1. Copyright © 2009 Pearson Education, Inc., publishing as Pearson Benjamin Cummings. BIOLOGY A GUIDE TO THE NATURAL WORLD FOURTH EDITION DAVID KROGH Life’s Border: The Plasma Membrane

2. Copyright © 2009 Pearson Education, Inc., publishing as Benjamin Cummings. 5.1 The Nature of the Plasma Membrane

3. Copyright © 2009 Pearson Education, Inc., publishing as Benjamin Cummings. The Nature of the Plasma Membrane The plasma membrane is a thin, fluid entity that manages to be very flexible and yet is stable enough to stay together despite being continually remade due to the constant movement of materials in and out of it.

4. Copyright © 2009 Pearson Education, Inc., publishing as Benjamin Cummings. The Plasma Membrane In animal cells, the plasma membrane has four principal components: 1. A phospholipid bilayer. 2. Molecules of cholesterol interspersed within the bilayer.

5. Copyright © 2009 Pearson Education, Inc., publishing as Benjamin Cummings. The Plasma Membrane 3. Proteins that are embedded in or that lie on the bilayer. 4. Short carbohydrate chains on the cell surface, collectively called the glycocalyx, that function in cell adhesion and as binding sites on proteins.

6. Copyright © 2009 Pearson Education, Inc., publishing as Benjamin Cummings. The Plasma Membrane Figure 5.1 glycocalyx proteins integral protein peripheral protein cytoskeleton cholesterol Phospholipid bilayer: a double layer of phospholipid molecules whose hydrophilic “heads” face outward, and whose hydrophobic “tails” point inward, toward each other. Glycocalyx: sugar chains that attach to proteins and phospholipids, serving as protein binding sites and as cell lubrication and adhesion molecules. Cholesterol molecules that act as a patching substance and that help the cell maintain an optimal level of fluidity. Proteins, which are integral, meaning bound to the hydrophobic interior of the membrane, or peripheral, meaning not bound in this way. phospholipids cell interior cell exterior

7. Copyright © 2009 Pearson Education, Inc., publishing as Benjamin Cummings. The Phospholipid Bilayer Phospholipids are molecules composed of two fatty acid chains linked to a charged phosphate group.

8. Copyright © 2009 Pearson Education, Inc., publishing as Benjamin Cummings. The Phospholipid Bilayer The fatty acid chains are hydrophobic, meaning they avoid water, while the phosphate group is hydrophilic, meaning it readily bonds with water.

9. Copyright © 2009 Pearson Education, Inc., publishing as Benjamin Cummings. The Phospholipid Bilayer Figure 5.2 polar head nonpolar tails watery extracellular fluid watery cytosol hydrophilic hydrophobic hydrophobic molecules pass through freely hydrophilic molecules do not pass through freely hydrophilic (a)(b)Phospholipid moleculePhospholipid bilayer –

10. Copyright © 2009 Pearson Education, Inc., publishing as Benjamin Cummings. The Phospholipid Bilayer Such phospholipids arrange themselves into bilayers—two layers of phospholipids in which the fatty acid “tails” of each layer point inward (avoiding water), while the phosphate “heads” point outward (bonding with it).

11. Copyright © 2009 Pearson Education, Inc., publishing as Benjamin Cummings. The Phospholipid Bilayer Phospholipids take on this configuration in the plasma membrane because a watery environment lies on either side of the membrane.

12. Copyright © 2009 Pearson Education, Inc., publishing as Benjamin Cummings. The Phospholipid Bilayer In animal cells, the cholesterol molecules that are interspersed between phospholipid molecules in the plasma membrane perform two functions: 1.They act as a patching material that helps keep some small molecules from moving through the membrane. 2.They keep the membrane at an optimal level of fluidity.

13. Copyright © 2009 Pearson Education, Inc., publishing as Benjamin Cummings. The Phospholipid Bilayer Some plasma membrane proteins are integral, meaning they are bound to the hydrophobic interior of the phospholipid bilayer. Others are peripheral, meaning they lie on either side of the membrane but are not bound to its hydrophobic interior.

14. Copyright © 2009 Pearson Education, Inc., publishing as Benjamin Cummings. Membrane Protein Functions In animal cells, the cholesterol molecules that are interspersed between phospholipid molecules in the plasma membrane perform two functions: 1.structural support 2.cell identification, by serving as external recognition proteins that interact with immune system cells

15. Copyright © 2009 Pearson Education, Inc., publishing as Benjamin Cummings. Membrane Protein Functions 3.communication, by serving as external receptors for signaling molecules 4.transport, by providing channels for the movement of compounds into and out of the cell

16. Copyright © 2009 Pearson Education, Inc., publishing as Benjamin Cummings. The Plasma Membrane Figure 5.3 (a)(b)(c)(d)Structural supportRecognitionCommunicationTransport Membrane proteins can provide structural support, often when attached to parts of the cell’s scaffolding or “cytoskeleton.” Binding sites on some proteins can serve to identify the cell to other cells, such as those of the immune system. Receptor proteins, protruding out from the plasma membrane, can be the point of contact for signals sent to the cell via traveling molecules, such as hormones. Proteins can serve as channels through which materials can pass in and out of the cell.

