15.2: Membrane Transport - Biology

15.2: Membrane Transport - Biology

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Section Overview

The chemistry of living things occurs in aqueous solutions, and balancing the concentrations of those solutions is an ongoing problem. Some materials diffuse readily through the membrane, but others are hindered, and their passage is made possible by specialized proteins, such as channels and transporters.

Transport across the membrane

One of the great wonders of the cell membrane is its ability to regulate the concentration of substances inside the cell. These substances include ions such as Ca2+, Na+, K+, and Cl; nutrients including sugars, fatty acids, and amino acids; and waste products, particularly carbon dioxide (CO2), which must leave the cell.

Design Challenge Subproblem:

Controlling what enters and exits the cell.

The membrane’s lipid bilayer structure provides the first level of control. The phospholipids are tightly packed together, and the membrane has a hydrophobic interior. This structure causes the membrane to be selectively permeable. A membrane that has selective permeability allows only substances meeting certain criteria to pass through it unaided. In the case of the cell membrane, only relatively small, nonpolar materials can move through the lipid bilayer at biologically relevant rates (remember, the lipid tails of the membrane are nonpolar). The rates of transport of various molecules is tabulated in the Membranes section. All substances that move through the membrane do so by one of two general methods, which are categorized based on whether or not the transport process is exergonic or endergonic. Passive transport is the exergonic movement of substances across the membrane. In contrast, active transport is the endergonic movement of substances across the membrane that is coupled to an exergonic reaction.

Passive Transport

Passive transport does not require the cell to expend energy. In passive transport, substances move from an area of higher concentration to an area of lower concentration, down the concentration gradient and energetically favorable. Depending on the chemical nature of the substance, different processes may be associated with passive transport.


Diffusion is a passive process of transport. A single substance tends to move from an area of high concentration to an area of low concentration until the concentration is equal across a space. You are familiar with diffusion of substances through the air. For example, think about someone opening a bottle of ammonia in a room filled with people. The ammonia gas is at its highest concentration in the bottle; its lowest concentration is at the edges of the room. The ammonia vapor will diffuse, or spread away, from the bottle, and gradually, more and more people will smell the ammonia as it spreads. Materials move within the cell’s cytosol by diffusion, and certain materials move through the plasma membrane by diffusion.

Figure 1. Diffusion through a permeable membrane moves a substance from an area of high concentration (extracellular fluid, in this case) down its concentration gradient (into the cytoplasm). Each separate substance in a medium, such as the extracellular fluid, has its own concentration gradient, independent of the concentration gradients of other materials. In addition, each substance will diffuse according to that gradient. Within a system, there will be different rates of diffusion of the different substances in the medium (Attribution: Mariana Ruiz Villareal, modified)

Factors That Affect Diffusion

If unconstrained, molecules will move through and explore space randomly at a rate that depends on their size, their shape, their environment, and their thermal energy. This type of movement underlies the diffusive movement of molecules through whatever medium they are in. The absence of a concentration gradient does not mean that this movement will stop, just that there may be no net movement of the number of molecules from one area to another, a condition known as dynamic equilibrium.

Factors influencing diffusion include:

  • Extent of the concentration gradient: The greater the difference in concentration, the more rapid the diffusion. The closer the distribution of the material gets to equilibrium, the slower the rate of diffusion becomes.
  • Shape, size and mass of the molecules diffusing: Large and heavier molecules move more slowly; therefore, they diffuse more slowly. The reverse is typically true for smaller, lighter molecules.
  • Temperature: Higher temperatures increase the energy and therefore the movement of the molecules, increasing the rate of diffusion. Lower temperatures decrease the energy of the molecules, thus decreasing the rate of diffusion.
  • Solvent density: As the density of a solvent increases, the rate of diffusion decreases. The molecules slow down because they have a more difficult time getting through the denser medium. If the medium is less dense, rates of diffusion increase. Since cells primarily use diffusion to move materials within the cytoplasm, any increase in the cytoplasm’s density will decrease the rate at which materials move in the cytoplasm.
  • Solubility: As discussed earlier, nonpolar or lipid-soluble materials pass through plasma membranes more easily than polar materials, allowing a faster rate of diffusion.
  • Surface area and thickness of the plasma membrane: Increased surface area increases the rate of diffusion, whereas a thicker membrane reduces it.
  • Distance travelled: The greater the distance that a substance must travel, the slower the rate of diffusion. This places an upper limitation on cell size. A large, spherical cell will die because nutrients or waste cannot reach or leave the center of the cell, respectively. Therefore, cells must either be small in size, as in the case of many prokaryotes, or be flattened, as with many single-celled eukaryotes.

Facilitated transport

In facilitated transport, also called facilitated diffusion, materials diffuse across the plasma membrane with the help of membrane proteins. A concentration gradient exists that allows these materials to diffuse into or out of the cell without expending cellular energy. In the case that the materials are ions or polar molecules, compounds that are repelled by the hydrophobic parts of the cell membrane, facilitated transport proteins help shield these materials from the repulsive force of the membrane, allowing them to diffuse into the cell.

Note: Possible Discussion

Compare and contrast passive diffusion and facilitated diffusion.


The integral proteins involved in facilitated transport are collectively referred to as transport proteins, and they function as either channels for the material or carriers. In both cases, they are transmembrane proteins. Different channel proteins have different transport properties. Some have evolved to be have very high specificity for the substance that is being transported while others transport a variety of molecules sharing some common characteristic(s). The interior "passageway" of channel proteins have evolved to provide a low energetic barrier for transport of substances across the membrane through the complementary arrangement of amino acid functional groups (of both backbone and side-chains). Passage through the channel allows polar compounds to avoid the nonpolar central layer of the plasma membrane that would otherwise slow or prevent their entry into the cell. While at any one time significant amounts of water crosses the membrane both in and out the rate of individual water molecule transport may not be fast enough to adapt to changing environmental conditions. For such cases Nature has evolved a special class of membrane proteins called aquaporins that allow water to pass through the membrane at a very high rate.

Figure 2: Facilitated transport moves substances down their concentration gradients. They may cross the plasma membrane with the aid of channel proteins. (Attribution: Mariana Ruiz Villareal, modified.)

Channel proteins are either open at all times or they are “gated.” The latter controls the opening of the channel. Various mechanisms may be involved in the gating mechanism. For instance, the attachment of a specific ion or small molecule to the channel protein may trigger opening. Changes in local membrane "stress" or changes in voltage across the membrane may also be triggers to open or close a channel.

Different organisms and tissues in multicellular species express different sets of channel proteins in their membranes depending on the environments they live in or specialized function they play in an organisms. This provides each type of cell with a unique membrane permeability profile that is evolved to complement its "needs" (note the anthropomorphism). For example, in some tissues, sodium and chloride ions pass freely through open channels, whereas in other tissues a gate must be opened to allow passage. This occurs in the kidney, where both forms of channels are found in different parts of the renal tubules. Cells involved in the transmission of electrical impulses, such as nerve and muscle cells, have gated channels for sodium, potassium, and calcium in their membranes. Opening and closing of these channels changes the relative concentrations on opposing sides of the membrane of these ions, resulting a change in electrical potential across the membrane that lead to message propagation in the case of nerve cells or in muscle contraction in the case of muscle cells.

