Pavlov's Dogs Experiment

From Salivating Dogs in St. Petersburg to Modern Associative Learning

Ivan Petrovich Pavlov was a physiologist, not a psychologist, and he resisted the label of psychologist for most of his life. By the time he stumbled upon what would become classical conditioning, he had already won the 1904 Nobel Prize in Physiology or Medicine for his rigorous, surgically demanding research on the digestive system of dogs. The conditioning work — the work that made him famous to undergraduates a century later — emerged as a side observation during that digestive program. His dogs began salivating not just at the sight of food but at the appearance of the lab assistants who fed them, and Pavlov, a serious scientist who had learned not to dismiss such observations, decided to take the phenomenon seriously.

What followed was a decades-long, methodologically meticulous research program at the Institute of Experimental Medicine in St. Petersburg. Pavlov and his collaborators isolated the conditions under which a previously neutral signal — a metronome, a tuning fork, a light — could come to elicit salivation when reliably paired with food. They mapped acquisition, extinction, spontaneous recovery, generalization, discrimination, and second-order conditioning. The data set they accumulated remains one of the foundations of learning theory, and the concepts they formalized are still taught in essentially the same vocabulary today, even as modern work has substantially expanded what we understand associative learning to be.

Quick Facts About Pavlov's Experiments

  • Ivan Pavlov (1849–1936) worked at the Institute of Experimental Medicine in St. Petersburg, Russia
  • Awarded the 1904 Nobel Prize in Physiology or Medicine for research on digestion
  • Conditioning research began around the turn of the twentieth century and continued into the 1930s
  • Conditioned stimuli included metronomes, tuning forks, buzzers, lights, and tactile signals — not, in Pavlov's reports, a bell of the popular cliche
  • The salivary fistula allowed quantitative measurement of saliva drop by drop
  • Core findings: acquisition, extinction, spontaneous recovery, generalization, discrimination, second-order conditioning
  • His Lectures on Conditioned Reflexes were translated into English in 1927 by G. V. Anrep
  • Modern theory (Rescorla and Wagner, 1972) replaced simple S-R associations with predictive-error learning

1. Historical and Intellectual Context

Pavlov's intellectual milieu was Russian physiological materialism of the late nineteenth century, a tradition deeply marked by Ivan Sechenov's 1863 monograph Reflexes of the Brain. Sechenov had argued, controversially for the time, that all psychological phenomena could be reduced to reflexes — that thought, emotion, and volition were the cerebral analogs of the more obvious spinal reflexes. Sechenov's work pulled Russian neurophysiology toward an objective, measurement-based approach to mind, and away from the introspective philosophical psychology that dominated much of Western Europe.

Pavlov was trained in this tradition. He arrived in St. Petersburg in the 1870s as a student of physiology, worked with the eminent physiologist Carl Ludwig in Leipzig, and built his career on careful surgical preparations of intact animals to study normal physiological function. His 1897 lectures on digestion, translated into English as The Work of the Digestive Glands, are still cited as a model of integrative physiological investigation. By 1904 he had won the Nobel Prize for that work, recognized internationally as one of the great physiologists of the era.

The shift to conditioning was incremental rather than abrupt. Pavlov noticed during his digestion experiments that his dogs salivated not only when food was placed in their mouths (an obvious reflex) but also when they heard the footsteps of the assistant who fed them, when they saw the food bowl, when they were brought into the experimental room. He initially called these "psychic secretions," a label he later rejected because it imported subjective vocabulary into what he wanted to be a purely physiological investigation. He came to call them conditional reflexes — usually translated as conditioned reflexes — because they depended on prior conditions of pairing with food.

The political environment of the work was complicated. Pavlov was a critic of the Bolshevik regime after 1917, was personally protected by Lenin and Stalin because of his international stature, and used his protected position to speak more frankly about Soviet conditions than was safe for most scientists. He kept his laboratory operating through revolution, civil war, famine, and Stalinist purges. The continuity of his research program across those decades is itself a remarkable historical fact.

2. Research Questions

Pavlov's program asked a structured set of questions that became more sophisticated over decades. The initial question was descriptive: under what conditions does a previously neutral stimulus come to elicit a response that was originally elicited only by another stimulus? Subsequent questions investigated the parameters of acquisition: how many pairings are needed, what is the optimal temporal interval between the neutral and reinforcing stimulus, how does the strength of the unconditioned stimulus matter?

