THE HUMAN I]ITAIIN

A GuidedTlrtrr

SUSAN GREENIIIIiI,I)

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P HCENIX

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Ffow does the brain work? What does it actually

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rrrade to 'enhance

public

more

life have led

such as depresdependence

\Me are faced with. an

as we carr about the

brain. On July L7, 1990, the then president

.io r'yt.r'?et.t

better. The

illnesses of old age that attack the brain, such as

to a huge increase in psychiatric

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in human understanding.

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do? These

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At last, however,

to tackle wh-at might

tl e final frontier

devastating

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\tr IT H IN

of the United

that every effort should be

awareness

of the benefits

derived from brain research.' \4/e are currently

to be

in the middle

of the 'Decade of the Brain.' A general inlerest in the brain is

official.
Remote from the rest of the bodv in its own custom-built

THE

H U MAN

BRAIN

B R A IN S

W ITI.' IIN

B R A IN S

casing of skull bone, the brain has a consistency similar to a

the body into the brain. Btrt if t-[re].rrain was the centre for

soft-boiled egg and has no intrinsic moving parts. Thus, it is

thinking, what about the soul?

obviously

not

participate

in

destined to take any physical

strain or

The Greek physician Galen (x.o. a29-199) pointed to a

actions. The

part of the brain that was the least solid, the most ethereal,

Greeks crme to the conclusion that this insubstantial and

and clearly discernible to the naked eye. Deep within the

brain is a labyrinth of interconnecting cavities, formed

any large-scale mechanical

secretive substance was the perfect site for the soul. Most
importantly, soul was immortal: it had nothing to do with

during development in the womb and containing a colour-

thinking.

In fact, all the abilities we now attri.bute to the

brain, the Greeks localized in the heart or the lungs (there

less fluid. This insubstantial-seeming fluid bathes the whole

of the outer surface of the brain and spinal cord and is

was never total agreement about the precise location). The

immortal 'soul' was of course so sacred and elusive t].at the

known as cerebrospinal fluid (CSF). It is often used in the.

diagnosis of various neurological problems when sampled

silent, remote grey home provided

from the lower portion

for it by the brain

presented a serious focus with alrnost mystical properties:

the Greeks imposed strict taboos against eating the brains of
any animal. The soul in this case was quite explicitly a

of the spinal cord in a lumbar

puncture. Norma}ly, however, the CSF is reabsorbed into the

blood supply, so that fresh fluid is constantly being produced (about o.2 rnillilitres

per minute in humans) and is

different entit5r from 'consciousness' and the 'mind' and all

the other interesting properties that we now associate with

thus constantly circulating.

our individuality

substance, as opposed to ttre sluggish slurry of brain, might

have been a good candidate for the substance of the soul. We

and personality.

Such quirky reasoning, where normal mental activities

were not associated at all with the brain, changed with a

It is easy to imagine

how

this

mysterious

swirling

great discovery made by Alcmaeon of Croton. Alcmaeon

now know tl at the CSF contains merely salts, sugars, and

certain proteins. Far from being the seat of the soui, it has

showed that there were actual connections leading from tle

eyes to the brain. Surely, he claimed, this area must be the

even been disparagingly referred to as 'the urine of the

brain.' No one, even those who believe in an immortal soul,

seat of thought.

now expects to find it in the brain. The mortal brain, which

This

revolutionary

idea tied

in

with

observations made by two Egyptian anatomists, Herophilus

and Erasistlatus, who managed to trace nerves - obviously

everyone regards as obviously

not identified as such at the time - leading from the rest of

of riddles.

responsible

for

all

our

thoughts and feelings, presents in itself the most tantaiizing

THE

HUMAN

BRAIN

B R A IN S

\^/ITH IN

B R A IN S

In this book ure are going to see how far we can progress
with

answering

the question: FIow does the brain work? But

this question is far too global and vague to have any meaning
of actual

in terms

experiments

or observations.

What

we

need to do instead is to tackle more specific sub-questions

that nonetheless

make a contribution

to a final understand-

ing of this secretive mass of tissue in which,

essence of our personalities
The first

topic

we shall

somehow,

the

is locked away.
explore

in this

chapter

is the

j$e

appearance of the brain. Imagine you were looking

physical

at a brain in your hands: what you would

be a creamy coloured, wrinkled
kilograrn,

be holding

object weighing

on average about 1.3 kilograms.

The first feature you would

would

just over 1

(See Figure 1-.)

notice is that the strange-looking

object, small enough to be supported in one cupped palm, is

made up of distinct

regions of a particular

that fold around each other and interlock

shape and texture

according

to discern.

grand scheme that we are only now beginning

The brain has the consistency
ground

plan that is always

to some

of a raw egg with an overall

the same. There are two clear

ii&*
/,,3,{i11'
,*rta$l

halves, called hemispheres, that seem to sit around a kind of

thick

stalk (brain stem). Tiris brain stem eventually

clown into

the spinal

tapers

cord. At the back is a cauliflower-

shapod extrusion, a 'little brai.n' (cerebellum) that protrudes

prer:ariously behind

the main brain (cerebrum).

If yotr vvere to look at the cerebellum,

the srrrface of these hemispheres,

the brain stern, and

you would

see that they

FISURE
'
A cross-section
of the human brain within the head. Note how the brain is
composed of different gross regions easily discernible to the naked eye, and
how it is continuous with the spinal cord. (From M. A. England andJ. wakely,
A colour Atlas of the Brain and spinal cord, wolfe publishing, 19s1.)

THE

H UM AN

B R A IT.I :

BRAIN

are ali diiTerent in surface texture, as weil as varying slightly

in colour

along the cream--pink-brown.

more,

you

if

underside,

turned

it would

the

brain

over

and

be easy to see still

regions again distinguishable

Further-

spectrum.
looked

further

at the
different

by colour, texture, and shape.

For the most part, each region is duplicated

on either side of

the brain so that you could draw a line down the middle
an axis, about which
The different

the brain would

ordered anatornical

these boundaries

the stalk-

up by neuroscientists

in an

scheme. One way to think of these brain

regions is as countries

filled ventricle

be syrnmetrical.

regions of ihe brain pile around

like brain stem and are divided

as

distinguished

are very

obvious:

by boundaries.
one rnight

Often

be a fluid-

where we saw the soul was once thought

to

vis;iou \tiir$ [haL thet nervous system rvas

cord' wilh
like an invertecl tr-ee. The trunk was in the spinal
the nerves
the roots in the brain ancl the branches in
the flrst part
extending throughout ttre bocly' A little later' in
Flourens also
of the eighteenth century, Jean-Pierre-Marie
the results
conclrrcled that the brain was homogeneous' frorn

gland. Malpigiii's

of his rather

experiments'

ghoulish

animals'

variety of laboratory

rather

than

impaired.

selective

With
functions

functions

undeniable

selective parts of the brain.

different

name, but we will

example, cerebellum

be gathering

or brain stem) only as we need them.

Rather than a cletailed ciocumentation

brain,

our prime

regions

contribute

such labels (for

concern

of the anatomy of the

here is to discover

to the feats of sun'ival

how

in the outside

world as well as to consciousness of that inner world,

most private
have tantalized

place of thoughts

certain

one's

and feelings. These issues

people since long before the dawning

of the

Decade of the Brain.

For some, such as Marcello
century,

the brain

functioned

Malpighi

in the seventeenth

homogeneously,

as a huge

logic,

becoming
Flourens

could. not be localized

According

has a

on a
more

andrnoreoftheirbrainsandobservingtheeffects'Whathe
weaker'
found. vr.as that all function$ grew progressively

distinct

scheme, each region

removing

methodically

employed

\rerysimplerationale:taremoveclifferentpartsofthebrain
and see what functiorrs remained' He experimented

lurk, another might be subtly changed in texture or colour.

to tkre recognized

Flourens

This scenario of a uniform

inspirecl the concept

brain with

of mass action'

specifically
decided

that

as such within

no specialist

parts

It is an idea that still

to explain a
persists today, in a less extreme form, in order
but fairly frequent occurrence: when
seeming miraculous
a
parts of the brain are destroyed, say by stroke' then,after
that at }east
while, other intact parts appear to take over' so
some of the original function is rq'stored'
contrast to this idea is tfre view that the brain
a highty"
can bo divicled into rigid compartments' each with
this,-vision
specific function' The most famous proponent of
In complete

Gall'yras
i /as Franz Gall, a cloctor born in Vienna in L758'

IO

THE

HUMAN

BRAIN

very interested

t,

time,

in the human mind, but he considered

to probe

delicate

B R A IN S

he was probably

GaIl hit upon

Instead,

right.

the skulls of the

the theory that if he studied

He developed

t,

characters of those people, then perhaps he could identify

I
I
I
I
I
I
I

up were the most easy feature to detect: the bumps

t
I \'

up with

dead and then saw how these matched

physical

trait

with

corresponded

that

the alleged
a

aspects of

certain

character. The aspects of the brain tl at Gail chose to match

on the

concluded

that

were

there

character traits. These purported

different

twenty-seven

blocks of person-

building

ality actually

turned

out to be rather sophisticated

of the human

mind:

instinct

and friendship,

attachment

offspring,

oneself and one's property, instinct

defensive

ing

of

to steal, pride and

individual.

nremory

for people, sense of words,

sense of the spoken

The nagging

were

small

and still

a specific

a physical

displaced
they

by the bumps

were

pushed

particular

pattern

somewhat

primitive

to pierce

of perforations
readout

Caspar Spurzheim,

|ohann
the

upward

r and sarcasm, poetic

irnit

umdfrllt'
,nfim-''"

God and religion,

e different

spirit, sense of

talent, goodness, faculty

of

steadfastness.

qualities

- which

were

eventually

mental

each

state might
let alone

on the skull.

the procedure

'the

study

at the brain,

on objective measurements,

true science - as such it rapidly

with

a new

The

character.

of the mind,'

and its underlying

became

paper.

one of Gall's colleagues, coined

a new way of looking

Phrenologv

through

of an individual's

Greek terrn pftrenology,

describe

pins were

in the paper thus gave a

captured

popular

basis for morality,

philosophy.

to
It

and because it

it had all the lr:stre of a

present people with a more 'scientific'

wisclom, depth of thought and metaphysical

in

on the surface of the skull so that

rolationship

comparative

the

question

infrastructure,

times.

sense of mechanics,

or large

unanswerable

worcl, sense of colour, sense of tonal relations, sense of the

of numbers,

where

of hat. When placed on the skull, movable

relied

and

- a

The apparatus that Gall used to make his enalyses was a

relations,

of things

for exarnple, banality

one as remote from brain tissue as a bump

facts, sense of spatial

memory

lumps

ever be associated with

offered

circumspection

the

B R A IN S

to greater or lesser degrees, accord-

was not even raised of how

foresight,

vanity,

to include,

were localized

to whether

and

authority,

of

instinct

for cruelty, cleverness,

feoling of possessiveness and inclination

love

features

love for one's

of reproduction,

to thirty-two

map of the surface of the head was produced,

kind

surface of the skull.

Gall

expanded

functions

the brain.

more subtle way of studying

seemingly

another,

quite

of the

the techniques

Given

surgically.

it too

W ITH l N

of the

because it seemed

to

approach as well as

something

measured and did not entail difficult

the spirit

that

could

be

and abstract ideas, like

soul. Seen as a secular, objective system, stripped

of any

need for bli.nd faith,

to the

phrenology

catered

superbly

tl

:,+19!&r:r,.!il

I7

THE

H U M At.!

growing

B i,-.i r' 1.;

f!R,AIN

number

of people disaffected

at tirat time with

the

church.

appeal. 'I'he problslrr lvas .roiirl)ounded wlien, a i.;w yoars

later, another

Anotller
of making

physician,

advantage was, of course, tl:at it was a new way

covered a different

large amounts

that Wernicke

of money: phrenology

books, and models all started to proliferate.

pamphlets,

Indeed phrenol-

ogy became an integral

:3

'. . : !-?-;i

different

dis-

Carl Wernicke,

the Austrian

In the patients

type of speech problem.