Carrie Proteins

Another type of protein embedded in the plasma membrane is a carrier protein. This aptly named protein binds a substance and, in doing so, triggers a change of its own shape, moving the bound molecule from the outside of the cell to its interior; depending on the gradient, the material may move in the opposite direction. Carrier proteins are typically specific for a single substance. This selectivity adds to the overall selectivity of the plasma membrane. The molecular-scale mechanism of function for these proteins remains poorly understood.

Figure 3: Some substances are able to move down their concentration gradient across the plasma membrane with the aid of carrier proteins. Carrier proteins change shape as they move molecules across the membrane. (Attribution: Mariana Ruiz Villareal, modified.)

Carrier proteins play an important role in the function of kidneys. Glucose, water, salts, ions, and amino acids needed by the body are filtered in one part of the kidney. This filtrate, which includes glucose, is then reabsorbed in another part of the kidney with the help of carrier proteins. Because there are only a finite number of carrier proteins for glucose, if more glucose is present in the filtrate than the proteins can handle, the excess is not reabsorbed and it is excreted from the body in the urine. In a diabetic individual, this is described as “spilling glucose into the urine.” A different group of carrier proteins called glucose transport proteins, or GLUTs, are involved in transporting glucose and other hexose sugars through plasma membranes within the body.

Channel and carrier proteins transport materials at different rates. Channel proteins transport much more quickly than do carrier proteins. Channel proteins facilitate diffusion at a rate of tens of millions of molecules per second, whereas carrier proteins work at a rate of a thousand to a million molecules per second.

Note: A note of appreciation

The rates of transport just discussed are astounding. Recall that these molecular catalysts are on the scale of 10s of nanometers (10-9 meters) and that they are composed of a self-folding string of 20 amino acids and the relatively small selection of chemical functional groups that they carry.


Osmosis is the movement of water through a semipermeable membrane according to the concentration gradient of water across the membrane, which is inversely proportional to the concentration of solutes. While diffusion transports material across membranes and within cells, osmosis transports only water across a membrane and the membrane limits the diffusion of solutes in the water. Not surprisingly, the aquaporins that facilitate water movement play a large role in osmosis, most prominently in red blood cells and the membranes of kidney tubules.


Osmosis is a special case of diffusion. Water, like other substances, moves from an area of high concentration to one of low concentration. An obvious question is what makes water move at all? Imagine a beaker with a semipermeable membrane separating the two sides or halves. On both sides of the membrane the water level is the same, but there are different concentrations of a dissolved substance, or solute, that cannot cross the membrane (otherwise the concentrations on each side would be balanced by the solute crossing the membrane). If the volume of the solution on both sides of the membrane is the same, but the concentrations of solute are different, then there are different amounts of water, the solvent, on either side of the membrane.

Figure 4: In osmosis, water always moves from an area of higher water concentration to one of lower concentration. In the diagram shown, the solute cannot pass through the selectively permeable membrane, but the water can.

To illustrate this, imagine two full glasses of water. One has a single teaspoon of sugar in it, whereas the second one contains one-quarter cup of sugar. If the total volume of the solutions in both cups is the same, which cup contains more water? Because the large amount of sugar in the second cup takes up much more space than the teaspoon of sugar in the first cup, the first cup has more water in it.

Returning to the beaker example, recall that it has a mixture of solutes on either side of the membrane. A principle of diffusion is that the molecules move around and will spread evenly throughout the medium if they can. However, only the material capable of getting through the membrane will diffuse through it. In this example, the solute cannot diffuse through the membrane, but the water can. Water has a concentration gradient in this system. Thus, water will diffuse down its concentration gradient, crossing the membrane to the side where it is less concentrated. This diffusion of water through the membrane—osmosis—will continue until the concentration gradient of water goes to zero or until the hydrostatic pressure of the water balances the osmotic pressure. Osmosis proceeds constantly in living systems.


Tonicity describes how an extracellular solution can change the volume of a cell by affecting osmosis. A solution's tonicity often directly correlates with the osmolarity of the solution. Osmolarity describes the total solute concentration of the solution. A solution with low osmolarity has a greater number of water molecules relative to the number of solute particles; a solution with high osmolarity has fewer water molecules with respect to solute particles. In a situation in which solutions of two different osmolarities are separated by a membrane permeable to water, though not to the solute, water will move from the side of the membrane with lower osmolarity (and more water) to the side with higher osmolarity (and less water). This effect makes sense if you remember that the solute cannot move across the membrane, and thus the only component in the system that can move—the water—moves along its own concentration gradient. An important distinction that concerns living systems is that osmolarity measures the number of particles (which may be molecules) in a solution. Therefore, a solution that is cloudy with cells may have a lower osmolarity than a solution that is clear, if the second solution contains more dissolved molecules than there are cells.

Hypotonic Solutions

Three terms—hypotonic, isotonic, and hypertonic—are used to relate the osmolarity of a cell to the osmolarity of the extracellular fluid that contains the cells. In a hypotonicsituation, the extracellular fluid has lower osmolarity than the fluid inside the cell, and water enters the cell. (In living systems, the point of reference is always the cytoplasm, so the prefix hypo- means that the extracellular fluid has a lower concentration of solutes, or a lower osmolarity, than the cell cytoplasm.) It also means that the extracellular fluid has a higher concentration of water in the solution than does the cell. In this situation, water will follow its concentration gradient and enter the cell.

Hypertonic Solutions

As for a hypertonic solution, the prefix hyper- refers to the extracellular fluid having a higher osmolarity than the cell’s cytoplasm; therefore, the fluid contains less water than the cell does. Because the cell has a relatively higher concentration of water, water will leave the cell.

Isotonic Solutions

In an isotonic solution, the extracellular fluid has the same osmolarity as the cell. If the osmolarity of the cell matches that of the extracellular fluid, there will be no net movement of water into or out of the cell, although water will still move in and out. Blood cells and plant cells in hypertonic, isotonic, and hypotonic solutions take on characteristic appearances.


Figure 5: Osmotic pressure changes the shape of red blood cells in hypertonic, isotonic, and hypotonic solutions. (credit: Mariana Ruiz Villareal)

A doctor injects a patient with what the doctor thinks is an isotonic saline solution. The patient dies, and an autopsy reveals that many red blood cells have been destroyed. Do you think the solution the doctor injected was really isotonic?

Link to Learning:

For a video illustrating the process of diffusion in solutions, visit this site.

Tonicity in Living Systems

In a hypotonic environment, water enters a cell, and the cell swells. In an isotonic condition, the relative concentrations of solute and solvent are equal on both sides of the membrane. There is no net water movement; therefore, there is no change in the size of the cell. In a hypertonic solution, water leaves a cell and the cell shrinks. If either the hypo- or hyper- condition goes to excess, the cell’s functions become compromised, and the cell may be destroyed.

A red blood cell will burst, or lyse, when it swells beyond the plasma membrane’s capability to expand. Remember, the membrane resembles a mosaic, with discrete spaces between the molecules composing it. If the cell swells, and the spaces between the lipids and proteins become too large, the cell will break apart.

In contrast, when excessive amounts of water leave a red blood cell, the cell shrinks, or crenates. This has the effect of concentrating the solutes left in the cell, making the cytosol denser and interfering with diffusion within the cell. The cell’s ability to function will be compromised and may also result in the death of the cell.

