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1.

- Diffusion
Diffusion is the movement of molecules via random motion from areas of high concentration to areas of low concentration.
KEY POINTS

Molecules move around rapidly, generating thermal energy. This causes them to diffuse randomly throughout a space. Molecules move randomly but may move directionally as a group across a concentration gradient. Dynamic equilibrium is achieved across a membrane when movement of molecules as a population are not directional - molecules move in both directions at the same rate. Diffusion always occurs from higher concentrated areas to lower concentrated areas of a molecule, down the "concentration gradient." Substances diffuse based on the concentration of their own molecules in the solution and are not affected by the concentration of other molecules. Passive diffusion is spontaneous and requires no energy. Diffusion across cell membranes does not require cell energy. The rate of diffusion of different molecules is determined by the selectiveness of the cell membrane. When molecules move across a membrane in both directions at the same rate, dynamic equilibrium is achieved.

KEY TERMS

Dynamic Equilibrium An equilibrium in which two reversible reactions occur at the same rate. Concentration Gradient A concentration gradient is present when a membrane separates two different concentrations of molecules. Diffusion the movement of molecules from regions of high concentration (high water vapor pressure) toward regions of lower concentration.

EXAMPLES

Figure 1 depicts diffusion across a membrane. The membrane is highly selective for certain molecules to pass through it. Molecules then move through the membrane along a concentration gradient.

FIG. 1 Diffusion across a membrane

FULL TEXT The Process Of Diffusion


Diffusion is the movement of molecules by random motion from regions of higher concentration to regions of lower concentration. Diffusion of molecules across biological membranes occurs only for molecules that are highly permeable to the membrane, which provides a means for selective regulation.Diffusion across the plasma membrane does not require any energy. Molecules move around rapidly, generating thermal energy. This causes them to diffuse randomly throughout space. Over time, this process results in the equilibration of molecules throughout space. When a concentration gradient is present,diffusion results in the movement of molecules directionally across the concentration gradient, provided that the membrane is permeable to the molecule. Dynamic equilibrium is achieved across a membrane when movement of molecules is not directional (i.e., molecules move in both directions at the same time). Figure 1 illustrates the diffusionof water and dye molecules across a permeable membrane. Diffusion always occurs from higher concentrated areas to lower concentrated areas of a molecule, down the "concentration gradient." Thediffusion of any given molecule is determined by its own concentration and is not affected by the concentration of other molecules. Diffusion is spontaneous and requires no energy; diffusion across cell membranes does not require cell energy. Diffusion is a time dependent process. In the context of biological membranes, the rate of diffusion of different molecules is also governed by the selectiveness of the cell membrane. The more permeable a molecule is, the more rapidly it will diffuse when there is a concentration gradient. Due to the inherent composition of cell membranes, only nonpolar molecules, such as oxygen and carbon dioxide, are highly permeable to it and, therefore, can diffuse very rapidly from areas of high concentration to areas of low concentration. This feature is exploited by cells to uptake oxygen rapidly when increased cellular demands from respiration require immediate oxygen.Likewise, the rapid diffusion of oxygen and carbon dioxide across the alveolar-capillary membrane is fundamentally important for gas exchange in mammalian lungs.

2. Osmosis
Osmosis is the movement of water across a membrane from a low solute concentration to a high solute concentration to equilibrium.

KEY POINTS

Osmosis involves the movement of water across a semi-permeable membrane that does not have pores large enough to allow large molecules to pass. Like diffusion, osmosis requires no energy input. Osmotic (turgor) pressure is the pressure required to maintain an equilibrium, with no net movement of solvent. Turgot pressure is the main cause of support in many plants. Osmosis is also responsible for the ability of plant roots to draw water from the soil.

KEY TERMS

Osmosis The net movement of solvent molecules from a region of high solvent potential to a region of lower solvent potential through a partially permeable membrane. Turgor The pressure produced by a solution in a space that is enclosed by a differentially permeable membrane. Osmosis the net movement of solvent molecules from a region of high solvent potential to a region of lower solvent potential through a partially permeable membrane.


Fig. 1 Water balance in animal and plant cells. Fig. 2 Water balance in animal and plant cells.


Fig. 3 Contractile vacuole of a paramecium.

FULL TEXT

Semi-permeable membranes, such as those in cells, do not have pores large enough to allow large molecules, such as sugar, to pass through. Osmosis is the movement of water across the membrane, driven by differences in the concentration of solutes (such as glucose) on either side of the membrane.The differential concentration of solutes results in different concentrations of free water molecules of either side of the membrane. Consequently, water traverses from the side with lower solute concentration to that of the higher solute concentration until the solute concentrations are equal. Osmosis is important for cellular survival in non-isotonic environments. In plants,osmosis provides for plant rigidity via turgor pressure. Osmosis is the movement of solvent molecules through a selectively permeable membrane into a region of higher solute concentration, aiming to equalize the solute concentrations on the two sides ([[fig:7888]]). It is used to describe a physical process in which any solvent moves, without input of energy, across a semipermeable membrane (permeable to the solvent but not the solute) separating two solutions of different concentrations.