In this case the patient,

The only problem

in

zodiac, so in the last century

Words are jumtrled together in an incoherent sequence and

would

have a tiny,

personalized

handle. But eventually

tive endeavour

phrenology

this fascinating,

painless, and lucra-

was to run into trouble.

In 1861, in France, the neuroanatomist

Paul Broca examined
man could
words,

bust on the

and anthropologist

pronounce

any other

hence he was ref'erred to as 'Tan', even though

real name was Leborgne.

his

Tan earned his place in history

because six days after the examination

he had the misfor-

is t}.at the speech is often gibberish.

no apparent mean-

with

fr-equently new words are invented

ing at all.
The discovery

a man who was unable to speak. This

only say 'tan': he couldn't

aphasia

words perfectly.

unlike

Tan, could articulate

canes, for example,

of yet another

associated with, but linked

that the problem

shows

area of the brain,

was not

for phrenology

of a single

speech

centre

functions.

brain. It turned

on the skull as an index of brain function,

completely

out that the area of the brain damaged was

different

some phrenology
in the lower

from tllat predicted

by phrenology.

On

busts, the centre for }anguage is localized

part of the left eye socket, whereas

in Tan,s

problem

Irrespective

of how

of the absurdity

a cohesive behaviour,

thought is translated into a physical

was a simple one-to-one mapping

of the left-hand side of the brain. Henceforth tJ'is part of the

product -

brain became knornrn as Broca's orea.

single, small region. In retrospect

observations

unequivocal

such as these, phrenology

clinical

started to lose its

is not valid.

a complex

function

of measuring

brain
bumps

there is still the

skill,

sensation,

event somewhere

brain, and vice versa. The phrenologists

brain, the damaged area was a small region toward the front

Because it did not match up with

of location,

Bumps on the skull clearly do not represent different

tune to die, thereby giving Broca the chance to examine his

even a

observations

raised the even deeper issue that, irrespective

even the concept

clearly

aspect of, speech,

to a different

of the speech centre: Wernicke's

mislocation

thought

or

in the

that there

of the complete finished

such as language - with

it is easy to see that they

centres

of memorY,

ernotion, and so forth still persists in folklore

views of the

were

wrong,

II

part of many people's lives. Just as

today items ranging from mugs to jewellery bear signs of the
walking

I
rl

.studied, there was damage to a completely

part of the brain.

tVernicke's

although

the

idea

of

l;

r;

It

1l

it
F
T

t
t

.4

THE

HUMAN

BRAIN

B R A IN S

brain. But if chunks

and directly

of the brain are not rnerely passively

corresponcling

to chunks of the outside wcrld,

or chunks

of our behavioural

what kind

of alternative

the brain

most primitive
ing functions
hs'ce

Abnorrnal
now

then

(i.835-j.9j.1),

a British

neurolo-

as organized into a hierarchy.

that were increasingly

more sophisticated,

in humans.

The

rnovements

be interpreted

resulting

from

as an unleashing

lnvoluntary

rnovements

inlluencos.

similarly,

and even sociology.

brain

damage could

of lower

functions,

from their normal restraining

sigmund

and

This idea was to have

for neurology, psychiatry,

of the bumps

higher

Freud was able to refer to

psychiatric

or

Paul

Maclean

viewed

the

in

the

brain

more advanced'old
mammalian.'

The

also be interpreted

havlng escaped from a 'higher' controlling

Alttrough
llrovides

Hughlings-|ackson's

an interesting

llsychiatry,

and

even

force.

spinal

framework
trehaviour,

nssrrrnption rnade by the phrenologists

the old mammalian

behaviour,
rnamrnalian

region

but

this

time

reptilian,'

brain,

which

interconnecting

brain

the

corresp-

behaviour.

was constituted

middle

brain-level

system, which

particularly
brain

for instinctive

was the

controlled

area for rational

in the outer layer

from

from

emotional
the new
pro-

ttrought

of ttre brain.

derived

By

structures

aggression and sex. Finally,

is kno'r,vn as the cortex,

This

outer

the Latin

for

fcrr neurology,

'bark,' sirrce it covers the outer surface of the brain as does

the

the bark on a tree.

erroneous

lurks here also. The

Maclean

r:orrr;opt of a lrierarchy implies something must be at the top,

maintained

llrnl tlrero rnust be some ultimate

condition

controller.

Maclean

and the most sophisticated

reptilian

cord), was responsible

cesses housed

idea is appealing in that it

common
crowd

as

by

onded to the brain stem (tlle central stalk arising from the

as the limbic

mob could

the

the function-

1.95Os. Again,

of hierarchy,

mammalian,'

known

lorrr of an ungoverned

a physical

brain within

of three tiors: the most 'prirnitive

composed

'ne'w

1-94Os and

as a kind

of

behav-

have

in

ing of gross brain regions to each other was developed

series

brain, the anarchic

not

attempt at a scherne for interrelating

<:onscionce of the 'superego.' Finallv,

an individual

does

understandable

all operations.

contrast,

{rrona, far boyond

superego, while
terms,

t5

B R A IN S

bust. Alternatively,

as such. There is no mini-super

brain directing
Another

moral

tho passionate drives of the 'id' as being restrained by tl'e

'ogo' (consciousness), which was kept in check by the
everl in t]le political

on a phrenology

the idea of an ultimate

counterpart

drives were kept in check by higher restrain-

most developed

inrplications

repertoire,

scenario can be envisaged?

John Hughlings-Jackson
gist, viewed

and mental

redolent

W ITH IN

Flowever, the

irlun of a single executive centre for memory or movement is

referred

to his concept

as the triune

brarn and

that much of the conflict arising from the human

resulted

three tiers. Although

from

a poor coordination

this theory might

between

the

help us understand

E*dlidr'*

16

THE

H U M AN

the literally
political

rnindless and unifbrm

rallies,

it does little

localized

Nonetheless,

a comparison

such

as

humans,

might

provide

from

on the central

in.the outside world

are

in the brain.

species,

brains

light

reptiles,

different

of the brains

of the diflbrent

non-hurnan

mammals,

some clues to the puzzle.

animals,

the most obvious

and
In the r

feature is

then is that the size ,

that the bigger the brain the :

is all important,

more intelligent
An

tle

elephant's

brain

is five times bigger than

brain: it weighs about B kilograms,

elephant

is fir'e

Presumably
a lot

important

times

more

but would

intelligent

than

hurnans,

on its own

but

it might
rather

and a bird

a human

format

we say that an ,
than

not

be size that

the percentage

are
is I

of body

weight that is made up of brain. The elephant brain is only

O.2 per cent of its body
brain,'which

weight

compared

to the human

brain

nonetheless

a basic

and consistent

starts to emerge'

Some

changed at all over time: for example'

brain stem' is
the stalk that rises from the spinal cord' the
there
recognizable as a landmark in most cases' llowever'
the
are variations on a theme: for example, in the cockerel'
cerebellum, the 'little brain,' is about half of the total brain
can actually

reach up to 9O per cent of the total mass of the brain.

cerebellum

is not the entire story: the

shrew's brain is about 3.33 per cent of its body weight yet no
one would claim that the shrew is particularly
intelligent in fact, the shrew is not at all famous for what it tliinks.

for the

as a reptile

as the cockerel'

such

such as the crocodile

mass. In contrast, in certain fish the cerebellum

is 2.33 per cent of body weight.

But percentage of body weight

regions have hardly

a person?

not: some people clairn that since elephants

bigger

In species as different

behaviour

must

of a wide

but is nonetheless
i
l

Perhaps the most celebrated fact about this little creature is :

what it needs to eat, namely its own body weight of insects

that is common

have a function

range of animals,

particularly

dominant

cockerels, and even more important

For animals

such as humans,

including

The

humans,

in the repertoire

more

t
t
;

of
;

sophisticated

lifestyles, the cerebellum constitutes a far smaller fraction of

the total brain.

I
I
I

in the

still in flsh.

with

I-l
;

animals.

animal.

I
I

rememberthecriticalandbasicfeatureofthebrainist}ratit
we are exploring the
is composed of different regions' If
it might be very
significance of different brain regions'
and see how
helpful to turn once more to evolution
with those of other
individual human brain regions compare

that they vary in size. An easy deduction

of the brain

BR'Arhls t't

arrcl ratio to body

every clay. Theref'ore' crthel thirn size
facts about the brain'
weight, there must be other critical
absolute brain size'
So far we have been consiclering only
mass' but
treating the brain as a single homogeneous

of masses at

behaviour

to throw

topic of this chapter: how functions

actually

.i1it'rri'r

::."I.

BRAIN

It seems reasonable to presume

that the

I
I

t
I
I
I
I
I
I
I
I
I
I
I

t8

THE

HUMAN

BRAIN

cerebellum

is not linked

idiosyncratic
capable

B R A IN S

and

compiicated

repertoire

closely
of

for which

to the moxe varied

behaviour

we rnust

of

which

presumal-rly

we

are

have more

brains. In contrast to the cerebellum,

region that has undergone

and

the brair_l

the most change during evolution

is the outer layer of the brain, the cortex.

An

important

clue

to brain

function

a clear correspondence

with

localized
contract

part of the cortex down through

is that

able

rcspecting the confines of a relatively

the spinal cord to

as

the rnotor cortex. At the sarne time there are other specific
in

more

areas of the cortex

for example,

eyes and ears respectively.

been

and outputs. For

muscles - hence this region of cortex is known

that

area has

brain inputs

example, the brain sends signals from nerves from a highly

auditory

surface

B R A IN S

certain areas of the cortex, but by no merns all" seem to have

sophisticated animals the cortex is folded - convoluted - so

its

W ITH IN

to

increase

while

smali skull. Flattened

the visual

and

cortex

cortex - that receive and process signals from the

skin carrying

In a similar fashion, rlerves in the

signals relating

to pain and touch are sent up

out, the rat cortex would be the size of a postage starnp, that

the spinal cord to the area of the cortex that responds to

of the chimp would be the size of a piece of standard typing

incoming

signals regarding touch, the somatosensory

l)aper, while the human brain would be four times greater

stilll Humans have the least stereotyped, most flexible

be so neatly

classified. For example,

lit'estyle of all animal species, and it is believed tJre cortex

back

head

urust therefore

receives input

individual

in sorne way

be related

from fixed" predetermined

to liberating

the

patterns of behaviour.

'I-he rnore extensive the cortex, the more an individual

be ;rble to react in
accordance with

a specific,

unpredictable

the dictates of a conrplex

will

fashion

situation.

in
The

However,

cortex.

there are other regions of the cortex that cannot

of the

at tle

parietal

the

cortex)

and domatosensory

of such a reg;ion is less obvious.

damage to the parietal

range of impairments,
of the lesion.

(posterior

frorn the visual, auditory,

systems. Thus, the function

Patients with

top

a region toward

cortex display

a rvide

according to the exact area and extent

These symptoms

can include

a failure

to

rnore extensive the cortex, the more an animal will be able to

recogriize objects by sight or touch, or a failure to recognize

think

with

for

itself.

But

what

is reallv

rneant bv

t]e

term

tltinking'!
'l'he
divided,

one sense -what has already

another: for example,

cortex

is about

2 millimetres

according to different

thick

conventions,

lhat cach belong to fifty to a hundred

and

into functions

completely

ar()as. [Jp to a point this type of classification

can be

separate

makes sense:

been experienced

someone with

might be unable to recognize

with

parietal Iobe damage

by sight a ball he or she had

previously

heLd wtrile blindfolded.

ders with

the senses, the output

As well as these disorof the brain,

the motor

system, is also disturbed. For exarnple, parietal patients can

t9

v.