Various living things have ways of controlling the effects of osmosis—a mechanism called osmoregulation. Some organisms, such as plants, fungi, bacteria, and some protists, have cell walls that surround the plasma membrane and prevent cell lysis in a hypotonic solution. The plasma membrane can only expand to the limit of the cell wall, so the cell will not lyse. In fact, the cytoplasm in plants is always slightly hypertonic to the cellular environment, and water will always enter a cell if water is available. This inflow of water produces turgor pressure, which stiffens the cell walls of the plant. In nonwoody plants, turgor pressure supports the plant. Conversly, if the plant is not watered, the extracellular fluid will become hypertonic, causing water to leave the cell. In this condition, the cell does not shrink because the cell wall is not flexible. However, the cell membrane detaches from the wall and constricts the cytoplasm. This is called plasmolysis. Plants lose turgor pressure in this condition and wilt.

Figure 6: The turgor pressure within a plant cell depends on the tonicity of the solution that it is bathed in. (credit: modification of work by Mariana Ruiz Villareal)
Figure 7: Without adequate water, the plant on the left has lost turgor pressure, visible in its wilting; the turgor pressure is restored by watering it (right). (credit: Victor M. Vicente Selvas)

Many marine invertebrates have internal salt levels matched to their environments, making them isotonic with the water in which they live. Fish, however, must spend approximately five percent of their metabolic energy maintaining osmotic homeostasis. Freshwater fish live in an environment that is hypotonic to their cells. These fish actively take in salt through their gills and excrete diluted urine to rid themselves of excess water. Saltwater fish live in the reverse environment, which is hypertonic to their cells, and they secrete salt through their gills and excrete highly concentrated urine.

In vertebrates, the kidneys regulate the amount of water in the body. Osmoreceptors are specialized cells in the brain that monitor the concentration of solutes in the blood. If the levels of solutes increase beyond a certain range, a hormone is released that retards water loss through the kidney and dilutes the blood to safer levels. Animals also have high concentrations of albumin, which is produced by the liver, in their blood. This protein is too large to pass easily through plasma membranes and is a major factor in controlling the osmotic pressures applied to tissues.

Exercise 1

A doctor injects a patient with what the doctor thinks is an isotonic saline solution. Do you think the solution the doctor injected was really isotonic?

Review Questions

Exercise 2

The principal force driving movement in diffusion is the __________.

  1. temperature
  2. particle size
  3. concentration gradient
  4. membrane surface area

Exercise 3

What problem is faced by organisms that live in fresh water?

  1. Their bodies tend to take in too much water.
  2. They have no way of controlling their tonicity.
  3. Only salt water poses problems for animals that live in it.
  4. Their bodies tend to lose too much water to their environment.

Free Response

Exercise 4

Why does water move through a membrane?

Active Transport

Section Summary

The combined gradient that affects an ion includes its concentration gradient and its electrical gradient. A positive ion, for example, might tend to diffuse into a new area, down its concentration gradient, but if it is diffusing into an area of net positive charge, its diffusion will be hampered by its electrical gradient. When dealing with ions in aqueous solutions, a combination of the electrochemical and concentration gradients, rather than just the concentration gradient alone, must be considered. Living cells need certain substances that exist inside the cell in concentrations greater than they exist in the extracellular space. Moving substances up their electrochemical gradients requires energy from the cell. Active transport uses energy stored in ATP to fuel this transport. Active transport of small molecular-sized materials uses integral proteins in the cell membrane to move the materials: These proteins are analogous to pumps. Some pumps, which carry out primary active transport, couple directly with ATP to drive their action. In co-transport (or secondary active transport), energy from primary transport can be used to move another substance into the cell and up its concentration gradient.

Active Transport

Active transport mechanisms require the use of the cell’s energy, usually in the form of adenosine triphosphate (ATP). If a substance must move into the cell against its concentration gradient—that is, if the concentration of the substance inside the cell is greater than its concentration in the extracellular fluid (and vice versa)—the cell must use energy to move the substance. Some active transport mechanisms move small-molecular weight materials, such as ions, through the membrane. Other mechanisms transport much larger molecules.

Moving Against a Gradient

To move substances against a concentration or electrochemical gradient, the cell must use energy. This energy is harvested from ATP generated through the cell’s metabolism. Active transport mechanisms, collectively called pumps, work against electrochemical gradients. Small substances constantly pass through plasma membranes. Active transport maintains concentrations of ions and other substances needed by living cells in the face of these passive movements. Much of a cell’s supply of metabolic energy may be spent maintaining these processes. (Most of a red blood cell’s metabolic energy is used to maintain the imbalance between exterior and interior sodium and potassium levels required by the cell.) Because active transport mechanisms depend on a cell’s metabolism for energy, they are sensitive to many metabolic poisons that interfere with the supply of ATP.

Two mechanisms exist for the transport of small-molecular weight material and small molecules. Primary active transport moves ions across a membrane and creates a difference in charge across that membrane, which is directly dependent on ATP. Secondary active transport describes the movement of material that is due to the electrochemical gradient established by primary active transport that does not directly require ATP.

Carrier Proteins for Active Transport

An important membrane adaption for active transport is the presence of specific carrier proteins or pumps to facilitate movement: there are three types of these proteins or transporters. A uniporter carries one specific ion or molecule. A symporter carries two different ions or molecules, both in the same direction. An antiporter also carries two different ions or molecules, but in different directions. All of these transporters can also transport small, uncharged organic molecules like glucose. These three types of carrier proteins are also found in facilitated diffusion, but they do not require ATP to work in that process. Some examples of pumps for active transport are Na+-K+ ATPase, which carries sodium and potassium ions, and H+-K+ ATPase, which carries hydrogen and potassium ions. Both of these are antiporter carrier proteins. Two other carrier proteins are Ca2+ ATPase and H+ ATPase, which carry only calcium and only hydrogen ions, respectively. Both are pumps.

Figure 9: A uniporter carries one molecule or ion. A symporter carries two different molecules or ions, both in the same direction. An antiporter also carries two different molecules or ions, but in different directions. (credit: modification of work by “Lupask”/Wikimedia Commons)

Primary Active Transport

In primary active transport, the energy is derived directly from the breakdown of ATP. Often times, primary active transport such as that shown below which functions to transport sodium and potassium ions allows secondary active transport to occur (discussed in the section below). The second transport method is still considered active because it depends on the use of energy from the primary transport.

Figure 10: Primary active transport moves ions across a membrane, creating an electrochemical gradient (electrogenic transport). (credit: modification of work by Mariana Ruiz Villareal)

One of the most important pumps in animal cells is the sodium-potassium pump (Na+-K+ ATPase), which maintains the electrochemical gradient (and the correct concentrations of Na+ and K+) in living cells. The sodium-potassium pump moves K+ into the cell while moving Na+ out at the same time, at a ratio of three Na+ for every two K+ ions moved in. The Na+-K+ ATPase exists in two forms, depending on its orientation to the interior or exterior of the cell and its affinity for either sodium or potassium ions. The process consists of the following six steps.

