Lehrstuhl für Genetik und Neurobiologie
Theodor-Boveri-Institut für Biowissenschaften
BIOZENTRUM - Universität Würzburg
  
Reflexology - a historical outline  

Reflexology - an historical outline

The origins of the concept of reflexes in behaviour can be traced back to "Discours de la Méthode" by René Descartes (1637). He asserted that the actions of animals (but not man) could be explained entirely in terms of physical principles. Largly through dissection studies, he investigated what functions the various bodily organs might serve in the operation of animals. Descartes was particularly scathing about the quality of medical knowledge in the seventeenth century, most of which was not based on empirical evidence and called for public support of experimental physiological research. Although Descartes held very firmly to the view that man, but not animals, had a soul, and that therefore the behaviour of animals, but not man, could be explained mechanistically, he did suggest that involuntary behaviours in both man and animals may be explained on a common basis. Descartes was inspired by animated statues in the palace at St.Germain which were controlled hydraulically. For example, a statue of the goddess Diana bathing naked would retreat into some rosebushes when an approaching art-lover trod on a panel concealed in the pathway. Pressure on this panel opened a valve which released a flow of water which caused the statue to move. Descartes suggested that 'animal spirits' flowing through the nerves of animals or humans served a similar function in automatic behavioural responses in man and animals or reflexes. The term 'reflex' is derived from the notion that the flow of animal spirits produced by a stimulus is somehow reflected by the brain into an outgoing flow which eventually produces some behaviour. However, the assumptions upon which this theory were based had been shown to be false experimentally within Descartes' lifetime - nerves are not hollow, and the contraction and expansion of muscles is not achieved through inflating or emptying them of some fluid. Nevertheless, Descartes' concept of the reflex and of the explicability of human and animal behaviour mechanistically, although not widely accepted at the time, shaped much following physiological research. 

Acceptance of the importance of reflexes was limited while the actions of nerves were still not understood. In the interim a wide range of behaviours were suggested to be reflexive in action, for example, digestion, coughing, sneezing, pupilliary reaction to light and so on. In addition, it was suggested by La Mettrie in his "Man a Machine" (1748), although it offended many deeply, that the behaviour man as well as that of animals could be explained entirely mechanistically. Nevertheless, it was hard to take these ideas seriously while there was no sensible mechanistic explanation of nervous action. The breakthrough came with flood of late eighteenth century work on electricity, e.g. Benjamin Franklin's accounts of his experiments with lightening published in 1751. Demonstrations of the effects of electricity, produced by generators of static electric charges, were also popular theatrical entertainments at this time. It had been suggested earlier in the century that electricity might form the basis of Descartes' elusive 'animal spirits', however, the absence of appropriate insulation in the nervous system seemed to rule the idea out. This all changed with Luigi Galvani's experiments with frogs and static electricity. He claimed to demonstrate that electrical energy was generated in the nervous systems of animals. As his experimental results did not justify his conclusions (something still not uncommon) a period of controversy during which many more experiments carried out in Italy by Allessandro Volta, Galvani's nephew Giovanni Aldini and others lead to much greater understanding of electricity and it's role in conducting signal in the nervous system (although precise understanding of this would not happen until the mid-twentieth century). Precise understanding was not, however, necessary for the theory of reflexes to be much more attractive now that a reasonable mechanism of nervous signal conduction had been suggested. The focus of work on neurophysiology and reflexes then moved to Germany, culminating in the measurement of the speed of nervous signal conduction in reflexes by Herman von Helmholtz (whose work on electricity also made an impact in physics). 

