Chapter 7 Plasma Membranes A. Fluid Mosaic model 1.

The plasma membrane is the selective permeability boundary that separates the living cell from its surroundings. 2. As we have seen previously, the plasma membrane is composed largely of phospholipids that are amphipathic molecules (molecules that have hydrophilic and hydrophobic regions).

3. The fluid mosaic model states that a membrane is a fluid structure with a mosaic of various proteins embedded in it.

4. Phospholipids and some of the proteins in the plasma membrane can drift laterally within 5. Membranes must be fluid to work properly; they are usually about as fluid as salad oil

6. The steroid cholesterol is sandwiched into the membrane and helps to maintain proper fluid properties despite environmental changes in temperature.

7. Proteins embedded in the membrane determine most of the membranes specific functions

8. Peripheral proteins are bound to the surface of the membrane 9. Integral proteins penetrate the hydrophobic core a. Integral proteins that span the membrane are called transmembrane proteins

b. The hydrophobic regions of an integral protein consist of one or more stretches of nonpolar amino acids, often coiled into alpha helices

1) Role of membrane bound proteins a) Transport

i. ii.

Transport proteins allow passage of hydrophilic substances across the membrane Some transport proteins, called channel proteins, have a hydrophilic channel that certain molecules or ions can use as a tunnel Channel proteins called aquaporins facilitate the passage of water

iii.

b) Enzymatic activity

I. II. III.

Enzyme built into the membrane Active site positioned to the area outside the cell or to the cytoplasm Sometimes arranged in series to help process a particular enzyme catalyzed pathway.

c) Signal transduction

I.

Membrane protein functions as a receptor with a specific site for a particular molecule, the signaling molecule. Example large protein hormones. When the signaling molecule binds to the receptor, it usually causes a shape change in the membrane protein. This shape change, in turn, causes a change in another molecule located on the inside of the membrane and triggers a series of events signal transduction.

II. III.

IV.

d) Cell-cell recognition

i.

Cells recognize each other by binding to surface molecules, often carbohydrates, on the plasma membrane Membrane carbohydrates may be covalently bonded to lipids (forming glycolipids) or more commonly, as we see above, to proteins (forming glycoproteins) Carbohydrates on the external side of the plasma membrane vary among species, individuals, and even cell types in an individual and this variation is important in the recognition between cells.

ii.

iii.

e) Intercellular joining

i)

Proteins in membranes of adjacent cell may hook together to form various kinds of junctions

f)

Attachment to the cytoskeleton and extracellular matrix (ECM)

i.

Cytoskeletal elements may form temporary associations with membrane protein on the cytoplasm side of plasma membrane helps stabilize cell shape and stabilizes proteins position within the membrane (remember these proteins can float in the membrane). ii. membrane proteins on the outer surface of the plasma membrane may bind to elements of the extra cellular matrix. Can be part of a pathway that transmits information from the exterior environment to the interior of the cell

2)

Cell permeability, Diffusion, and osmosis a) Nonpolar (Hydrophobic) molecules can pass through the membrane relatively easily b) Polar molecules such as sugars and charged amino acids as well as ions cannot pass through the hydrophobic layer of the plasma membrane. c) As we have already seen transport or carrier proteins within the membrane are required to move these molecules and ions. d) Material moves through the membrane either by diffusion down a concentration or pressure gradient or against the concentration gradient by active transport. i) Diffusion (1) Molecules tend to spread out evenly within an available space. (a) Molecules remain in motion (b) If there are initially more molecules in one area, random motion will result in a net into areas with fewer molecules. Eventually, a state will be reached in which for every movement in one direction there is a corresponding movement in the opposite direction and the solution will be said to be in equilibrium. (i) Equilibrium should be defined as net movement = 0 rather than equal concentration. (2) We are interested in diffusion across a selectively permeable membrane. (a) The membrane partitions the solution into two separate solutions one on each side of the membrane. (b) The substances will diffuse across the membrane from an area of high concentration to an area of low concentration. Keep in mind there are other factors beside concentration that can impact rates of molecular motion and therefore, diffusion.

(c) The energy for this movement has come from molecular motion. The cell has not had to expend any energy of its own. This type of movement across the plasma membrane is termed passive transport.