A second set of questions concerned the loss of conditioned responses. What happens when the conditioned stimulus is presented repeatedly without the unconditioned stimulus? Does the response disappear permanently, or does it recover with time? Can a previously extinguished response be re-acquired more easily than the original acquisition?

A third set concerned generalization and discrimination. Does the conditioned response transfer to stimuli similar to the original conditioned stimulus, and how does the strength of transfer vary with similarity? Can animals be trained to discriminate fine differences — to salivate to one tone but not to a slightly different one — and what happens when the discrimination is pushed beyond the animal's capacity?

A fourth set, less famous outside specialist circles, concerned higher-order conditioning, sensory preconditioning, and what Pavlov called the "investigation reflex" — an animal's tendency to orient toward novel stimuli. These extensions probed the limits of how complex associative structures could become from the simple pairing mechanism.

Across all these questions, Pavlov's deeper aim was to develop an objective, physiological account of higher nervous activity. He believed conditioning provided a tool for mapping the activity of the cerebral cortex, and he proposed neurological models — excitation, inhibition, irradiation, concentration — that he expected would eventually be confirmed by direct study of brain tissue. Some of his neurological speculation has aged poorly; the behavioral findings have aged remarkably well.

3. Method and Procedure

The Salivary Fistula

The methodological foundation of Pavlov's conditioning work was a surgical preparation he had refined for his digestion research. A small fistula was created in the dog's cheek, draining the duct of one salivary gland externally so that saliva could be collected drop by drop in a calibrated tube. The animals lived normally afterward, eating and drinking through their mouths in the ordinary way, but the experimenter had quantitative access to the salivary response. This was a genuine methodological breakthrough: it converted a vague behavioral observation into a continuous, measurable variable.

The Experimental Chamber

Pavlov designed sound-insulated experimental rooms — eventually a purpose-built "Tower of Silence" at the Institute of Experimental Medicine in St. Petersburg — to control extraneous stimuli. The dog stood on a small platform in a harness that prevented gross movement without causing distress, with the salivary tube positioned to collect drops. The experimenter operated from an adjacent room and observed through a periscope or one-way arrangement, presenting conditioned stimuli and food via remote-controlled devices.

The Conditioned Stimuli

Despite the textbook cliche, Pavlov's primary conditioned stimuli were not bells but metronomes, tuning forks, buzzers, lights, and tactile signals delivered through small mechanical devices. He chose these because they could be presented with precise onset and offset, at controlled intensities, and could be varied systematically — a metronome can be set to a particular rate, a tuning fork to a particular pitch — for discrimination work.

The Standard Procedure

In a basic acquisition trial, the experimenter presented the conditioned stimulus (the metronome) for a few seconds and then delivered food (the unconditioned stimulus). The dog salivated to the food reflexively. After repeated pairings — typically tens of trials, sometimes more — the dog began salivating to the metronome before the food arrived. The amount of saliva produced by the metronome alone, in the absence of food, was the conditioned response, and its volume across trials traced an acquisition curve.

Extinction

For extinction studies, the conditioned stimulus was presented repeatedly without food. The conditioned response decreased across trials, often following a relatively orderly decay curve. Pavlov observed that the decline was not symmetric with acquisition; extinction often progressed faster, and the response sometimes returned without further pairings after a period of rest — a phenomenon he called spontaneous recovery.

Generalization and Discrimination

To test generalization, Pavlov trained dogs to salivate to a particular tone, then probed them with tones of different pitches. Response strength varied systematically with similarity to the trained stimulus, producing what is now called a generalization gradient. To train discrimination, he reinforced one stimulus (a 60-beat-per-minute metronome) with food and presented a similar stimulus (a 40-beat-per-minute metronome) without food. With many trials, the dog learned to salivate to one but not the other. Pushing this discrimination to the limit of the dog's perceptual capacity sometimes produced what Pavlov called experimental neurosis, with whining, agitation, and breakdown of previously learned responses — an observation that influenced his later thinking about psychopathology.

Second-Order Conditioning

Once a stimulus had become a conditioned stimulus, Pavlov could use it as a substitute for the unconditioned stimulus in further conditioning. A new neutral stimulus (a light) paired with the established conditioned stimulus (the metronome), even without any food in the new pairings, could come to elicit salivation. This second-order conditioning showed that associative structures could be built on top of one another, extending the reach of the basic mechanism.

