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, by Campbell and Reece)
Membrane Structure and Function (Chapter Eight)
MEMBRANE STRUCTURE The selectively permeable plasma membrane allows certain substances easier passage than others. Lipids and proteins are the staple ingredients of membranes. Phospholipids are especially important. A phospholipid is an molecule, amphipathic molecule meaning it has hydrophilic and hydrophobic regions. The fluid mosaic model states that the membrane is a fluid structure with various proteins embedded in or attached to a bilayer of phospholipids. Membrane Models Have Evolved to Fit New Data Scientists worked to discover how the membrane was composed long before membranes could be seen with electron microscopes. The Dutch scientists E. Gorter and F. Grendel reasoned that cell membranes must be phospholipid bilayers, with the hydrophobic tails facing each other and the hydrophobic heads in contact with water. The Davson-Danielli model shows a phospholipid bilayer sandwiched in between two protein layers. It was also thought that all cell membranes were the same. However, scientists later recognized problems with these conclusions. With further usage of the electron microscope, it was discovered that not all membranes look alike. Membranes with different functions differ in chemical composition and structure. Additionally, membrane proteins are not very soluble in water, and are amphipathic as well. If they were layered on top of the phospholipid bilayer, their hydrophobic regions would be in contact with an aqueous environment. As a result, S. J. Singer and G. Nicolson advocated a revised model in which proteins were dispersed and inserted into the phospholipid bilayer. According to this model, the membrane is a mosaic of protein molecules bobbing in a fluid bilayer of phospholipids. Freeze-fracture is a method of preparing cells for electron microscopy that splits a membrane along the middle of the phospholipid bilayer. When the halves are viewed, small bumps are visible, with protein particles interspersed in the matrix. Membranes Are Fluid Membranes are held together by hydrophobic interactions, which are much weaker than covalent bonds. As a result, lipids and proteins can move around in the membrane. Some membrane proteins drift, while others move along with more purpose. Furthermore, other membrane proteins seem to be held immobile by their attachments to the cytoskeleton.
2 UNIT TWO: THE CELL Chapter Eight: Membrane Structure and Function (Text from Biology, 6th Edition, by Campbell and Reece) A membrane remains fluid as temperature decreases until the phospholipids settle into a closely packed arrangement and the membrane solidifies. The steroid cholesterol, wedged between phospholipid molecules in the plasma membranes of animal cells, has different effects on membrane fluidity. At human body temperature, it restrains phospholipid movement. However, it also lowers the temperature required for solidification. Membranes must be fluid to work properly. Membranes Are Mosaics of Structure and Function There are two major populations of membrane proteins. Integral proteins penetrate the hydrophobic core of the lipid bilayer. Many of them are transmembrane proteins, which means they completely cross the membrane. The hydrophobic regions of an integral protein consist of one or more stretches of nonpolar amino acids, usually coiled into α helices. The hydrophilic parts are exposed to the aqueous solution on either side of the membrane. Peripheral proteins are not embedded in the lipid bilayer, but are loosely bound to the surface of the membrane. Often, they are bound to the exposed parts of integral proteins. Some membrane proteins on the cytoplasmic side of the plasma membrane are held in place by attachment to the cytoskeleton. On the exterior side, certain membrane proteins are attached to fibers of the ECM. These attachments combine to give animal cells a strong external framework. Membranes have distinct inside and outside faces. The two lipid layers may differ in specific lipid composition, and each protein has directional orientation in the membrane. Carbohydrates are only found on the exterior surface. Membrane Carbohydrates Are Important For Cell-Cell Recognition Cells can recognize other cells by keying on surface molecules, often carbohydrates, on the plasma membrane. Membrane carbohydrates are usually branched oligosaccharides with fewer than 15 sugar units. Some of these oligosaccharides are covalently bonded to lipids, which forms molecules called glycolipids. Most are covalently bonded to proteins (glycoproteins). Oligosaccharides on the external side of the plasma membrane vary from species to species, individuals of the same species, and even from one cell type to another.