  1. With the enzyme oriented towards the interior of the cell, the carrier has a high affinity for sodium ions. Three ions bind to the protein.
  2. ATP is hydrolyzed by the protein carrier and a low-energy phosphate group attaches to it.
  3. As a result, the carrier changes shape and re-orients itself towards the exterior of the membrane. The protein’s affinity for sodium decreases and the three sodium ions leave the carrier.
  4. The shape change increases the carrier’s affinity for potassium ions, and two such ions attach to the protein. Subsequently, the low-energy phosphate group detaches from the carrier.
  5. With the phosphate group removed and potassium ions attached, the carrier protein repositions itself towards the interior of the cell.
  6. The carrier protein, in its new configuration, has a decreased affinity for potassium, and the two ions are released into the cytoplasm. The protein now has a higher affinity for sodium ions, and the process starts again.

Several things have happened as a result of this process. At this point, there are more sodium ions outside of the cell than inside and more potassium ions inside than out. For every three ions of sodium that move out, two ions of potassium move in. This results in the interior being slightly more negative relative to the exterior. This difference in charge is important in creating the conditions necessary for the secondary process. The sodium-potassium pump is, therefore, an electrogenic pump (a pump that creates a charge imbalance), creating an electrical imbalance across the membrane and contributing to the membrane potential.

Link to Learning

Visit the site to see a simulation of active transport in a sodium-potassium ATPase.

Secondary Active Transport (Co-transport)

Secondary active transport brings sodium ions, and possibly other compounds, into the cell. As sodium ion concentrations build outside of the plasma membrane because of the action of the primary active transport process, an electrochemical gradient is created. If a channel protein exists and is open, the sodium ions will be pulled through the membrane. This movement is used to transport other substances that can attach themselves to the transport protein through the membrane. Many amino acids, as well as glucose, enter a cell this way. This secondary process is also used to store high-energy hydrogen ions in the mitochondria of plant and animal cells for the production of ATP. The potential energy that accumulates in the stored hydrogen ions is translated into kinetic energy as the ions surge through the channel protein ATP synthase, and that energy is used to convert ADP into ATP.

Figure 11: An electrochemical gradient, created by primary active transport, can move other substances against their concentration gradients, a process called co-transport or secondary active transport. (credit: modification of work by Mariana Ruiz Villareal)

If the pH outside the cell decreases, would you expect the amount of amino acids transported into the cell to increase or decrease?


Exercise 5

Injection of a potassium solution into a person’s blood is lethal; this is used in capital punishment and euthanasia. Why do you think a potassium solution injection is lethal?

Exercise 6

If the pH outside the cell decreases, would you expect the amount of amino acids transported into the cell to increase or decrease?

Review Questions

Exercise 7

Active transport must function continuously because __________.

  1. plasma membranes wear out
  2. not all membranes are amphiphilic
  3. facilitated transport opposes active transport
  4. diffusion is constantly moving solutes in opposite directions

Exercise 8

How does the sodium-potassium pump make the interior of the cell negatively charged?

  1. by expelling anions
  2. by pulling in anions
  3. by expelling more cations than are taken in
  4. by taking in and expelling an equal number of cations

Exercise 9

What is the combination of an electrical gradient and a concentration gradient called?

  1. potential gradient
  2. electrical potential
  3. concentration potential
  4. electrochemical gradient

Free Response

Exercise 10

Where does the cell get energy for active transport processes?

Exercise 11

How does the sodium-potassium pump contribute to the net negative charge of the interior of the cell?


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Figure 1. Phospholipid Structure. A phospholipid molecule consists of a polar phosphate “head,” which is hydrophilic and a non-polar lipid “tail,” which is hydrophobic. Unsaturated fatty acids result in kinks in the hydrophobic tails.

The cell membrane is an extremely pliable structure composed primarily of back-to-back phospholipids (a “bilayer”). Cholesterol is also present, which contributes to the fluidity of the membrane, and there are various proteins embedded within the membrane that have a variety of functions. A single phospholipid molecule has a phosphate group on one end, called the “head,” and two side-by-side chains of fatty acids that make up the lipid tails (Figure 1). The phosphate group is negatively charged, making the head polar and hydrophilic—or “water loving.”

A hydrophilic molecule (or region of a molecule) is one that is attracted to water. The phosphate heads are thus attracted to the water molecules of both the extracellular and intracellular environments. The lipid tails, on the other hand, are uncharged, or nonpolar, and are hydrophobic—or “water fearing.”

A hydrophobic molecule (or region of a molecule) repels and is repelled by water. Some lipid tails consist of saturated fatty acids and some contain unsaturated fatty acids. This combination adds to the fluidity of the tails that are constantly in motion. Phospholipids are thus amphipathic molecules.

An amphipathic molecule is one that contains both a hydrophilic and a hydrophobic region. In fact, soap works to remove oil and grease stains because it has amphipathic properties. The hydrophilic portion can dissolve in water while the hydrophobic portion can trap grease in micelles that then can be washed away.

Figure 2. Phospolipid Bilayer. The phospholipid bilayer consists of two adjacent sheets of phospholipids, arranged tail to tail. The hydrophobic tails associate with one another, forming the interior of the membrane. The polar heads contact the fluid inside and outside of the cell.

The cell membrane consists of two adjacent layers of phospholipids. The lipid tails of one layer face the lipid tails of the other layer, meeting at the interface of the two layers. The phospholipid heads face outward, one layer exposed to the interior of the cell and one layer exposed to the exterior (Figure 2).

Because the phosphate groups are polar and hydrophilic, they are attracted to water in the intracellular fluid. Intracellular fluid (ICF) is the fluid interior of the cell. The phosphate groups are also attracted to the extracellular fluid. Extracellular fluid (ECF) is the fluid environment outside the enclosure of the cell membrane. Interstitial fluid (IF) is the term given to extracellular fluid not contained within blood vessels. Because the lipid tails are hydrophobic, they meet in the inner region of the membrane, excluding watery intracellular and extracellular fluid from this space. The cell membrane has many proteins, as well as other lipids (such as cholesterol), that are associated with the phospholipid bilayer. An important feature of the membrane is that it remains fluid the lipids and proteins in the cell membrane are not rigidly locked in place.

Membrane Proteins

The lipid bilayer forms the basis of the cell membrane, but it is peppered throughout with various proteins. Two different types of proteins that are commonly associated with the cell membrane are the integral proteins and peripheral protein (Figure 3). As its name suggests, an integral protein is a protein that is embedded in the membrane. A channel protein is an example of an integral protein that selectively allows particular materials, such as certain ions, to pass into or out of the cell.

Figure 3. Cell Membrane. The cell membrane of the cell is a phospholipid bilayer containing many different molecular components, including proteins and cholesterol, some with carbohydrate groups attached.

Another important group of integral proteins are cell recognition proteins, which serve to mark a cell’s identity so that it can be recognized by other cells. A receptor is a type of recognition protein that can selectively bind a specific molecule outside the cell, and this binding induces a chemical reaction within the cell. A ligand is the specific molecule that binds to and activates a receptor. Some integral proteins serve dual roles as both a receptor and an ion channel. One example of a receptor-ligand interaction is the receptors on nerve cells that bind neurotransmitters, such as dopamine. When a dopamine molecule binds to a dopamine receptor protein, a channel within the transmembrane protein opens to allow certain ions to flow into the cell.

Some integral membrane proteins are glycoproteins. A glycoprotein is a protein that has carbohydrate molecules attached, which extend into the extracellular matrix. The attached carbohydrate tags on glycoproteins aid in cell recognition. The carbohydrates that extend from membrane proteins and even from some membrane lipids collectively form the glycocalyx.

