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PID controller 1

PID controller
A proportional–integral–derivative
controller (PID controller) is a generic
control loop feedback mechanism
(controller) widely used in industrial control
systems – a PID is the most commonly used
feedback controller. A PID controller
calculates an "error" value as the difference
between a measured process variable and a
desired setpoint. The controller attempts to
minimize the error by adjusting the process A block diagram of a PID controller

control inputs.

The PID controller calculation (algorithm) involves three separate parameters, and is accordingly sometimes called
three-term control: the proportional, the integral and derivative values, denoted P, I, and D. Heuristically, these
values can be interpreted in terms of time: P depends on the present error, I on the accumulation of past errors, and
D is a prediction of future errors, based on current rate of change.[1] The weighted sum of these three actions is used
to adjust the process via a control element such as the position of a control valve or the power supply of a heating

In the absence of knowledge of the underlying process, a PID controller is the best controller.[2] By tuning the three
constants in the PID controller algorithm, the controller can provide control action designed for specific process
requirements. The response of the controller can be described in terms of the responsiveness of the controller to an
error, the degree to which the controller overshoots the setpoint and the degree of system oscillation. Note that the
use of the PID algorithm for control does not guarantee optimal control of the system or system stability.
Some applications may require using only one or two modes to provide the appropriate system control. This is
achieved by setting the gain of undesired control outputs to zero. A PID controller will be called a PI, PD, P or I
controller in the absence of the respective control actions. PI controllers are fairly common, since derivative action is
sensitive to measurement noise, whereas the absence of an integral value may prevent the system from reaching its
target value due to the control action.

Control loop basics

A familiar example of a control loop is the action taken when adjusting hot and cold faucet valves to maintain the
faucet water at the desired temperature. This typically involves the mixing of two process streams, the hot and cold
water. The person touches the water to sense or measure its temperature. Based on this feedback they perform a
control action to adjust the hot and cold water valves until the process temperature stabilizes at the desired value.
Sensing water temperature is analogous to taking a measurement of the process value or process variable (PV). The
desired temperature is called the setpoint (SP). The input to the process (the water valve position) is called the
manipulated variable (MV). The difference between the temperature measurement and the setpoint is the error (e)
and quantifies whether the water is too hot or too cold and by how much.
After measuring the temperature (PV), and then calculating the error, the controller decides when to change the tap
position (MV) and by how much. When the controller first turns the valve on, it may turn the hot valve only slightly
if warm water is desired, or it may open the valve all the way if very hot water is desired. This is an example of a
simple proportional control. In the event that hot water does not arrive quickly, the controller may try to speed-up
the process by opening up the hot water valve more-and-more as time goes by. This is an example of an integral
PID controller 2

Making a change that is too large when the error is small is equivalent to a high gain controller and will lead to
overshoot. If the controller were to repeatedly make changes that were too large and repeatedly overshoot the target,
the output would oscillate around the setpoint in either a constant, growing, or decaying sinusoid. If the oscillations
increase with time then the system is unstable, whereas if they decrease the system is stable. If the oscillations
remain at a constant magnitude the system is marginally stable.
In the interest of achieving a gradual convergence at the desired temperature (SP), the controller may wish to damp
the anticipated future oscillations. So in order to compensate for this effect, the controller may elect to temper their
adjustments. This can be thought of as a derivative control method.
If a controller starts from a stable state at zero error (PV = SP), then further changes by the controller will be in
response to changes in other measured or unmeasured inputs to the process that impact on the process, and hence on
the PV. Variables that impact on the process other than the MV are known as disturbances. Generally controllers are
used to reject disturbances and/or implement setpoint changes. Changes in feedwater temperature constitute a
disturbance to the faucet temperature control process.
In theory, a controller can be used to control any process which has a measurable output (PV), a known ideal value
for that output (SP) and an input to the process (MV) that will affect the relevant PV. Controllers are used in industry
to regulate temperature, pressure, flow rate, chemical composition, speed and practically every other variable for
which a measurement exists.

PID controller theory

This section describes the parallel or non-interacting form of the PID controller. For other forms please see the
section "Alternative nomenclature and PID forms".
The PID control scheme is named after its three correcting terms, whose sum constitutes the manipulated variable
(MV). Hence:

, , and are the contributions to the output from the PID controller from each of the three
terms, as defined below.
PID controller 3

Proportional term
The proportional term (sometimes
called gain) makes a change to the
output that is proportional to the current
error value. The proportional response
can be adjusted by multiplying the error
by a constant Kp, called the proportional

The proportional term is given by:

Plot of PV vs time, for three values of Kp (Ki and Kd held constant)

: Proportional term of output
: Proportional gain, a tuning parameter
: Setpoint, the desired value
: Process value (or process variable), the measured value
: Error
: Time or instantaneous time (the present)
A high proportional gain results in a large change in the output for a given change in the error. If the proportional
gain is too high, the system can become unstable (see the section on loop tuning). In contrast, a small gain results in
a small output response to a large input error, and a less responsive (or sensitive) controller. If the proportional gain
is too low, the control action may be too small when responding to system disturbances.

A pure proportional controller will not always settle at its target value, but may retain a steady-state error.
Specifically, the process gain - drift in the absence of control, such as cooling of a furnace towards room
temperature, biases a pure proportional controller. If the process gain is down, as in cooling, then the bias will be
below the set point, hence the term "droop".
Droop is proportional to process gain and inversely proportional to proportional gain. Specifically the steady-state
error is given by:

Droop is an inherent defect of purely proportional control. Droop may be mitigated by adding a compensating bias
term (setting the setpoint above the true desired value), or corrected by adding an integration term (in a PI or PID
controller), which effectively computes a bias adaptively.
PID controller 4

Despite droop, both tuning theory and industrial practice indicate that it is the proportional term that should
contribute the bulk of the output change.

Integral term
The contribution from the integral term
(sometimes called reset) is proportional
to both the magnitude of the error and
the duration of the error. Summing the
instantaneous error over time
(integrating the error) gives the
accumulated offset that should have
been corrected previously. The
accumulated error is then multiplied by
the integral gain and added to the
controller output. The magnitude of the
contribution of the integral term to the
overall control action is determined by
the integral gain, .

The integral term is given by:

Plot of PV vs time, for three values of Ki (Kp and Kd held constant)

: Integral term of output
: Integral gain, a tuning parameter
: Setpoint, the desired value
: Process value (or process variable), the measured value
: Error
: Time or instantaneous time (the present)
: a dummy integration variable
The integral term (when added to the proportional term) accelerates the movement of the process towards setpoint
and eliminates the residual steady-state error that occurs with a proportional only controller. However, since the
integral term is responding to accumulated errors from the past, it can cause the present value to overshoot the
setpoint value (cross over the setpoint and then create a deviation in the other direction). For further notes regarding
integral gain tuning and controller stability, see the section on loop tuning.
PID controller 5

Derivative term
The rate of change of the process error
is calculated by determining the slope
of the error over time (i.e., its first
derivative with respect to time) and
multiplying this rate of change by the
derivative gain . The magnitude of
the contribution of the derivative term
(sometimes called rate) to the overall
control action is termed the derivative
gain, .

The derivative term is given by:

Plot of PV vs time, for three values of Kd (Kp and Ki held constant)

: Derivative term of output
: Derivative gain, a tuning parameter
: Setpoint, the desired value
: Process value (or process variable), the measured value
: Error
: Time or instantaneous time (the present)
The derivative term slows the rate of change of the controller output and this effect is most noticeable close to the
controller setpoint. Hence, derivative control is used to reduce the magnitude of the overshoot produced by the
integral component and improve the combined controller-process stability. However, differentiation of a signal
amplifies noise and thus this term in the controller is highly sensitive to noise in the error term, and can cause a
process to become unstable if the noise and the derivative gain are sufficiently large. Hence an approximation to a
differentiator with a limited bandwidth is more commonly used. Such a circuit is known as a Phase-Lead
PID controller 6

The proportional, integral, and derivative terms are summed to calculate the output of the PID controller. Defining
as the controller output, the final form of the PID algorithm is:

where the tuning parameters are:

Proportional gain,
Larger values typically mean faster response since the larger the error, the larger the proportional term
compensation. An excessively large proportional gain will lead to process instability and oscillation.
Integral gain,
Larger values imply steady state errors are eliminated more quickly. The trade-off is larger overshoot: any
negative error integrated during transient response must be integrated away by positive error before reaching
steady state.
Derivative gain,
Larger values decrease overshoot, but slow down transient response and may lead to instability due to signal
noise amplification in the differentiation of the error

Loop tuning
Tuning a control loop is the adjustment of its control parameters (gain/proportional band, integral gain/reset,
derivative gain/rate) to the optimum values for the desired control response. Stability (bounded oscillation) is a basic
requirement, but beyond that, different systems have different behavior, different applications have different
requirements, and requirements may conflict with one another.
Some processes have a degree of non-linearity and so parameters that work well at full-load conditions don't work
when the process is starting up from no-load; this can be corrected by gain scheduling (using different parameters in
different operating regions). PID controllers often provide acceptable control using default tunings, but performance
can generally be improved by careful tuning, and performance may be unacceptable with poor tuning.
PID tuning is a difficult problem, even though there are only three parameters and in principle is simple to describe,
because it must satisfy complex criteria within the limitations of PID control. There are accordingly various methods
for loop tuning, and more sophisticated techniques are the subject of patents; this section describes some traditional
manual methods for loop tuning.

If the PID controller parameters (the gains of the proportional, integral and derivative terms) are chosen incorrectly,
the controlled process input can be unstable, i.e. its output diverges, with or without oscillation, and is limited only
by saturation or mechanical breakage. Instability is caused by excess gain, particularly in the presence of significant
Generally, stability of response (the reverse of instability) is required and the process must not oscillate for any
combination of process conditions and setpoints, though sometimes marginal stability (bounded oscillation) is
acceptable or desired.
PID controller 7

Optimum behavior
The optimum behavior on a process change or setpoint change varies depending on the application.
Two basic requirements are regulation (disturbance rejection – staying at a given setpoint) and command tracking
(implementing setpoint changes) – these refer to how well the controlled variable tracks the desired value. Specific
criteria for command tracking include rise time and settling time. Some processes must not allow an overshoot of the
process variable beyond the setpoint if, for example, this would be unsafe. Other processes must minimize the
energy expended in reaching a new setpoint.

Overview of methods
There are several methods for tuning a PID loop. The most effective methods generally involve the development of
some form of process model, then choosing P, I, and D based on the dynamic model parameters. Manual tuning
methods can be relatively inefficient, particularly if the loops have response times on the order of minutes or longer.
The choice of method will depend largely on whether or not the loop can be taken "offline" for tuning, and the
response time of the system. If the system can be taken offline, the best tuning method often involves subjecting the
system to a step change in input, measuring the output as a function of time, and using this response to determine the
control parameters.

Choosing a Tuning Method

Method Advantages Disadvantages

Manual Tuning No math required. Online method. Requires experienced personnel.

Ziegler–Nichols Proven Method. Online method. Process upset, some trial-and-error,

very aggressive tuning.

Software Tools Consistent tuning. Online or offline method. May include valve and sensor analysis. Some cost and training involved.
Allow simulation before downloading. Can support Non-Steady State (NSS) Tuning.

Cohen-Coon Good process models. Some math. Offline method. Only

good for first-order processes.

Manual tuning
If the system must remain online, one tuning method is to first set and values to zero. Increase the
until the output of the loop oscillates, then the should be set to approximately half of that value for a "quarter
amplitude decay" type response. Then increase until any offset is correct in sufficient time for the process.
However, too much will cause instability. Finally, increase , if required, until the loop is acceptably quick
to reach its reference after a load disturbance. However, too much will cause excessive response and overshoot.
A fast PID loop tuning usually overshoots slightly to reach the setpoint more quickly; however, some systems cannot
accept overshoot, in which case an over-damped closed-loop system is required, which will require a setting
significantly less than half that of the setting causing oscillation.
PID controller 8

Effects of increasing a parameter independently

Parameter Rise time Overshoot Settling time Steady-state error [3]

Decrease Increase Small change Decrease Degrade

[4] Increase Increase Decrease significantly Degrade


Minor decrease Minor decrease Minor decrease No effect in theory Improve if small

Ziegler–Nichols method
Another heuristic tuning method is formally known as the Ziegler–Nichols method, introduced by John G. Ziegler
and Nathaniel B. Nichols in the 1940s. As in the method above, the and gains are first set to zero. The P
gain is increased until it reaches the ultimate gain, , at which the output of the loop starts to oscillate. and
the oscillation period are used to set the gains as shown:

Ziegler–Nichols method
Control Type

P - -

PI -


These gains apply to the ideal, parallel form of the PID controller. When applied to the standard PID form, the
integral and derivative time parameters and are only dependent on the oscillation period . Please see the
section "Alternative nomenclature and PID forms".

PID tuning software

Most modern industrial facilities no longer tune loops using the manual calculation methods shown above. Instead,
PID tuning and loop optimization software are used to ensure consistent results. These software packages will gather
the data, develop process models, and suggest optimal tuning. Some software packages can even develop tuning by
gathering data from reference changes.
Mathematical PID loop tuning induces an impulse in the system, and then uses the controlled system's frequency
response to design the PID loop values. In loops with response times of several minutes, mathematical loop tuning is
recommended, because trial and error can literally take days just to find a stable set of loop values. Optimal values
are harder to find. Some digital loop controllers offer a self-tuning feature in which very small setpoint changes are
sent to the process, allowing the controller itself to calculate optimal tuning values.
Other formulas are available to tune the loop according to different performance criteria. Many patented formulas are
now embedded within PID tuning software and hardware modules.
Advances in automated PID Loop Tuning software also deliver algorithms for tuning PID Loops in a dynamic or
Non-Steady State (NSS) scenario. The software will model the dynamics of a process, through a disturbance, and
calculate PID control parameters in response.
PID controller 9

Modifications to the PID algorithm

The basic PID algorithm presents some challenges in control applications that have been addressed by minor
modifications to the PID form.
Integral windup
One common problem resulting from the ideal PID implementations is integral windup, where a large change in
setpoint occurs (say a positive change) and the integral term accumulates a significant error during the rise (windup),
thus overshooting and continuing to increase as this accumulated error is unwound. This problem can be addressed
• Initializing the controller integral to a desired value
• Increasing the setpoint in a suitable ramp
• Disabling the integral function until the PV has entered the controllable region
• Limiting the time period over which the integral error is calculated
• Preventing the integral term from accumulating above or below pre-determined bounds
Freezing the integral function in case of disturbances
If a PID loop is used to control the temperature of an electric resistance furnace, the system has stabilized and
then the door is opened and something cold is put into the furnace the temperature drops below the setpoint.
The integral function of the controller tends to compensate this error by introducing another error in the
positive direction. This can be avoided by freezing of the integral function after the opening of the door for the
time the control loop typically needs to reheat the furnace.
Replacing the integral function by a model based part
Often the time-response of the system is approximately known. Then it is an advantage to simulate this
time-response with a model and to calculate some unknown parameter from the actual response of the system.
If for instance the system is an electrical furnace the response of the difference between furnace temperature
and ambient temperature to changes of the electrical power will be similar to that of a simple RC low-pass
filter multiplied by an unknown proportional coefficient. The actual electrical power supplied to the furnace is
delayed by a low-pass filter to simulate the response of the temperature of the furnace and then the actual
temperature minus the ambient temperature is divided by this low-pass filtered electrical power. Then, the
result is stabilized by another low-pass filter leading to an estimation of the proportional coefficient. With this
estimation, it is possible to calculate the required electrical power by dividing the set-point of the temperature
minus the ambient temperature by this coefficient. The result can then be used instead of the integral function.
This also achieves a control error of zero in the steady-state, but avoids integral windup and can give a
significantly improved control action compared to an optimized PID controller. This type of controller does
work properly in an open loop situation which causes integral windup with an integral function. This is an
advantage if, for example, the heating of a furnace has to be reduced for some time because of the failure of a
heating element, or if the controller is used as an advisory system to a human operator who may not switch it
to closed-loop operation. It may also be useful if the controller is inside a branch of a complex control system
that may be temporarily inactive.
Many PID loops control a mechanical device (for example, a valve). Mechanical maintenance can be a major cost
and wear leads to control degradation in the form of either stiction or a deadband in the mechanical response to an
input signal. The rate of mechanical wear is mainly a function of how often a device is activated to make a change.
Where wear is a significant concern, the PID loop may have an output deadband to reduce the frequency of
activation of the output (valve). This is accomplished by modifying the controller to hold its output steady if the
change would be small (within the defined deadband range). The calculated output must leave the deadband before
the actual output will change.
PID controller 10

The proportional and derivative terms can produce excessive movement in the output when a system is subjected to
an instantaneous step increase in the error, such as a large setpoint change. In the case of the derivative term, this is
due to taking the derivative of the error, which is very large in the case of an instantaneous step change. As a result,
some PID algorithms incorporate the following modifications:
Derivative of output
In this case the PID controller measures the derivative of the output quantity, rather than the derivative of the
error. The output is always continuous (i.e., never has a step change). For this to be effective, the derivative of
the output must have the same sign as the derivative of the error.
Setpoint ramping
In this modification, the setpoint is gradually moved from its old value to a newly specified value using a
linear or first order differential ramp function. This avoids the discontinuity present in a simple step change.
Setpoint weighting
Setpoint weighting uses different multipliers for the error depending on which element of the controller it is
used in. The error in the integral term must be the true control error to avoid steady-state control errors. This
affects the controller's setpoint response. These parameters do not affect the response to load disturbances and
measurement noise.

PID controllers date to 1890s governor design.[2] [5] PID controllers
were subsequently developed in automatic ship steering. One of the
earliest examples of a PID-type controller was developed by Elmer
Sperry in 1911,[6] while the first published theoretical analysis of a PID
controller was by Russian American engineer Nicolas Minorsky, in
(Minorsky 1922). Minorsky was designing automatic steering systems
for the US Navy, and based his analysis on observations of a
helmsman, observing that the helmsman controlled the ship not only
based on the current error, but also on past error and current rate of PID theory developed by observing the action of
change;[7] this was then made mathematical by Minorsky. His goal was helmsmen.
stability, not general control, which significantly simplified the
problem. While proportional control provides stability against small disturbances, it was insufficient for dealing with
a steady disturbance, notably a stiff gale (due to droop), which required adding the integral term. Finally, the
derivative term was added to improve control.

Trials were carried out on the USS New Mexico, with the controller controlling the angular velocity (not angle) of
the rudder. PI control yielded sustained yaw (angular error) of ±2°, while adding D yielded yaw of ±1/6°, better than
most helmsmen could achieve.[8]
The Navy ultimately did not adopt the system, due to resistance by personnel. Similar work was carried out and
published by several others in the 1930s.

Limitations of PID control

While PID controllers are applicable to many control problems, and often perform satisfactorily without any
improvements or even tuning, they can perform poorly in some applications, and do not in general provide optimal
control. The fundamental difficulty with PID control is that it is a feedback system, with constant parameters, and no
direct knowledge of the process, and thus overall performance is reactive and a compromise – while PID control is
the best controller with no model of the process,[2] better performance can be obtained by incorporating a model of
PID controller 11

the process.
The most significant improvement is to incorporate feed-forward control with knowledge about the system, and
using the PID only to control error. Alternatively, PIDs can be modified in more minor ways, such as by changing
the parameters (either gain scheduling in different use cases or adaptively modifying them based on performance),
improving measurement (higher sampling rate, precision, and accuracy, and low-pass filtering if necessary), or
cascading multiple PID controllers.
PID controllers, when used alone, can give poor performance when the PID loop gains must be reduced so that the
control system does not overshoot, oscillate or hunt about the control setpoint value. They also have difficulties in
the presence of non-linearities, may trade off regulation versus response time, do not react to changing process
behavior (say, the process changes after it has warmed up), and have lag in responding to large disturbances.

Another problem faced with PID controllers is that they are linear, and in particular symmetric. Thus, performance
of PID controllers in non-linear systems (such as HVAC systems) is variable. For example, in temperature control, a
common use case is active heating (via a heating element) but passive cooling (heating off, but no cooling), so
overshoot can only be corrected slowly – it cannot be forced downward. In this case the PID should be tuned to be
overdamped, to prevent or reduce overshoot, though this reduces performance (it increases settling time).

Noise in derivative
A problem with the derivative term is that small amounts of measurement or process noise can cause large amounts
of change in the output. It is often helpful to filter the measurements with a low-pass filter in order to remove
higher-frequency noise components. However, low-pass filtering and derivative control can cancel each other out, so
reducing noise by instrumentation means is a much better choice. Alternatively, a nonlinear median filter may be
used, which improves the filtering efficiency and practical performance [9] . In some case, the differential band can
be turned off in many systems with little loss of control. This is equivalent to using the PID controller as a PI


The control system performance can be improved by combining the feedback (or closed-loop) control of a PID
controller with feed-forward (or open-loop) control. Knowledge about the system (such as the desired acceleration
and inertia) can be fed forward and combined with the PID output to improve the overall system performance. The
feed-forward value alone can often provide the major portion of the controller output. The PID controller can be used
primarily to respond to whatever difference or error remains between the setpoint (SP) and the actual value of the
process variable (PV). Since the feed-forward output is not affected by the process feedback, it can never cause the
control system to oscillate, thus improving the system response and stability.
For example, in most motion control systems, in order to accelerate a mechanical load under control, more force or
torque is required from the prime mover, motor, or actuator. If a velocity loop PID controller is being used to control
the speed of the load and command the force or torque being applied by the prime mover, then it is beneficial to take
the instantaneous acceleration desired for the load, scale that value appropriately and add it to the output of the PID
velocity loop controller. This means that whenever the load is being accelerated or decelerated, a proportional
amount of force is commanded from the prime mover regardless of the feedback value. The PID loop in this
situation uses the feedback information to change the combined output to reduce the remaining difference between
the process setpoint and the feedback value. Working together, the combined open-loop feed-forward controller and
closed-loop PID controller can provide a more responsive, stable and reliable control system.
PID controller 12

Other improvements
In addition to feed-forward, PID controllers are often enhanced through methods such as PID gain scheduling
(changing parameters in different operating conditions), fuzzy logic or computational verb logic [10] [11] . Further
practical application issues can arise from instrumentation connected to the controller. A high enough sampling rate,
measurement precision, and measurement accuracy are required to achieve adequate control performance.

Cascade control
One distinctive advantage of PID controllers is that two PID controllers can be used together to yield better dynamic
performance. This is called cascaded PID control. In cascade control there are two PIDs arranged with one PID
controlling the set point of another. A PID controller acts as outer loop controller, which controls the primary
physical parameter, such as fluid level or velocity. The other controller acts as inner loop controller, which reads the
output of outer loop controller as set point, usually controlling a more rapid changing parameter, flowrate or
acceleration. It can be mathematically proven that the working frequency of the controller is increased and the time
constant of the object is reduced by using cascaded PID controller..

Physical implementation of PID control

In the early history of automatic process control the PID controller was implemented as a mechanical device. These
mechanical controllers used a lever, spring and a mass and were often energized by compressed air. These pneumatic
controllers were once the industry standard.
Electronic analog controllers can be made from a solid-state or tube amplifier, a capacitor and a resistance.
Electronic analog PID control loops were often found within more complex electronic systems, for example, the
head positioning of a disk drive, the power conditioning of a power supply, or even the movement-detection circuit
of a modern seismometer. Nowadays, electronic controllers have largely been replaced by digital controllers
implemented with microcontrollers or FPGAs.
Most modern PID controllers in industry are implemented in programmable logic controllers (PLCs) or as a
panel-mounted digital controller. Software implementations have the advantages that they are relatively cheap and
are flexible with respect to the implementation of the PID algorithm.
Variable voltages may be applied by the time proportioning form of Pulse-width modulation (PWM) – a cycle time
is fixed, and variation is achieved by varying the proportion of the time during this cycle that the controller outputs
+1 (or −1) instead of 0. On a digital system the possible proportions are discrete – e.g., increments of .1 second
within a 2 second cycle time yields 20 possible steps: percentage increments of 5% – so there is a discretization
error, but for high enough time resolution this yields satisfactory performance.

Alternative nomenclature and PID forms

Ideal versus standard PID form

The form of the PID controller most often encountered in industry, and the one most relevant to tuning algorithms is
the standard form. In this form the gain is applied to the , and terms, yielding:

is the integral time
is the derivative time
PID controller 13

In this standard form, the parameters have a clear physical meaning. In particular, the inner summation produces a
new single error value which is compensated for future and past errors. The addition of the proportional and
derivative components effectively predicts the error value at seconds (or samples) in the future, assuming that
the loop control remains unchanged. The integral component adjusts the error value to compensate for the sum of all
past errors, with the intention of completely eliminating them in seconds (or samples). The resulting
compensated single error value is scaled by the single gain .
In the ideal parallel form, shown in the controller theory section

the gain parameters are related to the parameters of the standard form through and . This

parallel form, where the parameters are treated as simple gains, is the most general and flexible form. However, it is
also the form where the parameters have the least physical interpretation and is generally reserved for theoretical
treatment of the PID controller. The standard form, despite being slightly more complex mathematically, is more
common in industry.

Laplace form of the PID controller

Sometimes it is useful to write the PID regulator in Laplace transform form:

Having the PID controller written in Laplace form and having the transfer function of the controlled system makes it
easy to determine the closed-loop transfer function of the system.

PID Pole Zero Cancellation

The PID equation can be written in this form:

When this form is used it is easy to determine the closed loop transfer function.



This can be very useful to remove unstable poles

PID controller 14

Series/interacting form
Another representation of the PID controller is the series, or interacting form

where the parameters are related to the parameters of the standard form through
, , and


This form essentially consists of a PD and PI controller in series, and it made early (analog) controllers easier to
build. When the controllers later became digital, many kept using the interacting form.

Discrete implementation
The analysis for designing a digital implementation of a PID controller in a Microcontroller (MCU) or FPGA device
requires the standard form of the PID controller to be discretised [12] . Approximations for first-order derivatives are
made by backward finite differences. The integral term is discretised, with a sampling time ,as follows,

The derivative term is approximated as,

Thus, a velocity algorithm for implementation of the discretised PID controller in a MCU is obtained by
differentiating , using the numerical definitions of the first and second derivative and solving for and
finally obtaining:

Here is a simple software loop that implements the PID algorithm in its 'ideal, parallel' form:

previous_error = 0
integral = 0
error = setpoint - actual_position
integral = integral + (error*dt)
derivative = (error - previous_error)/dt
output = (Kp*error) + (Ki*integral) + (Kd*derivative)
previous_error = error
goto start
PID controller 15

PI controller
A PI Controller (proportional-integral
controller) is a special case of the PID
controller in which the derivative (D) of the
error is not used.
The controller output is given by

Basic block of a PI controller.

where is the error or deviation of actual measured value (PV) from the set-point (SP).
= SP - PV.
A PI controller can be modelled easily in software such as Simulink using a "flow chart" box involving Laplace

= proportional gain
= integral gain
Setting a value for is often a trade off between decreasing overshoot and increasing settling time.
The lack of derivative action may make the system more steady in the steady state in the case of noisy data. This is
because derivative action is more sensitive to higher-frequency terms in the inputs.
Without derivative action, a PI-controlled system is less responsive to real (non-noise) and relatively fast alterations
in state and so the system will be slower to reach setpoint and slower to respond to perturbations than a well-tuned
PID system may be.

[1] Araki, M.. "PID Control" (http:/ / www. eolss. net/ ebooks/ Sample Chapters/ C18/ E6-43-03-03. pdf). .
[2] Bennett, Stuart (1993). A history of control engineering, 1930-1955. IET. p.  p. 48 (http:/ / books. google. com/ books?id=VD_b81J3yFoC&
pg=PA48). ISBN 9-780863412998
[3] Ang, K.H., Chong, G.C.Y., and Li, Y. (2005). PID control system analysis, design, and technology, IEEE Trans Control Systems Tech, 13(4),
pp.559-576. http:/ / eprints. gla. ac. uk/ 3817/
[4] http:/ / saba. kntu. ac. ir/ eecd/ pcl/ download/ PIDtutorial. pdf
[5] Bennett, Stuart (November 1984). "Nicholas Minorsky and the automatic steering of ships" (http:/ / ieeexplore. ieee. org/ iel5/ 37/ 24267/
01104827. pdf?arnumber=1104827). IEEE Control Systems Magazine 4 (4): 10–15. doi:10.1109/MCS.1984.1104827. ISSN 0272-1708.
[6] http:/ / www. building-automation-consultants. com/ building-automation-history. html
[7] (Bennett 1993, p. 67 (http:/ / books. google. com/ books?id=VD_b81J3yFoC& pg=PA67))
[8] Bennett, Stuart (June 1986). A history of control engineering, 1800-1930. IET. pp.  142–148 (http:/ / books. google. com/
books?id=1gfKkqB_fTcC& pg=PA142). ISBN 978-0-86341047-5.
[9] Li, Y. and Ang, K.H. and Chong, G.C.Y. (2006) PID control system analysis and design - Problems, remedies, and future directions. IEEE
Control Systems Magazine, 26 (1). pp. 32-41. ISSN 0272-1708 (http:/ / eprints. gla. ac. uk/ 3815/ )
[10] Yang, T. (June 2005). "Architectures of Computational Verb Controllers: Towards a New Paradigm of Intelligent Control". International
Journal of Computational Cognition (Yang's Scientific Press) 3 (2): 74–101.
[11] Liang, Y.-L.(et al) (2009). "Controlling fuel annealer using computational verb PID controllers". Proceedings of the 3rd international
conference on Anti-Counterfeiting, security, and identification in communication (IEEE): 417–420.
[12] http:/ / www. scribd. com/ doc/ 19070283/ Discrete-PI-and-PID-Controller-Design-and-Analysis-for-Digital-Implementation
PID controller 16

• Minorsky, Nicolas (1922). "Directional stability of automatically steered bodies". J. Amer. Soc. Naval Eng. 34
(2): 280–309
• Liptak, Bela (1995). Instrument Engineers' Handbook: Process Control. Radnor, Pennsylvania: Chilton Book
Company. pp. 20–29. ISBN 0-8019-8242-1.
• Tan, Kok Kiong; Wang Qing-Guo, Hang Chang Chieh (1999). Advances in PID Control. London, UK:
Springer-Verlag. ISBN 1-85233-138-0.
• Van, Doren, Vance J. (July 1, 2003). "Loop Tuning Fundamentals" (
268148-Loop_Tuning_Fundamentals.php.html). Control Engineering (Reed Business Information).
• Sellers, David. "An Overview of Proportional plus Integral plus Derivative Control and Suggestions for Its
Successful Application and Implementation" ( (PDF). Archived from the original (http://www.peci.
org/library/PECI_ControlOverview1_1002.pdf) on March 7, 2007. Retrieved 2007-05-05.
• Graham, Ron (10/03/2005). "FAQ on PID controller tuning" ( Retrieved 2009-01-05.

External links

PID tutorials
• PID Tutorial (
• P.I.D. Without a PhD ( a beginner's
guide to PID loop theory with sample programming code
• What's All This P-I-D Stuff, Anyhow? (
Article in Electronic Design
• Shows how to build a PID controller with basic electronic components (
VtolIndoorFlying/rapports/rapportSemStauffer.pdf) (pg. 22)
• Virtual PID Controller Laboratory (
• PID Design & Tuning (
• Online PID Tuning applet from University of Texas Control Group (
• example PID implementation in 16F887 microchip (

Special topics and PID control applications

• Proven Methods and Best Practices for PID Control (
• PID Control Primer ( Article in Embedded Systems
Radial basis function network 17

Radial basis function network

A radial basis function network is an artificial neural network that uses radial basis functions as activation
functions. It is a linear combination of radial basis functions. They are used in function approximation, time series
prediction, and control.

Network architecture

Figure 1: Architecture of a radial basis function network. An input vector x is used as

input to all radial basis functions, each with different parameters. The output of the
network is a linear combination of the outputs from radial basis functions.

Radial basis function (RBF) networks typically have three layers: an input layer, a hidden layer with a non-linear
RBF activation function and a linear output layer. The output, , of the network is thus

where N is the number of neurons in the hidden layer, is the center vector for neuron i, and are the weights of
the linear output neuron. In the basic form all inputs are connected to each hidden neuron. The norm is typically
taken to be the Euclidean distance and the basis function is taken to be Gaussian

The Gaussian basis functions are local in the sense that

i.e. changing parameters of one neuron has only a small effect for input values that are far away from the center of
that neuron.
RBF networks are universal approximators on a compact subset of . This means that a RBF network with
enough hidden neurons can approximate any continuous function with arbitrary precision.
The weights , , and are determined in a manner that optimizes the fit between and the data.
Radial basis function network 18


Normalized architecture

In addition to the above unnormalized

architecture, RBF networks can be
normalized. In this case the mapping is

Figure 2: Two unnormalized radial basis functions in one input dimension. The basis
function centers are located at and .


is known as a "normalized radial basis function".

Theoretical motivation for


There is theoretical justification for

this architecture in the case of
stochastic data flow. Assume a
stochastic kernel approximation for the
joint probability density

Figure 3: Two normalized radial basis functions in one input dimension. The basis
function centers are located at and .
Radial basis function network 19

where the weights and are exemplars from the data and we require the kernels to be normalized


The probability densities in the input

and output spaces are

Figure 4: Three normalized radial basis functions in one input dimension. The additional
basis function has center at

The expectation of y given an input is


is the conditional probability of y given . The conditional probability is related to the joint probability through
Bayes theorem

which yields

This becomes

when the integrations are performed.

Radial basis function network 20

Local linear models

It is sometimes convenient to expand
the architecture to include local linear
models. In that case the architectures
become, to first order,

Figure 5: Four normalized radial basis functions in one input dimension. The fourth basis
function has center at . Note that the first basis function (dark blue) has
become localized.


in the unnormalized and normalized cases, respectively. Here are weights to be determined. Higher order linear
terms are also possible.
This result can be written



in the unnormalized case and

in the normalized case.

Here is a Kronecker delta function defined as
Radial basis function network 21

In a RBF network there are three types of parameters that need to be chosen to adapt the network for a particular
task: the center vectors , the output weights , and the RBF width parameters . In the sequential training of
the weights are updated at each time step as data streams in.
For some tasks it makes sense to define an objective function and select the parameter values that minimize its value.
The most common objective function is the least squares function


We have explicitly included the dependence on the weights. Minimization of the least squares objective function by
optimal choice of weights optimizes accuracy of fit.
There are occasions in which multiple objectives, such as smoothness as well as accuracy, must be optimized. In that
case it is useful to optimize a regularized objective function such as



where optimization of S maximizes smoothness and is known as a regularization parameter.

RBF networks can be used to interpolate a function when the values of that function are known on
finite number of points: . Taking the known points to be the centers of the radial
basis functions and evaluating the values of the basis functions at the same points the
weights can be solved from the equation

It can be shown that the interpolation matrix in the above equation is non-singular, if the points are distinct, and
thus the weights can be solved by simple linear algebra:
Radial basis function network 22

Function approximation
If the purpose is not to perform strict interpolation but instead more general function approximation or classification
the optimization is somewhat more complex because there is no obvious choice for the centers. The training is
typically done in two phases first fixing the width and centers and then the weights. This can be justified by
considering the different nature of the non-linear hidden neurons versus the linear output neuron.

Training the basis function centers

Basis function centers can be randomly sampled among the input instances or obtained by Orthogonal Least Square
Learning Algorithm or found by clustering the samples and choosing the cluster means as the centers.
The RBF widths are usually all fixed to same value which is proportional to the maximum distance between the
chosen centers.

Pseudoinverse solution for the linear weights

After the centers have been fixed, the weights that minimize the error at the output are computed with a linear
pseudoinverse solution:
where the entries of G are the values of the radial basis functions evaluated at the points :
The existence of this linear solution means that unlike Multi-Layer Perceptron (MLP) networks the RBF networks
have a unique local minimum (when the centers are fixed).

Gradient descent training of the linear weights

Another possible training algorithm is gradient descent. In gradient descent training, the weights are adjusted at each
time step by moving them in a direction opposite from the gradient of the objective function (thus allowing the
minimum of the objective function to be found),

where is a "learning parameter."

For the case of training the linear weights, , the algorithm becomes

in the unnormalized case and

in the normalized case.

For local-linear-architectures gradient-descent training is
Radial basis function network 23

Projection operator training of the linear weights

For the case of training the linear weights, and , the algorithm becomes

in the unnormalized case and

in the normalized case and

in the local-linear case.

For one basis function, projection operator training reduces to Newton's method.


