Biomembranes

An outline
Membrane Structure
science
A changing picture - shows the process of A fluid mosaic of lipids, proteins, carbohydrates

Membrane Function Selectively permeable

Passive transport Diffusion OSMOSIS = diffusion of water Active Transport Pumping of solutes AGAINST a gradient Requires energy

Biomembranes
A membrane is a fluid mosaic of lipids, proteins and carbohydrates

(Figure 3-32. from Molecular Cell Biology. Lodish)

Oil spreading on water

A Langmuir trough is a simple device for controlling the spreading of an oil or fat on a water surface. The molecules in the film become orientated so that their hydrophobic tails are in the air and their polar heads are in the water. A key part of this device is a method for moving the barrier to cause a defined lateral pressure against the oil layer.

Membrane structure
The model of the plasma membrane has changed over time - shows how science proceeds Example: even well before scientists could see the membrane, they knew it was made of lipids..how did they know that?

They could measure that lipids entered cells more rapidly than substance that were insoluble in lipids

Membrane structure
They eventually concluded that PHOSPHOLIPIDS were the main backbone of the membrane and when they measured the amount of phospholipids - they found 2X the amount to cover the cell?

Explanation ?

Membrane structure
The hydrophilic head of phospholipids are in water, and the hydrophobic tails are excluded from water

A bilayer of phospholipids form a stable boundery between two aqueous compartments

Phospholipids are the main lipid constituents of most biomembranes

Phosphate group

+ Glycerol + 2 Fatty Acids

Hydrophillic and hydrophobic regions in one molecule

AMPHIPATHIC

Phospholipid bilayer
Hydrocarbon side chains in each leaflet form a hydrophobic core that is 3-4 nm thick. Phospholipids differ in the charge carried by the polar head groups at neutral pH. phosphatidylcholine has no net electric charge. phosphatidylserine has a net negative charge. Polar head groups in all phospholipids can pack together into the characteristic bilayer structure.

(Figure 5-30. from Molecular Cell Biology. Lodish)

Steroids, another important class of membrane lipids

Basic structure is a 4 ring hydrocarbon. Cholesterol is the major steroidal constituent of animal tissues, with one hydroxyl substituent on one ring.
Abundant in plasma membrane (30-50% of total lipids).
(Figure 5-29. from Molecular Cell Biology. Lodish)

Absent in prokaryotic cells

Chemical analysis of membranes from red blood cells showed PROTEIN ! Proteins must be part of the membrane - but how? A protein coat? The Davson- Danielli Model

DAVSON-DANIELLI MODEL

FLUID MOSAIC MODEL

Membrane proteins
Classification: Integral:
Move across the membrane Embebed in the phospholipid bilayer Form channels or pores Structural proteins

Functions: Peripherical:

Anchoring to cytoskeletal proteins Do not interact with the hydrobobic core of the bilayer Function: Cell-cell signalling or interactions

Integral membrane proteins

Peripheral membrane proteins

Fluid mosaic model

Properties of biomembranes

Asymmetry Motility

Asymmetry Phospholipid composition of the two leaflets

Different abundance of phospholipid and cholesterol in both membrane leaflets. It is not clear how these differences in lipid composition of the two leaflets arise. One possibility is that certain lipids bind to specific protein domains that occur preferentially in one membrane leaflet. Lipids with positively charged head group are found in the exoplasmic leaflet. Lipids with neutral or negative polar head groups located in the cytosolic leaflet.

Biological membranes asymmetry

Essential for the structure and function
Each type of integral membrane protein has a single orientation to the cytosolic and exoplasmic faces of a cellular membrane. Proteins have never been observed to flip-flop across a membrane (unfavorable) Glycoproteins and glycolipids are abundant in the plasma membrane.
Glycolipids are attached to carbohydrates localized to the exoplasmic membrane face. Glycoproteins with their polar carbohydrate chains away from the cell.
(Figure 3-33. from Molecular Cell Biology. Lodish)

X-ray diffraction analysis of myelin membranes

Observations: Regular variation in density is consistent with a bilayer organization of each membrane unit. Protein is located mainly on the hydrophilic external faces or central region.
(Figure 5.33. from Molecular Cell Biology. Lodish)

Fluidity of biomembranes
1. Fatty acid chains
Long and saturated melt at higher temperatures form stronger van der Waals interactions one each other and gel state is stabilized . Short and unsaturated melt at lower temperatures less surface area to form van der Waals interactions the kinks in unsaturated forming less stable van der waals interactions.

