CHAPTER 17  SPECIFIC DEFENSES OF THE HOST: THE IMMUNE RESPONSE

 

 

·         It has long been recognized that surviving a case of certain diseases results in long-term or lifelong immunity. This chapter concentrates on the means of defense that provides this protection—the immune response.

 

 

IMMUNITY

 

This is a specific response to invasion by a particular foreign organism or other foreign substance. The immune system recognizes these as foreign—not belonging in the body—an d develops an immune response against them. Anything that provokes an immune response is called an antigen. The immune response involves the production of antibodies and specialized lymphocytes that work to destroy or inactivate that particular antigen. Organisms that act as antigens might be bacteria, viruses, protozoa, fungi or helminths. Substances include pollen, insect venom, and transplanted tissue. Body cells that change and become cancerous may also be perceived as foreign and act as antigens, so cancers may be eliminated in very early stages by our immune system.

 

 

TYPES OF ACQUIRED IMMUNITY

 

Acquired immunity is protection developed against certain microbes or foreign substances. It is developed during a person’s lifetime.

 

 

NATURALLY ACQUIRED

ACTIVE

NATURALLY ACQUIRED

PASSIVE

ARTIFICIALLY ACQUIRED

ACTIVE

ARTIFICIALLY ACQUIRED

PASSIVE

ANTIGENS ENTER THE BODY NATURALLY AND CAUSE THE PRODUCTION OF ANTIBODIES AND SPECIALIZED LYMPHOCYTES

ANTIBODIES PASS FROM MOTHER TO FETUS ACROSS THE PLACENTA OR BY BREAST MILK

ANTIGENS ARE INTRODUCED INTO THE BODY IN VACCINES---A MODIFIED VERSION THAT DOES NOT CAUSE DISEASE BUT STILL PRODUCES AN IMMUNE RESPONSE

PREFORMED ANTIBODIES MADE BY ANOTHER INDIVIDUAL ARE INJECTED INTO THE BODY

 

 

Note that active immunity involves a response by the person’s immune system. In passive immunity the antibodies are made by one individual and transferred to another. Passive immunity has a short-term effect, since antibodies remain effective for only weeks or months.

 

NATURALLY ACQUIRED IMMUNITY

 

   1. Naturally acquired active immunity is acquired as a person is exposed to various antigens in the course of normal life. In some cases, it is lifelong. In other cases, it may last only a few years. Subclinical infections (those that do not produce signs of illness) may occur and produce this type of immunity, but usually the person becomes ill with the disease and recovers, developing immunity in the process.

 

   2. Naturally acquired passive immunity involves the transfer of immunity from mother to baby. Some of the antibodies cross the placenta before birth (transplacental transfer). After birth, antibodies can also be passed on to the baby by breast milk, especially the first secretion produced which is called colostrum. This immunity is short-term, but it protects the baby while its own immune system is developing.

 

 

ARTIFICIALLY ACQUIRED IMMUNITY

 

   1. Artificially acquired active immunity occurs as a result of vaccines. Vaccination (immunization) introduces prepared antigens called vaccines into the body. The antigen in the vaccine has been modified in some way so that it will not cause disease, but will still stimulate an immune response. Modifications include:

·         Inactivated bacterial toxins (called toxoids)

·         Killed microorganisms

·         Living microorganisms that have been weakened (attenuated)

·         Possibly just parts of the disease-causing organism are included, not the whole thing

 

   2. Artificially acquired passive immunity involves injecting antibodies already made by another person or an animal already immune to the disease. The antibodies are found in the serum of the immune individual (this is the liquid that is left after clotting has occurred and the clot has been removed along with the blood cells). Serum obtained for this purpose is sometimes called antiserum. The study of reactions between antigens and antibodies is known as serology. The serum proteins that include most 7

of the antibodies are the globulins, which include alpha, beta, and gamma globulin. Gamma globulin is the fraction that contains the greatest number of antibodies.

 

When immune serum globulin is injected, it gives immediate protection against the disease, but this protection lasts only as long as the injected antibodies last. Typically, about half of the antibodies are gone after 3 weeks, and the rest will remain effective no longer than a few months.