Logistic map
The basic properties of radial basis
functions can be illustrated with a
simple mathematical map, the logistic
map, which maps the unit interval onto
itself. It can be used to generate a
convenient prototype data stream. The
logistic map can be used to explore
function approximation, time series
prediction, and control theory. The
map originated from the field of
Figure 6: Logistic map time series. Repeated iteration of the logistic map generates a
population dynamics and became the chaotic time series. The values lie between zero and one. Displayed here are the 100
prototype chaotic time series. The map, training points used to train the examples in this section. The weights c are the first five
in the fully chaotic regime, is given by points from this time series.

where t is a time index. The value of x at time t+1 is a parabolic function of x at time t. This equation represents the
underlying geometry of the chaotic time series generated by the logistic map.
Generation of the time series from this equation is the forward problem. The examples here illustrate the inverse
problem; identification of the underlying dynamics, or fundamental equation, of the logistic map from exemplars of
the time series. The goal is to find an estimate

for f.
Radial basis function network 24

Function approximation

Unnormalized radial basis functions

The architecture is

Figure 7: Unnormalized basis functions. The Logistic map (blue) and the approximation
to the logistic map (red) after one pass through the training set.


Since the input is a scalar rather than a vector, the input dimension is one. We choose the number of basis functions
as N=5 and the size of the training set to be 100 exemplars generated by the chaotic time series. The weight is
taken to be a constant equal to 5. The weights are five exemplars from the time series. The weights are trained
with projection operator training:

where the learning rate is taken to be 0.3. The training is performed with one pass through the 100 training points.
The rms error is 0.15.
Radial basis function network 25

Normalized radial basis functions

The normalized RBF architecture is

Figure 8: Normalized basis functions. The Logistic map (blue) and the approximation to
the logistic map (red) after one pass through the training set. Note the improvement over
the unnormalized case.



Again, we choose the number of basis functions as five and the size of the training set to be 100 exemplars generated
by the chaotic time series. The weight is taken to be a constant equal to 6. The weights are five exemplars
from the time series. The weights are trained with projection operator training:

where the learning rate is again taken to be 0.3. The training is performed with one pass through the 100 training
points. The rms error on a test set of 100 exemplars is 0.084, smaller than the unnormalized error. Normalization
yields accuracy improvement. Typically accuracy with normalized basis functions increases even more over
unnormalized functions as input dimensionality increases.
Radial basis function network 26

Time series prediction

Once the underlying geometry of the
time series is estimated as in the
previous examples, a prediction for the
time series can be made by iteration:

Figure 9: Normalized basis functions. The Logistic map (blue) and the approximation to
the logistic map (red) as a function of time. Note that the approximation is good for only a
few time steps. This is a general characterisitc of chaotic time series.

A comparison of the actual and estimated time series is displayed in the figure. The estimated times series starts out
at time zero with an exact knowledge of x(0). It then uses the estimate of the dynamics to update the time series
estimate for several time steps.
Note that the estimate is accurate for only a few time steps. This is a general characteristic of chaotic time series.
This is a property of the sensitive dependence on initial conditions common to chaotic time series. A small initial
error is amplified with time. A measure of the divergence of time series with nearly identical initial conditions is
known as the Lyapunov exponent.
Radial basis function network 27

Control of a chaotic time series

We assume the output of the logistic
map can be manipulated through a
control parameter such that

Figure 10: Control of the logistic map. The system is allowed to evolve naturally for 49
time steps. At time 50 control is turned on. The desired trajectory for the time series is
red. The system under control learns the underlying dynamics and drives the time series
to the desired output. The architecture is the same as for the time series prediction

The goal is to choose the control parameter in such a way as to drive the time series to a desired output . This
can be done if we choose the control paramer to be


is an approximation to the underlying natural dynamics of the system.

The learning algorithm is given by


• J. Moody and C. J. Darken, "Fast learning in networks of locally tuned processing units," Neural Computation, 1,
281-294 (1989). Also see Radial basis function networks according to Moody and Darken [1]
• T. Poggio and F. Girosi, "Networks for approximation and learning," Proc. IEEE 78(9), 1484-1487 (1990).
• Roger D. Jones, Y. C. Lee, C. W. Barnes, G. W. Flake, K. Lee, P. S. Lewis, and S. Qian, ?Function
approximation and time series prediction with neural networks [2],? Proceedings of the International Joint
Conference on Neural Networks, June 17-21, p. I-649 (1990).
• Martin D. Buhmann (2003). Radial Basis Functions: Theory and Implementations. Cambridge University. ISBN
• Yee, Paul V. and Haykin, Simon (2001). Regularized Radial Basis Function Networks: Theory and Applications.
John Wiley. ISBN 0-471-35349-3.
Radial basis function network 28

• John R. Davies, Stephen V. Coggeshall, Roger D. Jones, and Daniel Schutzer, "Intelligent Security Systems," in
Freedman, Roy S., Flein, Robert A., and Lederman, Jess, Editors (1995). Artificial Intelligence in the Capital
Markets. Chicago: Irwin. ISBN 1-55738-811-3.
• Simon Haykin (1999). Neural Networks: A Comprehensive Foundation (2nd edition ed.). Upper Saddle River,
NJ: Prentice Hall. ISBN 0-13-908385-5.
• S. Chen, C. F. N. Cowan, and P. M. Grant, "Orthogonal Least Squares Learning Algorithm for Radial Basis
Function Networks", IEEE Transactions on Neural Networks, Vol 2, No 2 (Mar) 1991.

[1] http:/ / www. ki. inf. tu-dresden. de/ ~fritzke/ FuzzyPaper/ node5. html
[2] http:/ / ieeexplore. ieee. org/ xpl/ freeabs_all. jsp?arnumber=137644

Control theory
Control theory is an interdisciplinary
branch of engineering and
mathematics, that deals with the
behavior of dynamical systems. The
desired output of a system is called the
reference. When one or more output
The concept of the feedback loop to control the dynamic behavior of the system: this is
variables of a system need to follow a negative feedback, because the sensed value is subtracted from the desired value to create
certain reference over time, a the error signal which is amplified by the controller.
controller manipulates the inputs to a
system to obtain the desired effect on the output of the system.

Control theory is
• a theory that deals with influencing the behavior of dynamical systems
• an interdisciplinary subfield of science, which originated in engineering and mathematics, and evolved into use by
the social sciences, like psychology, sociology and criminology.

An example
Consider a car's cruise control, which is a device designed to maintain vehicle speed at a constant desired or
reference speed provided by the driver. The controller is the cruise control, the plant is the car, and the system is the
car and the cruise control. The system output is the car's speed, and the control itself is the engine's throttle position
which determines how much power the engine generates.
A primitive way to implement cruise control is simply to lock the throttle position when the driver engages cruise
control. However, if the cruise control is engaged on a stretch of flat road, then the car will travel slower going uphill
and faster when going downhill. This type of controller is called an open-loop controller because no measurement of
the system output (the car's speed) is used to alter the control (the throttle position.) As a result, the controller can not
compensate for changes acting on the car, like a change in the slope of the road.
In a closed-loop control system, a sensor monitors the system output (the car's speed) and feeds the data to a
controller which adjusts the control (the throttle position) as necessary to maintain the desired system output (match
the car's speed to the reference speed.) Now when the car goes uphill the decrease in speed is measured, and the
Control theory 29

throttle position changed to increase engine power, speeding the vehicle. Feedback from measuring the car's speed
has allowed the controller to dynamically compensate for changes to the car's speed. It is from this feedback that the
paradigm of the control loop arises: the control affects the system output, which in turn is measured and looped back
to alter the control.

Although control systems of various types date back to antiquity, a
more formal analysis of the field began with a dynamics analysis of the
centrifugal governor, conducted by the physicist James Clerk Maxwell
in 1868 entitled On Governors.[1] This described and analyzed the
phenomenon of "hunting", in which lags in the system can lead to
overcompensation and unstable behavior. This generated a flurry of
interest in the topic, during which Maxwell's classmate Edward John
Routh generalized the results of Maxwell for the general class of linear
systems.[2] Independently, Adolf Hurwitz analyzed system stability
using differential equations in 1877, resulting in what is now known as
the Routh-Hurwitz theorem.[3] [4]

A notable application of dynamic control was in the area of manned

flight. The Wright Brothers made their first successful test flights on
December 17, 1903 and were distinguished by their ability to control
their flights for substantial periods (more so than the ability to produce
Centrifugal governor in a Boulton & Watt engine
lift from an airfoil, which was known). Control of the airplane was of 1788
necessary for safe flight.

By World War II, control theory was an important part of fire-control systems, guidance systems and electronics.
The Space Race also depended on accurate spacecraft control. However, control theory also saw an increasing use in
fields such as economics.

People in systems and control

Many active and historical figures made significant contribution to control theory, including, for example:
• Alexander Lyapunov (1857–1918) in the 1890s marks the beginning of stability theory.
• Harold S. Black (1898–1983), invented the concept of negative feedback amplifiers in 1927. He managed to
develop stable negative feedback amplifiers in the 1930s.
• Harry Nyquist (1889–1976), developed the Nyquist stability criterion for feedback systems in the 1930s.
• Richard Bellman (1920–1984), developed dynamic programming since the 1940s.
• Andrey Kolmogorov (1903–1987) co-developed the Wiener-Kolmogorov filter (1941).
• Norbert Wiener (1894–1964) co-developed the Wiener-Kolmogorov filter and coined the term cybernetics in the
• John R. Ragazzini (1912–1988) introduced digital control and the z-transform in the 1950s.
• Lev Pontryagin (1908–1988) introduced the maximum principle and the bang-bang principle.
Control theory 30

Classical control theory

To avoid the problems of the open-loop controller, control theory introduces feedback. A closed-loop controller uses
feedback to control states or outputs of a dynamical system. Its name comes from the information path in the system:
process inputs (e.g. voltage applied to an electric motor) have an effect on the process outputs (e.g. velocity or torque
of the motor), which is measured with sensors and processed by the controller; the result (the control signal) is used
as input to the process, closing the loop.
Closed-loop controllers have the following advantages over open-loop controllers:
• disturbance rejection (such as unmeasured friction in a motor)
• guaranteed performance even with model uncertainties, when the model structure does not match perfectly the
real process and the model parameters are not exact
• unstable processes can be stabilized
• reduced sensitivity to parameter variations
• improved reference tracking performance
In some systems, closed-loop and open-loop control are used simultaneously. In such systems, the open-loop control
is termed feedforward and serves to further improve reference tracking performance.
A common closed-loop controller architecture is the PID controller.

Closed-loop transfer function

The output of the system y(t) is fed back through a sensor measurement F to the reference value r(t). The controller
C then takes the error e (difference) between the reference and the output to change the inputs u to the system under
control P. This is shown in the figure. This kind of controller is a closed-loop controller or feedback controller.
This is called a single-input-single-output (SISO) control system; MIMO (i.e. Multi-Input-Multi-Output) systems,
with more than one input/output, are common. In such cases variables are represented through vectors instead of
simple scalar values. For some distributed parameter systems the vectors may be infinite-dimensional (typically

If we assume the controller C, the plant P, and the sensor F are linear and time-invariant (i.e.: elements of their
transfer function C(s), P(s), and F(s) do not depend on time), the systems above can be analysed using the Laplace
transform on the variables. This gives the following relations:

Solving for Y(s) in terms of R(s) gives:

Control theory 31

The expression is referred to as the closed-loop transfer function of the system.

The numerator is the forward (open-loop) gain from r to y, and the denominator is one plus the gain in going around
the feedback loop, the so-called loop gain. If , i.e. it has a large norm with each value of s, and if
, then Y(s) is approximately equal to R(s) and the output closely tracks the reference input.

PID controller
The PID controller is probably the most-used feedback control design. PID is an acronym for
Proportional-Integral-Differential, referring to the three terms operating on the error signal to produce a control
signal. If u(t) is the control signal sent to the system, y(t) is the measured output and r(t) is the desired output, and
tracking error , a PID controller has the general form

The desired closed loop dynamics is obtained by adjusting the three parameters , and , often
iteratively by "tuning" and without specific knowledge of a plant model. Stability can often be ensured using only
the proportional term. The integral term permits the rejection of a step disturbance (often a striking specification in
process control). The derivative term is used to provide damping or shaping of the response. PID controllers are the
most well established class of control systems: however, they cannot be used in several more complicated cases,
especially if MIMO systems are considered.
Applying Laplace transformation results in the transformed PID controller equation

with the PID controller transfer function

Modern control theory

In contrast to the frequency domain analysis of the classical control theory, modern control theory utilizes the
time-domain state space representation, a mathematical model of a physical system as a set of input, output and state
variables related by first-order differential equations. To abstract from the number of inputs, outputs and states, the
variables are expressed as vectors and the differential and algebraic equations are written in matrix form (the latter
only being possible when the dynamical system is linear). The state space representation (also known as the
"time-domain approach") provides a convenient and compact way to model and analyze systems with multiple inputs
and outputs. With inputs and outputs, we would otherwise have to write down Laplace transforms to encode all the
information about a system. Unlike the frequency domain approach, the use of the state space representation is not
limited to systems with linear components and zero initial conditions. "State space" refers to the space whose axes
are the state variables. The state of the system can be represented as a vector within that space.[5] )
Control theory 32

Topics in control theory

The stability of a general dynamical system with no input can be described with Lyapunov stability criteria. A linear
system that takes an input is called bounded-input bounded-output (BIBO) stable if its output will stay bounded for
any bounded input. Stability for nonlinear systems that take an input is input-to-state stability (ISS), which combines
Lyapunov stability and a notion similar to BIBO stability. For simplicity, the following descriptions focus on
continuous-time and discrete-time linear systems.
Mathematically, this means that for a causal linear system to be stable all of the poles of its transfer function must
satisfy some criteria depending on whether a continuous or discrete time analysis is used:
• In continuous time, the Laplace transform is used to obtain the transfer function. A system is stable if the poles of
this transfer function lie strictly in the open left half of the complex plane (i.e. the real part of all the poles is less
than zero).
• In discrete time the Z-transform is used. A system is stable if the poles of this transfer function lie strictly inside
the unit circle. i.e. the magnitude of the poles is less than one).
When the appropriate conditions above are satisfied a system is said to be asymptotically stable: the variables of an
asymptotically stable control system always decrease from their initial value and do not show permanent oscillations.
Permanent oscillations occur when a pole has a real part exactly equal to zero (in the continuous time case) or a
modulus equal to one (in the discrete time case). If a simply stable system response neither decays nor grows over
time, and has no oscillations, it is marginally stable: in this case the system transfer function has non-repeated poles
at complex plane origin (i.e. their real and complex component is zero in the continuous time case). Oscillations are
present when poles with real part equal to zero have an imaginary part not equal to zero.
Differences between the two cases are not a contradiction. The Laplace transform is in Cartesian coordinates and the
Z-transform is in circular coordinates, and it can be shown that:
• the negative-real part in the Laplace domain can map onto the interior of the unit circle
• the positive-real part in the Laplace domain can map onto the exterior of the unit circle
If a system in question has an impulse response of

then the Z-transform (see this example), is given by

which has a pole in (zero imaginary part). This system is BIBO (asymptotically) stable since the pole is
inside the unit circle.
However, if the impulse response was

then the Z-transform is

which has a pole at and is not BIBO stable since the pole has a modulus strictly greater than one.
Numerous tools exist for the analysis of the poles of a system. These include graphical systems like the root locus,
Bode plots or the Nyquist plots.
Mechanical changes can make equipment (and control systems) more stable. Sailors add ballast to improve the
stability of ships. Cruise ships use antiroll fins that extend transversely from the side of the ship for perhaps 30 feet
(10 m) and are continuously rotated about their axes to develop forces that oppose the roll.
Control theory 33

Controllability and observability

Controllability and observability are main issues in the analysis of a system before deciding the best control strategy
to be applied, or whether it is even possible to control or stabilize the system. Controllability is related to the
possibility of forcing the system into a particular state by using an appropriate control signal. If a state is not
controllable, then no signal will ever be able to control the state. If a state is not controllable, but its dynamics are
stable, then the state is termed Stabilizable. Observability instead is related to the possibility of "observing", through
output measurements, the state of a system. If a state is not observable, the controller will never be able to determine
the behaviour of an unobservable state and hence cannot use it to stabilize the system. However, similar to the
stabilizability condition above, if a state cannot be observed it might still be detectable.
From a geometrical point of view, looking at the states of each variable of the system to be controlled, every "bad"
state of these variables must be controllable and observable to ensure a good behaviour in the closed-loop system.
That is, if one of the eigenvalues of the system is not both controllable and observable, this part of the dynamics will
remain untouched in the closed-loop system. If such an eigenvalue is not stable, the dynamics of this eigenvalue will
be present in the closed-loop system which therefore will be unstable. Unobservable poles are not present in the
transfer function realization of a state-space representation, which is why sometimes the latter is preferred in
dynamical systems analysis.
Solutions to problems of uncontrollable or unobservable system include adding actuators and sensors.

Control specification
Several different control strategies have been devised in the past years. These vary from extremely general ones (PID
controller), to others devoted to very particular classes of systems (especially robotics or aircraft cruise control).
A control problem can have several specifications. Stability, of course, is always present: the controller must ensure
that the closed-loop system is stable, regardless of the open-loop stability. A poor choice of controller can even
worsen the stability of the open-loop system, which must normally be avoided. Sometimes it would be desired to
obtain particular dynamics in the closed loop: i.e. that the poles have , where is a fixed value
strictly greater than zero, instead of simply asking that .
Another typical specification is the rejection of a step disturbance; including an integrator in the open-loop chain (i.e.
directly before the system under control) easily achieves this. Other classes of disturbances need different types of
sub-systems to be included.
Other "classical" control theory specifications regard the time-response of the closed-loop system: these include the
rise time (the time needed by the control system to reach the desired value after a perturbation), peak overshoot (the
highest value reached by the response before reaching the desired value) and others (settling time, quarter-decay).
Frequency domain specifications are usually related to robustness (see after).
Modern performance assessments use some variation of integrated tracking error (IAE,ISA,CQI).

Model identification and robustness

A control system must always have some robustness property. A robust controller is such that its properties do not
change much if applied to a system slightly different from the mathematical one used for its synthesis. This
specification is important: no real physical system truly behaves like the series of differential equations used to
represent it mathematically. Typically a simpler mathematical model is chosen in order to simplify calculations,
otherwise the true system dynamics can be so complicated that a complete model is impossible.
System identification
The process of determining the equations that govern the model's dynamics is called system identification. This can
be done off-line: for example, executing a series of measures from which to calculate an approximated mathematical
model, typically its transfer function or matrix. Such identification from the output, however, cannot take account of
Control theory 34

unobservable dynamics. Sometimes the model is built directly starting from known physical equations: for example,
in the case of a mass-spring-damper system we know that . Even assuming that a "complete" m
designing the controller, all the parameters included in these equations (called "nominal parameters") are never
known with absolute precision; the control system will have to behave correctly even when connected to physical
system with true parameter values away from nominal.
Some advanced control techniques include an "on-line" identification process (see later). The parameters of the
model are calculated ("identified") while the controller itself is running: in this way, if a drastic variation of the
parameters ensues (for example, if the robot's arm releases a weight), the controller will adjust itself consequently in
order to ensure the correct performance.
Analysis of the robustness of a SISO control system can be performed in the frequency domain, considering the
system's transfer function and using Nyquist and Bode diagrams. Topics include gain and phase margin and
amplitude margin. For MIMO and, in general, more complicated control systems one must consider the theoretical
results devised for each control technique (see next section): i.e., if particular robustness qualities are needed, the
engineer must shift his attention to a control technique including them in its properties.
A particular robustness issue is the requirement for a control system to perform properly in the presence of input and
state constraints. In the physical world every signal is limited. It could happen that a controller will send control
signals that cannot be followed by the physical system: for example, trying to rotate a valve at excessive speed. This
can produce undesired behavior of the closed-loop system, or even break actuators or other subsystems. Specific
control techniques are available to solve the problem: model predictive control (see later), and anti-wind up systems.
The latter consists of an additional control block that ensures that the control signal never exceeds a given threshold.

System classifications

Linear Systems control

For MIMO systems, pole placement can be performed mathematically using a state space representation of the
open-loop system and calculating a feedback matrix assigning poles in the desired positions. In complicated systems
this can require computer-assisted calculation capabilities, and cannot always ensure robustness. Furthermore, all
system states are not in general measured and so observers must be included and incorporated in pole placement

Nonlinear Systems control

Processes in industries like robotics and the aerospace industry typically have strong nonlinear dynamics. In control
theory it is sometimes possible to linearize such classes of systems and apply linear techniques, but in many cases it
can be necessary to devise from scratch theories permitting control of nonlinear systems. These, e.g., feedback
linearization, backstepping, sliding mode control, trajectory linearization control normally take advantage of results
based on Lyapunov's theory. Differential geometry has been widely used as a tool for generalizing well-known linear
control concepts to the non-linear case, as well as showing the subtleties that make it a more challenging problem.
Control theory 35

Decentralized Systems
When the system is controlled by multiple controllers, the problem is one of decentralized control. Decentralization
is helpful in many ways, for instance, it helps control systems operate over a larger geographical area. The agents in
decentralized control systems can interact using communication channels and coordinate their actions.

Main control strategies

Every control system must guarantee first the stability of the closed-loop behavior. For linear systems, this can be
obtained by directly placing the poles. Non-linear control systems use specific theories (normally based on
Aleksandr Lyapunov's Theory) to ensure stability without regard to the inner dynamics of the system. The possibility
to fulfill different specifications varies from the model considered and the control strategy chosen. Here a summary
list of the main control techniques is shown:
Adaptive control
Adaptive control uses on-line identification of the process parameters, or modification of controller gains,
thereby obtaining strong robustness properties. Adaptive controls were applied for the first time in the
aerospace industry in the 1950s, and have found particular success in that field.
Hierarchical control
A Hierarchical control system is a type of Control System in which a set of devices and governing software is
arranged in a hierarchical tree. When the links in the tree are implemented by a computer network, then that
hierarchical control system is also a form of Networked control system.
Intelligent control
Intelligent control uses various AI computing approaches like neural networks, Bayesian probability, fuzzy
logic, machine learning, evolutionary computation and genetic algorithms to control a dynamic system.
Optimal control
Optimal control is a particular control technique in which the control signal optimizes a certain "cost index":
for example, in the case of a satellite, the jet thrusts needed to bring it to desired trajectory that consume the
least amount of fuel. Two optimal control design methods have been widely used in industrial applications, as
it has been shown they can guarantee closed-loop stability. These are Model Predictive Control (MPC) and
Linear-Quadratic-Gaussian control (LQG). The first can more explicitly take into account constraints on the
signals in the system, which is an important feature in many industrial processes. However, the "optimal
control" structure in MPC is only a means to achieve such a result, as it does not optimize a true performance
index of the closed-loop control system. Together with PID controllers, MPC systems are the most widely
used control technique in process control.
Robust control
Robust control deals explicitly with uncertainty in its approach to controller design. Controllers designed using
robust control methods tend to be able to cope with small differences between the true system and the nominal
model used for design. The early methods of Bode and others were fairly robust; the state-space methods
invented in the 1960s and 1970s were sometimes found to lack robustness. A modern example of a robust
control technique is H-infinity loop-shaping developed by Duncan McFarlane and Keith Glover of Cambridge
University, United Kingdom. Robust methods aim to achieve robust performance and/or stability in the
presence of small modeling errors.
Stochastic control
Stochastic control deals with control design with uncertainty in the model. In typical stochastic control
problems, it is assumed that there exist random noise and disturbances in the model and the controller, and the
control design must take into account these random deviations.
Control theory 36

[1] Maxwell, J.C. (1867). "On Governors" (http:/ / links. jstor. org/ sici?sici=0370-1662(1867/ 1868)16<270:OG>2. 0. CO;2-1). Proceedings of
the Royal Society of London 16: 270–283. doi:10.1098/rspl.1867.0055. . Retrieved 2008-04-14.
[2] Routh, E.J.; Fuller, A.T. (1975). Stability of motion. Taylor & Francis.
[3] Routh, E.J. (1877). A Treatise on the Stability of a Given State of Motion, Particularly Steady Motion: Particularly Steady Motion. Macmillan
and co..
[4] Hurwitz, A. (1964). "On The Conditions Under Which An Equation Has Only Roots With Negative Real Parts". Selected Papers on
Mathematical Trends in Control Theory.
[5] Donald M Wiberg. State space & linear systems. Schaum's outline series. McGraw Hill.

Further reading
• Levine, William S., ed (1996). The Control Handbook. New York: CRC Press. ISBN 978-0-849-38570-4.
• Karl J. Åström and Richard M. Murray (2008). Feedback Systems: An Introduction for Scientists and Engineers.
( Princeton
University Press. ISBN 0691135762.
• Christopher Kilian (2005). Modern Control Technology. Thompson Delmar Learning. ISBN 1-4018-5806-6.
• Vannevar Bush (1929). Operational Circuit Analysis. John Wiley and Sons, Inc..
• Robert F. Stengel (1994). Optimal Control and Estimation. Dover Publications. ISBN 0-486-68200-5, ISBN
• Franklin et al. (2002). Feedback Control of Dynamic Systems (4 ed.). New Jersey: Prentice Hall.
ISBN 0-13-032393-4.
• Joseph L. Hellerstein, Dawn M. Tilbury, and Sujay Parekh (2004). Feedback Control of Computing Systems. John
Wiley and Sons. ISBN 0-47-126637-X, ISBN 978-0-471-26637-2.
• Diederich Hinrichsen and Anthony J. Pritchard (2005). Mathematical Systems Theory I - Modelling, State Space
Analysis, Stability and Robustness. Springer. ISBN 0-978-3-540-44125-0.
• Andrei, Neculai (2005). Modern Control Theory - A historical Perspective (
pdf). Retrieved 2007-10-10.
• Sontag, Eduardo (1998). Mathematical Control Theory: Deterministic Finite Dimensional Systems. Second
Edition (
pdf). Springer. ISBN 0-387-984895.
• Goodwin, Graham (2001). Control System Design. Prentice Hall. ISBN 0-13-958653-9.

External links
• Control Tutorials for Matlab ( - A set of worked through control
examples solved by several different methods.
• Control Tuning and Best Practices (
Feedback 37

Feedback describes the situation when output from (or information about the result of) an event or phenomenon in
the past will influence an occurrence or occurrences of the same (i.e. same defined) event / phenomenon (or the
continuation / development of the original phenomenon) in the present or future. When an event is part of a chain of
cause-and-effect that forms a circuit or loop, then the event is said to "feed back" into itself.
Feedback is also a synonym for:
• Feedback signal - the information about the initial event that is the basis for subsequent modification of the event
• Feedback loop - the causal path that leads from the initial generation of the feedback signal to the subsequent
modification of the event
• Audio feedback - the special kind of positive feedback that occurs when a loop exists between an audio input and

Feedback is a mechanism, process or signal that is looped back to control a system within itself. Such a loop is called
a feedback loop. In systems containing an input and output, feeding back part of the output so as to increase the input
is positive feedback (regeneration); feeding back part of the output in such a way as to partially oppose the input is
negative feedback (degeneration).
In more general terms, a control system has input from an external signal source and output to an external load; this
defines a natural sense (or direction) or path of propagation of signal; the feedforward sense or path describes the
signal propagation from input to output; feedback describes signal propagation in the reverse sense. When a sample
of the output of the system is fed back, in the reverse sense, by a distinct feedback path into the interior of the
system, to contribute to the input of one of its internal feedforward components, especially an active device or a
substance that is consumed in an irreversible reaction, it is called the "feedback". The propagation of the signal
around the feedback loop takes a finite time because it is causal.
The natural sense of feedforward is defined chemically by some irreversible reaction, or electronically by an active
circuit element that has access to an auxiliary power supply, so as to be able to provide power gain to amplify the
signal as it propagates from input to output. For example, an amplifier can use power from its controlled power
reservoir, such as its battery, to provide power gain to amplify the signal; but the reverse is not possible: the signal
cannot provide power to re-charge the battery of the amplifier.
Feedforward, feedback and regulation are self related. The feedforward carries the signal from source to load.
Negative feedback helps to maintain stability in a system in spite of external changes. It is related to homeostasis.
For example, in a population of foxes (predators) and rabbits (prey), an increase in the number of foxes will cause a
reduction in the number of rabbits; the smaller rabbit population will sustain fewer foxes, and the fox population will
fall back. In an electronic amplifier feeding back a negative copy of the output to the input will tend to cancel
distortion, making the output a more accurate replica of the input signal[1]
Positive feedback amplifies possibilities of divergences (evolution, change of goals); it is the condition to change,
evolution, growth; it gives the system the ability to access new points of equilibrium.
For example, in an organism, most positive feedback provides for fast autoexcitation of elements of endocrine and
nervous systems (in particular, in stress responses conditions) and are believed to play a key role in morphogenesis,
growth, and development of organs, all processes that are, in essence, a rapid escape from the initial state.
Homeostasis is especially visible in the nervous and endocrine systems when considered at organism level. Chemical
potential energy for irreversible reactions or electrical potential energy for irreversible cell-membrane current powers
the feedforward sense of the process. However, in the case of morphogenesis, feedback may only be enough to
explain the increase in momentum of the system, and may not be sufficient in itself to account for the movement or
Feedback 38

direction of its parts.

When a public-address system is used with a microphone to amplify speech, the output from a random sound at the
microphone may produce sound at a loudspeaker that reaches the microphone such as to reinforce and amplify the
original signal (positive feedback), building up to a howl (of frequency dependent upon the acoustics of the hall). A
similar process is used deliberately to produce oscillating electrical signals.
Feedback is distinctly different from reinforcement that occurs in learning, or in conditioned reflexes. Feedback
combines immediately with the immediate input signal to drive the responsive power gain element, without changing
the basic responsiveness of the system to future signals. Reinforcement changes the basic responsiveness of the
system to future signals, without combining with the immediate input signal. Reinforcement is a permanent change
in the responsiveness of the system to all future signals. Feedback is only transient, being limited by the duration of
the immediate signal.

Types of feedback
When feedback acts in response to an
event/phenomenon, it can influence the
input signal in one of two ways:
1. An in-phase feedback signal, where a
positive-going wave on the input leads to
a positive-going change on the output,
will amplify the input signal, leading to
more modification. This is known as
positive feedback. Figure 1: Ideal feedback model. The feedback is negative if B < 0

2. A feedback signal which is inverted,

where a positive-going change on the input leads to a negative-going change on the output, will dampen the effect
of the input signal, leading to less modification. This is known as negative feedback.
• Note that an increase or decrease of the feedback signal here refers to the magnitude relative to the input signal's
absolute value, without regard to the polarity or sign of the feedback signal. For example if the input signal
changes by 100 then a change in feedback signal value from +5 to +10 is a positive feedback. If the input signal
changes by -100 then a change in the feedback signal from -5 to -10 is also a positive feedback.
Positive feedback tends to increase the event that caused it, such as in a nuclear chain-reaction. It is also known as a
self-reinforcing loop.[2] An event influenced by positive feedback can increase or decrease its output/activation until
it hits a limiting constraint. Such a constraint may be destructive, as in thermal runaway or a nuclear chain reaction.
Self-reinforcing loops can be a smaller part of a larger balancing loop, especially in biological systems such as
regulatory circuits.
Negative feedback, which tends to reduce the input signal that caused it, is also known as a self-correcting or
balancing loop.[2] Such loops tend to be goal-seeking, as in a thermostat, which compares actual temperature with
desired temperature and seeks to reduce the difference. Balancing loops are sometimes prone to hunting: an
oscillation caused by an excessive or delayed negative feedback signal, resulting in over-correction, wherein the
signal becomes a positive feedback.
The terms negative and positive feedback can be used loosely or colloquially to describe or imply criticism and
praise, respectively. This may lead to confusion with the more technically accurate terms positive and negative
reinforcement, which refer to something that changes the likelihood of a future behaviour. Moreover, when used
technically, negative feedback leads to stability that is, in general, considered good, whereas positive feedback can
lead to unstable and explosive situations that are considered bad. Thus, when used colloquially, these terms imply
the opposite desirability to that when used technically.
Feedback 39

Negative feedback was applied by Harold Stephen Black to electrical amplifiers in 1927, but he could not get his
idea patented until 1937.[3] Arturo Rosenblueth, a Mexican researcher and physician, co-authored a seminal 1943
paper Behavior, Purpose and Teleology[4] that, according to Norbert Wiener (another co-author of the paper), set the
basis for the new science cybernetics. Rosenblueth proposed that behaviour controlled by negative feedback,
whether in animal, human or machine, was a determinative, directive principle in nature and human creations. This
kind of feedback is studied in cybernetics and control theory.


In biological systems such as organisms, ecosystems, or the biosphere, most parameters must stay under control
within a narrow range around a certain optimal level under certain environmental conditions. The deviation of the
optimal value of the controlled parameter can result from the changes in internal and external environments. A
change of some of the environmental conditions may also require change of that range to change for the system to
function. The value of the parameter to maintain is recorded by a reception system and conveyed to a regulation
module via an information channel. An example of this is Insulin oscillations.
Biological systems contain many types of regulatory circuits, both positive and negative. As in other contexts,
positive and negative do not imply consequences of the feedback have good or bad final effect. A negative feedback
loop is one that tends to slow down a process, whereas the positive feedback loop tends to accelerate it. The mirror
neurons are part of a social feedback system, when an observed action is "mirrored" by the brain - like a
self-performed action.
Feedback is also central to the operations of genes and gene regulatory networks. Repressor (see Lac repressor) and
activator proteins are used to create genetic operons, which were identified by Francois Jacob and Jacques Monod in
1961 as feedback loops. These feedback loops may be positive (as in the case of the coupling between a sugar
molecule and the proteins that import sugar into a bacterial cell), or negative (as is often the case in metabolic
Any self-regulating natural process involves feedback and/or is prone to hunting. A well-known example in ecology
is the oscillation of the population of snowshoe hares due to predation from lynxes.
In zymology, feedback serves as regulation of activity of an enzyme by its direct product(s) or downstream
metabolite(s) in the metabolic pathway (see Allosteric regulation).
Hypothalamo-pituitary-adrenal and gonadal axis is largely controlled by positive and negative feedback, much of
which is still unknown.
In psychology, the body receives a stimulus from the environment or internally that causes the release of hormones.
Release of hormones then may cause more of those hormones to be released, causing a positive feedback loop. This
cycle is also found in certain behaviour. For example, "shame loops" occur in persons who blush easily. When they
realize that they are blushing, they become even more embarrassed, which leads to further blushing, and so on.[5]
Feedback 40

Climate science
The climate system is characterized by strong positive and negative feedback loops between processes that affect the
state of the atmosphere, ocean, and land. A simple example is the ice-albedo positive feedback loop whereby melting
snow exposes more dark ground (of lower albedo), which in turn absorbs heat and causes more snow to melt.

Control theory
Feedback is extensively used in control theory, using a variety of methods including state space (controls), full state
feedback (also known as pole placement), and so forth.
The most common general-purpose controller using a control-loop feedback mechanism is a
proportional-integral-derivative (PID) controller. Heuristically, the terms of a PID controller can be interpreted as
corresponding to time: the proportional term depends on the present error, the integral term on the accumulation of
past errors, and the derivative term is a prediction of future error, based on current rate of change.[6]

Mechanical engineering
In ancient times, the float valve was used to regulate the flow of water in Greek and Roman water clocks; similar
float valves are used to regulate fuel in a carburettor and also used to regulate tank water level in the flush toilet.
In 1745, the windmill was improved with by blacksmith Edmund Lee who added a fantail to keep the face of the
windmill pointing into the wind. In 1787, Thomas Mead regulated the rotation speed of a windmill by using a
centrifugal pendulum to adjust the distance between the bedstone and the runner stone (i.e., to adjust the load).
The use of the centrifugal governor by James Watt in 1788 to regulate the speed of his steam engine was one factor
leading to the Industrial Revolution. Steam engines also use float valves and pressure release valves as mechanical
regulation devices. A mathematical analysis of Watt's governor was done by James Clerk Maxwell in 1868.
The Great Eastern was one of the largest steamships of its time and employed a steam powered rudder with feedback
mechanism designed in 1866 by J.McFarlane Gray. Joseph Farcot coined the word servo in 1873 to describe
steam-powered steering systems. Hydraulic servos were later used to position guns. Elmer Ambrose Sperry of the
Sperry Corporation designed the first autopilot in 1912. Nicolas Minorsky published a theoretical analysis of
automatic ship steering in 1922 and described the PID controller.
Internal combustion engines of the late 20th century employed mechanical feedback mechanisms such as the vacuum
timing advance but mechanical feedback was replaced by electronic engine management systems once small, robust
and powerful single-chip microcontrollers became affordable.

