chapter 8 cell membrane
TRANSCRIPT
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Chapter 8: Cellular Membranes
Membrane Functions
1. Compartmentalization
• continuous unbroken sheets, enclosecompartments
• enclose entire cell
• allow specialized activities to occur withoutexternal interference
• regulate cellular activities
2. Scaffold for Biochemical Activities
• compartment themselves
• reactants present, relative position notstabilized; interaction dependent on random
collisions
• membrane provide cell with extensiveframework or scaffolding -> use for effectiveinteraction
3. Selectively permeable barrier
• restrict exchange of molecules from oneside to other
• means of communication bet. compartmentsthey separate
• like moat around castle with gated “bridges”promote movement of select elements in/outof cell
4. Transport Solutes
• machinery for physical transport ingsubstances
• from region of low concentration of solute toregion of higher concentration
• accumulate substances (sugars, aminoacids, macromolecules)
• transport specific ions -> ionic gradients
5. Respond to External stimuli
• signal transduction
• receptors combine with ligands (specificmolecules) or other stimuli (light, mechanicaltension)
• diff cells with diff receptors -> capable ofrecognizing/ respond to diff. environmentalstimuli
• interaction of membrane and ext. stimuli ->generate signal stimulates or inhibits internal
activity
6. Intercellular interaction
• mediate interaction between a cell andneighbor
• recognize to adhere and to exchangematerials and information
• proteins may facilitate interaction bet.extracellular and intracellular cytoskeleton
7. Energy Transduction
• energy transduction
• photosynthesis
• light absorbed by membraned boundpigments-> chemical energy stored incarbohydrates
• transfer of chemical energy to ATP
• in mitochondria/chloroplasts
Brief History of Plasma Membranes
- Ernst Overton (1890)• non polar solutes readily dissolve in
nonpolar solvents
• test permeability -> more lipid-soluble solute,more rapid entrance the root hair cells
- E. Gorter and F. Grendel (1925) proposed thatcell membrane contain lipid bilayer
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• extract lipid from RBC measure ant ofsurface area of lipid
• plasma membrane only lipid containingstructure
• contained bimolecular layer of lipid -> lipidbilayer
- polar head group face cytoplasm
- hyrdrophobic fatty acyl chains areprotected from contact with H2O
- decrease in surface tension -> presence ofPROTEINS
• Hugh Davson & James Danielli (1935)->plasma membrane lipid bilayer + lined byglobular proteins; (1950) revised: has outer
and inner protein layers and proteinpenetrate the membrane (provide polarsolutes and ions entrance to cell)
- Jonathan Singer & Garth Nicolson(1972) Fluid-mosaic model
• “central dogma”
• focused on physical state of lipid
• presented in fluid state; lipid molecules movelaterally within plane of membrane
• “mosaic” of discontinuous particles penetratelipid sheet
• dynamic structures-> mobile and can engagein va r ious t ypes o f t r ans ien t o r
semipermanent interactions
Chemical Composition of Membranes
- held together by non-covalent bonds
- lipid bilayer (structural backbone, preventrandom movement of water-soluble materials)
- protein (specific functions)
- ratio of lipid-protein depends on type of cellularmembrane, organism, cell; in other words,depends on the function of membrane in
certain organelle
• mitochondrial membrane - high protein/lipidratio for protein carriers of ETC
• myelin sheath- low concentration of protein,more lipid for electrical insulation
MEMBRANE LIPIDS
- amphipathic both hydrophilic and hydrophobic
- 3 types
1. Phosphoglycerides
• phospholipids (contain phosphate grp)
• b u i l t w i t h g l y c e r o l b a c k b o n e(phosphoglycerides)
• diglycerides (2 hydroxyl grps of glycerolesterified to fatty acid; 3rd is esterfied tohydrophilic P group) -> phosphatidic
acid
• a d d i t i o n a l g r o u p : c h o l i n e (phosphatidylcholine, PC), ethanolamine(PE), serine (PS), inositol (PI)
• small and hydrophilic + negativecharge P = highly water-soluble at oneend (head group)
• @ Physiological pH PS and PI (-); PC andPE (neutral)
• fatty acyl chains are hydrophobic,
unbranched hydrocarbons (16-22carbons)
• membrane FA may be fully saturated (nodouble bonds); monounsaturated (one
double bond); polyunsaturated (>1double)
• contain 1 unsaturated and 1 saturatedfatty acyl chain
• EPA and DHA (2 highly unsat. FA) contain5and 6 double bonds in PE and PC (brainor retina); omega-3 FA
2. Sphingolipids
• less abundant membrane lipid
• sphingosine - amino alcohol with longHC chain
• sphingosine + FA by its amino group -ceramide
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• sphingomyelin (phosphoryl choline)
• glycolipid - substitiution is a carbohydrate
• simple sugar -> cerebroside
• includes silica acid ->ganglioside
• amphiphatic
• FA chains are longer and highly saturated
• glycolipids- Nervous system, myelin sheath(galactocerebroside)
3. Cholesterol
• 50% of lipid molecules
• plant cells contain cholesterol like sterols
• small hydrophilic hydroxyl grip towardmembrane surface
• hydrophobic rings are flat and rigid ->interfere with FA tails of phospholipids
NATURE IMPORTANCE OF LIPID BILAYER
- important effects on biological properties ofmembrane
• composition determine physical state ofmembrane; influence activity of mem.