4. Participants and Setting

The Dogs

Pavlov's research used domestic dogs, typically mongrels obtained locally. Across his career and the work of his many collaborators, hundreds of dogs participated. Individual animals worked over months and sometimes years, and Pavlov kept records on each one as a research subject with its own temperament and conditioning history. He distinguished between "excitable" and "inhibited" dog types and incorporated these temperament differences into his theoretical models.

The dogs were housed at the Institute and were handled by lab staff who came to know them well. Pavlov insisted that the animals be treated calmly and humanely, partly out of welfare concern and partly out of methodological necessity: an agitated dog produces unreliable data. This is one of the reasons the sound-insulated chambers were built — to give the animal a calm, predictable environment in which conditioning trials could proceed without contamination by ambient stimuli.

The Institute of Experimental Medicine

The Institute of Experimental Medicine in St. Petersburg, founded in 1890, became the institutional home of Pavlov's research and one of the leading physiological laboratories in Europe. Pavlov directed the physiology department from 1891 until his death in 1936. The institute combined research with veterinary care, animal husbandry, and the training of large numbers of graduate students and visiting researchers, many of whom carried Pavlovian conditioning methods to laboratories in other countries.

The Collaborators

Although the work is universally identified with Pavlov, much of it was actually performed by a large team of assistants, graduate students, and visiting researchers. Names such as N. I. Krasnogorski, G. V. Anrep, and B. P. Babkin appear on the underlying experimental papers. Anrep was particularly important in transmitting Pavlov's work to English-speaking audiences; his 1927 translation, Conditioned Reflexes: An Investigation of the Physiological Activity of the Cerebral Cortex, brought Pavlov's lectures to readers like John B. Watson and influenced the development of American behaviorism.

5. Results

Acquisition

Across dogs and stimuli, the basic acquisition finding was robust. With consistent pairings of conditioned and unconditioned stimulus, conditioned responses appeared and grew across trials, typically reaching an asymptote after several dozen pairings. The shape of the acquisition curve depended on parameters such as the intensity of the conditioned stimulus, the size of the food reward, and the interval between conditioned and unconditioned stimulus. Short intervals (about half a second to several seconds, depending on the response) produced stronger conditioning than very long or very short ones.

Extinction

Repeated presentation of the conditioned stimulus alone reliably weakened the conditioned response. The number of unreinforced presentations needed to extinguish a response was usually much smaller than the number that had produced acquisition. However, extinction was not simply unlearning. Pavlov demonstrated several phenomena showing that the original learning was preserved: spontaneous recovery after a rest period, rapid reacquisition if pairing was resumed, and what was later called renewal — return of the conditioned response when the stimulus was presented in a different context than the one in which extinction had taken place.

Spontaneous Recovery

If an extinguished response was tested again after a rest period of hours or days, it often returned at partial strength without further pairings. Repeated extinction and rest cycles produced progressive weakening but rarely permanent abolition. This was one of Pavlov's most theoretically important findings: the original association was apparently preserved in some form, and extinction was an active inhibitory process layered on top of it rather than literal forgetting.

Generalization

Conditioned responses transferred to stimuli similar to the trained stimulus along the relevant sensory dimension. A tone of slightly different pitch produced a smaller but real conditioned response; a more dissimilar tone produced less. Generalization gradients were a stable, quantifiable feature of the system.

Discrimination and Experimental Neurosis

Animals could learn to discriminate between stimuli, with the unreinforced stimulus eventually producing no measurable response while the reinforced stimulus continued to elicit a strong response. When experimenters pushed discriminations to the limit of perceptual capacity — for example, training dogs to discriminate a perfect circle from a barely flattened ellipse — animals sometimes broke down. They became agitated, whined, refused food, lost previously learned discriminations, and showed what Pavlov labeled experimental neurosis. The observation influenced his later writing on the cerebral basis of human neurotic disorders.

Second-Order Conditioning

Second-order conditioning was demonstrable but weaker and more fragile than first-order conditioning. Conditioned responses based on second-order pairings were typically smaller in amplitude and extinguished more easily. Third-order conditioning was even harder to establish reliably. The implication was that complex chains of conditioned associations could be built but did not propagate indefinitely.