Electronic engineering
The main applications of feedback in electronics are in the designs of amplifiers, oscillators, and logic circuit
The processing and control of feedback is engineered into many electronic devices and may also be embedded in
other technologies.
If the signal is inverted on its way round the control loop, the system is said to have negative feedback; otherwise,
the feedback is said to be positive. Negative feedback is often deliberately introduced to increase the stability and
accuracy of a system by correcting unwanted changes. This scheme can fail if the input changes faster than the
system can respond to it. When this happens, the lag in arrival of the correcting signal results in unintended positive
feedback, causing the output to oscillate or hunt[7] Oscillation is usually an unwanted consequence of system
Harry Nyquist contributed the Nyquist plot for assessing the stability of feedback systems. An easier assessment, but
less general, is based upon gain margin and phase margin using Bode plots (contributed by Hendrik Bode). Design to
insure stability often involves frequency compensation, one method of compensation being pole splitting.
Feedback 41

The high-pitched squeal that sometimes occurs in audio systems, PA systems, and rock music is known as audio
feedback. If a microphone is in front of a speaker that it is connected to, the noise put into the microphone will come
out of the speaker. Since the microphone is in front of the speaker, the original sound (now coming from the speaker)
goes back into the microphone. This happens over and over, getting louder each time. This process produces the
Electronic feedback loops
They are used to control the output of electronic devices, such as amplifiers. A feedback loop is created when
all or some portion of the output from an electronic device is fed-back to the input. A device is said to be
operating open loop if no output feedback is being employed and closed loop if feedback is being used.
Electronic feedback loops take two forms: negative feedback loops and positive feedback loops.[8]
Negative feedback loops
They exist when the fed-back output signal is out of phase with the input signal. This occurs when the
fed-back signal is anywhere from 90° to 270° with respect to the input signal. Negative feedback is generally
used to correct output errors or to lower device output gain to a pre-determined level. In feedback amplifiers,
this correction is generally for waveform distortion reduction or to establish a specified gain level. A general
expression for the gain of a negative feedback amplifier is the asymptotic gain model.
Positive feedback loops
They occur when the fed-back signal is in phase with the input signal. Under certain gain conditions, positive
feedback reinforces the input signal to the point where the output of the device oscillates between its
maximum and minimum possible states. Positive feedback may also introduce hysteresis into a circuit. This
can cause the circuit to ignore small signals and respond only to large ones. It is sometimes used to eliminate
noise from a digital signal. Under some circumstances, positive feedback may cause a device to latch, i.e., to
reach a condition in which the output is locked to its maximum or minimum state.

Software engineering and computing systems

Feedback loops provide generic mechanisms for controlling the running, maintenance, and evolution of software and
computing systems.[9] Feedback-loops are important models in the engineering of adaptive software, as they define
the behaviour of the interactions among the control elements over the adaptation process, to guarantee system
properties at run-time. Feedback loops and foundations of control theory has been successfully applied to computing
systems.[10] In particular, they have been applied to the development of products such as IBM's Universal Database
server and IBM Tivoli. From a software perspective, the autonomic MAPE loop proposed by researchers of IBM is
another valuable contribution to the application of feedback loops to the control of dynamic properties and the design
and evolution of autonomic software systems.[11] [12]

Social sciences

Reflexive feedback
A sociological concept that states a feedback association is created within a certain relationship whereby the
subject/object that delivers a stimulus to a second subject/object, will in response receive the stimulus back. This first
impulse is reflected back and forth over and over again.

Economics and finance

A system prone to hunting (oscillating) is the stock market, which has both positive and negative feedback
mechanisms. This is due to cognitive and emotional factors belonging to the field of behavioural finance. For
Feedback 42

• When stocks are rising (a bull market), the belief that further rises are probable gives investors an incentive to buy
(positive feedback, see also stock market bubble); but the increased price of the shares, and the knowledge that
there must be a peak after which the market will fall, ends up deterring buyers (negative feedback).
• Once the market begins to fall regularly (a bear market), some investors may expect further losing days and
refrain from buying (positive feedback), but others may buy because stocks become more and more of a bargain
(negative feedback).
George Soros used the word reflexivity, to describe feedback in the financial markets and developed an investment
theory based on this principle.
The conventional economic equilibrium model of supply and demand supports only ideal linear negative feedback
and was heavily criticized by Paul Ormerod in his book "The Death of Economics", which, in turn, was criticized by
traditional economists. This book was part of a change of perspective as economists started to recognise that Chaos
Theory applied to nonlinear feedback systems including financial markets.

World-system development
The hyperbolic growth of the world population observed till the 1970s has recently been correlated to a non-linear
second-order positive feedback between the demographic growth and technological development that can be spelled
out as follows: technological growth - increase in the carrying capacity of land for people - demographic growth -
more people - more potential inventors - acceleration of technological growth - accelerating growth of the carrying
capacity - the faster population growth - accelerating growth of the number of potential inventors - faster
technological growth - hence, the faster growth of the Earth's carrying capacity for people, and so on.[13]

Young students will often look up to instructors as experts in the field and take to heart most of the things instructors
say. Thus, it is believed that spending a fair amount of time and effort thinking about how to respond to students may
be a worthwhile time investment. Sometimes the term "feedback" is used loosely or carelessly to refer to what is
more accurately called educational reinforcement. Some general types of reinforcement that can be used in many
types of student assessment are:

Confirmation Your answer was correct.

Corrective Your answer was incorrect. The correct answer was Jefferson.

Explanatory Your answer was incorrect because Carter was from Georgia; only Jefferson called Virginia home.

Diagnostic Your answer was incorrect. Your choice of Carter suggests some extra instruction on the home states of past presidents might be

Elaborative Your answer, Jefferson, was correct. The University of Virginia, a campus rich with Jeffersonian architecture and writings, is
sometimes referred to as "Mr. Jefferson's University".

(Adapted from Flemming and Levie.[14] )

A different application of feedback in education is the system for "continuous improvement" of engineering curricula
monitored by the Accreditation Board for Engineering and Technology (ABET)[15]
Feedback 43

Examples of feedback in government are:
• Elections
• Mass media
• Revolution
• Curfews
• Surveys

Email administration
A mechanism to alert the purported sender of an email with information about the email.

In organizations
As an organization seeks to improve its performance, feedback helps it to make required adjustments. Feedback
serves as motivation for many people in the work place. When one receives either negative or positive feedback,
they decide how they will apply it to his or her job. Joseph Folkman says that to find the greatest level of success in
an organization, working with other people, a person should learn how to accept any kind of feedback, analyze it in
the most positive manner possible, and use it to further impact future decision making.[16]
Examples of feedback in organizations:
• Financial audit
• Performance appraisal
• Shareholders' meeting
• Marketing research
• 360-degree feedback
• Walkout
• Lockout

[1] H.S. Black, "Stabilized feed-back amplifiers", Electrical Engineering, vol. 53, pp. 114-120, Jan. 1934.
[2] Peter M. Senge (1990). The Fifth Discipline: The Art and Practice of the Learning Organization. New York: Doubleday. pp. 424.
ISBN 0-385-260-946.
[3] Richard R Spencer & Ghausi MS (2003). Introduction to electronic circuit design (http:/ / worldcat. org/ isbn/ 0-201-36183-3). Upper Saddle
River NJ: Prentice Hall/Pearson Education. pp. 661. ISBN 0-201-36183-3. .
[4] Rosenblueth A, Wiener N & Bigelow J: Behavior, Purpose and Teleology (http:/ / links. jstor. org/
sici?sici=0031-8248(194301)10:1<18:BPAT>2. 0. CO;2-8)
[5] http:/ / www. psychologytoday. com/ blog/ lets-connect/ 200909/ the-emotionalrelational-world
[6] Araki, M., PID Control (http:/ / www. eolss. net/ ebooks/ Sample Chapters/ C18/ E6-43-03-03. pdf),
[7] With mechanical devices, hunting can be severe enough to destroy the device.
[8] P. Horowitz & W. Hill, The Art of Electronics, Cambridge University Press (1980), Chapter 3, relating to operational amplifiers.
[9] H. Giese, Y. Brun, J. D. M. Serugendo, C. Gacek, H. Kienle, H. Müller, M. Pezzè, and M. Shaw (2009). "Engineering self-adaptive and
self-managing systems". Springer-Verlag.
[10] J. L. Hellerstein, Y. Diao, S. Parekh, and D. M. Tilbury (2004). Feedback Control of Computing Systems. John Wiley & Sons.
[11] J. O. Kephart and D. M. Chess (2003). "The vision of autonomic computing".
[12] H. A. Müller, H. M. Kienle, and U. Stege, (2009). "Autonomic computing: Now you see it, now you don’t—design and evolution of
autonomic software systems".
[13] Introduction to Social Macrodynamics (http:/ / cliodynamics. ru/ index. php?option=com_content& task=view& id=172& Itemid=70) by
Andrey Korotayev et al.
[14] Fleming, M., & Levie, W.H. (1993). Instructional message design: principles from the behavioral and cognitive sciences (http:/ / books.
google. com/ books?id=vLxPAAAAMAAJ& dq=intitle:Instructional+ intitle:message+ intitle:design& lr=& as_brr=0& pgis=1) (Second
Edition ed.). Englewood Cliffs NJ: Educational Technology Publications. ISBN 0877782539. .
[15] Accreditation provides you with a structured mechanism to assess and improve the quality of your program: The two-loop feedback diagram
(http:/ / www. vanth. org/ curriculum/ twoloop. html)
Feedback 44

[16] Folkman, Joseph R. The Power of Feedback. 1st ed. New Jersey: John Wiley & Sons Inc., 2006. Print.

Further reading
• Katie Salen and Eric Zimmerman. Rules of Play. MIT Press. 2004. ISBN 0-262-24045-9. Chapter 18: Games as
Cybernetic Systems.
• Korotayev A., Malkov A., Khaltourina D. Introduction to Social Macrodynamics: Secular Cycles and Millennial
Trends. ( Moscow:
URSS, 2006. ISBN 5-484-00559-0
• Dijk, E., Cremer, D.D., Mulder, L.B., and Stouten, J. "How Do We React to Feedback in Social Dilemmas?" In
Biel, Eek, Garling & Gustafsson, (eds.), New Issues and Paradigms in Research on Social Dilemmas, New York:
Springer, 2008.

Instability in systems is generally characterized by some of the
outputs or internal states growing without bounds. Not all systems that
are not stable are unstable; systems can also be marginally stable or
exhibit limit cycle behavior.
In control theory, a system is unstable (or as sometimes referred to as
instable) if any of the roots of its characteristic equation has real part
greater than zero. This is equivalent to any of the eigenvalues of the A ball on the top of a hill is an unstable situation.

state matrix having real part greater than zero.

In structural engineering, a structure can become unstable when excessive load is applied. Beyond a certain
threshold, structural deflections magnify stresses, which in turn increases deflections. This can take the form of
buckling or crippling. The general field of study is called structural stability.

Fluid instabilities
Fluid instabilities occur in liquids, gases and plasmas, and are often
characterized by the shape that form; they are studied in fluid dynamics
and magnetohydrodynamics. Fluid instabilities include:
• Ballooning mode instability (some analogy to the Rayleigh–Taylor
instability); found in the magnetosphere
• Atmospheric instability
• Hydrodynamic instability or dynamic instability (atmospheric
• Inertial instability; baroclinic instability; symmetric
Hydrodynamics simulation of the
instability, conditional symmetric or convective symmetric Rayleigh–Taylor instability
instability; barotropic instability; Helmholtz or shearing
instability; rotational instability
• Hydrostatic instability or static instability/vertical instability (parcel instability), thermodynamic instability
(atmospheric thermodynamics)
Instability 45

• Conditional or static instability, buoyant instability, latent

instability, nonlocal static instability, conditional-symmetric
instability; convective, potential, or thermal instability,
convective instability of the first and second kind; absolute or
mechanical instability
• Bénard instability
• Drift mirror instability
• Kelvin–Helmholtz instability (similar, but different from the
diocotron instability in plasmas)
• Rayleigh–Taylor instability
• Plateau-Rayleigh instability (similar to the Rayleigh–Taylor
• Richtmyer-Meshkov instability (similar to the Rayleigh–Taylor

Plasma instabilities Unstable flow structure generated from the

collision of two impinging jets.
Plasma instabilities can be divided into two general groups (1)
hydrodynamic instabilities (2) kinetic instabilities. Plasma instabilities are also categorised into different modes - see
this paragraph in plasma stability.

Instabilities of stellar systems

Galaxies and star clusters can be unstable, if small perturbations in the gravitational potential cause changes in the
density that reinforce the original perturbation. Such instabilities usually require that the motions of stars be highly
correlated, so that the perturbation is not "smeared out" by random motions. After the instability has run its course,
the system is typically "hotter" (the motions are more random) or rounder than before. Instabilities in stellar systems
• Bar instability of rapidly-rotating disks
• Jeans instability
• Firehose instability[2]
• Gravothermal instability
• Radial-orbit instability
• Various instabilities in cold rotating disks

[1] Shengtai Li, Hui Li "Parallel AMR Code for Compressible MHD or HD Equations" (Los Alamos National Laboratory)
(http:/ / math. lanl. gov/ Research/ Highlights/ amrmhd. shtml)
[2] Merritt, D.; Sellwood, J. (1994), "Bending Instabilities of Stellar Systems" (http:/ / adsabs. harvard. edu/ abs/ 1994ApJ. . . 425. .
551M), The Astrophysical Journal 425: 551–567, doi:10.1086/174005,

External links
• eFluids Fluid Flow Image Gallery (
Oscillation 46

Oscillation is the repetitive variation, typically in time, of some measure about a
central value (often a point of equilibrium) or between two or more different states.
Familiar examples include a swinging pendulum and AC power. The term vibration
is sometimes used more narrowly to mean a mechanical oscillation but sometimes is
used to be synonymous with "oscillation". Oscillations occur not only in physical
systems but also in biological systems and in human society.

Simple harmonic oscillator

The simplest mechanical oscillating system is a mass attached to a linear spring
subject to no other forces. Such a system may be approximated on an air table or ice
surface. The system is in an equilibrium state when the spring is static. If the system
is displaced from the equilibrium, there is a net restoring force on the mass, tending
to bring it back to equilibrium. However, in moving the mass back to the equilibrium
position, it has acquired momentum which keeps it moving beyond that position,
establishing a new restoring force in the opposite sense. If a constant force such as
gravity is added to the system, the point of equilibrium is shifted. The time taken for
an oscillation to occur is often referred to as the oscillatory period.

The specific dynamics of this spring-mass system are described mathematically by

the simple harmonic oscillator and the regular periodic motion is known as simple
harmonic motion. In the spring-mass system, oscillations occur because, at the static
equilibrium displacement, the mass has kinetic energy which is converted into
potential energy stored in the spring at the extremes of its path. The spring-mass An undamped spring–mass
system illustrates some common features of oscillation, namely the existence of an system is an oscillatory system.

equilibrium and the presence of a restoring force which grows stronger the further
the system deviates from equilibrium.

Damped and driven oscillations

All real-world oscillator systems are thermodynamically irreversible. This means there are dissipative processes such
as friction or electrical resistance which continually convert some of the energy stored in the oscillator into heat in
the environment. This is called damping. Thus, oscillations tend to decay with time unless there is some net source
of energy into the system. The simplest description of this decay process can be illustrated by oscillation decay of the
harmonic oscillator.
In addition, an oscillating system may be subject to some external force, as when an AC circuit is connected to an
outside power source. In this case the oscillation is said to be driven.
Some systems can be excited by energy transfer from the environment. This transfer typically occurs where systems
are embedded in some fluid flow. For example, the phenomenon of flutter in aerodynamics occurs when an
arbitrarily small displacement of an aircraft wing (from its equilibrium) results in an increase in the angle of attack of
the wing on the air flow and a consequential increase in lift coefficient, leading to a still greater displacement. At
sufficiently large displacements, the stiffness of the wing dominates to provide the restoring force that enables an
Oscillation 47

Coupled oscillations
The harmonic oscillator and the systems it models have a single degree of freedom. More complicated systems have
more degrees of freedom, for example two masses and three springs (each mass being attached to fixed points and to
each other). In such cases, the behavior of each variable influences that of the others. This leads to a coupling of the
oscillations of the individual degrees of freedom. For example, two pendulum clocks (of identical frequency)
mounted on a common wall will tend to synchronise. This phenomenon was first observed by Christiaan Huygens in
1665.[1] The apparent motions of the compound oscillations typically appears very complicated but a more
economic, computationally simpler and conceptually deeper description is given by resolving the motion into normal
More special cases are the coupled oscillators where the energy alternates between two forms of oscillation.
Well-known is the Wilberforce pendulum, where the oscillation alternates between an elongation of a vertical spring
and the rotation of an object at the end of that spring.
Oscillation 48

Continuous systems – waves

As the number of degrees of freedom becomes arbitrarily large, a system approaches continuity; examples include a
string or the surface of a body of water. Such systems have (in the classical limit) an infinite number of normal
modes and their oscillations occur in the form of waves that can characteristically propagate.


Mechanical Economic and

• Double pendulum
Electro-mechanical social
• Foucault pendulum • Crystal oscillator • Business cycle
• Helmholtz resonator • Loudspeaker • Generation gap
• Neutron-star oscillations • Microphone • Malthusian economics
(helioseismology) and stars • News cycle
• Quantum harmonic oscillator Optical
• Playground swing
Climate and
• Laser (oscillation of electromagnetic field with frequency of
• String instruments geophysics
order 1015 Hz)
• Torsional vibration • Oscillator Toda or self-pulsation (pulsation of output power of • Atlantic multidecadal
• Tuning fork laser at frequencies 104 Hz – 106 Hz in the transient regime) oscillation
• Vibrating string • Quantum oscillator may refer to an optical local oscillator, as • Chandler wobble
• Wilberforce pendulum well as to a usual model in quantum optics. • El Niño-Southern
• Lever escapement
• Pacific decadal oscillation
Electrical • Quasi-biennial oscillation
• Circadian rhythm
• Alternating current • Circadian oscillator
• Armstrong (or Tickler or Meissner) Astrophysics
• Lotka–Volterra equation
oscillator • Neural oscillation • Neutron stars
• Astable multivibrator • Oscillating gene • Cyclic Model
• Blocking oscillator
• Butler oscillator
• Clapp oscillator Human Chemical
• Colpitts oscillator
• Neural oscillation • Belousov–Zhabotinsky
• Delay line oscillator
• Insulin release oscillations reaction
• Dow (or ultra-audion) oscillator
• gonadotropin releasing hormone pulsations • Mercury beating heart
• Electronic oscillator
• Pilot-induced oscillation • Briggs–Rauscher reaction
• Hartley oscillator
• Voice production • Bray–Liebhafsky reaction
• Oscillistor
• Pierce oscillator
• Relaxation oscillator
• RLC circuit
• Royer oscillator
• Vačkář oscillator
• Wien bridge oscillator
Oscillation 49

[1] Strogatz, Steven. Sync: The Emerging Science of Spontaneous Order. Hyperion, 2003, pp 106-109

External links
• Vibrations ( – a chapter from an online

Friction is the force resisting the relative motion of solid surfaces,
fluid layers, and/or material elements sliding against each other. It may
be thought of as the opposite of "slipperiness".
There are several types of friction:
• Dry friction resists relative lateral motion of two solid surfaces in
contact. Dry friction is subdivided into static friction between
non-moving surfaces, and kinetic friction between moving surfaces.
• Fluid friction describes the friction between layers within a viscous
fluid that are moving relative to each other.[1] [2]
• Lubricated friction is a case of fluid friction where a fluid
separates two solid surfaces.[3] [4] [5] Force diagram for block on ground. Arrows are
vectors indicating directions and magnitudes of
• Skin friction is a component of drag, the force resisting the motion
forces. W is the force of weight, N is the normal
of a solid body through a fluid. force, F is an applied force, and Ff is the force of
• Internal friction is the force resisting motion between the elements kinetic friction which is equal to the coefficient of
kinetic friction times the normal force. Since the
making up a solid material while it undergoes deformation.[2]
magnitude of the applied force is greater than the
When surfaces in contact move relative to each other, the friction magnitude of the force of kinetic friction
between the two surfaces converts kinetic energy into heat. This opposing it, the block is accelerating to the left.

property can have dramatic consequences, as illustrated by the use of

friction between pieces of wood to start a fire.
Another important consequence of many types of friction can be wear, which may lead to performance degradation
and/or damage to components. Friction is a component of the science of tribology.
Friction is not a fundamental force but occurs because of the electromagnetic forces between charged particles which
constitute the surfaces in contact. Because of the complexity of these interactions friction cannot be calculated from
first principles, but instead must be found empirically.
Friction 50

Several famous scientists and engineers contributed to our understanding of dry friction.[6] They include Leonardo da
Vinci, Guillaume Amontons, John Theophilus Desaguliers, Leonard Euler, and Charles-Augustin de Coulomb.
Nikolai Pavlovich Petrov and Osborne Reynolds later supplemented this understanding with theories of lubrication.

Basic properties
Basic properties of friction have been described as laws: [6]
• Amontons' 1st Law: The force of friction is directly proportional to the applied load.
• Amontons' 2nd Law: The force of friction is independent of the apparent area of contact.
• Coulomb's Law of Friction: Kinetic friction is independent of the sliding velocity.
Amontons' 2nd Law is an idealization assuming perfectly rigid and inelastic materials. For example, wider tires on
cars provide more traction than narrow tires for a given vehicle mass because of surface deformation of the tire.

Dry friction
Dry friction resists relative lateral motion of two solid surfaces in contact. The two regimes of dry friction are static
friction between non-moving surfaces, and kinetic friction (sometimes called sliding friction or dynamic friction)
between moving surfaces.
Coulomb friction, named after Charles-Augustin de Coulomb, is an approximate model used to calculate the force of
dry friction. It is governed by the equation:

• is the force exerted by friction (in the case of equality, the maximum possible magnitude of this force).
• is the coefficient of friction, which is an empirical property of the contacting materials,
• is the normal force exerted between the surfaces.
The Coulomb friction may take any value from zero up to , and the direction of the frictional force against
a surface is opposite to the motion that surface would experience in the absence of friction. Thus, in the static case,
the frictional force is exactly what it must be in order to prevent motion between the surfaces; it balances the net
force tending to cause such motion. In this case, rather than providing an estimate of the actual frictional force, the
Coulomb approximation provides a threshold value for this force, above which motion would commence. This
maximum force is known as traction.
The force of friction is always exerted in a direction that opposes movement (for kinetic friction) or potential
movement (for static friction) between the two surfaces. For example, a curling stone sliding along the ice
experiences a kinetic force slowing it down. For an example of potential movement, the drive wheels of an
accelerating car experience a frictional force pointing forward; if they did not, the wheels would spin, and the rubber
would slide backwards along the pavement. Note that it is not the direction of movement of the vehicle they oppose,
it is the direction of (potential) sliding between tire and road.
In the case of kinetic friction, the direction of the friction force may or may not match the direction of motion: a
block sliding atop a table with rectilinear motion is subject to friction directed along the line of motion; an
automobile making a turn is subject to friction acting perpendicular to the line of motion (in which case it is said to
be 'normal' to it). The direction of the static friction force can be visualized as directly opposed to the force that
would otherwise cause motion, were it not for the static friction preventing motion. In this case, the friction force
exactly cancels the applied force, so the net force given by the vector sum, equals zero. It is important to note that in
all cases, Newton's first law of motion holds.
Friction 51

The normal force

The normal force is defined as the net force compressing two parallel
surfaces together; and its direction is perpendicular to the surfaces. In the
simple case of a mass resting on a horizontal surface, the only component
of the normal force is the force due to gravity, where . In this
case, the magnitude of the friction force is the product of the mass of the
object, the acceleration due to gravity, and the coefficient of friction.
However, the coefficient of friction is not a function of mass or volume; it
depends only on the material. For instance, a large aluminum block has the
same coefficient of friction as a small aluminum block. However, the
magnitude of the friction force itself depends on the normal force, and
hence the mass of the block.

If an object is on a level surface and the force tending to cause it to slide is

horizontal, the normal force between the object and the surface is just
its weight, which is equal to its mass multiplied by the acceleration due to
earth's gravity, g. If the object is on a tilted surface such as an inclined
plane, the normal force is less, because less of the force of gravity is
perpendicular to the face of the plane. Therefore, the normal force, and
ultimately the frictional force, is determined using vector analysis, usually
via a free body diagram. Depending on the situation, the calculation of the
normal force may include forces other than gravity.

Coefficient of friction
Block on a ramp (top) and corresponding
The 'coefficient of friction' (COF), also known as a 'frictional coefficient' or free body diagram of just the block
'friction coefficient' and symbolized by the Greek letter µ, is a (bottom).

dimensionless scalar value which describes the ratio of the force of friction
between two bodies and the force pressing them together. The coefficient of friction depends on the materials used;
for example, ice on steel has a low coefficient of friction, while rubber on pavement has a high coefficient of friction.
Coefficients of friction range from near zero to greater than one – under good conditions, a tire on concrete may
have a coefficient of friction of 1.7.

For surfaces at rest relative to each other , where is the coefficient of static friction. This is usually
larger than its kinetic counterpart.
For surfaces in relative motion , where is the coefficient of kinetic friction. The Coulomb friction is
equal to , and the frictional force on each surface is exerted in the direction opposite to its motion relative to the
other surface.
The coefficient of friction is an empirical measurement – it has to be measured experimentally, and cannot be found
through calculations. Rougher surfaces tend to have higher effective values. Both static and kinetic coefficients of
friction depend on the pair of surfaces in contact; for a given pair of surfaces, the coefficient of static friction is
usually larger than that of kinetic friction; in some sets the two coefficients are equal, such as teflon-on-teflon.
Most dry materials in combination have friction coefficient values between 0.3 and 0.6. Values outside this range are
rarer, but teflon, for example, can have a coefficient as low as 0.04. A value of zero would mean no friction at all, an
elusive property – even magnetic levitation vehicles have drag. Rubber in contact with other surfaces can yield
friction coefficients from 1 to 2. Occasionally it is maintained that µ is always < 1, but this is not true. While in most
relevant applications µ < 1, a value above 1 merely implies that the force required to slide an object along the surface
is greater than the normal force of the surface on the object. For example, silicone rubber or acrylic rubber-coated
Friction 52

surfaces have a coefficient of friction that can be substantially larger than 1.

While it is often stated that the COF is a "material property," it is better categorized as a "system property." Unlike
true material properties (such as conductivity, dielectric constant, yield strength), the COF for any two materials
depends on system variables like temperature, velocity, atmosphere and also what are now popularly described as
aging and deaging times; as well as on geometric properties of the interface between the materials. For example, a
copper pin sliding against a thick copper plate can have a COF that varies from 0.6 at low speeds (metal sliding
against metal) to below 0.2 at high speeds when the copper surface begins to melt due to frictional heating. The latter
speed, of course, does not determine the COF uniquely; if the pin diameter is increased so that the frictional heating
is removed rapidly, the temperature drops, the pin remains solid and the COF rises to that of a 'low speed' test.

Approximate coefficients of friction

Materials Static friction,

Dry & clean Lubricated

Aluminum Steel 0.61

Copper Steel 0.53

Brass Steel 0.51

Cast iron Copper 1.05

Cast iron Zinc 0.85

Concrete (wet) Rubber 0.30

Concrete (dry) Rubber 1.0

Concrete Wood [7]


Copper Glass 0.68

Glass Glass 0.94

Metal Wood [7] [7]

0.2–0.6 0.2 (wet)

Polythene Steel [8] [8]

0.2 0.2

Steel Steel [8] [8]

0.80 0.16

Steel Teflon [8] [8]

0.04 0.04

Teflon Teflon [8] [8]

0.04 0.04

Wood Wood [7] [7]

0.25–0.5 0.2 (wet)

The slipperiest solid known, discovered in 1999, dubbed BAM (for the elements boron, aluminum, and magnesium),
has an approximate coefficient of friction of 0.02, about half that of Teflon.[9]
Friction 53

Static friction
Static friction is friction between two solid objects that are not moving relative to each other. For example, static
friction can prevent an object from sliding down a sloped surface. The coefficient of static friction, typically denoted
as μs, is usually higher than the coefficient of kinetic friction.
The static friction force must be overcome by an applied force before an object can move. The maximum possible
friction force between two surfaces before sliding begins is the product of the coefficient of static friction and the
normal force: . When there is no sliding occurring, the friction force can have any value from zero up to
. Any force smaller than attempting to slide one surface over the other is opposed by a frictional force
of equal magnitude and opposite direction. Any force larger than overcomes the force of static friction and
causes sliding to occur. The instant sliding occurs, static friction is no longer applicable—the friction between the
two surfaces is then called kinetic friction.
An example of static friction is the force that prevents a car wheel from slipping as it rolls on the ground. Even
though the wheel is in motion, the patch of the tire in contact with the ground is stationary relative to the ground, so
it is static rather than kinetic friction.
The maximum value of static friction, when motion is impending, is sometimes referred to as limiting friction,[10]
although this term is not used universally.[1] It is also known as traction.

Kinetic friction
Kinetic (or dynamic) friction occurs when two objects are moving relative to each other and rub together (like a sled
on the ground). The coefficient of kinetic friction is typically denoted as μk, and is usually less than the coefficient of
static friction for the same materials.[11] [12] In fact, Richard Feynman reports that "with dry metals it is very hard to
show any difference."[13]
New models are beginning to show how kinetic friction can be greater than static friction.[14] Contrary to earlier
explanations, kinetic friction is now understood not to be caused by surface roughness but by chemical bonding
between the surfaces.[15] Surface roughness and contact area, however, do affect kinetic friction for micro- and
nano-scale objects where surface area forces dominate inertial forces.[16]

Angle of friction
For certain applications it is more useful to define static friction in terms of the maximum angle before which one of
the items will begin sliding. This is called the angle of friction or friction angle. It is defined as:

where θ is the angle from horizontal and µ is the static coefficient of friction between the objects.[17] This formula
can also be used to calculate µ from empirical measurements of the friction angle.

Friction at the atomic level

Determining the forces required to move atoms past each other is a challenge in designing nanomachines. In 2008
scientists for the first time were able to move a single atom across a surface, and measure the forces required. Using
ultrahigh vacuum and nearly-zero temperature (5 K), a modified atomic force microscope was used to drag a cobalt
atom, and a carbon monoxide molecule, across surfaces of copper and platinum.[18]

Limitations of the Coulomb model

The Coulomb approximation mathematically follows from the assumptions that surfaces are in atomically close
contact only over a small fraction of their overall area, that this contact area is proportional to the normal force (until
saturation, which takes place when all area is in atomic contact), and that frictional force is proportional to the
applied normal force, independently of the contact area (you can see the experiments on friction from Leonardo Da
Friction 54

Vinci). Such reasoning aside, however, the approximation is fundamentally an empirical construction. It is a rule of
thumb describing the approximate outcome of an extremely complicated physical interaction. The strength of the
approximation is its simplicity and versatility – though in general the relationship between normal force and
frictional force is not exactly linear (and so the frictional force is not entirely independent of the contact area of the
surfaces), the Coulomb approximation is an adequate representation of friction for the analysis of many physical
When the surfaces are conjoined, Coulomb friction becomes a very poor approximation (for example, adhesive tape
resists sliding even when there is no normal force, or a negative normal force). In this case, the frictional force may
depend strongly on the area of contact. Some drag racing tires are adhesive in this way. However, despite the
complexity of the fundamental physics behind friction, the relationships are accurate enough to be useful in many

Fluid friction
Fluid friction occurs between layers within a fluid that are moving relative to each other. This internal resistance to
flow is described by viscosity. In everyday terms viscosity is "thickness". Thus, water is "thin", having a lower
viscosity, while honey is "thick", having a higher viscosity. Put simply, the less viscous the fluid is, the greater its
ease of movement.
All real fluids (except superfluids) have some resistance to stress and therefore are viscous, but a fluid which has no
resistance to shear stress is known as an ideal fluid or inviscid fluid.

Lubricated friction
Lubricated friction is a case of fluid friction where a fluid separates two solid surfaces. Lubrication is a technique
employed to reduce wear of one or both surfaces in close proximity moving relative to each another by interposing a
substance called a lubricant between the surfaces.
In most cases the applied load is carried by pressure generated within the fluid due to the frictional viscous resistance
to motion of the lubricating fluid between the surfaces. Adequate lubrication allows smooth continuous operation of
equipment, with only mild wear, and without excessive stresses or seizures at bearings. When lubrication breaks
down, metal or other components can rub destructively over each other, causing destructive damage, heat, and

Skin friction
Skin friction arises from the friction of the fluid against the "skin" of the object that is moving through it. Skin
friction arises from the interaction between the fluid and the skin of the body, and is directly related to the area of the
surface of the body that is in contact with the fluid. Skin friction follows the drag equation and rises with the square
of the velocity.
Skin friction is caused by viscous drag in the boundary layer around the object. There are two ways to decrease skin
friction: the first is to shape the moving body so that smooth flow is possible, like an airfoil. The second method is to
decrease the length and cross-section of the moving object as much as is practicable.
Friction 55

Internal friction
Internal friction is the force resisting motion between the elements making up a solid material while it undergoes
plastic deformation.
Plastic deformation in solids is an irreversible change in the internal molecular structure of an object. This change
may be due to either (or both) an applied force or a change in temperature. The change of an object's shape is called
strain. The force causing it is called stress. Stress does not necessarily cause permanent change. As deformation
occurs, internal forces oppose the applied force. If the applied stress is not too large these opposing forces may
completely resist the applied force, allowing the object to assume a new equilibrium state and to return to its original
shape when the force is removed. This is what is known in the literature as elastic deformation (or elasticity). Larger
forces in excess of the elastic limit may cause a permanent (irreversible) deformation of the object. This is what is
known as plastic deformation.

Other types of friction

Rolling resistance
Rolling resistance is the force that resists the rolling of a wheel or other circular object along a surface caused by
deformations in the object and/or surface. Generally the force of rolling resistance is less than that associated with
kinetic friction.[19] Typical values for the coefficient of rolling resistance are 0.001.[20] One of the most common
examples of rolling resistance is the movement of motor vehicle tires on a road, a process which generates heat and
sound as by-products.[21]

Triboelectric effect
Rubbing dissimilar materials against one another can cause a build-up of electrostatic charge, which can be
hazardous if flammable gases or vapours are present. When the static build-up discharges, explosions can be caused
by ignition of the flammable mixture.

Belt friction
Belt friction is a physical property observed from the forces acting on a belt wrapped around a pulley, when one end
is being pulled. The resulting tension, which acts on both ends of the belt, can be modeled by the belt friction
In practice, the theoretical tension acting on the belt or rope calculated by the belt friction equation can be compared
to the maximum tension the belt can support. This helps a designer of such a rig to know how many times the belt or
rope must be wrapped around the pulley to prevent it from slipping. Mountain climbers and sailing crews
demonstrate a standard knowledge of belt friction when accomplishing basic tasks.

Reducing friction

Devices such as wheels, ball bearings, roller bearings, and air cushion or other types of fluid bearings can change
sliding friction into a much smaller type of rolling friction.
Many thermoplastic materials such as nylon, HDPE and PTFE are commonly used in low friction bearings. They are
especially useful because the coefficient of friction falls with increasing imposed load. For improved wear resistance,
very high molecular weight grades are usually specified for heavy duty or critical bearings.
Friction 56

A common way to reduce friction is by using a lubricant, such as oil, water, or grease, which is placed between the
two surfaces, often dramatically lessening the coefficient of friction. The science of friction and lubrication is called
tribology. Lubricant technology is when lubricants are mixed with the application of science, especially to industrial
or commercial objectives.
Superlubricity, a recently-discovered effect, has been observed in graphite: it is the substantial decrease of friction
between two sliding objects, approaching zero levels. A very small amount of frictional energy would still be
Lubricants to overcome friction need not always be thin, turbulent fluids or powdery solids such as graphite and talc;
acoustic lubrication actually uses sound as a lubricant.
Another way to reduce friction between two parts is to superimpose micro-scale vibration to one of the parts. This
can be sinusoidal vibration as used in ultrasound-assisted cutting or vibration noise, known as dither.

Energy of friction
According to the law of conservation of energy, no energy is destroyed due to friction, though it may be lost to the
system of concern. Energy is transformed from other forms into heat. A sliding hockey puck comes to rest because
friction converts its kinetic energy into heat. Since heat quickly dissipates, many early philosophers, including
Aristotle, wrongly concluded that moving objects lose energy without a driving force.
When an object is pushed along a surface, the energy converted to heat is given by:

is the normal force,
is the coefficient of kinetic friction,
is the coordinate along which the object transverses.
Energy lost to a system as a result of friction is a classic example of thermodynamic irreversibility.

Work of friction
In the reference frame of the interface between two surfaces, static friction does no work, because there is never
displacement between the surfaces. In the same reference frame, kinetic friction is always in the direction opposite
the motion, and does negative work.[22] However, friction can do positive work in certain frames of reference. One
can see this by placing a heavy box on a rug, then pulling on the rug quickly. In this case, the box slides backwards
relative to the rug, but moves forward relative to the frame of reference in which the floor is stationary. Thus, the
kinetic friction between the box and rug accelerates the box in the same direction that the box moves, doing positive
The work done by friction can translate into deformation, wear, and heat that can affect the contact surface properties
(even the coefficient of friction between the surfaces). This can be beneficial as in polishing. The work of friction is
used to mix and join materials such as in the process of friction welding. Excessive erosion or wear of mating
surfaces occur when work due frictional forces rise to unacceptable levels. Harder corrosion particles caught between
mating surfaces (fretting) exacerbates wear of frictional forces. Bearing seizure or failure may result from excessive
wear due to work of friction. As surfaces are worn by work due to friction, fit and surface finish of an object may
degrade until it no longer functions properly.[24]
Friction 57

Friction is an important factor in many engineering disciplines.

