Membrane fluidity

In biology, membrane fluidity refers to the viscosity of made artificially. In such cases, one can still speak of
the lipid bilayer of a cell membrane or a synthetic lipid membrane fluidity. These membranes are supported by
membrane. Lipid packing can influence the fluidity of a flat surface, e.g. the bottom of a box. The fluidity of
the membrane. Viscosity of the membrane can affect the these membranes can be controlled by the lateral pressure
rotation and diffusion of proteins and other bio-molecules applied, e.g. by the side walls of a box.
within the membrane, there-by affecting the functions of
these molecules.[1]
2 Heterogeneity in membrane
physical property
1 Factors determining membrane
fluidity Discrete lipid domains with differing composition, and
thus membrane fluidity, can coexist in model lipid mem-
Membrane fluidity can be affected by a number of branes; this can be observed using fluorescence mi-
factors.[1] One way to increase membrane fluidity is to croscopy.[2] The biological analogue, 'lipid raft', is hy-
heat up the membrane. Lipids acquire thermal energy pothesized to exist in cell membranes and perform bi-
when they are heated up; energetic lipids move around ological functions.[3] Also, a narrow annular lipid shell
more, arranging and rearranging randomly, making the of membrane lipids in contact with integral membrane
membrane more fluid. At low temperatures, the lipids proteins have low fluidity compared to bulk lipids in
are laterally ordered and organized in the membrane, and biological membranes, as these lipid molecules stay stuck
the lipid chains are mostly in the all-trans configuration to surface of the protein macromolecules.
and pack well together. The composition of a membrane
can also affect its fluidity. The membrane phospholipids
incorporate fatty acids of varying length and saturation. 3 Measurement methods
Lipids with shorter chains are less stiff and less viscous
because they are more susceptible to changes in kinetic
energy due to their smaller molecular size and they have Membrane fluidity can be measured with electron spin
resonance (ESR), fluorescence, or deuterium nuclear
less surface area to undergo stabilizing van der Waals in-
teractions with neighboring hydrophobic chains. Lipid magnetic resonance spectroscopy (NMR). ESR mea-
chains with double bonds are more fluid than lipids that surements involve observing spin probe behaviour in
are saturated with hydrogen and thus have only single the membrane. Fluorescence experiments involve ob-
bonds. On the molecular level, unsaturated double bonds serving fluorescent probes incorporated into the mem-
make it harder for the lipids to pack together by putting brane. Solid state deuterium nuclear magnetic resonance
kinks into the otherwise straightened hydrocarbon chain. spectroscopy involves observing deuterated lipids.[1] The
Membranes made with such lipids have lower melting techniques are complementary in that they operate on dif-
points: less thermal energy is required to achieve the same ferent timescales.
level of fluidity as membranes made with lipids with satu- Membrane fluidity can be described by two different
rated chains.[1] Incorporation of particular lipids, such as types of motion: rotational and lateral. In ESR, rotational
sphingomyelin, into synthetic lipid membranes is known correlation time of spin probes is used to characterize
to stiffen a membrane. Such membranes can be de- how much restriction is imposed on the probe by the
scribed as “a glass state, i.e., rigid but without crystalline membrane. In fluorescence, steady-state anisotropy of
order”.[2] Cholesterol acts as a bidirectional regulator of the probe can be used, in addition to the rotation correla-
membrane fluidity because at high temperatures, it stabi- tion time of the fluorescent probe.[1] Fluorescent probes
lizes the membrane and raises its melting point, whereas show varying degree of preference for being in an envi-
at low temperatures it intercalates between the phospho- ronment of restricted motion. In heterogeneous mem-
lipids and prevents them from clustering together and branes, some probes will only be found in regions of
stiffening. Some drugs, e.g. Losartan, are also known higher membrane fluidity, while others are only found in
to alter membrane viscosity.[2] Another way to change regions of lower membrane fluidity.[4] Partitioning pref-
membrane fluidity is to change the pressure.[1] In the lab- erence of probes can also be a gauge of membrane flu-
oratory, supported lipid bilayers and monolayers can be idity. In deuterium NMR, the average carbon-deuterium