; ;..,....

a,s

*.*si-li4r*.rysr*w@ebElfryA$48&&.+r,

20

THE

HUI-{AN

(apraxia) in manipulating

be clumsy

ing. They can confilse

nbjecis o!: e\ren dress-

left and right and have disorders of

spatial skills. As well as these problems

sensory inputs
the parietal

and motor

outputs

involving

of the. brain,

patients

may

deny

that

h.alf of their

actually belongs to them. This phenonenon

problem

auditory

where

stimulation

It is irnportant
parietal

they neglect

operational

appears to lie in the massive

that we normally

sensory

in

visual,

some

system

v;ay

to another,

sensory systems and

well. Rather, the problem

coordination

of senses and

take for granted.

the parietal

cortex

or indeed

sensory

cortex. But a cortical

Because it
relates

one

systems to
as the

area such as the parietal

cortex does not just act as a simple crossroads for the inputs
and outputs
recognition

of the brain. In addition,

for parietal

there are problems

patients, possibly

of

leading to bizarre

denials of half their body: in turn, this 'neglect' can give rise
to even wilder
someone

claims that their arm, for example, belongs to

else. Clearly

'association'

then, the parietal

cortex, like other

areas of cortex, must be responsible

most sophisticated
as neuroscientists

and elusive functions

prefer, cognitive

association

sophisticated,

most

for the

of all: thinking,

processes.

marked

areas were most

our deoxyribonucleic
the

or

com-

Even

whorn

several

are indeed

times

that it is these aleas of cortex not

allocated to controlling

movement

our senses that are the most intriguing,

the harclest to understand

the

with

with

or to processing

and at the same time

in terms of exactly what they do

and how they do it.

For example, a large part of association cortex (see Figure
2) is found at the front of the brain, the prefrontal cortex. Of
ttre

all tl.e regions of cortex, this region has demonstrated

most spectacular

growth:

during

mammalian

evolution

it

has increased 3 per cent in cats, t7 pet cent in chimps, and a

staggering 29 per cent in humans. The first clue as to the
actual

function

of the prefrontal

cortex

is from

a chance

event that happened in 1B4B in Vermont.

At that time in the United

States there was an enormous

expansion of the railways. Phineas Gage was a foreman on a

railway gang ancl it was his job to push dynamite down a
hole in order to explode

differs by only 1 per cent,

cortex

greater. It is not surprising

in animals

the chimpanzee,

acid (IINA)

areas of association

expect that cortical

iifestyles.

individualistic

brain

specilrc

strategy of comparing
species, \Me might

regions in different

directly

motor systerns, this cortical region has become known

association

and

to realize that patients with damage to their

crn move their muscles perfectly

seems that

body

is part of a still

all tactile,

ti; the

pared to our nearest relative,

of that side of the body,

lobes have fully

rnovement

damage to

lobe can result in some very bizarre thinking.

For example,

wider

the main

Retulrririg

2l

B R A IN S

\^/ITI' tIN

B R A IN S

BRAT N

any obstacles that blocked

the

passage where the track was going to be laid. In orcler to

apply the clynamite, Phineas had to use a rod-like

object

Irl
I
I
I
;

I
I
I

22

THE

HUMAN

B RAIN

B R A IN S

knor,rrn as a tarnping

iron,

which

in this case tvas 3 feet Z

prefrontal

cortex. Arnazingly,

\^/ITH IN

after a brief period

B R A IN S

of uncon-

inches long, and at its widest point was i..25 inches across.
One day Phineas was pushing dynamite down the hole

sciousness, Phineas seemed to be remarkably

such a dramatic

colrrse of events. Once the infection

with his tamping iron, when a tragic accident happened. By

chance, a spark ignited the dynamite prematurely and it

cleared

up,

senses and

nothing

had ever happened.

exploded.

Although

there

was

Phineas survived, but not without

holdi^g
ignition

his

head

bone into

hefty

explosion,

some injurv. He had been

to one side, such

of the dynamite

the le{t-hand

a very

that the prernature

drove t} e tamping iron up through

side of his skull. The iron went through

the front

of the brain,

severely

damaging

his

movements

were

by
had

as though

But as tirne went on, people started to notice a difference.

Whereas

he had been a cooperative

before

and friendly

perscln, Phineas now became overbearing, indecisive,

gant, obstinate,

and uncaring

ally left his job on the railway

his

of his life as a fairground

iron still lodged
After

and ended up living

freak, touring

more startling

survival

skills

the tamping

cases of brain

were reported, al} rnore or less indicating

cortex

with

does not

appear

such as breathing

or to the processing

to be related

or regulating

to brute

temperature,

of any of the senses or the coordination

but rather wittr the most sophisticated

of our rnind,

the essence of our personality

as individuals

immediatelv

the

to

interesting

world.

Such

we think

aspects

and how

we

anecdotes

are

because in retrospect

that our cb.aracters, which

inviolate

damage

the same idea: the

of movernent,

react

the rest

in his brain.

this incident,

prefrontal

arro-

for ottrers. In fact, he eventu-

the

F IGURE 2
( i,rrr[)ilrib-()n.f thc r;ortex of the human andchirnpanzee.
Note thatrnore of the
r,or'lrrx of tho r;himpanzee is devoted to specific functions (shaded
areas),
wlttttrttls tlttt llttrrrail has mluy rnore parts of the cortex that are not allocated
to
r :1,irr l.l,s (trrrsh ircled areas) - association cortex - particularly in the
front of
llrrr lrlrrirr (lrrofrontal cortex). (Adapted frorn p. corsi
{ed.), The Enchanted
l,t n )tr r, ( )x li rld (Iniversitl' Press, 1.991.)

unaffected

they reveal

of as fairly

fixed and

aspects of ourselves, are really at the mercy of our

physical brain: they are our brain. For current purposes they
also raise less philosophical,
ing

the

prefrontal

function

of

the

more specifi.c questions regardarea of brain

in

question,

cortex. Is this single area that controls

the

character a

73

24

THE HUMAN

kjnd

of executive

tninibrain

within

a brain? Sur:]t a concept

was even too gross for phrenologists

who devised

sions

distributecl

of character

different

and

regions. What

erroneously

then is the function

subdiviin

them

of this frontal

brain area?

the Second International

that he heard a report

monkey

who

became much

Inspired,

approach

difficult

white

of leucotomy,

Moniz
humans.

from

the treatment

neurotic

after its frontal

proposed

a similar

He deveioped

the

the Greek for 'cu.tting

the

{nerve fibres)' that connected

rest of the brain. Until

attended

It rnras at this

of an a;lparcntly

more relaxed

lobes were lesioned.

to treating

Egas Moniz,

Congress in London.

meeting

techniqrre

the frontal lobes to the

t].e 196Os, frontal leucotornies were

of choice fol a whole range of very intense and

persistent emotirrnal responses such as depression, anxiety,

phobias,

Drrrini; their' lrelzillllr' |.,1,toltlrnics waie claimed to result

in t-ew side-effects. It gradually
that

and aggression. Between

35,OOOpeople in the United

1936 and tgZB, sorne

States underwent

the surgical

procedure. In order to appreciate jrrst how many individuals

were so treated, take a look at everyone listed as 'Smittr' in

sight

and

plans

performed

each year. The development

cated drugs as well

as a final realization

nJeficits that can result

hands of clinicians

from

for whom

the surgery

of leucotomies

of more sophisti-

In

unresponsive'

to be proactive,

from

information
behaviour;
This

or change

instead, ttrey perseverate.

profile

of dysfunction

on certain specific

studlziylg the performance

tasks of patients

has Lreen characterized

and indeed

by

experimental

rnonkeys who have damage in

the frontal lobes. For exarnple, such subjects cannot switch

rules if tley

are doing

something,

cards

such as sorting

according to colour of symbols, when subsequently

asked to

sort according to shape of symtrol. Some people ref'er to this

ability, which

we all normally

frarnewr:rk

have, as working

in r,vhich a task is performed

dubbed

the 'blackboard

memory, a
and which
With

of the mind.'

failures in warking memory, it is hard to remember events in

But

the

problem

of

context.

have stayed the

prefrontal

cortex is not just one of memory'

the only possible course of action a few decades earlier.

use

cannot

They

a proper

have seemed

strategies or

novel

of the cognitive

surgery would

damaged

to regulate

environment

their

this

wiih

line

patients with

problem.

a particular

to tackle

in fore-

characters, lacking

lobes are less able to develop

frontal

is sometimes

the number

becarne changed

apparent inability

there

in

and

beneit

therapeutic

arguable

ernotionally

working

a decline

became apparent, however,

indeed that the side-effects were severe. As r^/ith Phineas,

the New York City telephone directory! Since the Iate 196Os
has been

no net

was

there

these patients

In 1935, a Portuguese neurologist,

B R A Ii ' IS

W ITH IN I

B fi ,A IN S

BRA' / V

/)

damage

of this damage is loss of vertral spontaneity:

damage to the

prefrontal

cortex

tend

to

Another

the

result

patients with

to volunteer

less

rl

I
;

rlI
I
I
I

t
I
I
I
I
I
I
I

26

T HE

HUMAN

B R AIN

information,
behaviour

B R A IN S

as well

that we saw with

Despite this wealth

exactly what might
some

as displaying

the

be the function
have

between

patients

phrenia.

Schizophrenics

with

social

Phineas.
it is still hard to say
of the prefrontal

pointed

prefrontal

hard

to match

exciusively

of information,

neuroscientists

impaired

to

the

lobes.

similarities

lobe darnage and schizo-

also appear to have problems

on

Different

up

with

familiar

actual

events

events

in

in

W ITH IN

the

outside

world

brain

region.

a single

parts of the cortex, such as the motor

the somatosensory

cortex, clearly

of the parietal

cortex

must

cortex and

have different

and association areas such as the prefrontal

B R A IN S

functions,

cortex and parts

each have their

own

specialized roles. But contrary to the phrenologists'

type

of

vision,

the same working memory tasks as the neurological patients.

these roles do not correspond

I-lence schizophrenia

obvious aspects of our character and specific activities in the

matching

has been interpreted

up incoming

information

as a disorder

with internalized

ards, rules, or expectations. The schizophrenic

tal patient would both tre overwhelmed
sonsory input

of

stand-

and prefron-

and dominated by a

that they could not adequately categorize or

l2y memories that they could not fit into the correct temporal
soquence.

It

is

almost

as though

they

lack

the

inner

real world.
today

to

understand

aspects,

consequences,

and

corollaries,

to

be

The

cases of Phineas
one approach
the

role

erstwhile

function

lives. II we were phrenologists, it would be hard to think of a

that might

one-worcl label that could be appropriately

directly

is

used in brain study by those trying

of

a specific

example

have been thought

the function

patients

brain

region:

look

at

brain region and infer its

dysfunction

is now appa-

of selective

to indicate

brain

damage

irnmediately

of the area in question

and

is Parkinson's

disease.