• Rail adhesion refers to the grip wheels of a train have on the rails.
• Road slipperiness is an important design and safety factor for automobiles
• Split friction is a particularly dangerous condition arising due to varying friction on either side of a car.
• Road texture affects the interaction of tires and the driving surface.

• A tribometer is an instrument that measures friction on a surface.
• A profilograph is a device used to measure pavement surface roughness.

[1] Beer, Ferdinand P.; E. Russel Johnston, Jr. (1996). Vector Mechanics for Engineers (Sixth ed.). McGraw-Hill. p. 397. ISBN 0072976888.
[2] Meriam, J. L.; L. G. Kraige (2002). Engineering Mechanics (fifth ed.). John Wiley & Sons. p. 328. ISBN 0471602930.
[3] Ruina, Andy; Rudra Pratap (2002). Introduction to Statics and Dynamics (http:/ / ruina. tam. cornell. edu/ Book/ RuinaPratapNoProblems.
pdf). Oxford University Press. p. 713. .
[4] Hibbeler, R. C. (2007). Engineering Mechanics (Eleventh ed.). Pearson, Prentice Hall. p. 393. ISBN 0131271466.
[5] Soutas-Little, Robert W.; Inman, Balint (2008). Engineering Mechanics. Thomson. p. 329. ISBN 0495296104.
[6] "Introduction to Tribology – Friction" (http:/ / depts. washington. edu/ nanolab/ ChemE554/ Summaries ChemE 554/ Introduction Tribology.
htm). . Retrieved 2008-12-21.
[7] Engineers: Handbook Friction Coefficients (http:/ / www. engineershandbook. com/ Tables/ frictioncoefficients. htm)
[8] "The Engineering Toolbox: Friction and Coefficients of Friction" (http:/ / www. engineeringtoolbox. com/ friction-coefficients-d_778. html).
. Retrieved 2008-11-23.
[9] Kurt Kleiner (2008-11-21). "Material slicker than Teflon discovered by accident" (http:/ / www. newscientist. com/ article/
dn16102-material-slicker-than-teflon-discovered-by-accident. html). . Retrieved 2008-12-25.
[10] Bhavikatti, S. S.; K. G. Rajashekarappa (1994). Engineering Mechanics (http:/ / books. google. com/ ?id=4wkLl4NvmWAC& pg=PA112&
lpg=PA112). New Age International. p. 112. ISBN 9788122406177. . Retrieved 2007-10-21.
[11] Sheppard and Tongue (2005). Statics: Analysis and Design of Systems in Equilibrium. Wiley and Sons. p. 618. ISBN 0471372994. "In
general, for given contacting surfaces, μk < μs"
[12] Meriam and Kraige (2002). Engineering Mechanics: Statics. Wiley and Sons. p. 330. ISBN 0471406465. "Kinetic friction force is usually
somewhat less than the maximum static friction force."
[13] Feynman, Leighton, and Sands (1964). "The Feynman Lectures on Physics, Vol. I, p. 12-5" (http:/ / hyperphysics. phy-astr. gsu. edu/ hbase/
frict2. html). Addison-Wesley. . Retrieved 2009-10-16.
[14] Persson, B. N.; Volokitin, A. I (2002). "Theory of rubber friction: Nonstationary sliding" (http:/ / juwel. fz-juelich. de:8080/ dspace/
bitstream/ 2128/ 1343/ 1/ 17249. pdf). Physical Review B 65: 134106. doi:10.1103/PhysRevB.65.134106. .
[15] Beatty, William J.. "Recurring science misconceptions in K-6 textbooks" (http:/ / amasci. com/ miscon/ miscon4. html#fric). . Retrieved
[16] Persson, Bo N. J., Sliding Friction
[17] Nichols, Edward Leamington; Franklin, William Suddards (1898). The Elements of Physics (http:/ / books. google. com/
?id=8IlCAAAAIAAJ). 1. Macmillan. p. 101. .
[18] Ternes, Markus; Lutz, Christopher P.; Hirjibehedin, Cyrus F.; Giessibl, Franz J.; Heinrich, Andreas J. (2008-02-22). "The Force Needed to
Move an Atom on a Surface". Science 319 (5866): 1066–1069. doi:10.1126/science.1150288. PMID 18292336.
[19] Benjamin Silliman, Principles of Physics, Or Natural Philosophy, Ivison, Blakeman, Taylor & company publishers (1871)
[20] Hans-Jürgen Butt, Karlheinz Graf, Michael Kappl, Physics and Chemistry of Interfaces, Wiley, ISBN 3527404139 (2006)
[21] Hogan, C. Michael (1973). "Analysis of highway noise". Water, Air, and Soil Pollution 2: 387. doi:10.1007/BF00159677.
[22] Den Hartog, J. P. (1961). Mechanics (http:/ / books. google. com/ ?id=WRXrtu44W9UC). Courier Dover Publications. p. 142.
ISBN 0486607542. .
[23] Leonard, William J (2000). Minds-on Physics (http:/ / books. google. com/ ?id=t_AKvmza5s8C& pg=PA603). Kendall/Hunt. p. 603.
ISBN 0-7872-3932-1. .
[24] Bayer, Raymond George (2004). Mechanical wear (http:/ / books. google. com/ ?id=Q64Kq2HlyucC& pg=PA3& lpg=PA3& dq=Physical+
wear+ is+ associated+ with+ friction). CRC Press. pp. 1, 2. ISBN 0824746201. . Retrieved 2008-07-07.
Friction 58

External links
• Coefficients of Friction ( – tables of
coefficients, plus many links
• Physclips: Mechanics with animations and video clips ( from the
University of New South Wales
• CRC Handbook of Chemistry & Physics – Values for Coefficient of Friction (
• Characteristic Phenomena in Conveyor Chain (
• Atomic-scale Friction Research and Education Synergy Hub (AFRESH) ( an Engineering
Virtual Organization for the atomic-scale friction community to share, archive, link, and discuss data, knowledge
and tools related to atomic-scale friction.

Muscle (from Latin musculus, diminutive of
mus "mouse"[1] ) is the contractile tissue of
animals and is derived from the mesodermal
layer of embryonic germ cells. Muscle cells
contain contractile filaments that move past
each other and change the size of the cell.
They are classified as skeletal, cardiac, or
smooth muscles. Their function is to
produce force and cause motion. Muscles
can cause either locomotion of the organism
itself or movement of internal organs.
Cardiac and smooth muscle contraction
occurs without conscious thought and is
necessary for survival. Examples are the
contraction of the heart and peristalsis which A top-down view of skeletal muscle
pushes food through the digestive system.
Voluntary contraction of the skeletal muscles is used to move the body and can be finely controlled. Examples are
movements of the eye, or gross movements like the quadriceps muscle of the thigh. There are two broad types of
voluntary muscle fibers: slow twitch and fast twitch. Slow twitch fibers contract for long periods of time but with
little force while fast twitch fibers contract quickly and powerfully but fatigue very rapidly.

Muscles are predominately powered by the oxidation of fats and carbohydrates, but anaerobic chemical reactions are
also used, particularly by fast twitch fibers. These chemical reactions produce adenosine triphosphate (ATP)
molecules which are used to power the movement of the myosin heads.
Muscle 59

All muscles derive from paraxial mesoderm.[2] The paraxial mesoderm is divided along the embryo's length into
somites, corresponding to the segmentation of the body (most obviously seen in the vertebral column.[2] Each somite
has 3 divisions, sclerotome (which forms vertebrae), dermatome (which forms skin), and myotome (which forms
muscle).[2] The myotome is divided into two sections, the epimere and hypomere, which form epaxial and hypaxial
muscles, respectively.[2] Epaxial muscles in humans are only the erector spinae and small intervertebral muscles, and
are innervated by the dorsal rami of the spinal nerves.[2] All other muscles, including limb muscles, are hypaxial
muscles, formed from the hypomere, and inervated by the ventral rami of the spinal nerves.[2]
During development, myoblasts (muscle progenitor cells) either remain in the somite to form muscles associated
with the vertebral column or migrate out into the body to form all other muscles.[2] Myoblast migration is preceded
by the formation of connective tissue frameworks, usually formed from the somatic lateral plate mesoderm.[2]
Myoblasts follow chemical signals to the appropriate locations, where they fuse into elongate skeletal muscle cells.[2]

There are three types of muscle:
• Skeletal muscle or "voluntary
muscle" is anchored by tendons (or
by aponeuroses at a few places) to
bone and is used to effect skeletal
movement such as locomotion and Types of muscle (shown at different magnifications)
in maintaining posture. Though this
postural control is generally maintained as a subconscious reflex, the muscles responsible react to conscious
control like non-postural muscles. An average adult male is made up of 42% of skeletal muscle and an average
adult female is made up of 36% (as a percentage of body mass).[3]
• Smooth muscle or "involuntary muscle" is found within the walls of organs and structures such as the esophagus,
stomach, intestines, bronchi, uterus, urethra, bladder, blood vessels, and the arrector pili in the skin (in which it
controls erection of body hair). Unlike skeletal muscle, smooth muscle is not under conscious control.
• Cardiac muscle is also an "involuntary muscle" but is more akin in structure to skeletal muscle, and is found only
in the heart.
Cardiac and skeletal muscles are "striated" in that they contain sarcomeres and are packed into highly regular
arrangements of bundles; smooth muscle has neither. While skeletal muscles are arranged in regular, parallel
bundles, cardiac muscle connects at branching, irregular angles (called intercalated discs). Striated muscle contracts
and relaxes in short, intense bursts, whereas smooth muscle sustains longer or even near-permanent contractions.
Skeletal muscle is further divided into several subtypes:
• Type I, slow oxidative, slow twitch, or "red" muscle is dense with capillaries and is rich in mitochondria and
myoglobin, giving the muscle tissue its characteristic red color. It can carry more oxygen and sustain aerobic
• Type II, fast twitch muscle, has three major kinds that are, in order of increasing contractile speed:[4]
• Type IIa, which, like slow muscle, is aerobic, rich in mitochondria and capillaries and appears red.
• Type IIx (also known as type IId), which is less dense in mitochondria and myoglobin. This is the fastest
muscle type in humans. It can contract more quickly and with a greater amount of force than oxidative muscle,
but can sustain only short, anaerobic bursts of activity before muscle contraction becomes painful (often
incorrectly attributed to a build-up of lactic acid). N.B. in some books and articles this muscle in humans was,
confusingly, called type IIB.[5]
Muscle 60

• Type IIb, which is anaerobic, glycolytic, "white" muscle that is even less dense in mitochondria and
myoglobin. In small animals like rodents this is the major fast muscle type, explaining the pale color of their

The anatomy of muscles includes both gross anatomy, comprising all the muscles of an organism, and, on the other
hand, microanatomy, which comprises the structures of a single muscle.

Gross anatomy

Muscles, anterior view (See Gray's muscle Muscles, posterior view (See Gray's muscle
pictures for detailed pictures) pictures for detailed pictures)

The gross anatomy of a muscle is the most important indicator of its role in the body. The action a muscle generates
is determined by the origin and insertion locations. The cross-sectional area of a muscle (rather than volume or
length) determines the amount of force it can generate by defining the number of sarcomeres which can operate in
parallel. The amount of force applied to the external environment is determined by lever mechanics, specifically the
ratio of in-lever to out-lever. For example, moving the insertion point of the biceps more distally on the radius
(farther from the joint of rotation) would increase the force generated during flexion (and, as a result, the maximum
weight lifted in this movement), but decrease the maximum speed of flexion. Moving the insertion point proximally
(closer to the joint of rotation) would result in decreased force but increased velocity. This can be most easily seen
by comparing the limb of a mole to a horse - in the former, the insertion point is positioned to maximize force (for
digging), while in the latter, the insertion point is positioned to maximize speed (for running).
One particularly important aspect of gross anatomy of muscles is pennation or lack thereof. In most muscles, all the
fibers are oriented in the same direction, running in a line from the origin to the insertion. In pennate muscles, the
individual fibers are oriented at an angle relative to the line of action, attaching to the origin and insertion tendons at
each end. Because the contracting fibers are pulling at an angle to the overall action of the muscle, the change in
length is smaller, but this same orientation allows for more fibers (thus more force) in a muscle of a given size.
Pennate muscles are usually found where their length change is less important than maximum force, such as the
rectus femoris.
There are approximately 639 skeletal muscles in the human body. However, the exact number is difficult to define
because different sources group muscles differently and some muscles, such as palmaris longus, are variably present
in humans.
Muscle 61

Muscle is mainly composed of muscle cells. Within the cells are myofibrils; myofibrils contain sarcomeres, which
are composed of actin and myosin. Individual muscle fibres are surrounded by endomysium. Muscle fibers are
bound together by perimysium into bundles called fascicles; the bundles are then grouped together to form muscle,
which is enclosed in a sheath of epimysium. Muscle spindles are distributed throughout the muscles and provide
sensory feedback information to the central nervous system.
Skeletal muscle is arranged in discrete muscles, an example of which is the biceps brachii. It is connected by tendons
to processes of the skeleton. Cardiac muscle is similar to skeletal muscle in both composition and action, being made
up of myofibrils of sarcomeres, but anatomically different in that the muscle fibers are typically branched like a tree
and connect to other cardiac muscle fibers through intercalcated discs, and form the appearance of a syncytium.

The three types of muscle (skeletal, cardiac and smooth) have significant differences. However, all three use the
movement of actin against myosin to create contraction. In skeletal muscle, contraction is stimulated by electrical
impulses transmitted by the nerves, the motor nerves and motoneurons in particular. Cardiac and smooth muscle
contractions are stimulated by internal pacemaker cells which regularly contract, and propagate contractions to other
muscle cells they are in contact with. All skeletal muscle and many smooth muscle contractions are facilitated by the
neurotransmitter acetylcholine.
Muscular activity accounts for much of the body's energy consumption. All muscle cells produce adenosine
triphosphate (ATP) molecules which are used to power the movement of the myosin heads. Muscles conserve energy
in the form of creatine phosphate which is generated from ATP and can regenerate ATP when needed with creatine
kinase. Muscles also keep a storage form of glucose in the form of glycogen. Glycogen can be rapidly converted to
glucose when energy is required for sustained, powerful contractions. Within the voluntary skeletal muscles, the
glucose molecule can be metabolized anaerobically in a process called glycolysis which produces two ATP and two
lactic acid molecules in the process (note that in aerobic conditions, lactate is not formed; instead pyruvate is formed
and transmitted through the citric acid cycle). Muscle cells also contain globules of fat, which are used for energy
during aerobic exercise. The aerobic energy systems take longer to produce the ATP and reach peak efficiency, and
requires many more biochemical steps, but produces significantly more ATP than anaerobic glycolysis. Cardiac
muscle on the other hand, can readily consume any of the three macronutrients (protein, glucose and fat) aerobically
without a 'warm up' period and always extracts the maximum ATP yield from any molecule involved. The heart,
liver and red blood cells will also consume lactic acid produced and excreted by skeletal muscles during exercise.

Nervous control

Efferent leg
The efferent leg of the peripheral nervous system is responsible for conveying commands to the muscles and glands,
and is ultimately responsible for voluntary movement. Nerves move muscles in response to voluntary and autonomic
(involuntary) signals from the brain. Deep muscles, superficial muscles, muscles of the face and internal muscles all
correspond with dedicated regions in the primary motor cortex of the brain, directly anterior to the central sulcus that
divides the frontal and parietal lobes.
In addition, muscles react to reflexive nerve stimuli that do not always send signals all the way to the brain. In this
case, the signal from the afferent fiber does not reach the brain, but produces the reflexive movement by direct
connections with the efferent nerves in the spine. However, the majority of muscle activity is volitional, and the
result of complex interactions between various areas of the brain.
Muscle 62

Nerves that control skeletal muscles in mammals correspond with neuron groups along the primary motor cortex of
the brain's cerebral cortex. Commands are routed though the basal ganglia and are modified by input from the
cerebellum before being relayed through the pyramidal tract to the spinal cord and from there to the motor end plate
at the muscles. Along the way, feedback, such as that of the extrapyramidal system contribute signals to influence
muscle tone and response.
Deeper muscles such as those involved in posture often are controlled from nuclei in the brain stem and basal

Afferent leg
The afferent leg of the peripheral nervous system is responsible for conveying sensory information to the brain,
primarily from the sense organs like the skin. In the muscles, the muscle spindles convey information about the
degree of muscle length and stretch to the central nervous system to assist in maintaining posture and joint position.
The sense of where our bodies are in space is called proprioception, the perception of body awareness. More easily
demonstrated than explained, proprioception is the "unconscious" awareness of where the various regions of the
body are located at any one time. This can be demonstrated by anyone closing their eyes and waving their hand
around. Assuming proper proprioceptive function, at no time will the person lose awareness of where the hand
actually is, even though it is not being detected by any of the other senses.
Several areas in the brain coordinate movement and position with the feedback information gained from
proprioception. The cerebellum and red nucleus in particular continuously sample position against movement and
make minor corrections to assure smooth motion.

Exercise is often recommended as a means of improving motor skills, fitness, muscle and bone strength, and joint
function. Exercise has several effects upon muscles, connective tissue, bone, and the nerves that stimulate the
muscles. One such effect is muscle hypertrophy, an increase in size. This is used in bodybuilding.
Various exercises require a predominance of certain muscle fiber utilization over another. Aerobic exercise involves
long, low levels of exertion in which the muscles are used at well below their maximal contraction strength for long
periods of time (the most classic example being the marathon). Aerobic events, which rely primarily on the aerobic
(with oxygen) system, use a higher percentage of Type I (or slow-twitch) muscle fibers, consume a mixture of fat,
protein and carbohydrates for energy, consume large amounts of oxygen and produce little lactic acid. Anaerobic
exercise involves short bursts of higher intensity contractions at a much greater percentage of their maximum
contraction strength. Examples of anaerobic exercise include sprinting and weight lifting. The anaerobic energy
delivery system uses predominantly Type II or fast-twitch muscle fibers, relies mainly on ATP or glucose for fuel,
consumes relatively little oxygen, protein and fat, produces large amounts of lactic acid and can not be sustained for
as long a period as aerobic exercise. The presence of lactic acid has an inhibitory effect on ATP generation within
the muscle; though not producing fatigue, it can inhibit or even stop performance if the intracellular concentration
becomes too high. However, long-term training causes neovascularization within the muscle, increasing the ability to
move waste products out of the muscles and maintain contraction. Once moved out of muscles with high
concentrations within the sarcomere, lactic acid can be used by other muscles or body tissues as a source of energy,
or transported to the liver where it is converted back to pyruvate. The ability of the body to export lactic acid and use
it as a source of energy depends on training level.
Humans are genetically predisposed with a larger percentage of one type of muscle group over another. An
individual born with a greater percentage of Type I muscle fibers would theoretically be more suited to endurance
events, such as triathlons, distance running, and long cycling events, whereas a human born with a greater percentage
of Type II muscle fibers would be more likely to excel at anaerobic events such as a 200 meter dash, or weightlifting.
Muscle 63

Delayed onset muscle soreness is pain or discomfort that may be felt one to three days after exercising and subsides
generally within two to three days later. Once thought to be caused by lactic acid buildup, a more recent theory is
that it is caused by tiny tears in the muscle fibers caused by eccentric contraction, or unaccustomed training levels.
Since lactic acid disperses fairly rapidly, it could not explain pain experienced days after exercise.[6]
Muscular, spinal and neural factors all affect muscle building. Sometimes a person may notice an increase in strength
in a given muscle even though only its opposite has been subject to exercise, such as when a bodybuilder finds her
left biceps stronger after completing a regimen focusing only on the right biceps. This phenomenon is called cross

Symptoms of muscle diseases may include weakness, spasticity, myoclonus and myalgia. Diagnostic procedures that
may reveal muscular disorders include testing creatine kinase levels in the blood and electromyography (measuring
electrical activity in muscles). In some cases, muscle biopsy may be done to identify a myopathy, as well as genetic
testing to identify DNA abnormalities associated with specific myopathies and dystrophies.
Neuromuscular diseases are those that affect the muscles and/or their nervous control. In general, problems with
nervous control can cause spasticity or paralysis, depending on the location and nature of the problem. A large
proportion of neurological disorders leads to problems with movement, ranging from cerebrovascular accident
(stroke) and Parkinson's disease to Creutzfeldt-Jakob disease.
A non-invasive elastography technique that measures muscle noise is undergoing experimentation to provide a way
of monitoring neuromuscular disease. The sound produced by a muscle comes from the shortening of actomyosin
filaments along the axis of the muscle. During contraction, the muscle shortens along its longitudinal axis and
expands across the transverse axis, producing vibrations at the surface.[7]

There are many diseases and conditions which cause a decrease in muscle mass, known as muscle atrophy. Examples
include cancer and AIDS, which induce a body wasting syndrome called cachexia. Other syndromes or conditions
which can induce skeletal muscle atrophy are congestive heart disease and some diseases of the liver.
During aging, there is a gradual decrease in the ability to maintain skeletal muscle function and mass, known as
sarcopenia. The exact cause of sarcopenia is unknown, but it may be due to a combination of the gradual failure in
the "satellite cells" which help to regenerate skeletal muscle fibers, and a decrease in sensitivity to or the availability
of critical secreted growth factors which are necessary to maintain muscle mass and satellite cell survival.
Sarcopenia is a normal aspect of aging, and is not actually a disease state yet can be linked to many injuries in the
elderly population as well as decreasing quality of life.[8]
Atrophy is of particular interest to the manned spaceflight community, since the weightlessness experienced in
spaceflight results is a loss of as much as 30% of mass in some muscles.[9] [10]
Muscle 64

Physical inactivity and atrophy

Inactivity and starvation in mammals lead to atrophy of skeletal muscle, accompanied by a smaller number and size
of the muscle cells as well as lower protein content.[11] In humans, prolonged periods of immobilization, as in the
cases of bed rest or astronauts flying in space, are known to result in muscle weakening and atrophy. Such
consequences are also noted in small hibernating mammals like the golden-mantled ground squirrels and brown
Bears are an exception to this rule; species in the family Ursidae are famous for their ability to survive unfavorable
environmental conditions of low temperatures and limited nutrition availability during winter by means of
hibernation. During that time, bears go through a series of physiological, morphological and behavioral changes.[13]
Their ability to maintain skeletal muscle number and size at time of disuse is of a significant importance.
During hibernation, bears spend four to seven months of inactivity and anorexia without undergoing muscle atrophy
and protein loss.[12] There are a few known factors that contribute to the sustaining of muscle tissue. During the
summer period, bears take advantage of the nutrition availability and accumulate muscle protein. The protein balance
at time of dormancy is also maintained by lower levels of protein breakdown during the winter time.[12] At times of
immobility, muscle wasting in bears is also suppressed by a proteolytic inhibitor that is released in circulation.[11]
Another factor that contributes to the sustaining of muscle strength in hibernating bears is the occurrence of periodic
voluntary contractions and involuntary contractions from shivering during torpor.[14] The three to four daily episodes
of muscle activity are responsible for the maintenance of muscle strength and responsiveness in bears during

A display of "strength" (e.g. lifting a weight) is a result of three factors that overlap: physiological strength (muscle
size, cross sectional area, available crossbridging, responses to training), neurological strength (how strong or weak
is the signal that tells the muscle to contract), and mechanical strength (muscle's force angle on the lever, moment
arm length, joint capabilities). Contrary to popular belief, the number of muscle fibres cannot be increased through
exercise; instead the muscle cells simply get bigger. Muscle fibres have a limited capacity for growth through
hypertrophy and some believe they split through hyperplasia if subject to increased demand.

The "strongest" human muscle

Since three factors affect muscular strength simultaneously and muscles never work individually, it is misleading to
compare strength in individual muscles, and state that one is the "strongest". But below are several muscles whose
strength is noteworthy for different reasons.
• In ordinary parlance, muscular "strength" usually refers to the ability to exert a force on an external object—for
example, lifting a weight. By this definition, the masseter or jaw muscle is the strongest. The 1992 Guinness Book
of Records records the achievement of a bite strength of 4337 N (975 lbf) for 2 seconds. What distinguishes the
masseter is not anything special about the muscle itself, but its advantage in working against a much shorter lever
arm than other muscles.
• If "strength" refers to the force exerted by the muscle itself, e.g., on the place where it inserts into a bone, then the
strongest muscles are those with the largest cross-sectional area. This is because the tension exerted by an
individual skeletal muscle fiber does not vary much. Each fiber can exert a force on the order of 0.3 micronewton.
By this definition, the strongest muscle of the body is usually said to be the quadriceps femoris or the gluteus
• A shorter muscle will be stronger "pound for pound" (i.e., by weight) than a longer muscle. The myometrial layer
of the uterus may be the strongest muscle by weight in the human body. At the time when an infant is delivered,
the entire human uterus weighs about 1.1 kg (40 oz). During childbirth, the uterus exerts 100 to 400 N (25 to
100 lbf) of downward force with each contraction.
Muscle 65

• The external muscles of the eye are conspicuously large and strong in relation to the small size and weight of the
eyeball. It is frequently said that they are "the strongest muscles for the job they have to do" and are sometimes
claimed to be "100 times stronger than they need to be." However, eye movements (particularly saccades used on
facial scanning and reading) do require high speed movements, and eye muscles are exercised nightly during
rapid eye movement sleep.
• The statement that "the tongue is the strongest muscle in the body" appears frequently in lists of surprising facts,
but it is difficult to find any definition of "strength" that would make this statement true. Note that the tongue
consists of sixteen muscles, not one.
• The heart has a claim to being the muscle that performs the largest quantity of physical work in the course of a
lifetime. Estimates of the power output of the human heart range from 1 to 5 watts. This is much less than the
maximum power output of other muscles; for example, the quadriceps can produce over 100 watts, but only for a
few minutes. The heart does its work continuously over an entire lifetime without pause, and thus does "outwork"
other muscles. An output of one watt continuously for eighty years yields a total work output of two and a half

The efficiency of human muscle has been measured (in the context of rowing and cycling) at 18% to 26%. The
efficiency is defined as the ratio of mechanical work output to the total metabolic cost, as can be calculated from
oxygen consumption. This low efficiency is the result of about 40% effiency of generating ATP from food energy,
losses in converting energy from ATP into mechanical work inside the muscle, and mechanical losses inside the
body. The latter two losses are dependent on the type of exercise and the type of muscle fibers being used
(fast-twitch or slow-twitch). For an overal efficiency of 20 percent, one watt of mechanical power is equivalent to
4.3 kcal per hour. For example, a manufacturer of rowing equipment shows burned calories as four times the actual
mechanical work, plus 300 kcal per hour,[15] which amounts to about 20 percent efficiency at 250 watts of
mechanical output. The mechanical energy output of a cyclic contraction can depend upon many factors, including
activation timing, muscle strain trajectory, and rates of force rise & decay. These can be synthesized experimentally
using work loop analysis.

Density of muscle tissue compared to adipose tissue

The density of mammalian skeletal muscle tissue is about 1.06 kg/liter.[16] This can be contrasted with the density of
adipose tissue (fat), which is 0.9196 kg/liter.[17] This makes muscle tissue approximately 15% denser than fat tissue.

Resting energy expenditure of muscle

At rest, skeletal muscle consumes 54.4 kJ/kg (13.0 kcal/kg) per day. This is larger than adipose tissue (fat) at
18.8 kJ/kg (4.5 kcal/kg), and bone at 9.6 kJ/kg (2.3 kcal/kg).[18]

Muscle evolution
Evolutionarily, specialized forms of skeletal and cardiac muscles predated the divergence of the vertebrate/arthropod
evolutionary line.[19] This indicates that these types of muscle developed in a common ancestor sometime before 700
million years ago (mya). Vertebrate smooth muscle was found to have evolved independently from the skeletal and
cardiac muscles.
Muscle 66

See also
• Atrophy
• Bodybuilding
• Cross education
• Electroactive polymers (materials that behave like muscles, used in robotics research)
• Fascia
• Hand strength
• In vitro muscle testing
• List of muscles of the human body
• List of weight training exercises
• Muscle atrophy
• Muscle memory
• Muscle tone (residual muscle tension)
• Musculoskeletal system
• Muscular system
• Myopathy (pathology of muscle cells)
• Myotomy
• Phonomyography
• Preflexes
• Rapid plant movement
• Rohmert's law
• Soft tissue

[1] Definition and origin of the word 'muscle' (http:/ / www. askoxford. com/ concise_oed/ muscle)
[2] Basic Concepts in Embryology: A Student's Survival Guide (Paperback) Lauren Sweeney. 1997. McGraw-Hill Professional, 1st edition
[3] Marieb, Elaine; Katja Hoehn (2007). Human Anatomy & Physiology (7th ed.). Pearson Benjamin Cummings. p. 317. ISBN 0805353879.
[4] Larsson, L; Edström, L; Lindegren, B; Gorza, L; Schiaffino, S (July 1991). "MHC composition and enzyme-histochemical and physiological
properties of a novel fast-twitch motor unit type" (http:/ / ajpcell. physiology. org/ cgi/ reprint/ 261/ 1/ C93). The American Journal of
Physiology 261 (1 pt 1): C93–101. PMID 1858863. . Retrieved 2006-06-11.
[5] Smerdu, V; Karsch-Mizrachi, I; Campione, M; Leinwand, L; Schiaffino, S (December 1994). "Type IIx myosin heavy chain transcripts are
expressed in type IIb fibers of human skeletal muscle" (http:/ / ajpcell. physiology. org/ cgi/ reprint/ 267/ 6/ C1723). The American Journal of
Physiology 267 (6 pt 1): C1723–1728. PMID 7545970. . Retrieved 2006-06-11. Note: Access to full text requires subscription; abstract freely
[6] Robergs R, Ghiasvand F, Parker D (2004). "Biochemistry of exercise-induced metabolic acidosis.". Am J Physiol Regul Integr Comp Physiol
287 (3): R502–16. doi:10.1152/ajpregu.00114.2004. PMID 15308499.
[7] 'Muscle noise' could reveal diseases' progression (http:/ / www. undeadlinks. com/ s. php?link=http:/ / www. newscientisttech. com/ article.
ns?id=dn11885& feedId=online-news_rss20& l=1) 18 May 2007, news service, Belle Dumé
[8] http:/ / www3. interscience. wiley. com/ journal/ 110504447/ abstract?CRETRY=1& SRETRY=0
[9] Roy, R. R., Baldwin, K. M., and Edgerton, V. R. (1996) Response of the neuromuscular unit to spaceflight: What has been learned from the
rat model. Exerc. Sport Sci. Rev. 24, 399–425
[10] "NASA Muscle Atrophy Research (MARES) Website" (http:/ / www. nasa. gov/ mission_pages/ station/ science/ experiments/ MARES.
[11] Fuster G, Busquets S, Almendro V, López-Soriano FJ, Argilés JM (2007). "Antiproteolytic effects of plasma from hibernating bears: a new
approach for muscle wasting therapy?" (http:/ / linkinghub. elsevier. com/ retrieve/ pii/ S0261-5614(07)00124-0). Clin Nutr 26 (5): 658–61.
doi:10.1016/j.clnu.2007.07.003. PMID 17904252. .
[12] Lohuis TD, Harlow HJ, Beck TD (2007). "Hibernating black bears (Ursus americanus) experience skeletal muscle protein balance during
winter anorexia" (http:/ / linkinghub. elsevier. com/ retrieve/ pii/ S1096-4959(07)00053-X). Comp. Biochem. Physiol. B, Biochem. Mol. Biol.
147 (1): 20–8. doi:10.1016/j.cbpb.2006.12.020. PMID 17307375. .
[13] Carey HV, Andrews MT, Martin SL (2003). "Mammalian hibernation: cellular and molecular responses to depressed metabolism and low
temperature" (http:/ / physrev. physiology. org/ cgi/ pmidlookup?view=long& pmid=14506303). Physiol. Rev. 83 (4): 1153–81.
doi:10.1152/physrev.00008.2003 (inactive 2008-06-24). PMID 14506303. .
Muscle 67

[14] Harlow, H.J. (2004). "Body Surface Temperature Of Hibernating Black Bears May Be Related To Periodic Muscle Activity". Journal of
Mammalogy 85 (3): 414–419. doi:10.1644/1545-1542(2004)085< (inactive 2008-06-24).
[15] Concept II Rowing Ergometer, user manual (http:/ / www. concept2. com/ us/ support/ manuals/ pdf/ B_UsersManual. pdf). (1993)
[16] Urbancheka M, Pickenb E, Kaliainenc L, Kuzon W (2001). "Specific Force Deficit in Skeletal Muscles of Old Rats Is Partially Explained by
the Existence of Denervated Muscle Fibers. (http:/ / answers. google. com/ answers/ threadview?id=576481)". The Journals of Gerontology
Series A: Biological Sciences and Medical Sciences 56:B191-B197.
[17] Farvid, MS; Ng, TW; Chan, DC; Barrett, PH; Watts, GF (2005). "Association of adiponectin and resistin with adipose tissue compartments,
insulin resistance and dyslipidaemia". Diabetes, obesity & metabolism 7 (4): 406–13. doi:10.1111/j.1463-1326.2004.00410.x.
PMID 15955127.
[18] http:/ / ajpendo. physiology. org/ cgi/ reprint/ 282/ 1/ E132
[19] Evolution of muscle fibers (http:/ / www. umbi. umd. edu/ ~collins/ myoinformatics/ muscle-evolution. pdf)

External links
• Muslumova, Irada (2003). "Power of a Human Heart" (
shtml). The Physics Factbook. (Heart output 1.3 to 5 watts, lifetime output 2 to 3 ×109 joules)
• University of Dundee ( article on
performing neurological examinations (Quadriceps "strongest")
• Muscle efficiency in rowing (
• Human Muscle Tutorial ( (clear pictures of
main human muscles and their Latin names, good for orientation)
• Microscopic stains of skeletal (
skm02.32706.jpg) and cardiac (
mycard-positive.jpg) muscular fibers to show striations. Note the differences in myofibrilar arrangements.

Muscle tone
In physiology, medicine, and anatomy, muscle tone (residual muscle tension or tonus) is the continuous and
passive partial contraction of the muscles. It helps maintain posture, and it declines during REM sleep.

Unconscious nerve impulses maintain the muscles in a partially contracted state. If a sudden pull or stretch occurs,
the body responds by automatically increasing the muscle's tension, a reflex which helps guard against danger as
well as helping to maintain balance.
The presence of near-continuous innervation makes it clear that tonus describes a "default" or "steady state"
condition. There is, for the most part, no actual "rest state" insofar as activation is concerned.
In terms of skeletal muscle, both the extensor and flexor muscles, under normal innervation, maintain a constant tone
while "at rest" that maintains a normal posture.
Cardiac muscle and smooth muscle, although not directly connected to the skeleton, also have tonus in the sense that
although their contractions are not matched with those of antagonist muscles, the non-contractive state is
characterized by (sometimes random) enervation.
Muscle tone 68

Pathological tonus
Physical disorders can result in abnormally low (hypotonia) or high (hypertonia) muscle tone. Another form of
hypertonia is paratonia, which is associated with dementia. Hypotonia is caused by lower motor neuron disease like
poliomyelitis. Hypertonia is caused in upper motor neuron disease like lesion in pyramidal tract and extrapyramidal
tract. Hypertonia can be of clasp knife variety, in which there is increased resistance only at the beginning or at the
end of the movement, or lead pipe variety, in which there is resistance throughout to passive movement, or it may be
of cog wheel type, in which the resistance to passive movement is in jerky manner.

Tonus in surgery
In ophthalmology, tonus may be a central consideration in eye surgery, as in the manipulation of extraocular muscles
to repair strabismus. Tonicity aberrations are associated with many diseases of the eye (e.g. Adie syndrome).

External links
• BBC series on muscles [1]

[1] http:/ / www. bbc. co. uk/ science/ humanbody/ body/ factfiles/ tone/ adductor_animation. shtml

Muscle memory
Muscle memory, also known as motor learning, is a form of procedural memory that involves consolidating a
specific motor task into memory through repetition. When a movement is repeated over time, a long-term muscle
memory is created for that task, eventually allowing it to be performed without conscious effort. This process
decreases the need for attention and creates maximum efficiency within the motor and memory systems. Examples
of muscle memory are found in many everyday activities that become automatic and improve with practice, such as
riding a bicycle, typing on a keyboard, playing a melody or phrase on a musical instrument, playing video games[1] ,
or even solving a puzzle cube.