17. Copyright © 2009 Pearson Education, Inc., publishing as Benjamin Cummings. The Plasma Membrane The plasma membrane today is described by a conceptualization called the fluid-mosaic model It views the membrane as a fluid, phospholipid bilayer that has a mosaic of proteins either fixed within it or capable of moving laterally across it.

18. Copyright © 2009 Pearson Education, Inc., publishing as Benjamin Cummings. 5.2 Diffusion, Gradients, and Osmosis

19. Copyright © 2009 Pearson Education, Inc., publishing as Benjamin Cummings. Diffusion, Gradients, and Osmosis Diffusion is the movement of molecules or ions from a region of their higher concentration to a region of their lower concentration.

20. Copyright © 2009 Pearson Education, Inc., publishing as Benjamin Cummings. Diffusion, Gradients, and Osmosis A concentration gradient defines the difference between the highest and lowest concentrations of a solute within a given medium. Through diffusion, compounds naturally move from higher to lower concentrations, meaning down their concentration gradients.

21. Copyright © 2009 Pearson Education, Inc., publishing as Benjamin Cummings. Diffusion, Gradients, and Osmosis Figure 5.4 water molecules dye molecules (a)(b)(c)Dye is dropped inDiffusion beginsDye is evenly distributed

22. Copyright © 2009 Pearson Education, Inc., publishing as Benjamin Cummings. Diffusion, Gradients, and Osmosis Energy must be expended to move compounds against their concentration gradients, meaning from a lower to a higher concentration.

23. Copyright © 2009 Pearson Education, Inc., publishing as Benjamin Cummings. Diffusion, Gradients, and Osmosis A semipermeable membrane is one that allows some compounds to pass through freely while blocking the passage of others.

24. Copyright © 2009 Pearson Education, Inc., publishing as Benjamin Cummings. Diffusion, Gradients, and Osmosis Osmosis is the net movement of water across a semipermeable membrane from an area of lower solute concentration to an area of higher solute concentration.

25. Copyright © 2009 Pearson Education, Inc., publishing as Benjamin Cummings. Diffusion, Gradients, and Osmosis Because the plasma membrane is a semipermeable membrane, osmosis operates in connection with it. Osmosis is a major force in living things; it is responsible for much of the movement of fluids into and out of cells.

26. Copyright © 2009 Pearson Education, Inc., publishing as Benjamin Cummings. Diffusion, Gradients, and Osmosis Figure 5.5 (a) An aqueous solution divided by a semipermeable membrane has a solute —in this case, salt— poured into its right chamber. solute solvent semipermeable membrane pure water osmosis water bound to salt ions (b) As a result, though water continues to flow in both directions through the membrane, there is a net movement of water toward the side with the greater concentration of solutes in it. (c) Why does this occur? Water molecules that are bonded to the sodium (Na + ) and chloride (Cl – ) ions that make up salt are not free to pass through the membrane to the left chamber of the container.

27. Copyright © 2009 Pearson Education, Inc., publishing as Benjamin Cummings. Osmotic Imbalances Osmotic imbalances can cause cells either to dry out from losing too much water or, in the case of animal cells, to break from taking too much water in. Plant cells generally do not have this problem because their cell walls limit their uptake of water.

28. Copyright © 2009 Pearson Education, Inc., publishing as Benjamin Cummings. Solute Concentration Cells will gain or lose water relative to their surroundings in accordance with what the solute concentration is inside the cell as opposed to outside it.

29. Copyright © 2009 Pearson Education, Inc., publishing as Benjamin Cummings. Solute Concentration A cell will lose water to a surrounding solution that is hypertonic—a solution that has a greater concentration of solutes in it than does the cell’s cytoplasm. A cell will gain water when the surrounding solution is hypotonic to the cytoplasmic fluid.

30. Copyright © 2009 Pearson Education, Inc., publishing as Benjamin Cummings. Solute Concentration Figure 5.6 Balanced water movement Net movement of water into cell Animal cell: plasma membrane plasma membrane H2OH2O H2OH2O Plant cell: cell wall wilted turgid (a) (b)(c) Hypertonic surroundings Isotonic surroundings Hypotonic surroundings Net movement of water out of cell H2OH2O H2OH2O H2OH2O H2OH2O

31. Copyright © 2009 Pearson Education, Inc., publishing as Benjamin Cummings. Solute Concentration Water flow is balanced between the cell and its surroundings when the surrounding fluid and the cytoplasmic fluid are isotonic to each other—when they have the same concentration of solutes.

32. Copyright © 2009 Pearson Education, Inc., publishing as Benjamin Cummings. Plasma Membranes and Diffusion PLAY Animation 5.1: Plasma Membranes and Diffusion

33. Copyright © 2009 Pearson Education, Inc., publishing as Benjamin Cummings. 5.3 Moving Smaller Substances In and Out

34. Copyright © 2009 Pearson Education, Inc., publishing as Benjamin Cummings. Moving Smaller Substances In and Out Some compounds are able to cross the plasma membrane strictly through diffusion; others require diffusion and special protein channels; still others require protein channels and the expenditure of cellular energy.