Wo can say that a patient has social problems or problems

rn'itlr working

what

in outward

Gage and leucotomized

from whatever

rent. One well-known

l'rontal lobes.

events are reflected

examples of damage to a particular

summed up as a single and icientifi.able function in our daily

placed on the

between

behaviour.

to identify

clifferent

relationship

certain brain regions and how such

physiological

internalized

illustrate

and abstract a process, rsith far too rnany

the

actually going on within

happenstance of life. However,

it is too complex

basis with

It is one of the biggest challenges in neuroscience

resources that act for most of us as shock absorbers to the

if such a hypothesis is true,

on a one-to-one

rnemory, but it is verlz hard to find out what

Parkinson's
Parkinson,

disease

was

originally

who first reported

named

after James

the conctition

in L81-7. This

affects mainly

older people,

I lru r:ornrnon lactor is between these two disparate impair-

sevele disorder

rnorrls. Inrleed for many, if not most, areas of the brain, it is

although younger people can sometimes fall victim. Patients

of movement

27

THE

HUHAN

B F.A IN J

B R A IN

have great difficulty

when

a tremor

many

At the very

or

lies, in

exactly where the problem

in the middle

of the brain.
part of the brain

core of this middle

moustache-shaped,

such as depression

of the brain,

we know

an area deep down

disease is that

aspect of Parkinson's

disorders

schizophrenia,

of their

hands are at rest and stiffness

their

limbs. The fascinating

unlike

they might have

in moving; in addition,

area

black-coloured

lies a

consequently

more relevant
question

not directly

product

of an important

has undergone
firmly

various

established

normally

brain chemical,

cells

normal brain is compared

substantia

wittr a Parkinsonian

dying

is that the chemical

manufactured
given

a tablet

in this
of

region.

the

ttrat if a

one, then the

with

paler - the cells containing

have died. One of the important

are

Parkinson's
the pigment

consequences of these cells

dopamine

is no longer

If Parkinsonian

chemical

(L-DOPA)

being

patients
from

are

which

target brain region, the striatum'

then

would.

be: what

is t]-e

The imporof

function

of the brain

The anatomy

in the striatum?

to anotfrer

doparnine

does

of the brain: there

match up with the chemistry

exclusive to any one brain region. Rather,

over marry different brain
the same chemical is distributed
regions while each brain region makes and uses many
is no one chemical

what

when

is most important

the brain

It is therefore

chernicals.

brain

different

for a Iong time

nigra in the brain of a patient

disease is much

nigra

dopamine.

it has been known

Similarly,

after it

dopamine,

the substantia

in

makir-rg the chemical

end

reactions. Hence it is now

chernical

that

the

turn

cells deliver

the critical

from which

pigment

is in

Parkinson's

cal region but is also specific to the chemical dopamine'

some might view the substantia nigra as the mere location

dopamine

Melanin

the fact that

not just the substantia nigra as an anatomi-

disease involves

nigra appears black because the cells in tl:is region have the
thern.

ignore

cannot

wo

Moreover,

tant

in

B ' A .A II.IS

- that is, dopamine - rrcr orre has any precise idea as to the
function of the substantia nigra in normal movement'

named in Latin substantia nigra (black mass). The substantia

melanin

\^.' ITH I N

region

concerned

considering

very hard to say

brain

damage -

or the change in the chemical

balance in the brain'

There is an.other reason for being wary when trying to pin
down particnlar functions to particular brain areas: neuronal plasticity.
many

reasons

gunshot,

Brain

areas can, of course, be damaged

for

accident'

or

whether

br.rt a very

it

be

common

disease,
reason

car

is stroke'

A stroke

is in Parkinson's disease - that is, the substantia nigra - and

there is not enough oxygen in the brain' This

lack of oxygen could be due to a blood vessel that has
become blockecl, thus preventing access of the blood nor-

even though we know

mally

dopamine
movement.

is made, then there is a dramatic

improvement

in

Even though we know exactly where the damage

what particular

chemical

is deficient

occurs when

carrying

oxygen around the brain, or because there is

r ro

I
I
I
I
I
I
I
I
I
I

t
I
T

THE

HUMAN

B R A IN

a reduction

B R A IN S

in blood flow due to a namowing

of the blood

claim that one part of t-le brain definitely

\^/ITH IN

does one thing; if

vessels. If a stroke occurs, in the motor cortex for example,

other, albeit

then it is possible to trace a sequence of rather interesting

then clearly there is at least a degree of flexibility,

events that unfold.

neuronal

Initially

after such a stroke there may be no movement

all, not even reflexes: the limb

at

on the affected side of the

body just dangles loosely (flaccid paralysis). Then a seeming

miracle occurs over a period of days and weeks, although
the extent of the miracle will

vary enorrlously

from patient

to patient. First of all, reflexes can return, t-Len the arm will
start to be rigid
rnovements,

and the patient

and finally

will

the stroke victim

grasp something. In one study, a third

hatl stroke in the motor
spontaneously

be able to make limb

will

be able to

of the patients who

cortex were able to grasp objects

and hence reach this final stage of recovery.

adjacent,

brain

areas can take over that role,

known

How then rnight we study the function

of different

video, of tJ'e inside

talking,

of the brain as a person is thinking,

or performing

The story of how

this idea is actually

starts with a familiar

high-frequency,

any of a variety

procedure:

radiation

becoming

a reality

t_heuse of X rays. X rays are

short-duration

Because X-ray

of usual functions.

electromagnetic

is very

high

waves.

energy, it

readily

penetrates a test object: the atoms in this test object absorb

some of the radiation,
strike a photographic

leaving

the unabsorbed

plate, thereby

exposing

portion

whereas

population

r:orrlrl rocrovery of function

rlrrosliorr, with

their

of neurons - how
occur

exclusive

Inslonrl, it is as though

otherwise

if the original
monopoly,

other brain

cells in

were

dead?

cells were gradually

l,rrrnirrg to take over the role of the damaged cells. Indeed,

llro slngos of recovery

of the grasping movement

Irrrvo irrsl trirceiclfollowing

that we

the

more

radiodense

plate.

This

photographic
very well

at security

an object,

plate,

the whiter

process, as we all know,

checks in airports

to

it. Thus, the

rios. Brain functions

particular

brain

regions? What we really need is a snapshot, or better still a

less radiodense an object, the darker the photographic

Itl o'e

as

plasticity.

-l-here are also reports of recovery from brain clamage that

havo affected speech and memory after certain head injutherefore need not belong to one area,

3l

B R A IN S

for material

the

works
where

there is great contrast, such as a gun inside a suitcase, or in

hospitals to visualize
Although
happening

rays

broken bones in flesh.

are

effective

detecting

what

is

in most of the body, when it comes to the brain

stroke in the motor cortex are very

there is a problem.

Unlike

rrirrrilirr to tlro initial development of this same movement in

lrrlirrrls, irl; we "shall see in Chapter.l. Again, it is hard to

flesh, there is little

difference

region

for

compared

the contrast between

to another.

in the density
To overcome

bone and

of one brain
this

hurdle,

32

T HE

HUHAhJ

B RAIN

solution would
alternatively

ts R A IN S

be to rnake the brain more radio-paqrle, or

to make the technique

made to resemble the scenario of the gun in a case, how

compared

could be made to give greater contrast

rnrith the rest of the trrain. This objective

realized by injecting

injection
bone.

is not directly

Rather,

can be

into the brain a dye th:rt, is very opaque

in that it can absorb a lot

the

of the X rays. I{owever,

into

t}re brain

the dye is introduced

through

into

the skull

the artery

that

pumps blood into the brain. You can locate tJ'is artery (the
carotid artery)-if you place your hands on your neck, near
each side of the windpipe,
radio-paque

the brain very quickly.

obtained

and feel a pulse beating. Once the

dye enters the blood

is called

circulation,

The kind of picture

an angiogram.

readout

of the pattern

through

all the brain regions.

it is fed into

that can then be

Angiograms

of branching

blood

give a clear

vessels that go

for

example,

blockage
problem

if

someone

or a narrowing

of the blood

it will

away and the abnormal

will

has a stroke

valuable

circumventing

the problem

lies.

to making the brain more radio-paque

is to

is not where

circulation

Then angiograms will

The alternative

not be helpfui.

make the method of detection more sensitive. With norrnal

X rays there are about twenty

to thirty variations

there

vessel walls.

more than two hundred

variations

In CAT, brain X rays are taken in a series of sections or

his or her head in a cylinder

scans. The patient lies with

with

an X-ray tube on one side and an X-ray beam on the

other, and these two devices are placed around the head.
The X ray does not strike a photographic

plate but rather'a

sensor that is connected

this sensor is far

to a computer;

X rays. All the measurements

of the insensitivity

ere taken and assembled by

to give a scan. The tube moves along the axis'of

computer

This

the brody and this procedure is repeated eight or nine times.

eye. In this way, angiograms

offer

plate used in ordinary

is a

due to the displacernent

that

axial tomography

(CAT).

neurologists

tools

and has been routinely

used since the early 19BOs: computerized

sometimes push blood vessels

diagnostic

on the grey

scale; but a technique was developed in the early 1970s with

The kind

positioning

the problem

but the blood

if a

be detectable to a trained

are very

where

urill tl.en show up on the angiogram. Similarly,

patient has a tumour

brain tissuc. ijrrt r.rilrirtii' the blood vessels are functiclning

more sensitive than the photographic

Now imagine there is damage to the cerebral circulation

B R A IN S

normally? It coulrl be that there are problems with the brain,

of X rays more sensitive.

Let's look first at how the inside of the brain itself could be

certain components

W I"I H IN

a way

of

of X rays to

of pictures

that can be seen by CAT scans give

and brain surgeons a valuable indication

of the

location and extent of tumours arrd tissue loss. For example,

CAT scans have recently given a clue toward understanding
the degenerative

disorder

ihere is severe confusion

of Alzheimer's

disease, where

and loss of memorv.

A. D. Smith

33

34

I
I
I
I
I
I
I
I
I
I
I
l

t-

THE

B R A IN

HUMAN

B R A IN S

and K. A. Jobst found that in Alzheirner patients a certain

brain region (the medial temporal iobe) gradually decreased
over

time

to

half

about

the

width

that

of

in

of the same age ('Use of structural

volunteers

study the progression

of Alzheimer's

healthy

imaging

to

disease,' British Medi-

brain

uses up so much

for 2O per cent of energy consumption

When

has enormous

for tJlis debilitating

potential

diagnostic

brain damage before the clinical

disorder,

the start of

for revealing
symptoms

but it

of memory

loss

have become clearly apparent.

most of the twentieth

has

angiograms

there

abnormalities

in the anatomical

you have a CAT

scan it will

physically

wrong

brain, such as a tumour

tbr

are limits

that can be studied

dysfunction

something

their use in CAT scans and

invaluable

been

F{owever,

damage.

century,

to us for

investigating

to the type

brain
of brain

and

you whether
enduring

you have
youf

within

or a lesion. But if the problem

is

the actual operations of the brain - X rays will

rrot tell you

during

bits

of your

brain

a particular

are working

task.

How

might

at particular
this

tirnes

problem

be

it uses up much

Of all the organs of the body, the brain is the greediest in

It burns

oxygen and glucose at ten

more

in the food

that you eat and the oxygen in the air that you breathe: when
carbohydrates

react

with

olcygen,

water, and rnost important

they

generate

carbon

of all, heat. In the body,

all the energy from food is not released immediately

simple

combustion,

because it would

there was no energy left for any of the functions

and body. Thus, alttrough

Through

if

of the brain

some heat is needed to keep us

in the body

that prevents

the

release of all the energy from food we have eaten.

its formation

we are able to store up this energy for

t}.e rnechanical,

electrical,

and

has

the

in a

not be very helpful

brain

to

and chemical
do.

The

energy-storing

(ATP) is produced

adenosine triphosphate

work that the body

chemical

from the food we

eat for as long as we are alive. ATP stores energy and has the
potential

to

liberate

it

like

a compressed

spring

upon

release.
If brain regions are active during a particular

overcome'l

its fuol consumption.

at rest. But what

fuel. The fuel for the brain is the carbohydrates

irnmediate

If

to rlo with

what

is working,

properties
tell

rather than anatomical

region

\Marm, there is a chemical

- something

functional

a brain

in this way. X rays detect

of the brain.

it is responsible

to this energy? It enables the brain to 'work.'

dioxide,

Despite the fact that X rays have been familiar

of

Even though the brain is less

than 2.5 per cent of our total body weight,

indicate the brain region that should be targeted for developtherapies

energy that it dies if deprived

oxygen for only a few minutes.

happens

ing possible

35

times the rate of all other body tissues at rest. In fact, the

52, 575-5BO). Not only d.oes such an observation

cal Bulletin

B R A IN S

W ITH IN

working

task, they are

hard and using more energy; thev are making great

demand on ATP stores, and hence more carbohydrates, the

36

TH E

HUMAN

simplest

B RAIN

B R A IN S

is glucose, as well

fbrm of which

required.

It follows

demand

for oxygen

brain, we would

if we could

that

principle

two

of the

One technique

is known

as positron

an unstable

speed. Positrons

electrons

positrons

nucleus

ernission tomography

that ejects positrons

particles

are fundamental

by the blood

collide

to

atom, in the sense that it

into

urith electrons

the brain and mutually

at very

similar

to

annihilate

S peak i ng w ords

Generating verbs

charge. Radioac-

into either gluc.ose or water

are irrjected intravenously.

carried

used

so that it can be easily tracked. The

tive oxygen atoms incorporated

is then

techniques

except that they have a positive

molecules

task. This is

in PET is for either oxygen or

'label' in this case is a radioactive

high

parts of the

at work.