Movement and motor learning

Movement is a critical part of our life, and it is a major component of our evolutionary development; without it, we
could not survive.[2] It has been suggested that our developed cognitive capacities evolved so we could make
movements essential to our survival. For example, cognitive abilities evolved so we could use tools, build shelter,
and hunt for animals.
The origins of research for the acquisition of motor skills stem from philosophers such as Plato, Aristotle and Galen.
Friedrich Bessel is a philosopher who is especially noteworthy, as he was among the first to empirically observe
motor learning. Bessel tried to observe the difference in his colleagues with the method in which they recorded the
transit time of stars.[2] After the break from tradition of the pre-1900s view of introspection, psychologists
emphasized research and more scientific methods in observing behaviours.[3] Thereafter, numerous studies exploring
the role of motor learning were conducted. Such studies included the research of handwriting, and various practice
methods to maximize motor learning.[2]
Muscle memory 69

The retention of motor skills, now referred to as muscle memory, also began to be of great interest in the early
1900s. Most motor skills are thought to be acquired through practice; however, mere observation of the skill has led
to learning as well.[4] Research suggests we do not start off with a blank slate in regards to motor memory although
we do learn most of our motor memory repertoire during our lifetime.[5] Movements such as facial expressions,
which are thought to be learned, can actually be observed in children who are blind; thus there is some evidence for
motor memory being genetically pre-wired.[5]
In the early stages of empirical research of motor memory Edward Thorndike, a leading pioneer in the study of
motor memory, was among the first to acknowledge learning can occur without conscious awareness.[6] One of the
earliest and most notable studies regarding the retention of motor skills was by Hill, Rejall, and Thorndike, who
showed savings in relearning typing skills after a 25 year period with no practice.[2] Findings related to the retention
of learned motor skills have been continuously replicated in studies, suggesting that through subsequent practice,
motor learning is stored in the brain as memory. This is why performing skills such as riding a bike or driving a car
are effortlessly and ‘unconsciously’ executed, even if someone had not performed these skills in a long period of


Motor behavior
When first learning a motor task, movement is often slow, stiff and easily disrupted without attention. With practice,
execution of motor task becomes smoother, there is a decrease in limb stiffness, and muscle activity necessary to the
task is performed without conscious effort.[7]

Muscle memory encoding

The neuroanatomy of memory is widespread throughout the brain; however, the pathways important to motor
memory are separate from the medial temporal lobe pathways associated with declarative memory.[8] As with
declarative memory, motor memory is theorized to have two stages; a short term memory encoding stage that is
fragile and susceptible to damage, as well as a long term memory consolidation stage which is more stable.[9]
The memory encoding stage is often referred to as motor learning, and requires an increase in brain activity in motor
areas as well as an increase in attention. Brain areas active during motor learning include the motor and
somatosensory cortices; however these areas of activation decrease once the motor skill is learned. The prefrontal
and frontal cortices are also active during this stage due to the need for increased attention on the task being
The main area involved in motor learning is the cerebellum. Some models of cerebellar-dependent motor learning,
particularly the Marr-Albus model, propose a single plasticity mechanism involving the cerebellar long term
depression(LTD) of the parallel fiber synapses onto Purkinje cells. These modification in synapse activity would
mediate motor input with motor outputs critical to inducing motor learning.[10] However, conflicting evidence
suggests that a single plasticity mechanism is not sufficient and a multiple plasticity mechanism is needed to account
for the storage of motor memories over time. Regardless of the mechanism, studies of cerebellar-dependent motor
tasks show that cerebral cortical plasticity is crucial for motor learning, even if not necessarily for storage.[11]
The basal ganglia also play an important role in memory and learning; particularly in reference to stimulus-response
associations and the formation of habits. The basal ganglia-cerebellar connections are thought to increase with time
when learning a motor task.[12]
Muscle memory 70

Muscle memory consolidation

Muscle memory consolidation involves the continuous evolution of neural processes after practicing a task has
stopped. The exact mechanism of motor memory consolidation within the brain is controversial; however most
theories assume that there is a general redistribution of information across the brain from encoding to consolidation.
Hebb's rule states that “synaptic connectivity changes as a function of repetitive firing.” In this case, that would mean
that the high amount of stimulation coming from practicing a movement would cause the repetition of firing in
certain motor networks; presumably leading to an increase in the efficiency of exciting these motor networks over
Though the exact location of muscle memory storage isn’t known, studies have suggested that it is the inter-regional
connections that play the most important role in advancing motor memory encoding to consolidation, rather than
decreases in overall regional activity. These studies have shown a weakened connection from the cerebellum to the
primary motor area with practice; possibly because of a decreased need for error correction from the cerebellum.
Conversely, the connection between the basal ganglia and the primary motor area is strengthened, suggesting the
basal ganglia plays an important role in the motor memory consolidation process.[11]

Athletic training
When participating in any sport, new motor skills and movement combinations are frequently being used and
repeated. All sports require some degree of strength, endurance training and skilled reaching, in order to be
successful in the required tasks.
Evidence has shown that increases in strength occur well before muscle hypertrophy, and decreases in strength due
to detraining or ceasing to repeat the exercise over an extended period of time precede muscle atrophy.[13]
Specifically, strength training enhances motor neuron excitability and induces synaptogenesis, both of which would
help in enhancing communication between the nervous system and the muscles themselves.[13] .
However, neuromuscular efficacy is not altered within a 2 week time
period following cessation of the muscle usage; instead it is merely the
neuron`s ability to excite the muscle that declines in correlation with
the muscle's decrease in strength.[14] This confirms that muscle
strength is first influenced by the inner neural circuitry, rather than by
external physiological changes in the muscle size.

Reorganization of motor maps within the cortex are not altered in

either strength or endurance training. However, within the motor
cortex, endurance induces angiogenesis within as little as 3 weeks to
increase blood flow to the involved regions.[13] In addition, neurotropic factors within the motor cortex are
upregulated in response to endurance training to promote neural survival.[13]
Skilled motor tasks have been divided into two distinct phases; a fast learning phase, in which an optimal plan for
performance is established, and a slow learning phase, in which longer term structural modifications are made on
specific motor modules.[15] Even a small amount of training may be enough to induce neural processes that continue
to evolve even after the training has stopped, which provides a potential basis for consolidation of the task.
Additionally, studying mice while they are learning a new complex reaching task, has found that “motor learning
leads to rapid formation of dendritic spines (spinogenesis) in the motor cortex contralateral to the reaching
forelimb”.[16] However, motor cortex reorganization itself does not occur at a uniform rate across training periods. It
has been suggested that the synaptogenesis and motor map reorganization merely represent the consolidation, and
not the acquisition itself, of a specific motor task.[17] Furthermore, the degree of plasticity in various locations
(namely motor cortex versus spinal cord) is dependent on the behavioural demands and nature of the task (i.e. skilled
reaching versus strength training).[13]
Muscle memory 71

Whether strength or endurance related, it is plausible that the majority of motor movements would require a skilled
moving task of some form, whether it be maintaining proper form when paddling a canoe, or bench pressing a
heavier weight. Endurance training assists the formation of these new neural representations within the motor cortex
by up regulating neurotropic factors that could enhance the survival of the newer neural maps formed due to the
skilled movement training.[13] Strength training results are seen in the spinal cord well before any physiological
muscular adaptation is established through muscle hypertrophy or atrophy.[13] The results of endurance and strength
training, and skilled reaching, therefore combine to help each other maximize performance output.

Fine motor memory

Fine motor skills are often discussed in terms of transitive movements, which are those done when using tools
(which could be as simple as a tooth brush or pencil).[18] Transitive movements have representations that become
programmed to the premotor cortex, creating motor programs which result in the activation of the motor cortex and
therefore the motor movements.[18] In a study testing the motor memory of patterned finger movements (a fine motor
skill) it was found that retention of certain skills are susceptible to disruption if another task interferes with one’s
motor memory.[1] However, such susceptibility can be can be reduced with time. For example, if a finger pattern is
learned, then another is learned six hours later the original pattern will still be remembered, while learning such
patterns back to back may cause forgetting of the initial one.[1] Furthermore, the heavy use of computers by recent
generations has both positive and negative effects. It was found that one of the main positive effects is that it
enhances fine motor skills of children.[19] Repetitive behaviours, such as typing on a computer from a young age, can
enhance such abilities. Therefore, by beginning computer use at an early age muscle memory may be activated

Music memory
Fine motor skills are very important in playing musical instruments. It
was found that muscle memory is relied on when playing the clarinet,
specifically to help create special effects through certain tongue
movements when blowing air into the instrument.[20] Memorizing is
done by muscles as a note is seen and recalled, its auditory pair is
learned and is matched by fingers movements (a fine motor skill).[21]
When reproducing a motor action, you must have previous experience
with it to memorize set actions. If there is no previous experience there
will be no mental image of the motion, and therefore no actual

Certain human behaviours, especially actions like the fingering in

musical performances, are very complex and require many
interconnected neural networks where information can be transmitted
across multiple brain regions.[22] It has been found that there are often
functional differences in the brains of professional musicians, when
compared to other individuals. This is thought to reflect the musician’s
innate ability which may be fostered by an early exposure to musical Playing the piano requires complex actions
training.[22] An example of this is bimanual synchronized finger
movements which play an essential role in piano playing. It is suggested that bimanual coordination can only come
from years of bimanual training, where such actions become adaptations of the motor areas.[23] When comparing
professional musicians to a control group in complex bimanual movements, professionals are found to use an
Muscle memory 72

extensive motor network much less than those non-professionals.[23] This is because professionals rely on a motor
system that has increased efficiency, and therefore those who are less trained have a network which is more strongly
activated.[23] It is implied that the untrained pianists have to invest more neuronal activity to have the same level of
performance that is achieved by professionals.[23] This, yet again, is said to be a consequence of many years of motor
training and experience which helps form a fine motor memory skill of musical performance.
It is often reported that when a pianist hears a well-trained piece of music it can involuntarily trigger synonymous
fingering.[22] This implies there is a coupling between the perception of music and the motor activity of those
musically trained individuals.[22] Therefore, one’s muscle memory in the context of music can easily be triggered
when one hears certain familiar pieces. Overall, long-term musical fine motor training allows for complex actions to
be performed at a lower level of movement control, monitoring, selection, attention, and timing.[23] This leaves room
for musicians to focus attention synchronously elsewhere, such as on the artistic aspect of the performance, without
having to consciously control one’s fine motor actions.[23]

Gross motor memory

Gross motor skills are concerned with the movement of large muscles, or major body movements, such as those
involved in walking or kicking, and are associated with normal development.[24] The extent to which one exhibits
gross motor skills depends largely on their muscle tone and the strength.[24] In a study looking at people with Down
Syndrome it was found that the pre-existing deficits, with regards to verbal-motor performance, has an impact on
limiting the individuals transfer of gross motor skills following visual and verbal instruction to verbal instruction
only.[25] The fact that the individuals could still exhibit two of the three original motor skills may have been a result
of positive transfer in which previous exposure allows the individual to remember the motion, under the visual and
verbal trial, and then later perform it under the verbal trial.[25]

Learning in childhood
The way in which a child learns a gross motor skill can impact how long it takes to consolidate it and be able to
reproduce the movement. In a study with preschoolers, looking at the role of self-instruction on acquiring complex
gross motor chains using ballet positions, it was found that the motor skills were better learned and remembered,
with the self-instruction procedure, over the no self-instruction procedure.[26] This suggests that the use of
self-instruction will increase the speed with which a preschooler will learn and remember a gross motor skill. It was
also found that once the preschoolers learned and mastered the motor chain movements, they ceased the use of
self-instruction. This suggests that the memory for the movements became strong enough that there was no longer a
need for self-instruction and the movements could be reproduced without it.[26]

Impact of Alzheimer's disease

It has been suggested that consistent practice of a gross motor skill can help a patient with Alzheimer’s Disease learn
and remember that skill. It was thought that the damage to the hippocampus may result in the need for a specific type
of learning requirement.[27] A study was created to test this assumption in which the patients were trained to throw a
bean bag at a target.[27] It was found that the Alzheimer’s patients performed better on the task when learning
occurred under constant training as opposed to variable. Also, it was found that gross motor memory in Alzheimer’s
patients was the same as healthy adults when learning occurs under constant practice.[27] This suggests that damage
to the hippocampal system does not impair an Alzheimer’s patient from retaining new gross motor skills; implying
that motor memory for gross motor skills is stored elsewhere in the brain.
Muscle memory 73

It is difficult to display cases of “pure” motor memory impairment because the memory system is so widespread
throughout the brain that damage isn’t often isolated to one specific type of memory. Similarly, diseases commonly
associated with motor deficits, such as Huntington's and Parkinson's disease, have a wide variety of symptoms and
associated brain damage that make it impossible to pinpoint whether or not motor memory is in fact impaired. Case
studies have provided some examples of how motor memory has been implemented in patients with brain damage.

Consolidation deficit
A recent issue in motor memory is whether or not it consolidates in a manner similar to declarative memory; a
process that involves an initial fragile learning period that eventually becomes stable and less susceptible to damage
over time.[1] An example of stable motor memory consolidation in a patient with brain damage is the case of Clive
Wearing. Clive has severe antrograde and retrograde amnesia due to damage in his temporal lobes, frontal lobes and
his hippocampi; which prevents him from storing any new memories and making him aware of only the present
moment. However, Clive still retains access to his procedural memories, specifically the motor memories involved in
playing the piano. This could be because motor memory is demonstrated through savings over several trials of
learning, whereas declarative memory is demonstrated through recall of a single item[1] . This suggests that lesions in
certain brain areas normally associated with declarative memory would not affect motor memory for a well learned

Dysgraphia for the alphabet

Case study: 54 year old woman with known history of epilepsy This patient was diagnosed with a pure form of
dysgraphia of letters, meaning she had no other speech or reading impairments.[28] Her impairment was specific to
letters in the alphabet. She was able to copy letters from the alphabet, but she was not able to write these letters.[28]
She had previously been rated average on the Wechsler Adult Intelligence Scale's vocabulary subtest for writing
ability comparative to her age before her diagnosis.[28] Her writing impairment consisted of difficulty remembering
motor movements associated with the letters she was supposed to write.[28] She was able to copy the letters, and also
form images that were similar to the letters.[28] This suggests that dysgraphia for letters is a deficit related to motor
memory.[28] Somehow there is a specific portion of the brain related to writing letters, which is dissociated from
copying and drawing letter-like items.

See also
• Procedural memory
• Motor learning
• Motor coordination
• Muscle
• Memory consolidation
Muscle memory 74

[1] Krakauer, J.W., & Shadmehr, R. (2006). Consolidation of motor memory. Trends in Neurosciences, 29: 58-64.
[2] Lee, D.T., & Schmidt, A.R. (2005). Motor Control and Learning: A Behavioural Emphasis. (4th ed). Windsor, ON: Human Kinetics
[3] Adams, A.J. (1987). Historical Review and Appraisal of Research on the Learning, Retention, and Transfer of Human Motor Skills.
Psychological Bulletins, 101(1), 41-74
[4] Celnik, P., Classen, J., Cohen, G.L., Duque, J., Mazzocchio, R., Sawaki, L., Stephan, K., & Ungerleider, L. (2005). Formation of a Motor
Memory by Action Observation. The Journal of Neuroscience, 25(41), 9339-9346
[5] Flanagan, R.J., Ghahramani, Z., & Wolpert, M.D. (2001). Perspectives and Problems in Motor Learning. Trends in Cognitive Sciences, 5(11),
[6] Shanks, D.R. & St. John, M.F. (1994). Characteristics of Dissociable Human Learning Systems. Behavioural and Brain Sciences, 17(3),
[7] Shadmehr R, Holcomb HH. (1997). Neural correlates of motor memory consolidation. Science 227: 821-25.
[8] Brashers-Krug, T, Shadmehr, R. and Bizzi, E. (1996). Consolidation in human motor memory. Nature. 382.
[9] Atwell P.,Cooke S.,Yeo C. (2002). Cerebellar function in consolidation of motor memory. Neuron. 34:1011-1020.
[10] Boyden, E., Katoh, A., Raymond, J.(2004). Cerebellum-dependent learning: the role of multiple plasticity mechanisms. Annu. Rev.
[11] Ma, L., et al,. (2010). Changes in regional activity are accompanied with changes in inter-regional connectivity during 4 weeks motor
learning. Brain Res. doi: 10.1016/j.brainres.2009.12.073
[12] Packard, M., Knowlton, B. 2002. Learning and memory functions of the basal ganglia. Annu. Rev. Neuroscience. 25: 563-93.
[13] Adkins, DeAnna L., Boychuck, Jeffery. 2006. Motor training induces experience specific patterns of plasticity across motor cortex and
spinal cord. Journal of Applied Physiology. 101: 1776-1782.
[14] Deschenes Michael R., Giles Jennifer A. 2002. Neural factors account for strength decrements observed after short-term muscle unloading.
The American Journal of Physiology - Regulatory, Integrative and Comparative Physiology. 282: R578-R583.
[15] Karni, Avi, Meyer, Gundela. 1998. The acquisition of skilled motor performance: Fast and slow experience-driven changes in primary motor
cortex. Proceedings of the National Academy of Sciences. 861-868.
[16] Xu, Tonghui, Perlik, Andrew J. 2009. Rapid formation and selective stabilization of synapses for enduring motor memories. Nature. 915-20.
[17] Kleim Jerrery L., Hogg Theresa M. 2004. Cortical Synaptogenesis and Motor Map Reorganization Occur during Late, But not Early, Phase
of Motor Skill Learning. The Journal of Neuroscience. 24:629-633.
[18] Dowell, L. R., Mahone, E. M., & Mostofsky, S. H. (2009). Associations of postural knowledge and basic motor skill with dyspraxia in
autism: Implication for abnormalities in distributed connectivity and motor learning. Neuropsychology, 23, 563-570.
[19] Straker, L., Pollock, C., & Maslen, B. (2009). Principles for the wise use of computers by children. Ergonomics [Ergonomics], 52,
[20] Fritz, C., & Wolfe, J. (2005). How do clarinet players adjust the resonances of their vocal tracts for different playing effects? Journal of the
Acoustical Society of America,118, 3306-3315.
[21] Smith, T. L. (1896). On muscular memory. American Journal of Psychology, 7, 453-490.
[22] Kim, D., Shin, M., Lee, K., Chu, K., Woo, S., Kim, Y., Song, E., Lee, Jun., Park, S., & Roh, J. (2004). Musical Training-Induced Functional
Reorganization of the Adult Brain: Functional Magnetic Resonance Imaging and Transcranial Magnetic Stimulation Study on Amateur String
Players. Human Brain Mapping, 23, 188-199.
[23] Haslinger, B., Erhard, P., Altenmüller, E., Hennenlotter, A., Schwaiger, M., von Einsiedel, H. G., Rummeny, E., Conrad, B., &
Ceballos-Baumann, A. O. (2004). Reduced Recruitment of Motor Association Areas During Bimanual Coordination in Concert
PianistsHuman Brain Mapping, 22, 206-215.
[24] “Gross motor Skills – What are Gross Motor Skills (http:/ / learningdisabilities. about. com/ od/ gi/ p/ grossmotorskill. htm)”
[25] Meegan, S., Maraj, B. K.V., Weeks, D., Chua, R. (2006). Gross Motor Skill Acquisition in Adolescents With Down Syndrom.
DownSyndrome and Practice 9(3), 75-80
[26] Vintere, P., Hemmes, N. S., Brown, B. L., Poulson, C. L. (2004) Gross-Motor Skill Acquisition by Preschool Dance Stoudents Under
Self-Instruction Procedures. Journal of Applied Behaviour Analysis 37(3), 305-322
[27] Dick, M. B., Shankle, R. W., Beth, R. E., Dick-Muehlke, C., Cotman, C. W., Kean, M. L. (1996). Acquisition and long-term retention of a
gross motor skill in Alzheimer's disease patients under constant and varied practice conditions. The Journals of Gerontology: Series B
Psychological sciences and social sciences, 51B(2), 103-111
[28] Kapur, N., & Lawton, N.F. (1983). Dysgraphia for Letters: a Form of Motor Memory Deficit? Journal of Neurological Psychiatry, 46(6),
Muscle atrophy 75

Muscle atrophy
Muscle atrophy
Classification and external resources

ICD-10 [1]
M 62.5

ICD-9 [2]

DiseasesDB [3]

MedlinePlus [4]

MeSH [5]

Muscle atrophy is defined as a decrease in the mass of the muscle; it can be a partial or complete wasting away of
muscle. When a muscle atrophies, this leads to muscle weakness, since the ability to exert force is related to mass.
Muscle atrophy results from a co-morbidity of several common diseases, including cancer, AIDS, congestive heart
failure, COPD (chronic obstructive pulmonary disease), renal failure, and severe burns; patients who have
"cachexia" in these disease settings have a poor prognosis. Moreover, starvation eventually leads to muscle atrophy.

Clinical settings of atrophy

There are many diseases and conditions which cause a decrease in muscle mass, known as atrophy, including:
Dejerine Sottas syndrome (HSMN Type III), inactivity, as seen when a cast is put on a limb, or upon extended
bedrest (which can occur during a prolonged illness); cachexia - which is a "body-wasting" syndrome that is a
co-morbidity of cancer and Congestive Heart Failure; Chronic Obstructive Pulmonary Disease; burns, liver failure,
etc. Other syndromes or conditions which can induce skeletal muscle atrophy are liver disease, and starvation.

Quality of life
Muscular atrophy decreases quality of life as the sufferer becomes unable to perform certain tasks or worsen the risks
of accidents while performing those (like walking). Muscular atrophy increases the risks of falling in conditions such
as IBM (inclusion body myositis). Muscular atrophy affects a major number of elderly.

Other muscles diseases, distinct from atrophy

During aging, there is a gradual decrease in the ability to maintain skeletal muscle function and mass. This condition
is called "sarcopenia". The exact cause of sarcopenia is unknown, but it may be due to a combination of the gradual
failure in the "satellite cells" which help to regenerate skeletal muscle fibers, and a decrease in sensitivity to or the
availability of critical secreted growth factors which are necessary to maintain muscle mass and satellite cell
In addition to the simple loss of muscle mass (atrophy), or the age-related decrease in muscle function (sarcopenia),
there are other diseases which may be caused by structural defects in the muscle (muscular dystrophy), or by
inflammatory reactions in the body directed against muscle (the myopathies).
Muscle atrophy 76

Muscle atrophy occurs by a change in the normal balance between protein synthesis and protein degradation. During
atrophy, there is a down-regulation of protein synthesis pathways, and an activation of protein breakdown
pathways[6] . The particular protein degradation pathway which seems to be responsible for much of the muscle loss
seen in a muscle undergoing atrophy is the ATP-dependent ubiquitin/proteasome pathway. In this system, particular
proteins are targeted for destruction by the ligation of at least four copies of a small peptide called ubiquitin onto a
substrate protein. When a substrate is thus "poly-ubiquitinated", it is targeted for destruction by the proteasome.
Particular enzymes in the ubiquitin/proteasome pathway allow ubiquitination to be directed to some proteins but not
others - specificity is gained by coupling targeted proteins to an "E3 ubiquitin ligase". Each E3 ubiquitin ligase binds
to a particular set of substrates, causing their ubiquitination.

Potential treatment
Muscle atrophy can be opposed by the signaling pathways which induce muscle hypertrophy, or an increase in
muscle size. Therefore one way in which exercise induces an increase in muscle mass is to downregulate the
pathways which have the opposite effect.
One important rehabilitation tool for muscle atrophy includes the use of functional electrical stimulation to stimulate
the muscles. This has seen a large amount of success in the rehabilitation of paraplegic patients. [7]
Since the absence of muscle-building amino acids can contribute to muscle wasting (that which is torn down must be
rebuilt with like material), amino acid therapy may be helpful for regenerating damaged or atrophied muscle tissue.
The branched-chain amino acids or BCAAs (leucine, isoleucine, and valine) are critical to this process, in addition to
lysine and other amino acids.

A CT scan can distinguish muscle tissue from other tissues and thereby estimate the amount of muscle tissue in the
Fast loss of muscle tissue (relative to normal turnover), can be approximated by the amount of urea in the urine. The
equivalent nitrogen content (in gram) of urea (in mmol) can be estimated by the conversion factor 0.028 g/mmol.[8]
Furthermore, 1 gram of nitrogen is roughly equivalent to 6 gram of protein, and 1 gram of protein is roughly
equivalent to 4 gram of muscle tissue. Subsequently, in situations such as muscle wasting, 1 mmol of excessive urea
in the urine (as measured by urine volume in litres multiplied by urea concentration in mmol/l) roughly corresponds
to a muscle loss of 0,67 gram.

[1] http:/ / apps. who. int/ classifications/ apps/ icd/ icd10online/ ?gm60. htm+ m625
[2] http:/ / www. icd9data. com/ getICD9Code. ashx?icd9=728. 2
[3] http:/ / www. diseasesdatabase. com/ ddb29472. htm
[4] http:/ / www. nlm. nih. gov/ medlineplus/ ency/ article/ 003188. htm
[5] http:/ / www. nlm. nih. gov/ cgi/ mesh/ 2010/ MB_cgi?field=uid& term=D009133
[6] Sandri M. 2008. Signaling in Muscle Atrophy and Hypertrophy. Physiology 23: 160-170. (http:/ / physiologyonline. physiology. org/ cgi/
content/ short/ 23/ 3/ 160)
[7] D.Zhang et al., Functional Electrical Stimulation in Rehabilitation Engineering: A survey, Nenyang technological University, Singapore
[8] Section 1.9.2 (page 76) in: Jacki Bishop; Thomas, Briony (2007). Manual of Dietetic Practice. Wiley-Blackwell. ISBN 1-4051-3525-5.
Muscle atrophy 77

External links
• MeSH Muscular+atrophy (

Hand strength
Hand strength measurements are of interest to study
pathology of the hand that involves loss of muscle
strength. Examples of these pathologies are carpal
tunnel syndrome, nerve injury, tendon injuries of the
hand, and neuromuscular disorders. Hand strength
testing is frequently used for clinical decision-making
and outcome evaluation in evidence based medicine. It
is used to diagnose diseases, to evaluate and compare
treatments, to document progression of muscle
strength, and to provide feedback during the
rehabilitation process. In addition, strength testing is
often used in areas such as sports medicine and
ergonomics. In general, hand strength measurements
can be divided into manual muscle testing and
Two Tai Chi practitioners participate in Pushing hands, an exercise
dynamometry. particularly involving the use of hand strength and flexibility

Manual muscle strength testing of

the hand muscles
In clinical practice, hand muscles are most often evaluated using manual muscle strength testing using the Medical
Research Council (MRC) Scale.[1] In this scale, muscle strength is graded on a scale from 0 to 5. For evaluating the
strength of the intrinsic hand muscles, a small modification to the standard MRC grading has been made so that
grade 3 indicates ‘full active range of motion’ as compared to ‘movement against gravity’[2] :

Modified Medical Research Council Scale for measuring hand muscles

Grade 5: full active range of motion & Normal muscle resistance
Grade 4: full active range of motion & Reduced muscle resistance
Grade 3: full active range of motion & No muscle resistance
Grade 2: Reduced active range of motion & No muscle resistance
Grade 1: No active range of motion & Palpable muscle contraction only
Grade 0: No active range of motion & No palpable muscle contraction
Manual muscle testing, however, has a number of limitations. One limitation is that the MRC scale is an ordinal
scale with disproportional distances between grades. Another limitation of the MRC scale is that the scoring depends
on the judgment of the examiner. Finally, with the 6-point ordinal MRC scale, it is difficult to identify relatively
small but clinically relevant changes in muscle strength.
Hand strength 78

Grip and pinch dynamometry

To create more quantitative assessments of hand muscle strength, dynamometers have been developed. These
dynamometer measurements are more sensitive to change compared to manual muscle testing and render outcome on
a continuous scale. In clinical evaluation and research studies on patients with hand problems, muscle strength
measurements are usually based on grip strength and pinch strength dynamometry. The most commonly used grip
and pinch dynamometers are the Jamar dynamometers and similar devices by other manufacturers. In several
patients groups, these measurements have a good reliability and validity. In addition, grip- and pinch strength are
functionally relevant to measure the combined action of a large number of intrinsic and extrinsic hand muscles as
well as the combined action of a number of different joints. By comparing outcome with normative data,[3] the
amount of muscle strength loss can be determined.

Dynamometry of the intrinsic hand muscles

For more specific dynamometry of the intrinsic muscles, intrinsic hand dynamometers have been developed. The
advantage of these dynamometers is that they to do not measure a large number of muscles in combined action, but
can measure single actions such as thumb opposition of index finger abduction.[4] One such dynamometer is the
Rotterdam Intrinsic Hand Myometer (RIHM).[5] Reliability and validity of this dynamometer is comparable to grip-
and pinch dynamometers.[6] [7]

[1] Neurological examination
[2] Brandsma JW, Schreuders TA (2001). "Sensible manual muscle strength testing to evaluate and monitor strength of the intrinsic muscles of
the hand: a commentary". J Hand Ther 14 (4): 273–8. PMID 11762727.
[3] Mathiowetz V, Kashman N, Volland G, Weber K, Dowe M, Rogers S (Feb 1985). "Grip and pinch strength: normative data for adults". Arch
Phys Med Rehabil 66 (2): 69–74. PMID 3970660.
[4] Geere J, Chester R, Kale S, Jerosch-Herold C (2007). "Power grip, pinch grip, manual muscle testing or thenar atrophy - which should be
assessed as a motor outcome after carpal tunnel decompression? A systematic review" (http:/ / www. pubmedcentral. nih. gov/ articlerender.
fcgi?tool=pmcentrez& artid=2213649). BMC Musculoskelet Disord 8: 114. doi:10.1186/1471-2474-8-114. PMID 18028538. PMC 2213649.
[5] RIHM (http:/ / www. handweb. eu/ RIHM/ )
[6] Selles RW, van Ginneken BT, Schreuders TA, Janssen WG, Stam HJ (Dec 2006). "Dynamometry of intrinsic hand muscles in patients with
Charcot-Marie-Tooth disease". Neurology 67 (11): 2022–7. doi:10.1212/01.wnl.0000247272.96136.16. PMID 17159111.
[7] Schreuders TA, Selles RW, Roebroeck ME, Stam HJ (2006). "Strength measurements of the intrinsic hand muscles: a review of the
development and evaluation of the Rotterdam intrinsic hand myometer". J Hand Ther 19 (4): 393–401; quiz 402.
doi:10.1197/j.jht.2006.07.024. PMID 17056399.
Atrophy 79

Classification and external resources

Mice with spinal muscular atrophy

MeSH [1]

Atrophy is the partial or complete wasting away of a part of the body. Causes of atrophy include mutations (which
can destroy the gene to build up the organ), poor nourishment, poor circulation, loss of hormonal support, loss of
nerve supply to the target organ, disuse or lack of exercise or disease intrinsic to the tissue itself. Hormonal and
nerve inputs that maintain an organ or body part are referred to as trophic [noun] in medical practice. Trophic
describes the trophic condition of tissue. A diminished muscular trophic is designated as atrophy.
Atrophy is the general physiological process of reabsorption and breakdown of tissues, involving apoptosis on a
cellular level. When it occurs as a result of disease or loss of trophic support due to other disease, it is termed
pathological atrophy, although it can be a part of normal body development and homeostasis as well.

Atrophy examples

Normal development
Examples of atrophy as part of normal development include shrinking and involution of the thymus in early
childhood, and the tonsils in adolescence.

Muscle atrophies
Disuse atrophy of muscles (muscle atrophy) and bones, with loss of mass and strength, can occur after prolonged
immobility, such as extended bedrest, or having a body part in a cast (living in darkness for the eye, bedridden for
the legs etc.). This type of atrophy can usually be reversed with exercise unless severe. Astronauts in microgravity
must exercise regularly to minimize atrophy of their limb muscles.
There are many diseases and conditions which cause atrophy of muscle mass. For example diseases such as cancer
and AIDS induce a body wasting syndrome called "cachexia", which is notable for the severe muscle atrophy seen.
Other syndromes or conditions which can induce skeletal muscle atrophy are congestive heart failure and liver
During aging, there is a gradual decrease in the ability to maintain skeletal muscle function and mass. This condition
is called "sarcopenia", and may be distinct from atrophy in its pathophysiology. While the exact cause of sarcopenia
is unknown, it may be induced by a combination of a gradual failure in the "satellite cells" which help to regenerate
Atrophy 80

skeletal muscle fibers, and a decrease in sensitivity to or the availability of critical secreted growth factors which are
necessary to maintain muscle mass and satellite cell survival.[2]

Dystrophies, myosities, and motor neuron conditions

Pathologic atrophy of muscles can occur due to diseases of the motor nerves, or due to diseases of the muscle tissue
itself. Examples of atrophying nerve diseases include CMT (Charcot Marie Tooth syndrome) poliomyelitis,
amyotrophic lateral sclerosis (ALS or Lou Gehrig's disease), and Guillain-Barré syndrome. Examples of atrophying
muscle diseases include muscular dystrophy, myotonia congenita, and myotonic dystrophy.
Changes in Na+ channel isoform expression and spontaneous activity in muscle called fibrillation can also result in
muscle atrophy.

Gland atrophy
The adrenal glands atrophy during prolonged use of exogenous glucocorticoids like prednisone. Atrophy of the
breasts can occur with prolonged estrogen reduction, as with anorexia nervosa or menopause. Atrophy of the testes
occurs with prolonged use of enough exogenous sex steroid (either androgen or estrogen) to reduce gonadotropin

Vaginal atrophy
In post-menopausal women, the walls of the vagina become thinner (Atrophic vaginitis). The mechanism for the
age-related condition is not yet clear, though there are theories that the effect is caused by decreases in estrogen
levels.[3] This atrophy, and that of the breasts concurrently, is consistent with the homeostatic (normal development)
role of atrophy in general, as after menopause the body has no further functional biological need to maintain the
reproductive system which it has permanently shut down.

It has been reported that some drugs might prevent the loss of the muscle wasting that occurs in immobile, bedridden
patients.[4] Testing upon mice showed that it blocked the activity of a protein present in the muscle that is involved
in muscle atrophy.[5] However the concerns for the drug's longterm effects on the heart preclude its routine use in
humans for this indication, and further alternative drugs are being sought.[4] Not exercising is the main cause for
muscle atrophy.

[1] http:/ / www. nlm. nih. gov/ cgi/ mesh/ 2010/ MB_cgi?field=uid& term=D001284
[2] Campellone, Joseph V. (2007-05-22). "Muscle atrophy" (http:/ / www. nlm. nih. gov/ medlineplus/ ency/ article/ 003188. htm) (html).
MedlinePlus. . Retrieved 2007-10-02.
[3] "Types of Atrophy" (http:/ / medicine. science-tips. org/ health/ diseases-and-conditions/ types-of-atrophy. html) (html). . Retrieved
[4] "Drug could stop muscle wasting'" (http:/ / www2. netdoctor. co. uk/ news/ index. asp?y=2006& m=5& d=25& id=123026). 2006-05-25. . Retrieved 2006-05-27.
[5] Wang X, Hockerman GH, Green Iii HW, Babbs CF, Mohammad SI, Gerrard D, Latour MA, London B, Hannon KM, Pond AL (May24
2006). "Merg1a K+ channel induces skeletal muscle atrophy by activating the ubiquitin proteasome pathway". FASEB J 20 (9): 1531.
doi:10.1096/fj.05-5350fje. PMID 16723379.
Rohmert's law 81

Rohmert's law
Widely used in the human factors and ergonomics field, Rohmert's law states that the maximum force one's muscles
can exert decreases exponentially from the time one begins continuously exerting the said force. It is commonly used
to calculate "maximum holding time" for any particular task.
Maximum force decays exponentially due to the amount of energy (in the form of oxygen and ATP) the body is able
to supply to the muscles. The circulatory systems keeps muscles flooded in nutrients at all times, so that muscles
have a supply of fuel ready to burn at any given moment. A task requiring maximum force burns a large amount of
those nutrients at the onset of the task; the circulatory system is then unable to replenish the nutrients at a rate fast
enough to maintain maximum force for long. As a result, the maximum force the muscle is capable of producing is
limited by the bottleneck in nutrient availability, and decreases exponentially.
Imagine a theoretical arm wrestling match with two perfectly matched opponents, each exactly as strong as each
other. They both begin the match by exerting maximum force on each other's hands, but very soon, their arms get
fatigued and the actual force being exerted on each others' hands drops off quickly. They are still exerting as much
force as they can, but their muscles are burning energy faster than can be replenished, and their maximum force is
decreasing exponentially. Eventually their arms are completely fatigued; they are basically just holding hands and
applying what little force their muscles can muster, wondering when the other will give up.
While Rohmert's law applies to maximum force, the inverse is true as well; the less force one is asked to exert, the
longer one will be able to exert that force before their muscles become fatigued. If one is asked to exert zero force,
they can theoretically hold the position indefinitely.
Rohmert's law has been found to be true across all humans. While everyone has a different initial maximum force
they can apply, their maximum force will decrease according to the same exponential curve as everyone else.
Muscular system 82

Muscular system
Muscular system

Muscles anterior labeled

Muscle posterior labeled

Latin systema musculare

The muscular system is the anatomical system of a species that allows it to move. The muscular system in
vertebrates is controlled through the nervous system, although some muscles (such as the cardiac muscle) can be
completely autonomous.

