A motor control study supporting hierarchical control?

[Martin Taylor 2013.02.01.23.08]

[From Bill Powers (2013.02.01.1635 MST)]

  Adam Matic 2013.1.31.2100 cet 00

Rick Marken (2013.01.31.1100)

[…]
So my conclusion is that this paper, that is presumably
“supportive” of PCT is, at best, irrelevant to PCT and, at
worst, completely misleading about the nature of purposeful
(control)
behavior. In my estimation this paper is worth less than
nothing.

    AM:

    After the tone of your previous post, I expected something like

this, so
I went over the paper a few more times and… yeah… they don’t
look like
they understand control theory let alone PCT. There are really
no control
systems modeled, of any kind.

  BP: I ran across a paper about mice or rats using vibrissae

(movable
whiskers) to sense the positions of things – didn’t save it, but
I think
the Saig et. al, paper is specifically about that kind of sensing.
The
two arms sweeping the hands back and forth until they touch
something are
like what the whiskers do.

  I concur with your judgement, and Rick's, about the relevance this

paper
to PCT. Zero, or perhaps a little less.

I don't know if the paper I attach is the paper to which you refer,

but it uses the mouse vibrissae. It’s one I had intended to bring up
in this forum anyway, for two reasons. Firstly, from time to time
questions have been raised about the computational abilities of
neurons, and this paper seems to show that individual dendrites do
their own computations, and secondly because the HPCT structure
shows only one place in which sensory and motor “input” come
together, and that is the comparator, where input comes from the
higher level outputs in the form of a reference signal. Could this
kind of dendritic computation be some element of comparator
function? It’s not easy for me to see how, but if that’s not it,
then there must be another place in some revised HPCT circuitry
where motor and sensory inputs are used in coordination.

Here's the abstract. The paper is attached.

Martin

--------------Nature 13 Decembeer 2012, 492, 247-251---------
  • Ning-long Xu,

  • Mark T. Harnett,

  • Stephen R. Williams,

  • Daniel Huber,

  • Daniel H. O’Connor,

  • Karel Svoboda

  • & Jeffrey C. Magee
    Active dendrites provide neurons with powerful processing
    capabilities. However, little is known about the role of
    neuronal dendrites in behaviourally related circuit
    computations. Here we report that a novel global dendritic
    nonlinearity is involved in the integration of sensory and
    motor information within layer 5 pyramidal neurons during
    an active sensing behaviour. Layer 5 pyramidal neurons
    possess elaborate dendritic arborizations that receive
    functionally distinct inputs, each targeted to spatially
    separate regions1, 2 . At the
    cellular level, coincident input from these segregated
    pathways initiates regenerative dendritic electrical
    events that produce bursts of action potential output3, 4 and circuits featuring
    this powerful dendritic nonlinearity can implement
    computations based on input correlation5 . To
    examine this in vivo we recorded dendritic
    activity in layer 5 pyramidal neurons in the barrel cortex
    using two-photon calcium imaging in mice performing an
    object-localization task. Large-amplitude, global calcium
    signals were observed throughout the apical tuft dendrites
    when active touch occurred at particular object locations
    or whisker angles. Such global calcium signals are
    produced by dendritic plateau potentials that require both
    vibrissal sensory input and primary motor cortex activity.
    These data provide direct evidence of nonlinear dendritic
    processing of correlated sensory and motor information in
    the mammalian neocortex during active sensation.

    Nonlinear dendritic integration of sensory and motor input during an
    active sensing task

dendriticIntegration.pdf (856 KB)

···

[Martin Taylor 2013.02.02.09.58]

Did others get two copies of [Martin Taylor 2013.02.01.23.08]? If so, I apologise. I don't know how it happened, since as far as I know, I sent it only once. If not, the problem must be local.

Martin

[From Rick Marken (2013.02.02.1200)]

Martin Taylor (2013.02.01.23.08)

MT: I don't know if the paper I attach is the paper to which you [BP] refer,

but it uses the mouse vibrissae. It’s one I had intended to bring up
in this forum anyway, for two reasons. Firstly, from time to time
questions have been raised about the computational abilities of
neurons, and this paper seems to show that individual dendrites do
their own computations, and secondly because the HPCT structure
shows only one place in which sensory and motor “input” come
together, and that is the comparator, where input comes from the
higher level outputs in the form of a reference signal. Could this
kind of dendritic computation be some element of comparator
function? It’s not easy for me to see how, but if that’s not it,
then there must be another place in some revised HPCT circuitry
where motor and sensory inputs are used in coordination.