FIGURE 3Contractile vacuole of a paramecium

Net movement of solvent takes place from the less concentrated (hypotonic) to the more concentrated (hypertonic) solution, which tends to reduce the difference in concentrations between the two compartments (Figure 3). This effect can be countered by increasing the pressure of the hypertonic solution, with respect to the hypotonic.The osmotic pressure is defined to be the pressure required to maintain an equilibrium, with no net movement of solvent. Osmotic pressure depends on the molar concentration of the solute but not on its identity. Organisms contain many biological membranes, which act as semipermeable membranes, permitting osmosis. In general, these membranes are impermeable to organic solutes with large molecules, such as polysaccharides, while permeable to water and small, uncharged solutes [[fig:394]]. Permeability may depend on solubility properties, charge, or chemistry, as well as solute size. Water molecules travel through the plasma cell wall, tonoplast (vacuole), or protoplast in two ways, either by diffusing across the phospholipid bilayer directly or via aquaporins (small transmembrane proteins which form water-permeables pores). Osmosis provides the primary means by which water is transported into and out of cells. Osmosis is the main cause of support in many plants. The osmotic entry of water raises the turgor pressure exerted against the cell wall, until it equals the osmotic pressure, creating a steady state. When a plant cell is placed in a hypertonic solution, the water in the cells moves to an area higher in solute concentration and the cell shrinks and, in doing so, becomes flaccid. This means the cell has become

plasmolysedthe cell membrane has completely left the cell wall due to lack of water pressure on it, the opposite of turgid.Also, osmosis is responsible for the ability of plant roots to draw water from the soil. Since there are many fine roots, they have a large surface area, and water enters the roots by osmosis. In unusual environments, osmosis can be very harmful to organisms. For example, freshwater and saltwater aquarium fish, placed in water of a different salinity than that to which they are adapted to, will quickly die.


FIGURE 2Water balance in animal and plant cells

If an animal or a plant cell is placed in a solution of sugar or salt in water and if the medium is hypotonic (a dilute solution with a higher water concentration than the cell), the cell will gain water through osmosis (Figure 2). If the medium is isotonic (a solution with exactly the same water concentration as the cell), there will be no net movement of water across the cell membrane. If the medium is hypertonic (a concentrated solution with a lower water concentration than the cell), the cell will lose water by osmosis. Essentially, this means that if a cell is put in a solution which has a solute concentration higher than its own, then it will shrivel up, and if it is put in a solution with a lesser solute concentration than its own, the cell will expand and burst.


Osmostic pressure (also named Turgor) represents the force per unit area (pressure) required to prevent the passage of water through a semipermeable (selective) membrane, into a solution of higher concentration. Osmotic pressure is thus a measure of the tendency of water to move into one solution from another by osmosis. Osmotic pressure is a colligative property, meaning that the property depends on the concentration of the solute but not on its identity. Osmotic gradient is the difference in concentration between two solutions on either side of a semipermeable membrane.

3. Water Balance
A solution's tonicity relative to a cell will determine whether the cell will gain or lose water when immersed in that solution.

KEY POINTS

Animal cells are stable in isotonic environments: the solute concentrations inside and outside are the same and there is no net movement of water. Animal cells will gain water, swell and possibly burst in hypotonic environments: the solute concentrations inside the cell are higher than outside and the net movement of water is into the cell. In hypertonic and hypotonic environments, cells must be adapted to osmoregulate to prevent shrinking or swelling. Cells with cell walls (plants, prokaryotes, fungi, some protists) function differently in response to tonicity than animal cells. Cells with cell walls function best in hypotonic environments. The cell swells with water, but the rigid cell wall exerts turgor pressure on the inside of the cell preventing it from taking up too much water and bursting. There are three classifications of tonicness that one solution can have relative to another: hypertonic, hypotonic, and isotonic.

KEY TERMS

Plasmolysis The shrinking of protoplasm away from the cell wall of a plant or bacterium due to water loss. Osmoregulation The homeostatic regulation of osmotic pressure in the body in order to maintain a constant water content. Hypotonic Having a lower osmotic pressure than another. Tonicity the ability of a solution to exert an osmotic pressure upon a membrane

EXAMPLES

Figure 1 depicts red blood cells in three different environments. In hypertonic conditions, the cells shrink as water diffuses out of the cell while in hypotonic conditions water diffuses into the cells causing them to swell. In isotonic conditions, there is no net movement of water.

Fig. 1 Water balance in animal and plant cells.

Fig. 2 Water balance in animal and plant cells. Fig. 3 Contractile vacuole of a paramecium.