(PET). The basic requirernent

glucose to be labelled

any particular

particular

the brain actually

contains

trace the increased

be able to say what brain areas were most

hardest during

visualize

B R A IN S

as oxygen are

or for glucose by certain

active or working
the

W i TI."I IN

The radioactive
the brain.

label

The ernitted

in other molecules

within

each other. The burst of

FIGURE 3
(PET) ecan of a
The interactive
brain. A positron emission tornography
conscious human subject performing similar but subtly different tasks, Note
that although the tasks are all linked to language, different brain regions will
be involved according to the precise nature of the undertaking, Note also that
at no time is just one brain region active completely on its own. (Adapted from
M. I. Posner and M. E. Raichle, Images of Mind, Scientific American Library
S e r i e s , 1 9 9 4 .J

energy that results forms a gamma ray that is of sufficiently

high energy to penetrate
outside

through

the skull

and be detected

of t]:e head.

Because these high-energy

gamma rays can travel a long

way, they pass right out of the head and strike sensors, the
signal frorn which
brain at work.

is then used to build

up an image of tl:e

The glucose or oxygen accumulates

brain regions that need it most, namely

in the

whose working

the

hardest.

With

areas according
compared

PET, it is possible

to show

to tasks as subtly different

different

active

as saying words

with reading words. (See Figure 3.)

A second imaging
nance imaging

technique,

functional

magnetic

reso-

{MRI), is like PET in that it relies on the

THE

BRAIN

HUMAN

of energy by whatever

expenditure

differential
are working

however,

hardest;

exactly when

brain regions

ttris time no injections

Because there is no protrlem

involved.

with

B R A IN S

with

are

ascertaining

MR[, like

PBT, also measures changes in blood o>cygen concentration

serving

brain

method

of detection

protein

haemogl.obin.

amount

of oxygen present affects the magnetic

haemoglobin:

areas tJ:at are rnore

these

is different.

Oxygen

MRI exploits

properties

presence of a magnetic

active;

bombarded

is carried

by the

properties

can be monitored

field, where the nuclei

and pushed

the

the fact that the actual

in

of
the

of the atoms

line. The radio

signal is unique

carried by haemoglobin

and

to the arnount

of different

of oxygen

regions of

an area as small as l-

events

that during

brain regions are working

a specific

taking

place

ilppear to contribute

it is becoming

over

increas-

task several different

simultaneously.

clno brain area for one function

magnetoencephalography

(MEG), measures the magnetic field generated by differential

electrical

activity

resolution,

but

of the brain
is only

future,

when

commensurate

the

space

with

and

currently

regions of the brain. Although

has a superior

accurate

There is not just

but rather several brain areas

to a particular

function.

Moreover,

if

for

their true potential

and

tirne

resolutions

time

the

outer

lies in the
. are more

the scale of real brain cells, techniques

such as PET, MR[, and MEG are already offering windows

at work.

Perhaps the most

obvious

in

the

regions
different

phrenologists'
cornbine

in

scenario.
some

way

of one

function,

Instead,

different

to work

in

as

brain

parallel

for

functions

The brain is made up of anatomically

ttrese regions

tJ e most

to

lesson they

to think

brain region as having one specific, autonomous

constitute

the use of these techniques

apparent

rnethod,

have taught us so far is that it is misleading

seconcls.

ingly

metre of tissue. Another

by radio

can pinpoint
measure

over a time scale exceeding

out of alignment

very sensitive measure of the activity

to 2 millirnetres

events are monitored

the brain

in the sample and therefore gives a

the brain. This technique

Brain

magnets.

waves, these atoms ernit radio signals as they spin back into

With

lation of brain regions appears.

miniature

they were themselves

line up as though

however,

constel-

several seconds and averaged over, at best, a cubic milli-

to give an even more faithful

r:eflection of what is going on at a given moment.

B R A IN S

such as hearing

words rather than speaking words, then a different

the injected. label reaches the brain, imaging

MRI has the potential

When

sorne aspect of the task changes slightly,

\^r' ITH IN

are not autonomous

a cohesive

minibrains;

and integrated

part in a mysterious

distinct

way.

regions, but
rather, they

system organized

for

Therefore,

be

it *"ill

almost impossible

to learn how the brain works by studying

just one particular

region at a time. Insteacl, let us start with

specific and familiar

functions

and trace how their process-

ing is parcelled out between multiple

areas in the brain.

39

;;;;;;;*T;ffi:
Every waking moment we are bombarded with sights,
sounds,and smells as we touch, run, climb, and speakour
way through life. Animal existence is one of constant
dialogue with the outside world. The brain is vital for
processingand coordinating the information that floods in
through the senses,with the outputs of the brain expressed
as movements. But how is it all achieved?In the previous
chapter we saw that there was no single centre in the brain
directly correspondingto each and every function or behaviour in the outside world. In this chapter we shall explore
the riddle of how function is localized in the brain by
starting not with brain regions but with the functions
themselves.
Charles Sherrington, one of the greatest pioneers of
physiology during the first half of this century, summed up
the all-pervasive contribution of movement to our lives:
'From a whisper in the forest to a felling of a tree, 'tis all
movement.' From the subtleties of body language to the
precision of the spoken word to the unambiguity of a simple
hrg, virtually all communication relies on movement.
Howeverglobal or imperceptible,all movementdependson

.mls
.2

THE HUMAN BRAI N

SYSTEH S OF SYSTEM S 1 3

the contraction of some muscle group somewhere

in the
body. If contraction of all muscle is defunct, all
that is left is
the ability to drool or shed tears.
Although plants can move in the sensethat they

may turn

to the light, they cannot generate movements

as we do.
Outside the realm of science fiction, no plant
locomotes
from one place to another.In clear contrast,
all animals are
on the move - that is, they are antmofed.
Interestingly
enough, the Latin onjmus means ,consciousness.,
If you move about and you are a multicellular
organism,
then you have, at the very least, a primitive kind
of brain.
The importance for moving creaturesof having
somekincl

of
brain is best illustrated by an observationinitially
made by
the late Emperor Hirohito gf
Japan,for whom the study of
marine life was a passionatehobby. The tunicatein
question
is known as a sea squirt. When it is an immature
larva the
sea squirt spends its time swimming around: not
only is it
capableof coordinatedmovementbut it alsohas
a primitive
vibration-sensitivedevice,crudery comparablewith
an ear,
and a primitive light-sensitivedevice,roughly analogous
to
an eye.In fact, the seasquirtcould be said to have
a modest
brain. However, when it becomes mature the sea
squirt
changesits lifestyle and attachesto a rock. It no ronger
has
to swim around anymore,becauseit now lives by
filtering
seawater.At this .stagethe sea squirt actually performs
the
remarkable act of consumingits own brain.
The clue to brain functionprovidedby this taleis that you

Iil

only need a brain when you are moving. For stationarylife

forms, a brain is no longer necessary.The whole point is that

for an animal moving around, there is an interaction with an

environmentthat is incessantlychanging.you needa device
to tell you very quickly what is happening and, most
importantly, to enableyou to respond to what is happening,
to get out of the way of predatorsor to chaseafter prey. So
the brain, in whatevershape,size, and degreeof sophistication, is somehowconnectedin a very basicway to ensuring
survival as both a consequenceand a causeof movement.
According to the lifestyle of the animal, there are different
types of movement. The trapeze artistry of the swinging
monkey,the precisiongliding of the eagle,and the coordination of the legs on a millipede are examplesof specialized
movement accommodatingparticular lifestyles.
How is movement of any,sort actually achieved?The
contraction of the appropriate muscle occurs following a
signal sent down from the brain, along the spinal cord.
Nervescontrolling all the different musclesleavethe spinal
cord in an ordered fashion,accordingto the location of the
musclein the body. Peoplewith injuries to their spine are
unable to move to varying extents accordingto the level at
which the spinal cord is damaged.
Sometimes the spinal cord can function more or less
autonomously,without descendinginstructionsor control
from the brain. Such movementsare reflexes.A reflexcan be
clefinedas a fixed responseto a particular trigger- the most

I
I
Ir

II

t
t
t
;

THE HUHAN BRAI N

OF SYSTEM S
SYSTET.,IS

obvious example is the knee jerk. The knee jerk reflex is

is, however, yet a further type of movement that we tend to

triggered when the knee is tapped and in responsethe lower

take for granted and that is not controlled by any of these

leg shoots out. Neuroscientists refer to this familiar

four systems:fine movement of the fingers. Dexterity with

sequence of events as the 'stretch reflex' becausethe tap at

our hand distinguishesprimates from all other animals. It

the critical point on the knee in effect compressesa tendon

enables us to fashion and use tools and thus to attain a

by which the muscle in the lower leg is suspended,thereby

lifestyle that other specieswill never be able to realize.The

exerting extra pressure on the muscle and stretching it. In

dexterity of a violinist, for example, where fast, controlled,

order to compensate for this lengthening, the muscle con-

independent movement of the digits is critical, is a spectacu-

tracts, so that the leg shoots forward.

Our normal repertoire of movement is not one of fixed
rsponses to rather artificial triggers like the neurologist's
delicate hammer. We do not wait for someone to tap our
knee so that we might jerk our leg. Many of the movements
that we make - such as wa[king, swimming, and running involve a moie complex coordinationof muscle groups.But
even these movements are, in a sense, semi-automatic.
These kind of rhythmic, subconscious movements are
caused by signals coming down from the brain stem (see
Chapter 1). Different groups of neurons in this brain stem

Iar feat of evolution.

Unlike the other descending spinal routes used for the
contractionof muscle,the messagesinstigatingand controlling fine finger movement do not originate from the brain
stem at the top of the spinal cord but from the very top of the
brain, a striplike region of the cortex fitting acrossthe brain a
Iittle like a hairband, and known as the motor cortex (see
Chapter 1). The motor cortex controls fine movementsby
sending signals directly to the digits in question. It also
indirectly influences movements by sending other signals to

region send signals down the spinal cord to cause the

the four motorway centresin the brain stem, which in turn

appropriate contraction of musclesin a repeatingsequence.

activate the appropriate contraction of muscle. Different

There are four such brain motorways coming down the

parts of the motor cortex are allocated to controlling

spinal cord from the brain stem.One is responsiblefor semi-

different parts of the body. It might be assumedthat such an

reflex rhythmic movements,like swimming, lvhile another

allocation would correspondto the size of the body part in

coordinatesmovement with visual and sensoryinformation;

question - that is, a tiny area such as the hand would be

yet another is important for balance, while the fourth

controlled by only a tiny part of motor cottex, whereas a

nrotorway mediatesthe moving of individual limbs. There

largearealike the back would have the lion's shareof motor

46

THE HUMAN B RAI N

cortex to control its movements. However, nothing could be

further from the truth.

#:l

SYSTEM S OF SYSTEM S

brain, was seen in Chapter 1 to have a function that

presumablydominatesthe lifestyle of cockerelsand fish far

The critical factor turns out to be the precision of the

more than it does our own. Pecking at food or swimming

movement that the body part needs to make. The more

through the sea necessitatesan ability to coordinate the

precise the movementsgenerated,the larger the area of the

information constantly coming in through the senseswith

brain that is devoted to them. Hence the hands and the

appropriate movements. There is no time for thinking or

mouth have an enormousallocation within the motor cortex

planning a movement as other animals loom close to the

compared with the upper arm and the small of the back,

crumbs in the barnyard. Perhaps then the cerebellum is

which do not seem to have much representationat all. The

important for automated movements triggered by outside

kind of movements you make with your back are not that

events rather than by internalized proactive thought pro-

fine or precise. (SeeFigure 4.)

cesses.

