Chapter 7 MEMBRANE STRUCTURE AND FUNCTION

Summary of Chapter 7, BIOLOGY, 9

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ED Campbell, by J.B. ee!e et al. "#$$.
The plasma membrane separates the cell (living) from the outside world (non-living) and defines
the cell as a distinct entity.
The plasma membrane is selectively permeable; it shows selective permeability.
It allows some substances to pass through and prevents others from passing through.
The main functions of the plasma membranes.
. The plasma membrane helps maintain a life-supporting internal environment by
regulating the passage of materials in and out of the cell.
!. "lasma membranes receive information that permits the cells to sense changes in the
environment and respond to them.
#. Communication between cells ta$es place through the plasma membrane.
%. &iochemical reactions occur on their surface.
MEMBRANE STRUCTURE
&iological membranes are made of lipids and proteins forming a double layer or bilayer. There
are also carbohydrates.
The most common lipids are phospholipids.
"hospholipids are amphipathic molecules; they have a hydrophobic and a hydrophilic
region.
'embrane proteins also have hydrophobic and hydrophilic regions.
istory
()* - Charles +verton hypothesi,ed that membranes were made of lipids.
)7 - Irving -angmuir made artificial membranes by adding phospholipids dissolved in
ben,ene.
)!* - .. /orter and 0. /rendel proposed that membranes were a phospholipid bilayer
two molecules thic$.
)#* - 1ugh 2avson and 3ames 2anielli proposed a sandwich model4 a phospholipid
bilayer between two layers of globular proteins.
In )*56s - electron microscopy seem to support the 2avson-2anielli model and by...
&y )756s - the 2avson-2anielli had become widely accepted as the structure of the
plasma membrane.
&y the end of )756s - 8uestions about all cell membranes being identical began to arise.
2ifferences in thic$ness and appearance between plasma and mitochondrial
membranes.
2ifferences in phospholipid and protein content and type.
"osition of proteins and their hydrophobic parts.
)7! - 9. 3. 9inger and /. :icolson proposed that the proteins are embedded in the
bilayer with only the hydrophilic region e;posed to the li8uid medium. This is called the !l"id
mosaic model.
Membra#es are !l"id
'embrane proteins are mostly mobile and do not remain static in one location.
$hospholipids move laterally in a membrane.
They may flip-flop from the outer layer to the inner layer or vice-versa. This is a rare event.
<nsaturated hydrocarbon tails have $in$s that prevent phospholips from pac$ing together.
This helps the fluidity of the membrane.
$rotei#s are much larger than phospholipids and some drift along the membrane.
The movement of some proteins is very directed probably due to connections to the
cytos$eleton.
'any proteins are= however= anchored to the cytos$eleton and remain fi;ed.
Cholesterol molecules are part of the plasma membrane.
>t high temperatures= #7?C in humans= it interferes with phospholipid drifting and membrane
fluidity.
>t low temperatures interferes with the pac$ing of phospholipids and the free,ing of the
membrane. It lowers the solidification point of the membrane.
Changes in the fluidity of the membrane will affect its function.
@ead4 http4AAwww.mmeade.comAcheatAcholesterol.html
0ishes that live in e;treme cold water have membranes with a high proportion of unsaturated
hydrocarbon tails= enabling their membranes to remain fluid.
9ome bacteria and >rchaea live in very hot water= up to )5BC; their membranes contains
unusual lipids that prevent e;cessive fluidity.