 

 

THE TWO BRANCHES OF THE IMMUNE RESPONSE

 

Successful study of immunology began in the late 1800’s. Emil van Behring received a Nobel Prize in  1901 for proving that immunity to diphtheria could be transferred from an immune animal to a susceptible animal by serum. This was called humoral immunity. Almost forty years later it was discovered that the actual factors involved were antibodies. Cell-mediated immunity (production of specialized lymphocytes) was discovered later. The two branches of the immune response, humoral immunity and cell-mediated immunity, work together to give specific protection against foreign cells and substances.

HUMORAL (ANTIBODY-MEDIATED) IMMUNITY

 

This involves the production of antibodies, which are found in extracellular fluids such as blood plasma, lymph, and mucus secretions. Antibodies are produced by B lymphocytes  (B cells) which have changed in response to the presence of foreign material and become plasma cells. This type of defense works best against:

 

·         Most bacteria

·         Bacterial toxins

·         Viruses that have not yet invaded cells

·         Part of the response against transplanted tissue

 

 

CELL-MEDIATED IMMUNITY

 

This involves specialized lymphocytes called T lymphocytes  (T cells) that act directly against foreign organisms or tissues. T cells also regulate the activity of other parts of the immune system. T cells work directly against:

·         Bacteria or viruses inside infected host cells or inside phagocytic cells

·         Fungi

·         Protozoa

·         Helminths

·         Primary response against transplanted tissue

·         Certain cancers

 

 

 

ANTIGENS

 

The immune system of normal individuals is able to recognize parts of the body as “self” and foreign matter as “nonself.” This is why we normally do not produce an immune response against our own tissues. (Unfortunately, this sometimes does happen when things go wrong, and results in what we call an autoimmune disease.)

 

Antigens are any foreign matter that can provoke an immune response. Typically antigens are proteins, although some large polysaccharides also act as antigens. Lipids and nucleic acids combined with proteins and polysaccharides may also be antigenic. Common antigens include:

·         Components of invading microbes such as capsules, cell walls, flagella, and fimbriae

·         Toxins produced by bacteria

·         Outer coats of viruses

·         Outer surfaces of other types of microbes

·         Pollen

·         Egg white

·         “Wrong type” blood cells

·         Serum proteins from other individuals or species

·         Surface molecules of transplanted tissues and organs

 

Specific regions of the antigen are recognized by antibodies. These special regions are called epitopes or antigenic determinants. Most antigens are large (as molecules go). Sometimes a foreign substance that has a low molecular weight on its own will combine with proteins within the body and then the combination acts as an antigen. The small molecule on its own is called a hapten. Once the combination has caused the production of antibodies, the antibodies can react with the hapten portion alone. Allergies to penicillin are good examples of this.

 

 

ANTIBODIES (IMMUNOGLOBULINS)

 

These are defensive proteins that are made in response to the presence of an antigen. They are able to recognize and bind to that same antigen and somehow inactivate or destroy it. Each antibody molecule has at least two areas called antigen-binding sites that bind to the antigenic determinant areas  of the antigen. The number of antigen-binding sites on the antibody molecule is called the valence of that antibody. Most human antibodies have 2 binding sites and are known as bivalent antibodies. Antibodies are members of the group of soluble proteins known as immunoglobulins (Igs).

 

A bivalent antibody has the simplest molecular structure and is called a monomer. Typically it has four polypeptide chains, two light chains that are identical to each other and two heavy chains which are also identical to each other.  The chains are joined to each other by disulfide links and other bonds, and the resulting complete antibody molecule has a Y shape. This Y-shaped molecule can bend in two areas called the hinge region, and take on a T shape.

 

 

 

 

 

 

 

 

 

 

 

 

The two sections located at the ends of the arms of the Y are called the variable (V) regions. The amino acid sequences (remember these are protein molecules) are identical all antibody molecules made in response to the same antigen. This will be the part that fits that particular antigen, so it is the part called the antigen-binding site. The stem of the Y and the lower part of each arm are called the constant  ( C or Fc ) regions. There are only five different structures for the Fc regions (while the variable regions can have infinite numbers of different structures). Which of these five possible structures the Fc region of the antibody molecule has is what places it in one of five major classes of immunoglobulins. 