FULL TEXT
Tonicity is a measure of the osmotic pressure of two solutions (as defined by the water potential of each) separated by a semipermeable membrane. There are three classifications of tonicity that one solution can have relative to another: hypertonic, hypotonic and isotonic. Isotonicity is condition or property of a solution in which its solute concentration is the same as the solute concentration of another solution with which it is compared.

Hypertonic Solution
Hypertonic solution has a greater solute concentration than the cytosol. When a cells cytoplasm is bathed in a hypertonic solution, the water will be drawn into the solution and out of the cell by osmosis. If water molecules continue to diffuse out of the cell, the cell will crenate (shrink). A hypertonic solution is used in osmotherapy to treat cerebral hemorrhage. If a plant cell is placed in a hypertonic solution, the plant cell loses water and hence turgor pressure (see below), making the plant cell flaccid. Plants with cells in this condition wilt. Further water loss causes plasmolysis.

Hypotonic Solution And Turgor Pressure


Hypotonic solution is a solution having a lesser solute concentration than the cytosol. When a cells cytoplasm is bathed in a hypotonic solution, the water will be drawn out of the solution and into the cell by osmosis. If water molecules continue to diffuse into the cell, the cell will swell to the point that cytolysis (rupture) may occur. In plant cells, the cell will not always rupture.When placed in a hypotonic solution, the cell will achieve Turgor Pressurethe main pressure of the cell content against the cell wall in plant cells and bacteria cells, as determined by the water content of the vacuole, resulting from osmotic pressure. The cell then proceeds with its normal functions. Plant cells tend to resist bursting, however, due to the reinforcement of their cell wall, which provides effective osmolarity or osmolality. In some cases of suspensions intended for intramuscular injection, a slightlyhypotonic solution is preferred in order to increase the dissolution and absorption of the drug by absorbing water from the surrounding tissues. A plant cell in a more hypotonic solution will absorb water by endosmosis, so that the increased volume of water in the cell, will increase pressure, making the protoplasm push against the cell wall. Plant cell walls resist further water entry after a certain point, known as full turgor.

Osmotic Pressure And Osmoregulation

FIGURE 3

Contractile vacuole of a paramecium

In hyper- or hypotonic environments, cells must adapt to prevent shrinkage or swelling. This is done by osmoregulation mechanisms. As an example, paramecium constantly pump water out of the cell to maintain water balance (Figure 3). Some organisms have evolved intricate methods of circumventing hypertonicity. For example, saltwater is hypertonic to the fish that lives in it. The fish need a large surface area in their gills in contact with seawater for gas exchange, resulting in osmotic water loss from gill cells to the seawater. The fish respond to the loss of water by drinking large amounts of saltwater, and actively excreting the excess salt. This process is called osmoregulation.Likewise, fresh water fish urinate constantly to prevent cytolysis.

FIGURE 1Water

balance in animal and plant cells

For the effect of osmosis, osmotic pressure and tonicity on cells, see Figure 1. Cells with cell walls (plants, fugi, some protests, and prokaryotes) respond to osmotic pressure in a different way than cells without cell walls, such as that from animals. Cells with cell walls give rigidity to the plant and have a higher resistance in hypotonic environments, where the cell wall can prevent cellular bursting. In eukaryotic animal cells, a hypertonic environment forces water to leave the cell and the shape of the cell becomes distorted and wrinkled, a state known as crenation. In plant cells, the effect is more dramatic. The flexible cell membrane pulls away from the rigid cell wall but remains joined to the cell wall at points called plasmodesmata. The cell takes on the appearance of a pincushion. Plasmolysis is the process in plant cells where the plasma membrane pulls away from the cell wall due to the loss of water through osmosis. The reverse process, cytolysis, can occur if the cell is in a hypotonic solution, resulting in a higher external osmotic pressure and a net flow of water into the cell.Through observation of plasmolysis and deplasmolysis, it is possible to determine the tonicity of the cell's environment, as well as the rate solute molecules cross the cellular membrane. The terms "isotonic", "hypotonic", and "hypertonic" cannot always be applied strictly, or accurately, given that the pressure exerted by the cell wall significantly affects the osmotic equilibrium point.

4. Facilitated diffusion
Facilitated diffusion is spontaneous passage of molecules or ions across a biological membrane passing through specific transmembrane integral proteins.

KEY POINTS

Channel proteins are hydrophilic corridors for specific molecules to diffuse into and out of the cell. Aquaporins are channel proteins that allow for rapid diffusion of water into and out of cells. Ion channels are integral membrane proteins that allow the passage of charged particles (ions) into and out of the cell. Sometimes, they are gated channels and open or close to control the movement of the ions. Carrier proteins change shape in response to the interception of the molecule to be transported. The shape change causes the molecule to be released on the other side. Neither channels nor carrier proteins require energy to facilitate transport and molecules move down their concentration gradients.