The motor cortex is pivotal to the generation of move-

It is quite remarkable that as long ago as 1664 the

ment: not only does it have direct control of some of the

physician Thomas Willis had also formed this view of the

muscles controlling the \ands, and hence of precision

movements,ibut it also exerts a hierarchical influence over

cerebellum, which he refened to as the cerebel. Willis

thelother four movement motorways. In the previous chap-

rest of the brain, responsiblefor unconsciousmovements:

viewed the cerebellumas a structure truly isolatedfrom the

ter the idea of a single command centre for any one function
was rejectedrbut surely here it seemsthat the motor cortex

The Cerebelis a peculiarfountainof animalSpiritsdesignedfor

is well qualified to be the 'Movement Centre' of the brain.

someworks,wholly distincthom the Brain.Within theBrain..'

Not quite. Although the motor cortex plays a critical role

motions,to wit, of which we areknowing

all the spontaneous

in the control of movement,it does not have a monopoly.

and willing, are performed.... But the spirits inhabitingthe

Two other areas would also be worthy contenders for the

Cerebelperform unperceivedlyand silently their works of

title of movement centre:the basal ganglia and the cerebel-

anatome:
or carelWillis,Cerebic
Naturewithoutour knowledge

lum. If either of theseregions,which are far away from the

rrviduessitnervorumdescriptioof unes(London:1664),p' 1111.

motor cortex, is damaged,ttren movement is dramatically

impaired in various ways.
The cerebellum. the little brain at the back of the main

Three hundred years later, this description could still

apply. Patientswith damageto the cerebellumcan move,

i I7

48

SYSTEM S OF SYSTEM S

THE HUMAN B' RAI N

senses,which in turn will trigger or influence the nelrt type

of movement. Imagine, for example, that you have to trace a
complex pattern onto paper, Your hand is under constant
surveillance from your eyes. People with cerebellar damage
find this tracking movement particularly hard.
Humans engagein many more sophisticatedactivitiesnot
dependent on the immediate triggers in the environment.
Our more flexible and versatile repertoire of movement
reduces the centrality of the cerebellum in terms of the
fraction of our brain it constitutes compared to the cockerel
or the fish. The cerebellum is nonethelessof vital importance since the sensory motor coordination it generates
underpins skilled movements that are also the type of
movements not requiring conscious thought. These movements improve with practice to become almost subconscious.For this reasonthe cerebellumhas been dubbed the
F I G URE 4
Reconstructionsofthe human body in accordancewith the spacein the cortex
devoteclto eachbody part for moving.Notehow the mouth and the handshave
thogreatestnumberof braincellscontrollingthem.(Fromthe BritishMuseum
of Natural History.)

'autopilot' of the brain. This epithet fits closely with the

description long ago formulated by Willis.
There is another type of subsconsciousmovement that is
not modified by updated information from the senses.
Unlike movements controlled by the cerebellum, those

but in a clumsy way. They have particular difficulty with

associatedwith the basal ganglia cannot be changed once

movementsrequiring the type of sensorymotor coordination

they have been initiated. These 'ballistic' movements re-

that characterizesskilled movements such as playing the

semblea cannonballexplodingout of a cannonmouth: once

piano or dancing. The cerebellum is important for move-

started,it

and its trajectory cannot be

ments where there is a continuous feedback from your

modified.

a golf swing, the ball might

UNAFT

49

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THE HUMAN. BRAI N

SYSTEM S OF SYSTEM S

stay mockingly on the tee becausethe movement cannot be

corrected at the last moment: it is, literally, hit or miss.

moustache-shaped,black-pigmented region toward the back

of the brain, the substantia nigra, that is less dominant.

The area of the brain associated wiih these ballistic

Becausethese two regions are normally so closely linked

movements, the basal ganglia, is really a group of various

lvith each other, any drug that restoresthe balanceof power

interconnectedbrain regions,When any of theseregions are

betweenthem will be effective.In Parkinson'sdisease,drugs

damaged,there are devastatingconsequencesfor movement.

that dampen activity in the striatum have a similar effect to

According to the part of the basal ganglia that is impaired,

thoseenhancingactivity in the substantianigra. Conversely,

there can be wilcl, involuntary movements (Huntington's

any drug that reduces activity in the substantia nigra or

chorea), or the exact opposite, difficulty in moving at all,

enhancesactivity in the striatum is pernicious in Parkin-

combined with muscle rigidity and tremor (Parkinson's

son's diseasebut highly beneficial in Huntington's chorea.

disease).Huntington's choreaand Parkinson'sdiseaseaffect

Even within one generalbrain region, the basal ganglia,the

two different parts of the basal ganglia (the striatum and

component parts themselvesare not autonomous but are

substantianigra respectively)that seem to normally work in

functioning in an incessantdialogue with each other.

a kind of power-balancin! act, Iocked together so that the

Thus, there is no single movement centre after all. Rather,

the second,a little like a seesawor

first region'counteracts

movementcan be split up - although we are not consciously

arm wrestling. Normally, as with a seesaw or with arrn

aware of it happening - into different types that are in turn

wrestling between two equally matched individuals, one

controlled by different basic brain areas. However, even

brain region keeps the other in check.

thesedifferent brain areas,such as the cerebellumand basal

But imagine a scenarioin wlilCh one person on a seesawis

ganglia,do not function as autonomousunits, but are in turn

much lighter, or an arm-wrestlingopponent is much weaker,

in dialogue rvith different parts of the outer layer of the

the bilance c o lla p s e sHe

. n c eif o n e
tha n his o r her colleague:

brain, the cortex.The cerebellum,for example,has strong

brain region is underactive,the other becomestoo active,It

connectionswith a part of the cortex that lies distinct from,

is this imbalancein activity that appearsto lead to abnormal

and in front of, the moto.rcortcx (lateral premotor area),

movements.In the caseof Huntington'schorea,the deficient

while the basal ganglia are in intimate contact rvith yet

region in the dialogue is toward the front of the brain, the

anotherarea of cortex known as thc supplementarvmotor

striatum; on the other hand, in Parkinson's disease,it is the

area. Indeed, damage to the strplllcnrcntarymotor area

5l

"i;r I

52

THE HUHAN

BRAI N

tr' {
I
{

SYSTEM S OF SYSTEM S

can lead to impairments strikingly similar to Parkinson's

a symphony. The type of movement being made, and

disease.

whether it requires conscious control, determines exactly

In the normal situation, an attractive though speculative

which brain regionsare involved,.while pathologiessuch as

scenario is to view the subcorticalregionsas controlling the

Parkinson's diseasehave highlighted the consequencesof

movements that do not rely on any contributior of con-

dialoguesbetween brain regions becoming too one-sided.

scious thought. For example, pressing the brake when the

But the idea of a 'centre' for different functions in the

traffic lights are red seemsto be an automatic movement,

brain is so intuitively appealing,it is hard to relinquish. We

which is in fact associatedwith the cerebellum.In contrast,

might have more luck with the senses.Unlike with move-

if you finally decide to dragyourself out of the armchair on a

ment, the sensesoffer us a clear stimulus - be it a light, a

Sunday afternoon, the actual movement requires very little

bang, a pinch, or the taste of raspberries- where we can

conscious planning. There is no immediate sensory trigger,

tracethe fate of a 'signal' as it is processedat different stages

but the standing up is automatic nonetheless.Some neuro-

in the brain. Perhaps such a clear path will lead us,

scientists even go so far as to refer to this type of movement

naturally, to a final vision centre,hearing centre,and so on.

as a 'mdtoq program.' lVhatever the label, this type of

'\
internally ftrii;gered movement that most of us take for
granted is controlled by the basal ganglia. It is particularly

Just as there are motorways leaving the brain via the

spinal cord to control musqles and hence movement, so

hard, however, for a Parkinsonianpatient. The basal ganglia

sent up the spinal cord into the brain. Thesesignalsrelateto

and cerebellum in these casesare freeing up the cortex for

touch and pain, and are referred to as the somatosensory

other roles beyond the minute-by-minute task of motor

system. Triggered by the point of contact where, for ex-

control. On the other hand, some movements, be they

ample, a pin pierces the skin, local nerves within the skin

ballistic or sensorytriggered,might require different degrees

transmit signals to the spinal cord. These signals are then

of conscious control. In this case the supplementary motor

relayed up from the spinal cord and finally arrive at the

area and Iateral premotor area dominate more fully in the

outermost reachesof the brain, in an area of cortex just

respective dialogues over their subcortical partners, the

behind the motor cortexknown as the somatosensorycortex.

basal ganglia or cerebellum.

there are incoming signalsthat, as we saw in Chapter 1, are

There are two major motorways heading up the spinal

The generation of movement is the net result of many

cord for the somatosensorycortex; one, the evolutionary

brain regions acting togetheras individual instruments do in

system,is chiefly relatedto pain and temperature,rvhile the

53

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THE HUMAN

SYSTEM S OF SYSTEM S

newer system carries precise signals relating to touch. This

the body can be demonstrated very easily. If a pair of

arrangement has an intuitive appeal in that it makes sense

compasspoints are set relatively near each other and placed

for the more basic, establishedsystem to be concernedwith

lightly on different parts of the body, they will be perceived

basic survival factorssuch as pain and temperature,whereas

either as one point or two according to where they are

the more refined skills involving precision of touch would

placed, even though the distance between them remains

become increasingly important as the organism evolved.

constant.in the small of the back, for example, where the

Different neurons in the somatosensorycortex correspond

sensitivity is modest due to the modest allocation of cells in

to touch in different parts of the body. You might expect that

the cortex, tr,vopoints relatively close are felt as just one. By

your hand, which is a relatively small part of your body,

contrast, when the poinis are placed on the tips of the

would have neurons that register impulses in a very small

fingers, there is a sufficient number of corresponding

part of the cortex. However, just as we saw for the motor

neurons in the cortex to relay an adequately sensitive

cortex, there is no direct matching of an areaof your body to

messagethat there are two points, The allocation of the

an area of the somatosensorycortex. The hands and the

brain to coordinating parts of the body depends on the

mouth have an enormousrvastly disproportionaterepresen-

importanceof that part of the body to the,task at hand. But

tation.

horv does the brain deal with inputs t{rough specialized

This biased allocation of neurons makes sense.Just as the

organs that do not rely on messagesbding passed up the

hands and mouth claim a large allocation of neurons in the

spinal cord or on different contributions from different parts

motor cortex to enableviolin playing and speaking,so those

of the body? How do we even begin to see and hear?

same parts of the body lionize ljrrge proportions of neurons

In simpler nervous systemswith much simpler lifestyles

it important for the mouth

I!
and hands to be most sensitiveabtouch becauseeating and

there is no need for a rich tapestry of visual scenes.A frog

r,vouldbenefit little from being able to distinguish the fine

feeling things with the hands are among the most basic

detail of the Mona Lisa, for example. In the frog world, all a

human hehaviours.If you have had a local anaestheticat the

frog wants to know is if there are predatorsor prey: its retina

dentist, you know how debilitated you feel at not being

has accordingly become sensitive only to shadows, which

sensitive to movement or lgyclt in even a small area of the

mou th.

would be cast by predatorsor prey that it can eat, namely

This difference in sensitivity to touch on different parts of

irrelevant and thereforesimply not registeredby the frog's

in the somatosensorycortex

flies moving backwardand forward. Fine detail of objectsis

56

THE HUHA N

BRAI N

eyes. When presented with a piece of cork dangling on a

string, crudely resemblinga fly flitting past, a frog will make
all the predatory and gustatory movements not only of
sticking out its tongueto catch the fly but of licking its lips

SYSTEM S OF SYSTEM S

lens is a transparent, elastic convex structure that is

suspendedby ligamentsto control its shape,and this shape
can change from one moment to the next according to
whether you rvant to look a long way off or a short distance.

as well.