 

The stem of the Y is called the Fc region, and is important because, after the antigen-binding sites on the arms of the Y are attached to antigens, this Fc region can bind complement and enhance the destruction of the bacterium. The Fc region can also bind to a cell, leaving the antigen-binding sites projecting out and ready to bind with antigens.

 

 

FIVE CLASSES OF IMMUNOGLOBULINS

 

These are:

·         IgG

·         IgM

·         IgA

·         IgD

·         IgE

 

Each class has a different role in the immune response. IgGs, IgDs, and IgEs have the Y-shaped structure. IgMs and IgAs are usually two or more monomers joined together.

 

See TABLE 17.1 P 507  for a sketch of the structures and a summary of the characteristics.

 

   1. IgG—these antibodies account for about 80% of the antibodies circulating in blood. They can cross the walls of blood vessels and enter tissue fluids. They can cross the placenta to give passive immunity to the fetus. Functions:

·         Protects against circulating bacteria and viruses

·         Neutralizes bacterial toxins

·         Triggers the complement system

·         Enhances phagocytosis

 

   2. IgM—5 - 10% of circulating antibodies. One IgM molecule consists of five monomers held together by a polypeptide called a J chain. This produces a large molecule which generally is too big to leave blood vessels. Functions:

·         Type involved in reactions to “wrong” ABO group blood cells

·         Aggregates antigens (lumps them together)

·         Triggers complement system

·         Enhances phagocytosis

Overall, these antibodies are less effective than IgG antibodies, but IgMs can be produced quickly in response to a first encounter with an antigen.

 

   3. IgA---10 - 15% of circulating antibodies (in blood) but found in large numbers in mucous membranes and body secretions such as mucus, saliva, tears, and breast milk. Considering the body as a whole, IgAs are the most abundant antibodies. Considering only the blood (serum) IgGs are the most abundant. When found in serum, this class is most in the form of a monomer and is called serum IgA. In secretions, two monomers are joined by a J chain and this form is called secretory IgA. This form is produced by plasma cells in the mucous membrane, and then passes through a mucosal cell on its way to become part of the secretion. As it passes through the mucosal cell, it acquires a polypeptide called a secretory component that protects it from enzymes that may also be found in the secretion. Functions:

·         Prevents attachment of pathogens to the mucous membranes (viruses and certain bacteria)

·         Very important in resistance to respiratory pathogens. Unfortunately, IgA immunity is short-lived and this is why our resistance to respiratory infections is usually not longterm.

·         Found in large numbers in colostrum and functions in preventing gastrointestinal infections in infants

 

·         Act as antigen receptors on the surface of B cells

·         No known function in serum

 

   5. IgE---these also are found only in tiny traces in serum (0.002% of circulating antibodies). These molecules bind tightly by their Fc regions to receptors on the surface of mast cells and basophils. These are the antibodies responsible for allergic reactions, which often do more harm than good. When an antigen binds to an IgE antibody, this causes the associated mast cell or basophil to release histamine and other chemicals, which produces symptoms ranging from mild hay fever to severe, fatal allergic reactions. This type of reaction can help protect against parasitic worms. Functions:

·         Beneficial---when bound to antigen, attracts IgGs, complement, and phagocytes to combat parasites

·         Harmful---allergic reactions to substances which otherwise are harmless to the body

 

 

 

B CELLS AND HUMORAL (ANTIBODY-MEDIATED) IMMUNITY

 

This response is carried out by antibodies, which are produced by B lymphocytes activated into plasma cells. B cells (B lymphocytes) develop from stem cells located in red bone marrow. After maturation in the bone marrow, the B cells migrate to lymphoid tissue such as lymph nodes or the spleen. Each B cell is programmed to recognize a particular antigen and has specific antigen receptors located on the surface of its plasma membrane. When the antigen for which the B cell is specific comes along, with the assistance of T cells, the B cell will react in two ways:

·         It will divide into a clone of many cells specific to the same antigen as the original B cell

·         The clone of cells will change into antibody-secreting plasma cells

 

The body is constantly producing lymphocytes, about 100 million each day, and that means about the same number must die. This is done by means of apoptosis, or programmed destruction. B cells that do not soon meet their antigen will undergo apoptosis, and the remains will be cleaned up by phagocytes.  Fortunately, the body is constantly producing more, so we always have at least a few B cells specific to any antigen that we might encounter.