KEY TERMS

Gated Channels Gated channels are ion channels that can be opened or closed by chemical or electrical signal in order to control the movement of the ions. Aquaporins Aquaporins are proteins embedded in the cell membrane that regulate the flow of water. Carrier Protein Any protein whose function is to transport small molecules (or other proteins) through biological membranes

Fig. 1 Facilitated Diffusion.

FULL TEXT

Facilitated Diffusion

FIGURE 1Facilitated

Diffusion

Facilitated diffusion is the spontaneous passage of molecules or ions across a biological membrane passing through specific transmembrane integral proteins. Facilitated diffusion is not a type of diffusion (thermal motion) but a type of transport process that is passive; it does not require any energy input. The molecules that are transported via facilitated diffusion are not permeable to the plasma membrane. They include ions (highly charged), as well as numerous nonpolar molecules that are poorly soluble in water. Facilitated diffusion is mediated by protein channels andcarrier proteins (Figure 1). Most transport proteins that mediate facilitated diffusion are very selective and only transport certain molecules. The major classes of proteins involved in facilitated diffusion are aquaporins, ion channels, and carrier proteins. Importantly, neither channels nor carrier proteins require energy to facilitate the transport of molecules; they enable molecules to move down their concentration gradients by either providing an exclusive corridor in which to pass through (a pore with specificity for the type of molecule to be transported) or via specific binding sites on carrier proteins.

AQUAPORINS
Aquaporins are proteins embedded in the cell membrane that regulate the flow of water. They are integral membrane proteins which form pores in the membrane of biological cells. Genetic defects involving aquaporin genes have been associated with several human diseases. Aquaporins selectively conduct water molecules in and out of the cell, while preventing the passage of ions and other solutes. Also known as water channels, aquaporins are integral membrane pore proteins. Some of them, known as "aquaglyceroporins", also transport other small uncharged solutes, such as glycerol, CO2, ammonia and urea across the membrane, depending on the size of the pore. However, the water pores are completely impermeable to charged species, such as protons, a property critical for the conservation of the membrane's electrochemical potential. Water molecules traverse the pore of the channel in single file. The presence of water channels increases membrane permeability to water. Many human cell types express aquaporins; red blood cells and kidney cells express them at high levels. They are also expressed in certain bacteria and many other organisms, such as plants for which it is essential for the water transport system.

ION CHANNELS
Ion channels are integral membrane proteins or, more typically, an assembly of several proteins that transport ions. They are present on all membranes of cell (plasma membrane) and intracellular organelles (nucleus, mitochondria, endoplasmic reticulum, golgi apparatus, and so on). Channels differ with respect to the ion they let pass (e.g., Na+, K+, Cl), the ways in which they may be regulated, the number of subunits of which they are composed, and other aspects of structure. Channels belonging to the largest class, which includes the voltage-gated channels that underlie the nerve impulse, consists of four subunits with six transmembrane helices each. On activation, these helices move about and open the pore, forming a hydrophilic corridor for ions to pass through. There are over 300 types of ion channels in a living cell. Ion channels may be classified by the nature of their gating, the species of ions passing through those gates, the number of gates (pores), and localization of proteins. Further heterogeneity of ion channels arises when channels with different constitutive subunits give rise to a specific kind of current. Some ion channels require an electrical stimulus. Because "voltage-activated" channels underlie the nerve impulse and because "transmitter-activated" channels mediate conduction across the synapses, channels are especially prominent components of the nervous system. Indeed, most of the offensive and defensive toxins that organisms have evolved for shutting down the nervous systems of predators and prey (e.g., the venoms produced by spiders, scorpions, snakes, fish, bees, sea snails, and others) work by modulating ion channel conductance and/or kinetics. Other channels respond to different stimuli. For example, mechanosensitive ion channels are opening under the influence of stretch, pressure, shear, and displacement. Cyclic nucleotide-gated channels are characterized by activation due to the binding of intracellular cAMP or cGMP, with specificity varying by member. These channels are primarily permeable to monovalent cations, such as K+ and Na+. `Carrier proteins are proteins involved in the movement of ions, small molecules, or macromolecules, such as another protein, across a biological membrane.Carrier proteins are integral membrane proteins; that is they exist within and span the membrane across which they transport substances. Each carrier protein, even within the same cell membrane, is specific to one type or family of molecules. For example, GLUT1 is a named carrier protein found in almost all animal cell membranes that transports glucose across the bilayer. Other specific carrier proteins also help the body function in important ways.Hemoglobin, a carrier protein, transports oxygen through the blood.Cytochromes operate in the electron transport chain as carrier proteins for electrons.

5. Energy use in active transport


Active transport allows cells to move molecules against a concentration gradient using energy.

KEY POINTS

Cells expend energy to transport molecules against the concentration gradient using carrier proteins. Active transport allows cells to maintain non-equilibrial concentrations of solutes inside and outside of the cell. ATP powers active transport, sometimes by transferring a phosphate group to the protein, which induces a shape change. The sodium-potassium pump in animal cells requires ATP.

