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Molecular Biology of the Cell by Bruce Alberts, Alexander Johnson, Julian Lewis, David Morgan, Martin Raff, Keith Roberts, Peter Walter by by Bruce Alberts, Alexander Johnson, Julian Lewis, David Morg

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OVERVIEW OF THE ADAPTIVE IMMUNE SYSTEM

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(A) resting T or B cell

1 µm

(C) effector T cell

1 µm

(B) effector B cell (plasma cell)

hematopoietic system, including erythrocytes, leukocytes, and platelets (see Figure

22–32).

Because they are sites where lymphocytes develop from lymphoid progenitor

cells, the thymus and bone marrow are referred to as central (primary) lymphoid

organs (see Figure 24–12). As we discuss later, most B and T cells die in

the central lymphoid organs soon after they develop, without ever functioning.

Others, however, mature and migrate via the blood to the peripheral (secondary)

lymphoid organs—mainly the lymph nodes, spleen, and epithelium-associated

lymphoid tissues in the gastrointestinal tract, respiratory MBoC6 tract, m25.07/24.15 and skin. It is in

these peripheral lymphoid organs that foreign antigens activate B and T cells (see

Figure 24–13).

B and T cells become morphologically distinguishable from each other only

after antigen has activated them: resting B and T cells look very similar, even in

an electron microscope (Figure 24–14A). After activation by an antigen, both proliferate

and mature into effector cells. Effector B cells secrete antibodies; in their

most mature form, called plasma cells, they are filled with an extensive rough

endoplasmic reticulum that is busily making antibodies (Figure 24–14B). In contrast,

effector T cells (Figure 24–14C) contain very little endoplasmic reticulum

and secrete a variety of cytokines rather than antibodies. Whereas B‐cell-derived

antibodies are widely distributed by the bloodstream, T‐cell-derived cytokines

mainly act locally on neighboring cells, although some are carried via the blood

and act on distant host cells.

1 µm

Figure 24–14 Electron micrographs of

resting and effector lymphocytes.

(A) This resting lymphocyte could be

either a B cell or a T cell, as these cells

are difficult to distinguish morphologically

until antigen activates them to become

effector cells. (B) An effector B cell (a

plasma cell). It is filled with an extensive

rough endoplasmic reticulum (ER), which is

distended with antibody molecules that are

secreted in large amounts. (C) An effector

T cell, which has relatively little rough ER

but is filled with free ribosomes; it secretes

cytokines, but in relatively small amounts.

The three cells are shown at the same

magnification. (A, courtesy of Dorothy

Zucker-Franklin; B, courtesy of Carlo

Grossi; A and B, from D. Zucker-Franklin

et al., Atlas of Blood Cells: Function and

Pathology, 2nd ed. Milan, Italy: Edi. Ermes,

1988; C, courtesy of Stefanello de Petris.)

Immunological Memory Depends On Both Clonal Expansion and

Lymphocyte Differentiation

The most remarkable feature of the adaptive immune system is that it can respond

to millions of different foreign antigens in a highly specific way. Human B cells, for

example, collectively, can make more than 10 12 different antibody molecules that

react specifically with the antigen that induced their production. How can B cells

and T cells respond specifically to such an enormous diversity of foreign antigens?

The answer for both B and T cells is the same. As each lymphocyte develops in

a central lymphoid organ, it becomes committed to react with a particular antigen

before ever being exposed to the antigen. It expresses this commitment in

the form of cell-surface receptors that specifically bind the antigen. When a lymphocyte

encounters its antigen in a peripheral lymphoid organ, the binding of the

antigen to the receptors (with help from co-stimulatory signals, discussed later)

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