There are three distinct types of muscles: skeletal muscles, cardiac or heart muscles, and smooth (non-striated)
muscles. Muscles provide strength, balance, posture, movement and heat for the body to keep warm.
Upon stimulation by an action potential, skeletal muscles perform a coordinated contraction by shortening each
sarcomere. The best proposed model for understanding contraction is the sliding filament model of muscle
contraction. Actin and myosin fibers overlap in a contractile motion towards each other. Myosin filaments have
club-shaped heads that project toward the actin filaments.
Larger structures along the myosin filament called myosin heads are used to provide attachment points on binding
sites for the actin filaments. The myosin heads move in a coordinated style, they swivel toward the center of the
sarcomere, detach and then reattach to the nearest active site of the actin filament. This is called a rachet type drive
system. This process consumes large amounts of adenosine triphosphate (ATP).
Energy for this comes from ATP, the energy source of the cell. ATP binds to the cross bridges between myosin
heads and actin filaments. The release of energy powers the swiveling of the myosin head. Muscles store little ATP
and so must continuously recycle the discharged adenosine diphosphate molecule (ADP) into ATP rapidly. Muscle
tissue also contains a stored supply of a fast acting recharge chemical, creatine phosphate which can assist initially
producing the rapid regeneration of ADP into ATP.
Calcium ions are required for each cycle of the sarcomere. Calcium is released from the sarcoplasmic reticulum into
the sarcomere when a muscle is stimulated to contract. This calcium uncovers the actin binding sites. When the
Muscular system 83

muscle no longer needs to contract, the calcium ions are pumped from the sarcomere and back into storage in the
sarcoplasmic reticulum.

There are approximately 639 skeletal muscles in the human body.
The following are some major muscles[1] and their basic features:

Muscle Origin Insertion Artery Nerve Action Antagonist

gastrocnemius femur calcaneus sural arteries tibial nerve plantarflexion, Tibialis anterior
flexion of knee muscle

tibialis posterior tibia, fibula Foot posterior tibial tibial nerve inversion of the foot, Tibialis anterior
artery plantar flexion of the muscle
foot at the ankle

soleus fibula, medial calcaneus sural arteries tibial nerve plantarflexion Tibialis anterior
border of tibia muscle

tibialis anterior tibia foot anterior tibial Fibular nerve dorsiflex and invert Fibularis longus,
artery the foot Gastrocnemius,
Soleus, Plantaris,
Tibialis posterior

longus fibula Foot fibular artery Superficial fibular plantarflexion, Tibialis anterior
nerve eversion muscle

brevis fibula Foot, eversion peroneal artery superficial peroneal


gluteus maximus ilium, sacrum, Gluteal gluteal arteries inferior gluteal nerve external rotation and Iliacus, Psoas
muscle sacrotuberous tuberosity of the extension of the hip major, Psoas minor
ligament femur joint

biceps femoris ischium, femur fibula inferior gluteal tibial nerve, common flexes and laterally Quadriceps muscle
artery, popliteal peroneal nerve rotates knee joint,
artery extends hip joint

semitendinosus ischium tibia inferior gluteal sciatic flex knee, extend hip Quadriceps muscle
artery joint

semimembranosus ischium tibia profunda femoris, sciatic nerve Hip extension, Knee Quadriceps muscle
gluteal artery flexion

iliopsoas ilium femur medial femoral femoral nerve, lumbar flexion of hip Gluteus maximus,
circumflex artery, nerves posterior
iliolumbar artery compartment of

quadriceps femoris combined rectus femoral artery Femoral nerve Knee extension; Hip Hamstring
femoris and flexion
vastus muscles

adductor muscles pubis femur, tibia obturator nerve adduction of hip

of the hip

levator scapulae vertebral column scapula dorsal scapular cervical nerve, dorsal Elevates scapula,
artery scapular nerve tilts its glenoid
cavity inferiorly

trapezius the rear of the clavicle, scapula cranial nerve XI, retraction of scapula Serratus anterior
skull, vertebral cervical nerves muscle
Muscular system 84

rectus abdominis pubis Costal cartilage inferior epigastric segmentally by flexion of Erector spinae
of ribs 5-7, artery thoraco-abdominal trunk/lumbar
sternum nerves vertebrae

transversus ribs, ilium pubic tubercle lower intercostal compress the ribs
abdominis nerves, iliohypogastric and viscera, thoracic
nerve and the and pelvic stability
ilioinguinal nerve

Abdominal Lower 8 costae Crista iliaca, lower 6 intercostal Rotates torso

external oblique ligamentum nerve, subcostal nerve
muscle inguinale

Abdominal internal Inguinal Linea alba, Compresses

oblique muscle ligament, Iliac sternum and the abdomen and rotates
crest and the inferior ribs. vertebral column.

erector spinae on the spines of both the spines lateral sacral posterior branch of extends the vertebral Rectus abdominis
the last four of the most artery spinal nerve column muscle
thoracic vertebræ cranial thoracic
vertebrae and
the cervical

pectoralis major clavicle, humerus thoracoacromial lateral pectoral nerve Clavicular head:
sternum, costal trunk and medial pectoral flexes the humerus
cartilages nerve Sternocostal head:
extends the humerus
As a whole, adducts
and medially rotates
the humerus. It also
draws the scapula
anteriorly and

biceps brachii scapula radius brachial artery Musculocutaneous flexes elbow and Triceps brachii
nerve supinates forearm muscle

triceps brachii scapula and ulna deep brachial radial nerve extends forearm, Biceps brachii
humerus artery caput longum muscle
adducts shoulder

brachialis humerus ulna radial recurrent musculocutaneous flexion at elbow joint

artery nerve

pronator teres humerus, ulna radius ulnar artery and median nerve pronation of forearm, Supinator muscle
radial artery flexes elbow

brachioradialis humerus radius radial recurrent radial nerve Flexion of forearm


rhomboids nuchal scapula dorsal scapular dorsal scapular nerve Retracts the scapula Serratus anterior
ligaments, artery and rotates it to muscle
spinous depress the glenoid
processes of the cavity. fixes the
C7 to T5 scapula to the
vertebrae thoracic wall.

deltoid clavicle, deltoid primarily posterior Axillary nerve shoulder abduction, Latissimus dorsi
acromion, tuberosity of circumflex flexion and
scapula humerus humeral artery extension
Muscular system 85

latissimus dorsi vertebral humerus subscapular artery, thoracodorsal nerve pulls the forelimb deltoid, trapezius
column, ilium dorsal scapular dorsally and caudally
and inferior 3 or artery
4 ribs

Aerobic and anaerobic muscle activity

At rest, the body produces the majority of its ATP aerobically in the mitochondria[2] without producing lactic acid or
other fatiguing byproducts.[3] During exercise, the method of ATP production varies depending on the fitness of the
individual as well as the duration, and intensity of exercise. At lower activity levels, when exercise continues for a
long duration (several minutes or longer), energy is produced aerobically by combining oxygen with carbohydrates
and fats stored in the body. Activity that is higher in intensity, with possible duration decreasing as intensity
increases, ATP production can switch to anaerobic pathways, such as the use of the creatine phosphate and the
phosphagen system or anaerobic glycolysis. Aerobic ATP production is biochemically much slower and can only be
used for long-duration, low intensity exercise, but produces no fatiguing waste products that can not be removed
immediately from sarcomere and body and results in a much greater number of ATP molecules per fat or
carbohydrate molecule. Aerobic training allows the oxygen delivery system to be more efficient, allowing aerobic
metabolism to begin quicker.[3] Anaerobic ATP production produces ATP much faster and allows near-maximal
intensity exercise, but also produces significant amounts of lactic acid which render high intensity exercise
unsustainable for greater than several minutes.[3] The phosphagen system is also anaerobic, allows for the highest
levels of exercise intensity, but intramuscular stores of phosphocreatine are very limited and can only provide energy
for exercises lasting up to ten seconds. Recovery is very quick, with full creatine stores regenerated within five

Cardiac muscle
Heart muscles are distinct from skeletal muscles because the muscle fibers are laterally connected to each other.
Furthermore, just as with smooth muscles, they are not controlling themselves. Heart muscles are controlled by the
sinus node influenced by the autonomic nervous system.

Smooth muscle
Smooth muscles are controlled directly by the autonomic nervous system and are involuntary, meaning that they are
incapable of being moved by conscious thought. Functions such as heart beat and lungs (which are capable of being
willingly controlled, be it to a limited extent) are involuntary muscles but are not smooth muscles.

Control of muscle contraction

Neuromuscular junctions are the focal point where a motor neuron attaches to a muscle. Acetylcholine, (a
neurotransmitter used in skeletal muscle contraction) is released from the axon terminal of the nerve cell when an
action potential reaches the microscopic junction, called a synapse. A group of chemical messengers cross the
synapse and stimulate the formation of electrical changes, which are produced in the muscle cell when the
acetylcholine binds to receptors on its surface. Calcium is released from its storage area in the cell's sarcoplasmic
reticulum. An impulse from a nerve cell causes calcium release and brings about a single, short muscle contraction
called a muscle twitch. If there is a problem at the neuromuscular junction, a very prolonged contraction may occur,
tetanus. Also, a loss of function at the junction can produce paralysis.
Skeletal muscles are organized into hundreds of motor units, each of which involves a motor neuron, attached by a
series of thin finger-like structures called axon terminals. These attach to and control discrete bundles of muscle
fibers. A coordinated and fine tuned response to a specific circumstance will involve controlling the precise number
of motor units used. While individual muscle units contract as a unit, the entire muscle can contract on a
Muscular system 86

predetermined basis due to the structure of the motor unit. Motor unit coordination, balance, and control frequently
come under the direction of the cerebellum of the brain. This allows for complex muscular coordination with little
conscious effort, such as when one drives a car without thinking about the process.

[1] List of major muscles of the human body (http:/ / www. cylive. com/ viewContent. do?id=192& vt=pub)
[2] ABERCROMBIE M; HICKMAN C.J; JOHNSON M.L, 1973, A Dictionary of Biology, Page 179, Middlesex (England), Baltimore (U.S.A),
Ringwood (Australia): Penguin Books
[3] "St Paul’s College Stage 2 EXERCISE PHYSIOLOGY Energy Systems Part 5" (http:/ / intranet. spc. sa. edu. au/ usermedia/ curr/ pe/ ep5.
ppt) (ppt). . Retrieved 2007-10-16.

• Online Muscle Tutorial (
Medical and Health Encyclopedia, chapter 1

External links
• GetBody Smart ( Muscle system
tutorials and quizzes
• ( 6/muscle_metabolism_march_2007.htm) Use and
formation of ATP in muscle

Human musculoskeletal system

A musculoskeletal system (also known as the locomotor system) is an organ system that gives animals (including
humans) the ability to move using the muscular and skeletal systems. The musculoskeletal system provides form,
support, stability, and movement to the body.
It is made up of the body's bones (the skeleton), muscles, cartilage,[1] tendons, ligaments, joints, and other connective
tissue that supports and binds tissues and organs together. The musculoskeletal system's primary functions include
supporting the body, allowing motion, and protecting vital organs.[2] The skeletal portion of the system serves as the
main storage system for calcium and phosphorus and contains critical components of the hematopoietic system.[3]
This system describes how bones are connected to other bones and muscle fibers via connective tissue such as
tendons and ligaments. The bones provide the stability to a body in analogy to iron rods in concrete construction.
Muscles keep bones in place and also play a role in movement of the bones. To allow motion, different bones are
connected by joints. Cartilage prevents the bone ends from rubbing directly on to each other. Muscles contract
(bunch up) to move the bone attached at the joint.
There are, however, diseases and disorders that may adversely affect the function and overall effectiveness of the
system. These diseases can be difficult to diagnose due to the close relation of the musculoskeletal system to other
internal systems. The musculoskeletal system refers to the system having its muscles attached to an internal skeletal
system and is necessary for humans to move to a more favorable position. Complex issues and injuries involving the
musculoskeletal system are usually handled by a physiatrist (specialist in Physical Medicine and Rehabilitation) or
an orthopaedic surgeon.
Human musculoskeletal system 87


The Skeletal System serves many important functions; it provides the
shape and form for our bodies in addition to supporting, protecting,
allowing bodily movement, producing blood for the body, and storing
minerals.[4] The number of bones in the human skeletal system is a
controversial topic. Humans are born with about 300 to 350 bones,
however, many bones fuse together between birth and maturity. As a
result an average adult skeleton consists of 206 bones. The number of
bones varies according to the method used to derive the count. While
some consider certain structures to be a single bone with multiple
parts, others may see it as a single part with multiple bones.[5] There
are five general classifications of bones. These are Long bones, Short
bones, Flat bones, Irregular bones, and Sesamoid bones. The human
skeleton is composed of both fused and individual bones supported by
ligaments, tendons, muscles and cartilage. It is a complex structure
with two distinct divisions. These are the axial skeleton and the
appendicular skeleton.[6]


The Skeletal System serves as a framework for tissues and organs to

attach themselves to. This system acts as a protective structure for vital
organs. Major examples of this are the brain being protected by the
Front view of a skeleton of an adult human
skull and the lungs being protected by the rib cage.
Located in long bones are two distinctions of bone marrow (yellow and
red). The yellow marrow has fatty connective tissue and is found in the marrow cavity. During starvation, the body
uses the fat in yellow marrow for energy.[7] The red marrow of some bones is an important site for blood cell
production, approximately 2.6 million red blood cells per second in order to replace existing cells that have been
destroyed by the liver.[4] Here all erythrocytes, platelets, and most leukocytes form in adults. From the red marrow,
erythrocytes, platelets, and leukocytes migrate to the blood to do their special tasks.

Another function of bones is the storage of certain minerals. Calcium and phosphorus are among the main minerals
being stored. The importance of this storage "device" helps to regulate mineral balance in the bloodstream. When the
fluctuation of minerals is high, these minerals are stored in bone; when it is low it will be withdrawn from the bone.

There are three types of muscles—cardiac, skeletal, and smooth.
Smooth muscles are used to control the flow of substances within the
lumens of hollow organs, and are not consciously controlled. Skeletal
Types of muscle and their appearance
and cardiac muscles have striations that are visible under a microscope
due to the components within their cells. Only skeletal and smooth
muscles are part of the musculoskeletal system and only the skeletal muscles can move the body. Cardiac muscles
are found in the heart and
Human musculoskeletal system 88

are used only to circulate blood; like the smooth muscles, these
muscles are not under conscious control. Skeletal muscles are attached
to bones and arranged in opposing groups around joints.[8] Muscles are
innervated, to communicate nervous energy to,[9] by nerves, which
conduct electrical currents from the central nervous system and cause
the muscles to contract.[10]

Contraction initiation

In mammals, when a muscle contracts, a series of reactions occur.

Muscle contraction is stimulated by the motor neuron sending a
message to the muscles from the somatic nervous system.
Depolarization of the motor neuron results in neurotransmitters being
released from the nerve terminal. The space between the nerve terminal
and the muscle cell is called the neuromuscular junction. These
neurotransmitters diffuse across the synapse and bind to specific
receptor sites on the cell membrane of the muscle fiber. When enough
receptors are stimulated, an action potential is generated and the
permeability of the sarcolemma is altered. This process is known as


A tendon is a tough, flexible band of fibrous connective tissue that

connects muscles to bones.[12] The extra-cellular connective tissue
between muscle fibers binds to tendons at the distal & proximal ends,
and the tendon binds to the periosteum of individual bones at the
muscle's origin & insertion. As muscles contract, tendons transmit the
forces to the rigid bones, pulling on them and causing movement.
Tendons can stretch substantially, allowing them to function as springs
during locomotion, thereby saving energy.

Joints, ligaments, and bursae

Human musculoskeletal system 89


Joints are structures that connect individual bones and may allow
bones to move against each other to cause movement. There are two
divisions of joints, diarthroses which allow extensive mobility between
two or more articular heads, and false joints or synarthroses, joints that
are immovable, that allow little or no movement and are predominantly
fibrous. Synovial joints, joints that are not directly joined, are
lubricated by a solution called synovial Fluid that is produced by the
synovial membranes. This fluid lowers the friction between the
articular surfaces and is kept within an articular capsule, binding the
joint with its taut tissue.[6]


A ligament is a small band of dense, white, fibrous elastic tissue.[6] Human synovial joint composition
Ligaments connect the ends of bones together in order to form a joint.
Most ligaments limit dislocation, or prevent certain movements that may cause breaks. Since they are only elastic
they increasingly lengthen when under pressure. When this occurs the ligament may be susceptible to break resulting
in an unstable joint.
Ligaments may also restrict some actions: movements such as hyper extension and hyper flexion are restricted by
ligaments to an extent. Also ligaments prevent certain directional movement.[13]

A bursa is a small fluid-filled sac made of white fibrous tissue and lined with synovial membrane. Bursa may also be
formed by a synovial membrane that extends outside of the joint capsule.[7] It provides a cushion between bones and
tendons and/or muscles around a joint; bursa are filled with synovial fluid and are found around almost every major
joint of the body.

Diseases and disorders

Because many other body
systems, including the
vascular, nervous, and
integumentary systems,
are interrelated, disorders
of one of these systems Disability-adjusted life year for musculoskeletal diseases per 100,000 inhabitants in 2004. "WHO
may also affect the Disease and injury country estimates". World Health Organization. 2009. . Retrieved Nov. 11, 2009.  no
musculoskeletal system data  less than
400  400-450  450-500  500-550  550-600  600-650  650-700  700-850  850-900  900-925  925-950  more
and complicate the
than 950
diagnosis of the disorder's
origin. Diseases of the
musculoskeletal system mostly encompass functional disorders or motion discrepancies; the level of impairment
depends specifically on the problem and its severity. Articular (of or pertaining to the joints)[15] disorders are the
most common. However, also among the diagnoses are: primary muscular diseases, neurologic (related to the
medical science that deals with the nervous system and disorders affecting it)[16] deficits, toxins, endocrine

abnormalities, metabolic disorders, infectious diseases, blood and vascular disorders, and nutritional imbalances.
Disorders of muscles from another body system can bring about irregularities such as: impairment of ocular motion
Human musculoskeletal system 90

and control, respiratory dysfunction, and bladder malfunction. Complete paralysis, paresis, or ataxia may be caused
by primary muscular dysfunctions of infectious or toxic origin; however, the primary disorder is usually related to
the nervous system, with the muscular system acting as the effector organ, an organ capable of responding to a
stimulus, especially a nerve impulse.[3] One understated disorder that begins during pregnancy is Pelvic girdle pain),
it is complex and multi-factorial and likely to be also represented by a series of sub-groups driven by pain varying
from peripheral or central nervous system,[17] altered laxity/stiffness of muscles,[18] laxity to injury of
tendinous/ligamentous structures [19] to ‘mal-adaptive’ body mechanics.[17]

[1] MeSH Musculoskeletal+System (http:/ / www. nlm. nih. gov/ cgi/ mesh/ 2011/ MB_cgi?mode=& term=Musculoskeletal+ System)
[2] Mooar, Pekka (2007). "Introduction" (http:/ / www. merck. com/ mmhe/ sec05/ ch058/ ch058a. html). Merck Manual. . Retrieved
[3] Kahn, Cynthia; Scott Line (2008). Musculoskeletal System Introduction: Introduction (http:/ / www. merckvetmanual. com/ mvm/ index.
jsp?cfile=htm/ bc/ 90100. htm). NJ, USA: Merck & Co., Inc.. .
[4] Applegate, Edith; Kent Van De Graaff. "The Skeletal System" (http:/ / www. mnsu. edu/ emuseum/ biology/ humananatomy/ skeletal/
skeletalsystem. html). . Retrieved 2009-01-03.
[5] Engelbert, Phillis; Carol DeKane Nagel (2009). "The Human Body / How Many Bones Are In The Human Body?" (http:/ / www. enotes.
com/ science-fact-finder/ human-body/ how-many-bones-human-body). U·X·L Science Fact Finder., Inc.. . Retrieved 2009-01-24.
[6] Gary, Farr (2002-06-25). "The Musculoskeletal System" (http:/ / www. becomehealthynow. com/ category/ bodymusculo/ ). . Retrieved
[7] "Skeletal System" (http:/ / www. besthealth. com/ besthealth/ bodyguide/ reftext/ html/ skel_sys_fin. html). 2001. . Retrieved 2009-01-08.
[8] Mooar, Pekka (2007). "Muscles" (http:/ / www. merck. com/ mmhe/ sec05/ ch058/ ch058c. html). The Merck Manuals Online Medical
Library. . Retrieved 2008-11-16.
[9] "innervated" (http:/ / dictionary. reference. com/ browse/ innervated)., LLC.. 2008. . Retrieved 2009-01-03.
[10] Bárány, Michael (2002). "SMOOTH MUSCLE" (http:/ / www. uic. edu/ classes/ phyb/ phyb516/ index. htm). . Retrieved 2008-11-19.
[11] "The Mechanism of Muscle Contraction" (http:/ / meat. tamu. edu/ muscontract. html). Principles of Meat Science (4th Edition). . Retrieved
[12] Jonathan, Cluett (2008). "Tendons" (http:/ / orthopedics. about. com/ cs/ sportsmedicine/ g/ tendon. htm). . Retrieved 2008-11-19.
[13] Bridwell, Keith (06/07/2008). "Ligaments" (http:/ / www. spineuniverse. com/ displayarticle. php/ article1268. html). . Retrieved
[14] "WHO Disease and injury country estimates" (http:/ / www. who. int/ healthinfo/ global_burden_disease/ estimates_country/ en/ index.
html). World Health Organization. 2009. . Retrieved Nov. 11, 2009.
[15] "articular" (http:/ / dictionary. reference. com/ browse/ articular). Random House Unabridged Dictionary. Random House, Inc.. 2006. .
Retrieved 2008-11-15.
[16] "neurologic" (http:/ / dictionary. reference. com/ browse/ neurologic). The American Heritage Dictionary of the English Language, Fourth
Edition. Houghton Mifflin Company. 2006. . Retrieved 2008-11-15.
[17] Diagnosis and classification of pelvic girdle pain disorders— Part 1: A mechanism based approach within a bio psychosocial framework.
Manual Therapy, Volume 12, Issue 2, May 2007, PB. O’Sullivan and DJ Beales.
[18] European guidelines for the diagnosis and treatment of pelvic girdle pain. Eur Spine J. 2008 Feb 8, A Vleeming, HB Albert, HC Ostgaard, B
Sturesson, B Stuge.
[19] Possible role of the long dorsal sacroiliac ligament in women with peripartum pelvic pain. Acta Obstetricia et Gynecologica Scandinavica,
Volume 81, Issue 5 , Page 430-436, May 2002, A Vleeming, HJ de Vries, JM Mens, J-P van Wingerden
Muscle spindle 91

Muscle spindle
Muscle spindles are sensory receptors
within the belly of a muscle, which
primarily detect changes in the length
of this muscle. They convey length
information to the central nervous
system via sensory neurons. This
information can be processed by the
brain to determine the position of body
parts. The responses of muscle
spindles to changes in length also play
an important role in regulating the
contraction of muscles, by activating
motoneurons via the stretch reflex to Mammalian muscle spindle showing typical position in a muscle (left), neuronal
connections in spinal cord (middle) and expanded schematic (right). The spindle is a
resist muscle stretch.
stretch receptor with its own motor supply consisting of several intrafusal muscle fibres.
The sensory endings of a primary (group Ia) afferent and a secondary (group II) afferent
coil around the non-contractile central portions of the intrafusal fibres. Gamma
Anatomy motoneurons activate the intrafusal muscle fibres, changing the resting firing rate and
stretch-sensitivity of the afferents. Animated version: [1]
Muscle spindles are found within the
belly of muscles, embedded in
extrafusal muscle fibers. Note that "fusus" is the Latin word for spindle. Muscle spindles are composed of 3-12
intrafusal muscle fibers, of which there are three types:
• dynamic nuclear bag fibers (bag1 fibers)
• static nuclear bag fibers (bag2 fibers)
• nuclear chain fibers and the axons of sensory neurons.
Axons of gamma motoneurons also terminate in muscle spindles; they make synapses at either or both of the ends of
the intrafusal muscle fibers and regulate the sensitivity of the sensory afferents, which are located in the
non-contractile central (equatorial) region[2] .
Muscle spindles are encapsulated by connective tissue, and are aligned parallel to extrafusal muscle fibers, unlike
Golgi tendon organs, which are oriented in series.
The muscle spindle has both sensory and motor components.
• Primary and secondary sensory nerve fibers spiral around and terminate on the central portions of the intrafusal
muscle fibers, providing the sensory component of the structure via stretch-sensitive ion-channels of the axons.
• In mammals including humans, the motor component is provided by up to a dozen gamma motoneurons and to a
lesser extent by one or two beta motoneurons. Gamma and beta motoneurons are called fusimotor neurons,
because they activate the intrafusal muscle fibers. Gamma motoneurons only innervate intrafusal muscle fibers,
whereas beta motoneurons innervate both extrafusal and intrafusal muscle fibers and so are referred to as
skeletofusimotor neurons.
• Fusimotor drive causes a contraction and stiffening of the end portions of the intrafusal muscle fibers.
Fusimotor neurons are classified as static or dynamic according to the type of intrafusal muscle fibers they innervate
and their physiological effects on the responses of the Ia and II sensory neurons innervating the central,
non-contractile part of the muscle spindle.
• The static axons innervate the chain or bag2 fibers. They increase the firing rate of Ia and II afferents at a given
muscle length (see schematic of fusimotor action below).
Muscle spindle 92

• The dynamic axons innervate the bag1 intrafusal muscle fibers. They increase the stretch-sensitivity of the Ia
afferents by stiffening the bag1 intrafusal fibers.

Sensitivity modification
The function of the gamma motoneurons is not to supplement the force of muscle contraction provided by the
extrafusal fibers, but to modify the sensitivity of the muscle spindle sensory afferents to stretch. Upon release of
acetylcholine by the active gamma motoneuron, the end portions of the intrafusal muscle fibers contract, thus
elongating the non-contractile central portions (see "fusimotor action" schematic below). This opens stretch-sensitive
ion channels of the sensory endings, leading to an influx of sodium ions. This raises the resting potential of the
endings, thereby increasing the probability of action potential firing, thus increasing the stretch-sensitivity of the
muscle spindle afferents. For an interactive animation created by Jan Kowalczewski at the University of Alberta,
demonstrating spindle afferent responses to muscle stretch with and without gamma (fusimotor) action, go to: [1].
How does the central nervous system control gamma fusimotor neurons? It has been difficult to record from gamma
motoneurons during normal movement because they have very small axons. Several theories have been proposed,
based on recordings from spindle afferents.
• 1) Alpha-gamma coactivation. Here it is posited that gamma motoneurons are activated in parallel with alpha
motoneurons to maintain the firing of spindle afferents when the extrafusal muscles shorten.[3]
• 2) Fusimotor set: gamma motoneurons are activated according to the novelty or difficulty of a task. Whereas
static gamma motoneurons are continuously active during routine movements such as locomotion, dynamic
gamma motoneoruns tend to be activated more during difficult tasks, increasing Ia stretch-sensitivity[4] .
• 3) Fusimotor template of intended movement. Static gamma activity is a "temporal template" of the expected
shortening and lengthening of the receptor-bearing muscle. Dynamic gamma activity turns on and off abruptly,
sensitizing spindle afferents to the onset of muscle lengthening and departures from the intended movement
trajectory[5] .

Stretch reflex
When a muscle is stretched, primary sensory fibers (Group Ia afferent neurons) of the muscle spindle respond to both
changes in muscle length and velocity and transmit this activity to the spinal cord in the form of changes in the rate
of action potentials. Likewise, secondary sensory fibers (Group II afferent neurons) respond to muscle length
changes (but with a smaller velocity-sensitive component) and transmit this signal to the spinal cord. The Ia afferent
signals are transmitted monosynaptically to many alpha motor neurons of the receptor-bearing muscle. The
reflexly-evoked activity in the alpha motoneurons is then transmitted via their efferent axons to the extrafusal fibers
of the muscle, which generate force and thereby resist the stretch. The Ia afferent signal is also transmitted
polysynaptically through interneurons (Renshaw_cells) which inhibit alpha motoneurons of antagonist muscles,
causing them to relax.
After stroke or spinal cord injury in humans, spastic hypertonus often develops, whereby the stretch reflex in flexor
muscles of the arms and extensor muscles of the legs is overly sensitive. This results in abnormal postures, stiffness
and contractures. Hypertonus may be the result of over-sensitivity of alpha motoneurons and interneurons to the Ia
and II afferent signals[6] .
PNF stretching, or proprioceptive neuromuscular facilitation, is a method of flexibility training that can reduce
hypertonus, allowing muscles to relax and lengthen.
Muscle spindle 93

It is also believed that muscle spindles play a critical role in sensorimotor development.

See also
• Type Ia sensory fiber
• Type II sensory fiber
• Alpha motor neuron
• Gamma motor neuron
• Beta motor neuron
• Intrafusal muscle fibre
• Extrafusal muscle fiber

Additional images

Muscle spindle Gamma fiber 1A fiber Alpha


schematic of fusimotor action

[1] http:/ / www. ualberta. ca/ ~aprochaz/ research_interactive_receptor_model. html
[2] Hulliger M. The mammalian muscle spindle and its central control. Reviews of Physiology Biochemistry & Pharmacology 101: 1-110, 1984.
[3] Vallbo AB, and al-Falahe NA. Human muscle spindle response in a motor learning task. J Physiol (Lond) 421: 553-568, 1990
[4] Prochazka A. Proprioceptive feedback and movement regulation. In: Exercise: Regulation and Integration of Multiple Systems, edited by
Rowell L, and Sheperd JT. New York: American Physiological Society, 1996, p. 89-127.
[5] Taylor A, Durbaba R, Ellaway PH, and Rawlinson S. Static and dynamic gamma-motor output to ankle flexor muscles during locomotion in
the decerebrate cat. J Physiol 571: 711-723, 2006.
[6] Heckmann CJ, Gorassini MA, and Bennett DJ. Persistent inward currents in motoneuron dendrites: implications for motor output. Muscle
Nerve 31: 135-156, 2005.
Muscle spindle 94

External links
• MeSH Muscle+Spindles (
(http:/ / www. kcl. ac. uk/ teares/ gktvc/ vc/ lt/ mspindle/ ) (http:/ / www. ualberta. ca/ ~aprochaz/

Type Ia sensory fiber

Type Ia Sensory Fiber also called Primary
Afferent Fiber is a type of sensory fiber. It is a
component of a muscle fiber's muscle spindle which
keeps track of how fast a muscle stretch changes
(the velocity of the stretch).

Function of muscle spindles

A muscle spindle, with γ motor and Ia sensory fibers
In order to control movements, the nervous system
must receive continuous sensory information from muscles and joints. For this purpose the body has specialized
sensory receptors called proprioceptors. Muscle spindles are a type of proprioceptor, and they are located inside the
muscle itself. They are sensitive to muscle length because they are in parallel with the contractile fibers.

Types of sensory fibers

This change in length of the spindle is transduced (transformed into electric membrane potentials) by two types of
sensory afferents, whose cell bodies are located in dorsal root ganglia located next to the spinal cord.
The two kinds of sensory fibers are different in respect to the kind of potentials they generate:

Type Primary/secondary Response

Type Ia primary Respond to the rate of change in muscle length, as well to change in length

Type Ib N/A in Golgi tendon organ

Type II secondary Respond only to changes in length

The first of the two main groups of stretch receptors wrapping the intrafusal fibers are the Ia fiber, which are the
largest and fastest fibers, and they fire when the muscle is stretching. They are characterized by their rapid
adaptation, because as soon as the muscle stops changing length, the Ia stop firing and adapt to the new length. Ia
fibers essentially supply proprioceptive information about the rate of change of its respective muscle: the derivative
of the muscle's length (or position).
Type Ia fibers connect to both nuclear bag fibers and nuclear chain fibers. These connections are also called
"annulospiral endings ".
Type Ia sensory fiber 95

Efferent innervation
In addition, the spindle also has a motor efferent innervation carried by gamma motor neurons, which is used by the
nervous system to modify the spindle's sensitivity.

Termination of afferents
Ia afferents from the muscle spindle terminate on the proximal dendrites of motor neurones.

External links
• Lecture notes [1] from John D.C. Lambert on neurophysiology.

[1] http:/ / www. fi. au. dk/ jl/ lect. html

Type II sensory fiber

Type II sensory fiber (group Aβ) is a type of sensory fiber, the second of the two main groups of stretch receptors.
They are non-adapting, meaning that they keep responding even when the muscle has stopped changing its length.
They are the second most highly myelinated fibers in the body.
Their firing rate is directly related to the muscle's instantaneous length, or position. This information would indicate
the position of one's leg once it has stopped moving. They do not respond to rate of length changes as do the Ia
Type II fibers connect to nuclear chain fibers, but not to nuclear bag fibers. These connections, referred to as "flower
spray endings" due to their appearance, embed into the poles (ends) of the fibre. It is thought that the relative
position of the equatorial regions of the spray when stretched determines the action potential output.
Alpha motor neuron 96

Alpha motor neuron

Neuron: Alpha motor neuron

Alpha motor neurons are derived from the basal plate (basal lamina) of the developing embryo.
Gray's [1]
subject #190

NeuroLex ID [2]

Dorlands/Elsevier [3]

Alpha motor neurons (α-MNs) are large lower motor neurons of the brainstem and spinal cord. They innervate
extrafusal muscle fibers of skeletal muscle and are directly responsible for initiating their contraction. Alpha motor
neurons are distinct from gamma motor neurons, which innervate intrafusal muscle fibers of muscle spindles.
While their cell bodies are found in the central nervous system (CNS), alpha motor neurons are also considered part
of the somatic nervous system—a branch of the peripheral nervous system (PNS)—because their axons extend into
the periphery to innervate skeletal muscles.
An alpha motor neuron and the muscle fibers it innervates is a motor unit. A motor neuron pool contains the cell
bodies of all the alpha motor neurons involved in contracting a single muscle.

Alpha motor neurons innervating the head and neck are found in the brainstem; the remaining α-MNs innervate the
rest of the body and are found in the spinal cord. Because there are fewer muscles in the head and neck than in the
rest of the body, there are more α-MNs in the spinal cord than in the brainstem.
In general, α-MNs on one side of the brainstem or spinal cord innervate muscles on that same side of body. The one
exception is the trochlear nucleus in the brainstem, which innervates the superior oblique muscle of the eye on the
opposite side of the face.

In the brainstem, α-MNs and other neurons reside within clusters of cells called nuclei, some of which contain the
cell bodies of neurons belonging to the cranial nerves. Not all cranial nerve nuclei contain α-MNs; those that do are
motor nuclei, while others are sensory nuclei. Motor nuclei are found throughout the brainstem—medulla, pons, and
midbrain—and for developmental reasons are found near the midline of the brainstem.
Generally, motor nuclei found higher in the brainstem (ie, more rostral) innervate muscles that are higher on the face.
For example, the oculomotor nucleus contains α-MNs that innervate muscles of the eye, and is found in the
midbrain, the most rostral brainstem component. By contrast, the hypoglossal nucleus, which contains α-MNs that
innervate the tongue, is found in the medulla, the most caudal (ie, towards the bottom) of the brainstem structures.
Alpha motor neuron 97

Spinal cord
In the spinal cord, α-MNs are located within the gray matter that
forms the ventral horn. These α-MNs provide the motor
component of the spinal nerves that innervate muscles of the body.

The corticospinal tract is one of the major descending

pathways from the brain to the α-MNs of the spinal

As in the brainstem, higher segments of the spinal cord contain

α-MNs that innervate muscles higher on the body. For example,
the biceps brachii muscle, a muscle of the arm, is innervated by
α-MNs in spinal cord segments C5, C6, and C7, which are found
rostrally in the spinal cord. On the other hand, the gastrocnemius
muscle, one of the muscles of the leg, is innervated by α-MNs
within segments S1 and S2, which are found caudally in the spinal

Alpha motor neurons are located in a specific region of the spinal

Alpha motor neurons are located in lamina IX cord's gray matter. This region is designated lamina IX in the
according to the Rexed lamina system. Rexed lamina system, which classifies regions of gray matter
based on their cytoarchitecture. Lamina IX is located
predominantly in the medial aspect of the ventral horn, although there is some contribution to lamina IX from a
collection of motor neurons located more laterally. Like other regions of the spinal cord, cells in this lamina are
somatotopically organized, meaning that the position of neurons within the spinal cord is associated with what
muscles they innervate. In particular, α-MNs in the medial zone of lamina IX tend to innervate proximal muscles of
the body, while those in the lateral zone tend to innervate more distal muscles. There is similar somatotopy
associated with α-MNs that innervate flexor and extensor muscles: α-MNs that innervate flexors tend to be located
in the dorsal portion of lamina IX; those that innervate extensors tend to be located more ventrally.
Alpha motor neuron 98

Like other neurons, lower motor neurons have both afferent (incoming) and efferent (outgoing) connections. Alpha
motor neurons receive input from a number of sources, including upper motor neurons, sensory neurons, and
interneurons. The primary output of α-MNs is to extrafusal muscle fibers. This afferent and efferent connectivity is
required to achieve coordinated muscle activity.