6. The Researchers' Interpretation

Pavlov interpreted his findings as a window onto the activity of the cerebral cortex. His central theoretical claim was that conditioned reflexes are produced by the temporary formation of functional connections in the cortex between two centers of excitation: the center activated by the conditioned stimulus and the center activated by the unconditioned stimulus. With repeated co-activation, the connection becomes strong enough that activation of the first center alone produces the response that previously required activation of the second.

He developed a vocabulary of cortical processes — excitation, inhibition, irradiation (the spread of an excitatory state to neighboring cortical areas), concentration (the localization of such a state), and the balance between excitatory and inhibitory tone — that he believed captured the fundamental dynamics of higher nervous activity. Some of this vocabulary survived in modified form into later neuroscience; much of it was speculative and is no longer used.

Pavlov was emphatic that his work was physiology, not psychology. He resisted any explanation that appealed to the dog's beliefs, expectations, or knowledge. The dog salivated because the conditioned stimulus had become functionally equivalent to the unconditioned stimulus through cortical connection, not because the dog "expected" food. This insistence was partly a matter of methodological discipline — Pavlov did not want to import vocabulary he could not measure — and partly a matter of intellectual politics in distinguishing his work from the psychology of his day.

He extended his findings to human pathology cautiously but increasingly in later years. Experimental neurosis in dogs suggested that excessive demands on the discriminative capacity of the nervous system could produce maladaptive behavior. Pavlov speculated that human neuroses might similarly reflect a breakdown of the normal balance between excitation and inhibition in the cortex, and that conditioning principles might inform their treatment. This speculative extension was taken up by his students and indirectly shaped Soviet psychiatry through much of the twentieth century.

7. Modern Reanalyses and Criticisms

From Stimulus Substitution to Predictive Cues

The most consequential modern revision is the move away from Pavlov's view of conditioning as stimulus substitution — the conditioned stimulus comes to evoke a response because it has become functionally equivalent to the unconditioned stimulus — toward a view of conditioning as the learning of predictive relationships. The decisive work was done in the 1960s by Robert Rescorla and Allan Wagner, whose 1972 model treated conditioning as a process by which an organism learns the predictive value of cues with respect to outcomes. Crucially, the strength of conditioning depends not on the number of pairings but on the prediction error generated by each pairing — the difference between what the organism expects and what actually happens.

Rescorla's earlier demonstration of blocking (Leon Kamin) and his "truly random control" work showed that simple co-occurrence of conditioned and unconditioned stimulus was insufficient for conditioning. If the conditioned stimulus did not provide new predictive information — because the unconditioned stimulus was equally likely whether the conditioned stimulus appeared or not — conditioning did not occur. This finding undermined Pavlov's simple pairing account.

Predictive Cues and Occurrence Cues

Modern associative learning theory distinguishes between cues that signal that an outcome is about to occur (predictive cues) and cues that merely co-occur with the outcome (occurrence cues). Only predictive cues — those that change the organism's estimate of what is coming — produce strong conditioning. This refinement preserves the empirical phenomena Pavlov documented while replacing the underlying mechanism.

Garcia Effect and Biological Preparedness

John Garcia's work in the 1950s and 1960s on conditioned taste aversion showed that not all stimuli are equally conditionable. Rats readily learn to associate a taste with subsequent illness even when the gap between taste and illness is many hours. They do not readily learn to associate the same illness with a light or sound. The same rats readily learn to avoid a sound that predicts shock, but struggle to associate a taste with shock. The pattern shows that associative learning is selectively prepared by evolution: certain cue-outcome relationships are easy to learn because they correspond to relationships that mattered in the ancestral environment, while others are difficult or impossible even with many pairings. This was a major qualification of Pavlov's view that any neutral stimulus could be conditioned to any unconditioned stimulus.

Context and Renewal

Modern extinction research has substantially elaborated Pavlov's observation of spontaneous recovery. Mark Bouton and colleagues have shown that extinction is highly context-dependent: a response extinguished in one context returns when the conditioned stimulus is presented in a different context. This finding has direct clinical implications. Exposure therapy that works perfectly in the therapist's office may fail in the patient's daily environment because the extinction learning is bound to the therapy context. Modern protocols attempt to generalize extinction by conducting exposure in multiple contexts and by combining it with cognitive elements.