With the corneaat the very front of the eye,the lens helps us

As a general rule in the animal kingdom, the more

complex or relatively big the eye in relation to the body,
then the smaller the rest of the brain. More processingwill

to focus. The coloured iris, which varies greatly between

individuals, regulateslight by constricting or dilating the

go on at the earlieststages,in the peripheral organ, than in

part of the eye is filled with a watery fluid, By contrast,in

more sophisticated brains where the input will not have

already been heavily biased. Insects have compound eyes
that look a little like geodesicdomes arrangedon each side
of the head. Each eye consistsof some ten thousand clistinct
rnodules amounting to ten thousand facets all angled in
different d]rections.someoinsects
ha'e up to thirty thousand
such facets. tight is funnelled through each module, so that
thore is a huge magnification.However, in human terms the
rosults would be far from ideal as the lensesof these facets
cannot be focused.The huge advantagefor the insect is that
a large visual field is projected onto a small number of cells
withorrt the insect moving i-tshead. The more facets,the
rnolc dr,'tailedthe picture.Thh type of eye is very sensitive
tu nny change in tlie visual sceneand to the planes of light
polirrization;howe',,er,
the compoundeye cannotgive a high
tlogrocol' resolution.
'l'lru hrrman eye is very different:
it is ball shapecland
r;orrsist.s
of two main sections,separatedby the lens. The

pupil. The gap betweenthe cornea and the lens in this front
the second compartment comprising the main body of the
eye, the cavity is filled with a jelly-like substance.
At the verl' back of the eye is the retina, rvhich is the
image-detectingzone. If you look at the retina under a
microscope,there is a tangle of cells seemingly forming a
jumbled mass that looks a little like a i*

-hence the name

retina,from the Latin retusfor'net.' Thesecells in the retina

respond to changesof light by a change in an electrical
signal passed on to two more relays of cells before being
transmitted into the brain proper, via the bundle of fibres
known as the optic nerve.
The exit point where this nerve leaves the retina and
burrows into the brain is the 'blind spot' where there is
obviously no room for light-sensitivecells. The blind spot is
just to the side of the middle of the eye, near the nose.On
the other side of the middle of the eye, near the ear, is an
area of the retina called the fovea. The fovea is a small
indent where there is a high concentrationof a certain type

58

T HE I + UI . I A N BR AIN

SYSTEM S OF SYSTEM S

of cells that are sensitive to light. If light strikes this area,

be converted by the retina into electrical impulses. In the

vision is optimal as there are more cells here to do the job.

dark there is a steadyreleaseof a chemical messengerfrom

Birds of prey can have up to five times more concentrated

the rods onto the next relay of cells within the retina. When

cells in their fovea than humans. In addition, unlike

light first strikes, it is absorbed by a special chemical

(rhodopsin) within the rod. The ensuing change in this

humans, eagleshave two foveas.One, the searchfovea,is for

sideways vision, whereas the pursuit fovea judges depth,
which is done with both eyes.
Unlike human eyes,all bird eyesare fixed in their sockets.

chemical causedby the absorption of light then triggers a

cascadeof chemical reactionsinside the cell. The end result
of thesereactionswithin the rod is a changein its electrical

In order to turn their eyes, birds have to turn their entire

properties.

head and neck. Our lifestyle would be drastically compro-

It is this change in electrical properties, namely the

voltage normally generated by the rod, that changes the

mised if we were unable to move our eyes back and forth

without moving our head - imagine reading, for example!
For both the eagleand humans, however, light (electromagnetic waves) travels through the eyeball and penetratesthe
outer two layers of the retina to be processedby the lighf
sensitive cells. The particular cells sensitive to colour are
known as cones, and the other type of light-sensitivecells

message it had been transmitting as long as there was

darkness.In the caseof the other type of light-sensitivecells,
the cones, we start to'process colour_by the selectivity of
different cones responding to certaii ranges of light with
peak sensitivities for red, greeri, or blue wavelengths'
Different colours excite different combinations of these

conditions, leaving three types gl cone to respond princi-

cones in different proportions. For example, a certain

wavelength excites red and green cones in equal numbers

pally to one of the three primoryiolor-os: red, green,or blue.

and is perceivedas yellow.

Within the electromagnetic ,pr.trurr,, our human eyes

We have seenthat electromagneticwaves are convertedby

cells in the retina into electrical signals.However, the retina

are referred to as rods. Theserods are for vision in darkened

detect only a very small sectionas visible light: between400

and 200 nanometres on a scale varying between 10 metres,
the wavelengthof an AM radio, and less than a nanometre,
the wavelength range in which we find X rays and gamma
rays.
FIow is light actually registere-d
in the brain? It must first

does not just signal uniformly and equally everything in

your visual field. The image is relayed into the brain with an
enormousbias.For example,if there is a largeuniform area
within an object, then only weak signals are passed on,
whereas if there is contrast the visual signals are most

5'

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THE HU},IAN BRAI N

SYSTEM S OF SYSTEM S

vigorous. The retina is really only concerned with detecting

change. But change does not just occur in space, with

for example, she poured tea, it apparently seemedfrozen

contrasting edges; there is also change in time, namely

movement. The retina can adapt so that it no longer
responds to stationary objectswhile still retaining the ability

activity becauseshe could not ptop pouring: she could not

like a glacier. Indeed she was unable to engage in this

seethe level of the fluid in the cup rising sufficiently well to
know when to stop. This patient also said that when she

to signal for movement. To appreciate the preference of the

nervous system for statesof change, think of how a flashing

spoketo people,conversationproved a problem becauseshe

light is more noticeable than a constant one. our survival

may well depend on a change in the surrounding situation
much more than if everything remains the same.

Even worse,and more dangerous,she was unable to monitor

The eyeball itself is not a self-contained centie for visiou

rather, it is the gateway,by which the all-important signals

woman could detect movement through her senseof sound

gain accessto the brairl for further processingbefore we can

actually see. From the retina, cells send out electrical signals

Comparablesituations have been reported since World

was unaware of the movements of the mouth of the speaker'

the progressof a car: first it was in one place and then
suddenly it was almost on top of her. On the other hand, this

or touch.
War I, when people came under the scrutiny of doctors

along the fipres, exiting via the blind spot, deep into the
brain, to the thalamus,meaning'room'in Greek.This brain

following head injuries due to the wounds of battle. A

structure, which occupies a substantialpart of the middle

section of the brain (the diencephalon), then relays the

patients:he reported that there were people who could see

signalsto the visual cortex,the outer rayer at the back of the

not shape or colour. Often anyone with normal vision can

head. studies of people who have lost parts of the visual

cortex have given neuroscientistssome very helpful and

experiencethis phenomenon: if something moves in your

intriguing insights into underslanding what might happen

in this area that enablesus to-see.

you then need to turn your head to see exactly what it was

For example, a stroke victim in her forties had cells

damaged in a highly localized region within the visual

Similarly, there are those who can see form and move-

cortex. Although she could see ali stationaryobjectsas well

as anyone else, she was unablu to ,"" objectsin motion, If,

physician of the time, George Riddoch, studied these

movement,unlike the lady we have just been discussing,but

extremeperipheral vision, you are aware of movement,but

that moved.
ment but cannot experience colour. A world composed
entirely of shadesof grey would be the fate of people who
had either a deficit of cones in the retina or damage to

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THE HUMAN B RAI N

SYSTEM S OF SYSTEM S

critical regions on both sides of their head. If the brain is

only damaged on one side, however, then half the world

process vision at least partly in parallel, that is, we are

appearsin colour and half the world is in black and white.

Finally, some patients with damageto the visual system

parts of the brain. Different aspects of our vision, form,

can see movement and colour but not form. Agnosia, from
the Greek meaning 'failure to recognize,, is a condition

actually processed,at least in part, by different svstems

connectingthrough relays from the r,)tina to the back of the

characterizedby the ability to see objectsr,vithoutidentify-

head.Thus, as we saw in the caseof movement,lve can see

ing them. Agnosia can vary in its severity from patient to

patient, and even the same patient may have better form

that different regions of the brain are working together to

vision from time to time. The vision expert semir Zeki

suggestsone reason this particular condition can be so

caseseeing.The big mystery is how doesit all come together

variable:if complex forms were to be gradually assembledin

signalsconvergeinto a single entity?

our brains from less complex patterns, then perhaps this

gradual processof constructioncould be arrestedat different

processingvisual signals simultaneously but in different

colour,and rnovementseemto us a cohesive',vhole,'butare

contribute to what we regard as a single function, in this

again?Where in the brain do we make all the parallel visual
Somepeople have suggestedthat there is a convergenceof
these different pathways in certain parts of the brain, like

stagesin different people. some patients would thus have a

more extensive visual repertoire than others. Zeki suggests

railway tracks leading into Grand.-Central Station. This

that understanding and seeing are not two separat'epro_

cessesbut rather they are inextricably linked: if you see

the doctrine of phrenology discussedin the previous chap-

something you will automatically identify it. on the other

hand, if you do not seean object in front of you, Zeki argues

one or two Grand Central Stationsin our heads:if such an

that it is because there has been a collapse of the higher

integrative processesfor complex form recognition in the

completelylost. But this scenario never occurs.Thus, we

visual cortex. obviously, you will not recognizethe object.

You will be to a greateror lesserextent ,form blind.,
After considering the casesjust presented, it is evident

as a simple bundle of minibrains.The connectionsbetween

that vision of form, movement, and colour can occur

i.dependently of each other. current thinking is that we

scenariois, in a sense,a late-twentieth century version of

ter. Just imagine that we possessedthe brain equivalent of
area were damaged, then it follows that vision would be
have yet a further example of how the brain is not organized
brain regionsare not directed to convergeinto an executive
centre but are more likely to take the form of balanced
dialoguesbetweenkey brain regions,comparableto what we

t
;

saw for the control of movement.

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THE HUMAN

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Yet in itself this scenario of interactive, parallel brain

regions does not solve one of the greatest mysteries of
neuroscience:how do we actually see?On the one hand,
great strides have been made in understandingthe complex
steps in the visual processingof an object: it is now known
what parts of the brain, during vision, are active at what
times and under what conditions. But such responsescan

SYSTEM S OF SYSTEM S

interaction, is more critical than any individual region on its

own. Harth suggeststhat not only are signals relating to the
sensesconveyedfor processingto the cortex but in turn the
cortex can send signals that intercept this incoming stream
of information, to modify it. The more vigorous the cortical
interception, the more idiosyncratic and remote from objec-

persist even when the brain is anaesthetized,when there is

tive external events will be the final conscious experience'

Zeki also makes uses of these feedback pathways in his

no consciousnessat all. No one has yet pointed to a single

interpretation of blind sight.

event that occurs in awake but not anaesthetized brains

where the intervention,.of consciousnessinto the visual

Zeki suggeststhat blind sight is due to a rupture in this

balancedcircuitry. The processingof the signalscan still be

process could be identified as an unambiguous physiologi-

relayed into the brain, but conscious vision is abolished

cal/anatomical mechanism or event.

since certain pathways normally sustaining dialogue upon

The riddle is compounded by observationswhere there is

a conspicucius dissociation of brain visual processes and

dialogue between brain regions are no longer operational.

However, this idea does not square entirely with a particu-

conscious awareness in fully awake patients. The first

larly interesting observation concerning blind-sight patients:

condition was again reported initially as a result of head

the actual degreeof physical damage to certain brain regions

wounds in l,Vorld War I and was later dubbed hlind sight in

the 1970s.Blind-sight patients are blind in a certain part of

need not be the only factor in determining how a blind-sight

patient will respond. In certain casesblind sight can be

their visual field, but if asked to 'guess',they can neverthe-

reversed by other factors: for example, if an otherwise

less point to objects, which they claim they cannot see,

stationaryobject is made to move. Perhapsthen, for a final

placed in that area.Clearly the brain is still functioning, but

consciousnessof seeing an object, not only are intact

the consciousnessis lost of actually seeingthe object.Some,

neuronal circuits important but also the particular proper-

such as Zeki and the physicist Eric Harth, have argued that

ties of the object in question.

the anatomical integrity of neuronal circuits is all-important.