proteins
• provide precursors for highly active chemicalmessengers regulate cellular function
• flexibility -> membranes are deformable,shape changee (locomotion, cell div)
• facilitate regulated fusion or budding
• maintain proper internal composition,separate charges
• self assemble (liposomes)
ASYMMETRY OF MEMBRANE LIPIDS
- lipid bilayer has 2 distinct leafletswith differentlipid composition
• lipid digesting enzyme cannot fully penetrate
• bilayer contains 2 more/less stableindependent monolayers with diff physical
and chemical properties
• glycolipids -> outer leaflet serve as receptors
• PE -> inner leaflet for curvature (for budding/ fusion)
• PS -> inner “ binding (+) lysine/arginine
• PI -> inner “ phosphorylated on inositol ring -
> phosphoinositides (role in transfer ofstimuli)
MEMBRANE CARBOHYDRATES
- 2-10% by weight (90%) covalently linked toproteins -> glycoproteins; (10%) linked to
lipids -> glycolipids
- glycosylation - addition of carbohydrates mostcomplex modifications
- glycoprotein carbohydrate - short branchedhydrophilic oligosaccharides (15 sugars/
chain)
- carbohydrate projection -> mediate interactionof cell with environment; sorting of membraneproteins to diff cellular compartments
• glycolipids of RBC determine A, B, AB, O
Structure and Function of Membrane
Proteins
- grouped into 3 distinct classes:
1. Intergral proteins
• transmembrane protein; penetrate lipid
bilayer
2. Peripheral Proteins
• entirely outside; either cytoplasmic orextracellular by noncovalent bonds
3. Lipid-anchored proteins
• outside lipid bilayer but are covalently linked to lipid molecule within bilayer
INTERGRAL MEMBRANE PROTEINS
- receptors, channels or transporters, agents thattransfer electrons (photosynthesis/ respiration)
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- amphipathic
- amino acid residues in transmembranedomains form van der Waals with fatty acyl
chains -> seal protein and anchor within bilayer
- DISTRIBUTION OF INTEGRAL PROTEINS:Freeze- Fracture Analysis
• freeze-fracture replication
- structure and properties
• difficult to isolate in soluble form
• use of detergent (SDS -> denature protein;Triton X-100 -> tertiary structure)
PERIPHERAL MEMBRANE PROTEINS
- weak electrostatic bonds
- solubilized by extraction with high conc salsolutions (weaken bond)
- best studied are cytosolic peripheral proteins ->form fibrillar network as “skeleton”
- mechanical support; anchor for integral proteins
- function as enzyme, specialized coats, transmittransmembrane signals
LIPID-ANCHORED MEMBRANE PROTEINS
- smal l , complex o l igosacch l inked tophosphatidylinositol in outer leaflet
- G P I - a n c h o r e d p r o t e i n s - g l y c o s y l -phosphatidylinositol linkagecontained byperipheral proteins
- released by phospholipase (cleave inositolphospholipid)
Membrane Lipids and membrane fluidity
- physical state described by fluidity (viscosity)
- temperature- at which change occur
• Saturated FA
- straight flexible rod
- more compact
• Cis - unsaturated FA
- crooks in chain where there are doublebonds
• greater degree of unsaturation: lowertemperature before bilayer gels
- fatty acid chain length• shorter: lower melting temperature
- by cholesterol
• disrupts close packing of fatty
• abolish sharp transition temperatures;regulate intermediate fluidity
• increase durability while decreasingpermeability of membrane
IMPORTANCE OF MEMBRANE FLUIDITY
- perfect compromise between rigid, orderedstructure (mobility is absent) and completely
fluid (mobility present)
- allow interaction; assemble proteins atparticular sites to form specialized structures(junctions, light capturing, synapses)
- membrane growth accomplished by insertion oflipids and proteins into fluid matrix
- allow cell movement, growth, division,formation of intercellular junctions, secretion,
endocytosis
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MAINTAIN MEMBRANE FLUIDITY