Reconsolidation

More recent work suggests that retrieval of a conditioned memory briefly returns it to a labile state during which it can be modified or, in some preparations, erased. Reconsolidation-based interventions, in which a conditioned response is reactivated and then either weakened pharmacologically or replaced behaviorally, are a current frontier of translational research. The phenomena are real but more complex than initial enthusiastic reports suggested.

The Bell Myth

One smaller modern correction is worth noting. The popular image of Pavlov ringing a bell, while not entirely fictional, is somewhat misleading. Pavlov's primary conditioned stimuli were metronomes, tuning forks, buzzers, and lights, chosen for their precise controllability. Bells appear occasionally in the secondary literature and in some translations, but the bell of the textbook cliche is a simplification of a more varied research program.

8. Ethical Considerations

Pavlov's research was conducted before any formal regulation of animal experimentation, and judged by contemporary standards it raises real concerns. The salivary fistula was a surgical alteration of a healthy animal. Dogs were restrained in harnesses for hours of conditioning trials. The experimental neurosis induced by impossibly fine discriminations produced visible distress: whining, agitation, refusal of food, breakdown of previously stable behavior. Other dogs were subjected to studies involving electrical shock as unconditioned stimulus, which Pavlov's group used in some defensive-reflex conditioning work.

It would be incorrect to portray Pavlov as indifferent to animal welfare. By the standards of his era he was attentive: he insisted on careful surgical technique, recovery time, calm handling, predictable routine, and the use of food rewards in a substantial portion of his work. He treated his dogs as research subjects whose long-term cooperation mattered, and many animals served across years of work and were retired rather than discarded. He was famously distressed by the loss of dogs during the 1924 St. Petersburg flood, when several drowned, and the incident is sometimes cited as evidence of his attachment to the animals.

Modern animal-research ethics — the three Rs of replacement, reduction, and refinement — would constrain elements of Pavlov's program. The most invasive surgeries would require strong justification today. The experimental-neurosis studies would face questions about whether the scientific gain justified the induced distress. Pavlov's program could be largely replicated under modern oversight, but with refinements and limitations he did not consider.

There is also a softer ethical question about the use of Pavlov's findings in human research. The first major human application — Watson and Rayner's conditioning of fear in Little Albert — was an example of the principles being deployed without the kind of protections Pavlov himself provided to his dogs. The history of behavior therapy includes both the principled, careful work of researchers who used conditioning ethically and the cautionary work that prompted the development of human-subjects regulations.

9. Influence on Psychology

Behaviorism

Pavlov's work, translated into English in 1927 by Anrep, became one of the principal scientific foundations of American behaviorism. John B. Watson explicitly built his behaviorist programme on Pavlovian conditioning, and his 1920 study of Little Albert was meant to demonstrate that the mechanism applied to human emotional learning. B. F. Skinner's later operant work, while focused on a different form of conditioning, took for granted the Pavlovian framework as part of the broader landscape of associative learning.

Behavior Therapy

The direct line from Pavlov's laboratory to the modern clinic runs through Mary Cover Jones's 1924 work with Peter, through Joseph Wolpe's systematic desensitization in the 1950s, and through the development of modern exposure-based therapies. Exposure therapy for phobias, panic disorder, post-traumatic stress disorder, and obsessive-compulsive disorder is built on the principle that conditioned fear responses can be extinguished through repeated unreinforced presentations of the conditioned stimulus — exactly the procedure Pavlov used to extinguish salivary responses. The clinical sophistication added over the past half-century includes attention to context generalization, prediction error in exposure design, and the integration of cognitive elements, but the basic mechanism is Pavlovian.

Learning Theory

The Rescorla-Wagner model and its successors — Mackintosh, Pearce-Hall, Wagner's SOP, and modern Bayesian and reinforcement-learning frameworks — all build on the empirical foundation Pavlov established. Modern reinforcement learning, the algorithmic framework that powers many current artificial-intelligence systems, owes a direct conceptual debt to the Pavlovian and operant learning traditions. The temporal-difference learning rule of Richard Sutton and Andrew Barto, fundamental to modern AI, is mathematically related to the Rescorla-Wagner update equation.

