Animals within zoo environments are learning continuously: they learn signals that predict when food is going to arrive or that the presence of a certain person means that something unpleasant may happen. They may learn to control their environment and caregivers: for example, they may learn that if they perform a particular behaviour (eg repetitive behaviour) they will receive a reward (ie food or attention from a caregiver). Using standard operant conditioning and classical conditioning techniques we can easily modify the behaviour of animals in zoos. Animals can be trained to comply with almost all minor veterinary procedures and examinations, such as injection, the measurement of heart rate, the cleaning of teeth and the treatment of superficial injuries. Compliance can be achieved using standard animal learning abilities without the need for punishment type I (ie physical punishment) or immobilisation (chemical or physical). In this paper we discuss how we apply learning theory to such procedures, the dangers associated with such programs (eg injury to the trainer) and the benefits (eg the treatment of large endangered animal species without the use of anaesthetic drugs). Additionally, we briefly discuss the selection and management of animal trainers. The methods we describe here are equally applicable to laboratory, farm and pet animals. Finally, as with all management processes applied to animals, a written policy on animal training needs to be produced by any institution training animals.

The aim of this paper was to develop a protocol to study the anticipatory response in cats as a measure of welfare. Seven experimental cats were trained in a classical conditioning paradigm to associate a sound with food arrival, while sound and food were presented without contingency in four control cats. Increasing the interval between sound and food up to 60 s allowed a detailed description of cats’ anticipatory response. Compared to control animals, experimental cats showed significantly shorter latencies to orient towards (average 2.96 s) and approach the source of the sound (12.98 s) as well as longer durations of exploring and standing by the source of the sound (namely 69.97 and 52.32%, respectively of the interval sound-food). Experimental cats also exhibited behaviours that may derive from predation patterns, eg short pauses and predatory crouch while approaching the source of the sound (namely in 28.93 and 29.64% of trials), rapid head movements while watching it (55.36% of trials) and pouncing on the food (9.29% of trials). This protocol should be further studied to assess its effectiveness in highlighting differences according to the welfare of individual cats.

A number of studies have investigated anticipatory behaviour in animals as a measure of sensitivity to reward or as an expression of emotional state. A common feature of many studies is that they base inferences on seemingly arbitrary measures, for example, the frequency of behavioural transitions (ie number of times an animal switches between different behaviours). This paper critically reviews the literature and discusses various hypotheses for why specific behavioural responses occur in the anticipatory period between the signal and reward in conditioned animals. We argue that the specific behaviours shown may be the result of superstitious learning and thus highly variable, leaving behavioural transitions as the only response that can be scored consistently, and that sometimes these responses may relate more to frustration than to a positive emotional state. Finally, we propose new research approaches to avoid potential confounds and improve future studies on this topic.

Previous research on electrodermal conditioning suggests that the conditioned diminution of the unconditioned response (UR) has an associative basis. The aim of this experiment was to test whether this phenomenon also occurs in heart rate (HR) classical conditioning. For this purpose, a differential classical conditioning was performed. The conditioned stimuli (CSs) were geometrical shapes (the CS+ was a square and the CS− was a triangle) displayed on a computer screen and a burst of white noise was used as unconditioned stimulus (US). For analysis of the conditioned response (CR) components, an interval between CS+ and US of 8 seconds was used. After the acquisition phase, participants were tested using trials with the US preceded either by a CS+, a CS−, or a neutral stimulus (a circle). The results showed conditioned diminution of the UR and suggest that the second heart rate deceleration component (D2) is responsible for the occurrence of this phenomenon.

Using a conditioned suppression task, we examined the minimal conditions to establish context conditioning as induced by unpredictability of an unconditioned stimulus (US). We investigated whether a biologically significant US is necessary to produce such context conditioning effects. In this between-subjects experiment, we manipulated the nature of the US and US-unpredictability. In the Paired condition, the conditioned stimulus (CS) was always followed by the US, whereas in the Unpaired condition, the CS and the US were presented explicitly unpaired, that is, the CS was never followed by the US. Half of the participants received an aversive, biologically significant human scream, and the other half received a more neutral, biologically non-significant sound as US. Results show more contextual suppression in the Unpaired condition than in the Paired condition. We conclude that in an expectancy-based conditioning task, US-unpredictability, but not a biologically potent US, is crucial to establish context conditioning.

