Bistability

For electronics, see Flip-op (electronics) and of semiconductor memory. A bistable device can store
Multivibrator. one bit of binary data, with one state representing a 0
and the other state a 1. It is also used in relaxation os-
cillators, multivibrators, and the Schmitt trigger. Optical
bistability is an attribute of certain optical devices where
two resonant transmissions states are possible and sta-
E ble, dependent on the input. Bistability can also arise in
2 biochemical systems, where it creates digital, switch-like
outputs from the constituent chemical concentrations and
activities. It is often associated with hysteresis in such
systems.
1 3

X
A graph of the potential energy of a bistable system; it has two
local minima. A surface shaped like this with two low points
can act as a bistable system; a ball resting on the surface can only 1 Mathematical modelling
be stable at those two positions. Balls marked 1 and 3 are in
the two stable positions, while ball 2 is at the point of unstable
equilibrium between them. In the mathematical language of dynamic systems analy-
sis, one of the simplest bistable systems is
In a dynamical system, bistability means the system has dy
= y(1 y 2 ).
dt
two stable equilibrium states. Something that is bistable
can be resting in either of two states. These rest states This system describes a ball rolling down a curve with
4 2
need not be symmetric with respect to stored energy. In shape y4 y2 , and has three steady-states: y = 1 ,
terms of potential energy, a bistable system has two local y = 0 , and y = 1 . The middle steady-state y = 0
minima of potential energy separated by a peak (local is unstable, while the other two states are stable. The di-
maximum). An example of a mechanical device which rection of change of y(t) over time depends on the ini-
is bistable is a light switch. The switch lever is designed tial condition y(0) . If the initial condition is positive
to rest in the on or o position, but not between the ( y(0) > 0 ), then the solution y(t) approaches 1 over
two. time, but if the initial condition is negative ( y(0) < 0 ),
then y(t) approaches 1 over time. Thus, the dynamics
In a conservative force eld, bistability stems from the
are bistable. The nal state of the system can be either
fact that the potential energy has three equilibrium points. y = 1 or y = 1 , depending on the initial conditions.
Two of them are minima and one is a maximum. By
For the detail techniques of mathematical modelling of
mathematical arguments, the maximum must lie between bistability, you are encouraged to read an accessible tu-
the two minima. At rest, a particle will be in one of torial by Chong et al. (2015).[1] The tutorial provides a
the minimum equilibrium positions, because that corre- simple example illustration of bistability using a synthetic
sponds to the state of lowest energy. The maximum can toggle switch proposed by Gardner et al. (2000).[2] The
be visualized as a barrier between them. tutorial also use a dynamical system software XPPAUT
[3]
A system can transition from one state of minimal en- to show practically how to see bistability captured by
ergy to the other if it is given enough activation energy a saddle-node bifurcation diagram and the hysteresis be-
to penetrate the barrier (compare activation energy and haviours when the bifurcation parameter is increased or
Arrhenius equation for the chemical case). After the bar- decreased slowly over the tipping points and a protein gets
rier has been reached, the system will relax into the other turned 'On' or turned 'O'.
minimum state in a time called the relaxation time. The appearance of a bistable region can be understood
Bistability is widely used in digital electronics devices to for the model system dy dt = y(r y ) which undergoes
2

store binary data. It is the essential characteristic of the a supercritical pitchfork bifurcation with bifurcation pa-
ip-op, a circuit widely used in latches and some types rameter r .

1
2 2 IN BIOLOGICAL AND CHEMICAL SYSTEMS

2 In biological and chemical sys- saddle-node bifurcation gives rise to a pair of new xed
points emerging, one stable and the other unstable, at a
tems critical value of the parameter. The unstable solution can
then form another saddle-node bifurcation with the initial
stable solution at a higher value of the parameter, leaving

P
only the higher xed solution. Thus, at values of the pa-
rameter between the two critical values, the system has
two stable solutions. An example of a dynamical system
that demonstrates similar features is
45
dx x5
40 dt =r+ 1+x5 x
35 where x is the output, and r is the parameter, acting as
30 the input.[9]
z
25
Bistability can be modied to be more robust and to tol-
O
20