35. Copyright © 2009 Pearson Education, Inc., publishing as Benjamin Cummings. Transport Through the Plasma Membrane Figure 5.7 Passive transportActive transport simple diffusionfacilitated diffusion Materials move down their concentration gradient through the phospholipid bilayer. The passage of materials is aided both by a concentration gradient and by a transport protein. Molecules again move through a transport protein, but now energy must be expended to move them against their concentration gradient. ATP

36. Copyright © 2009 Pearson Education, Inc., publishing as Benjamin Cummings. Transport Through the Plasma Membrane Active transport is any movement of molecules or ions across a cell membrane that requires the expenditure of energy. Passive transport is any movement of molecules or ions across a cell membrane that does not require the expenditure of energy.

37. Copyright © 2009 Pearson Education, Inc., publishing as Benjamin Cummings. Types of Passive Transport There are two forms of passive transport: simple diffusion and facilitated diffusion. For either form of transport to bring about a net movement of materials into or out of a cell, a concentration gradient must exist.

38. Copyright © 2009 Pearson Education, Inc., publishing as Benjamin Cummings. Types of Passive Transport A concentration gradient is all that is required for simple diffusion to operate. Facilitated diffusion, however, requires both a concentration gradient and a protein channel.

39. Copyright © 2009 Pearson Education, Inc., publishing as Benjamin Cummings. Facilitated Diffusion In facilitated diffusion, transport proteins function as channels for larger hydrophilic substances—substances that, because of their size and electrical charge, cannot diffuse through the hydrophobic portion of the plasma membrane.

40. Copyright © 2009 Pearson Education, Inc., publishing as Benjamin Cummings. Facilitated Diffusion Figure 5.8 inside cell outside cell plasma membrane glucose The transport pro- tein has a binding site for glucose that is open to the outside of the cell. Glucose binds to the binding site. This binding causes the protein to change shape, exposing glucose to the inside of the cell. Glucose passes into the cell and the protein returns to its original shape. 1.2.3.4.

41. Copyright © 2009 Pearson Education, Inc., publishing as Benjamin Cummings. Active Transport Cells cannot rely solely on passive transport to move substances across the plasma membrane. A cell may need to maintain a greater concentration of a given substance on one side of its membrane. Yet, passive transport equalizes concentrations of substances on both sides of the plasma membrane.

42. Copyright © 2009 Pearson Education, Inc., publishing as Benjamin Cummings. Active Transport To deal with such needs, cells use active transport. Chemical pumps move compounds across the plasma membrane against their concentration gradients.

43. Copyright © 2009 Pearson Education, Inc., publishing as Benjamin Cummings. Active Transport One example of such transport is the pumping of glucose into cells that line the small intestines.

44. Copyright © 2009 Pearson Education, Inc., publishing as Benjamin Cummings. 5.4 Getting the Big Stuff In and Out

45. Copyright © 2009 Pearson Education, Inc., publishing as Benjamin Cummings. Getting the Big Stuff In and Out Larger materials are brought into the cell through endocytosis and moved out through exocytosis.

46. Copyright © 2009 Pearson Education, Inc., publishing as Benjamin Cummings. Exocytosis and Endocytosis Both mechanisms employ vesicles, the membrane-lined enclosures that alternately bud off from membranes or fuse with them.

47. Copyright © 2009 Pearson Education, Inc., publishing as Benjamin Cummings. Exocytosis In exocytosis, a transport vesicle moves from the interior of the cell to the plasma membrane and fuses with it, at which point the contents of the vesicle are released to the environment outside the cell.

48. Copyright © 2009 Pearson Education, Inc., publishing as Benjamin Cummings. Exocytosis Figure 5.9 (a)(b)ExocytosisMicrograph of exocytosis extracellular fluid protein cytosol transport vesicle

49. Copyright © 2009 Pearson Education, Inc., publishing as Benjamin Cummings. Endocytosis There are two principal forms of endocytosis: pinocytosis and phagocytosis.

50. Copyright © 2009 Pearson Education, Inc., publishing as Benjamin Cummings. Endocytosis 1.Pinocytosis is the movement of moderate- sized molecules into a cell by means of the creation of transport vesicles produced through an infolding or “invagination” of a portion of the plasma membrane.

51. Copyright © 2009 Pearson Education, Inc., publishing as Benjamin Cummings. Endocytosis 2.Phagocytosis is when certain cells use pseudopodia or “false feet” to surround and engulf whole cells, fragments of them, or other large organic materials.

52. Copyright © 2009 Pearson Education, Inc., publishing as Benjamin Cummings. Endocytosis Figure 5.10 receptors captured molecules coated pit vesicle bacterium (or food particles) pseudopodium (a)Pinocytosis (b)Phagocytosis Formation of a pinocytosis vesicle. A human macrophage (colored blue) uses phagocytosis to ingest an invading yeast cell.

53. Copyright © 2009 Pearson Education, Inc., publishing as Benjamin Cummings. Endocytosis In pinocytosis, materials are brought into the cell inside vesicles that bud off from the plasma membrane.