Converging data from different disciplines are showing the role of classical conditioning processes in the elaboration of human and animal behavior to be larger than previously supposed. Restricted views of classically conditioned responses as merely secretory, reflexive, or emotional are giving way to a broader conception that includes problem-solving, and other rule-governed behavior thought to be the exclusive province of either operant conditiońing or cognitive psychology. These new views have been accompanied by changes in the way conditioning is conducted and evaluated. Data from a number of seemingly unrelated phenomena such as relapse to drug abuse by postaddicts, the placebo effect, and the immune response appear to involve classical conditioning processes. Classical conditioning, moreover, has been found to occur in simpler and simpler organisms and recently even demonstrated in brain slices and in utero. This target article will integrate the several research areas that have used the classical conditioning process as an explanatory model; it will challenge teleological interpretations of the classically conditioned CR and offer some basic principles for testing conditioning in diverse areas.

Most phobias are focussed on a small number of fear-inducing stimuli (e.g., snakes, spiders). A review of the evidence supporting biological and cognitive explanations of this uneven distribution of phobias suggests that the readiness with which such stimuli become associated with aversive outcomes arises from biases in the processing of information about threatening stimuli rather than from phylogenetically based associative predispositions or “biological preparedness.” This cognitive bias, consisting of a heightened expectation of aversive outcomes following fear-relevant stimuli, generates and maintains robust learned associations between them. Some of the features of such stimuli which determine this expectancy bias are estimates of how dangerous they are, the semiotic similarity between them and their aversive outcomes, and the degree of prior fear they elicit. Ontogenetic and cultural factors influence these features of fear-relevant stimuli and are hence important in determining expectancy bias. The available evidence does not exclude the possibility that both expectancy biases and specific evolved predispositions coexist, but the former can explain a number of important findings that the latter cannot.

The sway that the response-reinforcement framework (Spencer, Thorndike, Hull, Skinner) has held on the behavioral sciences for nearly a hundred years is finally ending. The strength of this framework lay in providing concepts and methods for studying the effects of hedonic (reinforcing) stimuli on the repetition of specified responses acquired in instrumental training situations of various kinds. Its weakness lay in the invalidity of its central assumptions, stimulus-response association and response-reinforcement, which could not deal with motor-equivalence and flexibility (or “intelligence”) in behavior. To the four decades of incisive criticism on particular theoretical and empirical grounds, a more comprehensive challenge to the response-reinforcement framework is now added by the newer ideas about the nature of cognitive, motivational, and response-production processes that have emerged from the work of ethologists, neuroscientists, and cognitive psychologists. An alternative framework, incorporating the newer ideas, is clearly needed.

The particular framework proposed here is based on the ideas of perceptual learning of stimulus-stimulus correlations and of a motivational (rather than reinforcing) role of hedonic (incentive) stimuli. According to it, an act is produced when its act-assembly is activated by a pexgo (perceptual representation) of a certain eliciting stimulus complex (ES). When certain eliciting stimuli are correlated with incentive stimuli, they acquire motivational properties that serve to strengthen the pexgos generated by those eliciting stimuli and thereby increase the probability of activation of the corresponding act-assemblies. Motivation thus influences response production, not by directly instigating “existing” responses, but by modulating the strength of pexgos of eliciting stimuli for the succession of acts that comprise a response. Therefore, a response is always constructed afresh on the basis of current perceptions; not even a stable and stereotyped response occurs as a mere activation of a preformed motor program. The topography of any response that emerges is determined by the nature of the motivational state and the momentary spatiotemporal distribution of eliciting stimuli of changing motivational valence.

By suggesting that the animal learns the overlapping and nested correlations between the stimulus events that commonly occur in a given situation, and by separating what is learned from the processes of response production, the proposed perceptual-motivational framework seems capable of dealing with the problems of motor equivalence and flexibility in adaptive behavior. Some implications of this approach for further behavioral and brain research on such problems as behavior modification, learning by observation of models, analysis of causality, and search for neural substrates of learning and response production, are outlined.

Behavioral science has long been puzzled by the experience of temptation, the resulting impulsiveness, and the variably successful control of this impulsiveness. In conventional theories, a governing faculty like the ego evaluates future choices consistently over time, discounting their value for delay exponentially, that is, by a constant rate; impulses arise when this ego is confronted by a conditioned appetite. Breakdown of Will (Ainslie 2001) presents evidence that contradicts this model. Both people and nonhuman animals spontaneously discount the value of expected events in a curve where value is divided approximately by expected delay, a hyperbolic form that is more bowed than the rational, exponential curve.