Here's the abstract. The paper is attached.

RM: Thanks for the paper, Martin. I don’t think can understand it but I’ll give it a try.

But I am puzzled by your question about the neural basis of the comparator. I think there is pretty strong neural evidence that supports the control model of the operation of a comparator, which computes the difference between sensory and reference input… The evidence is that motor (corresponding to reference) and sensory (corresponding to perceptual) neurons often synapse with the same motor (o) neuron in opposite ways; one inhibitory and the other excitatory, so that the net level of output (neural firing rate) from the o neuron is proportional to the difference in the level of excitatory and inhibitory input to the neuron. So if r, p and o are neural signals measured in terms of firing rate, then the combination of excitatory and inhibitory neural signals synapsing at the cell body of an output neural would be approximately o = r - p. Of course, there will be non-linearities in this relationship when it’s implemented in neurally. It might be interesting to see what the behavioral implications of this non-linearity might be. I don’t know of any studies related to behavioral implications of the possible non-linearity of the comparator function, although the universal error curve (http://www.mindreadings.com/ControlDemo/ErrorCurve.html) could be considered one. This might be a fertile area for the kind of research you seem to like.

Best

Rick

···


Richard S. Marken PhD
rsmarken@gmail.com
www.mindreadings.com

[Martin Taylor 2013.02.02.16.28]

[From Rick Marken (2013.02.02.1200)]

But I am puzzled by your question about the neural basis of the comparator.

I had no such question. I had a question about what in the HPCT model corresponds to the computation within a single dendrite that includes input both from sensory and from motor systems. The only place in HPCT that this happens is the comparator, but the nature of the dendritic computation doesn't seem to me to be what one would expect of a comparator.

Martin

[From Rick Marken (2013.02.02.1450)]

Martin Taylor (2013.02.02.16.28)–

RM: But I am puzzled by your question about the neural basis of the comparator.

MT: I had no such question. I had a question about what in the HPCT model corresponds to the computation within a single dendrite that includes input both from sensory and from motor systems. The only place in HPCT that this happens is the comparator, but the nature of the dendritic computation doesn’t seem to me to be what one would expect of a comparator.

RM: Well that’s much easier to answer. The answer is… “Nothing”.

Best

Rick

···


Richard S. Marken PhD
rsmarken@gmail.com

www.mindreadings.com

[From Adam Matic 2013.2.3.0010 cet]

Wow. This one is hard to understand. I cound barely visualise what the title is about.

Adam

[From Bill Powers (2012.02.03.0845 MST)]

Martin Taylor 2013.02.01.23.08

I don’t know if the paper I
attach is the paper to which you refer, but it uses the mouse vibrissae.
It’s one I had intended to bring up in this forum anyway, for two
reasons. Firstly, from time to time questions have been raised about the
computational abilities of neurons, and this paper seems to show that
individual dendrites do their own computations, and secondly because the
HPCT structure shows only one place in which sensory and motor
“input” come together, and that is the comparator, where input
comes from the higher level outputs in the form of a reference signal.
Could this kind of dendritic computation be some element of comparator
function? It’s not easy for me to see how, but if that’s not it, then
there must be another place in some revised HPCT circuitry where motor
and sensory inputs are used in coordination.