%&embra'e l(p()* of +r!haea are u'(,ue a') )(*t('!t from tho*e fou') (' Eu-arya a') Ba!ter(a. The polar l(p()* !o'*(*t of
(*opre'o() !ha('*, "#./# !arbo'* (' le'0th a') u*ually *aturate), 1h(!h are atta!he) 2(a *table ether bo')* to the 0ly!erol !arbo'*
at the *'3",4 po*(t(o'*. 5olar hea) 0roup* )(ffer at the 0e'u* le2el of )(2er*(ty a') !o'*(*t of m(6ture* of 0ly!o 0roup* 7ma('ly
)(*a!!har()e*8, a')9or pho*pho 0roup* pr(mar(ly pho*pho0ly!erol, pho*pho*er('e, pho*phoetha'olam('e or pho*pho('o*(tol:;
Archaeal Membra#e %ipids a#d Applicatio#s& /. 2ennis 9prott= Institute for &iological 9ciences= +ttawa= +:= Canada
http4AAwww.els.netACileyC2>A.ls>rticleArefId-a5555#(*.html
See f(0* 7.", 4,/ a') < o' pa0e* $"= a') $"7.
MEMBRANE $ROTEINS
"roteins are embedded in a matri; of phospholipids.
The proteins determine most of the specific functions of the membranes.
I:T./@>- "@+T.I:9 are embedded in the bilayer with the hydrophilic side e;posed to the
a8ueous environment and the hydrophobic side inside the bilayer.
T@>:9'.'&@>:. "@+T.I:9 are integral proteins that pass through the bilayer from side to
side.
Transmembrane proteins may function as channels or pores through which small ions and
water-soluble molecules can pass; others act as carriers that bind to a substance and move it
through the membrane.
".@I"1.@>- "@+T.I:9 are not embedded in the bilayer but are usually bonded to integral
proteins by non-covalent interactions.
The outer and inner sides of the plasma membrane have different arrangement of proteins.
"roteins are arranged asymmetrically and each side of the membrane has different
characteristics.
+n the cytoplasmic side of the plasma membrane= some proteins are held in place by the
cytos$eleton.
+n the e;terior side of the plasma membrane= some proteins are attached to fibers of the
e;tracellular matri;.
The two layers of the membrane may differ in phospholipid composition and the proteins have a
directional orientation in the membrane.
The membrane also has carbohydrates= which are restricted to the e;terior surface. 9ome
carbohydrates are attached to membrane proteins.
The asymmetrical distribution of phospholipids= proteins and carbohydrates is determined as the
membrane is being built by the endoplasmic reticulum.
F"#ctio# o! the membra#e protei#s
. Transport across the membrane.
!. .n,ymatic activity.
#. 9ignal transduction.
%. Intercellular Doining.
*. Cell-cell recognition.
7. >ttachment to the cytos$eleton and .C'
See *ummary o' pa0e $"9, f(0.7.$#.
Role o! carbohydrates i# cell'to'cell reco(#itio#
Cells have the ability to distinguish one type of neighboring cell from another.
This is important during embryonic development when the sorting out of cells into tissues occur.
It is the basis for the reDection of foreign cells by the immune system.
Carbohydrates are important in cell-cell recognition.
'embrane carbohydrates are usually short-chained saccharides called oli(osaccharides=
consisting of less than * sugar units.
'embrane carbohydrates on the e;ternal side of the plasma membrane vary from species to
species= among individuals of the same species and from one cell type to another in the same
individual.
9ome are covalently bonded to phospholipids forming (lycolipids; most are bonded to proteins
forming (lycoprotei#s.
'embrane proteins and lipids are synthesi,ed in the .@. Carbohydrates are added here to
ma$e glycoproteins and glycolipids. The carbohydrate portion may then be modified.
The glycoproteins are transported from the .@ to the /olgi comple; in small vesicles and
released in the cis portion of the /olgi facing the same way as in the .@4 the side that was
facing the lumen in the .@ also faces the lumen in the /olgi apparatus.
.n,ymes in the /olgi comple; lumen further modify the carbohydrate branch of the
glycoproteins and incorporate it into secretory vesicles.
The secretory vesicles move to the plasma membrane and fuse with it e;posing the
carbohydrate chain of the glycoprotein to the outside of the cell.
'embrane proteins are involved in the transport of materials in and out of the cell= act as
en,ymes= receive stimuli and transmit information= function in cell recognition and lin$ cells
together.