The glycocalyx is a fuzzy-appearing coating around the cell formed from glycoproteins and other carbohydrates attached to the cell membrane. The glycocalyx can have various roles. For example, it may have molecules that allow the cell to bind to another cell, it may contain receptors for hormones, or it might have enzymes to break down nutrients. The glycocalyces found in a person’s body are products of that person’s genetic makeup. They give each of the individual’s trillions of cells the “identity” of belonging in the person’s body. This identity is the primary way that a person’s immune defense cells “know” not to attack the person’s own body cells, but it also is the reason organs donated by another person might be rejected.

Peripheral proteins are typically found on the inner or outer surface of the lipid bilayer but can also be attached to the internal or external surface of an integral protein. These proteins typically perform a specific function for the cell. Some peripheral proteins on the surface of intestinal cells, for example, act as digestive enzymes to break down nutrients to sizes that can pass through the cells and into the bloodstream.

Transport across the Cell Membrane

One of the great wonders of the cell membrane is its ability to regulate the concentration of substances inside the cell. These substances include ions such as Ca 2+ , Na + , K + , and Cl – nutrients including sugars, fatty acids, and amino acids and waste products, particularly carbon dioxide (CO2), which must leave the cell. The membrane’s lipid bilayer structure provides the first level of control. The phospholipids are tightly packed together, and the membrane has a hydrophobic interior. This structure causes the membrane to be selectively permeable.

Selective Permeability

A membrane that has selective permeability allows only substances meeting certain criteria to pass through it unaided. In the case of the cell membrane, only relatively small, nonpolar materials can move through the lipid bilayer (remember, the lipid tails of the membrane are nonpolar). Some examples of these are other lipids, oxygen and carbon dioxide gases, and alcohol. However, water-soluble materials—like glucose, amino acids, and electrolytes—need some assistance to cross the membrane because they are repelled by the hydrophobic tails of the phospholipid bilayer. All substances that move through the membrane do so by one of two general methods, which are categorized based on whether or not energy is required.

Passive and Active Passage Through the Cell Membrane

Passive transport is the movement of substances across the membrane without the expenditure of cellular energy. In contrast, active transport is the movement of substances across the membrane using energy from adenosine triphosphate (ATP).

Passive Transport

Passive processes do not use ATP but do need some sort of driving force. It is usually from kinetic energy in the form of a concentration gradient. Molecules will tend to move from high to low concentrations by the random movement of molecules. There are 3 main types of passive processes.

In order to understand how substances move passively across a cell membrane, it is necessary to understand concentration gradients and diffusion. A concentration gradient is the difference in concentration of a substance across a space. Molecules (or ions) will spread/diffuse from where they are more concentrated to where they are less concentrated until they are equally distributed in that space. (When molecules move in this way, they are said to move down their concentration gradient.) Diffusion is the movement of particles from an area of higher concentration to an area of lower concentration. A couple of common examples will help to illustrate this concept. Imagine being inside a closed bathroom. If a bottle of perfume were sprayed, the scent molecules would naturally diffuse from the spot where they left the bottle to all corners of the bathroom, and this diffusion would go on until no more concentration gradient remains. Another example is a spoonful of sugar placed in a cup of tea. Eventually the sugar will diffuse throughout the tea until no concentration gradient remains. In both cases, if the room is warmer or the tea hotter, diffusion occurs even faster as the molecules are bumping into each other and spreading out faster than at cooler temperatures. Having an internal body temperature around 98.6 ° F thus also aids in diffusion of particles within the body.

Whenever a substance exists in greater concentration on one side of a semipermeable membrane, such as the cell membranes, any substance that can move down its concentration gradient across the membrane will do so. Consider substances that can easily diffuse through the lipid bilayer of the cell membrane, such as the gases oxygen (O2) and CO2. O2 generally diffuses into cells because it is more concentrated outside of them, and CO2 typically diffuses out of cells because it is more concentrated inside of them. Neither of these examples requires any energy on the part of the cell, and therefore they use passive transport to move across the membrane. Before moving on, you need to review the gases that can diffuse across a cell membrane. Because cells rapidly use up oxygen during metabolism, there is typically a lower concentration of O2 inside the cell than outside. As a result, oxygen will diffuse from the interstitial fluid directly through the lipid bilayer of the membrane and into the cytoplasm within the cell. On the other hand, because cells produce CO2 as a byproduct of metabolism, CO2 concentrations rise within the cytoplasm therefore, CO2 will move from the cell through the lipid bilayer and into the interstitial fluid, where its concentration is lower. This mechanism of molecules spreading from where they are more concentrated to where they are less concentration is a form of passive transport called simple diffusion (Figure 4).

Figure 4. Simple Diffusion across the Cell (Plasma) Membrane. The structure of the lipid bilayer allows only small, non-polar substances such as oxygen and carbon dioxide to pass through the cell membrane, down their concentration gradient, by simple diffusion.

Solutes dissolved in water on either side of the cell membrane will tend to diffuse down their concentration gradients, but because most substances cannot pass freely through the lipid bilayer of the cell membrane, their movement is restricted to protein channels and specialized transport mechanisms in the membrane. Facilitated diffusion is the diffusion process used for those substances that cannot cross the lipid bilayer due to their size and/or polarity (Figure 5). A common example of facilitated diffusion is the movement of glucose into the cell, where it is used to make ATP. Although glucose can be more concentrated outside of a cell, it cannot cross the lipid bilayer via simple diffusion because it is both large and polar. To resolve this, a specialized carrier protein called the glucose transporter will transfer glucose molecules into the cell to facilitate its inward diffusion.

Figure 5. Facilitated Diffusion. (a) Facilitated diffusion of substances crossing the cell (plasma) membrane takes place with the help of proteins such as channel proteins and carrier proteins. Channel proteins are less selective than carrier proteins, and usually mildly discriminate between their cargo based on size and charge. (b) Carrier proteins are more selective, often only allowing one particular type of molecule to cross.

Even though sodium ions (Na + ) are highly concentrated outside of cells, these electrolytes are polarized and cannot pass through the nonpolar lipid bilayer of the membrane. Their diffusion is facilitated by membrane proteins that form sodium channels (or “pores”), so that Na + ions can move down their concentration gradient from outside the cells to inside the cells. There are many other solutes that must undergo facilitated diffusion to move into a cell, such as amino acids, or to move out of a cell, such as wastes. Because facilitated diffusion is a passive process, it does not require energy expenditure by the cell. Water also can move freely across the cell membrane of all cells, either through protein channels or by slipping between the lipid tails of the membrane itself.

Osmosisis the diffusion of water through a semipermeable membrane (Figure 6).

Figure 6. Osmosis. Osmosis is the diffusion of water through a semipermeable membrane down its concentration gradient. If a membrane is permeable to water, though not to a solute, water will equalize its own concentration by diffusing to the side of lower water concentration (and thus the side of higher solute concentration). In the beaker on the left, the solution on the right side of the membrane is hypertonic.