KEY TERMS

Sodium-Potassium Pump An enzyme (an electrogenic transmembrane ATPase) located in the plasma membrane of all animal cells. Active Transport Active transport is the movement of a substance across a cell membrane against its concentration gradient (from low to high concentration). In all cells, this is usually concerned with accumulating high concentrations of molecules that the cell needs, such as ions, glucose and amino acids. Active Transport the movement of a substance across a cell membrane against its concentration gradient Active Site the small portion of an enzyme where substrate molecules bind and undergo a chemical reaction

EXAMPLES

The sodium-potassium pump also regulates cell volume. Many proteins inside a cell are negatively charged and this attracts positively charged molecules. These interactions cause the osmosis of water into the cell. When a cell starts to swell, the sodium-potassium pump begins moving sodium ions outside the cell and in turn, causes water to flow out of the cell.

FIGURES

FIG. 1 The sodium-potassium pump

FULL TEXT Active transport is the movement of molecules up their concentration gradient, which requires the expenditure of energy. It is mediated only by carrier proteins and is fundamentally important to maintain concentration gradients across plasma membranes.

FIGURE 1The

sodium-potassium pump

The active transport represents the movement of molecules against the concentration gradient.The active transport maintains the concentration gradient between the extracellular and cytoplasmic compartments, requires energy consumption, and is accomplished by carrier proteins. For example, in animals cells, K+ and Na+ ions have steep concentration gradients across the cell membrane.Figure 1 shows a schematic of K+ gradients in a plasma membrane. ABC transporters are transmembrane proteins that utilize the energy of ATP hydrolysis to function. ABC transporters are the largest family of transporters of various substrates across membranes, and also play a role in critical non-transport related functions, such as translation of RNA and/or DNA repair. They transport a large variety of substrates across membranes; molecules such as metabolic products, lipids, sterols, and drugs. Genes of various members of the ATP binding cassette transporters are abundant in the genomes of all vertebrates. These members are associated with human diseases, involving almost all organ systems of the human body, and at least some of them play an important role in cancer resistance to therapy, bacterial resistance and cystic fibrosis. Na+/K+ ATPase is also known as the Na+/K+ pump, sodium-potassium pump, or sodium pump, and it is an enzyme and active carrier located in the plasma membrane (to be specific, an electrogenic transmembrane ATPase) in all animals. Active transport is responsible for cells containing relatively high concentrations of potassium ions, but low concentrations of sodium ions. The mechanism responsible for this is the sodium-potassium pump, which moves these two ions in opposite directions across the plasma membrane. This was investigated by following the passage of radioactively labeled ions across the plasma membrane of certain cells. It was found that the concentrations of sodium and potassium ions on the two other sides of the membrane are interdependent, suggesting that the same carrier transports both ions. It is now known that the carrier is an ATP-ase, and that it pumps three sodium ions out of the cell for every two potassium ions pumped in.

THE SODIUM-POTASSIUM PUMP


The sodium-potassium pump was discovered in the 1950s by a Danish scientist, Jens Christian Skou, who was awarded a Nobel Prize in 1997. It marked an important step forward in our understanding of how ions get into and out of cells, and it has a particular significance for excitable cells, such as nervous cells, which depend on it for responding to stimuli and transmitting impulses. Export of sodium from the cell provides the driving force for several secondary active transporters: membrane transport proteins, which import glucose, amino acids, and other nutrients into the cell by use of the sodium gradient. Another important task of the Na+-K+ pump is to provide a Na+ gradient that is used by certain carrier processes. In the gut, for example, sodium is transported out of the reabsorbing cell on the blood (interstitial fluid) side via the Na+-K+ pump, whereas, on the reabsorbing (luminal) side, the Na+-Glucose symporter uses the created Na+ gradient as a source of energy to import both Na+ and glucose, which is far more efficient than simple diffusion. Similar processes are located in the renal tubular system. The pump, with bound ATP, binds three intracellular Na+ ions. ATP is hydrolyzed, leading to phosphorylation of the pump at a highly conserved aspartate residue and subsequent release of ADP. A conformational change in the pump exposes the Na+ ions to the outside. The phosphorylated form of the pump has a low affinity for Na+ ions, so they are released. The pump binds two extracellular K+ ions. This causes the dephosphorylation of the pump, reverting it to its previous conformational state, transporting the K+ ions into the cell. The unphosphorylated form of the pump has a higher affinity for Na+ ions than K+ ions, so the two bound K+ ions are released.ATP binds, and the process starts again.

6. Ion pumps maintain membrane potential


The membrane potential is the difference in voltage between the exterior and interior of a cell.