 

Remember, each B cell is specific to some particular antigen, and can respond only to that one antigen. Each mature B cell has a large number of antigen receptors attached to its surface in the form of IgMs and IgDs all specific to a certain antigen. When the specific antigen comes along and binds to the antigen receptors on the surface of the B cell, the B cell responds by dividing and producing a large number of identical cells. This increase in number is proliferation. The group of cells is called a clone and they are all specific to the same antigen. These B cells next differentiate into plasma cells and start secreting antibodies against that specific antigen. If we ever encounter an antigen for which we do not have B cells (and also T cells) preprogrammed, we would be unable to produce an immune response.

 

Before going in to the steps involved in activation of B cells, we need to take a look at 2 things involved in the immune response. One is MHC antigens. These are proteins unique to each person. They are used by the immune system to distinguish self from non-self. They come in 2 classes:

     MHC-I—these are found on the outside of all nucleated body cells

     MHC-II—these are found only on the outside of antigen-presenting cells

The other thing is cytokines. These are small protein hormones involved in regulation of various activities within the body. Many of them are used for communication between white blood cells. One group especially important for this purpose is the interleukins.

 

Steps in activation of B cells:

 

Activated B cells which have differentiated into plasma cells begin to secrete large numbers of antibody molecules, living only a few days but producing as many as 2000 antibody molecules per second while active. A few B cells of the clone do not become active plasma cells, but instead become memory B cells which can recognize and respond to the same antigen if it comes along again, producing antibodies much faster and more effectively with a subsequent encounter, and providing long-term immunity.

 

This series of steps describes the processing of antigens that originate from outside the cell—exogenous antigens. It is also a description of production of antibodies against T-dependent antigens. Most antigens are of this type, and these are always proteins.

 

A few antigens are T-independent antigens. These are polysaccharides or lipopolysaccharides, and bacterial capsules would be an example. With these, the antigen binds directly to receptors on the B cell and does not enter the cell. This produces a weaker response and the antibodies are IgM’s. No memory cells are formed. This type of response seems not to occur in babies under the age of 2.

 

It could be said that the antigen selects the lymphocyte that responds to it, and the process is called clonal selection. It is believed that we are all prepared to respond to as many as 100 million different antigens, which means that we have that many different types of B cells, probably just a few specific to each antigen, waiting for it to come along.

 

Since human cells have the ability to act as antigens, we must have self-tolerance to our own cells and macromolecules. That means, in practical terms, that we do not produce an immune response to parts of ourselves, but if that same part were removed and put into another person, the recipient’s body would respond and try to destroy the foreign tissue. The immune system must be able to differentiate between self and nonself. Although we are not entirely sure how all this works, this is the current theory. Apparently we all start out with B and T cells that would react to our own tissues, but during fetal development, all B and T cells programmed to respond to self are destroyed and never reach maturity, probably in the thymus gland. This is called clonal deletion---we delete all cells that would otherwise produce an immune response against our own tissues.

 

 

ANTIGEN-ANTIBODY REACTION

 

When a matching antigen and antibody come together, an antigen-antibody complex forms. The antibody binds to the antigen at the antigen-binding site and marks the antigen for destruction by phagocytes and complement. The antibody itself does not damage the antigen, but because of the bonding the end result, by one means or another, should be the lysis of the foreign cell and inflammation. Here are the ways this may come about:

   1. Agglutination---antibodies cause antigens to clump together, making it possible for phagocytes to take in more than one of the antigen at a time. This reaction is also used in diagnosis of some diseases and blood typing.

   2. Opsonization---the antigen is coated with antibodies that enhance ingestion and lysis by phagocytic cells.

   3. Neutralization---IgG antibodies inactivate viruses by blocking their attachment to host cells, which is a necessary step in invasion of a cell by a virus. IgGs can also neutralize bacterial toxins by blocking their active sites.