+ligosaccharides on the e;ternal side of the membrane vary from species to species= individual
to individual and from cell to cell. They act as mar)ers than distinguish one cell from another.
In si(#al tra#sd"ctio#= a receptor protein converts an e;tracellular signal into an intracellular
signal that causes some change in the cell using a series of molecules that relay information
from one to another.
TRAFFIC ACROSS TE MEMBRANE
Cells are bathed in and e;tracellular fluid called the i#terstitial !l"id that is derived from the
blood.
This fluid contains thousands of ingredients li$e amino acids= carbohydrates= fatty acids=
vitamins= hormones= neurotransmitters= salts= and waste products.
The cell must e;tract from this soup the e;act amounts of the substances it needs at specific
time and reDect the rest.
The membrane has the ability to regulate traffic across its phospholipid bilayer.
Selective permeability
9electively permeable membranes allow the passage of some substances and prevent others
from passing through.
&iological membranes are usually permeable to small molecules and nonpolar lipid-soluble
substance.
Cater= gases (+!= :!= C+!= C+)= small polar molecules (glycerol)= larger non-polar
molecules (hydrophobic substances li$e hydrocarbons and some fats)= alcohol.
&iological membranes are impermeable to and use proteins to transport the following types of
molecules=
Ions= water= amino acids and sugars= e. g. glucose.
The hydrophobic center of the bilayer prevents the direct passage of ions and polar molecules=
which are hydrophilic= through the membrane.
TRANS$ORT $ROTEINS
1ydrophilic substances enter the cell through transport proteins thus avoiding contact with the
hydrophobic core of the membrane.
These transport proteins span the membrane.
. Cha##el protei#s are li$e tunnels used by ions and polar molecules to enter the cell=
e.g. a8uaporins.
!. Carrier protei#s change shape in a way that pushes the hydrophilic substance to the
other side of the membrane.
&oth transport proteins are specific for the substance that is to be transported.
$ASSI*E TRANS$ORT
"assive transport is the diffusion of a substance through a biological membrane and it does not
re8uire energy.
Di!!"sio# is the movement of molecules that causes them to spread to occupy evenly the entire
available space.
It depends on the concentration gradient. 9ubstances will diffuse from the area of high
concentration to the area of lower concentration.
The concentration gradient represents the potential energy.
2iffusion is a spontaneous process that does not re8uire energy.
TRANS$ORT MECANISMS
2iffusion and active transport re8uire energy.
Co#ce#tratio# (radie#t provides the energy for diffusion= also called passive
tra#sport.
>T" provides the energy for active transport.
+& Simple di!!"sio#
2iffusion is movement of molecules down the co#ce#tratio# (radie#t from the area of high
concentration to the area of low concentration using the $inetic energy of the molecules. This
results in even scattering of molecules throughout the environment.
The greater the difference in concentration between two areas of the environment= the faster
diffusion will occur.
Dialysis is the diffusion of a substance through a membrane.
Osmosis is the diffusion of water= a solvent= through a membrane from the region of high
water concentration to the region of low water concentration.
Osmotic press"re of a solution is the tendency of water to move from the area of high
concentration to the area of low concentration.
'uch of the traffic across the membrane occurs by diffusion. This is called passive tra#sport.
The concentration gradient represents potential energy and drives diffusion.
2iffusion is unaffected by the presence of other substances.
> solution with high solute concentration (e.g. salt) has in effect low water concentration and
high osmotic pressure.
Isoto#ic sol"tio#s have the same osmotic pressure. ..g. the cell has the same solute
concentration as its environment.
yperto#ic sol"tio#s have higher solute concentration than the other solution. ..g. the
environment has greater concentration than the cell= so the environment is hypertonic to the
cell. The cell loses water and becomes plasmoly,ed (plasmolysis).
ypoto#ic sol"tio#s have lower solute concentration than the other solution. ..g. the
environment has lower concentration than the cell= so the environment is hypotonic to the cell.
