# Comments on Bolles

**URL:** <http://discourse.iapct.org/t/comments-on-bolles/2688>\
**Category:** CSG1995\
**Created:** [September 16, 1995, 2:23am UTC](http://discourse.iapct.org/t/comments-on-bolles/2688 "1995-09-16T02:23:26Z")\
**Posts on this page:** 2\
**Page:** 1

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**Author:** ![Bruce\_Abbott](http://discourse.iapct.org/letter_avatar_proxy/v4/letter/b/ba8739/32.png) [@Bruce\_Abbott](http://discourse.iapct.org/u/Bruce_Abbott)\
**Post date:** [September 16, 1995, 2:23am UTC](http://discourse.iapct.org/t/comments-on-bolles/2688/1 "1995-09-16T02:23:26Z")

</div>

[From Bruce Abbott (950915.2125 EST)]

My thanks to (in order of receipt) Avery Andrews, Rick Marken, and Bill  
Powers for their comments on the Bolles quote on regulation I posted  
yesterday (950914.1840 EST). One thing I've been trying to do lately is  
find out what happened to the intense interest in regulatory (control)  
models involved in feeding, drinking, and other activities which seemed  
aimed at controlling some physiological quantity or quantities such as  
cellular hydration, blood glucose level, or body weight, which was in  
evidence in the late 70's. After a brief "heyday," this approach seems to  
have withered under intense criticism levied against it, such as Bolles  
expresses. I thought posting Bolles's critique would provide an opportunity  
to deal publicly with the issues Bolles (and others) have raised. In so  
doing I am neither espousing nor defending the position Bolles takes.  
However, given that this kind of reasoning seems to have led to the  
dismissal of control theory as an adequate model of physiologically-based  
motivation (hunger, thirst, etc.), it would seem essential that one be able  
to answer these criticisms when they are raised.

One of the difficulties Bolles (and others) apparently had with applying  
control theory seems to be that they expected the elements of the model to  
be represented somewhere in the body literally as diagrammed. If the block  
diagram shows a comparator, then one must be able to find the comparator,  
the mechanism in the body where the reference signal is physically  
subtracted from the perceptual signal to produce the error signal. If the  
diagram of the standard control system shows a proportional compensator able  
to oppose errors from both above and below, then the organism had better  
contain such a mechanism. If there is a set point in the system, there must  
be a physical signal, being generated by something, which represents the  
desired state of the controlled variable. When these physical entities were  
not discovered, this evidence was taken to repudiate the control-system model.

Bolles is not the only writer to complain about the problem. After  
describing temperature regulation by the gound squirrel (which forages in  
the desert sun until it gets hot and then dives into its burrow and lays  
flat on the cold floor until it looses the extra heat) and the bee (which  
regulates the hive temperature through a set of fairly complex interactions  
among individual bees), J. D. Davis in the same volume says ". . .  
although one might want to posit the existence of a set-point somewhere in  
the brain of the ground squirrel it is very difficult to imagine what a  
set-point for bee hive temperature would be like, where it would be located,  
or how it would work." D. A. Booth, after outlining his marvelously  
detailed Mark 3 computer model of feeding, has the following to say under  
the subtitle "Regulation - a Pernicious Concept":