   4. Antibody-dependent cell-mediated cytotoxicity---antibodies coat the outside of an organism and attract nonspecific immune cells that attack the organism from the outside. This is used against a large organism such as a parasitic worm, that cannot be phagocytized.

   5. Inflammation

   6. Activation of the complement system---both IgGs and IgMs can trigger the complement system. This contributes to inflammation, which tends to coat microbes in the area with reactive proteins which further encourage the activity of MACs. As the microbes are lysed, this attracts more phagocytes and immune cells to the area.

 

Unfortunately, the action of antibodies can also damage the host.

·         Immune complexes of antibody, antigen, and complement can damage host tissue, even when the antigen has been inactivated

·         Allergic reactions can be anything from unpleasant to fatal

·         Somehow, in spite of safeguards, antibodies that react with host tissue are produced (autoimmune diseases) and result in destruction of that tissue

 

 

IMMUNOLOGICAL MEMORY

 

The level of antibodies in the serum against a particular antigen can be measured---this is called antibody titer. If a person has never encountered an antigen, there are no antibodies, and for several days after the first encounter this is still the case. Then antibodies begin to appear and the level slowly rises. The first antibodies are IgMs, then IgGs begin to appear. This is called the primary response to the antigen.

 

If the same antigen is encountered again, the response will be quite different. This is called the secondary response, the memory response, or the anamnestic response. It is due to memory B cells specific to the antigen and causes a sharp, rapid rise in the level of IgG antibodies, which often prevents the development of the disease at all.

 

 

 

 

MONOCLONAL ANTIBODIES

 

In addition to body defense, antibodies are also very helpful in diagnosis of disease. For some time, the procedure for obtaining them involved injection of the antigen into an animal, which stimulated the animal to produce antibodies which could be recovered from the serum. This did not result in great amounts of the antibody, and it was difficult to isolate one particular antibody from other antibodies in the serum.

 

In 1984, three immunologists received the Nobel Prize for their discovery of a method by which a B cell could be treated so that it would divide indefinitely when removed from the body and grown in tissue culture. This involved combining a cancerous B cell, which would divide indefinitely, and a normal B cell which would produce the antibody desired. This resulted in a hybridoma, which would grow and produce antibodies indefinitely. Since all the antibodies come from clones of a single cell, they are called monoclonal antibodies. They are important because:

·         They are all identical

·         They are highly specific

·         They can be produced in very large numbers

 

This leads to the following uses:

·         Readily available for use in diagnostic tests, including tests for “strep throat, “ pregnancy tests, and tests for the presence of chlamydia

·         Used to suppress T cell activity and control rejection of transplanted tissue---antibodies that combine with the specific T cells fighting the transplant reduce their activity

·         Monoclonal antibodies targeted against cells of a cancerous tumor might attack the tumor

·         Also there is the idea that antibodies specific to the cells of a cancerous tumor might be combined with a toxin. The combination might selectively kill the cancer cells with the toxin instead of or combined the immune response, with few side effects.

 

 

CHEMICAL MESSENGERS: THE CYTOKINES

 

In the middle of the last century (1900s) immunologists began to realize that fluids containing immune cells also contained some sort of soluble chemicals that allowed the various cells to communicate with each other. These factors are now better understood (but still not completely), and they have been given the name cytokines. We have identified more than 60 of these. Interleukin is a name for cytokines that allow white blood cells to communicate with each other.