The cell swells.
T"r(or press"re is the pressure caused by the cell of plants against the cell wall when the cell
swells with water.
See f(0. 7.$/.
Cells lac$ing walls are isotonic to their environment or have adaptations for osmore("latio#=
the control of water balance.
In a hypertonic solution= these cells lose water= shrivel and die. They become plasmoly,ed.
In a hypotonic solution= these cells will ta$e in water= swell and eventually burst.
Cells with walls are those of plants= fungi and some protists.
In a hypertonic solution= these cells will also become plasmoly,ed.
In a hypotonic solution= the cells will swell until they begin to e;ert pressure against the cell
wall. The cell wall will e;ert a bac$pressure that will counteract the pressure of the swollen
cell. >t this point= the volume will not increase any more and water will leave the cell in the
same amount as it enters. The cell is t"r(id (very firm) and has reached a point of
e8uilibrium.
,& Carrier'mediated tra#sport.
9peciali,ed integral membrane proteins move ions or molecules across the membrane.
A& Facilitated di!!"sio#.
<ses the concentration gradient as the energy source. Concentration gradient must be
maintained.
9ome integral proteins provide a corridor through which ions and molecules can pass
through the membrane following the concentration gradient.
'olecule binds to integral protein. It cannot wor$ against the gradient.
Transmembrane protein changes shape and open a channel through the membrane.
9hape change allows the release of the molecule into the cytoplasm4 (ated cha##els&
Transmembrane protein reverts to its original shape when the molecule is released.
> stimulus (electrical= chemical) is necessary for the gated channels to open.
See f(0. 7.$7.
B& Carrier'mediated active tra#sport.
The cell spends energy from >T" to move ions or molecules across the membrane
against the concentration gradient. This is called active tra#sport&
Ions bind to the transmembrane protein= the pump.
"hosphate group is transferred from >T" to the transport protein.
Transport protein undergoes a conformational change and ions are released to the other
side of the membrane.
See f(0. 7.$>.
Membra#e pote#tial
>ll cells have voltages across the membrane.
Eoltage is electrical potential energy due to the separation of charges= positive from
negative.
The cytoplasm of the cell tends to be negative due to an une8ual distribution of charges
on both sides of the membrane.
The voltage across the membrane is called the membra#e pote#tial.
The membrane potential is measured in volts. The potential across cell membranes
ranges fro -*5 to -!55 volts.
This membrane potential favors the passage of cations to the inside of the cell and of
anions out of the cell.
>n electrochemical (radie#t is created when ions are stored against the concentration
gradient.
Two forces drive the diffusion of ions across a membrane4 the ionFs concentration
gradient and the electrical force that attracts to the side of the membrane with the
opposite charge.
Ions diffuse down its electrochemical gradient.
Lear' the e6ample of the *o)(um3pota**(um pump o' pa0e $4=.
> transport protein that generates a membrane potential is called an electro(e#ic
p"mp. "roton pumps are electrogenic pumps.
-& Cotra#sport.
In cotransport= and >T"-powered system transports ions or molecules and indirectly powers the
movement of other solutes by maintaining a concentration gradient.
>T" is used to create a gradient of ions or molecules.
Chen these ions or molecules move bac$ to the lower concentration area= they carry with it
the molecules of a solute against the solute concentration gradient.
>T" energy is used indirectly.
/radient energy is used directly.
Lear' the e6ample of the *u!ro*e3proto' !otra'*port *y*tem o' pa0e $47.
.& E/ocytosis&
The cell releases metabolic products to the outside through the fusion of a vesicle with the
plasma membrane.
0& E#docytosis.
In endocytosis materials are ta$en into the cell by engulfing the material with a portion of the
plasma membrane and forming a vesicle or vacuole that is released inside the cell. See f(0.