KEY POINTS

Cells have voltages across plasma membranes created by the separation of opposite charges, with the cytoplasmic side negatively charged and ranging from -50 to - 200mV. The membrane potential forces the transport of ions across the membrane since they are attracted to the oppositely charged area. The electrochemical gradient summarizes the two forces acting on transmembrane transport: ionic concentration and membrane potential. In passive transport, ions diffuse down BOTH the ionic concentration and the electrochemical gradient. Active transport also contributes to the membrane potential. The sodium-potassium pump is the major electrogenic pump of animal cells. Proton gradients are used to carry out cellular functions like synthesizing ATP.

KEY TERMS

Electrochemical Gradient An electrochemical gradient is a gradient of electrochemical potential, usually for an ion that can move across membrane. The gradient consist of two parts, the electrical potential and a difference in the chemical concentration across a membrane. Action Potential A short term change in the electrical potential that travels along a cell such as a nerve or muscle fiber.
EXAMPLES Certain toxins can block ion and voltage gated channels. Venom from scorpions has been found to block certain potassium channels in neurons. Moreover, snake venom contains toxins that block voltage-gated potassium channels in motor neurons.

FIGURES

1.

FIG. 1 Proton

pump and membrane potentials

2.

FIG. 2 Facilitated

Transport

3.

FIG. 3 The

sodium-potassium pump

FULL TEXT

The Function Of Ion Pumps


Cells have differences in concentrations of ions on either side of the plasma membrane that are maintained by both active and passive transport processes. Ion channels and ion pumps are the major classes of proteins that control the transport of ions across cell membranes. These ionic differentials are exploited as a means to store energy to carry out numerous cellular processes, including the synthesis of ATP. Membrane potential is the difference in voltage (or electrochemical difference) between the interior and exterior of a cell (Figure 1). The membrane potential arises from the net actions of ion channels and ion pumps embedded in the membrane, which produces different concentrations of ions (and therefore electrical charge) on the intracellular and extracellular sides of the membrane. This creates an electrochemical gradient. The ion channels, when active, partially discharge the membrane potential, while the ion pumps restore and maintain it. The membrane potential that is maintained in the absence of fluctuations (e.g., action potentials) is called the "resting membrane potential". The ions that contribute the most significantly to membrane potential are sodium (Na+) and chloride (Cl) ions which have high concentrations in the extracellular region, and potassium (K+) ions that have high concentrations in the intracellular region. In a typical cell, the cytoplasmic side of the plasma membrane is negatively charged, ranging from -50 to -200 mV. The membrane potential is held at a relatively stable value, called the resting potential. For many cells, typical values of the resting potential range from 70 to 80 millivolts; that is, the interior of a cell has a negative baseline voltage of a bit less than one-tenth of a volt. Opening and closing of ion channels can induce a departure from the resting potential, called adepolarization. if the interior voltage rises (say from 70 mV to 65 mV), or ahyperpolarization, if the interior voltage becomes more negative (e.g., changing from 70 mV to 80 mV). In excitable cells, a sufficiently large depolarizations can evoke a short-lasting all-or-nothing event called an action potential, in which the membrane potential

very rapidly undergoes a large change, often briefly reversing its sign. Action potentials are generated by special types of voltage-dependent ion channels. An electrochemical gradient has two components that act on transmembrane transport: ionic concentration and membrane potential.Indeed, the electrical component is caused by a charge difference across the lipid membrane. The chemical component is caused by a differential concentration of ions across the membrane. The combination of these two factors determines the thermodynamically favorable direction for an ion's movement across a membrane. In biological membranes, the direction of an ion moves by passive transport (i.e., diffusion) or active transport across a membrane is determined by the electrochemical gradient. In passive transport, ions diffuse both down in the ionic concentration and theelectrochemical gradient. In mitochondria and chloroplasts, proton gradients are used to generate a chemiosmotic potential that is also known as a proton motive force. This potential energy is used for the synthesis of ATP by oxidative phosphorylation. The transport of ions is required to generate membrane potentials. The two types of structures that play the largest roles are ion channels and ion pumps, both usually formed from assemblages of protein molecules. Ion channels form selective pores through which ions can move. In most cases an ion channel is only permeable to specific types of ions (e.g., sodium and potassium but not chloride or calcium) and, sometimes, the permeability varies depending on the direction of ion movement. Ion pumps, also known as "ion transporters" or "carrier proteins", actively transport specific types of ions from one side of the membrane to the other, sometimes, using energy derived from metabolic processes to do so. The other major contributor in establishing the membrane potential is the process of active transport that is mediated by a variety of pumps/exchangers. For example, the sodiumpotassium exchange pump is a complex of proteins embedded in the membrane that derives energy from ATP in order to transport sodium and potassium ions across the membrane.On each cycle, the pump exchanges three Na+ ions from the intracellular space for two K+ ions from the extracellular space. If the numbers of each type of ion were equal, the pump would be electrically neutral, but because of the three-for-two exchange, it gives a net movement of one positive charge from intracellular to extracellular for each cycle, thereby contributing to a positive voltage difference. The pump has three effects:

Makes the sodium concentration high in the extracellular space and low in the intracellular space (Figure 3). Makes the potassium concentration high in the intracellular space and low in the extracellular space Gives the extracellular space a positive voltage with respect to the intracellular space.