With hyperbolic discounting, options that pay off quickly will be temporarily preferred to richer but slower-paying alternatives, a phenomenon that, over periods from minutes to days, can account for impulsive behaviors, and over periods of fractional seconds can account for involuntary behaviors. Contradictory reward-getting processes can in effect bargain with each other, and stable preferences can be established by the perception of recurrent choices as test cases (precedents) in recurrent intertemporal prisoner's dilemmas. The resulting motivational pattern resembles traditional descriptions of the will, as well as of compulsive phenomena that can now be seen as side-effects of will: over-concern with precedent, intractable but circumscribed failures of self-control, a motivated (“dynamic”) unconscious, and an inability to exploit emotional rewards. Hyperbolic curves also suggest a means of reducing classical conditioning to motivated choice, the last necessary step for modeling many involuntary processes like emotion and appetite as reward-seeking behaviors; such modeling, in turn, provides a rationale for empathic reward and the “construction” of reality.

Many neuroscientists and philosophers endorse a view about the explanatory reach of neuroscience (which we will call the neuron doctrine) to the effect that the framework for understanding the mind will be developed by neuroscience; or, as we will put it, that a successful theory of the mind will be solely neuroscientific. It is a consequence of this view that the sciences of the mind that cannot be expressed by means of neuroscientific concepts alone count as indirect sciences that will be discarded as neuroscience matures. This consequence is what makes the doctrine substantive, indeed, radical. We ask, first, what the neuron doctrine means and, second, whether it is true. In answer to the first question, we distinguish two versions of the doctrine. One version, the trivial neuron doctrine, turns out to be uncontroversial but unsubstantive because it fails to have the consequence that the nonneuroscientific sciences of the mind will eventually be discarded. A second version, the radical neuron doctrine, does have this consequence, but, unlike the first doctrine, is highly controversial. We argue that the neuron doctrine appears to be both substantive and uncontroversial only as a result of a conflation of these two versions. We then consider whether the radical doctrine is true. We present and evaluate three arguments for it, based either on general scientific and philosophical considerations or on the details of neuroscience itself, arguing that all three fail. We conclude that the evidence fails to support the radical neuron doctrine.

A recent positron emission tomography (PET) study on brain mechanisms in classical, or Pavlovian, conditioning is reviewed. The PET data were compared with skin conductance data recorded off-line. The participants were five men who participated in three different experimental phases. In the first (habituation) phase, a tone was repeated 24 times at random intervals. In the second (acquisition) phase, the tone was paired with a brief shock to the wrist. In the third (extinction) phase, the tone was presented alone again. Statistical parametric mapping analysis of the PET data showed significantly increased activation of the right hemisphere in the orbitofrontal cortex, dorsolateral prefrontal cortex, inferior and superior frontal cortices, and inferior and middle temporal cortices. In the left hemisphere, only Area 19 and the superior frontal cortex showed significant activation. The findings are discussed within a theoretical framework that argues for different brain mechanisms in acquisition and extinction in classical conditioning.

Long-term potentiation (LTP) is operationally defined as a long-lasting increase in synaptic efficacy following high-frequency stimulation of afferent fibers. Since the first full description of the phenomenon in 1973, exploration of the mechanisms underlying LTP induction has been one of the most active areas of research in neuroscience. Of principal interest to those who study LTP, particularly in the mammalian hippocampus, is its presumed role in the establishment of stable memories, a role consistent with “Hebbian” descriptions of memory formation. Other characteristics of LTP, including its rapid induction, persistence, and correlation with natural brain rhythms, provide circumstantial support for this connection to memory storage. Nonetheless, there is little empirical evidence that directly links LTP to the storage of memories. In this target article we review a range of cellular and behavioral characteristics of LTP and evaluate whether they are consistent with the purported role of hippocampal LTP in memory formation. We suggest that much of the present focus on LTP reflects a preconception that LTP is a learning mechanism, although the empirical evidence often suggests that LTP is unsuitable for such a role. As an alternative to serving as a memory storage device, we propose that LTP may serve as a neural equivalent to an arousal or attention device in the brain. Accordingly, LTP may increase in a nonspecific way the effective salience of discrete external stimuli and may thereby facilitate the induction of memories at distant synapses. Other hypotheses regarding the functional utility of this intensely studied mechanism are conceivable; the intent of this target article is not to promote a single hypothesis but rather to stimulate discussion about the neural mechanisms underlying memory storage and to appraise whether LTP can be considered a viable candidate for such a mechanism.