The movement of water molecules is not itself regulated by cells, so it is important that cells are exposed to an environment in which the concentration of solutes outside of the cells (in the extracellular fluid) is equal to the concentration of solutes inside the cells (in the cytoplasm). Two solutions that have the same concentration of solutes are said to be isotonic (equal tension). When cells and their extracellular environments are isotonic, the concentration of water molecules is the same outside and inside the cells, and the cells maintain their normal shape (and function). Osmosis occurs when there is an imbalance of solutes outside of a cell versus inside the cell. A solution that has a higher concentration of solutes than another solution is said to be hypertonic, and water molecules tend to diffuse into a hypertonic solution (Figure 7). Cells in a hypertonic solution will shrivel as water leaves the cell via osmosis. In contrast, a solution that has a lower concentration of solutes than another solution is said to be hypotonic, and water molecules tend to diffuse out of a hypotonic solution. Cells in a hypotonic solution will take on too much water and swell, with the risk of eventually bursting. A critical aspect of homeostasis in living things is to create an internal environment in which all of the body’s cells are in an isotonic solution. Various organ systems, particularly the kidneys, work to maintain this homeostasis.

Figure 7. Concentration of Solutions. A hypertonic solution has a solute concentration higher than another solution. An isotonic solution has a solute concentration equal to another solution. A hypotonic solution has a solute concentration lower than another solution.

Another mechanism besides diffusion to passively transport materials between compartments is filtration. Unlike diffusion of a substance from where it is more concentrated to less concentrated, filtration uses a hydrostatic pressure gradient that pushes the fluid—and the solutes within it—from a higher pressure area to a lower pressure area. Filtration is an extremely important process in the body. For example, the circulatory system uses filtration to move plasma and substances across the endothelial lining of capillaries and into surrounding tissues, supplying cells with the nutrients. Filtration pressure in the kidneys provides the mechanism to remove wastes from the bloodstream.

Active Transport

You have just finished investigating the passive methods of transport, now let’s look at active methods. In active methods the cell must expend energy (ATP) to do the work of moving molecules. Active transport often occurs when the molecule is being moved against its concentration gradient or when moving very large molecules into our out of the cell. There are 3 main types of active processes.

During active transport, ATP is required to move a substance across a membrane, often with the help of protein carriers, and usually against its concentration gradient. One of the most common types of active transport involves proteins that serve as pumps. The word “pump” probably conjures up thoughts of using energy to pump up the tire of a bicycle or a basketball. Similarly, energy from ATP is required for these membrane proteins to transport substances—molecules or ions—across the membrane, usually against their concentration gradients (from an area of low concentration to an area of high concentration). The sodium-potassium pump, which is also called N + /K + ATPase, transports sodium out of a cell while moving potassium into the cell. The Na + /K + pump is an important ion pump found in the membranes of many types of cells. These pumps are particularly abundant in nerve cells, which are constantly pumping out sodium ions and pulling in potassium ions to maintain an electrical gradient across their cell membranes. An electrical gradient is a difference in electrical charge across a space. In the case of nerve cells, for example, the electrical gradient exists between the inside and outside of the cell, with the inside being negatively-charged (at around -70 mV) relative to the outside. The negative electrical gradient is maintained because each Na + /K + pump moves three Na + ions out of the cell and two K + ions into the cell for each ATP molecule that is used (Figure 8). This process is so important for nerve cells that it accounts for the majority of their ATP usage.

Figure 8. Sodium-Potassium Pump. The sodium-potassium pump is found in many cell (plasma) membranes. Powered by ATP, the pump moves sodium and potassium ions in opposite directions, each against its concentration gradient. In a single cycle of the pump, three sodium ions are extruded from and two potassium ions are imported into the cell.

Other forms of active transport do not involve membrane carriers. Endocytosis (bringing “into the cell”) is the process of a cell ingesting material by enveloping it in a portion of its cell membrane, and then pinching off that portion of membrane (Figure 9). Once pinched off, the portion of membrane and its contents becomes an independent, intracellular vesicle. A vesicle is a membranous sac—a spherical and hollow organelle bounded by a lipid bilayer membrane. Endocytosis often brings materials into the cell that must to be broken down or digested. Phagocytosis (“cell eating”) is the endocytosis of large particles. Many immune cells engage in phagocytosis of invading pathogens. Like little Pac-men, their job is to patrol body tissues for unwanted matter, such as invading bacterial cells, phagocytize them, and digest them. In contrast to phagocytosis, pinocytosis (“cell drinking”) brings fluid containing dissolved substances into a cell through membrane vesicles.

Figure 9. Three Forms of Endocytosis. Endocytosis is a form of active transport in which a cell envelopes extracellular materials using its cell membrane. (a) In phagocytosis, which is relatively nonselective, the cell takes in a large particle. (b) In pinocytosis, the cell takes in small particles in fluid. (c) In contrast, receptor-mediated endocytosis is quite selective. When external receptors bind a specific ligand, the cell responds by endocytosing the ligand.

Figure 10. Exocytosis. Exocytosis is much like endocytosis in reverse. Material destined for export is packaged into a vesicle inside the cell. The membrane of the vesicle fuses with the cell membrane, and the contents are released into the extracellular space.

Phagocytosis and pinocytosis take in large portions of extracellular material, and they are typically not highly selective in the substances they bring in. Cells regulate the endocytosis of specific substances via receptor-mediated endocytosis. Receptor-mediated endocytosis is endocytosis by a portion of the cell membrane that contains many receptors that are specific for a certain substance. Once the surface receptors have bound sufficient amounts of the specific substance (the receptor’s ligand), the cell will endocytose the part of the cell membrane containing the receptor-ligand complexes. Iron, a required component of hemoglobin, is endocytosed by red blood cells in this way. Iron is bound to a protein called transferrin in the blood. Specific transferrin receptors on red blood cell surfaces bind the iron-transferrin molecules, and the cell endocytoses the receptor-ligand complexes. In contrast with endocytosis, exocytosis (taking “out of the cell”) is the process of a cell exporting material using vesicular transport (Figure 10).

Many cells manufacture substances that must be secreted, like a factory manufacturing a product for export. These substances are typically packaged into membrane-bound vesicles within the cell. When the vesicle membrane fuses with the cell membrane, the vesicle releases it contents into the interstitial fluid. The vesicle membrane then becomes part of the cell membrane. Cells of the stomach and pancreas produce and secrete digestive enzymes through exocytosis (Figure 11). Endocrine cells produce and secrete hormones that are sent throughout the body, and certain immune cells produce and secrete large amounts of histamine, a chemical important for immune responses.

Figure 11. Pancreatic Cells’ Enzyme Products. The pancreatic acinar cells produce and secrete many enzymes that digest food. The tiny black granules in this electron micrograph are secretory vesicles filled with enzymes that will be exported from the cells via exocytosis. LM × 2900. (Micrograph provided by the Regents of University of Michigan Medical School © 2012)

Diseases of the Cell: Cystic Fibrosis

Cystic fibrosis (CF) affects approximately 30,000 people in the United States, with about 1,000 new cases reported each year. The genetic disease is most well known for its damage to the lungs, causing breathing difficulties and chronic lung infections, but it also affects the liver, pancreas, and intestines. Only about 50 years ago, the prognosis for children born with CF was very grim—a life expectancy rarely over 10 years. Today, with advances in medical treatment, many CF patients live into their 30s.

