Sample Lecture Bio 361

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    Phospholipid structure

    Polar headCholine+

    Phosphate-

    Glycerol

    Non-polarfatty acidchains

    Nonpolar tails

    Hydrophilic

    Hydrophobic

    Many positively chargedgroups can occupy this

    position.

    Bends symbolize doublebonds less densely

    packed lipids.

    Water

    Micelle

    Phospholipidbilayer

    Hydrophobictails

    Hydrophillicheads

    Phospholipid behavior in water

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    Fluidity of membranes

    Frequent lateral movement oflipids; infrequent flip-flopsacross membrane leaflets.

    Flip-flops(rare)

    LateralMovement(frequent)

    (a)

    (b)

    Fluid Viscous

    Unsaturatedfatty acidtails with kinks

    Saturatedfatty acidtails

    Degree of unsaturated/saturatedfatty acid tails (hydrocarbon)determines membrane fluidity.

    Phospholipid composition and habitat temperature

    Phosphatidyl-ethanolamines

    Phosphatidyl-cholines

    Adaptation temperature (C)%o

    fhydrocarbontailsunsaturated

    Fatty acid chains vary in chemical unsaturation =

    double bonds that produce a bend in the chain.Unsaturated phospholipids are less densely packed

    and increase membrane fluidity. To maintainmembrane fluidity at low temperatures, cold adaptedspecies incorporate more unsaturated lipids into their

    membrane.

    after Logue et al. (2000), J Exp Biol 203, 2105-2115

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    PASSIVE: Diffusion as passive solute transport

    Diffusion of glucose toward equilibrium

    High Low

    glucose concentration

    Membrane

    Due to molecular movement glucose passes through the membrane passively.Because there is more glucose on the left side, on average and by chance moremolecules pass from left to right. This leads to equal concentrations on bothsides the equilibrium for this system.

    PASSIVE: Solute diffusion

    High C1 Low C2solute concentration

    J=net rate of diffusion

    X=length

    J= D A C1 C2X

    J ~ C1 C2: Diffusion rate is proportionalto concentration gradient

    Ficks diffusion equation(Adolf Fick, 1829-1901)

    J ~ 1/X: Diffusion rate is greater whendiffusion distance is shorter

    JY ~ C solute Y : Diffusion rate for solute Ydepends on concentration gradientof the solute Y

    J ~ D: Diffusion rate is proportional

    to the diffusion coefficient

    D is a measure of the ease with which asolute moves through a medium separatingthe two concentrations; temperaturedependent; defines permeability.

    How do membranes affect diffusion?

    J ~ A: Diffusion rate is greater whendiffusion area (surface) is greater

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    Outward diffusion from the animal orcell increases the concentration in the

    environmental solution next to theouter surface. The boundary layer

    thus created may be very thin yet stillhave a substantial impact on the rate

    of diffusion.

    Boundary layers and diffusion

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    PASSIVE: Permeability of a membrane for a solute

    Diffusion of glucose

    toward equilibrium

    High Low

    glucose concentration

    Membrane

    J= P A C1

    C2

    P = Dm

    KX

    The permeability of a membrane to a solute isthe ease with which it can move through themembrane by simple diffusion.

    Membrane permeability is based on

    Dm, the diffusion coefficient:rate of diffusion of a substance through a

    membrane

    K, the partition coefficient:

    an indicator for the ratio between dissolvedand undissolved solute in the membrane

    PASSIVE: Membranes and the diffusion of solutes

    The diffusion of a solute across membranes depends on thehydrophobic or hydrophilic nature of the solute.

    1. Lipid solutes (steroids and fatty acids): enter the interior ofthe cell membrane because of their hydrophobic nature.

    2. Molecular oxygen (O2): enter interior of membrane because of theirsmall and nonpolar nature.

    3. Inorganic ions: low permeability across membrane, but can diffusepassively through cell membranes at rapid rates due to ion channels.

    Ion channels allow only passive transport across membranes.They do not bind the ions that pass through them.They are selective in determining which ion can pass.Types include: voltage-gated, stretch-gated, phosphorylation-gated

    and ligand-gated channels.