Yes, that’s the paper I saw in Nature, to which I subscribe.
What do you mean by a “motor input?” More to the point, what
did the authors mean? I think they were talking about an input that was
produced by a motor action, namely, moving the arms so contact was
made with an object (and, of course, sensed). In PCT, that input,
therefore, qualifies as either a controlled variable or a variable
associated with one. Aside from that, it’s just another perception like
all the others we control, important and unimportant. I’d say it’s a
lower-order controlled variable, with a copy being sent to a higher-order
control system.
I’m getting tired of the way so many neuroscientists puff up their
knowledge when all they have are some vague guesses. What is
“powerful” about the “nonlinear” computations carried
out in the dendrites? That would be easier to judge if the authors had
told us what those computations are. As it is, I don’t think they have
any clue about those computations, or about analog computations in
general. They’re just throwing important-sounding words around. I am
unimpressed. I think we will eventually find that the computations are
pretty simple.
I would make guesses similar to the authors’ about how mice use vibrissae
to detect nearby objects. It doesn’t surprise me at all to see (if it’s
true) that the position perception is computed from both the sensed
vibrissa angle and the sense of contact – though it seems more likely to
be a direction perception, since distance of the point of contact along
the whisker can’t be detected at least not by that means. And of course
it’s no surprise to read that the computations are carried out in the
dendrites, though I don’t quite see how the authors rule out the cell
body or the axon hillock as the place where signal differences are
computed. I would suspect that the hillock would be where signals add and
subtract to produce the net depolarization that represents the spatial
relationship. It is, however, also known that interactions take place in
the dendrites, as for the effect of dopamine on the synaptic sensitivity
to other signals.
In fact, little about the details of signal interactions in neurons
surprises me. I don’t mean I know what they all all (far from it), but
simply that there are many possibilities and I don’t think any of them
are ruled out. Everything the nervous system does has to be accomplished
somehow. In stretch receptors embedded in muscles, for example,
there are little muscles at ends of each spindle which are innervated by
gamma efferents from elsewhere. When these muscles are active, the
central part of the spindle is stretched and sensory nerves wrapped
around that part generates signals that activate motor neurons. The
spindle is also stretched when the muscle as a whole contracts or
shortens for any reason, and that produces the same kind of signal from
the annulospiral sensory nerves. When you work out all the relationships
and the loops of which the spindles are part, it becomes evident that the
spindles are comparators, and the signals they send to the motor neurons
are error signals in the “stretch reflex”, which is really a
control system one level above the Golgi tendon reflex.

The stretch-reflex-controller is a Rube Goldberg device retained because
it works, and its comparator is a mish-mash of electronic, chemical, and
mechanical functions. I wouldn’t have designed it that way, but that’s
the way it evolved. So we just have to accept it and not get too excited
about it.

My negative reaction to this article comes simply from the hype. There
are some useful facts in it, and if anyone wants to know how the
vibrissae are used to detect objects this could prove somewhat helpful.
But the facts are presented as if they were understood, which is not true
– they are just there, undigested. Yuk.

Best,

Bill P.

[From Rick Marken (2013.02.06.1000)]

Bill Powers (2012.02.03.0845 MST)–

RM: Hope the move went (or is going) well.

BP: What do you mean by a “motor input?” More to the point, what
did the authors mean?

RM: My impression from both of the neurophysiology papers I’ve read is that “motor input” is presumed to be another sensory input that comes from a motor command. When these folks talk about the effect of motor output on perception they, I believe,are not talking about the ones we talk about in PCT: either a reference input from a higher level control system of the effect on a perceptual variable, such as the visual perception of the position of you hand, that can be varied by varying the motor inputs to the muscles of your arm. I think they are talking about a direct effect of motor neural signals on a perceptual (sensory) signal. The give away is that they talk about motor inputs as being involved in the “creation” of perceptions. This is like the “motor theory” of speech perception; the idea that what is perceive is the neural signals that “command” the the muscle movements that produce the speech sounds.

BP: My negative reaction to this article comes simply from the hype.

RM: Mine comes from that as well as the poorly thought out model of perception.

Best

Rick

···


Richard S. Marken PhD
rsmarken@gmail.com
www.mindreadings.com

[From Chad Green (2013.02.07.1139 EST)]

RM: Well that's much easier to answer. The answer is... "Nothing".

CG: Rick, are you familiar with Quine's notion of confirmation holism? The authors of this study could very well say the same thing about PCT.

Best,
Chad

Chad Green, PMP
Program Analyst
Loudoun County Public Schools
21000 Education Court
Ashburn, VA 20148
Voice: 571-252-1486
Fax: 571-252-1633

"If you want sense, you'll have to make it yourself." - Norton Juster

Richard Marken <rsmarken@GMAIL.COM> 2/2/2013 5:49 PM >>>

[From Rick Marken (2013.02.02.1450)]

Martin Taylor (2013.02.02.16.28)--

RM: But I am puzzled by your question about the neural basis of the

comparator.

