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Fluctuation in excitability
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Verveen, A.A., 1960. On the fluctuation of threshold of the nerve fibre.
In: Structure and Function of the Cerebral Cortex. D.B. Tower and J.P. Schadé, eds. Proceedings of the Second International Meeting of Neurobiologists (Amsterdam 1959). Amsterdam,Elsevier: 282-288.
In this paper, I report on the (re)discovery of threshold fluctuations of nerve fibres.
The paper summarizes chapter 3 of my 1961 thesis.
The studies of Pecher (La fluctuation d'excitabilite de la fibre nerveuse, Arch Intern Physiol, 1939 and his earlier papers, see below) and, earlier, Blair and Erlanger (1932-1936) and Monnier and Jasper (1932) had been utterly forgotten during "the reign of the Hodgkin-Huxley (H.H.) model", but  for Warren S. McCulloch (the father of probabilistic neurons) (1958), Lawrence S. Frishkopf (1956) and Walter A. Rosenblith (1959), as I was to learn much later. All three were at that time working at the Massachussetts Institute of Technology, MIT, Cambridge, USA.   

I confirmed Blair & Erlanger's (1933) and Pecher's (1937) conclusion that the stochastic behaviour of nerve fibres is due to an inherent property of the fibre.  People considered  the H.H.-model being deterministic, so my reports on the stochastic behaviour of nerve fibres were for a long time thought to be just a description of artifacts, despite proof of the reverse, so I "had been stumbling into the pitfall of instrumental noise". This attitude changed, gradually, only after the appearance of our 1966 paper in Science. I am, therefore, not impressed by the peer referee system. It is the scientific community as such who, in the long run, decides what is worth knowing. It is, by the way, also possible that people are reluctant to accept that nervous systems including our own are probabilistic.   

Only two people, Warren S. McCulloch in Cambridge, Mass. and Henk van der Tweel in Amsterdam, did in the mean time consider my measurements and results to be of a true property of neural tissue.

The blue-green background indicates that the paper is still cited occasionally,
more than 30 years after the original date of publication.



Verveen, A.A., 1961. Fluctuation in Excitability.
Research report on signal transmission in nerve fibers.
Amsterdam: Netherlands Central Institute for Brain Research. 76 pages.
PhD-thesis, July 6th, 1961
     The intensity of the threshold fluctuations relative to the threshold, the "relative spread" ( RS ), appeared to be a stimulus independent parameter of stochastic nerve fiber behaviour. Chemical substances could influence its intensity.
This study was motivated by the thought that stochastic behaviour plays a functional role in the function of the nervous system, even if only by statistically conveying information about stimulus intensities of about threshold size.

Verveen, A.A., 1962.
The influences of certain conditions on the fluctuation in excitability of the frog node of Ranvier.
Acta Physiol. Pharmacol., Neerlandica
10: 294-295.  
     (Summary of part of the 1961 work)
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Verveen, A.A., 1962. Probability phenomena in unmyelinated crayfish axon.
Acta Physiol. Pharmacol., Neerlandica
11: 268-269.  
     (Summary of part of the 1961 work)

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Verveen, A.A., 1962. A probability phenomenon in peripheral axon.
Proc. XXII International Congress. Physiol. (Leiden) 2: 788. (Abstract)
     Of historical importance only, since reading this paper and the the next one by Derksen brought us into contact and started our quest into the underlying process in the nerve membrane.

28 pdf
Derksen, H.E. and F. van der Mark, 1962.
Influence of resting activity on sensory threshold.

Proc. XXII International Congress Physiol. (Leiden)
2: 950.
(Abstract, see Derksen, 1964)
     See above.

68 pdf
Verveen, A.A., 1962. Axon diameter and fluctuation in excitability.
Acta Morphologica Neerlando-Scandinavica 5 (1): 79-85.
     An inverse relationship was found to exist between the  size of the fluctuations as characterized by the relative spread (RS) and nerve fibre diameter.
194 pdf
Ten Hoopen, M. and  A.A. Verveen, 1963.
Nerve-Model Experiments on Fluctuation in Excitability.

Progress in Brain Research
2: 8-21.
     The injection of noise into a neuron model reproduced the properties of fluctuation in excitability.
Intensity as well as bandwidth of the injected noise influenced the size of the fluctuations.
The stochastic response generated by the injection of noise did then contain information about the intensity of the stimulus.
519 pdf
Verveen, A.A. and J.P. Schadé. 1964. Excitability fluctuation in axons of Sepia officinalis. Netherlands Journal of Sea Research 2 (2): 145-154.
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Derksen, H.E., 1964. Discussion.         
In: Information processing in the nervous system. R.W. Gerard and J.W. Duyff, eds. Proceedings of the International Union of Physiological Sciences, 22d International Congress, Leiden, vol. 3: 384-394.
     On page 392 of this paper Derksen showed that the discontinuous step-response of a neuron-model aggregate to a continuously varying intensity of the stimulus transforms upon the injection of noise into a stochastically smooth response range.
This paper is forgotten and needs to be brought back into attention.
170 pdf
Verveen, A.A. and H.E. Derksen, 1965.
Fluctuations in membrane potential of axons and the problem of coding.

Biological Cybernetics
2: 152-160. (Kybernetik 2: 152-160).
     This journal was originally called Kybernetik.
     The paper discusses thoughts about a possible function of the irregular fluctuations in the membrane potential of neurons and, hence, in excitability.
582 pdf
Pecher, C., 1936.
Étude statistique des variations spontanées de l’excitabilité d’une fibre nerveuse.

C. R. Soc. Biol. 122: 87-91.
(Statistical study of the spontaneous variations in excitability of a nerve fibre).
2.125 pdf
Pecher, C., 1937.
Fluctuations indépendantes de l’excitabilité de deux fibres d’un même nerf.
C. R. Soc. Biol. 124: 839-842.
(Independent fluctuations in excitability of two fibres in the same nerve).
1.652 pdf
Pecher, C., 1939.
La fluctuation d’excitabilité de la fibre nerveuse.
Arch. Intern. Physiol. 49 (2): 129-152.
(Fluctuation in excitability of the nerve fibre).
This paper counts 9.484 kB and has been divided into three parts
For my personal account on Charles Pecher's contributions, click here
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