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uorllnporlul

plement

also

provides

a means

of attracting

macrophages, which scavenge

the target cells

or their

products.

Alternatively, the antigen-

antibody

complex may

be taken up directly by

macrophages

(scavenger

cells)

and destroyed.

The cell-mediated response is executed

by a class of T lymphocytes called

cytotoxic

T

cells

(also

called

killer T

cells). The basic func-

tion of the T cell

in

recognizing a target antigen

is indicated in r;r..,iri'rir

l:;r.li.

A

cell-mediated

response typically is elicited by an intracellular

parasite,

such as a virus that infects the body's

own cells.

As

a

result

of the viral infection, frag-

ments of foreign

(viral)

antigens are displayed

on the surface

of the

cell.

These

fragments are

recognized by the T cell receptor

(TCR),

which

is the T cells' equivalent of the antibody

pro-

ducedbyaBcell.

A

crucial

feature

of this

recognition reac-

tion

is

that

the antigen must

bepresented by a cel-

lular

protein

that is a member of the IVIHC

(maj

or

histocompatibility complex). The MHC

pro-

tein has a

groove

on

its

surface that binds a

pep-

tide

fragment derived from the foreign antigen.

The combination of

peptide

fragment

and

MHC

protein

is recognized by the T cell receptor. Every

individual has a characteristic

set of

MHC

pro-

teins. They are

important

in

graft

reactions; a

graft

of tissue

from

one

individual

to another

is

rejected because of the difference in MHC

pro-

teins between

the donor and recipient, an issue

of

major medical importance. The

demand

that

the T lymphocytes

recognize

both foreign anti-

gen

and

MHC

protein

ensures

that

the cell-

mediated response acts only on host cells that

have

been

infected with a foreign antigen.

(We

discuss the division of

MHC

proteins

into the

general

types of class

I

and class

II later in Sec-

tion

2).20, The Maj or Histocompatibility Locus

Codes for

Many

Genes of the

Immune

System.)

The

purpose

of each type of immune

response is to attack a foreign target.

Target

recognition is the

prerogative

of B-cell

im-

munoglobulins

and T

cell

receptors. A crucial

aspect of their

function lies in

the ability

to dis-

tinguish

"self"

from

"nonself."

Proteins and cells

of the body

itself must never be attacked. For-

eign targets

must

be destroyed entirely.TLre

prop-

erty

of failing to attack

"self"

is

called tolerance.

Loss

of

this ability results in an autoimmune

disease,

in

which

the immune system attacks

its

own

body, often with disastrous conse-

quences.

What

prevents

the

lymphocyte

pool

from

responding to

"self"

proteins?

Tolerance

prob-

ably arises early

in

lymphocyte

cell develop-

ment when

B cells and

T cells that

recognize

"self"

antigens

are

destroyed.

This

is called

clonal deletion.

In addition

to this

negative

selection, there

is also

positive

selection

for

T

cells carrying

certain

sets

of T cell

receptors.

A

corollary

of tolerance

is that

it

can

be dif-

ficult to obtain

antibodies

against

proteins that

are closely related

to those

of the organism

itself.

As

a

practical

matter,

therefore,

it

may be diffi-

cult to use

(for

example)

mice

or rabbits

to

obtain antibodies

against

human

proteins

that

have been

highly conserved

in mammalian

evo-

lution. The tolerance

of

the mouse

or

rabbit for

its

own

protein may extend

to the

human

pro-

tein in such cases.

Each of

the three

groups

of

proteins

required for

the

immune

response-im-

munoglobulins,

T cell

receptors,

and

MHC

proteins-is

diverse.

Examining

a Iarge

num-

ber of

individuals,

we

find many

variants

of

each

protein.