1
2 8 REFERENCES

bond orientation of the deuterated lipid gives rise to spe- 4.1 Diffusion coefficients
cific spectroscopic features. All three of techniques can
give some measure of the time-averaged orientation of Diffusion coefficients of fluorescent lipid analogues are
the relevant (probe) molecule, which is indicative of the about 10−8 cm2 /s in fluid lipid membranes. In gel lipid
rotational dynamics of the molecule.[1] membranes and natural biomembranes, the diffusion co-
−11 2 −9 2 [1]
Lateral motion of molecules within the membrane can efficients are about 10 cm /s to 10 cm /s.
be measured by a number of fluorescence techniques:
fluorescence recovery after photobleaching (FRAP) in-
volves photobleaching a uniformly labelled membrane 5 Charged lipid membranes
with an intense laser beam and measuring how long it
takes for fluorescent probes to diffuse back into the pho-
The melting of charged lipid membranes, such as 1,2-
tobleached spot.[1] Fluorescence correlation spectroscopy
dimyristoyl-sn-glycero-3-phosphoglycerol (DMPG), can
(FCS) monitors the fluctuations in fluorescence intensity
take place over a wide range of temperature. Within this
measured from a small number of probes in a small space.
range of temperatures, these membranes become very
These fluctuations are affected by the mode of lateral
viscous.[2]
diffusion of the probe. Single particle tracking involves
following the trajectory of fluorescent molecules or gold
particles attached to a biomolecule and applying statisti-
cal analysis to extract information about the lateral diffu- 6 Biological relevance
sion of the tracked particle.[5]
Microorganisms subjected to thermal stress are known to
alter the lipid composition of their cell membrane (see
homeoviscous adaptation). This is one way they can ad-
just the fluidity of their membrane in response to their
environment.[1] Membrane fluidity is known to affect the
4 Phospholipid-deficient bio- function of biomolecules residing within or associated
membranes with the membrane structure. For example,the binding
of some peripheral proteins is dependent on membrane
fluidity.[7] Lateral diffusion (within the membrane ma-
A study of central linewidths of electron spin resonance trix) of membrane-related enzymes can affect reaction
spectra of thylakoid membranes and aqueous disper- rates.[1] Consequently, membrane-dependent functions,
sions of their total extracted lipids, labeled with stearic such as phagocytosis and cell signalling, can be regulated
acid spin label (SASL) (having spin or doxyl moiety at by the fluidity of the cell-membrane.[8]
5,7,9,12,13,14 and 16th carbons, with reference to car-
bonyl group), reveals a fluidity gradient. Decreasing
linewidth from 5th to 16th carbons represents increas-
ing degree of motional freedom (fluidity gradient) from 7 See also
headgroup-side to methyl terminal in both native mem-
branes and their aqueous lipid extract (a multilamellar li- • Lipid bilayer phase behavior
posomal structure, typical of lipid bilayer organization).
This pattern points at similarity of lipid bilayer organi- • Saffman–Delbrück model
zation in both native membranes and liposomes. This
observation is critical, as thylakoid membranes compris- • Homeoviscous adaptation
ing largely galactolipids, contain only 10% phospholipid,
unlike other biological membranes consisting largely of • Lipid bilayer
phospholipids. Proteins in chloroplast thylakoid mem-
branes, apparently, restrict lipid fatty acyl chain segmen- • Liposome
tal mobility from 9th to 16th carbons vis a vis their li-
• Annular lipid shell
posomal conterparts. Surprisingly, liposomal fatty acyl
chains are more restricted at 5th and 7th carbon posi-
tions as compared at these positions in thylakoid mem-
branes. This is explainable as due to motional restricting 8 References
effect at these positions, because of steric hindrance by
large chlorophyll headgroups, specially so, in liposomes. [1] Gennis, R. B. (1989) Biomembranes: Molecular Structure
However, in native thylakoid membranes, chlorophylls and Function. Springer, ISBN 0387967605.
are mainly complexed with proteins as light-harvesting
complexes and may not largely be free to restrain lipid [2] Heimburg, T. (2007) Thermal Biophysics of Membranes.
fluidity, as such.[6] Wiley-VCH, ISBN 3527404716.
 3

[3] Simons K, Vaz WL (2004). “Model systems, lipid

rafts, and cell membranes”. Annual Review of Bio-
physics and Biomolecular Structure. 33: 269–95.
doi:10.1146/annurev.biophys.32.110601.141803. PMID
15139814.

[4] Baumgart, Tobias; Hunt, Geoff; Farkas, Elaine R.;

Webb, Watt W.; Feigenson, Gerald W. (2007).
“Fluorescence probe partitioning between Lₒ/L
phases in lipid membranes”. Biochimica et Biophysica
Acta (BBA) - Biomembranes. 1768 (9): 2182–94.
doi:10.1016/j.bbamem.2007.05.012. PMC 2702987 .
PMID 17588529.

[5] Almeida, P. and Vaz, W. (1995). “Lateral diffusion in

membranes”, Ch. 6, pp. 305–357 in: Lipowsky, R. and
Sackmann, E. (eds.) Handbook of biological physics. El-
sevier Science B.V. doi:10.1016/S1383-8121(06)80023-
0, ISBN 978-0-444-81975-8

[6] YashRoy R C (1990) Magnetic resonance studies

of dynamic organisation of lipids in chloroplast
membranes. Journal of Biosciences, vol. 15(4), pp.
281-288.http://www.researchgate.net/publication/
225688482_Magnetic_resonance_studies_of_dynamic_
organisation_of_lipids_in_chloroplast_membranes?ev=
prf_pub

[7] Heimburg, Thomas & Marsh, Derek (1996). “Thermo-

dynamics of the Interaction of Proteins with Lipid Mem-
branes”. In Kenneth M. Merz Jr. & Benoît Roux. Bio-
logical Membranes. Boston: Birkhäuser. pp. 405–462.
doi:10.1007/978-1-4684-8580-6_13. ISBN 978-1-4684-
8580-6.

[8] Helmreich EJ (2003). “Environmental influences on sig-

nal transduction through membranes: A retrospective
mini-review”. Biophysical chemistry. 100 (1–3): 519–34.
doi:10.1016/S0301-4622(02)00303-4. PMID 12646388.
4 9 TEXT AND IMAGE SOURCES, CONTRIBUTORS, AND LICENSES

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