of each of the comparison stimuli in training and the number of reinforcements associated with each of the comparison stimuli is equated. In the present experiment we found that the probability of choosing each of the comparison stimuli in training can affect comparison choice when delays are introduced, independently of sample frequency and when the number of reinforcements associated with each of the comparison stimuli is equated. Together, these experiments suggest that when the sample is not available, there is a partial dissociation between comparison choice and the probability of reinforcement associated with each of the comparison stimuli.     

Prospective timing, attention and the switch. A response to 'Gating or switching? gating is a better model of prospective timing' by Zakay

In response to the 'Switching or gating' paper (Lejeune, H., 1998. Switching or gating? The attentional challenge in cognitive models of psychological time. Behav. Proc. 44, 127-145), Zakay argued that attention allocation to time should reflect attentional processes in general and suggested that the attentional gate model (AGM) has more explanatory power than the temporal information processing model (TIP) of Church (Church, R.M., 1984. Properties of the internal clock. In: Gibbon, J., Allan, L., (Eds.), Timing and Time Perception, vol. 423. Annals of the New York Academy of Sciences, New York, pp. 566-582). The first point might not be challenged, provided that the specificity of the temporal stimulus is taken into account. Concerning the second point, we argue that the TIP model can account for human prospective timing and discuss differences between attention versus expectancy or motivation. We prefer a 'satellite' attention allocation process, targeting the switch and reference memory (Meck, W.H., 1983. Selective adjustment of the speed of the internal clock and memory processes. J. Exp. Psychol.: Anim. Behav. Proc. 9, 171-201) to an attentional gate serially included in the TIP model of Church.     

Pitch detection of dynamic iterated rippled noise by humans and a modified auditory model

Iterated ripple noise (IRN) is a broadband noise with temporal regularities, which can give rise to a perceptible pitch. Since the perceptual pitch to noise ratio of these stimuli can be altered without substantially altering their spectral content, they have been useful in exploring the role of temporal processing in pitch perception [Yost, W.A., 1996. Pitch strength of iterated rippled noise, J. Acoust. Soc. Am. 100 (5), 3329-3335; Patterson, R.D., Handel, S.,Yost, W.A., Datta, A.J., 1996. The relative strength of the tone and noise components in iterated rippled noise, J. Acoust. Soc. Am. 100 (5), 3286-3294]. A generalised IRN algorithm is presented, in which multiple time varying temporal correlations can be defined. The resulting time varying pitches are perceptually very salient. It is also possible to segregate and track multiple simultaneous time varying pitches in these stimuli. Temporal auditory models have previously been shown to account for the perception of IRNs with static delays [Patterson, R.D., Handel, S.,Yost, W.A., Datta, A.J., 1996. The relative strength of the tone and noise components in iterated rippled noise, J. Acoust. Soc. Am. 100 (5), 3286-3294]. Here we show that some simple modifications to one such model [Meddis R., Hewitt, M.J., 1991. Virtual pitch and phase sensitivity of a computer model of the auditory periphery I. Pitch identification, J. Acoust. Soc. Am. 89, 2866-2882] allow it to track moving correlations, and also improve its performance in response to static correlations.     

Negative anticipatory contrast: does it involve anticipation of an impending reward?

Negative anticipatory contrast (NAC) corresponds to the suppression in consumption of a first rewarding substance (e.g., saccharin 0.15%) when it is followed daily by a second preferred substance (e.g., sucrose 32%). The NAC has been interpreted as resulting from anticipation of the impending preferred reward and its comparison with the currently available first reward [Flaherty, C.F., Rowan, G.A., 1985. Anticipatory contrast: within-subjects analysis. Anim. Learn. Behav. 13, 2-5]. In this context, one should expect that devaluation of the preferred substance after the establishment of the NAC would either reduce or abolish the contrast effect. However, contrary to this prediction, the results of the present study show that the NAC is insensitive to devaluation of the second, preferred, substance. This allows one to question that interpretation. The results reported in this study support the view that the NAC effect is controlled by memory of the relative value of the first solution, which is updated daily by means of both a gustatory and/or post-ingestive comparison of the first and second solutions, and memory of past pairings.     

Time left in the mouse

Evidence suggests that the online combination of non-verbal magnitudes (durations, numerosities) is central to learning in both human and non-human animals [Gallistel, C.R., 1990. The Organization of Learning. MIT Press, Cambridge, MA]. The molecular basis of these computations, however, is an open question at this point. The current study provides the first direct test of temporal subtraction in a species in which the genetic code is available. In two experiments, mice were run in an adaptation of Gibbon and Church's [Gibbon, J., Church, R.M., 1981. Time left: linear versus logarithmic subjective time. J. Exp. Anal. Behav. 7, 87-107] time left paradigm in order to characterize typical responding in this task. Both experiments suggest that mice engaged in online subtraction of temporal values, although the generalization of a learned response rule to novel stimulus values resulted in slightly less systematic responding. Potential explanations for this pattern of results are discussed.     