(3) Osmosis the diffusion water across a selectively permeable membrane. (a) Water diffuses across the membrane from an area of high water concentration or LOW SOLUTE concentration to an area of low water concentration or HIGH SOLUTE concentration. (b) Water concentration in a solution is a function of solute concentration. If there is a higher concentration of solute in the solution on one side of the membrane, then the concentration of water is going to be lower. (i) For example: inside outside 10% Na+ 10% Cl5 % Glucose 25 % 75% 15 % Na+ 15 % Cl0 % Glucose 30% 70%

Total solute? Water Concentration?

(ii) When discussing osmosis, then, we have to have a measure of TOTAL solute concentration 1. Tonicity is the term that is used to describe the cells ability to gain or lose water a. Isotonic solution: Solute concentration outside the cell is the same as that inside the cell; no net water movement across the plasma membrane b. Hypotonic solution: Solute concentration outside the cell is less than that inside the cell; Therefore, the water concentration is greater outside the cell than inside the cell and, therefore, the cell gains water. c. Hypertonic solution: Solute concentration outside the cell is greater than that inside the cell; Therefore, the water concentration is less outside the cell than inside the cell and, therefore, the cell loses water (c) Thus a cell can find itself in one of three situations with respect to its external environment. This external environment can be an actual aquatic environment for organisms living in water or it can be the tissue fluids in the extracellular matrix surrounding cells in tissues in multicellular organisms. (d) The external environment is isotonic most marine invertebrates, plants live in this kind of environment. The tissue fluids of most multicellular animals are maintained in an isotonic state with the cells they are associated with. (i) Cells that are isotonic to the environment do not have to regulate their water content. (ii) Like all organisms, they do have to regulate their individual solute components however

(e) Hypertonic or hypotonic environments create problems for organisms and require osmoregulatory mechanisms (i) The second situation a cell can find itself in is one in which the environment is hypotonic to the cell and the cell has to deal with the gain of water in the cells that are in contact with this environment. All freshwater organisms have to deal with this as do the cartilaginous fish. 1. Plants actually turn this gain of water into a plus. In this example, a red blood cell which lacks any osmoregulatory abilities is placed in fresh water. Red blood cell builds up turgor pressure as water flows down the concentration gradient. As the turgor pressure rises the cell swells (no cell wall to prevent swelling). At some point the internal turgor pressure exceeds the strength of the plasma membrane and the cell bursts in a process termed cytolysis. Plant cells build turgor pressure as water flows into the cell down the concentration gradient. As the turgor pressure rises, the rate at which water molecules leave because of the pressure gradient out rises. The cell wall prevents cytolysis and eventually equilibrium is reached when turgor pressur out = osmotic pressure in and the net flow of water = 0)! 2. Animal cells or multicellular animals living in freshwater have had to evolve osmoregulatory mechanisms that invest energy to remove water at a rate equal to the rate at which it enters passively by osmosis. a. Example 1 Freshwater Protozoans such as paramecia the contractile vacuole.

b. Example 2 The vertebrate kidney. Freshwater fish have minimize their water loss by i. Lowering their total solute concentration an adaptation the evolved very early as the ancestors of all fish moved into freshwater from the oceans of the ancient world. ii. Waterproofing their bodies with scales and layers of mucus. iii. Cells of gills cannot be waterproofed however why not? iv. Large quantities of water diffuse across the gills. There is also a loss of salts at the gills v. Kidney is capable of removing enormous amounts of water from the blood producing a copious dilute urine. (ii) The third situation a cell can find itself in is one in which the environment is hypertonic to the cell and the cell has to deal with the loss of water in the cells that are in contact with this environment. Because of their evolutionary history, bony fish (osteichthyes) have this problem in the ocean.

In this example, both a red blood cell and a plant cell which lack any osmoregulatory abilities in hypertonic environments are placed in salt water. The cells loose turgor pressure as water flows out along the concentration gradient. As the turgor pressure falls the cell shrinks and at some point the plasma membrane is compromised and the cell is said to be plasmolyzed.

1. Because of their evolutionary history, bony fish (osteichthyes) have this problem in the ocean because of the lower their total solute concentration their ancestors brought with them from freshwater when the modern fish moved back into the (remember most marine organisms are isotonic to the ocean). i. Waterproofing their bodies with scales and layers of mucus. ii. Cells of gills cannot be waterproofed however why not? iii. Large quantities of water diffuse across the gills. They also gain salts at the gills. iv. They drink ocean water. Leaves a great big problem with salt intake. v. Kidney produces very concentrated urine (just enough to remove excess ions).