MT: I had no such question. I had a question about what in the HPCT model
corresponds to the computation within a single dendrite that includes input
both from sensory and from motor systems. The only place in HPCT that this
happens is the comparator, but the nature of the dendritic computation
doesn't seem to me to be what one would expect of a comparator.

RM: Well that's much easier to answer. The answer is... "Nothing".

Best

Rick

···

--
Richard S. Marken PhD
rsmarken@gmail.com
www.mindreadings.com

[From Rick Marken (2013.02.07.1325)]

Chad Green (2013.02.07.1139 EST)–

RM: Well that’s much easier to answer. The answer is… “Nothing”.

CG: Rick, are you familiar with Quine’s notion of confirmation holism? The authors of this study could very well say the same thing about PCT.

RM: I was not clear when I answered “Nothing”. It was in reply to Martin Taylor’s question:

MT: “… what in the HPCT model corresponds to the computation within a single dendrite that includes input both from sensory and from motor systems”.

RM: My answer to that was “Nothing”. But if by “input from motor system” Martin was referring to efferent neurons functioning as reference inputs to a control loop then my answer should have been “Comparator”.

No, I’ve never heard of Quine’s notion of confirmation holism. As far as the authors of the study conclude that PCT Has nothing to do with their work I’d be relieved. I’ve been reading too many papers lately which do refer to PCT as “compatible” with their theory when it really isn’t. It gievs me the creeps when that happens. Maybe Quine’s confirmatoin holism can explain why;-)

Best

Rick

···


Richard S. Marken PhD
rsmarken@gmail.com

www.mindreadings.com

[Martin Taylor 2013.02.08.12.38]

I had not realized that your "Nothing" answer was in response to my

question. I guess you must have the same question in your mind as I
do, then.
If I read the paper correctly, the observation is that the mouse has
cells whose dendrites receive input from the motor cortex and from
the sensory system, both being related to the whiskers. The motor
cortical signals are supposed to be “driving” the whisker
(presumably providing reference signals for lower-level control
units), while the sensory signals come from whatever it is at the
base of the whisker that senses forces on the whisker. When either
the motor signal or the sensory signal exists alone at the dendrite,
the cell doesn’t fire. When they are both there, the cell does fire.
Maybe I read it wrong, not being too well acquainted with
neurophysiology, but that is how I understand the observations, done
in a live mouse capable of voluntary action.
As I said, when I asked the question originally, firing only when
both sensory and motor signals are present doesn’t seem like the
action of a comparator, which is the only place in the strict HPCT
model where motor output and sensory input come together. So I asked
whether there might be some other place where this kind of
convergence would make sense. Now I understand that Rick said that
there isn’t and that this IS the way a comparator is expected to
perform, but I don’t believe him about the comparator. Bill’s approach to an answer was a bit different. To paraphrase, I
read Bill as saying that he didn’t understand what was meant by
“computation in a dendrite” and that the experimenters must have
been mistaken when they said that the signals to the dendrite came
from motor output and sensory input, because they were clearly both
sensory signals. Bill may well be correct, but if the experimenters
actually knew what they were doing and I read the paper correctly,
it is possible that he is wrong. So my question stands.
Part of the background to my question (and now I suppose Rick’s as
well), is that we know HPCT doesn’t cover everything, at least in
the form in which we customarily display it. In particular, there
are two areas we know to be important but that are not included in
the standard hierarchy (including the reorganization system). One is
the energy system (which is what fMRI observes), and the other is
gain control. I wondered whether there was any obviously reasonable
place in these systems where this kind of convergence might make
sense. Having thought about it a little, I think there might be a place for
it in a gain-changing system. If the animal is searching around for
a whisker touch (changing motor reference signals) and finds none,
there is no reason for anything outside the "control-of-searching’
part of the hierarchy to do anything. Likewise, if there is an
adventitious whisker-touch when the animal is not currently
controlling for anything associated with a whisker touch, all that
might happen would be within the classic HPCT hierarchy, if the
whisker-touch disturbed something the animal was controlling. However, if the animal is actively searching as part of controlling
for something that involves a whisker touch, then the coincidence of
motor output (reference signal) and sensory signal does make sense.
One way that it could be used might be to reduce the gain in the
part of the hierarchy that was performing the search, so that the
search stops and whatever was being controlled that needed the
whisker-touch could now be effectively controlled. That kind of
effect might be expected even if this paper didn’t exist.
It’s just a thought, but at the moment is seems to me that some such
explanation is needed, since I assume that the neurophysiological
team are competent, that my reading of their observations is
somewhere near correct, and that Rick’s “Nothing” answer is correct.
Martin