The symptoms of CF result from a malfunctioning membrane ion channel called the cystic fibrosis transmembrane conductance regulator, or CFTR. In healthy people, the CFTR protein is an integral membrane protein that transports Cl – ions out of the cell. In a person who has CF, the gene for the CFTR is mutated, thus, the cell manufactures a defective channel protein that typically is not incorporated into the membrane, but is instead degraded by the cell. The CFTR requires ATP in order to function, making its Cl – transport a form of active transport. This characteristic puzzled researchers for a long time because the Cl – ions are actually flowing down their concentration gradient when transported out of cells. Active transport generally pumps ions against their concentration gradient, but the CFTR presents an exception to this rule. In normal lung tissue, the movement of Cl – out of the cell maintains a Cl – -rich, negatively charged environment immediately outside of the cell. This is particularly important in the epithelial lining of the respiratory system.

Respiratory epithelial cells secrete mucus, which serves to trap dust, bacteria, and other debris. A cilium (plural = cilia) is one of the hair-like appendages found on certain cells. Cilia on the epithelial cells move the mucus and its trapped particles up the airways away from the lungs and toward the outside. In order to be effectively moved upward, the mucus cannot be too viscous rather it must have a thin, watery consistency. The transport of Cl – and the maintenance of an electronegative environment outside of the cell attract positive ions such as Na + to the extracellular space. The accumulation of both Cl – and Na + ions in the extracellular space creates solute-rich mucus, which has a low concentration of water molecules. As a result, through osmosis, water moves from cells and extracellular matrix into the mucus, “thinning” it out. This is how, in a normal respiratory system, the mucus is kept sufficiently watered-down to be propelled out of the respiratory system.

If the CFTR channel is absent, Cl – ions are not transported out of the cell in adequate numbers, thus preventing them from drawing positive ions. The absence of ions in the secreted mucus results in the lack of a normal water concentration gradient. Thus, there is no osmotic pressure pulling water into the mucus. The resulting mucus is thick and sticky, and the ciliated epithelia cannot effectively remove it from the respiratory system. Passageways in the lungs become blocked with mucus, along with the debris it carries. Bacterial infections occur more easily because bacterial cells are not effectively carried away from the lungs.

Membranes are semi-permeable which means that they allow certain molecules through but not others. The molecules can move in and out through passive transport which is a method that does not require any input of outside energy. It can either be done by simple diffusion or facilitated diffusion. Molecules will go from a region of high concentration to a region of low concentration as they move randomly and eventually become evenly distributed within the system if they are permeable to the membrane. Simple diffusion involves the diffusion of molecules through the phospholipid bilayer while facilitated diffusion involves the use of channel proteins embedded in the membrane. The cell membrane is hydrophobic inside so hydrophobic (lipid soluble) molecules will pass through by simple diffusion whereas hydrophilic molecules and charged particles will use facilitated diffusion. Water moves through by osmosis which is also by passive transport. Osmosis involves the movement of water molecules from a region of low solute concentration, to a region of high solute concentration. So if the solute concentration is higher inside the cell than outside the cell, water will move in and vice versa.

Active transport involves the movement of substances through the membrane using energy from ATP. The advantage of active transport is that substances can be moved against the concentration gradient, meaning from a region of low concentration to a region of high concentration. This is possible because the cell membrane has protein pumps embedded it which are used in active transport to move substances across by using ATP. Each protein pump only transports certain substances so the cell can control what comes in and what goes out.

15.2: Membrane Transport - Biology

Suitable for A/AS-level and Higher biology students.
Scroll down for answers.

1. Which of these is part of the cell membrane?
a. triglycerides
b. phospholipids
c. ATP
d. more than one of these

2. How do fat-soluble molecules normally get into a cell?
a. they dissolve in the fat layers of the membrane and enter the cell by diffusion
b. they pass through protein pores in the cell membrane
c. they are absorbed by phagocytosis
d. they never get in

3. The phospholipids are unusual molecules because:
a. they have hydrophilic regions
b. they have hydrophobic regions
c. they are triglycerides
d. both A and B

4. Which of the following statements best describes the "fluid mosaic model" of the structure of the cell membrane?
a. two layers of protein with lipid layers between the protein layers
b. two layers of lipid with proteins between the lipid layers
c. a double layer of lipid molecules with protein molecules suspended in the layer
d. A single layer of protein on the outside and a single layer of lipids on the inside

5. The movement of chloride ions from an area where chloride is concentrated to an area where chloride is less concentrated is which of these?
a. diffusion
b. active transport
c. osmosis
d. exocytosis

6. If a cell has a solute concentration of 0.07% which of the solutions would be hypotonic to the cell?
a. 0.01% solute
b. 0.1% solute
c. 1% solute
d. 10% solute

7. Which of the following is necessary in order for osmosis to occur?
a. a permeable membrane
b. a semi-permeable membrane
c. an isotonic solution
d. ATP

8. Which of these are passive transport mechanisms?
a. osmosis
b. diffusion
c. phagocytosis
d. both A and B

9. In an isotonic solution there would be:
a. no net movement of water
b. net movement of water into the cell
c. net movement of water out of the cell
d. bursting of the cell

10. The sodium-potassium pump (which carries sodium out of a cell and potassium into a cell) is an example of:
a. active transport
b. endocytosis
c. exocytosis
d. passive transport

11. The process of a cell engulfing a solid object is:
a. phagocytosis
b. exocytosis
c. pinocytosis
d. diffusion

12. What is likely to happen to a plant cell that is placed in pure water?
a. it becomes turgid
b. it becomes flaccid
c. it undergoes plasmolysis
d. it bursts

13. When a cell bursts due to osmosis, it is in a solution that is:
a. hypertonic
b. isotonic
c. hypotonic
d. either A or C

14. Why do plant cells behave differently to animal cells when placed in a hypotonic solution?
a. Plant cells are permeable to water
b. Plant cells do not carry out active transport
c. Plant cells contain a vacuole
d. Plant cells have a cell wall

15. Which of these equations is correct?
a. ATP + inorganic phosphate --> ADP
b. ADP + inorganic phosphate --> ATP
c. ATP + ADP --> inorganic phosphate

Answers: 1b, 2a, 3b, 4c, 5a, 6a, 7a, 8d, 9a, 10a, 11a, 12a, 13c, 14d, 15b

Structural Biology of Membrane Transport

T he Alam lab uses a combination of biochemical and structural (Cryo-EM and X-ray crystallography) techniques to primarily study macromolecules and macromolecular complexes involved in membrane transport, homeostasis, and biogenesis, with a particular focus on pinpointing lipid/protein interactions in context of membrane protein function. Maintenance of membrane bilayer integrity and tight control over material transfer across cellular and organellar membranes is central to proper physiological functioning. Dysfunction of these systems lies at the heart of several devastating, often fatal pathologies ranging from rare inherited diseases such as Zellweger’s syndrome and adrenoleukodystrophy (both stemming from peroxisomal dysfunction) to a range of neurodegenerative disorders, diabetes, and cancer.

The Hormel Institute ranks among a handful of elite institutions housing a Titan Krios microscope combined with a direct electron detector, a setup designed to obtain near-atomic resolution (routinely achieving better than 0.5 nanometer resolution). This allows for accurate visualization of the molecular details underlying the functioning of macromolecules and has revolutionized structural biology and allowed for targeting of complex biological problems previously considered offlimits to high resolution visualization. Such insight is invaluable in obtaining molecular details of membrane protein interactions with drugs, transport substrates, and inhibitors and can aid in the design of novel therapeutics targeting diseases stemming from membrane protein dysfunction.