(4) Role of transport proteins in the movement of material across membrane. (a) Facilitated diffusion - transport proteins speed the passive movement of molecules across the plasma membrane (i) Channel proteins provide corridors that allow a specific molecule or ion to cross the membrane

Water aquaporin Ion channels May be static allowing continuous diffusion Maybe capable of changing shape in response to some stimulus to open or close the channel. These channels are said to gated channels (ii) Carrier proteins - undergo a subtle change in shape that translocates the solute-binding site across the membrane assisting in the movement of solute across the membrane 1. 2. a. b.

1. still passive because the solute moves down its concentration gradient 2. no energy expended (b) Active transport (i) movement against the concentration gradient (ii) requires energy, usually in the form of ATP (iii) allows cells to maintain concentration gradients that differ from their surroundings (iv) Especially important in ionoregulation remember we discussed that cells not only have to deal with gain or loss of water through osmosis but also with gain or loss of solute.

(v) One of the best studied active transport mechanisms is the sodium-potassium pump.

1. In this example, the concentration of sodium (Na+)on inside of cell lower than outside; while the concentration of potassium (K+). The cell needs to maintain this relationship. 2. In what direction does sodium want to diffuse ( in or out of the cell)? How about potassium ? 3. To maintain a steady concentration of these two ions, the cell must actively transport using a sodium potassium pump to compensate for diffusion. 4. First the pump picks up 3 sodium ions 5. Then the pump is phosphorylated by ATP ADP and charges up with energy by the loading the phosphate. 6. The pump changes shape an translocates the three sodium ions to the outside. 7. The pump then loads with two potassium ions 8. The phosphate is released and the pump changes shape to translocate the two potassium ions to the cytoplasm. (vi) Active transport and membrane potential 1. Membrane potential is the voltage difference across a membrane 2. Voltage is created by differences in the distribution of positive and negative ions 3. Two combined forces, collectively called the electrochemical gradient, drive the diffusion of ions across a membrane: a. A chemical force (the ions concentration gradient) b. An electrical force (the effect of the membrane potential on the ions movement)

4. An electrogenic pump is a transport protein that generates voltage across a membrane a. The sodium-potassium pump is the major electrogenic pump of animal cells b. The main electrogenic pump of plants, fungi, and bacteria is a proton pump

c. We will also see a series of special proton pump in mitochondria and chloroplasts that do not run on ATP. 5. Cotransport occurs when active transport of a solute indirectly drives transport of another solute a. Plants commonly use the gradient of hydrogen ions generated by proton pumps to drive active transport of nutrients into the cell

(5) Bulk flow across the membrane -Large molecules, such as polysaccharides and proteins, cross the membrane in bulk via vesicles (a) Bulk transport requires energy (b) Bulk flow from cytoplasm to the exterior of the cell - exocytosis, transport vesicles migrate to the membrane, fuse with it, and release their contents. (i) Many secretory cells use exocytosis to export their products (c) Bulk flow from the exterior of the cell into cytoplasm - endocytosis, membrane invagenates and surround a portion of the exterior fluid forming a vesicle which becomes incorporated into the cytoplasm. Two types. (i) Phagocytosis 1. In phagocytosis a cell engulfs a particle in a vacuole 2. The vacuole fuses with a lysosome to digest the particle

(ii) Pinocytosis 1. In pinocytosis, molecules are taken up when extracellular fluid is gulped into tiny vesicles

(d) receptor-mediated endocytosis, binding of ligands to receptors triggers vesicle formation (i) A ligand is any molecule that binds specifically to a receptor site of another molecule (ii) Receptors on the surface of the cell. 1. Usually clustered in areas of the cell membrane called coated pits which are lined on the cytoplasm side by a fuzzy coat of protein (iii) When receptor binds to ligand the coated pit forms into a vesicle bringing the ligand molecule any other molecules in the area into the cytoplasm of the cell (iv) Vesicle empties content into cytoplasm and vesicle reunites with membrane returning receptors to the surface of the cell. (v) Example LDL receptors in humans. Part of mechanism by which cells take up cholesterol for use within the cell for use in the synthesis of membrane components and other steroid based molecules such as steroid hormones.