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A Glossary for Systems Biology
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CONTROL
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ADAPTATION
robustness , integral control, differentiation, system (taxonomy)
- The response to an extracellular stimulus returns to its pre-stimulus
value even in the continued presence of the signal. [63]
- Adaptation to changes in environmental conditions is the
driving force behind the evolutionary development
of all species from a common predecessor. [37]
- The ability of a system to compensate for changes in its environment.
[6,5]
- Adaptive control,
where controller parameters are constantly adjusted to accommodate
changes in process dynamics, process parameters or disturbances. [8]
- After reacting to a persisting stimulus, the output of the systems
returns to its pre-stimulus value.
In systems theory there are a number of ways to interpret adaptation.
The first two sound very similar: to compensate for changes. The difference
is in where the changes take place and what has to adapt to them.
In the first case the changes are external to the system, and the
systems adapts to them. In the second case they are internal to the
system, and a control scheme has to adapt to them in order to be able
to continue controlling the system.
In both cases the commencing change itself activates a mechanism that
opposes the external disturbance. This is done to ensure that the
essential variables of the system stay within acceptable (i.e. stable/non-dangerous/uncritical)
limits. These limits are much narrower than they would be if the disturbance
were unopposed. The narrowing is the objective manifestation of the
mechanism's adaptation [6].
The third possibility concerns the reaction to a stimulus that lasts
longer than a certain interval, out-lasting the 'sensitivity period'
of the system. This interpretation coincides with one of the meanings
of adaptation in biology.
``A hallmark of many biological sensing devices
is the ability to adapt to a persistent input stimulus, thereby increasing
the range of sensitivity.'' [63] To counter such persistent
stimuli, adaptation
can be implemented through integral feedback control
(integral control), in biological
as well as in technical systems. The term used for adaptation
in technical systems is 'asymptotic tracking'
[63]. In biology, this form of adaptation
is also called desensitization [63].
It is argued that integral control is not only sufficient but also
necessary for robust perfect adaptation
[12].
The second biological interpretation puts adaptation
in an evolutionary context. Adaptation to changing environmental
conditions - also called adaptive radiation
or acclimatization - resulted in
the development of new species which were adapted to those conditions,
thus being able to withstand selection. Successful species incorporated
the adapted characteristics into their genetic code
to pass it on to their offspring [37].
Regardless of context, adaptation is one possibility to make
a system robust against changes in its environment. (robustness)
- bacterial chemotaxis: changes in concentration
of a substrate in the surrounding media influence propulsion activity
of the cell, allowing it to move to the location of the highest (attractor)/lowest
(repellent) concentration [39,63]; see also Chapter 'Recommended Readings'.
- eye: The retina works best at a certain intensity of illumination.
In bright light the nervous system contracts the pupil, and in dim
relaxes it. Thus the amount of light entering the eye is maintained
within limits. [6,5]
- all homeostasis systems: ``For example, within the cell, the levels
of important second messenger molecules such as calcium and of key
metabolites such as ATP fluctuate dramatically in response to both
internal and external events. Integral control operating through the
enzymes that create or remove these molecules can provide a robust
mechanism for restoring the concentrations of these species to their
optimal steady-state level.''
[15]
- adaptive control (see Explanation), e.g. course control for a rocket
which burns off fuel, thus changing its mass [20].
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