···

[From Rick Marken (2013.02.07.1325)]

      Chad Green

(2013.02.07.1139 EST)–

      RM: Well that's much easier to answer. The answer is...

“Nothing”.

      CG: Rick, are you familiar with Quine's notion of confirmation

holism? The authors of this study could very well say the
same thing about PCT.

      RM: I was not clear when I answered "Nothing".  It was in

reply to Martin Taylor’s question:

        MT:

“… what in the HPCT model corresponds to the computation
within a single dendrite that includes input both from
sensory and from motor systems”.

      RM: My answer to that was "Nothing". But if by "input from

motor system" Martin was referring to efferent neurons
functioning as reference inputs to a control loop then my
answer should have been “Comparator”.

[From Bill Powers (2013.02.12.2019 MST)]

Martin Taylor 2013.02.01.23.08

I don’t know if the paper I
attach is the paper to which you refer, but it uses the mouse vibrissae.
It’s one I had intended to bring up in this forum anyway, for two
reasons. Firstly, from time to time questions have been raised about the
computational abilities of neurons, and this paper seems to show that
individual dendrites do their own computations

BP: If you think of the computations in the analog computer context,
“doing computations” just means physical/chemical processes
occurring according to the applicable laws. If excitatory ions are
released by one neurotransmitter, and inhibitory ions by a different
neurotransmitter, the result is that the net depolarization of the cell
membrane corresponds to the difference between the concentrations of the
two ions, for positive net concentrations. Hence the rate at which the
cell body fires will be some function of the rates at which the two kinds
of ion are being released, which depends in turn on the frequency at
which neural impulses are arriving at the two kinds of synapse. That is
all there is to the “computation.” Of course computations are
performed by the neurons – they are simply the way a neuron works. There
is no separate computational machinery – just the physical
processes.

We see the computation in the way the output frequency from the cell
depends on the frequencies of inputs to the cell. That is what we try to
represent by writing mathematical functions. But the cell does not write
mathematical functions or solve equations: it simply acts out the
physical effects, while we try to approximate them on paper.

As to the “computational abilities” of neurons, they are
whatever can be done to turn input streams into output streams. If we
have addition and subtraction of postsynaptic potentials, modulation of
one potential by another (multiplication), accumulation of ion
concentrations (integration), and other basic signal interactions as
sketched in by chapter 2 of B:CP, we can expect to see input-output
functions of all the kinds we describe with differential equations. The
best way to understand this is to study a text on analog computing.
Analog computing is very different from symbolic computing.

This is what I would much prefer to be doing with my time instead of
agonizing over where and how I will live.

Besg,

Bill P.

[From Rick Marken (2013.02.12.2205)]

Bill Powers (2013.02.12.2019 MST)–

Thanks for this (and everything) Bill. Sorry about the problems with the move. Must be awful. But I look forward to seeing you back and comfortable in the old place soon!

Love

Rick

···

Martin Taylor 2013.02.01.23.08

I don’t know if the paper I
attach is the paper to which you refer, but it uses the mouse vibrissae.
It’s one I had intended to bring up in this forum anyway, for two
reasons. Firstly, from time to time questions have been raised about the
computational abilities of neurons, and this paper seems to show that
individual dendrites do their own computations

BP: If you think of the computations in the analog computer context,
“doing computations” just means physical/chemical processes
occurring according to the applicable laws. If excitatory ions are
released by one neurotransmitter, and inhibitory ions by a different
neurotransmitter, the result is that the net depolarization of the cell
membrane corresponds to the difference between the concentrations of the
two ions, for positive net concentrations. Hence the rate at which the
cell body fires will be some function of the rates at which the two kinds
of ion are being released, which depends in turn on the frequency at
which neural impulses are arriving at the two kinds of synapse. That is
all there is to the “computation.” Of course computations are
performed by the neurons – they are simply the way a neuron works. There
is no separate computational machinery – just the physical
processes.

