Motor Integration in
the Central Nervous System
doctoral research was a study of the vocal behaviour of baby chicks, how they
twitter and peep as they run around together and with their mother hen. I would
put electrodes into their brains, send a little current that would make them call
(while they were anesthetized), and this work lent some interesting insight into
how motor behaviours are integrated in the nervous system. This work, conducted
from 1965 through 1970, is still fresh and cutting edge (I believe). There is
an introduction to it below, and/or you can consult the thesis itself at Sussex
University: "The Central Control of Vocalization in the Chick (of the domestic
fowl)"; the doctoral thesis of Norman Allan, 1971:
the cheeping of a chick:
of information and action
emerging out of the nervous system
I first encounter the
concept of non-linear oscillation
in the 1960s during research into
the neural organization
of an expressive behaviour -
to wit, the cherpings
of the chick.
for my sins I
tortured baby chicks
to see what makes them tick
see how they make their noises
I jolted their brains with electricity
and took them apart.
Physically, for the chick,
three things are involved in making a sound.
1 ~ it has to blow (expire
over a membrane which then vibrates.
this membrane is
called the syrinx;
2 ~ it has to maintain air-pressure
in the thoracic air-sac in which the syrinx
It does this by opening and closing a valve; and
3 ~ it modulates
the tension of the syrinx and hence
the pitch/tone of the called note
by the contraction on a pair of muscles
chick has the following repertoire of calls:
If you startle it
if you grab it, it trills (because it's startled)
it "shrieks"and then it peeps.
If it is (happily) interested in something
it does calls an excited
and if it is just tootleing along
it does another sort of twittering
if it is distressed it peeps
In its twittering
we see that it is calling portions of its
and we see that when it is happily interested
it calls the ascending
portions of the trill,
when it is distressed it calls the descending portion.
To do this the oscillating
motor out put to
the syringeal muscles and to the air-sac valve
be more or less matched for frequency
but must vary/shift in phase.
are occasions on which we can see
these two forms/signals manifest in the
It appears as if the air-valve.signal on occasion can
spill over into
the syringeal muscle signal.
this spill over is quite regularly seen
the excited tweet-tweet
So a continuum from approach/interest/(pleasure?)
through to avoidance/distress
is manifest in the chicks calls
as a shift in relative phase
the two cycles,
one driving the syringeal muscles
and another driving
We see a second continuum in the chick
When we evoke calls by electrically stimulating
area of the brain
(the Call Motor Area of the Optic Tectum overlaps,
but is not confined to, the Lateral Mesencephalic Nucleus)
get a series of calls that segregates into bouts
(this is in the anaesthetised animal)
and we see the peep emerge
the same peep we see in the unhappy chick
so here we see breath
and we see "quenching"
(as breath grows large the trill modulation
is blown out)
the repertoire maps out in two dimensions
(plus a single step seen in the startle:trill)
y) force of expiration / exertion
x) approach / avoid,
pleasure / distress, nice / nasty
z) rate of arousal (the
single step seen in the startle:trill)
I brought my data, the sound graphs,
to the mathematician biologist Brian
he said, "this looks like non-linear system."
shifts: there aren't many central nervous system phase shifts
in the literature.
The only one I'm familiar with originates around
the same era as my chick
call work, late 1960s:
Adey reports that when you are teaching a cat a task
during the learning a part of the hippocampus leads
(is phased in front of)
activity in the adjacent cortex
but after the task is learned it lags.
Judgement: does approach / avoid, pleasure
imply, represent, reflect
(here in our baby chick) manifest in a phase shift?
judgement a quality solely of mind?
don't machines make judgements and
In approach (pleasure) we
see a lifting, a rising tone
with avoidance, distress we see a falling, a
I am reminded of What's His Name,
"the baby watcher",
who speaks of "energy envelopes" (sic)
and how these give a qualitive tone to an event
is it building, bursting,
part of happening emotional tone come from its
reminded of Madame Blavatsky.
In my doctoral thesis I ventured (in a footnote)
that the chick's repertoire smacked of Madame B.
and her good and bad vibrations
see: Norman Allan's doctoral
"The Central Control of Vocalization
in the chick of
the Domestic Fowl."
Sussex University: 1971
4: neural nets