Afferent input

Selected pathways between upper motor neurons and alpha motor neurons
UMN origin α-MN target Tract name

Cerebral cortex Brainstem Corticonuclear tract

Cerebral cortex Spinal cord Corticospinal tract

Red nucleus Spinal cord Rubrospinal tract

Vestibular nuclei Spinal cord Vestibulospinal tract

Midbrain tectum Spinal cord Tectospinal tract

Reticular formation Spinal cord Reticulospinal tract

Upper motor neurons (UMNs) send input to α-MNs via several pathways, including (but not limited to) the
corticonuclear, corticospinal, and rubrospinal tracts. The corticonuclear and corticospinal tracts are commonly
encountered in studies of upper and lower motor neuron connectivity in the control of voluntary movements.
The corticonuclear tract is so named because it connects the cerebral cortex to cranial nerve nuclei. (The
corticonuclear tract is also called the corticobulbar tract, as the brainstem is sometimes called the "bulb" of the
brain.) It is via this pathway that upper motor neurons from the cortex descend from the cortex and synapse on
α-MNs of the brainstem. Similarly, UMNs of the cerebral cortex are in direct control of α-MNs of the spinal cord via
the lateral and ventral corticospinal tracts.
The sensory input to α-MNs is extensive and has its origin in Golgi tendon organs, muscle spindles,
mechanoreceptors, thermoreceptors, and other sensory neurons in the periphery. These connections provide the
structure for the neural circuits that underlie reflexes. There are several types of reflex circuits, the simplest of which
consists of a single synapse between a sensory neuron and a α-MNs. The knee-jerk reflex is an example of such a
monosynaptic reflex.
The most extensive input to α-MNs is from local interneurons, which are the most numerous type of neuron in the
spinal cord. Among their many roles, interneurons synapse on α-MNs to create more complex reflex circuitry. One
type of interneuron is the Renshaw cell, discussed later.

Efferent output
Alpha motor neurons send fibers that mainly synapse on extrafusal muscle fibers. Other fibers from α-MNs synapse
on Renshaw cells, inhibitory interneurons that synapse on the α-MN and limit its activity in order to prevent muscle

Like other neurons, α-MNs transmit signals as action potentials, rapid changes in electrical activity that propagate
from the cell body to the end of the axon. To increase the speed at which action potentials travel, α-MN axons have
large diameters and are heavily myelinated by both oligodendrocytes and Schwann cells. Oligodendrocytes
myelinate the part of the α-MN axon that lies in the central nervous system (CNS), while Schwann cells myelinate
Alpha motor neuron 99

the part that lies in the peripheral nervous system (PNS). The transition between the CNS and PNS occurs at the
level of the pia mater, the innermost and most delicate layer of meningeal tissue surrounding components of the
The axon of an α-MN connects with its extrafusal muscle fiber via a neuromuscular junction, a specialized type of
chemical synapse that differs both in structure and function from the chemical synapses that connect neurons to each
other. Both types of synapses rely on neurotransmitters to transduce the electrical signal into a chemical signal and
back. One way they differ is that synapses between neurons typically use glutamate or GABA as their
neurotransmitters, while the neuromuscular junction uses acetylcholine exclusively. Acetylcholine is sensed by
nicotinic acetylcholine receptors on extrafusal muscle fibers, causing their contraction.
Like other motor neurons, α-MNs are named after the properties of their axons. Alpha motor neurons have Aα
axons, which are large-caliber, heavily myelinated fibers that conduct action potentials rapidly. By contrast, gamma
motor neurons have Aγ axons, which are slender, lightly myelinated fibers that conduct less rapidly.

Role in disease
Injury to α-MNs is the most common type of lower motor neuron
lesion. Damage may be caused by trauma, ischemia, and infection,
among others. In addition, certain diseases are associated with the
selective loss of α-MNs. For example, poliomyelitis is caused by a
virus that specifically targets and kills motor neurons in the ventral
horn of the spinal cord. Amyotropic lateral sclerosis likewise is
associated with the selective loss of motor neurons.

Paralysis is one of the most pronounced effects of damage to

α-MNs. Because α-MNs provide the only voluntary innervation to
extrafusal muscle fibers, losing α-MNs effectively severs the
connection between the brainstem and spinal cord and the muscles
they innervate. Without this connection, voluntary and involuntary
(reflex) muscle control is impossible. Voluntary muscle control is
lost because α-MNs relay voluntary signals from upper motor Poliomyelitis, caused by the poliovirus seen here, is
neurons to muscle fibers. Loss of involuntary control results from associated with the selective loss of cells within the
interruption of reflex circuits such as the tonic stretch reflex. A ventral horn of the spinal cord, where α-MNs are
consequence of reflex interruption is that muscle tone is reduced,
resulting in flaccid paresis. Another consequence is the depression
of deep tendon reflexes, causing hyporeflexia.

Muscle weakness and atrophy are inevitable consequences of α-MN lesions as well. Because muscle size and
strength are related to the extent of their use, denervated muscles are prone to atrophy. A secondary cause of muscle
atrophy is that denervated muscles are no longer supplied with trophic factors from the α-MNs that innervate them.
Alpha motor neuron lesions also result in abnormal EMG potentials (eg, fibrillation potentials) and fasciculations,
the latter being spontaneous, involuntary muscle contractions.
Diseases that impair signaling between α-MNs and extrafusal muscle fibers, namely diseases of the neuromuscular
junction have similar signs to those that occur with α-MN disease. For example, myasthenia gravis is an autoimmune
disease that prevents signaling across the neuromuscular junction, which results in functional denervation of muscle.
Alpha motor neuron 100

Alpha motor neurons originate in the basal plate, the ventral portion of the neural tube in the developing embryo.
Sonic hedgehog (Shh) is secreted by the nearby notochord and other ventral structures (eg, the floor plate),
establishing a gradient of highly concentrated Shh in the basal plate and less concentrated Shh in the alar plate.
Under the influence of Shh and other factors, some neurons of the basal plate differentiate into α-MNs.
Like other neurons, α-MNs send axonal projections to reach their target extrafusal muscle fibers via axon guidance,
a process regulated in part by neurotrophic factors released by target muscle fibers. Neurotrophic factors also ensure
that each muscle fiber is innervated by the appropriate number of α-MNs. As with most types of neurons in the
nervous system, α-MNs are more numerous in early development than in adulthood. Muscle fibers secrete a limited
amount of neurotrophic factors capable of sustaining only a fraction of the α-MNs that initially project to the muscle
fiber. Those α-MNs that do not receive sufficient neurotrophic factors will undergo apoptosis, a form of programmed
cell death.
Because they innervate many muscles, some clusters of α-MNs receive high concentrations of neurotrophic factors
and survive this stage of neuronal pruning. This is true of the α-MNs innervating the upper and lower limbs: these
α-MNs form large cell columns that contribute to the cervical and lumbar enlargements of the spinal cord. In
addition to receiving neurotrophic factors from muscles, α-MNs also secrete a number of trophic factors to support
the muscle fibers they innervate. Reduced levels of trophic factors contributes to the muscle atrophy that follows an
α-MN lesion.

• John A. Kiernan (2005). Barr's the Human Nervous System: An Anatomical Viewpoint (8th edition ed.).
Hagerstown, MD: Lippincott Williams & Wilkins. ISBN 0-7817-5154-3.
• Duane E. Haines (2004). Neuroanatomy: An Atlas of Structures, Sections, and Systems (6th edition ed.).
Hagerstown, MD: Lippincott Williams & Wilkins. ISBN 0-7817-4677-9.

External links
• NIF Search - Alpha Motor Neuron [4] via the Neuroscience Information Framework

[1] http:/ / education. yahoo. com/ reference/ gray/ subjects/ subject?id=190#p
[2] http:/ / www. neurolex. org/ wiki/ sao1154704263
[3] http:/ / www. mercksource. com/ pp/ us/ cns/ cns_hl_dorlands_split. jsp?pg=/ ppdocs/ us/ common/ dorlands/ dorland/ m_20. htm
[4] http:/ / www. neuinfo. org/ nif/ nifgwt. html?query=%22Alpha%20Motor%20Neuron%22
Gamma motoneuron 101

Gamma motoneuron
Neuron: Gamma motorneuron

A muscle spindle, with γ motor and Ia sensory fibers

NeuroLex ID [1]

Gamma motoneurons (γ-motoneurons), also called gamma motor neurons, are the efferent component of the
fusimotor system, the system by which the central nervous system controls and modifies muscle spindle sensitivity.
The fusimotor system refers to the combination of muscle spindles and γ-motoneurons. The function of the muscle
spindle is to provide proprioceptive feedback for the movement, position and extension of muscles.
γ-motoneurons are located in the brainstem and spinal cord and are smaller than their α-motoneuron counterparts,
which are responsible for controlling skeletal muscle.

Conduction velocity
The axon from a γ-motoneuron is myelinated and has a slower conduction velocity than α-motoneuron axons. The
typical conduction velocity for a γ-motoneuron axon is in the region of 4 to 24 m/s.[2] [3]

Gamma-motoneurons regulate the gain of the stretch reflex by adjusting the level of tension in the intrafusal muscle
fibers of the muscle spindle. This mechanism sets the baseline level of activity in α-motoneurons and helps to
regulate muscle length and tone. For example, stimulation of a γ-motoneuron from higher centers contracts the ends
of the intrafusal fibres and consequently stretches the middle part of the muscle spindle (where Type Ia sensory
fibers are located). These afferent neurons are therefore innervated and go on to synapse with alpha-motoneurons,
This is called the gamma-loop.

Populations of neurons
There are two distinct populations of γ-motoneuron: dynamic γ-motoneurons and static γ-motoneurons.
• Dynamic γ-motoneurons have axons that innervate only dynamic nuclear bag fibres (bag1) .
• Static γ-motoneurons innervate both nuclear chain fibres and static nuclear bag (bag2) fibres.

Effects of nuclear chain fibers

The effect of nuclear chain fibres on primary endings is to drive the discharge up to a frequency of around 60 Hz in a
linear fashion, above which the discharge can become irregular. The activities of bag2 fibres show an initial sharp
peak in discharge, which gets less as the receptor adapts. Bag2 fibres also reduce the dynamic sensitivity of the Ia
afferent and sometimes also reduce the length sensitivity. Activation of bag1 fibres has the effect of increasing both
the length sensitivity and the dynamic sensitivity of the primary ending.[4]
It is believed that the secondary sensory endings serve to measure length and muscle contractions of nuclear chain
fibres at the pole via the static γ-motoneurons both excite the ending and increase its length sensitivity. Bag1 and
bag2 fibres receive very little innervation from secondary endings, and activation of these fibres has a minimal effect
Gamma motoneuron 102

on the discharge of the secondary ending.[4]

[1] http:/ / www. neurolex. org/ wiki/ sao1438006234
[2] Andrew BL, Part NJ (1972) Properties of fast and slow motor units in hind limb and tail muscles of the rat. Q J Exp Physiol Cogn Med Sci
[3] Russell NJ (1980) Axonal conduction velocity changes following muscle tenotomy or deafferentation during development in the rat. J Physiol
[4] Boyd I (1980) The action of the three types of intrafusal fibre in isolated cat muscle spindles on the dynamic and length sensitivities of
primary and secondary sensory endings. In: Muscle Receptors and Movement (Taylor A, Prochazka A, eds), pp 17 - 32. London: MacMillan.

External links
• MeSH Motor+Neurons,+Gamma (
• NIF Search - Gamma Motor Neuron ("gamma Motor
Neuron") via the Neuroscience Information Framework

Beta motor neuron

A beta motor neuron (or beta motoneuron) as alpha motor neuron and gamma motor neuron is a kind of lower motor
neuron. These motor neurons (β-MNs) innervate intrafusal fibers of muscle spindles with collaterals to extrafusal
fibers (type of slow twitch fibers). Axons of beta motor neurons (as alpha and gamma motoneuron) are myelinated.
Alpha, gamma and beta motor neurons originate from the anterior horn of spinal cord and travel to skeletal muscles
(efferent neurons). Alpha motor fibers have a larger diameter and higher conduction velocity than beta and gamma
motor fibers.

Types of beta motoneuron

There are two kinds of beta motoneuron (as gamma motoneuron) that include:
• Static beta motor neurons. These motor neurons innervate nuclear chain fibers of muscle spindles, with collaterals
to extrafusal muscle fibers.
• Dynamic beta motor neurons. The dynamic type innervates nuclear bag fibers of muscle spindles, with collaterals
to extrafusal muscle fibers.
Gamma motoneurons innervate only intrafusal fibers of muscle spindles, but extrafusal fibers (i.e. slow and fast
fibers) are innervated by alpha motoneurons.

• Williams & Warwick. Gray's Anatomy. Thirty-seventh edition.Churchill Livingstone. ISBN: 0-443-04177-6
Intrafusal muscle fiber 103

Intrafusal muscle fiber

Intrafusal muscle fibers (or intrafusal muscle
fibres) are skeletal muscle fibers that comprise the
muscle spindle and are innervated by gamma motor
neurons. These fibers are a proprioceptor that detect
the amount and rate of change of length in a
muscle.[1] These fibers are walled off from the rest
of the muscle by a collagen sheath. This sheath has a
spindle or "fusiform" shape, hence the name A muscle spindle, with γ motor and Ia sensory fibers

There are two types of intrafusal muscle fibers: nuclear bag and nuclear chain fibers. They bear two types of sensory
ending, known as annulospiral and flower-spray endings.
Intrafusal muscle fibers are not to be confused with extrafusal muscle fibers, which are innervated by alpha motor
neurons and contract, generating skeletal movement.
It is by the sensory information from these two intrafusal fiber types that one is able to judge the position of their
muscle, and the rate at which it is changing.

[1] Casagrand, Janet (2008) Action and Movement: Spinal Control of Motor Units and Spinal Reflexes. University of Colorado, Boulder.

External links
• Histology at (
• Histology at (
Extrafusal muscle fiber 104

Extrafusal muscle fiber

Extrafusal muscle fiber is a term given to standard muscle fibers as to distinguish them from intrafusal muscle
fibers. Extrafusal muscle fibers are innervated by alpha motor neurons and generate tension by contracting, thereby
allowing for skeletal movement. Extrafusal muscle fibers are not to be confused with intrafusal muscle fibers which
are innervated by gamma motor neurons and thus serve as a sensory proprioceptor.
The alpha motor neuron and the extrafusal muscle fibers it innervates make up the motor unit. The connection
between the alpha motor neuron and the extrafusal muscle fiber is a neuromuscular junction, where the neuron's
signal, the action potential, is transduced to the muscle fiber by the neurotransmitter acetylcholine.
Basal ganglia 105

Basal ganglia
Brain: Basal ganglia

Basal ganglia labeled at top right.

Latin nuclei basales

NeuroNames hier-206 [1]

MeSH [2]

NeuroLex ID birnlex_826 [3]

The basal ganglia (or basal nuclei) are a group of nuclei of varied origin (mostly telencephalic embryonal origin,
with some diencephalic and mesencephalic elements) in the brains of vertebrates that act as a cohesive functional
unit. They are situated at the base of the forebrain and strongly connected with the cerebral cortex, thalamus and
other brain areas. The basal ganglia are associated with a variety of functions, including voluntary motor control,
procedural learning relating to routine behaviors or "habits," eye movements, and cognitive,[4] emotional
functions.[5] Currently popular theories implicate the basal ganglia primarily in action selection, that is, the decision
of which of several possible behaviors to execute at a given time.[4] [6] Experimental studies show that the basal
ganglia exert an inhibitory influence on a number of motor systems, and that a release of this inhibition permits a
motor system to become active. The "behavior switching" that takes place within the basal ganglia is influenced by
signals from many parts of the brain, including the prefrontal cortex, which plays a key role in executive functions.[5]

The main components of the basal ganglia are the striatum (also called neostriatum) composed of caudate and
putamen, globus pallidus or pallidum composed of globus pallidus externa (GPe) or globus pallidus interna (GPi),
substantia nigra composed of both substantia nigra pars compacta (SNc) & substantia nigra pars reticulata (SNr), and
the subthalamic nucleus (STN).[8] The largest component, the striatum, receives input from many brain areas but
sends output only to other components of the basal ganglia. The pallidum receives its most important input from the
striatum (either directly or indirectly), and sends inhibitory output to a number of motor-related areas, including the
part of the thalamus that projects to the motor-related areas of the cortex. One part of substantia nigra, the reticulata
(SNr), functions similarly to the pallidum, and another part (compacta or SNc) provides the source of the
neurotransmitter dopamine's input to the striatum. The subthalamic nucleus (STN) receives input mainly from the
striatum and cortex, and projects to the a portion of the pallidum (interna portion or GPi). Each of these areas has a
complex internal anatomical and neurochemical organization.
The basal ganglia play a central role in a number of neurological conditions, including several movement disorders.
The most notable are, Parkinson's disease, which involves degeneration of the melanin-pigmented
dopamine-producing cells in the substantia nigra par compacta (SNc), and Huntington's disease, which primarily
involves damage to the striatum.[4] [8] Basal ganglia dysfunction is also implicated in some other disorders of
behavior control such as the Tourette's syndrome, ballismus (particularly hemibalismus), obsessive–compulsive
disorder (OCD), and Wilson's disease (Hepatolenticular degeneration); except for Wilson's disease and
hemiballismus, the neuropathological mechanisms underlying diseases of ganglia such as Parkinsons' and
Basal ganglia 106

Huntington's are not very well understood or are at best still developing theories.
The basal ganglia have a limbic sector whose components are assigned distinct names: the nucleus accumbens (NA),
ventral pallidum, and ventral tegmental area (VTA). VTA efferents provide dopamine to the nucleus accumbens
(ventral striatum) in the same way that the substantia nigra provides dopamine to the dorsal striatum. Because there
is much evidence that it plays a central role in reward learning, the VTA→NA dopaminergic projection has attracted
a great deal of attention. For example, a number of highly addictive drugs, including cocaine, amphetamines, and
nicotine, are thought to work by increasing the efficacy of the VTA→NA dopamine signal. There is also evidence
implicating overactivity of the VTA dopaminergic projection in schizophrenia.[9]

Developmentally, the human nervous system is often classified based on the original 3 primitive vesicles from which
it develops: These primary vesicles form in the normal development of the neural tube of the human fetus and
initially include prosencephalon, mesencephalon, and rhombencephalon, in rostral to caudal (from head to toe)
orientation. Later in development of the nervous system each section itself turns into smaller components. The
following table demonstrates this developmental classification and traces it to the anatomic structures found in the
basal ganglia[4] [8] [10] (the structures relevant to the basal ganglia are shown in bold):

Primary division of the neural Secondary Final segments in a human adult

tube subdivision

Prosencephalon 1. Telencephalon 1. The human cortex (both hemispheres), Caudate, Putamen, Globus pallidus
2. Diencephalon
2. Thalamus, hypothalamus, subthalamus, epithalamus, subthalamic nuclei

Mesencephalon 1. Mesencephalon 1. Mesencephalon (Midbrain), Substantia nigra pars compacta (SNc), Substantia
pars reticulata (SNr)

Rhombencephalon 1. Metencephalon 1. Pons and cerebellum

2. Myelencephalon 2. Medulla

The basal ganglia forms a fundamental

component of the telencephalon
(forebrain). In contrast the cortical
layer that lines the surface of the
forebrain, the basal ganglia is a
collection of distinct masses of gray
matter lying in deep in the brain not far
from the junction of the thalamus. Like
most parts of the brain, the basal
ganglia consist of left and right sides Coronal slices of human brain showing the basal ganglia. White matter is shown in dark
that are virtual mirror images of each gray, gray matter is show in light gray.
Anterior: striatum, globus pallidus (GPe and GPi)
Posterior: subthalamic nucleus (STN), substantia nigra (SN)
Anatomically, the basal ganglia is
divided by the anatomists into four distinct structures depending on how superior or rostral they are (in other words
depending on how close to the top of the head they are): Two of them, the striatum and the pallidum, are relatively
large; the other two, the substantia nigra and the subthalamic nucleus, are smaller. In the illustration to the right, two
coronal sections of the human brain show the location of the basal ganglia components. Of note, and not seen in this
section, the subthalamic nucleus and substantia nigra lie farther back (posteriorly) in the brain than the striatum and
Basal ganglia 107

The striatum is the largest component of the basal ganglia. The term "striatum" comes from the observation that this
structure has a striped appearance when sliced in certain directions, arising from numerous large and small bundles
of nerve fibers (white matter) that traverse it. Early anatomists, examining the human brain, perceived the striatum as
two distinct masses of gray matter separated by a large tract of white matter called the internal capsule. They named
these two masses the "caudate nucleus" and "putamen". More recent anatomists have concluded, on the basis of
microscopic and neurochemical studies, that it is more appropriate to consider these masses as two separated parts of
a single entity, the "striatum", in the same way that a city may be separated into two parts by a river. Numerous
functional differences between the caudate and putamen have been identified, but these are taken to be consequences
of the fact that each sector of the striatum is preferentially connected to specific parts of the cerebral cortex.
The internal organization of the striatum is extraordinarily complex. The great majority of neurons (about 96%) are
of a type called "medium spiny neurons".[4] These are GABAergic cells (meaning that they inhibit their targets) with
small cell bodies and dendrites densely covered with dendritic spines, which receive synaptic input primarily from
the cortex and thalamus. Medium spiny neurons can be divided into subtypes in a number of ways, on the basis of
neurochemistry and connectivity. The next most numerous type (around 2%) are a class of large cholinergic
interneurons with smooth dendrites. There are also several other types of interneurons making up smaller fractions of
the neural population.
Numerous studies have shown that the connections between cortex and striatum are generally topographic; that is,
each part of the cortex sends stronger input to some parts of the striatum than to others. The nature of the topography
has been difficult to understand, however—perhaps in part because the striatum is organized in three dimensions
whereas the cortex, as a layered structure, is organized in two. This dimensional discrepancy entails a great deal of
distortion and discontinuity in mapping one structure to the other. Interestingly, the same topography applies to the
striatal connections to the thalamus.[11]

The pallidum consists of a large structure called the globus pallidus ("pale globe") together with a smaller ventral
extension called the ventral pallidum. The globus pallidus appears as a single neural mass, but can be divided into
two functionally distinct parts, called the internal (sometimes "medial") and external (sometimes "lateral") segments,
abbreviated GPi and GPe.[4] Both segments contain primarily GABAergic neurons, which therefore have inhibitory
effects on their targets. The two segments participate in distinct neural circuits. The external segment, or GPe,
receives input mainly from the striatum, and projects to the subthalamic nucleus. The internal segment, or GPi,
receives signals from the striatum via two pathways, called "direct" and "indirect". Pallidal neurons operate using a
"disinhibition" principle. These neurons fire at steady high rates in the absence of input, and signals from the
striatum cause them to "pause". Because pallidal neurons themselves have inhibitory effects on their targets, the net
effect of striatal input to the pallidum is a reduction of the tonic inhibition exerted by pallidal cells on their targets.

Substantia nigra
Substantia nigra is a mesencephalic gray matter portion of the basal ganglia that is divided into SNr (reticulata) and
SNc (compacta). SNr often works in unison with GPi, and the SNr-GPi complex, inhibits thalamus. Substantia nigra
pars compacta (SNc) however, produces the neurotransmitter dopamine which is very significant in maintaining
balance in the striatal pathway. The circuit portion below explains the role and cirtuit connections of each of the
components of the basal ganglia.
Basal ganglia 108

Subthalamic nucleus
Subthalamic nucleus is a diencephalic gray matter portion of the basal ganglia, and the only portion of the ganglia
that actually produces an "excitatory," Glutamic acid neurotransmitter. The role of the subthalamic nucleus (STN) is
to stimulate the SNr-GPi complex and it is part of the "indirect pathway."

Circuit Connections
In order to understand the complex circuitry of the basal ganglia,
one has to first understand the important participants in this circuit.
The basal ganglia is in direct communication with the thalamus
and the cortex. Cortex, thalamus, and the basal ganglia are
therefore the three main participants in the circuit created by the
basal ganglia.

In the highest position of authority, and responsible for conscious

perception of the universe, lies the human cerebral cortex. All
actions undertaken by the nervous system directly or indirectly
relate to the cortex. Cortex has many different areas with different
functions. One such cortical area is called the pre-central gyrus,
also known as the "motor cortex." Specialized neurons from the
cortex in the motor cortex region (precentral gyrus) extend their
Connectivity diagram showing excitatory
axons all the way to the striatum portion of the basal ganglia.
glutamatergic pathways as red, inhibitory GABAergic
These cortical neurons release the neurotransmitter glutamate, pathways as blue, and modulatory dopaminergic as
which is excitatory in nature. Once excited by glutamate, the cells magenta.
in the striatum project in two different directions giving rise to two
major pathways: The "direct" and the "indirect" pathway:

In the direct pathway, once activated by the cortex, the cells of striatum, project inhibitory neurons (producing
GABA, an inhibitory neurotransmitter) onto the cells of the "SNr-GPi complex" (SNr and GPi are separate spatially
but due to similar function it is correct to think of them as a complex). SNr and GPi, are constantly in connection
with the thalamus through the ansa lenticularis pathway, producing GABA, in an attempt to inhibit the thalamus.
Due to the inhibition (via the striatal inhibition) of "SNr-GPi" (inhibitors of thalamus themselves), the end result is
lack of inhibition of the thalamus. Thalamus interestingly projects to the cortex itself, constantly stimulating the
cortex via glutamate. The direct pathway therefore results in the thalamus being allowed to stimulate the cortex.
Once stimulated the cortex will then send this message of "stimulation" down its motor pathway via the lateral
corticospinal tract to the muscles, resulting in a hyper-kinetic behavior (meaning increased motion). The following
diagram depicts the "direct" pathway:

Cortex -(stimulates)-> Striatum -(inhibits)-> "SNr-GPi" complex -(inhibits)-> Thalamus -(stimulates)-> Cortex
-(stimulates)-> Muscles, etc. -> (hyperkinetic state)
In the opposite end, is the indirect pathway which also starts from the neurons of the striatum. Once stimulated by
the cortex, a group of the neurons from the striatum, now send inhibitory axons onto the cells of the globus pallidus
externa (GPe). GPe is constantly inhibiting the subthalamic nucleus (STN). This inhibition (by the striatum) of an
inhibitor (GPe), results in the actual stimulation of STN! STN in return is also set to always stimulate the "SNr-GPi
complex", whose main task is to inhibit the thalamus. The end result is an actual inhibition of the thalamus and
therefore decreased stimulation of the cortex by the thalamus. This results in the cortex stimulating the muscles less
through the lateral corticospinal tract and favoring a hypo-kinetic state or decreased motion. In essence, the direct
and the indirect pathway are antagonist in function.
Basal ganglia 109

Cortex -(stimulates)-> Striatum -(inhibits)-> GPe -(inhibits)-> STN -(stimulates)-> "SNr-GPi" complex
-(inhibits)-> thalamus -(is stimulating less)-> Cortex -(is stimulating less)-> Muscles, etc. -> (hypokinetic state)
But if the "direct" pathway favors "motion" and the "indirect" pathway
does not, then there is an inherent contradiction in the human body in
terms of voluntary motion. This paradox is solved due to the function
of substantia nigra pars compacta (SNc) which produces "dopamine."
Special D1-receptors in the basal ganglia are excited by dopamine,
favoring the "direct" pathway, whereas specialized D2-receptors in the
basal ganglia are inhibited in presence of dopamine and actually favor
the "indirect pathway."[4] In this fashion, the body is able to maintain
balance between motion and lack thereof, to create a state of balance.
Lack of balance in this delicate system leads to pathologies such as
Parkinson's disease.

A note concerning terminology

The nomenclature of the basal ganglia system and its components has
always been problematic. Early anatomists, seeing the macroscopic
anatomical structure but knowing nothing of the cellular architecture or Main circuits of the basal ganglia. This diagram
neurochemistry, grouped together components that are now believed to shows 2 coronal slices that have been
have distinct functions (such as the internal and external segments of superimposed to include the involved basal
ganglia structures. + and - signs at the point of the
the globus pallidus), and gave distinct names to components that are
arrows indicate respectively whether the pathway
now thought to be functionally parts of a single structure (such as the is excitatory or inhibitory in effect. Green arrows
caudate nucleus and putamen). refer to excitatory glutamatergic pathways, red
arrows refer to inhibitory GABAergic pathways
The term "basal" comes from the fact that most of its elements are
and turquoise arrows refer to dopaminergic
located in the basal part of the forebrain. The term ganglia is a pathways that are excitatory on the direct
misnomer: in modern usage, neural clusters are only called "ganglia" in pathway and inhibitory on the indirect pathway.
the peripheral nervous system; in the central nervous system they are
called "nuclei". For this reason, the basal ganglia are also occasionally known as the "basal nuclei".[12] Terminologia
anatomica (1998), the international authority for anatomical naming, retained "nuclei basales", but this is not
commonly used.

The International Basal Ganglia Society (IBAGS) informally considers the basal ganglia to be made up of the
striatum, the pallidum (with two nuclei), the substantia nigra (with its two distinct parts) and the subthalamic
nucleus. Percheron et al. in 1991 and Parent and Parent in 2005 included the central region (centre
median-parafascicular) of the thalamus as part of the basal ganglia,[13] [14] while Mena-Segovia et al. in 2004
included the pedunculopontine complex as well.[15]
Also, the names given to the various nuclei of the basal ganglia are different in different species. In particular, the
internal segment of the globus pallidus in primates is called the entopeduncular nucleus in rodents. The "striatum"
and "external segment of the globus pallidus" in primates are called the "paleostriatum augmentatum" and
"paleostriatum primitivum" respectively in birds.
Basal ganglia 110

Information about the functions of the basal ganglia comes from anatomical studies, from physiological studies
carried out mainly in rats and monkeys, and from the study of diseases that damage them.
The greatest source of insight into the functions of the basal ganglia has come from the study of two neurological
disorders, Parkinson's disease and Huntington's disease. For both of these disorders, the nature of the neural damage
is well understood and can be correlated with the resulting symptoms. Parkinson's disease involves major loss of
dopaminergic cells in the substantia nigra; Huntington's disease involves massive loss of medium spiny neurons in
the striatum. The symptoms of the two diseases are virtually opposite: Parkinson's disease is characterized by gradual
loss of the ability to initiate movement, while Huntington's disease is characterized by an inability to prevent parts of
the body from moving unintentionally. It is noteworthy that although both diseases have cognitive symptoms,
especially in their advanced stages, the most salient symptoms relate to the ability to initiate and control movement.
Thus, both are classified primarily as movement disorders. A different movement disorder, called hemiballismus,
may result from damage restricted to the subthalamic nucleus. Hemiballismus is characterized by violent and
uncontrollable flinging movements of the arms and legs.

Eye movements
One of the most intensively studied functions of the basal ganglia (BG) is their role in controlling eye
movements.[16] Eye movement is influenced by an extensive network of brain regions that converge on a midbrain
area called the superior colliculus (SC). The SC is a layered structure whose layers form two-dimensional retinotopic
maps of visual space. A "bump" of neural activity in the deep layers of the SC drives an eye movement directed
toward the corresponding point in space.
The SC receives a strong inhibitory projection from the BG, originating in the substantia nigra pars reticulata
(SNr).[16] Neurons in the SNr usually fire continuously at high rates, but at the onset of an eye movement they
"pause", thereby releasing the SC from inhibition. Eye movements of all types are associated with "pausing" in the
SNr; however, individual SNr neurons may be more strongly associated with some types of movements than others.
Neurons in some parts of the caudate nucleus also show activity related to eye movements. Since the great majority
of caudate cells fire at very low rates, this activity almost always shows up as an increase in firing rate. Thus, eye
movements begin with activation in the caudate nucleus, which inhibits the SNr via the direct GABAergic
projections, which in turn disinhibits the SC.

Role in motivation
Although the role of the basal ganglia in motor control is clear, there are also many indications that it is involved in
the control of behavior in a more fundamental way, at the level of motivation. In Parkinson's disease, the ability to
execute the components of movement is not greatly affected, but motivational factors such as hunger fail to cause
movements to be initiated or switched at the proper times. The immobility of Parkinsonian patients has sometimes
been described as a "paralysis of the will".[17] These patients have occasionally been observed to show a
phenomenon called kinesia paradoxica, in which a person who is otherwise immobile responds to an emergency in a
coordinated and energetic way, then lapses back into immobility once the emergency has passed.
The role in motivation of the "limbic" part of the basal ganglia—the nucleus accumbens (NA), ventral pallidum, and
ventral tegmental area (VTA)—is particularly well established. Thousands of experimental studies combine to
demonstrate that the dopaminergic projection from the VTA to the NA plays a central role in the brain's reward
system. Animals with stimulating electrodes implanted along this pathway will bar-press very energetically if each
press is followed by a brief pulse of electrical current. Numerous things that people find rewarding, including
addictive drugs, good-tasting food, and sex, have been shown to elicit activation of the VTA dopamine system.
Damage to the NA or VTA can produce a state of profound torpor.
Basal ganglia 111

Although it is not universally accepted, some theorists have proposed a distinction between "appetitive" behaviors,
which are initiated by the basal ganglia, and "consummatory" behaviors, which are not. For example, an animal with
severe basal ganglia damage will not move toward food even if it is placed a few inches away, but if the food is
placed directly in the mouth, the animal will chew it and swallow it.

Comparative anatomy and naming

The basal ganglia form one of the basic components of the forebrain, and can be recognized in all species of
vertebrates.[18] Even in the lamprey (generally considered one of the most primitive of vertebrates), striatal, pallidal,
and nigral elements can be identified on the basis of anatomy and histochemistry.[19]
A clear emergent issue in comparative anatomy of the basal ganglia is the development of this system through
phylogeny as a convergent cortically re-entrant loop in conjunction with the development and expansion of the
cortical mantle. There is controversy, however, regarding the extent to which convergent selective processing occurs
versus segregated parallel processing within re-entrant closed loops of the basal ganglia. Regardless, the
transformation of the basal ganglia into a cortically re-entrant system in mammalian evolution occurs through a
re-direction of pallidal (or "paleostriatum primitivum") output from midbrain targets such as the superior colliculus,
as occurs in sauropsid brain, to specific regions of the ventral thalamus and from there back to specified regions of
the cerebral cortex that form a subset of those cortical regions projecting into the striatum. The abrupt rostral
re-direction of the pathway from the internal segment of the globus pallidus into the ventral thalamus—via the path
of the ansa lenticularis--could be viewed as a footprint of this evolutionary transformation of basal ganglia outflow
and targeted influence. The evolutionary emergence of cortical re-entrant systems in the brain has been postulated by
Gerald Edelman as a critical basis for the emergence of primary consciousness in the theory of Neural Darwinism.

In most regions of the brain, the predominant classes of neurons use glutamate as neurotransmitter and have
excitatory effects on their targets. In the basal ganglia, however, the great majority of neurons use GABA as
neurotransmitter and have inhibitory effects on their targets. The inputs from the cortex and thalamus to the striatum
and STN are glutamatergic, but the outputs from the striatum, pallidum, and substantia nigra pars reticulata all use
GABA. Thus, following the initial excitation of the striatum, the internal dynamics of the basal ganglia are
dominated by inhibition and disinhibition.
Other neurotransmitters have important modulatory effects. The most intensively studied is dopamine, which is used
by the projection from the substantia nigra pars compacta to the striatum, and also in the analogous projection from
the ventral tegmental area to the nucleus accumbens. Acetylcholine also plays an important role, being used both by
several external inputs to the striatum, and by a group of striatal interneurons. Although cholinergic cells make up
only a small fraction of the total population, the striatum has one of the highest acetylcholine concentrations of any
brain structure.
Basal ganglia 112

Disorders associated with the basal ganglia

• Attention-deficit hyperactivity disorder (ADHD)
• Athymhormic syndrome (PAP syndrome)
• Athetosis
• Cerebral palsy: basal ganglia damage during second and third trimester of pregnancy
• Chorea
• Dystonia
• Fahr's disease
• Foreign accent syndrome (FAS)
• Huntington's disease
• Lesch-Nyhan syndrome
• Obsessive-compulsive disorder[20] [21]
• Other anxiety disorders [21]
• Parkinson's disease
• Sydenham's chorea
• Tourette's disorder
• Tardive dyskinesia, caused by chronic antipsychotic treatment
• Stuttering[22]
• Spasmodic dysphonia
• Wilson's disease
• Blepharospasm

The acceptance that the basal ganglia system constitutes one major cerebral system took long to arise. The first
anatomical identification of distinct subcortical structures was published by Thomas Willis in 1664.[23] For many
years, the term corpus striatum[24] was used to describe a large group of subcortical elements, some of which were
later discovered to be functionally unrelated.[25] For many years, the putamen and the caudate nucleus were not
associated with each other. Instead, the putamen was associated with the pallidum in what was called the nucleus
lenticularis or nucleus lentiformis.
A thorough reconsideration by Cécile and Oskar Vogt (1941) simplified the description of the basal ganglia by
proposing the term striatum to describe the group of structures consisting of the caudate nucleus, the putamen and
the mass linking them ventrally, the nucleus accumbens. The striatum was named on the basis of the striated
(striped) appearance created by radiating dense bundles of striato-pallido-nigral axons, described by anatomist
Samuel Alexander Kinnier Wilson (1912) as "pencil-like".
The anatomical link of the striatum with its primary targets, the pallidum and the substantia nigra was discovered
later. The name globus pallidus was attributed by Déjerine to Burdach (1822). For this, the Vogts proposed the
simpler "pallidum". The term "locus niger" was introduced by Félix Vicq-d'Azyr as tache noire in (1786), though
that structure has since become known as the substantia nigra, due to contributions by Von Sömmering in 1788. The
structural similarity between the substantia nigra and globus pallidus was noted by Mirto in 1896. Together, the two
are known as the pallidonigral ensemble, which represents the core of the basal ganglia. Altogether, the main
structures of the basal ganglia are linked to each other by the striato-pallido-nigral bundle, which passes through the
pallidum, crosses the internal capsule as the "comb bundle of Edinger", then finally reaches the substantia nigra.
Additional structures that later became associated with the basal ganglia are the "body of Luys" (1865) (nucleus of
Luys on the figure) or subthalamic nucleus, whose lesion was known to produce movement disorders. More recently,
other areas such as the central complex (centre médian-parafascicular) and the pedunculopontine complex have been
Basal ganglia 113

thought to be regulators of the basal ganglia.