&nbsp;&nbsp;&nbsp;&nbsp;There is the seductive convenience of comparison between thermostats  
&nbsp;&nbsp;&nbsp;&nbsp;which everyone knows about and homeostats which we hope to learn about.  
&nbsp;&nbsp;&nbsp;&nbsp;In control theory, the value of the set-point is not identical to the  
&nbsp;&nbsp;&nbsp;&nbsp;value of the controlled variable when the system has steadied, but it  
&nbsp;&nbsp;&nbsp;&nbsp;can be made similar; thus it is easy to skip the distinction and read  
&nbsp;&nbsp;&nbsp;&nbsp;the value acheived by homeostasis into the setting of the homeostat.  
&nbsp;&nbsp;&nbsp;&nbsp;Most serious of all, the set-point formalism can be interpreted as a  
&nbsp;&nbsp;&nbsp;&nbsp;physical mechanism and then what may be equilibration between body,  
&nbsp;&nbsp;&nbsp;&nbsp;environment and brain is attributed to a mechanistic marvel within the  
&nbsp;&nbsp;&nbsp;&nbsp;brain. Some have claimed to resist this temptation and to intend their  
&nbsp;&nbsp;&nbsp;&nbsp;invocation of regulatory set-points merely as description of the system's  
&nbsp;&nbsp;&nbsp;&nbsp;performance characteristics. However, as equilibrium or unreferenced  
&nbsp;&nbsp;&nbsp;&nbsp;feedback control on the one hand and regulation on the other hand are  
&nbsp;&nbsp;&nbsp;&nbsp;intertranslatable formalisms for describing performance, the only point  
&nbsp;&nbsp;&nbsp;&nbsp;in invoking set-point regulation would appear to be the implication that  
&nbsp;&nbsp;&nbsp;&nbsp;a physical monitor-comparitor mechanism exists which explians the  
&nbsp;&nbsp;&nbsp;&nbsp;observed stabilising achievements. However, such inference is  
&nbsp;&nbsp;&nbsp;&nbsp;fallacious. The postulate is arbitrary unless and until independent  
&nbsp;&nbsp;&nbsp;&nbsp;evidence for a physical comparator mechanism is obtained by anatomical  
&nbsp;&nbsp;&nbsp;&nbsp;observation and physiological or biochemical measurement at the cellular  
&nbsp;&nbsp;&nbsp;&nbsp;level in the relevant location(s).

&nbsp;&nbsp;&nbsp;&nbsp;Thus, the result of this definition of regulation (even though not its  
&nbsp;&nbsp;&nbsp;&nbsp;logical requirement) is that stabilities and their defense that can be  
&nbsp;&nbsp;&nbsp;&nbsp;generated by equilibrium processes which are abundantly evidenced are  
&nbsp;&nbsp;&nbsp;&nbsp;being "explained" by physical mechanisms of much greater complexity than  
&nbsp;&nbsp;&nbsp;&nbsp;need to be invoked and are of a type for which there remains no direct  
&nbsp;&nbsp;&nbsp;&nbsp;physiological evidence in the whole of biology (not even for  
&nbsp;&nbsp;&nbsp;&nbsp;thermoregulation, or for metabolic biochemistry or genetic expression).  
&nbsp;&nbsp;&nbsp;&nbsp;The genes, with or without environmental aid, would have to specify a  
&nbsp;&nbsp;&nbsp;&nbsp;cellular arrangement to produce an extremely precise and unvarying signal  
&nbsp;&nbsp;&nbsp;&nbsp;of a particular strength throughout an animal's life; this would be the  
&nbsp;&nbsp;&nbsp;&nbsp;set-point. A monitor has to produce a graded signal of comparable  
&nbsp;&nbsp;&nbsp;&nbsp;accuracy and a comparator has to subtract signals and emit another graded  
&nbsp;&nbsp;&nbsp;&nbsp;signal which is converted into physiological or behavioral activity.  
&nbsp;&nbsp;&nbsp;&nbsp;Mechanical and electronic engineers find such material systems easy to  
&nbsp;&nbsp;&nbsp;&nbsp;construct but living matter does not have the properties that they exploit.

Whether this argument is well framed or not, it appears to have been  
excepted among the majority of researchers in this area. I am reminded of  
our own little e. coli nutrient-control model in comparison to Koshland's  
description of the physical arrangement actually found. In real e. coli,  
apparently, the rate of change of nutrient concentration as the organism  
swims about simply alters the rate at which tumbles are initiated via some  
chemical mediator. All that is required is that a direct relationship be  
established between rate of nutrient concentration increase and time to next  
tumble. No reference signal, no comparator, no error signal. Just greater  
rates of increase lead to longer intervals between tumbles. Our  
control-system model of e. coli, however, contains an explicit reference  
signal, comparator, and error signal. Anyone looking into e. coli's system  
for the physical representations of these entities is not going to find them  
there; logically, the system merely acts AS IF they are. Perhaps that is  
the important thing.

