1 A philosophical introduction

Chapter Outline
1.1 Adequate language 1
1.2 Specific features of biological objects 2
Slow muscles produce forces that depend on muscle length and velocity 2
Confusing sensory receptors 3
Long conduction delays 3
Neurons—threshold elements 4

1.1 Adequate language

The Ancient Egyptians built pyramids, perfect geometrical structures. However, we
associate the birth of the science of geometry not with those anonymous Egyptian
architects but with the name of Euclid. Why? This name was not selected by pure
chance. Euclid did turn a set of rules that had been known for hundreds of years before
him into a science. This required making a very bold step: Introducing a definition for
an object that did not exist—a point. Euclid defined a point as an object that had no
length, no width, and no height (look around and you will find no such objects).
Despite the fact that points did not exist, this was a crucial step that later allowed for
the introduction of definitions for a line, a plane, and a sphere, and geometry became
a science. The step made by Euclid may be called “introducing an adequate language”
for spatial relations among objects. This language permitted questions to be formu-
lated in a way that allowed rigorous, exact scientific inquiry.
For any area of science, questions have to be formulated in an adequate language.
As of now, unfortunately, no such language has been introduced for biology.
Motor control is a relatively young field of research in biology. Let me define it as an
area of science exploring natural laws that define how the nervous system interacts with
other body parts and the environment to produce purposeful, coordinated movements. It
is very hard to look for an adequate set of notions in an area that does not have them, but
it is also very challenging and exciting! It is much more simple and tempting to borrow
one of the developed approaches from another field that shares “key words” with motor
control, for example basic mechanics, control theory, or engineering. One should keep
in mind, however, that such an approach has strict limitations. It can provide tools that
help find answers to questions after the questions have been formulated. As of now, the
main problems of motor control are not in finding answers but in formulating questions.

Fundamentals of Motor Control. DOI: 10.1016/B978-0-12-415956-3.00001-4

Copyright Ó 2012 Elsevier Inc. All rights reserved.
2 A philosophical introduction

To achieve a scientific, physical understanding of the system for movement

production, researchers have to use adequate methods. Using methods developed for
systems that have very little in common with the human body (for example, control
theory was developed for ballistic missiles and other human-built objects) may lead to
important progress in building artificial systems that imitate aspects of human
behavior, with substantial impact in such areas as prosthetics and robotics. But this is
not motor control as defined above. Such approaches cannot offer an adequate
formulation of questions in a field that differs from the area for which those
approaches have been developed.

1.2 Specific features of biological objects

Let us start with a few basic features of biological objects. Such objects belong to the
physical world and they are alive. So, help with formulating questions may be expected
to come from natural sciences such as physics, chemistry, and biology (physiology).
Consider the following major difference between living and inanimate (including
man-made) objects. The behavior of living objects is intentional, that is, typically it has
a purpose. This is a major difference from the inanimate world. If one knows the initial
state of an inanimate object and the forces acting on the object, its behavior can be
predicted using the laws of classical mechanics. For example, a stone never rolls uphill
unless an external force pushes it up. In contrast, animals can run uphill using forces that
they themselves bring about. Sophisticated man-made systems, for example cars, can
also move uphill, but they are designed by humans to behave in this way. So, we can view
them as extensions of biological objects for as long as they are able to perform their
programmed functions. A car that has run out of gas or had one of its major components
broken immediately starts to behave as a member of the class of inanimate objects.
The fact that biological objects are intentional does not mean that they violate the
laws of physics. Living beings differ from the inanimate in their ability to turn the
laws of physics to their advantage.
If we now turn to the system for movement production, there are a few features in
this system that would make an engineer think that it was designed by someone with
no knowledge of basic engineering (which is very likely true!). Certain features of the
neuro-sensory-motor system look, to put it mildly, suboptimal, particularly when
considered by an expert in designing artificial moving systems. We will consider basic
properties of the system for movement production in one of the next sections; here, let
me emphasize only a few important points.

Slow muscles produce forces that depend on muscle length and velocity
Consider our motors, the muscles. Motors built by engineers are powerful, quick, and
can produce pre-computed forces and/or moments of force. In contrast, muscles are
sluggish, not very powerful, and generate forces that depend on muscle length and its
rate of change (velocity). Let me focus on this last feature. It means that when you send
a neural command to a muscle, the force transmitted by its tendons to the points of their
A philosophical introduction 3

attachment to bones will depend not only on the magnitude of the command but also on
the actual muscle length and velocity. Since motion of body segments depends not only
on muscle forces but also on forces from the environment, actual muscle forces become
unpredictable, unless the central nervous system possesses an ability to predict changes
in the environment perfectly, with 100% confidence. The environment is typically
unpredictable (at least, not predictable perfectly). This means that the central nervous
system is in principle unable to prescribe time patterns of muscle forces, except in very
special laboratory conditions with perfectly reproducible external force fields (actually,
this is impossible even in the best laboratory environments). This feature of the system
for movement production was emphasized by Nikolai Bernstein, a great Russian
physiologist who is sometimes considered the father of contemporary motor control.