Each

protein is coded

by a large

family

of

genes; in the case

of antibodies

and

the T cell

receptors,

the

diversity

of the

i:il.i"l[{f

i]jl.i.r

In cet[-mediated

immunity,

k'itter

T

cel.l.s

use the

T ce[[ receptor

to

recognize

a

fragment of

the

for-

eign antigen

that

is

presented

on

the surface

of the

tar-

get

cetl

by the

MHC

protein.

23.1

Introduction

573

population

is increased

by DNA

rearrangements

that

occur in the relevant

lymphocytes.

Immunoglobulins

and T

cell

receptors

are

direct

counterparts,

each

produced

by its own

type

of lymphocyte.

The

proteins

are related

in

structure,

and their

genes

are related in

organization.

The

sources of variability

are sim-

ilar.

The MHC

proteins

also

share some

com-

mon

features

with the

antibodies,

as do other

lymphocyte-specific

proteins.

In

dealing with

the

genetic

organization

of the immune

sys-

tem,

we

are therefore

concerned

with a series

of related gene

families,

indeed

a superfam-

ily

that may have

evolved

from some

common

ancestor representing

a

primitive

immune

resDonse.

:.:i.r-:i;ir

.l:i-:i

The

pooL

of immature

[ymphocytes

contains

B

ce[[s

and T

cetLs making

antjbodies

and receptors

with

a

variety

of specificities.

Reaction

with

an antigen

leads

to

clonaI

expansion

of the

lymphocyte

with the

antibooy

(B

cett)

or receptor (T

cetl) that can recognize

the antigen.

CHAPTER

23 Immune

Diversitv

C[onaL

Selection Am

plifies

Lymphocytes

That

Respond to IndividuaL

Antigens

r

Each B

lymphocyte expresses

a single

immunoglobutin

and each T

lymphocyte

expresses

a singte T

cetl receptor.

.

There

is a very [arge variety

of immunoglobutins

and

T

ce[[ receptors.

e

Antigen

binding to an immunogtobutin

or T ce[[

receptor

triggers cLonal. multiptication

of the cet[.

The

name of

the

immune

response

describes

one of its

central features. After

an

organism

has

been exposed

to an antigen,

it becomes

immune

to the effects of a new

infection.

Before

exposure

to a

particular

antigen,

the organism

lacks

adequate

capacity to

deal with

any toxic

effects. This

ability is acquired

during the

immune response.

After

the infection

has

been

defeated, the

organism retains

the

ability to

respond

rapidly in

the event

of a reinfection.

These features

are

accommodated

by

the

clonal selection

theory illustrated

in

F:Iili-Jiis f

i"5. The

pool

of lymphocytes

contains

B

cells and T

cells carrying

a large

variety

of

immunoglobulins

or T

cell receptors.

Any

indi-

vidual B

lymphocyte

produces

one immunoglobulin,

however,

which

is capable of recognizing

only a

sin-

gle

antigen;

similarly, any individual

T

lymphoqtte

produces

only

one

particular

T

cell receptor.

In

the

pool

of immature

lymphocytes,

the

unstimulated

B cells and

T cells

are morpho-

Iogically

indistinguishable.

On exposure

to anti-

gen,

though,

a B cell whose

antibody

is able

to

bind the antigen,

or a T

cell whose

receptor

can

recognize

it,

is stimulated

to divide, probably

by some feedback

from

the surface

of

the cell,

where the

antibody/receptor-antigen

reaction

occurs.

The stimulated

cells

then

develop into

mature

B or T lymphocytes;

this includes

mor-

phological

changes

involving

(for

example)

an

increase

in cell

size

(especially

pronounced

for

B

cells).

The initial

expansion

of a

specific

B or T

cell

population

upon first

exposure

to

an antigen

is

called

the

primary

immune

response.

Large

numbers

of B

or

T

lymphocytes

with

specificity

for

the target

antigen are

produced.

Each

pop-

ulation

represents

a clone

of the

original

responding

cell.

Antibody

is secreted

from

the

B cells in

large

quantities,

and it

may even

come

to dominate

the

antibody

population.

@ws

574

919

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