C
erate signicant changes in concentrations of reactants,
15
while still maintaining its switch-like character. Feed-
10
back on both the activator of a system and inhibitor make
5
the system able to tolerate a wide range of concentra-
0 00 tions. An example of this in cell biology is that activated
10 10
20 CDK1 (Cyclin Dependent Kinase 1) activates its activa-
y 30
40
30
20
x tor Cdc25 while at the same time inactivating its inacti-
40
vator, Wee1, thus allowing for progression of a cell into
Three-dimensional invariant measure for cellular-dierentiation mitosis. Without this double feedback, the system would
featuring a two-stable mode. The axes denote cell counts for still be bistable, but would not be able to tolerate such a
[10]
three types of cells: progenitor ( z ), osteoblast ( y ), and chondro- wide range of concentrations.
cyte ( x ). Pro-osteoblast stimulus promotes PO transition.[4]
Bistability has also been described in the embryonic de-
velopment of Drosophila melanogaster (the fruit y). Ex-
Bistability is key for understanding basic phenomena of amples are anterior-posterior [11] and dorso-ventral [12][13]
cellular functioning, such as decision-making processes axis formation and eye development.[14]
in cell cycle progression, cellular dierentiation,[5] and A prime example of bistability in biological systems
apoptosis. It is also involved in loss of cellular home- is that of Sonic hedgehog (Shh), a secreted signaling
ostasis associated with early events in cancer onset and molecule, which plays a critical role in development. Shh
in prion diseases as well as in the origin of new species functions in diverse processes in development, including
(speciation).[6] patterning limb bud tissue dierentiation. The Shh sig-
Bistability can be generated by a positive feedback loop naling network behaves as a bistable switch, allowing the
with an ultrasensitive regulatory step. Positive feedback cell to abruptly switch states at precise Shh concentra-
loops, such as the simple X activates Y and Y activates tions. gli1 and gli2 transcription is activated by Shh, and
X motif, essentially links output signals to their input their gene products act as transcriptional activators for
signals and have been noted to be an important regula- their own expression and for targets downstream of Shh
tory motif in cellular signal transduction because positive signaling.[15] Simultaneously, the Shh signaling network
feedback loops can create switches with an all-or-nothing is controlled by a negative feedback loop wherein the Gli
decision.[7] Studies have shown that numerous biologi- transcription factors activate the enhanced transcription
cal systems, such as Xenopus oocyte maturation,[8] mam- of a repressor (Ptc). This signaling network illustrates the
malian calcium signal transduction, and polarity in bud- simultaneous positive and negative feedback loops whose
ding yeast, incorporate temporal (slow and fast) positive exquisite sensitivity helps create a bistable switch.
feedback loops, or more than one feedback loop that oc- Bistability can only arise in biological and chemical sys-
curs at dierent times.[7] Having two dierent temporal tems if three necessary conditions are fullled: positive
positive feedback loops or dual-time switches allows feedback, a mechanism to lter out small stimuli and a
for (a) increased regulation: two switches that have in- mechanism to prevent increase without bound.[6]
dependent changeable activation and deactivation times;
and (b) linked feedback loops on multiple timescales can Bistable chemical systems have been studied extensively
lter noise.[7] to analyze relaxation kinetics, non-equilibrium ther-
modynamics, stochastic resonance, as well as climate
Bistability can also arise in a biochemical system only for change.[6] In bistable spatially extended systems the onset
a particular range of parameter values, where the param- of local correlations and propagation of traveling waves
eter can often be interpreted as the strength of the feed- have been analyzed.[16][17]
back. In several typical examples, the system has only
one stable xed point at low values of the parameter. A Bistability is often accompanied by hysteresis. On a pop-
 3

ulation level, if many realisations of a bistable system are

considered (e.g. many bistable cells (speciation)[18] ), one
typically observes bimodal distributions. In an ensemble
average over the population, the result may simply look
like a smooth transition, thus showing the value of single-
cell resolution.

3 In mechanical systems
Bistability is the ability of a material to present in two
stable phases that can both exist within a given range of
temperatures but above and below that range only one or
the other phase exists.
Bistability as applied in the design of mechanical systems A ratchet in action. Each tooth in the ratchet together with the
is more commonly said to be over centrethat is, work regions to either side of it constitutes a simple bistable mechanism.
is done on the system to move it just past the peak, at
which point the mechanism goes over centre to its sec-
ondary stable position. The result is a toggle-type action- Interferometric modulator display, a bistable reec-
work applied to the system below a threshold sucient tive display technology found in mirasol displays by
to send it 'over center' results in no change to the mecha- Qualcomm
nisms state.
Springs are a common method of achieving an over cen-
tre action. A spring attached to a simple two position 5 References
ratchet-type mechanism can create a button or plunger
that is clicked or toggled between two mechanical states. [1] Chong et al. (2015) http://www.mssanz.org.au/
Many ballpoint and rollerball retractable pens employ this modsim2015/C2/chong.pdf%5B%5D
type of bistable mechanism.
An even more common example of an over-center device [2] Collins, James J.; Gardner, Timothy S.; Cantor, Charles
R. (2000). Construction of a genetic toggle switch in
is an ordinary electric wall switch. These switches are of-
Escherichia coli. Nature. 403 (6767): 33942. PMID
ten designed to snap rmly into the on or o position 10659857. doi:10.1038/35002131.
once the toggle handle has been moved a certain distance
past the center-point. [3] http://www.math.pitt.edu/~{}bard/xpp/xpp.html%5B%
A ratchet-and-pawl is an elaborationa multi-stable 5D
over center system used to create irreversible motion.
[4] Kryven, I.; Rblitz, S.; Schtte, Ch. (2015). Solution
The pawl goes over center as it is turned in the forward
of the chemical master equation by radial basis functions
direction. In this case, over center refers to the ratchet approximation with interface tracking. BMC systems bi-
being stable and locked in a given position until clicked
ology (9.1).
forward again; it has nothing to do with the ratchet being
unable to turn in the reverse direction. [5] Ghaarizadeh A, Flann NS, Podgorski GJ (2014).
Multistable switches and their role in cellular dieren-
tiation networks. BMC Bioinformatics. 15: S7+. PMC
4 See also 4110729 . PMID 25078021. doi:10.1186/1471-2105-
15-s7-s7.