7."" o' pa0e $49.
Types of endocytosis4
A& $ha(ocytosis. > particle or cell is engulfed and a vacuole is formed.
B& $i#ocytosis. 2issolved materials are ta$en into the cell by forming a vesicle around the
droplets of fluid trapped in folds of the plasma membrane.
C& Receptor'mediated e#docytosis. 9pecific molecules called li(a#ds bind with receptor
molecules in depressions of the plasma membrane called coated pits. The pits are coated with
a layer of protein called clathri#& The receptor molecule bind with the ligand then forms a
coated vesicle that is released to the inside of the cell.
Coating detaches from the vesicle= leaving an uncoated vesicle. The vesicle is now called an
e#dosome. .ndosome divides into a vesicle that returns receptors to plasma membrane= and a
second vesicle that fuses with a lysosome. The contents are digested and returned to the
cytosol. http4AAmicro.magnet.fsu.eduAcellsAendosomesAendosomes.html
.;ample4 Cholesterol is removed from the blood in the form -2- (low-density lipoprotein) by
receptor- mediated endocytosis. If the receptors are defective= -2- remains in the blood leading
to atherosclerosis= the formation of pla8ue on the blood vessel inner surface.
1ere is some additional information about -2- and 12-= and their removal from the blood
stream.
What are the differences in the structures and effects of "good" and "bad" cholesterol?
5ro2()e) by +')rea La)), po*t)o!toral fello1, Baylor Colle0e of &e)(!('e, Hou*to' 7former
HH&I pre)o!toral fello1.8
Chole*terol (* a 1a6y, fatty *ub*ta'!e fou') (' all the !ell* of your bo)y. L(-e other l(p()* 7*u!h
a* fat*8, !hole*terol (* 'ot *oluble (' 1ater 7or (' bloo), 1h(!h (* lar0ely ma)e of 1ater8. To be
tra'*porte), therefore, !hole*terol 'ee)* to be helpe) by *pe!(al !arr(er mole!ule* !alle)
l(poprote('*. The*e !arr(er* are 1hat are be('0 referre) to 1he' you hear about ?0oo)? a')
?ba)? !hole*terol. H(0h3)e'*(ty l(poprote(', or HDL, (* the 0oo) -('), a') the ba) -(') (* lo13
)e'*(ty l(poprote(', or LDL. H(0h !o'!e'trat(o'* of HDL ha2e bee' *ho1' to lo1er your r(*- of
heart atta!-.
It (* thou0ht that HDL !arr(e* !hole*terol a1ay from the arter(e* a') to the l(2er, 1h(!h brea-* (t
)o1'. O' the other ha'), lot* of LDL !hole*terol (' your bloo)*tream lea)* to the )epo*(t(o' of
!hole*terol (' your arter(e*. To0ether 1(th other *ub*ta'!e*, the !hole*terol the' form* artery3
!lo00('0 pla,ue* that@(' the arter(e* fee)('0 your bra(' or heart@!a' lea) to *tro-e or heart
atta!-.
The maAor !ompo'e't of both HDL a') LDL !hole*terol (* a prote(' !alle) apol(poprote('. It
a**o!(ate* 1(th !hole*terol by form('0 a prote(' *hell arou') the ('*oluble !hole*terol, !reat('0
a )(*!o() or *pher(!al part(!le. The *tru!ture* of the HDL3 a') LDL3!hole*terol mole!ule*
them*el2e* are e**e't(ally the *ame, but HDL !hole*terol ha* a h(0her )e'*(ty of apol(poprote('
relat(2e to the amou't of a**o!(ate) !hole*terol tha' LDL )oe*, he'!e the 'ome'!lature. There
are, ho1e2er, )(ffere't form* of apol(poprote(', a') the e6a!t !ompo*(t(o' of )(ffere't
!hole*terol3!o'ta('('0 part(!le* !a' )(ffer. The apol(poprote('* o' the *urfa!e* of the part(!le*
('tera!t 1(th re!eptor* o' the *urfa!e* of other !ell*, lea)('0 to upta-e (' the l(2er 7for HDL
part(!le*8 or )epo*(t(o' (' bloo) 2e**el* 7for LDL part(!le*8.