- lower temperature, response is mediated byenzymes that remodel membranes make itcold resistant
• desaturating single bonds; form doublebonds (by desaturases - enzymes catalyzesingle to double bond)
• r e s h u f fl i n g c h a i n s b e t . d i f f e r e n tphospholipids to form 2 unsaturated FA(lower temp) (by phospholipases - split FAfrom glycerol backbone; acyltransferases -transfer FA)
Lipid Rafts
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- distinctive physical properties, micro domainstend to float within the more fluid and
disordered environment of artificial bilayer
Dynamic Nature of Plasma Membrane
- Diffusion of Membrane Proteins after CellFusion
• cell fusion- technique 2 different types ofcells fused to produce one cell with commoncytoplasm and single continuous plasma
membrane
• fuse by making surface “sticky” for adhesion;by addition of inactivated viruses attach tosurface; adding polyethylene glycol, or mild
electric shock
- Restrictions on Protein and Lipid Mobility
• fl u o r e s c e n c e r e c o v e r y a f t e rphotobleaching (FRAP)
- if proteins are mobile; random movementproduce gradua l appearance o ffluorescence in irradiated circle
• single-particle tracking (SPT)
• antibody coated gold particles
- Control of Membrane Mobility
- Membrane Lipid Mobility
• lipid diffuse freely before jumping toneighboring compartment
Movement o f Subs tances across ce l lmembranes
- net flux indicate movement of substance influxand efflux not balanced
- Diffusion
• substance move from region of higherconcentration to lower to regulate overall
concentration
• depends on random thermal motion ofsolutes; exergonic with increase in entropy
• electrochemical gradient
- solute is an electrolyte (charged)
- charges are opposite, moving process isfavorable
- tendency of electrolyte to diffuse dependson 2 gradients:
• chemical gradient - concentrationdifference
• electric potential gradient- difference incharge (ex: K+ higher concentration
inside cell)
- Diffusion of Substances through membranes
- Diffusion of Water through membrane
• semipermeable
• osmosis- lower solute concentration tohigher solute concentration
• hypertonic - compartment with highsolute concentration; shrink; plasmolysis
• hypotonic- compartment with low soluteconcentration; swell;
- Diffusion of Ions through Membranes
• highly impermeable to charged substance
(Na, K, Ca, Cl)
• conductance- rapid movement play role information of nerve impulse, secretion of
substances, muscle contraction, regulationof cell volume, opening of stomatal pores
• Ion channels - permeable to certain ions
1. voltage-gated- difference in ioniccharge on two sides of membrane
2. Ligand-gated channels- binding ofs p e c i fi c m o l e c u l e ( e x .
neurotransmitters)3. M e c h a n o - g a t e d c h a n n e l s -
mechanical forces(stretch tension)
applied
- KcsA (bacterial K+ ion channel
• Facilitated Diffusion
• Glucose transporter
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- phosphorylating sugar after enteringcytoplasm, lowering intracellular
glucose conc.
- GLUT1 to GLUT5 (isoforms)
- Insulin secretion increase glucosetransporters
• Active Transport
- input of energy
Membrane Potentials and Nerve Impulses
- irritability
- nerve cells (neurons) specialized for collection,conduction, transmission of information - formof fast moving electrical impulses
- cell body,dendrites, axon, myelin sheath
- Resting potential
• membrane potential
• resting potential
• action potential -
- Neurotransmission
• synapses- link between neurons
• synaptic cleft - narrow gap in synapse
• presynaptic cell(receptor) conduct impulsetowards synapse
• postsynaptic cell (neuron, muscle, gland cell)lie on receiving side of synapse
• neuromuscular junction- axon and skeletalmuscle ell
• synaptic vesicles
• neurotransmitters
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