Scaling relative incentive value: different adjustments to incentive downshift in pigeons and rats

Previous research suggests that pigeons and rats show differences in their behavioral adjustments in spaced-trial, incentive-downshift situations. Also, Papini and Pellegrini [Papini, M.R., Pellegrini, S., 2006. Scaling relative incentive value in consummatory behavior. Learn. Motiv. 37, 357-378] and Pellegrini and Papini [Pellegrini, S., Papini, M.R., 2007. Scaling relative incentive value in anticipatory behavior. Learn. Motiv. 38, 128-154] showed that changes in the rat's lever-pressing performance, runway running, and consumption of sucrose solutions after downshifts in incentive magnitude were a function of the ratio of postshift/preshift incentive magnitudes. Here, two experiments using a Pavlovian autoshaping procedure studied the adjustment of pigeons and rats to changes in incentive magnitude. In Experiment 1, pigeons received light-food pairings, whereas in Experiment 2, rats received lever-sucrose pairings. As a result, key-pecking and lever-pressing developed in each experiment, respectively. Preshift incentive magnitudes were downshifted so as to obtain postshift/preshift ratios of 0.125 and 0.25. Pigeons responded during the postshift phase according to the preshift incentive value and independently of the ratio value. However, rats showed ratio constancy, responding during the postshift in accordance with the postshift/preshift ratio, rather than with the absolute magnitudes of either the preshift or postshift incentives. These results support the comparative hypothesis that the mechanisms underlying ratio constancy during incentive downshifts are unique to mammals.     

A fuzzy-neural system for identification of species-specific alarm calls of Gunnison's prairie dogs

In this study we describe the design and application of an automated classification system that utilizes artificial intelligence to corroborate the finding that Gunnison's prairie dogs have different alarm calls for different species of predators. This corroboration is strong because it utilizes an entirely different analysis technique than that used in the original research by Slobodchikoff et al. [Slobodchikoff, C.N., Fischer, C., Shapiro, J., 1986. Predator-specific alarm calls of prairie dogs. Am. Zool. 26, 557] or in subsequent study done by Slobodchikoff et al. [Slobodchikoff, C.N., Kiriazis, J., Fischer, C., Creef, E., 1991. Semantic information distinguishing individual predators in the alarm calls of Gunnison's prairie dogs. Anim. Behav. 42, 713-719]. The study described here also is more completely automated than earlier study in this area. This automation allowed a large volume of field data to be processed where all measurements of relevant parameters were performed through software control. Previous study processed a smaller data set and utilized manual measurement techniques. The new classification system, which combines fuzzy logic and an artificial neural network, classified alarm calls correctly according to the eliciting predator species, achieving accuracy levels ranging from 78.6 to 96.3% on raw field data digitized with low quality audio equipment.     

Dissociation between the Espinet and perceptual learning effects in flavour aversion conditioning

In three experiments rats were given pre-exposure to two compound flavours, AX and BX, the two compounds being presented for some subjects on alternate trials (the intermixed schedule) and, for others, in separate blocks of trials (the blocked schedule). After aversion conditioning with A (in Experiments 1 and 2), the inhibitory properties of B were tested using both retardation (Experiment 1) and summation tests (Experiment 2). The results failed to support the proposal [Anim. Learn. Behav. 23 (1995) 361] that B should acquire inhibitory properties in the intermixed condition (the "Espinet effect"). Experiment 3 demonstrated that generalisation to BX after conditioning with AX was attenuated by intermixed pre-exposure (a perceptual learning effect). This pattern of results challenges the hypothesis that inhibitory learning during intermixed pre-exposure to AX and BX can account for both the Espinet and the perceptual learning effects.     

Differences between simultaneous and blocked training detected by a transfer test

Secondary data analysis was used to compare responding early on a transfer test from rats previously trained simultaneously or successively on multiple temporal discriminations for the same number of trials [Guilhardi, P., Church, R.M., 2005 a. Dynamics of temporal discrimination. Learn. Behav., 33, 399-416]. Three fixed intervals (30, 60, and 120 s) were signaled by three stimuli (light, noise, and clicker). Twelve rats were trained with the three stimulus-interval pairs intermixed on each experimental session (simultaneous condition); 12 other rats were trained in successive blocks of 10 sessions on each pair (blocked condition). Then, all rats had a transfer test in which all three stimulus-interval pairs were presented intermixed on each session. Rats in the simultaneous and blocked condition responded similarly during training, but differently during early stages of the transfer test. One possibility is that rats in the blocked condition were controlled by the previous interval, not by the current stimulus. These results challenge the usual assumptions from models of timing and conditioning that both simultaneous and blocked training produce learning of the associations between stimulus and interval in a multiple interval training task.     