http122333&hhmi&or(2c(ibi#2as)ascie#tist2hi(hli(ht&pl4)356!ile5a#s3ers7,Fmolec"lar
7,Fa#s89+,&html :this li#) does #ot 3or) a#ymore; < ,99. o3ard "(hes Medical
I#stit"te&
http4AAusers.rcn.comAD$imball.ma.ultranetA&iology"agesA.A.ndocytosis.html Galternate lin$H
H(0h3)e'*(ty l(poprote(' 7HDL8 !(r!ulate* (' the bloo)*tream, e6tra!t('0 !hole*terol from bo)y
t(**ue* a') tra'*port('0 (t to the l(2er for e6!ret(o' or re!y!l('0. I'!rea*e) le2el* of HDL ha2e
bee' !orrelate) 1(th a )e!rea*e) r(*- of athero*!lero*(*333a pr(mary !au*e of !ar)(o2a*!ular
)(*ea*e. Ba*!e't HDL part(!le* are )(*!o()al, !o'*(*t('0 of a pho*phat()yl!hol('e b(layer a') a
prote(' *hell, 1h(!h *h(el)* the hy)rophob(! l(p(), ta(l* from the a,ueou* e'2(ro'me't. +* (t
!(r!ulate* (' the bo)y, HDL !olle!t* !hole*terol, 1h(!h (* the' *tore) (' the l(p() b(layer.
I'!rea*e) eff(!(e'!y (* a!h(e2e) thou0h the a!t(2at(o' of the le!(th('3!hole*terol a!yl tra'*fera*e
7LC+T8 e'Cyme that !o'2ert* the amph(path(! !hole*terol *tore) (' the b(layer ('to hy)rophob(!
!hole*terol e*ter* 1h(!h !olle!t amo'0 the l(p() ta(l*. Th(* (')u!e* a tra'*format(o' of the HDL
)(*- to a *pher(!al form (' 1h(!h a hy)rophob(! !ore of !hole*terol e*ter* (* *h(el)e) by a
!omb('at(o' of l(p() a') prote('. +t th(* *ta0e !hole*terol !olle!t(o' !ea*e* a') the mature HDL
part(!le (* re!o0'(Ce) by the l(2er.
http122333&)s&"i"c&ed"2Research2apoa+2
=ames C& $hillips> ?illy ?ri((ers> @hi(a#( %i> A#a =o#as> a#d Ala"s Sch"lte#& $redicti#(
the str"ct"re o! apolipoprotei# A'I i# reco#stit"ted hi(h'de#sity lipoprotei# dis)s&
Biophysical =o"r#al> B-1,--B'',-.C> +DDB&
>lso4 http4AAwww.hhmi.orgAresearchAinvestigatorsAhobbs.html
S"mmary1
. 0unctions of the plasma membrane
!. 'embrane structure4
fluid-mosaic model
phospholipids= proteins= cholesterol
#. 'embrane proteins
Integral
Transmembrane
"eripheral
/lycoproteins
/lycolipids
0unction of membrane proteins
@ole of carbohydrates
%. Traffic across the plasma membrane
"ermeability and impermeability
Transport proteins4 carrier and channel proteins
*. Transport mechanisms
"assive transport and concentration gradient
9imple diffusion4 dialysis= osmosis= osmotic pressure
9olutions4 hypotonic= hypertonic= isotonic
Carrier mediated transport4
o 0acilitated diffusion
o Carrier-mediated active transport
o 'embrane potential= electrochemical gradient
Cotransport
.ndocytosis
.;ocytosis
"hagocytosis
"inocytosis
@eceptor-mediated endocytosis