Near the beginning of the 20th century, the basal ganglia system was first associated with motor functions, as lesions
of these areas would often result in disordered movement in humans (chorea, athetosis, Parkinson's disease).

[1] http:/ / braininfo. rprc. washington. edu/ Scripts/ hiercentraldirectory. aspx?ID=206
[2] http:/ / www. nlm. nih. gov/ cgi/ mesh/ 2007/ MB_cgi?mode=& term=Basal+ Ganglia
[3] http:/ / www. neurolex. org/ wiki/ birnlex_826
[4] Stocco, Andrea; Lebiere, Christian; Anderson, John R. (2010). "Conditional routing of information to the cortex: A model of the basal
ganglia’s role in cognitive coordination". Psychological Review 117 (2): 541–74. doi:10.1037/a0019077. PMID 20438237.
[5] Weyhenmeyer, James A.; Gallman, Eve. A. (2007). Rapid Review of Neuroscience. Mosby Elsevier. p. 102. ISBN 0-323-02261-8.
[6] Chakravarthy, V. S.; Joseph, Denny; Bapi, Raju S. (2010). "What do the basal ganglia do? A modeling perspective". Biological Cybernetics
103 (3): 237–53. doi:10.1007/s00422-010-0401-y. PMID 20644953.
[7] Cameron IG, Watanabe M, Pari G, Munoz DP. (2010 June). Executive impairment in Parkinson's disease: response automaticity and task
switching (http:/ / www. ncbi. nlm. nih. gov/ pubmed/ 20303998). 48. Neuropsychologia. PMID 20303998. .
[8] Fix, James D. (2008). "Basal Ganglia and the Striatal Motor System". Neuroanatomy (Board Review Series) (4th ed.). Baltimore: Wulters
Kluwer & Lippincott Wiliams & Wilkins. pp. 274–281. ISBN 0-7817-7245-1.
[9] Inta, D.; Meyer-Lindenberg, A.; Gass, P. (2010). "Alterations in Postnatal Neurogenesis And Dopamine Dysregulation in Schizophrenia: A
Hypothesis". Schizophrenia Bulletin. doi:10.1093/schbul/sbq134.
[10] Regina Bailey. "Divisions of the Brain" (http:/ / biology. about. com/ library/ organs/ brain/ blprosenceph. htm). . Retrieved
[11] Kamishina, H; Yurcisin, G; Corwin, J; Reep, R (2008). "Striatal projections from the rat lateral posterior thalamic nucleus". Brain Research
1204: 24–39. doi:10.1016/j.brainres.2008.01.094. PMID 18342841.
[12] Soltanzadeh, Akbar (2004). Neurologic Disorders. Tehran: Jafari. ISBN 964-6088-03-1.
[13] Percheron, G; Filion, M (1991). "Parallel processing in the basal ganglia: up to a point". Trends in Neurosciences 14 (2): 55–9.
doi:10.1016/0166-2236(91)90020-U. PMID 1708537.
[14] Parent, Martin; Parent, Andr� (2005). "Single-axon tracing and three-dimensional reconstruction of centre m�dian-parafascicular thalamic
neurons in primates". The Journal of Comparative Neurology 481 (1): 127–44. doi:10.1002/cne.20348. PMID 15558721.
[15] Menasegovia, J; Bolam, J; Magill, P (2004). "Pedunculopontine nucleus and basal ganglia: distant relatives or part of the same family?".
Trends in Neurosciences 27 (10): 585–8. doi:10.1016/j.tins.2004.07.009. PMID 15374668.
[16] Hikosaka, O; Takikawa, Y; Kawagoe, R (2000). "Role of the basal ganglia in the control of purposive saccadic eye movements.".
Physiological reviews 80 (3): 953–78. PMID 10893428.
[17] Niv, Y.; Rivlin-Etzion, M. (2007). "Parkinson's Disease: Fighting the Will?". Journal of Neuroscience 27 (44): 11777–9.
doi:10.1523/JNEUROSCI.4010-07.2007. PMID 17978012.
[18] Parent A (1986). Comparative Neurobiology of the Basal Ganglia. Wiley. ISBN 9780471803485.
[19] Grillner, S; Ekeberg, O; Elmanira, A; Lansner, A; Parker, D; Tegner, J; Wallen, P (1998). "Intrinsic function of a neuronal network — a
vertebrate central pattern generator1". Brain Research Reviews 26 (2-3): 184–97. doi:10.1016/S0165-0173(98)00002-2. PMID 9651523.
[20] Radua, Joaquim; Mataix-Cols, David (November 2009). "Voxel-wise meta-analysis of grey matter changes in obsessive–compulsive
disorder". British Journal of Psychiatry 195 (5): 393–402. doi:10.1192/bjp.bp.108.055046. PMID 19880927.
[21] Radua, Joaquim; van den Heuvel, Odile A.; Surguladze, Simon; Mataix-Cols, David (5 July 2010). "Meta-analytical comparison of
voxel-based morphometry studies in obsessive-compulsive disorder vs other anxiety disorders". Archives of General Psychiatry 67 (7):
701–711. doi:10.1001/archgenpsychiatry.2010.70. PMID 20603451.
[22] Alm, Per A. (2004). "Stuttering and the basal ganglia circuits: a critical review of possible relations". Journal of communication disorders
37 (4): 325–69. doi:10.1016/j.jcomdis.2004.03.001. PMID 15159193.
[23] Andrew Gilies, A brief history of the basal ganglia (http:/ / www. anc. ed. ac. uk/ ~anaru/ research/ history/ ), retrieved on 27 June 2005
[24] Vieussens (1685)
[25] Percheron, G; Fénelon, G; Leroux-Hugon, V; Fève, A (1994). "History of the basal ganglia system. Slow development of a major cerebral
system". Revue neurologique 150 (8-9): 543–54. PMID 7754290.
Basal ganglia 114

External links
• Basal+ganglia ( at
eMedicine Dictionary
• Imaging of Basal Ganglia (
srchstr=basal ganglia&srch_type=all) at USUHS
• Scholarpedia article on Basal ganglia (
• NIF Search - Basal Ganglia ("Basal Ganglia") via the
Neuroscience Information Framework
• The International Basal Ganglia Society (
Article Sources and Contributors 115

Article Sources and Contributors

PID controller  Source:  Contributors: A. Carty, A3RO, AaronMK, Abarry, Acdx, Ademkader, Aeolian, Alexius08, Alll, Altzinn, Aluvus,
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ego vos et irrumabo, Penubag, Persian Poet Gal, Petergress, Peterlewis, Peterlin, Phil Boswell, Philcohen, Philip Trueman, PhilippWeissenbacher, Physicistjedi, Pinethicket, Pinkadelica,
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RainbowOfLight, Ram-Man, Randy Johnston, Rat at WikiFur, Raul654, Raymondwinn, Rdsmith4, Reaper Eternal, RedHillian, Rehman faiz11, Retired username, RexNL, Rich Farmbrough,
Richard Stephens, Rick MILLER, Rigadoun, Rjwilmsi, Rockoprem, Romain.guises, Rory096, Rotje66, RoyBoy, Rracecarr, Rror, Rtdrury, RyanCross, Sajt, Sampson, Samtheboy, Sanpaz,
Scientizzle, Sdman923, Seanjp8329, Sentriclecub, Sgstarling, Shakejunt420, Shamilton, Shanes, SheikYerBooty, Shellreef, Shoefly, Shoeofdeath, SieG KilleR, Silly rabbit, Sir Lewk, Sir Stig,
Sirex98, Sketchee, Skittleskittle, Smalljim, Smurrayinchester, Snigbrook, Soliloquial, Some fat dude, Spiel496, Spoon!, Squiddy, Srikeit, Starwiz, Stephen J. Brooks, Stephenb, Stevertigo,
Article Sources and Contributors 116

Strikehold, Strongbad1982, StuRat, StuTheSheep, Subversive.sound, SuperColbertFan, Sverdrup, Swamp Ig, Swerdnaneb, Swpb, Syedrazahabbasjilani, Synergy, Syrthiss, T-borg, THEN WHO
WAS PHONE?, Tasc, Tatari-kun, Tawker, Tbhotch, TenPoundHammer, Thatkiwiboy, The Anome, The Firewall, The Thing That Should Not Be, TheDoctor10, TheKMan, Thedjatclubrock,
Thingg, Thu, Tiddly Tom, Tide rolls, Titoxd, Tivedshambo, TomasBat, Tombomp, TotoBaggins, Traveler100, Trevor MacInnis, Tslocum, Tylerni7, Tyrol5, Ukexpat, UniEmpPime, User A1,
VDZ, Vary, Verethor, Viskonsas, VivaEmilyDavies, Voyagerfan5761, Vrenator, Vsmith, Waggers, Waninge, Ward20, WatermelonPotion, Wavelength, Wayne Slam, WaysToEscape, Wayward,
Wazzzup7up, Where, Whiskey in the Jar, Wimt, Wingman358, Winstonio, Wisdom89, Wizard191, Wk muriithi, Wolfkeeper, Wolfmankurd, Wtfily, XB-70, Xakuzzah, Yamamoto Ichiro,
Yelyos, Yettie0711, Youngrod123, Zachplaysguitar5, Zanhsieh, Zaphod Beeblebrox, Zayzya, Zephyr9, Zetawoof, Zgystardst, Zzuuzz, Zzyzx11, ‫ملاع بوبحم‬, 1513 anonymous edits

Muscle  Source:  Contributors: 168..., 2pac 2007, A1z26b2y25, A314268, AS, AZ.spartan118, Achilles.g, Adambiswanger1, Addshore,
Ageekgal, Ahoerstemeier, Aitias, Ajaxkroon, Al.locke, Alai, Aldie, Alex.tan, Alias Flood, Alison, Allstarecho, Alm93, Alowrun, Anders Torlind, Andre Engels, AndrewHowse, Antandrus,
AnthonyQBachler, Antifamilymang, Antiuser, Anxietycello, Apotheosis247, Aramis1250, Arcadian, Arch dude, Ascidian, Askari Mark, Asterion, AstroNomer, Aviturner2, AxelBoldt,
Axeman89, Az1568, BANZ111, BMT, BRileyPTA, BW52, Barzkar, Beland, Bethenco, Bhadani, BiT, Bibliomaniac15, Bike4life15, Bird, Bj,jfkfjnhmgh, Bkkbrad, Blamblamblam, Bobblewik,
Bobianite, Bobo192, Boijunk, Bomac, Bongwarrior, Boomheadshot14, Bovlb, Boxing245, Bratsche, Brianmarx, Brighterorange, Burnedthru, CDN99, CRGreathouse, CWY2190, Caltas, Calvin
trailor, CambridgeBayWeather, Can't sleep, clown will eat me, Candorwien, Captain-n00dle, Captin Shmit, Carlwev,, Cburnett, CerealBabyMilk, Ceyockey, Chalom,
CharlesHoffman, Charley white, Chenfuyu, Chet nc, Chrischung28, Chrisklerkx, Chrislk02, Ciar, Cjwright79, Cleanupman, Clemwang, Clicketyclack, Clovis Sangrail, Cocoa amr, Conversion
script, Cremepuff222, Cutler, Cvagheei, Cyrius, Daarznieks, Dak, Dan Guan, Dan100, Darkfrog24, Dave6, DaveK@BTC, Dcfleck, DeadEyeArrow, Deli nk, Delta G, Demyx9, DennisDaniels,
DennyColt, Dinferno, Discospinster, Dj Capricorn, Dlohcierekim, DocWatson42, Dookaboo02, Dpbsmith, Drektor2oo3, Drex3ech, Dysepsion, Dysprosia, Dzubint, ESkog, Ecolatur, Edgar181,
Edward, Ekton, Electron9, Elert, Ember of Light, EncycloPetey, Epg, Epolk, Equendil, Everyking, Ewlyahoocom, Excirial, Fangusu, Felizdenovo, Fewpasture, Finlay McWalter, Fish and karate,
Fletcherkasmer, Former user, Frazzydee, Frivvy89, Frymaster, Furrykef, Fvasconcellos, Gacggt, Gaius Cornelius, Gamer007, Garfield226, Gciriani, Gene Nygaard, GeorgeStepanek, Ghakko,
Giftlite, Gigemag76, Gilliam, Gimboid13, Giraffedata, Gludwiczak, Gogo Dodo, Gongoozler123, Goochelaar, Googie man, Graham87, GrapeSteinbeck, GregorB, Grstain, Guardian of Light,
Gurch, Gwenmn, Gwernol, Hadseys, Hagerman, Hankwang, Hard Sin, Hdt83, Helenginn, Heron, Hiddenfromview, Honguy86, Hopetocome, Hu, Hu12, Huffers, Iambk, Icemaja, Iceman111111,
Iketsi, Imnotminkus, Instinct, Ixfd64, J. Spencer, J.delanoy, J04n, JDCMAN, JFreeman, Jack Daw, Jag123, Jahiegel, Jake harwood, Jakethewall, Jclemens, Jdlambert, Jeff G., Jeffhoy, Jemijohn,
Jeremykemp, Jerith, Jethero, Jfdwolff, Jhinman, Jiddisch, JinJian, JoanneB, Joehall45, John254, Johntherip, Jonathan Karlsson, Jose77, Jrockley, Jsmith86, Julesd, Jumbo Whales, Junglecat,
Juranas, KJ Sam, Kablammo, Kaisershatner, Kalamkaar, Kaobear, KaragouniS, Katalaveno, Kinhew04, Klenod, Knightshade, Knutux, Kochipoik, Kreachure, Krellis, Krich, Kseferovic,
Kunnskap, Kuru, LAX, Lamchops 25, Latexcat, Lexor, Liftarn, Limegreen, Lir, Lisatwo, LittleHow, LittleOldMe, Lo2u, Lordmetroid, Luk, Luna Santin, MONGO, Mac, MacGyverMagic,
Madhero88, Mahq, Maniwar, MarkN, Massagenj, Mat8989, Mattinbgn, Mayooranathan, Measure, MeekSaffron, MegaSloth, Meol, Miaowmiaowmiaow, Michael Hardy, Michael Rowe, Michael
Shields, Michaelbusch, Michaelkemp, Mightyzero, Mikael Häggström, Mike Dillon, Mild Bill Hiccup, Minisarm, Mintc0ndition, Mintleaf, Mj455972007, Mjpieters, Mokele, Mosmof, Mperkins,
Mporch, Mr Stephen, MrArt, MrBell, Mukkakukaku, Munita Prasad, Mwanner, Mxn, Nagara373, Nate1481, NawlinWiki, NeilN, Neitherday, Neli Georgieva, Nguyen Thanh Quang, Nick.alex1,
Nick123, NickBush24, NickGorton, Nicks221, Nixeagle, Nlu, Nmnogueira, Nono64, Nuttycoconut, ONEder Boy, ONUnicorn, Oddity-, Olivierd, Oxymoron83, P. Gollakota, PFHLai,
PHDrillSergeant, PTSE, Paolo.dL, Patrick, Patxi lurra, Paul Stansifer, Pb30, Pedro, Pentawing, Pgk, Philippe, PierreAbbat, Postglock, Profoss, Pumpie, Quangbao, QueQue81, QuietJohn,
Qwertying, RFerreira, RG2, Radon210, Raetzsch, Rajkiandris, Ranveig, Raul654, Rearete, RedWolf, Reddevil1213, Reedy, Rjwilmsi, RodC, Rodsan18, Romanm, Rory096, Rossenglish,
Rtcpenguin, Rudolf.hellmuth, Ruudse, SGFF, SRMN2, SU Linguist, SWAdair, Saadkamal, Saber girl08, Samuell, Sarranduin, Satori Son, Scarian, Scott MacLean, Seatdistrict, Selket,
Shadow5678910, Shanes, Shemadeit, Sherberton, Shoessss, Sidsel Sørendatter, Silversmith, SimonP, Sjakkalle, Sjhunt, Skittleys, Slakr, Slavik0329, Slicky, SmileToday, Smyth, Snobordingdani,
Snoopydawg, Snowmanradio, Sodium, Sohale, SolomonRR, Someguydude, SpuriousQ, St4mann, StAkAr Karnak, Stephenb, Steve Pucci, Stupid Corn, Suicidal9, Sulc, Super-Magician,
SweetNeo85, Switchercat, T456, TJJFV, TOR, Talking, Tb, Tbush, The Anome, The Rambling Man, TheGrandMaster1, Thedarkestclear, Theresa knott, Thingg, Til Eulenspiegel, Timir2, Tobias
Bergemann, Tony K10, Treesoulja, Trevor MacInnis, Tristanb, Tviscool3, Ufwuct, UltimateDestroyerOfWorlds, Urod, Veghead, Vera.tetrix, Versageek, Virtualken, Vranak, Vsion, Vulc4n,
WJBscribe, WLU, Waldir, Wavelength, Wes Hermann, Where, Wik, WikHead, Wiki alf, Wikidudeman, Wimt, Wine Guy, Wolfkeeper, Xcentaur, Yamamoto Ichiro, Yash Vaidya, Yashgaroth,
YoavD, Yosri, Ziphon, Zzorse, ‫ينام‬, ‫لیقع فشاک‬, 780 anonymous edits

Muscle tone  Source:  Contributors: AED, Aitias, Andres, Arcadian, Barzkar, Beland, Calaka, Can't sleep, clown will eat me, Crotalus
horridus, DeadEyeArrow, Ed Poor, Eddieleung, EncycloPetey, Erik456, Espoo, Eternauta3k, Everyking, Ewlyahoocom, Fama Clamosa, Gornosk, Holme053, Jeetnagda, Jeff G., Julesd, Karada,
Kauczuk, Lab-oratory, Lenoxus, MamaGeek, Midway, Mikael Häggström, NawlinWiki, Neutrality, NickelShoe, One Alternate, Ph.eyes, Phobius, Quartet, Rajah, Rd232, Schnolle, Shawnc,
SineWave, Snowolf, Stevegallery, Taral, The Thing That Should Not Be, Tyciol, WLU, Weregerbil, Wikidudeman, 65 anonymous edits

Muscle memory  Source:  Contributors: Alsopen, Amalas, Amore proprio, Andycjp, ArphaxadHunter, B9 hummingbird hovering, Beland,
BigrTex, Bleedingcherub, Blmpxcvd, Bobblehead, Busybox, CWenger, Can't sleep, clown will eat me, Capedia, Cometstyles, Darkfrog24, Daveswagon, David from Downunder, Digfarenough,
Dominic Hardstaff, Drmadskills, Dwanyewest, Espoo, Frap, Fredmv, Gary King, Graham87, HaeB, Induality, J04n, Jdauie, Jmswisher, Joel7687, John Riemann Soong, JubalHarshaw, Karlhahn,
Kencf0618, Keraunoscopia, Life of Riley, LilHelpa, LiquidEyes, Macslacker, MartinPoulter, Mattisse, Mindeye, Mossmaal, Nighthawkzx, Oliver.nissen, Pearle, PhilKnight, Preslethe, Psy3330
W10, Ptmis350, QueenCake, Raven4x4x, SE7, Shadowmaster13, SpaceFlight89, Tacllews, The monkeyhate, Therealintellectual, Tobias Bergemann, Twocs, Wdfarmer, WittyBeagle, Шизомби,
98 anonymous edits

Muscle atrophy  Source:  Contributors: Abdull, Aiko, Alex Pascual, Arcadian, BureX, Capricorn42, CerealBabyMilk, Clicketyclack,
Cosmo Notch, Dr.rapist69, Ecs-ajb405, Enigma55, Gacggt, GenOrl, Leviel, Lisaharvey5, Mikael Häggström, Mister Goodtimes, Muhandes, N1RK4UDSK714, Neilc, Nkayesmith, Nono64,
Nutriveg, Redvers, Rich Farmbrough, Rustavo, Stemonitis, Wavelength, WikHead, ‫דוד‬55, 42 anonymous edits

Hand strength  Source:  Contributors: Arcadian, Blanchardb, Blehfu, HelloAnnyong, IRushnikov, IceCreamAntisocial, J04n, RDBrown,
Ruudse, Steven Walling, Tide rolls, Woohookitty, 6 anonymous edits

Atrophy  Source:  Contributors: Alex.tan, Alexandrov, Alteripse, Animum, Anna Frodesiak, Arcadian, ArmadilloFromHell, Axl, Ben Ben,
Borgx, Bubba hotep, CDN99, Chillroy, Cmichael, Conty, Danicalove79, Daniel Case, Davidruben, Drgarden, Eras-mus, Erich gasboy, Ethii, Extropian314, FT2, Gacggt, Geogre, Hielor, Historia
Hater, Ignignot, Igoldste, Irdepesca572, Jauhienij, Keitei, Ketiltrout, LIC Habeeb, Loupeter, MER-C, MPerel, Marlowe, Maurreen, Momoricks, My Core Competency is Competency, Neqq,
Noface64, Oxtoby, Person112, Plasticup, Poccil, Polyparadigm, Rettetast, Rjwilmsi, Romero.josh, Rustavo, SCEhardt, Sam Hocevar, Scottalter, Selket, ShaunMacPherson, Siroxo, Spoon From
Atrophy, Stewartadcock, Sunray, Tbone55, The Thing That Should Not Be, WLU, WhatamIdoing, Winnebagan, Xezbeth, Xichael, Youwillbefoiledbyboardnoble, 106 anonymous edits

Rohmert's law  Source:  Contributors: Alvin Seville, CanisRufus, Edgar181, JIP, Michael Hardy, Mokele, No Guru, Oracleofottawa,
RHaworth, Rasinj, Stemonitis, The wub, Thiseye, 7 anonymous edits

Muscular system  Source:  Contributors: 21655, 7, AVand, Abb615, AdamSamuel, Addihockey10, Addshore, Agent Smith (The Matrix),
AgentCDE, Ahoerstemeier, Aitias, Aka042, Alansohn, Alexius08, Alexmoorehead42, Ali'i, Alpha Quadrant (alt), Anjelelsy, AnmaFinotera, Anna Lincoln, Ano-User, Antonio Lopez, Arakunem,
Arakunem (usurped), Arcadian, ArielGold, Arienh4, Arjun01, Armoreno10, Arthena, AtaruMoroboshi, Aua, AubreyEllenShomo, Aude, Avoided, Avs5221, Baa, Banime, Barliman Butterbur,
Beland, Belovedfreak, Bfigura's puppy, Bhadani, Billy joe mcgruffy, Blanchardb, Bob horn, Bobandjerry, Bobianite, Bobo192, Bongwarrior, BrianGV, Brianga, Bunnyhop11, Bücherwürmlein,
CWY2190, CWii, CZmarlin, CapitalR, Capricorn42, Captain panda, CardinalDan, Cat Whisperer, Catgut, Cathardic, Ccwaters, Cenarium, Centurioncode, Ch'marr, Charitwo, Chris tennis6569,
Chromaticity, CinchBug, Closedmouth, Cocoaguy, Cometstyles, Cotvc89, Countcake, Courcelles, D6, DanielDeibler, Darkwind, Darth Panda, Dauden, DennyColt, Deor, DerHexer, Dfrg.msc,
Dhp1080, Diberri, Dirkbb, Dizzydopeweed, Doctormatt, Dontknowwhatyoureexpecting, DoubleBlue, Dougofborg, Doulos Christos, Drahreg01, Dwayne, ERcheck, Echinesedog3, Echinesedog4,
EconoPhysicist, Edgarde, Editan, Eep², Egmontaz, El aprendelenguas, Enviroboy, Epbr123, Erich gasboy, Erkan Yilmaz, Euchiasmus, EverettColdwell, Excirial, Fabiom, Faradayplank,
Favonian, Ferkelparade, Feroxide, Fieldday-sunday, Finngall, Flubeca, Flyaway1111, FrankCostanza, Frehley, GB fan, GDonato, Gaff, Gary King, Gdavidp, Giftlite, Glane23, Gogo Dodo,
Goodnightmush, Gpkrules, Grafen, Grammer & spelling stickler, Gregory Shantz, Grimmygrammy, Gurch, Gyapronzhur, Gökhan, Hac13, Hadal, Hd21, Hello32020, Hersfold tool account,
Hertz1888, Igoldste, Imasleepviking, Immunize, Ixfd64, J. Spencer, J.delanoy, JFreeman, JHunterJ, JaGa, JamieS93, Jayk123x, Jdixon5318, Jeff G., Jfdwolff, Jh51681, Jhenderson777, Jketola,
Jobui19, Joehall45, JohnCD, Joingies, Jojansuckssucks, Jovianeye, Juliancolton, JuneGloom07, Jusdafax, KGasso, Kbdank71, Kehrbykid, Khukri, King of ice, Kingerties, Kingpin13, Kirill
Lokshin, KyraVixen, Lance Firestarter, Lankiveil, Lazylaces, Lespaul1959x, Lights, LittleOldMe, Lova Falk, Luigifan, MER-C, Maddman89, Madhero88, Magister Mathematicae, Malo,
Mannafredo, Marek69, Markjohndaley, MassimoAr, Matt Deres, Maxis ftw, McSly, Meddog, Mentifisto, Mercury, Metju, Michael Devore, Midgey34, Mikael Häggström, Minimac, Mokele,
MonkMan56, Moreschi, Mperkins, Mr Stephen, Myaca, Myanw, Mygerardromance, NERIC-Security, NJGW, NawlinWiki, Newbyguesses, Nivix, Nono64, Nummer29, Nuno Tavares, Nuttah,
Obli, Onorem, Orange Suede Sofa, Orangutan, Ottawa4ever, OverlordQ, Oxymoron83, P. Gollakota, PFHLai, Parutakupiu, Paul August, Pb30, Pedro, Pepper, Persian Poet Gal, Phantomsteve,
Pharaoh of the Wizards, Philip Trueman, Piano non troppo, Pinethicket, Postglock, Prashanthns, Princessinnocence, Pristontale111, Proquence, RadiantRay, RadioFan, Radioactivebloke, Ransid,
Raven in Orbit, Reach Out to the Truth, RedHillian, Redfarmer, RexNL, RobHyman, Rodsan18, Ronosher, Rory096, RoyBoy, Ryanjunk, SMC, ST47, SU Linguist, SahirShah, Sam
Coutu-Oughton, Samuell, Sango123, Sayden, Sbluen, Sdlwatson, Sealor21, Sean Whitton, Seaphoto, Selket, Seraphim, Shadowjams, Shane grindey, Shanes, Shappy, Skater, Skittleys, Slakr,
Smilieman, Snowolf, Soakologist, Someguydude, SpaceFlight89, Spaully, SpuriousQ, Stephenb, Storm Rider, Strom, Stroppolo, Sxc lucy 9, THEN WHO WAS PHONE?, TOR, Takowl,
Tangent747, Teeheeheeheehaha, Telamamas276, Tellyaddict, Template namespace initialisation script, Tempodivalse, The All-Traq, The Anome, The Rambling Man, The Thing That Should Not
Be, TheDJ, TheIguana, Thingg, Think outside the box, Tide rolls, Tommy2010, Trakesht, Traxs7, Tristanb, Truebluepassword, Tylerdeng, Ulric1313, Uncle Dick, Unionhawk, Unschool, User
A1, User27091, Versus22, Vincegoodrum, Vrenator, WLU, WadeSimMiser, Weregerbil, White wolf EN, WhyBeNormal, WikiFew, Wikidudeman, Wikinonymous, William Avery,
Williamma323, Wimt, Wperdue, Wyatt915, Yamamoto Ichiro, Yansa, Yekrats, Zarrinnis, Zijderveld, ‫دمحأ‬.‫يدماغ‬.24, 1184 anonymous edits
Article Sources and Contributors 117

Human musculoskeletal system  Source:  Contributors: Adrian.benko, Alansohn, AnOddName, Anatomicus, Arcadian, Arfgab, Arthena,
Axl, BinaryTed, Blablableep, Boijunk, Brianboulton, Bsadowski1, Bucephalus, Caerwine, CambridgeBayWeather, Camw, Capricorn42, Captain Weirdo the Great, CardinalDan, Caugustineb,
CommonsDelinker, Courcelles, D6, DVD R W, Damicatz, Darrien, DerHexer, Dondevoy01, Dr. ambitious, Drc79, DreamHaze, Ec5618, EoGuy, Eog1916, Ettrig, Faustnh, Favonian, Gaius
Cornelius, Gary King, Gavlyons86, Getwood, Glane23, Golbez, Grander13, Greatgabriel, H.M.S Me, Hetar, Iridescent, J04n, JackSchmidt, Jauhienij, Jfdwolff, JimmyButler, Jmh649, Juliepatt,
Jusdafax, KParks7, KalebOrtega, Kukini, Kumioko, Kupirijo, Kuru, Largoplazo, LindsayH, LizardJr8, Matthew Yeager, Mokele, Nephron, Nono64, NuclearWarfare, Nunh-huh, Opinionjockey,
Pdcook, PeterSymonds, PumbaLV, Pyrrhus16, R, Random user 39849958, RussellMcKenzie, Shameerbabu986, Shantavira, Ssp0062, Stuwy30, Subversive.sound, The Thing That Should Not
Be, The undertow, Time9, Tommy2010, Tyrol5, VMS Mosaic, Venirxy, Voyagerfan5761, WLU, Wassupwestcoast, Wendell, ZamorakO o, ‫לוכי לכ‬, 221 anonymous edits

Muscle spindle  Source:  Contributors: Afristelic, Alexay, Animeronin, Arcadian, Archibald Tuttle, Barzkar, Beeshop, Dave.Dunford,
Diberri, GUllman, Giraffedata, Heron, Jfdwolff, Jon scott1, LittleOldMe, MaestroGenius, Mbarden, Mick c, MindstormsKid, Mporch, Mrwojo, Neuromechanics, Orvilleduck, Rsabbatini, Sayeth,
Sbmehta, Sigma 7, Spagator, Spaully, The Thing That Should Not Be, Trixt, Uxh, Vojtech.dostal, Ziphon, ‫راکزرب میهاربا تسیپارتویزیف‬, ‫ينام‬, 34 anonymous edits

Type Ia sensory fiber  Source:  Contributors: Arcadian, Barzkar, CDN99, GraemeL, Jfdwolff, Mikael Häggström, Rhetth, Rsabbatini,
Thewayforward, Wyvyrn, 15 anonymous edits

Type II sensory fiber  Source:  Contributors: Arcadian, C39, Fram, Jpecino, Mikael Häggström, 7 anonymous edits

Alpha motor neuron  Source:  Contributors: Arcadian, Barzkar, Dantheman531, Delldot, Diberri, Dicklyon, ELLusKa 86, Freekozak,
J.smith, J04n, NifCurator1, Redtigr, ‫ينام‬, 8 anonymous edits

Gamma motoneuron  Source:  Contributors: Arcadian, Barzkar, C39, CanisRufus, Clicketyclack, Delldot, Diberri, Dicklyon,
EncycloPetey, Freekozak, Looie496, Mick c, NifCurator1, Nmg20, SCEhardt, Schmloof, Sgamer1770, Sonic is Cool!!, Timbom3, ‫راکزرب میهاربا تسیپارتویزیف‬, 11 anonymous edits

Beta motor neuron  Source:  Contributors: Barzkar, Tide rolls, ‫راکزرب میهاربا تسیپارتویزیف‬, 1 anonymous edits

Intrafusal muscle fiber  Source:  Contributors: Arcadian, Barzkar, Bearcat, Diberri, EncycloPetey, Gaius Cornelius, InvictaHOG,
Jmjanzen, Jukyuhn, Katiegan, Malcolma, Martinwardle, Miserlou, Murdochious, Poobarb, Some standardized rigour, Spaully, Steinwnj, TexasAndroid, Wdustbuster, 20 anonymous edits

Extrafusal muscle fiber  Source:  Contributors: Arcadian, Barzkar, Billyb, Delldot, Diberri, Dtsang, Ignisdu, Jburman, Lynntyler,
MNewnham, Mikael Häggström, Miserlou, Wdustbuster, 16 anonymous edits

Basal ganglia  Source:  Contributors: (jarbarf), A314268, Adam78, Alansohn, Alex.tan, Amatulic, Anaru, Andre Engels, Arcadian, Attys,
AzadMashari, BjarteSorensen, Brain-mapper, Bryan Derksen, Bubbachuck, CMBJ, Cacycle, CanisRufus, Chinasaur, Choosebrad, Chris Capoccia, Clicketyclack, Commander Keane, D666D,
DabMachine, Dan aka jack, DanielCD, Danierrr, David Buglar, Diberri, Digfarenough, Discospinster, Dr. Persi, Drisar, Drphilharmonic, Ec5618, Eleassar, EoGuy, Everyking, Fnielsen,, Gbin2000, General Tojo, Gerard.percheron, Gfmer, Gilgamesh he, Gleng, Hadal, Hehelol, Hermant patel, Hu12, Huji, Icairns, Jackfirst, Jackhynes, Jaksmith, Janelle91165,
Japanese Searobin, JeremyA, Jfdwolff, Jfurr1981, JohnnyTremolo, JustAGal, JustVisiting, Kelly Martin, Kpmiyapuram, Leevanjackson, Looie496, Ludovich-eng, Makeemlighter, Manco Capac,
MarkS, Mattashner, Mdd, Michal Nebyla, Mikael Häggström, Mmoneypenny, Modify, NifCurator1, Niubrad, Nrets, Nucleophilic, Octavio L, OldakQuill, Pearle, Per Alm, Ph.eyes, Pjf, Qradua,
R.Tempest, R500Mom, Ram-Man, RexNL, Rich Farmbrough, Scottishmatt, Scubert1, Sema nini, Sergeyf1, Shameerbabu986, Silvonen, Someone else, Sonjaaa, Squiggle, Sulbactam, Tarek, The
Anome, Torzsmokus, Tryptofish, Ulflarsen, Ulmo, Uwe Gille, Vaughan, Viral T, William Ortiz, Wisdom89, WriterHound, Xabian40409, Yakuzai, ‫راکزرب میهاربا تسیپارتویزیف‬, 228
anonymous edits
Image Sources, Licenses and Contributors 118

Image Sources, Licenses and Contributors

Image:Pid-feedback-nct-int-correct.png  Source:  License: GNU Free Documentation License  Contributors:
Original uploader was SilverStar at en.wikipedia
Image:Change with Kp.png  Source:  License: Public Domain  Contributors: User:Skorkmaz
Image:Change with Ki.png  Source:  License: Public Domain  Contributors: User:Skorkmaz
Image:Change with Kd.png  Source:  License: unknown  Contributors: User:Skorkmaz
File:Scross helmsman.jpg  Source:  License: Creative Commons Attribution-Sharealike 2.5  Contributors: Original uploader
was Fishdecoy at en.wikipedia
Image:PI controller.png  Source:  License: Public Domain  Contributors: User:Email4mobile
Image:Radial funktion network.svg  Source:  License: Public Domain  Contributors: SebDE
Image:Unnormalized radial basis functions.svg  Source:  License: Public Domain  Contributors:
Image:060803 normalized radial basis functions.png  Source:  License: Public Domain
 Contributors: BenFrantzDale, CommodiCast
Image:060804 3 normalized basis functions.png  Source:  License: Public Domain  Contributors:
Image:060804 4 normalized basis functions.png  Source:  License: Public Domain  Contributors:
Image:060731 logistic map time series 2.png  Source:  License: Public Domain  Contributors:
BenFrantzDale, CommodiCast
Image:060728b unnormalized basis function phi.png  Source:  License: Public Domain
 Contributors: BenFrantzDale, CommodiCast
Image:Normalized basis functions.png  Source:  License: unknown  Contributors: Original uploader was
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