3 UNIT TWO: THE CELL Chapter Eight: Membrane Structure and Function (Text from Biology, 6th Edition, by Campbell and Reece) TRAFFIC ACROSS MEMBRANES A Membrane’s Molecular Organization Results in Selective Permeability
The Role of Membrane Proteins
Proteins that span the membrane can provide a hydrophilic channel for certain solutes. Some transport proteins hydrolyze ATP so they can pump substances across the membrane even if it is against the concentration gradient. Proteins built into the membrane may be enzymes with the active site exposed to substances in the adjacent solution. These proteins can be adjacent to each other and work as a metabolic pathway. Membrane proteins may have a binding site with a specific shape that fits the shape of the chemical messenger, such as a hormone. The signal may cause a change in the shape of the protein that relays the message to the inside of the cell. Membrane proteins can be hooked together to form junctions. Some glycoproteins serve as identification tags. Other proteins can be bonded to the cytoskeleton, which helps maintain cell shape.
Permeability of the Lipid Bilayer
Hydrophobic molecules can dissolve in the lipid bilayer and easily cross. However, ions and polar molecules, which are hydrophilic, have a harder time crossing.
Transport proteins that span the membrane can provide channels through which hydrophilic substances can cross. These transport proteins are highly specific. Passive Transport is Diffusion Across a Membrane Molecules have intrinsic kinetic energy called heat. One result of thermal motion is diff usion the diffusion usion, tendency for molecules to spread out into available space. Although molecules themselves move randomly, diffusion of a population may be directional. A substance will diffuse from where it is more concentrated to where it is less concentrated: substances diffuse down their concentration gradient This is a spontaneous process because it gradient. decreases the free energy in a system by increasing entropy (disorder). Diffusion of substances across a biological membrane is passive transport because the cell does not have to expend energy to make it happen. The concentration gradient itself is potential energy and drives diffusion.
4 UNIT TWO: THE CELL Chapter Eight: Membrane Structure and Function (Text from Biology, 6th Edition, by Campbell and Reece) Osmosis is the Passive Transport of Water In comparing two solutions of unequal solute concentration, the solution with higher concentration of solute is said to be hypertonic Conversely, the solution with lower solute concentration is hypotonic hypertonic. hypotonic. These terms only make sense used comparatively. Solutions of equal solute concentration are isotonic. isotonic The direction of osmosis the diffusion of water, is determined by a difference in total solute osmosis, concentration. Water moves from a hypotonic to a hypertonic solution. It is helpful to remember that were there is more solute, there is less water and where there is less solute, there is more water; thus, water moves from areas of less solute to more solute. When two solutions are isotonic, there is still movement, but it is at an equal rate, so there is no net movement. Cell Survival Depends on Balancing Water Uptake and Loss
Water Balance of Cells Without Walls
Cells placed in isotonic solutions are stable. When they are placed in a hypertonic environment, water will move out of the cell into the environment and the cell will crenate as a result. However, when placed into a hypotonic solution, water moves into the cell and may result in lysis (bursting). Animals living in hypertonic or hypotonic environments must have special adaptations for osmoregulation the control of osmoregulation, water balance. Paramecium have a contractile vacuole that pumps out excess water.