 

 

CYTOKINE

ACTIVITY

INTERLEUKIN-1 (IL-1)

STIMULATES T CELLS IN PRESENCE OF ANTIGENS; ATTRACTS PHAGOCYTES IN INFLAMMATORY RESPONSE

INTERLEUKIN-2 (IL-2)

PROLIFERATION OF HELPER T CELLS; PROLIFERATION AND DIFFERENTIATION OF B CELLS; ACTIVATION OF CYTOTOXIC T CELLS AND NATURAL KILLER CELLS

INTERLEUKIN-8 (IL-8)

ATTRACTS PHAGOCYTIC AND IMMUNE CELLS TO AREAS OF INFLAMMATION

INTERLEUKIN-12 (IL-12)

DIFFERENTIATION OF HELPER T CELLS

GAMMA INTERFERON (g-IFN)

INHIBITS VIRAL REPLICATION; INCREASES ACTIVITY OF MACROPHAGES AGAINST MICROBES AND TUMOR CELLS

TUMOR NECROSIS FACTOR (IFN-b)

KILLS TUMOR CELLS; ENHANCES ACTIVITY OF PHAGOCYTES

GRANULOCYTE-MACROPHAGE COLONY-STIMULATING FACTOR (GM-CSF)

STIMULATES FORMATION OF RED AND WHITE BLOOD CELLS FROM STEM CELLS

 

 

 

T CELLS AND CELL-MEDIATED IMMUNITY

 

This is based on the activity of certain specialized T cells. This type of immunity is not transferred across the placenta, and occurs mostly in response to intracellular pathogens. T cells (T lymphocytes) are the key to cell-mediated immunity. They develop in the bone marrow, but leave it while still in an immature form and migrate to the thymus gland. There, influenced by thymic hormones, they reach maturity. They then migrate to lymphoid organs of the body.

 

Like B cells, each T cell is programmed to respond only to one certain antigen because it has antigen receptors on its surface only for that particular antigen. When activated by contact with its antigen, the T cell also proliferates and differentiates. This is also known as clonal selection. Also like B cells, some of the activated T cells become memory cells, able to respond quickly to a subsequent encounter with their particular antigen.

 

 

TYPES OF T CELLS

 

There seem to be 3 main types of functional T cells:

·         Helper T cells (TH cells)

·         Cytotoxic T cells (TC cells)

·         Regulatory T cells (TR cells)

 

T cells can also be classified according to which type of cell-surface receptor of the group known as the CD group is found on the outer surface of the plasma membrane. All T cells will have either CD4 proteins or CD8 porteinss. It appears these proteinss are added while the T cell is maturing in the thymus gland.  Cells with CD4 proteins (sometimes called CD4 T cells) become helper T cells. CD8 T cells become cytotoxic and suppressor T cells. The AIDS virus devastates the immune system by destruction of the CD4+ T cells. As we will soon see, this type of T cell is required for both antibody-mediated and cell-mediated immunity to proceed.

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

T cells do not recognize their particular antigen unless it is displayed on the surface of an antigen-presenting cell (APC). The main APCs are macrophages and dendritic cells. The APC ingests the antigen, breaks it up into pieces inside the cell, and combines the pieces of antigen with proteins belonging to the cell. The cellular proteins are components of the major histocompatibility complex (MHC) proteins. Every individual has his own unique MHC proteins, which are the part of the cell that the immune system recognizes as self (or nonself, in the case of transplanted tissues). They are displayed on the outside of the plasma membrane of all nucleated cells of the body, (which means that red blood cells do not have them).

 

   1. Helper T cells---these are of major importance to the immune response. By releasing cytokines, they:

·         Induce the formation of cytotoxic T cells

·         Activate macrophages

·         Stimulate B cells to become plasma cells and secrete antibodies

 

These are the steps in activation of helper T cells:

      1) A macrophage or dendritic cell takes in the antigen and processes it. The antigen is broken into bits, and some of the bits are combined with cellular proteins called MHC proteins. The combination of antigen/MHC protein starts to leave the cell by exocytosis, but remains stuck to the outside surface of the plasma membrane. The macrophage or dendritic cell is now an antigen-presenting cell (APC).

      2) A CD4 T cell recognizes its antigen-MHC-II complex as presented by the APC—this is the initial signal for activation of the T cell

      3) A second signal called a costimulator is required for full activation—this is often in the form of a cytokine released by the APC

      4)Activated helper T cell begins to proliferate and secrete cytokines of its own

     5) It differentiates into clones of TH 1 and TH2 cells as well as forming memory cells

      6) All of these are specific to the original antigen

 

TH1 lymphocytes mainly activate macrophages, CD8 cells, and NK cells.