Regards,

Bruce

---

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**Author:** ![William\_T\_Powers4](http://discourse.iapct.org/letter_avatar_proxy/v4/letter/w/977dab/32.png) [@William\_T\_Powers4](http://discourse.iapct.org/u/William_T_Powers4)\
**Post date:** [September 16, 1995, 4:53am UTC](http://discourse.iapct.org/t/comments-on-bolles/2688/2 "1995-09-16T04:53:23Z")

</div>

[From Bill Powers (950915.2140 MDT)]

Bruce Abbott (950915.2125 EST) --

Bruce, this is great work you're doing. All these things you're citing  
are exactly the reasons for which PCT has not been accepted.

&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;J. D. Davis in the same volume says ". . . although one might want  
&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;to posit the existence of a set-point somewhere in the brain of the  
&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;ground squirrel it is very difficult to imagine what a set-point  
&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;for bee hive temperature would be like, where it would be located,  
&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;or how it would work."

This manner of speaking uses words that say "I have difficulty  
understanding how ..." but which really mean "It is ridiculous to  
propose ...". Clearly, the idea of controlling a \_perception\_ of beehive  
temperature would solve Davis's problem, because that would tell us  
where the set point is located (inside the bee), and a little education  
in control theory would explain how it would work. But many people have  
taken Davis' attitude of "If I don't understand it, there can't be  
anything to it."

Booth has a similar problem:

&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;There is the seductive convenience of comparison between  
&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;thermostats which everyone knows about and homeostats which we hope  
&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;to learn about. In control theory, the value of the set-point is  
&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;not identical to the value of the controlled variable when the  
&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;system has steadied, but it can be made similar; thus it is easy to  
&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;skip the distinction and read the value acheived by homeostasis  
&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;into the setting of the homeostat.

This word "similar" implies "mere" similarity, hiding the fact that the  
actual temperature can be maintained within a fraction of a degree of  
the set point. If he had said "almost identical" the whole meaning of  
the sentence would have been changed. But that isn't Booth's only  
problem. He says

&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;The genes, with or without environmental aid, would have to specify  
&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;a cellular arrangement to produce an extremely precise and  
&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;unvarying signal of a particular strength throughout an animal's  
&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;life; this would be the set-point.

The term "extremely precise and unvarying signal of a given strength"  
reveals a misconception, because there is no "set point" that is  
invariant for the life of the organism or even for one whole day. A good  
antidote for this misconception is Myrsovski's \_Homeorhesis\_, in which  
he shows that (a) the set points of all known biological regulatory  
systems are variable, and (b) these set points can be proven to change  
by showing that the system defends different levels of the variable  
against disturbances at different times.

Perhaps someone influential in biology once misinterpreted a "set" point  
to mean a point that was fixed forever in some optimum state. It's that  
sort of pompous ignorance that I have been fighting for 40 years.

I do get hot under the collar at statements like this:

&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;A monitor has to produce a graded signal of comparable  
&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;accuracy and a comparator has to subtract signals and emit another  
&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;graded signal which is converted into physiological or behavioral  
&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;activity. Mechanical and electronic engineers find such material  
&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;systems easy to construct but living matter does not have the  
&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;properties that they exploit.

This is merely a pretense at expertise on a subject of which the author  
knows nothing. One of the main reasons for which control systems are  
used in engineering is that they do NOT require precise or stable  
components to achieve precise and stable results. One of the main  
reasons for which control theory is an ideal theory of real human  
systems is that it describes an organization that can work with the  
sorts of sloppy components found in living systems, yet achieve quite  
accurate results. It is the theories that do NOT use control principles  
that run into problems of precision and stability.

What elevates the temperature of my neck protector is these authors'  
totally unwarranted claim to understanding. The anti-control-theory  
literature in the behavioral sciences is full of this sort of thing:  
announcements of how control systems work and what they can or can't do  
that are based on a few minutes of reasoning and an abysmal lack of  
actual knowledge. This is how all the myths about control theory have  
got started; some amateur thinks that he's got it all figured out, but  
never having studied the subject he makes all the elementary mistakes  
that any sophomore undergraduate engineering student gets cured of in  
the first six weeks of an introductory course. Unfortunately, the people  
who make these mistakes have big names in their fields, so those who  
read their words naturally assume they know what they're talking about.  
So the myths just spread and spread.