ferroelectric, ferromagnetic, hysteresis, bistable per- [6] Wilhelm, T (2009). The smallest chemical reaction sys-
ception tem with bistability. BMC Systems Biology. 3: 90. PMC
2749052 . PMID 19737387. doi:10.1186/1752-0509-
astable multivibrator, monostable multivibrator. 3-90.

Schmitt trigger [7] O. Brandman, J. E. Ferrell Jr., R. Li, T. Meyer, Science

310, 496 (2005)
strong Allee eect
[8] Ferrell JE Jr.; Machleder EM (1998). The biochem-
Multistable perception describes the spontaneous or ical basis of an all-or-none cell fate switch in Xenopus
exogenous alternation of dierent percepts in face oocytes.. Science. 280 (5365): 8958. PMID 9572732.
of the same physical stimulus. doi:10.1126/science.280.5365.895.
4 6 EXTERNAL LINKS

[9] Angeli, David; Ferrell, JE; Sontag, Eduardo D (2003). BiStable Reed Sensor
Detection of multistability, bifurcations, and hystere-
sis in a large calss of biological positive-feedback sys-
tems. PNAS. 101 (7): 18227. PMC 357011 . PMID
14766974. doi:10.1073/pnas.0308265100.

[10] Ferrell JE Jr. (2008). Feedback regulation of

opposing enzymes generates robust, all-or-none
bistable responses (PDF). Current Biology. 18 (6):
R244R245. PMC 2832910 . PMID 18364225.
doi:10.1016/j.cub.2008.02.035.

[11] Lopes, Francisco J. P.; Vieira, Fernando M. C.; Holloway,

David M.; Bisch, Paulo M.; Spirov, Alexander V.; Ohler,
Uwe (26 September 2008). Spatial Bistability Generates
hunchback Expression Sharpness in the Drosophila Em-
bryo. PLOS Computational Biology. 4 (9): e1000184.
doi:10.1371/journal.pcbi.1000184.

[12] Wang, Yu-Chiun; Ferguson, Edwin L. (10 March 2005).

Spatial bistability of Dppreceptor interactions during
Drosophila dorsalventral patterning. Nature. 434
(7030): 229234. doi:10.1038/nature03318.

[13] Umulis, D. M.; Mihaela Serpe; Michael B. OConnor;

Hans G. Othmer (1 August 2006). Robust, bistable
patterning of the dorsal surface of the Drosophila em-
bryo. Proceedings of the National Academy of Sciences.
103 (31): 1161311618. PMC 1544218 . PMID
16864795. doi:10.1073/pnas.0510398103.

[14] Graham, T. G. W.; Tabei, S. M. A.; Dinner, A. R.; Rebay,

I. (22 June 2010). Modeling bistable cell-fate choices
in the Drosophila eye: qualitative and quantitative per-
spectives. Development. 137 (14): 22652278. PMC
2889600 . PMID 20570936. doi:10.1242/dev.044826.

[15] Lai, K., M.J. Robertson, and D.V. Schaer, The sonic
hedgehog signaling system as a bistable genetic switch.
Biophys J, 2004. 86(5): p. 2748-57.

[16] Elf, J.; Ehrenberg, M. (2004). Spontaneous separation

of bi-stable biochemical systems into spatial domains of
opposite phases. Systems Biology (Stevenage). 1 (2):
230236. PMID 17051695. doi:10.1049/sb:20045021.

[17] Kochanczyk, M.; Jaruszewicz, J.; Lipniacki, T. (Jul

2013). Stochastic transitions in a bistable reaction sys-
tem on the membrane. Journal of the Royal Society In-
terface. 10 (84): 20130151. PMC 3673150 . PMID
23635492. doi:10.1098/rsif.2013.0151.

[18] Nielsen; Dolganov, Nadia A.; Rasmussen, Thomas; Otto,

Glen; Miller, Michael C.; Felt, Stephen A.; Torreilles,
Stphanie; Schoolnik, Gary K.; et al. (2010). Isberg,
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6 External links
http://www.answers.com/topic/optical-bistability
 5

7 Text and image sources, contributors, and licenses

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Bistability Source: https://en.wikipedia.org/wiki/Bistability?oldid=776940515 Contributors: Andre Engels, Heron, Edward, Glenn, GRA-
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