Over the last year since its inception, our lab has determined the cryo-EM structures of several human membrane transporters at near atomic resolutions. Along with their biochemical characterization, these new structures will be part of several upcoming publications we are in the process of preparing. We have added a new postdoc to our roster, Dr. Le Thi My Le, who also successfully applied for and is partially funded by the 2020 Eagles Postdoctoral fellowship award. We have co-authored two recent publications with a fellow Hormel Institute collaborator, Dr. Rick Brown (Gao et al, Anal Chem, 2020), as well with Dr. Alam’s postdoctoral lab on the structure of human ABCB4 / MDR3 (Olsen et al, NSMB 2020). Finally, Dr. Alam was awarded a research grant from the United Leukodystrophy Foundation to support his work on peroxisomal biogenesis and dysfunction.

The Alam lab has made significant progress in deciphering the molecular details that govern lipid metabolism and homeostasis by studying the detailed 3D structures of an important class of membrane proteins called ABC transporters. Specifically, the Alam lab has determined the high-resolution structures of ABC transporters involved in the movement of cholesterol and phospholipids across cellular membranes, revealing hitherto unseen details about how this essential step in formation of high density lipoprotein (HDL) particles form and how their dysfunction leads to a range maladies ranging from various brain disorders including Alzheimer’s disease, to various neurodegenerative diseases, as well as cancer.

The Alam lab uses a combination of biochemistry, cell biology, and state of the art Cryo-EM tools to study this group of human proteins that has for decades proven extremely difficult to study.

Cell Membrane And Transport Worksheet Biology Answer Key

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What is Membrane Transport? (with pictures)

Animal cells have a selectively permeable membrane surrounding them that separates the interior contents of the cell from the exterior environment. The process by which ions and small soluble molecules, or solutes, pass through the cell membrane is known as membrane transport. These molecules are usually substances vital to the function and maintenance of the cell, such as glucose and amino acids. There are four main types of membrane transport: passive diffusion, or simply diffusion facilitated diffusion primary active transport and secondary active transport. Many of these transport mechanisms involve the use of specialized protein molecules located in the cell membrane called membrane transport proteins.

Passive diffusion occurs spontaneously, and is driven by the random activity of molecules in a solution. Molecules move from an area of high concentration, where there are many of them densely packed together, to an area of low concentration, where there are fewer molecules spaced further apart. Small molecules may achieve membrane transport by diffusing through the cell membrane. Rate of diffusion can be affected by many things, including the composition of the cell membrane and the size and charge of the molecule. The most well-known kind of passive diffusion is osmosis, a process involving the movement of water molecules from an area of high concentration to an area of lower concentration.

Facilitated diffusion involves the use of membrane transport proteins within the cell membrane called channel proteins. These proteins act like pores in the cell membrane, allowing water soluble particles to pass through, but barring the passage of lipophilic, or "fat-loving", molecules. Diffusion follows the same mechanism of action, with molecules moving from areas of high concentration to areas of low concentration.

Primary active transport uses energy to move ions and molecules from areas of high concentration to areas of low concentration. The energy required for primary active transport to take place is usually in the form of a nucleotide called adenosine triphosphate (ATP). One of the most commonly occurring forms of active transport is the sodium-potassium pump, which helps cells to maintain an electrical charge known as the resting potential, and also controls cell volume. The sodium-potassium pump moves sodium ions to the exterior of the cell and releases potassium ions into the cell's cytoplasm.

Secondary active transport uses membrane transport proteins called antiporters and symporters. Antiporters move ions and molecules by transporting one type of particle against its usual concentration gradient, from low to high concentration, while transporting the other type of particle in the normal way, from high to low concentration. Symporters transport two different types of molecule or ion across the cell membrane at the same time and in the same direction.

Active Transport of Amino Acids

The active transport of amino acids is also mediated by Na + cotransport.

How many sodium ions are needed to provide the free energy to transport a molecule of glutamic acid from a concentration of 0.1 mM outside the cell to 20 mM inside the cell?

Again, assume a temperature of 37°C (310°K).

7, glutamic acid molecules carry a net charge of minus 1 [View].

  • a concentration gradient (20/0.1 = 200) and a
  • electrostatic gradient (moving a negative charge against a voltage of &minus 70 mV).

Because sodium ions release only 3.3 kcal/mole (above), at least 2 Na + are needed to cotransport one molecule of glutamic acid.

Substances are transported to and from cells by the following methods:

Passive transport

This is the movement substances through the cell membrane where the cell doesn’t consume any energy. Passive transport has the following methods:


  • The movement of molecules through the membrane from a region of high concentration to a region of low concentration, which makes the concentration of molecules on both sides equal.
    For example, the exchange of carbon dioxide and oxygen gases between the inner and outer mediums of the cell during the respiration process.
  • If the concentration of molecules in a cell is greater than that of the external medium, molecules move from the cell to the external medium to balance the concentration of both sides.


  • Osmosis is the movement of pure water molecules through a membrane from a region of low concentration to a region of high concentration, which makes the concentration of both sides equal.
  • Osmotic pressure is the pressure that causes the diffusion of water through semi-permeable membranes. It increases the concentration of solutes in the solution, i.e. a directly proportional relationship.
  • Water transports from and to the cells due to the difference between the concentration of cytoplasm (fluid inside the cell) and the external medium.

1- When we put cells in a solution of low concentration, water transports from the external medium (low concentration) into the cell (high concentration), which tears up cells.

2- When we put cells in a solution of equal concentration, water transports equally between the two mediums, and cells sizes don’t change.

3- When we put cells in a solution of higher concentration, water transports from the cell (low concentration) to the external medium (high concentration), which makes cells shrink.

Facilitated transport

  • Facilitated transport is the transport of molecules through a membrane by means of a carrier protein (which carries molecules) where the cell doesn’t consume any energy.
    Forexample, glucose is transported to cells carried by carrier proteins.

Active transport

  • Active transport is the transport of big molecules and ions through the cell membrane against their concentration gradient (from low to high concentration) using energy. Active transport balance the concentration of ions inside cells.
  • Active transport allows nerve cells to control the concentration of sodium and potassium ions in them, which allows them to send nerve impulses to muscle cells.

Active transport and plant cells

  • This allows root cells to absorb the ions of soil salts (though the concentration of ions in these cells is lower than that of soil).

Bulk movement

  • Bulk movement is the transporting of relatively big molecules (such as wastes and protein particles) through cell membranes. There are two kinds of bulk movements, which are:

Transporting substances out of the cell through the plasma membrane.

How does Exocytosis take place?

  1. Golgi bodies store wastes in their vacuoles (Golgi vacuoles).
  2. They move through cytoplasm to the plasma membrane to fuse with it.
  3. They empty these wastes outside the cell.

Transporting substances to the cell through the plasma membrane.

How does Endocytosis take place?

1- A part of the plasma membrane bends to surround the particle, forming a sac.

2- The sac containing the particle moves to the cytoplasm.

  • If the solid substances enter the cell by means of endocytosis process, we call it a phagocytosis process.
  • If liquid substances enter the cell by means of the endocytosis process, we call it a pinocytosis process.
  • Solid particles, especially colloidal ones, have the ability to absorb liquids, swell, and increase in volume. For example, when a piece of wood is placed in water, it imbibes water. Imbibition extends through the piece of wood until it reaches the parts which are not submerged in water. Save

Watch the video: Fluid Mosaic Model of the Cell Membrane (June 2022).


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