We see the computation in the way the output frequency from the cell
depends on the frequencies of inputs to the cell. That is what we try to
represent by writing mathematical functions. But the cell does not write
mathematical functions or solve equations: it simply acts out the
physical effects, while we try to approximate them on paper.

As to the “computational abilities” of neurons, they are
whatever can be done to turn input streams into output streams. If we
have addition and subtraction of postsynaptic potentials, modulation of
one potential by another (multiplication), accumulation of ion
concentrations (integration), and other basic signal interactions as
sketched in by chapter 2 of B:CP, we can expect to see input-output
functions of all the kinds we describe with differential equations. The
best way to understand this is to study a text on analog computing.
Analog computing is very different from symbolic computing.

This is what I would much prefer to be doing with my time instead of
agonizing over where and how I will live.

Besg,

Bill P.


Richard S. Marken PhD
rsmarken@gmail.com
www.mindreadings.com

[Martin Taylor 2013.02.13.23.08]

Yes, indeed.

You may be amused by a small segment of an extended conversation I
was having a couple of weeks ago with someone who contacted me on
another matter but who had heard of PCT and wanted to know something
about it. In messages that preceded the section I will quote, I had
talked about e-coli control. Here’s part of his response and my
answer:
----quote----
Yes, that’s exactly what it is. The same is true when you talk about
humans, too. Bill Powers’ insight was simply that in order to
maintain internal stability, the organism, whether bacteria or
human, must act so that the effects of the environment on the
internal system are minimized. If the internal system is too salty,
move to where the environment is less salty. If your blood has too
much CO2, breathe. If your government is doing things that make you
uncomfortable, vote for the other guys. Not all actions are equally
effective, but they are (according to PCT) all performed in order to
stabilize some internal state that we call a perception.
—end quote----
Yes. Like you, I spent quite a bit of time wiring up op-amps and the
rest in my earlier days. I can’t remember the machine name, but at
one time we had quite a big hybrid computer that mixed analogue and
digital computation. Wolfram, as I understand it, would argue the
reverse of you, and say that the appearance of analogue computation
was the result of digital operations in the Universe. But who cares?
The result of the computation is what allows control to occur, not
the underlying technology.
Anyway, my interest in this thread was not particularly on whether
the computation was done in the dendrite, as the authors claim, but
in the claim that the requirement for cell firing was that both
motor output and sensory input co-exist, a property that seemed not
to conform to the operation of a comparator, but that might make
sense in other parts of the apparatus such as the energy system or
the control of gain in parts of the hierarchy.

···

[From Bill Powers (2013.02.12.2019 MST)]

    Martin Taylor

2013.02.01.23.08

    I don't know if the

paper I
attach is the paper to which you refer, but it uses the mouse
vibrissae.
It’s one I had intended to bring up in this forum anyway, for
two
reasons. Firstly, from time to time questions have been raised
about the
computational abilities of neurons, and this paper seems to show
that
individual dendrites do their own computations

  BP: If you think of the computations in the analog computer

context,
“doing computations” just means physical/chemical processes
occurring according to the applicable laws.

      When you talk about cell

membranes or the e-coli bacteria, I wonder how much of that
isn’t an effect of just the chemistry doing what it naturally
does.

  We see the computation in the way the output

frequency from the cell
depends on the frequencies of inputs to the cell. That is what we
try to
represent by writing mathematical functions. But the cell does not
write
mathematical functions or solve equations: it simply acts out the
physical effects, while we try to approximate them on paper.

  As to the "computational abilities" of neurons, they are

whatever can be done to turn input streams into output streams. If
we
have addition and subtraction of postsynaptic potentials,
modulation of
one potential by another (multiplication), accumulation of ion
concentrations (integration), and other basic signal interactions
as
sketched in by chapter 2 of B:CP, we can expect to see
input-output
functions of all the kinds we describe with differential
equations. The
best way to understand this is to study a text on analog
computing.
Analog computing is very different from symbolic computing.

  This is what I would much prefer to be doing with my time instead

of
agonizing over where and how I will live.

  Besg,



  Bill P.