Water Balance of Cells with Walls
Cells with cell walls cannot lyse, because the elastic wall only expands so much before it exerts a back pressure that prevents further water uptake. Plants are healthy in a hypotonic solution, which results in turgidity When plant cells are placed in an isotonic solution, they become flaccid and wilt. turgidity. In an hypertonic environment, plant cell membranes will shrink away from the wall in plasmolysis plasmolysis, which usually lethal. Specific Proteins Facilitate the Passive Transport of Water and Selected Solutes: A Closer Look In facilitated diffusion polar molecules and ions pass through the lipid bilayer with the help of diffusion, transport proteins. These proteins are specialized for the solute it transports and may even have a specific binding site similar to the active site of enzymes. Transport proteins can also be saturated, when they are transporting passengers at maximum rate. They can also be inhibited by molecules that
5 UNIT TWO: THE CELL Chapter Eight: Membrane Structure and Function (Text from Biology, 6th Edition, by Campbell and Reece) resemble the normal passenger and binds to the protein. Unlike enzymes, proteins catalyze a physical
process. Channel proteins allow water molecules or small ions to flow quickly from one side of the membrane to
the other. Water channel proteins are aquaporins Some channel proteins are gated channels stimuli aquaporins. channels; causes the gate to pen or close. Other proteins can change shape to translocate the solute-binding site across the membrane. Active Transport is the Pumping of Solutes Against Their Gradient Active transport requires work, since a molecule is being pumped across a membrane against its gradient. This is a major factor in the ability of a cell to maintain internal concentrations of small molecules that differ from concentrations in its environment. ATP supplies the energy for most active transport. This can occur through phosphorylation. In the sodium - potassium pump sodium ions are sodiumpump, exchanged for potassium ions. Every cycle of the pup moves three sodium ions out of the cell and brings two potassium ions in. Some Ion Pumps Generate Voltage Across Membranes Voltage is electrical potential energy. The cytoplasm of a cell is negative in charge compared to the extracellular fluid because of an unequal distribution of anions and cations on opposite sides of the membrane. The membrane potential voltage across a membrane, ranges from about -50 to -200 potential, millivolts. The membrane potential acts as an energy source that affects traffic of charges substances. Since the inside of the cell is negative compared to the outside, the membrane favors passage of cations inside the cell and anions outside the cell. Thus, ions are driven by a chemical force (concentration gradient) and electrical force. This combination is the electrochemical gradient An ion diffuses down its gradient. electrochemical gradient. With each cycle of the sodium-potassium pump, there is a net transfer of one positive charge from the cytoplasm to the extracellular fluid. A transport protein that stores energy in the form of voltage is an pump. pump, electrogenic pump The main electrogenic pump of plants, bacteria, and fungi is a proton pump which actively transports hydrogen ions (protons) out of the cell. This transfers positive charge from the cytoplasm to the extracellular solution. In Cotransport, a Membrane Protein Couples the Transport of Two Solutes A single ATP-powered pump that transports a specific solute can indirectly drive the active transport to several other solutes in a mechanism called cotransport A substance that has been cotransport. pumped across a membrane can do work as it leaks back by
6 UNIT TWO: THE CELL Chapter Eight: Membrane Structure and Function (Text from Biology, 6th Edition, by Campbell and Reece) diffusion. This is like pumping water downhill and then having it power a windmill as it flows downward. Another specialized transport protein can couple the “downhill” diffusion to the “uphill” transport against its own concentration gradient. For example, the gradient of hydrogen ions generated by proton pumps can be used to drive the active transport of amino acids, sugars, and several other nutrients into the cell. The protein can move sucrose into the cell only if the sucrose is accompanied by a hydrogen ion. Thus, the sucrose will follow the hydrogen ion into a transport protein. Exocytosis and Endocytosis Transport Large Molecules Larger molecules must travel through the cell membrane via vesicles. Exocytosis is the secretion of molecules by sending vesicles outwards to the plasma membrane. A transport vesicle can bud off the Golgi apparatus then merge with the plasma membrane. Exocytosis is used to export products out of a cell. In endocytosis the cell takes in macromolecules and particulate matter by forming vesicles from the endocytosis, plasma membrane. There are three types of endocytosis. In phagocytosis a cell engulfs particle by phagocytosis, wrapping pseudopodia around it and packaging it into a vacuole. This particle can then be digested when fused with a lysosome. In pinocytosis droplets of extracellular fluid are put into tiny vesicles. pinocytosis, ReceptorReceptor-mediated endocytosis is very specific. Proteins with specific receptor sites are exposed to the extracellular fluid. Ligands are any molecule that binds specifically to the receptor site of another molecule. The receptor proteins are usually clustered in regions called coated pits, which are lined by a layer of protein. When ligands bind to receptor proteins, a vesicle will break off and transport materials into the cell. Cells can acquire bulk quantities of specific substances through this form of endocytosis. For example, cholesterol is taken into the cell through this process.