Comparing the generalized matching law and contingency discriminability model as accounts of concurrent schedule performance using residual meta-analysis

To compare the generalized matching law (Baum, W.M., 1974b. On two types of deviation from the matching law: bias and undermatching. J. Exp. Anal. Behav. 22, 231-242) and contingency discriminability model (Davison, M., Jenkins, P.E., 1985. Stimulus discriminability, contingency discriminability, and schedule performance. Anim. Learn. Behav. 13, 77-84) as accounts of concurrent schedule performance, we conducted a residual meta-analysis of response- and time-allocation data from 20 studies (n's=886 and 774, respectively). Both models were fitted to the individual-subject data from each study, and residuals were obtained. Polynomial regressions were then performed on the pooled residuals to determine whether systematic trends were present as a function of predicted values. For the contingency discriminability model, the cubic coefficients were positive and statistically significant for both response- and time-allocation data. By contrast, no statistically significant systematic trend was obtained in the residuals for the generalized matching law. These results suggest that the relationship between log response allocation and log reinforcer allocation does not deviate significantly from linearity over an approximate range of +1.25 to -1.25 log units, consistent with the generalized matching law. Although qualitative criteria are also important in comparing models of behavioral phenomena, residual meta-analysis provides a powerful quantitative methodology for model selection and should prove useful in future research.     

Phosphorylation in sealed rod outer segments: effects of cyclic nucleotides

Rod outer segments (ROS) from rat were purified on Percoll gradients. These ROS had intact plasma membranes since they were impermeable to small molecules. Protein phosphorylation in the purified ROS was studied after the plasma membrane was disrupted by freeze/thawing. [gamma-32P]ATP was used as phosphate donor. ATP concentration, time, temperature, and light or dark adaptation were varied in the assays. The 32P-labeled proteins were separated by polyacrylamide gel electrophoresis and autoradiographed. Rhodopsin was the dominant phosphorylated protein, and the addition of adenosine cyclic 3',5'-phosphate (cAMP) or guanosine cyclic 3',5'-phosphate (cGMP) (10(-4) M) did not qualitatively alter the ROS phosphorylation pattern. The only cyclic nucleotide effect we could establish in these experiments was the inhibition of rhodopsin phosphorylation by cGMP. This inhibition did not appear to be competitive with ATP since cAMP was much less inhibitory than cGMP and the phosphorylation in the presence of cGMP reached a plateau at a much lower level than in control conditions. Hypotheses implying an involvement of protein phosphorylation/dephosphorylation in dark adaptation have been formulated [Miller, J. A., & Paulsen, R. (1975) J. Biol. Chem. 250, 4427-4432; Kuhn, H., McDowell, J. H., Leser, K. H., & Bader, S. (1977) Biophys. Struct. Mech. 3, 175-180]; we suggest that cGMP may control this process through the modulation of the extent of inhibition of phosphorylation of the visual pigment.     

Effect of amount of context extinction on revaluation of a target CS

In two experiments using rats in a conditioned lick suppression paradigm the effects of posttraining extinction of the conditioning context on responding to the target conditioned stimulus (CS) which had previously been paired with a footshock was examined in a neutral context. Experiment 1 replicated Marlin's [Learn. Motiv. 13 (1981) 526] observation that this treatment can reduce responding to the target CS. This finding is contrary to other reports that context extinction enhances responding to the target CS. Experiment 2 examined the amount of posttraining context extinction as a variable potentially controlling this effect. It found the mediated extinction effect following one, but not 10 h of context extinction.     

Conditioned taste avoidance induced by forced and voluntary wheel running in rats

Voluntary exercise by rats running in a freely rotating wheel (free wheel) produces conditioned taste avoidance (CTA) of a flavored solution consumed before running [e.g., Lett, B.T., Grant, V.L., 1996. Wheel running induces conditioned taste aversion in rats trained while hungry and thirsty. Physiol. Behav. 59, 699-702]. Forced exercise, swimming or running, also produces CTA in rats [e.g., Masaki, T., Nakajima, S., 2006. Taste aversion induced by forced swimming, voluntary running, forced running, and lithium chloride injection treatments. Physiol. Behav. 88, 411-416]. Energy expenditure may be the critical factor in producing such CTA. If so, forced running in a motorized running wheel should produce CTA equivalent to that produced by a similar amount of voluntary running. In two experiments, we compared forced running in a motorized wheel with voluntary running in a free wheel. Mean distance run over 30 min was equated as closely as possible in the two apparatuses. Both types of exercise produced CTA relative to sedentary, locked-wheel controls. However, voluntary running produced greater CTA than forced running. We consider differences between running in the free and motorized wheels that may account for the differences in strength of CTA.