&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;I am reminded of our own little e. coli nutrient-control model in  
&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;comparison to Koshland's description of the physical arrangement  
&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;actually found. In real e. coli, apparently, the rate of change of  
&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;nutrient concentration as the organism swims about simply alters  
&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;the rate at which tumbles are initiated via some chemical mediator.  
&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;All that is required is that a direct relationship be established  
&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;between rate of nutrient concentration increase and time to next  
&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;tumble. No reference signal, no comparator, no error signal. Just  
&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;greater rates of increase lead to longer intervals between tumbles.

Actually, if you read Koshland carefully, you'll see that there is  
evidence for variable reference signals. There are "mutants" which  
always tumble, and others that never tumble. A sufficiently rapid change  
in the concentration, however, can make the always-tumblers stop  
tumbling or (in the other direction) the never-tumblers start tumbling.  
Among normal bacteria, there are different biases on the rate of change  
of concentration that will begin to suppress tumbling, and of course any  
such bias is evidence of a reference signal, and by implication some  
comparison mechanism. The operational definition of a reference level,  
which is quite independent of the block diagram by which you choose to  
model the system, is "that level of input at which the output just  
becomes zero." There is certainly such a level of rate of change of  
concentration that can be determined for any individual bacterium. And  
any way you choose to explain this effect is exactly equivalent,  
mathematically, to the standard PCT diagram.

&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;Our control-system model of e. coli, however, contains an explicit  
&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;reference signal, comparator, and error signal. Anyone looking  
&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;into e. coli's system for the physical representations of these  
&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;entities is not going to find them there; logically, the system  
&nbsp;&nbsp;&nbsp;&nbsp;&nbsp;merely acts AS IF they are. Perhaps that is the important thing.

The important thing is not the particular block diagram, but the  
relationships observed among the variables. What we see is that

Output = k \* (RefLevel - Input),

Where RefLevel is simply that magnitude of input at which Output = 0. If  
the Output becomes zero as Input approaches 10 units, then Reflevel is  
10 and the equation is

Output = k \* (10 - Input).

Then by observing how the Output changes as Input departs from the value  
of 10 units, you can determine the value of k. This is all purely  
empirical, no theory involved.

There is absolutely nothing about this analysis that depends on the  
particular way you propose that the nervous system creates this  
relationship. We use the simple PCT diagram because it shows all the  
basic operations needed to create such a relationship, and its parts  
correspond to known components of some actual physiological systems. The  
Input Function converts the physical variable outside the system into an  
internal signal; the comparator implements the subtraction; the output  
function implements the amplification and conversion of the difference  
into a corresponding amount of output. Any two adjacent operations or  
all three of them could be combined in a single physical element of the  
system. A single neuron can function as a complete control system.

Koshland was actually quite interested in the 'signaling system', which  
is the Input Function part of the standard model. Once you have split  
out the input processes, you are left only with the comparison and  
output processes to explain, and you can model them separately or as a  
combined function -- until and unless further research enables you to  
show that these two functions are carried out by different means in  
different places.

This equation and the definition of RefLevel do not depend on the block  
diagram. The PCT diagram is simply one straightforward way of  
implementing the equation to produce the same input-output relationship.  
All of the principles of control can be expressed entirely in terms of  
observable variables, without ever trying to guess how the nervous  
system is internally organized. But we -- or at least I -- want to try  
to understand how the nervous system is organized, which is why I keep  
looking for physical evidence about the actual wiring, and keep  
proposing possible block diagrams.

Also, a block diagram communicates relationships which are hard for  
people who are not comfortable with simultaneous equations to  
understand. The other equation in the simultaneous pair is, of course,  
the equation that describes how the Input depends on the Output via the  
environment. The diagram of a control loop makes the solution of the  
simultaneous equations obvious, or at least more obvious.

> **···**
>
> -----------------------------------------------------------------------  
> Best to all,
> 
> Bill P.