7. Coupled transport
Coupled transport is the simultaneous passive transport of molecules across the plasma membrane in a fixed ratio.

KEY POINTS

A protein pumping molecules across a membrane can indirectly power the active transport of other molecules. It does this by utilizing the energy created by the diffusion of the first molecule back down its concentration gradient. A cotransporter is an integral membrane protein that moves two or more different molecules across the plasma membrane at the same time. Coupled transport is the entry of a molecule into a cell against a concentration gradient powered by the free energy liberated by the movement of another molecule down its concentration gradient.

KEY TERMS

Co-Transport Co-transport, also known as coupled transport or secondary active transport, refers to the simultaneous or sequential passive transfer of molecules or ions across biological membranes in a fixed ratio.

EXAMPLES

The sodium/glucose cotransporter has led diabetes researchers to develop drugs that target this coupled transporter. For instance, phlorizin, a drug derived from apple tree bark, inhibits glucose transport in the kidneys and has been used in the past to treat patients with diabetes.

FIGURES

1.

FIG. 1

Coupled transport

Coupled transport
In coupled transport, the required energy is derived from energy stored in the form of concentration differences in a second solute. Typically, the concentration gradient of the second solute was created by primary active transport, and the diffusion of the second solute across the membrane drives coupled transport.

FULL TEXT Coupled transport is the entry of a molecule into a cell against a concentration gradient powered by the free energy liberated by the movement of another molecule down its concentration gradient. These reactions are mediated by integral membrane proteins that bind to both sets of molecules and couple the transport of each in and out of the cell. One of the most important examples of co-transport is the sodium-glucose co-transporter, which is responsible for glucose uptake by intestinal cells. The discovery of this mode of co-transport led to the development of oral rehydration therapy to treat chronic diarrhea. Coupled transport (also known as "co-transport" or "secondary active transport") is the simultaneous passive transport of molecules or ions across the plasma membrane in a fixed ratio with that of an active transport process. This type of transport is mediated by a cotransporter, an integral membrane protein that moves two or more different molecules or ions across a phospholipid membrane at the same time. It is sometimes referred to interchangeably with symporter (molecules going in the same direction across the membrane), but the term "cotransporter" refers both to symporters and antiporters (molecules going across the membrane in an opposing direction). In most forms of coupled transport, facilitated diffusion of an ion(s) down the electrochemical gradient is exploited to move another type of molecule(s) against a concentration gradient. It is the free energy stored in an electrochemical ion gradient (created by the diffusion of the first molecule down its concentration gradient) that powers the active transport of another substrate. In many higher plants, sucrose is the dominant migrant carbohydrate source. Proton- pump driven sucrose transport is a critically important means by which sucrose is distributed throughout the plant. In this method of transport, the proton pump creates a gradient of protons so that there are many more on one side of the membrane than another. As the protons diffuse back across the membrane, the free energy liberated by this diffusion is utilized to co-transport sucrose. In mammalian cells, sodium-dependent glucose transporters use the energy from a downhill sodium gradient to transport glucose across membrane. Note that these are also known as "symporters," since both sodium and glucose are transported in the same direction across the membrane. In the 1950's, Robert K. Crane hypothesized that glucose transport into the cell is the first critical step in glucose metabolism. In August 1960, in Prague, Robert K. Crane presented data in support of his seminal discovery that sodium- glucose cotransport is the mechanism that underlies intestinal glucose absorption. Crane's discovery of cotransport was the first ever demonstration of this fundamentally key mechanism of transport in biology. It is viewed by some to be the most important discovery of carbohydrate absorption in the 20th century. Cranes discovery of cotransport provided the basis for the treatment of severe diarrhea, and led directly to the development of oral rehydration therapy. This treatment counter balances the loss of water and electrolytes caused by cholera via a glucose containing salt solution that accelerates water and electrolyte absorption. This is possible because cholera does not interfere with sodium-glucose cotransport. Oral rehydration therapy has saved the lives of millions of cholera patients in underdeveloped countries since the 1980's. In 1978, The Lancet wrote, "The discovery that sodium transport and glucose transport are coupled in the small intestine, so that glucose accelerates absorption of solute and water, was potentially the most important medical advance this century." Cranes discovery is also used in blockbuster drugs, such as the SSRI Prozax, which treats depression by inhibiting the Na/serotonin cotransporters in the brain. Furthermore, major pharmaceutical companies are developing inhibitors of the Na/glucose cotransporters to treat diabetes and obesity.

8. Endocytosis and Exocytosis


Exocytosis and endocytosis are processes by which material is encapsulated by a vesicle and either secreted by a cell or brought into a cell.

