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 |