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    PASSIVE: Selectively permeable membranescan create electrical gradients

    000

    000

    K+

    A1-

    A2-

    Na+

    Membrane isonly permeable

    to K+.

    +++

    ---

    A1-

    A2-

    Na+K+

    K+

    Cross membrane diffusion of K+

    creates positive charges on theright side, preventing K+ from

    further diffusing.

    At the beginningthere is zerocharge across

    the membrane.

    Donnanequilibrium

    PASSIVE: Selectively permeable membranes can

    create electrical gradients: Nernst equation

    Ex = RT ln CoutzF Cin

    R: Gas constantT: absolute temp in Kelvin (18C)F: Faraday constantz: charge on the permeant ion

    Ex = 0.058 log Coutz Cin

    Nernstequation

    electrical gradient = chemical gradient

    Substituting the constants results in:

    Consequences:

    Unequal concentrations of ions across a membranecan create a membrane potential = resting potential.

    A membrane potential can affect the concentrationgradient across a membrane.

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    PASSIVE: Diffusion of ions is affected bya chemical and an electrical gradient

    Electricalgradient

    moves ions insame

    direction asconc.

    gradient =fast

    diffusion.

    Electricalgradient

    opposes conc.gradient =

    slowingdiffusion orreversingdirection.

    High Low

    Na+ concentration

    Diffusion (slow)

    Membrane

    Concentration gradient

    Electrical gradient

    - +

    +++

    ---

    Diffusion (fast)

    High Low

    Na+ concentration

    Membrane

    Concentration gradient

    Electrical gradient

    + -

    +++

    ---

    PASSIVE: An electrochemical view of a single cell

    Cl-

    K+

    A-

    Extracellularfluid

    Na+

    Cell membraneAnionicproteins

    Intra-cellular

    fluid

    K+Cl-

    Na+

    (a) Ion concentration inside asingle animal cell

    ++

    ++

    ++

    ++

    ++

    ++

    --

    ----

    --

    -

    Cl- is near electrochemicalequilibrium because the

    concentration gradient andelectrical effect oppose each

    other.

    Na+ is far fromelectrochemical equilibrium,because the conc. gradient

    and electrical effect move itinto the cell.

    K+: The conc. effect isgreater than the

    electrical effect, but notas much as for Na+.

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    PASSIVE: Facilitated diffusion

    Polar organic solutes (glucose, aminoacids) are hydrophilic: diffuse through

    membrane with the help of carrierproteins: In direction of electrochemical

    gradient Facilitated because faster than simple

    diffusion Reversible and non-covalent binding

    PASSIVE SOLUTE TRANSPORT - SUMMARY

    Passive transport moves towards the electrochemical equilibrium.

    Simple diffusion depends on the concentration gradient for an unchargedsolute. In case of a charged solute, conc. gradients and electrical effectscontribute to diffusion.

    Polar organic solutes (glucose, amino acids) move across membranes withthe help of transporter proteins in the direction of the electrochemicalequilibrium (facilitated diffusion).

    The permeability of a membrane for a lipid solute depends on how readily thethe solute enters into and moves across the membrane lipid bilayer simplediffusion.

    For inorganic ions, the permeability depends on the number of ion channels.

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    Primary and secondary ACTIVE TRANSPORT

    Na+-K+-ATPase is an example of a primary active transport mechanism:

    it draws energy directly from ATP. An ATPase is a transporter and anenzyme.

    Other important ATPases:Ca2+-ATPase in the sarcoplasmic reticulum of muscle cells.H+-K+-ATPase is the proton-pump that acidifies stomach content.vesicular or v-type H+-ATPase in gills and kidneys.

    Secondary active transport draws energy from an electrochemicalgradient of a solute, and therefore indirectly from ATP.

    ACTIVE TRANSPORT

    Primary active-transportmechanisms.

