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Action potentials and frequency of firing Let us look at the myth of action potentials and firing patterns more in detail. This is a myth that the authors are upholding without any compunction ("we can see that the information about stimulus strength is now encoded in action potential frequency", p.342) in their otherwise excellent study book. The problem with such an affirmation is that it is undoubtedly true. A scientist can deduce the stimulus strength just by looking at the firing pattern or frequency! So the latter (frequency) can be construed as being the code for the former (strength). But is that also true for the brain? Doe the brain use really frequency codes, or firing patterns as a code? Psychophysics data indicate that the strength of a stimulus is experienced as the intensity of a sensation. The question is whether the strength of our sensations is retained in our brain, and if it is, whether the mnemonic trace of this strength is as detailed as the sensation itself. That could of course indicate a form of coarse coding, but of coding nonetheless. Let us follow the path firing patterns take to see if we can answer this question. The intracellular space (and/or the inner and outer membranes) of a neuron can be made more positive (depolarizing), or more negative (hyper-polarizing), with adequate electrical stimuli. Apparently, the strength of negative electrical currents has another effect on the neurons than positive electrical currents. In the first case, "The size of the change simply mirrors the applied amount of current stimulus" (p.50). These so-called "passive membrane properties" are also observed when using small positive currents. The difference in effect between (high) negative and (high) positive currents is certainly interesting, but remains unexplained. At least, the authors do not offer any explanation for this phenomenon, so that I can only, for now, register this mystery. It takes positive current to depolarize (make more positive) the cell and create an action potential. I think we should stop a moment and ponder this fact. An action potential means that whatever is happening in the brain does not stop with the current neuron we are studying, but is being propagated, in whatever form, to other neurons. The absence of an action potential in a hyper-polarized state certainly does not mean that nothing is happening to the organism, and that there are, therefore, no sensations involved. This consideration puts the functional distinction between On and Off neurons in a completely different light (no pun intended). Some authors, following Kuffler (1938), interpret the difference as that between light and dark, usually black (Peter Gouras thinks it is rather Blue, see his chapter in Helga Kolb's site Webvision). I think that both views indicate a rather naive interpretation of an apparent dichotomy which, let us not forget, is based on the concept of receptive fields. This is of course only in the cases where Off neurons are identified with a hyper-polarized state, not when they do produce an action potential. In the latter case I will give an everyday example to show that the dichotomy probably does not hold either. Imagine entering a not very brightly lit room (maybe the curtains are drawn, or it is a cloudy day, or dusk is approaching, or all of the above), which prompts you to turn on the lights. You then decide that it is too bright, and turn them back off. Your retinal receptors and neurons will certainly react to the change in illumination in both cases, but will that turn them into On and Off neurons? Or will it just show the reactions of the same neurons, once to the lights turned on, then to the lights turned off? The fact that there are no apparent differences between different types of neurons (On, Off, reacting to different features, etc.), seems to reinforce this view.
The idea that only reactions that produce an action potential are worth further study is illustrated by the concept of threshold. As Levitan and Kaczmarek put it:"The threshold is essential to ensure that small, random depolarizations of the membrane do not generate action potentials." They continue brazenly: "Only stimuli of sufficient importance (reflected by their larger amplitude) result in information transfer via action potentials in the axon." p.53 The threshold is, just like firing frequency, something that scientists can determine in an objective manner. It is therefore tempting to attribute to it, as inherent properties, those that are useful to us. It remains a question whether such a concept makes any sense when considering the brain as such. It could as well be the case that different amplitudes mean different inner experiences, with some pouring over their border as it were, while others just die out with the stimuli that triggered them.
Refractory Period Let us keep following the authors in the quest for neural coding: "Let us now examine the way the refractory period contribute to neuronal information coding". p.55 The concept of refractory period is very easily understood if you compare it with the need of recovering your breath after shouting. The faster you shout, the faster you will have to recover your breath. Up to a point, because however fast you would like to shout, there is a limit that you cannot ignore. Imagine now that it is the neuron doing the shouting in reaction to the electrical torture you are subjecting it. Which message do you suppose you are sending to the neuron, besides the strength of the electrical stimulus itself? What kind of message would the neuron need to encode in such a situation? The strength of the stimulus? It is already reacting to it by shouting accordingly. This is not the conclusion our authors think they can draw from the study of the refractory period: "Thus, even though action potential amplitude obeys the all-or-none law [The curve of action potential remains the same for every stimulus above threshold] and does not reflect stimulus intensity, the phenomena of threshold latency, and refractory period do indeed allow the encoding of stimulus intensity as a frequency code in the action." p.56. (original emphasis) This, once again, only proves that we, as external observers, can reconstruct the intensity of a stimulus from the frequency of the firing in the axon. But to be a neural code, it must be usable, and used, by the target neuron. It is evident that the target neuron will react differently to a single electrical burst, than to a continuous current. But how could that be construed as the reaction to a code instead of a direct reaction to a stimulus? Besides, we are the one sending the message, and it is not a very complicated message. We turn the power up if we want a higher current to flow through the neuron, and down if we don't. No other kinds of messages can be sent to a neuron, since neurotransmitters do nothing but make such a stimulation possible. Let us stand still by this point: all the possible messages we can send a neuron are fluctuations between a minimum and a maximum electrical current, and for these fluctuations to become a code, neurons must react differently to different fluctuations. But the only difference that we have been able to observe are different reactions to - sub or supra threshold, - speed or firing rate, in the limited sense that to every release of neurotransmitters corresponds a spike in the firing rate. All neurons react the same general way to a release of neurotransmitters or electrical stimulation (with a corresponding action potential). Unless we could refine the reactions of neurons to the point where we would find that some neurons react to, say, 5 spikes, but not to 3 or 7, while others do exactly the opposite, we must conclude that the frequency of firing is neither a message (unless we understand with message a mere stimulus) nor a code, and that, if it is a code, then it is a code that the brain has already cracked, since a higher stimulus gives rise to an intenser sensation.
Accommodation, habituation, and other mental phenomena We all have heard of the difference in reaction to different stimuli when considering behavior in general, but is particularly edifying when linked with the intrinsic properties of neurons, or what are thought to be such properties. Stimulating lightly a sea snail like Aplysia, will provoke first a withdrawal reaction, but if we keep it up, authors in general, and not only Levitan and Kaczmarek, suddenly change their language and revert to psycho-babble. They call it habituation, which comes not from the English word habit, but more from the French adjective habitue [with an accent on the final e], meaning that the snail has gotten used to the stimulation, and does not feel threatened by it anymore. But such a hybrid concept hides the fact that we have left the field of chemistry and entered the field of (animal) psychology. It tells nothing new about the neurons involved, and all about the way the animal reacts to different situations.
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