Classical conditioning
Classical conditioning (also respondent conditioning and Pavlovian conditioning) is a behavioral procedure in which a biologically potent stimulus (e.g. food, a puff of air on the eye, a potential rival) is paired with a neutral stimulus (e.g. the sound of a musical triangle). The term classical conditioning refers to the process of an automatic, conditioned response that is paired with a specific stimulus.[1]
"Pavlov's dog" and "Pavlovian" redirect here. For the Pavlovian Upper Paleolithic culture, see Pavlovian culture. For other uses, see Pavlov's dog (disambiguation).
The Russian physiologist Ivan Pavlov studied classical conditioning with detailed experiments with dogs, and published the experimental results in 1897. In the study of digestion, Pavlov observed that the experimental dogs salivated when fed red meat.[2] Pavlovian conditioning is distinct from operant conditioning (instrumental conditioning), through which the strength of a voluntary behavior is modified, either by reinforcement or by punishment. However, classical conditioning can affect operant conditioning; classically conditioned stimuli can reinforce operant responses.
Classical conditioning is a basic behavioral mechanism, and its neural substrates are now beginning to be understood. Though it is sometimes hard to distinguish classical conditioning from other forms of associative learning (e.g. instrumental learning and human associative memory), a number of observations differentiate them, especially the contingencies whereby learning occurs.[3]
Together with operant conditioning, classical conditioning became the foundation of behaviorism, a school of psychology which was dominant in the mid-20th century and is still an important influence on the practice of psychological therapy and the study of animal behavior. Classical conditioning has been applied in other areas as well. For example, it may affect the body's response to psychoactive drugs, the regulation of hunger, research on the neural basis of learning and memory, and in certain social phenomena such as the false consensus effect.[4]
Definition[edit]
Classical conditioning occurs when a conditioned stimulus (CS) is paired with an unconditioned stimulus (US). Usually, the conditioned stimulus is a neutral stimulus (e.g., the sound of a tuning fork), the unconditioned stimulus is biologically potent (e.g., the taste of food) and the unconditioned response (UR) to the unconditioned stimulus is an unlearned reflex response (e.g., salivation). After pairing is repeated the organism exhibits a conditioned response (CR) to the conditioned stimulus when the conditioned stimulus is presented alone. (A conditioned response may occur after only one pairing.) Thus, unlike the UR, the CR is acquired through experience, and it is also less permanent than the UR.[5]
Usually the conditioned response is similar to the unconditioned response, but sometimes it is quite different. For this and other reasons, most learning theorists suggest that the conditioned stimulus comes to signal or predict the unconditioned stimulus, and go on to analyze the consequences of this signal.[6] Robert A. Rescorla provided a clear summary of this change in thinking, and its implications, in his 1988 article "Pavlovian conditioning: It's not what you think it is".[7] Despite its widespread acceptance, Rescorla's thesis may not be defensible.[8]
Classical conditioning differs from operant or instrumental conditioning: in classical conditioning, behaviors are modified through the association of stimuli as described above, whereas in operant conditioning behaviors are modified by the effect they produce (i.e., reward or punishment).[9]
Phenomena observed[edit]
Acquisition[edit]
During acquisition, the CS and US are paired as described above. The extent of conditioning may be tracked by test trials. In these test trials, the CS is presented alone and the CR is measured. A single CS-US pairing may suffice to yield a CR on a test, but usually a number of pairings are necessary and there is a gradual increase in the conditioned response to the CS. This repeated number of trials increase the strength and/or frequency of the CR gradually. The speed of conditioning depends on a number of factors, such as the nature and strength of both the CS and the US, previous experience and the animal's motivational state.[6][9] The process slows down as it nears completion.[16]
External inhibition[edit]
External inhibition may be observed if a strong or unfamiliar stimulus is presented just before, or at the same time as, the CS. This causes a reduction in the conditioned response to the CS.
Theories[edit]
Data sources[edit]
Experiments on theoretical issues in conditioning have mostly been done on vertebrates, especially rats and pigeons. However, conditioning has also been studied in invertebrates, and very important data on the neural basis of conditioning has come from experiments on the sea slug, Aplysia.[6] Most relevant experiments have used the classical conditioning procedure, although instrumental (operant) conditioning experiments have also been used, and the strength of classical conditioning is often measured through its operant effects, as in conditioned suppression (see Phenomena section above) and autoshaping.
Applications[edit]
Neural basis of learning and memory[edit]
Pavlov proposed that conditioning involved a connection between brain centers for conditioned and unconditioned stimuli. His physiological account of conditioning has been abandoned, but classical conditioning continues to be used to study the neural structures and functions that underlie learning and memory. Forms of classical conditioning that are used for this purpose include, among others, fear conditioning, eyeblink conditioning, and the foot contraction conditioning of Hermissenda crassicornis, a sea-slug. Both fear and eyeblink conditioning involve a neutral stimulus, frequently a tone, becoming paired with an unconditioned stimulus. In the case of eyeblink conditioning, the US is an air-puff, while in fear conditioning the US is threatening or aversive such as a foot shock.
The American neuroscientist David A. McCormick performed experiments that demonstrated "...discrete regions of the cerebellum and associated brainstem areas contain neurons that alter their activity during conditioning – these regions are critical for the acquisition and performance of this simple learning task. It appears that other regions of the brain, including the hippocampus, amygdala, and prefrontal cortex, contribute to the conditioning process, especially when the demands of the task get more complex."[29]
Fear and eyeblink conditioning involve generally non overlapping neural circuitry, but share molecular mechanisms.
Fear conditioning occurs in the basolateral amygdala, which receives glutaminergic input directly from thalamic afferents, as well as indirectly from prefrontal projections. The direct projections are sufficient for delay conditioning, but in the case of trace conditioning, where the CS needs to be internally represented despite a lack of external stimulus, indirect pathways are necessary. The anterior cingulate is one candidate for intermediate trace conditioning, but the hippocampus may also play a major role. Presynaptic activation of protein kinase A and postsynaptic activation of NMDA receptors and its signal transduction pathway are necessary for conditioning related plasticity. CREB is also necessary for conditioning related plasticity, and it may induce downstream synthesis of proteins necessary for this to occur.[30] As NMDA receptors are only activated after an increase in presynaptic calcium(thereby releasing the Mg2+ block), they are a potential coincidence detector that could mediate spike timing dependent plasticity. STDP constrains LTP to situations where the CS predicts the US, and LTD to the reverse.[31]