    Cl-

    K+

    A-

    Extracellularfluid

    Na+

    Cell membraneAnionicproteins

    Intra-cellular

    fluid

    K+Cl-

    Na+

    (a) Ion concentration inside asingle animal cell

    ++

    ++

    ++

    ++

    ++

    ++

    --

    ----

    --

    -

    Bloodcapillary

    Intestinalepithelial cells

    Glucose frommeal

    (c) Glucose transport across intestinalepithelium into the blood system

    Cross section ofsmall intestine

    Intestinallumen

    Secondary active-transport mechanisms.

    Blood plasmaPondwater

    Gillepithelial

    cell

    Cl-

    Na+

    Na+

    Cl-

    (b) Ion concentration across gillepithelium of a freshwater fish

    Primary and secondary active-transport mechanisms.

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    ACTIVE: Primary active transport mechanisms

    Cl-

    K+

    A-

    Extracellularfluid

    3Na+

    Anionicproteins

    Intra-cellular

    fluid

    2K+

    Cl-

    Na+

    (a) Ion concentration inside asingle animal cell

    ++

    +

    +

    +

    +

    ++

    +

    --

    --

    -

    -

    ATP

    ADP

    Active transportDiffusion

    Cl- is close to electrochemicalequilibrium.

    Electrochemical gradients favorsteady diffusion of Na+ and K+

    through resting channels in thecell membranes.

    The Na+-K+ pump maintains Na+

    and K+ ions out of equilibrium byusing ATP-bond energy.

    Anionic proteins are trappedinside the cell.

    Bloodcapillary

    Intestinalepithelial cells

    Glucose frommeal

    (c) Glucose transport across intestinalepithelium into the blood system

    Cross section ofsmall intestine

    Intestinallumen

    Apical Basolateralmembrane

    ++

    +

    --

    -

    ACTIVE: Secondary active transport mechanisms

    3Na+

    2K+ATP

    ADP

    Na+ Na+

    GluGlu

    Glu

    2 Na+

    Cotransporter

    Transport against andin direction of electrochem.gradient

    The Na+-K+-ATPase in thebasolateral membrane

    pumps Na+ outside the cell,

    and thereby maintains theNa+ gradient across the

    apical membrane.

    A transporter carriesNa+ towards its EC

    equilibrium, and thereby

    forces the cotransportof glucose across themembrane.

    Na+

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    Apical Basolateral

    membrane

    ++

    ++

    --

    --

    3Na+

    2K+ATP

    ADP

    Na+

    ACTIVE: Ion movements in fresh water fish gills

    Blood plasmaPondwater

    Gillepithelial

    cell

    Cl-

    Na+

    Na+

    Cl-

    (b) Ion concentration across gillepithelium of a freshwater fish

    H+ATP

    ADP

    Transport against andin direction of electrochem.gradient

    Countertransporter

    Channel

    Na+

    A H+-ATPase generates anegative charge inside Na+ follows towards its

    EC equilibrium.

    Cl-

    HCO3-+H+HCO3

    - CO2 CO2 metabolic waste

    A gradient favors the

    outward diffusion ofbicarbonate ions (HCO3

    -),which drives the influx of Cl-

    against the Cl- EC gradient.

    ACTIVE SOLUTE TRANSPORT - SUMMARY

    Solute transport is active if it can move solutes away from electrochemicalequilibrium.

    Active transport is primary if the transporter is an ATPase (energy comesdirectly from ATP), secondary if the energy comes from a solute electro-chemical gradient. Organic solutes are pumped by secondary-activetransport mechanism.

    The transport of ions can create voltage differences (electrogenic) or not(electroneutral).

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    OSMOSIS An exampleIsotonic solution:

    empirical evaluation if

    cell loses or gains water

    Transport across membranes summary II

    The transport of water depends on the colligative properties of a solvent.

    Water moves from solution of lower osmotic pressure (more abundant water)to the solution of higher osmotic pressure (less abundant water).

    If two solutions have the same osmotic pressure they are called iso-osmotic.A solution with higher osmotic pressure is hyper-osmotic relative to one withlower pressure (hypo-osmotic).