The solid conceptual status of the reflex as a neurally mediated automatic response to a stimulus now inspired some German scientists to suggest that all behaviour was in fact automatic - spontaneous activity was impossible - all behaviour was the result of reflexes, however, some reflexes were clearly simpler than others. The major problem facing this argument was evidence (initially from experiments on brainless and sometimes legless frogs) that should one reflex fail to be effective for some reason the stimulus inducing it would, after some time, begin to elicit a different behaviour. This is very difficult to explain if all nerves can do is excite activity in muscles, glands or other nerves. Luckily, the ability of the vagus nerve to decrease heart-rate - an inhibitory action, had recently been discovered. The validity of the suggestion that all behaviour was automatic therefore rested on discovering whether (and by what means) reflexes could be inhibited. A Russian, Ivan Sechenov, was the first person to appreciate the significance of inhibition and reflex action. During work in Germany and France Sechenov showed that by placing salt crystals in certain parts of a frogs brain he could reversibly inhibit its leg-withdrawal reflex. He returned to St. Petersburg and in 1863 published a monograph, 'Reflexes and the Brain', describing this work. Sechenov's description of reflex had begun to diverge from Descartes' notion of simple fixed 'reflections' of stimuli. First, Sechenov suggested that the strength of stimuli and the responses they elicited need not be similar - very weak stimuli might trigger quite intense reactions. Second, Sechenov suggested that reflexes are ubiquitous and malleable, for example, he suggested that, as it was his habit to think of politics before going to bed each night it might happen that if he were to lie down in the daytime the properties of his bedroom might elicit thoughts of politics in him. Sechenov felt that inhibition played a significant role in both these extensions of Descartes' concept of the reflex. Having demonstrated the existence of centrally mediated inhibition of reflexes, however, Sechenov did not go on to test these later inferences, perhaps because of the political and social conditions in Russia towards the end of the nineteenth century. Nevertheless, Russian experimental physiology thrived. Sergei Botkin, who had accompanied Sechenov on some of his studies in Germany, became Professor of Clinical Medicine at the Military- Medial Academy in Moscow where he maintained an animal laboratory for the experimental study of physiology. In 1878, Botkin appointed a highly recommended young physiology graduate from St. Petersburg, Ivan Pavlov to be its director. 

Pavlov's early work, for which he was eventually gained the directorship of the Oldenburgski Institute of Experimental Medicine in St. Petersburg - intended to be a Russian equivalent of the Pasteur Institute, along with other high posts, was not aimed at understanding the mental role of the reflex. Rather, it was in the physiology of the digestive system. He was interested in the way in which particular foodstuffs might elicit the release of gut chemicals particularly suited to digesting them and how these reflexes integrated with other muscular reflexes involved in digestion to produce a balanced and integrated process. In order to pursue this study Pavlov pioneered procedures in which careful surgery was carried out on animals, usually dogs, from which they recovered to lead near-normal lives (at least physiologically normal ones) in contrast to the more extreme acute studies on frogs which had typified experimental physiology up to that point. Two techniques Pavlov perfected were first the insertion of tubes into an animals throat so that, when the experimenter desired, food could be eaten by the animal without reaching its stomach, and second, the insertion of a tube into the stomach so that gastric secretions could be collected for subsequent analysis. This second tube had an important intellectual and financial role in Pavlov's laboratory. Financially it was important because drinking the gastric juices of animals had become a popular cure for stomach ailments around St. Petersburg and Pavlov's laboratory was able to supply large quantities of this hard to obtain commodity. Intellectually the gastric tube was important because it allowed people to observe that Pavlov's dogs would start secreting gastric juices not just when food was present in their mouths, but at the mere sight of food in the test room, or indeed at the sight of their regular feeder. This secretion, elicited by psychological expectations of food rather than food itself, became termed 'psychic secretion' and was assumed to be quite different from food-induced gastric secretions. Although the phenomenon was observed regularly in Pavlov's laboratory it was basically disregarded until nearly the end of the century - 1897. At this time Pavlov, now in his 50's, realised that physiological methods could be used to study psychological phenomena and that these phenomena must be described and explained in physiological terms if they are to be understood. He had, to some extent, been anticipated by some of his student who had already begun to work on psychic secretions. Stefan Wolfsohn had discovered that the amount and nature of saliva produced by different foods varies according to the type of food. The same was true of sights of different foods. Another student, Anton Snarsky, went on to show that apparently arbitrary signals could elicit the same effects. Dilute acids produce a copious amount of a particular type saliva. Snarsky found that if he coloured this acid solution black then soon the sight of any black liquid would induce large amounts of this sort of salivation. This is probably the first recorded attempt intentional classical-conditioning experiment; all the crucial features are present - a pre-existing reflex and an arbitrary stimulus which eventually becomes capable of eliciting the same response produced in that reflex. Work in Pavlov's laboratory was now firmly directed at understanding classical-conditioning and related phenomena. Pavlov strongly rejected mentalist explanations of conditioning couched in terms of an animals 'expectations' or 'desires' , in the process falling out with Snarsky, much of his work was concerned with pinning down the neural basis of classical-conditioning - a task made somewhat more difficult by the discovery by two British workers, Starling and Bayliss, who showed that food- related information may be transmitted hormonally rather than neurally. 

This document was restructured from a lecture kindly provided by R.W.Kentridge

 
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