KEY POINTS

Transport vesicles for exporting molecules originate in the golgi complex and are moved along microtubules through the cytosol. In exocytosis, the transport vesicle lipid bilayer merges with the plasma membrane and the contents of the vesicle are released to the outside. Secretory cells use exocytosis to export molecules. In endocytosis, the transport vesicles are made from the plasma membrane surrounding molecules to be imported. There are three types of endocytosis: phagocytosis, pinocytosis, and receptor- mediated endocytosis. Endocytosis and exocytosis also act in the continual replacement of the cell membrane.

KEY TERMS

Pinocytosis A form of endocytosis in which material enters a cell through its membrane and is incorporated in vesicles for digestion. Phagocytosis The process where a cell incorporates a particle by extending pseudopodia and drawing the particle into a vacuole of its cytoplasm. Exocytosis the secretion of substances through cellular membranes, either to excrete waste products or as a regulatory function Endocytosis the process by which the plasma membrane of a cell folds inwards to ingest material Vesicle a membrane-bound compartment found in a cell

EXAMPLES

Another example of receptor-mediated endocytosis is with the EGF receptor. The EGF:EGF-R complex is brought in via clathrin-coated pits and then fused with endosomes. This internalization of the receptor creates a feedback loop on EGF signaling.

FIG. 1

The three types of endocytosis

FULL TEXT

Exocytosis and endocytosis are reciprocal processes by which material is either secreted by a cell (exocytosis) or is brought into a cell (endocytosis) without passing through the plasma membrane. In each case, vesicles are formed that encapsulate the material to be transported. In exocytosis, vesicles fuse with the cell membrane, which increases the surface area of the plasma membrane. In endocytosis, a part of the plasma membrane pinches off, decreasing the surface area of the cell membrane. All cells undergo these processes, which are fundamental for shuttling molecules in and out of cells in a highly coordinated manner. Exocytosis and endocytosis also work in concert to continually replace the plasma membrane which is required for the regulation of cell size during cell growth.

Exocytosis
Exocytosis is the secretion of cellular molecules generated inside the cell into the extracellular space. This process is fundamental to many diverse biological functions, such as secretion of preformed inflammatory mediators in allergic reactions, the secretion of insulin into the bloodstream by pancreatic islet cells, the secretion of neurotransmitters by neurons, and the export of cell wall components by plant cells. Exocytosis is a highly coordinated process that originates in the golgi complex where, in the molecules to be exported transport vesicles are contained. These vesicles are moved along microtubules through the cytosol. When the vesicles reach the plasma membrane, the vesicle lipid bi- layer merges with the plasma membrane and the contents of the vesicle are released to the outside.

Endocytosis
Endocytosis is the process by which a substance enters into a cell without passing through the plasma membrane. Transport vesicles are made from the plasma membrane surrounding the molecules to be imported. The material to be taken into the cells is first enclosed in a portion of the cell membrane that invaginates, forming a vesicle containing the material that is released into the cytoplasm. Together with exocytosis, endocytosis is fundamentally important in the maintenance and replacement of the cell membrane components. Endocytosis is currently divided in four categories (Figure 1):

1. Receptor Mediated endocytosis (clathrin dependent endocytosis) 2. Caveole 3. Pinocytosis 4. Phagocytosis The major route for endocytosis in most cells - and the best-understood - is that mediated by the molecule clathrin. This large protein assists in the formation of a coated pit on the inner surface of the plasma membrane of the cell. This pit then buds into the cell to form a coated vesicle in the cytoplasm of the cell. In doing so, it brings into the cell not only a small area of the surface of the cell but also a small volume of fluid from outside the cell.

Pinocytosis
Pinocytosis is a form of endocytosis in which small particles are brought into the cell suspended within small vesicles that subsequently fuse with lysosomes to hydrolyze, or to break down, the particles. It usually occurs within highly ruffled regions of the plasma membrane. It is this form of endocytosis, invagination of the cell membrane forming a pocket, which then pinches off into the cell to form a vesicle (0.55 m in diameter) filled with large volume of extracellular fluid and molecules (equivalent to approximately 100 Clathrin-coated vesicles). The filling of the pocket occurs in a non-specific manner. The vesicle then travels into the cytosol and fuses with other vesicles such as endosomes and lysosomes.

Phagocytosis
Phagocytosis is a specific form of endocytosis involving the vesicular internalization of solid particles, such as bacteria. It is, therefore, distinct from other forms of endocytosis, such as the vesicular internalization of various liquids. Phagocytosis is involved in the acquisition of nutrients for some cells, and, in the immune system, it is a major mechanism used to remove pathogens and cell debris. Bacteria, dead tissue cells, and small mineral particles are all examples of objects that may be phagocytosed. Sitio Online Free Boundless Textbooks. localizado en https://www.boundless.com/ Boundless. Biologa. Unidad 9. Consultado el 3/10/2013 desde https://www.boundless.com/biology/?app-id=118433-520