(Figure 2-18. from Molecular Cell Biology. Lodish)

Fluidity of biomembranes
2. Cholesterol
Cholesterol: is too hydrophobic to form a sheet structure. The net effect depends on the lipid composition. Restricts the random movement of the polar heads of the fatty acyl chains. Separates and disperses their tails, bilayer to become slightly more fluid.
(Figure 5-29. from Molecular Cell Biology. Lodish)

The fluidity of the plasma membrane

Lipids and proteins drifting laterally

Unsaturated Fatty Acids = more fluid

Cholesterol makes membranes less fluid

Lipids and proteins are laterally mobile in biomembranes

Thermal motion permits phospholipid and glycolipid molecules:
to rotate freely around their long axes and to diffuse laterally within the membrane leaflet

A typical lipid molecule :

exchanges places about 107 times per second. diffuses several micrometers per second (37 C).
(Figure 5-35. from Molecular Cell Biology. Lodish)

Todays picture

Typical roles of cell membranes

Compartmentalization of cells
Separation of the interior of the cell from its exterior environment Establishment of subcellular organelles Selective transport of ions and metabolites cell metabolism protein biosynthesis enzymatic regulatory processes Oxidative phosphorylation Photosynthesis Neural transmission Cell-cell communication

Semipermeable barrier

Site of complex enzymatic reaction

Energy production and transduction Cell communication and regulation

Traffic across membranes

Not all traffic is equal Membrane are SELECTIVELY PERMEABLE DEPENDS ON Selective Permeability: Some substances cross more easily than others - Membrane solubility characteristics - Specific integral transport proteins

Passive transport
Passive transport across a membrane is by DIFFUSION DIFFUSION ? Net movement of a substance down a concentration gradient. Your examples of diffusion? Smells?

(Figure 8-10 from Campbell)

Diffusion of water?

Hypertonic = a solution with greater solute concentration Hypotonic = a solution with a lower solute concentration
(Figure 8-11 from Campbell)

How osmosis can affect cells?

(Figure 8-12 from Campbell)

Facilited diffusion across a membrane

The role of:

A passive process.? Yes Assists molecules that are impeded by the phospholipids e.g., Polar molecules and ions

SIMILARITIES WITH ENZYMES

Solute specific Can be saturated with solute i.e. a maximum transport rate Subject to inhibition by similar molecules (competitive inhibition)

A model of facilitated diffusion

(Figure 8-14b from Campbell)

Transport proteins alternate betwen 2 shapes called CONFORMATIONS

Passive vs active transport

(Figure 8-16 from Campbell)

The sodium potassium pump

Per cycle: OUT: 3 Na+ INTO: 2 K+

(Figure 8-16 from Campbell)

Against a concentration gradient = A requirement for energy (ATP) = ACTIVE TRANSPORT!

Membrane potential
Some ion pumps create an electrical potential (voltage) across the membrane from -50 to -200 mv (inside is negative)

A transport protein that generates voltage across a membrane

Transport of large molecules?

Large molecules = proteins and polysaccharides

IN
ENDOCYTOSIS (3 types) Phagocytosis Pinocytosis Receptormediated endocytosis IN

OUT
EXOCYTOSIS Fusion of vesicles with the plasma membrane Vesicles from the ER and/or Golgi OUT

Endocytosis
1. Phagocytosis 2. Pinocytosis 3. Receptormediated endocytosis

Receptor-mediated endocytosis
Importation of specific macromolecules by the inward budding of vesicles Enables cells to import bulk quantities of specific substances even if they are in low concentration in the extracellular fluid!

Every cellular membrane forms a closed compartment

Phospholipid bilayer membranes spontaneously seal to form closed structures that separate two aqueous compartments.

(Figure 2-20. from Molecular Cell Biology. Lodish)

(Figure 5-31. from Molecular Cell Biology. Lodish)