Confusing sensory receptors

To produce coordinated purposeful movements it is necessary for animals to get
information on the environment and on the current state of body segments. This
information is used for planning actions as well as for correcting ongoing move-
ments if they happen to be inaccurate or if a major perturbation acts on the body
from the environment. Our body is equipped with a large number of sensors that
supply such information. In particular, we have sensors of muscle length and
velocity (muscle spindles), of tendon force (Golgi tendon organs), of the state of
joints (articular receptors), and of pressure on and motion of the skin (cutaneous and
subcutaneous receptors). At first glance, these sensors are sufficient to provide all
the necessary information. However, when one starts to consider their design and
properties, a few apparent problems emerge. The sensors seem to be noisy and
information from them can be confusing, at least as compared to nearly perfect
sensors that measure the variables of interest in artificial moving systems.
For example, muscle spindles produce signals related to muscle length and
velocity, but their gain is modulated by a special system of neurons in the spinal cord,
the gamma-motoneurons. So, one and the same signal may be generated at different
positions and velocities depending on the gamma-motoneuronal activity.
Golgi tendon organs are accurate tendon force sensors. Note, however, that our joint
movements are rotations. This means that adequate mechanical variables for joint
motion are moments of force, not forces. One and the same tendon force produces
different moments of force with respect to a joint at different joint positions because the
moment arm of the force changes with joint position. So, information from Golgi
tendon organs is by itself insufficient to know moments of force. Articular receptors
look even less reliable in providing information about variables that seem to be most
relevant for joint action: Joint angle, its angular velocity, and moment of force. More
information on the features of different sensors will be presented in Chapter 3.

Long conduction delays

As already mentioned, unexpected changes in the environment happen rather
commonly. To avoid movement disruptions by such changes, commands to muscles
4 A philosophical introduction

have to be corrected appropriately. Such corrections are made based on sensory

signals reporting that something unexpected has happened. A major problem is that
conduction pathways in our bodies are relatively long (typical distances are on the
order of 1 m), and the speed of conduction is relatively low (at best, about 100 m/s,
but frequently much lower). As a result, delays of a few tens of milliseconds are
common even for the quickest motor reactions to peripheral signals, and such delays
can easily reach 0.1 s and more, if they are computed from a peripheral signal to
a useful motor reaction produced by muscles. This is a very long time for some of the
quickest voluntary movements. A top-level sprinter covers 1 m over this time, while
a well-served tennis ball flies over 5 m. So, the central nervous system always deals
with outdated information on the state of the peripheral organs, and this information
becomes even more outdated when signals generated by the neural controller reach
their target muscles.

Neurons—threshold elements
The main element of the central nervous system is the neural cell, the neuron. Indi-
vidual neurons and neural networks are commonly assumed to perform all the
necessary computations to allow the organism to function adequately. Neurons
possess a feature that looks like a source of major computational problems: They are
threshold elements. This means that small changes in the input signals into a neuron,
that is, changes in the potential on its membrane, do not lead to any changes in its
output until these changes reach a certain threshold value. Then, the neuron generates
a standard signal, an action potential, with characteristics that do not depend on the
strength of the input. So, if one knows the output of a neuron (a time series of its
action potentials), it is in principle impossible to reconstruct the input into the neuron.
For example, if someone turned on the light in a room, you would have no idea with
what force that person pressed on the switch, only that the force was sufficiently high.
In other words, if you want a neuron to generate a particular output signal, there are an
infinite number of combinations of input signals into the neuron that can achieve this
result. This feature of neurons creates an insurmountable problem for so-called
inverse problems (see Section 5.1), that is, problems of computing a neural command
(somewhere in the brain) that would produce a desired motor output. As we will see
further in the book, this is a problem for scientists, not for the system for movement
production.
To summarize this section, our bodies were definitely not designed by well-
educated twenty-first century engineers or experts in control theory. So, engineering
and control theory will not help us understand how the structures within the body
interact among themselves and with the environment to bring about purposeful,
coordinated movements. We have to take a closer look at specific features of human
(and animal) movements and at the physiology of the main structures that participate
in movement production. First, however, let us consider the history of movement
science, a few facts about the system for movement production, and a few examples
from everyday life and laboratory experiments that emphasize several basic features
of typical human movements and suggest principles of their construction.