 

TH2 lymphocytes are mainly involved in allergic reactions and response to certain parasites.

 

   2. Cytotoxic T LYMPHOCYTES (ctls) are what cytotoxic T cells (CD8 cells) become when they are activated.

 

Steps in activation:

     1) CD8 cell recognizes its specific antigen combined with MHC-I proteins. (Review: the MHC-1 proteins are found on all nucleated body cells, not just the APCs as MHC-II proteins are. The fact that the antigen is combined with MHC-I proteins is a sign that the antigen is endogenous, not exogenous, so dealing with it is a job for the T cells.)

     2) Receptors on T cell combine with antigen-MHC-I complex

     3) Helper T cells, also specific to the same antigen, release cytokines required for activation of CD8 T cell

     4) Activated cytotoxic T cell is now called a cytotoxic T lymphocyte (CTL)  and goes to work

 

They are able to destroy target cells on contact. Intracellular pathogens cannot be attacked by antibodies. This includes all viruses, except during the time they are free in tissues or fluids while preparing to invade a cell, and some bacteria.   (Most bacteria cause their harm from outside the cell, but some go inside.)  The T cell operates as follows:

      1) A cytotoxic T lymphocyte specific to the antigen binds to the MHC-I-antigen complex on the surface of the infected cell

      2) Cytotoxic T lymphocyte releases a chemical called perforin

      3) Perforin forms a pore (opening) in the plasma membrane of the target cell

      4) CTL next releases granzymes, which enter the target cell through the pores and induce “cell suicide” (apoptosis)

      5) Remains are cleaned up by a phagocyte

      6) Cytotoxic T cells also attack some cancerous cells in the same way

 

Remember, once a cell has been invaded by the pathogen, that cell is generally doomed. Particularly if the pathogen is a virus, it will cause the death of the cell anyway. If the cell can be destroyed before a large number of new viruses are produced inside and released, that is to our advantage.

 

   3. REGULATORY T cells—these are not well understood. Many immunologists believe that these cells turn off the immune response when it is no longer needed. Others think they don’t even exist, and that TH cells and TC cells change and suppress the immune response at the appropriate time. However they originate, these cells release the cytokine IL-10

 

 

NONSPECIFIC CELLULAR COMPONENTS

 

There are other cells that enter into cell-mediated defense of the body. These include activated macrophages and natural killer cells.

 

   1. DENDRITIC CELLS—these are formed in red bone marrow and travel to locations such as lymph nodes, the spleen, and the skin. They are probably the main antigen-presenting cells. 

 

   2. Activated macrophages---although macrophages are always at least  somewhat phagocytic, this tendency can be greatly increased when they become activated. Activation may occur due to ingestion of antigenic material, or by the release of cytokines from helper T cells. Activated macrophages enlarge and become ruffled. They are especially important because of:

·         Ability to eliminate certain virus-infected cells

·         Ability to eliminate pathogenic intracellular bacteria

·         Ability to attack and destroy many cancer cells

·         Ability to act as APCs

 

   3. Natural killer cells (NK cells)—these are lymphocytes that are neither B cells nor T cells. They are not specific to particular antigens. They check body cells to see that they display the correct MHC-I proteins and no abnormal proteins. If abnormal proteins are present, NK cells are able to attack and destroy the same types of cells as the cytotoxic T cells of the immune response.

 

 

RELATIONSHIP BETWEEN THE TWO BRANCHES OF THE IMMUNE RESPONSE

 

The two branches of the immune response are closely interrelated, in several ways:

 

   1. Production of antibodies---the production of antibodies against many antigens requires the assistance of helper T cells. These antigens are called T-dependent antigens. -producing  plasma cells.

 

   2. Antibody-dependent cell-mediated cytotoxicity—defensive cells can be stimulated to kill organisms such as protozoa and helminths that are too large for phagocytosis. Steps:

      1) Target cell is coated with antibodies, arranged so that their Fc regions (stem of Y)  are pointed outwards

      2) Natural killer cells, macrophages, neutrophils, and eosinophils bind to these Fc regions

      3) All of these defensive cells secrete substances that lyse the target cell