[From Bill Powers (2012.09.29.0830 MDT)]
Rick Marken (2012.09.28.1100) --
I have added noise to my simulations and found that it had more effect
on the input than on the output, but really not too much effect on
either.. As you demonstrated to me (or, actually, had me demonstrate
to myself) an integral control system acts like a linear low-pass
filter, smoothing the output (and the input -- controlled variable)
compared to what would result if the same noise were added to an
equivalent open loop (and unfiltered) system.
BP: Most human control systems have leaky integrator outputs; the leak reduces the high-pass filtering. Also, if you add low-frequency noise (say 1 Hz bandwidth) to the comparator or input function, so it is in the same range as the bandwidth of good control, you will see even more effect on the output. The latter would be like driving in a gusty crosswind or steering a small motorboat through choppy seas, or an airplane through turbulent air. My first student takeoff in an airplane was through extremely gusty air and I was working very hard to keep the wings level (you've heard this story, I think). My instructor Ernie finally asked, "Bill, what are you doing?". I explained that the turbulence was pretty bad. "No, it isn't," he said. "Relax." I relaxed and the airplane steadied right down. Was that too much gain or not enough dead zone? Or as Kent McClelland has suggested, a nonlinear blend of the two?
RM: I can see that a narrow dead zone might be necessary for a high gain proportional control system but it seems like it wouldn't improve things much for an
integral control system.
BP: You're right in general, except for the effects of low-frequency disturbances.
RM: I guess I can test this using simulation but
just for the sake of simplicity it seems like the filtering
characteristics of closed loop (integral) control would have made it
unnecessary for the neurons to have developed a dead zone
characteristic.
BP: That all depends on the bandwidth of noise or disturbances.
RM: Is there any evidence that neurons do have a dead zone
(in terms of firing rate)?
BP: Yes. All neurons do. A single excitatory impulse entering a synapse raises the membrane potential by a small amount partway toward the threshold for producing an action potential, but in a few milliseconds it has decayed back toward the resting potential being maintained by the ion pumps in the membrane. At a very low rate of incoming impulses, that is all that happens for each incoming impulse and the neuron never fires. As the rate of incoming impulses increases, the decay back to resting potential becomes incomplete and the peak changes in potential start accumulating and becoming larger, until finally the peaks just cross the threshold potential for generating an output impulse or action potential from the neuron. The neuron then begins firing at a low rate which increases as the input rate increases still further. Each time the neuron fires, the membrane potential is reset to a low value (high negative value) and has to build up again over some number of input impulses (if they occur fast enough) until it reaches the action potential threshold again. That is why the output rate of firing is zero until the incoming rate rises above the dead zone.
Some "electrical" neurons have a resting potential that is not far below the threshold for firing of the neuron. In these, a single input impulse can produce an output impulse. These neurons do not have a dead zone. They are not common.
Some neurotransmitters, like dopamine, affect the sensitivity of synapses to other neurotransmitters. This effectively changes the gain at the synapse. Henry Yin has been studying this effect for some time, in connection with the basal ganglia and their role in Parkinson's disease. Chloride ions released by other neurotransmitters subtract fron the postsynaptic potential (or drive it more negative), increasing the size of the dead zone for all excitatory inputs. So both gain and dead zone are adjustable by neural signals just about everywhere in the brain.
An adjustable dead zone is a result of the inhibitory effect of some inputs. Inhibitory signals make the membrane potential even more negative than the resting potential. When the neuron is firing at some rate, the inhibitory input impulses just subtract from that rate. But since rates can't be less than zero, enough inhibition creates a dead zone that has to be overcome by excitation before further increases in excitation can cause any output impulses at all.
Best,
Bill P.