Taste Aversion and Biological Preparedness

Garcia's taste-aversion work and Martin Seligman's framework of biological preparedness reshaped how psychology thinks about the relationship between learning mechanisms and evolutionary history. Phobias of snakes and spiders, for instance, are more easily acquired and more resistant to extinction than phobias of electrical outlets or modern dangers, consistent with the idea that the fear-learning system was shaped by ancestral threats.

Immune and Physiological Conditioning

Robert Ader's work in the 1970s demonstrated that immune responses themselves can be classically conditioned. Pairing a flavored drink with an immunosuppressive drug led, after several pairings, to immune suppression in response to the flavor alone. This and subsequent findings opened the field of psychoneuroimmunology and showed that Pavlovian principles extend well beyond simple behavioral responses into autonomic, endocrine, and immune systems. Similar conditioning of placebo and nocebo effects has clinical implications across medicine.

Addiction

Drug tolerance, craving, and relapse have been understood in part through Pavlovian conditioning. Environmental cues that have been reliably paired with drug administration acquire the capacity to elicit compensatory or anticipatory physiological responses, contributing to tolerance when the drug is taken in familiar contexts and to overdose when it is taken in unfamiliar ones. Cue-exposure therapies for substance use disorder are built on this analysis.

10. What the Experiment Means Today

Pavlov's conditioning research is one of the few experimental programs in psychology where the basic empirical findings are essentially uncontested more than a century later. Acquisition, extinction, spontaneous recovery, generalization, discrimination, and second-order conditioning have been demonstrated repeatedly across species, sensory modalities, response systems, and laboratories. The mathematical structure of these phenomena, refined by Rescorla-Wagner and subsequent models, has been integrated with modern neuroscience and computational learning theory.

What has changed is the theoretical interpretation. Pavlov saw conditioning as stimulus substitution mediated by cortical connections. Modern researchers see it as the learning of predictive relationships, governed by prediction error and shaped by evolutionary preparedness, anchored in specific brain circuits (the amygdala for fear conditioning, the cerebellum for motor conditioning, the dopaminergic midbrain for reward prediction). The Pavlov of the textbooks is a useful first approximation. The Pavlov of the modern literature is a more sophisticated mechanism that includes attention, expectation, and context.

For the clinic, the Pavlovian inheritance is concrete and growing. Exposure therapies remain among the most strongly supported psychological treatments in psychiatry. The mechanistic understanding of extinction — including its context-dependence, its reliance on prediction error, and its potential modulation by pharmacological agents such as D-cycloserine — has driven a new generation of more targeted protocols. The same framework underlies emerging interventions in addiction, chronic pain, and traumatic memory.

For the student, Pavlov's work is now taught with a layered respect. The basic phenomena are reliable and worth memorizing. The simple S-R interpretation is a useful introduction but should be followed quickly with the predictive-cue framework. The history of the work — a Nobel-winning physiologist following an unexpected observation into a new field, building methods that allowed quantitative measurement of an unobservable internal process, and changing the trajectory of psychology in the process — is itself a model of how good science actually unfolds.

Conclusion

Ivan Pavlov's experiments on conditioned reflexes are among the most generative pieces of empirical work in the history of psychology, even though Pavlov himself rejected the discipline as poorly defined. By turning the salivary response of a restrained dog into a quantitative behavioral variable, he made it possible to study learning with the precision previously reserved for physiology. The empirical phenomena he documented — acquisition, extinction, generalization, discrimination, spontaneous recovery, second-order conditioning — have been replicated countless times and form the bedrock of modern associative learning theory.

The theoretical interpretation has changed substantially. The simple model of stimulus substitution by cortical connection has given way to a richer framework in which organisms learn the predictive structure of their environment, in which prediction error governs the rate of learning, in which biological preparedness constrains what can be learned, and in which extinction is an active, context-dependent inhibitory process rather than mere forgetting. The modern picture is more complex than Pavlov's, but it is built on his foundation.

Clinically, the descendants of Pavlov's program are everywhere. Exposure therapy for anxiety disorders, cue-exposure for addiction, the analysis of placebo and nocebo effects, the conditioning of immune responses, and the temporal-difference learning rules at the heart of modern artificial intelligence all owe a measurable debt to the work that began with dogs salivating at the footsteps of laboratory assistants in St. Petersburg. The unexpected observation that the careful physiologist refused to dismiss has, in the long run, turned out to be one of the most consequential side findings in the history of science.