Another example of brain damage that leads to vision

As we saw earlier, brain regions can be viewed as partici-

deficit is a condition that is the reverse of blind-sight:

pants on a seesaw,where the balance between them, their

prcsopggesill from the Greek meaning 'failure to recognize

iii::f
i

I
.i

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ir

66

THE HUMAN

BRAI N

SYSTEM S OF SYSTEM S

faces.' whereas blind sight entails recognition without

that we learn through experience to distinguish sound from

awareness, this condition is one of awareness without

sight; another idea is that each sensory system is linked

recognition. Patients can see faces as faces, but cannot

recognize anyone, even themselves.There can be a marked
difference if a face is made psychologically ,stronger' by
presenting faces that are linked. For example,if a picture of

preferentially in some way to certain types of movement,

which emphasizesthe distinction.
However, there are well-known examples of where this
distinction between the sensesfalls down, a mixing of the

Princess Diana is presentedafter that of her ex-husband,

sensesknown as synaesthesia.People displaying synaesthe-

Prince Charles, a patient can often recognizeDiana,s face.

sia may claim to 'see' certain musical notes in certain

Again we have an example of consciousnessbeing depend-

colours.Virtually any combination of two of the five senses

ent on more than one factor, but we still have no idea how
such factors lead to the seemingly magic step within the

is possibie,although it is the experienceof different colours

upon hearing different sounds that is the most common.

brain that enablesus not only to prbcessan object registered

Synaesthesiatends to occur more in childhood, but can

by our retina but to consciouslysee it as well.

often be triggeredin adults with psychotic disorderssuch as

For all the sensesthere.isalso the enigma of the nature of

schizophreniaor by hallucinatory drugs. The division of the

the first-pqson subjectiveconsciouselement.There is much

more to hearing, for example,then mere vibrations. We do

sensesthen is clearly attributable to some aspectof normal

67

I
I

,
I
I
I

brain organization, but an aspect that is not immune to

not hear a symphony as vibrations any more than we see a

face as lines and contrast. Rather, our perceptions are

individual perturbation. One poss'ibility is that there are

unified wholes, shot through with memories,hopes,prejudices, and other internalized cognitive idiosyncrasies.

extend not only from the sense organ in question to the

II

Another tantalizing and related mystery of the brain is

why electricalsignalsarrivingin the visual cortexshould be

another cortical sensearea as well. This idea, horvever,is

experienced as vision, while exactly the same kind of

electrical signals,arriving in anotherpart of the brain such

synaestheticexperiences,namely that such statesonly occur

as the somatosensorycortex or the auditory cortex,should

the cortex not allocatedto the primary processingof each of

be perceived as touch and hearing respectively.No one has

the individual senses,namelv the associationcortex, some-

yet given a satisfactoryexplanation,although one idea is

how play a part.

additional connectionsin the brain of the synaestheticthat

cortex appropriate for that modality but also innervate
not very likely as it would not account for the variability in
under certain conditions. It is more likely that the areasof

T
t
t
t

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THE HUMAN BRAI N

SYSTEM S OF SYSTEM S

We saw in Chapter 1 that even compared to the brain of

we moving about; at the other end of the spectrum, high

our nearest relative, the chimpanzee, the a-reasof human

arousal leads to distracted behaviour where we overreact to

brain classified as associationcortex are vast, It is possible

a minor occurrence and move about in a restless,prupose-

that inputs from association cortex into the areas of cortex

less way. Psychologists have long found that we are most

devoted to particular sensesmight in some way be aberrant.

efficient at performing tasks when we are in the middle

Certainly such a scenario would account for the greater

range level of arousal. Arousal then is an important consid-

predominance of synaesthesiain children, before distinc-

eration to our prevailing state of mind.

tions betweenthe sensesare learnedand where the neurons

We are familiar with the extremes of arousal: when we

of the brain are less hard wired and thus more flexible and

sleep, arousal levels are low; if we are over-activeand easily

versatilein their operations,as we shall seein Chapter 4. A

distracted, with a racing heart and sweaty palms, we are

malfunctioning of plfsiology

(the working of neurons)

over-excited or over-aroused.Arousal is with us all the time,

rather than anatomy (their physical connections)would also

to varying degrees. It is governed by distinct groups of

explain why synaesthesiacan suddenly appear in the brain

different chemicals in the brain stem that predominate at

of a schizophrenic. On th.eother hand, any real explanation

different times of the day and night, or with emotions or

of synaestfresiais impossible as it hinges on a subjective

illness, and that send signals up to the large regions of cortex

perspective: the first-hand experience of an individual.

to modulate, on a global scale, the workings of many circuits

Synaesthesiais a facet of consciousness,

that ultimate riddle

of brain cells. One measureof arousal can be monitored as

of the brairr.

changesin the ensuingelectrical activity averagedover large

So far we have seenhow that private inner world of one's

areas of the cortex.

own consciousnessis influenced by sensoryinput pouring

As long ago as 7875, weak electrical currents were

in and reflected in an outpu_!movement. By being able to

recorded from the brains of rabbits and monkeys by an

receive detailed and incessdntinformation about our envi-

English physiologist, Richard Caton. The finding triggered

ronment, and respondingquickly and appropriatelyto each

little attention. Fifteen years later, however, controversy

individual situation, we are in a constantdialogue with the

flared between a Polish physiologist, A. Beck, and an

outside world. A further factor in determininghow vigorous

Austrian, E. FleischelVon Marxow, who each laid claims to

and effectivethis dialogue may be is level of arousal.When

discoveringelectricalactivity in the brain. It was only when

asleep,we perceivenothing of the world around us, nor are

Caton settled the matter by pointing to his much older

59

'r;i

70

THE HUMAN BRAI N

SYSTEM S OF SYSTEM S

publication in the British'Medical lournal that the finding

was duly recognized. The clinical implications of this

method has involved connecting a person's EEG to a toy

discovery were not to be realizeduntil fifty yearslater, when

when alpha waves are generated.People can learn to put

in 1929 a German psychiatrist,Hans Berger,first attempted

themselvesin the correct state for making the train move. In

to record the electrical currents in the human brain.

contrast,if you are excited and aroused,then the pattern of

electric train, which only actually travels along the track

When electrodeswere placed on the surfaceof the scalp,

electricalwaves changesto one where the neuron is working

it caused no pain whatsoever - the person was totally

less as part of a whole group and instead acting more

conscious and yet different types of brain waves could be

autonomously.

detected.Berger was convinced that these signalswere due

The pattern of EEG can also vary according to age.

to psychological energy, r,vhichhe referred to as p-energy.

Electrical activity has been recorded from the mot}er's

This was the start of a technique that is still widely used

womb as early as the third fetal month. It is only in the sixth

today in neurology: the electroencephalogram(EEG).Con-

fetal month that the EEG changesto one of distinct, slow,

trary to Berger's vision of some special sort of brain/mind

regular waves. Until children are ten years old, two very

energy, the EEG records Wavesof electricity generatedfrom

slow rhythms can be detected: one is 4 to 7 waves per

hundreds df thousandsof brain cells iust beneaththe surface

second ('!bg!glhy!h-');

of the brairt.

second, but this 'delta rhythm' is never seen in healthy,

Not only does the EEG show what brain waves look like,
but it shows how they vary: the pattern can actually change

the other is as slow as L to 4 per

awake adults.

according to different arousalstates.If you are relaxed and

The EEG is important for studying not iust normal brain

activity but brain disorders as well, such as epilepsy. In

conscious,then slolv \,vaveswi4be generated,mainly at the

epilepsythere is a miniature brain storm of certain groupsof

back of your head. This p.oc-e&.iscall"dglpha .hyth-; yo.,

brain cells that can lead to convulsions. This virtual

can actually make yourself generate alpha rhythms by

explosion of electricity can be detectedon an EEG used

relaxing. Increasing numbers of people have problems

subsequentlyby neurologiststo locate where damagedbrain

relaxing, giving rise to one of the greatest scourges of

tissuelies.

modern times: chronic stress,.One way of helping such

Another use of the EEG is to provide a fascinatingwindow

individuals to learn to relax is _tolet them know when they

into what bqBIerE in the bfeg when we go to sleep. There

are generating alpha rhythm. A particularly ingenious

are four stagesof sleep, distinguished by different patterns

7I

72

THE HUM AN BRAI N

SYSTEM S OF SYSTEM S

of electricity recorded from the scalp. When we fall asleep

we descend very rapidly from level 1 through these four

sleeping time, it presumably has some value. There are

several theories on whY we dream.

stages,down to level 4. Throughout the night, we gradually

One theolcyis that since the. brain is no longer tied down

surface and descendagain through these four stages.

and restricted by the apparent reality of the messages

As well as the four stagesof sleepthrough which we cycle

severaltimes a night, there is also anotherstageof sleep that

coming from the outside world, it startsto, in a sense,freewheel. This situation may be analogousto a day off from

is totally different. It is in this stageof sleep that our eyes

move rapidly backwardand forward - henceits name,3gg(!

work with no structure to our activities. However, thete

must be more to dreaming than the brain just playing

elremovement or &EM sleep.Ifpeople are awakenedduring

around; there is evidence suggestingthere is some definite

REM sleep, they usually report that they have been dream-

benefit for the dreamer. If people are woken up when their

ing. It is easyto imagine that the darting eye movementsare

EEG shows that they are in REM sleep, and thus probably

a result of looking at imagesthat move about in our dream

world. Interestingly,during this dreamingstateof sleep our

dreaming,they try and compensatethe foilowing night: the

amount of REM sleep that they undergo increases.In one

EEG is just the sameas wlren we are awake,unlike when we

are asleep,in dreamlesssleep. However, in normal sleep,

registeredREM sleep. They were woken up to ten times on

when we are not dreaming,we might be tossingand turning,

the first night, but by the sixth night they were woken up as

but in REM sleep our muscles become paralysed. This

many as thirty-three times as their brains tried time and

immobility is important becauseit prevents us from acting

out our dreams.

again, in vain, to plunge into the dream world.

Different animals display different amounts of REM.

Reptilesdo not displayit at all, birds do occasionally,but all

with problems and consolidate whatever has happened

mammals, at least accordifg to their EEG, would seem

capable of dreaming. In an averagenight,s sleep of some

dreams may have come to serve this purpose, it seems

unlikely that it is their prime purpose.The fetus at twenty-

sevenand a half hours,humanscan spenda total of one and

six weeks spends all its time in REM sleep, yet has no

a half to two hours dreaming. The longest recorded single

experiencesto consolidate or resolve. Dreaming time then

period of continuous REM sleep is about two hours. Given

declines gradually through childhood. This observation

that REM sleep clearlyoccupiesa significantportion of our

suggests that dreaming represents more a state of the

experiment, people were woken up every time their EEG

Another idea is that dreams enable us to come to terms

during the day. Although in adults it is easy to see that

73

:1gii

$ql

71

THE HUMAN,BRAI N

functioning of an immature brain, where neuronal circuits

SYSTEI.IS OF SYSTEM S

are still very modest. Perhaps dreaming is a type of

world into which our species evolved thirty thousand years

ago,the sleeperwas highly vulnerable to attack from passing

consciousness resulting from a less vigorous dialogue

predators.Sleepmust thereforehave some enormousbenefit

between brain regions, caused in turn by the fact that the

in order to debilitate us in this way for up to eight or so

connecting fibres are still becoming established.

hours per night. It is now known that during sleep the brain

If true, this idea could have two very interestingimplica-

makes proteins at a much faster rate than when we are

tions. First, it suggeststhat when we are in REM sleep, the

awake. Proteins are large molecules that are essentialfor

degree of comrnunication between regions in our brain

maintaining the structure and which underpin the func-

becomes far less. Second, it has been noted that the

unconsciousnessof schizophrenicsis frequently very simi-

tion of all the cells of the body, including neurons. Sleep

gives us a chanceto stockpile the chemicalsthat seemto be

lar to the illogical but very real consciousnessof our dreams.

vital for our brain to function properly. But functioning

Hence it might be the case that a central problem in

properly does not refer only to the processesof which

schizophrenia is a reversal to a reduced communication

we are conscious,such as learning and memory, but to the

btween regions of neuronq,leading to a dreamlike view of

the world. Although the function of dreaming might end up

unconscious processesas well, such as those regulating

being one of consolidationof our problems,it is more Iikely

We normally use only a part oy'the energy defived from

that dreaming arises as a result of certain states of brain

food and oxygen for immediaib conversion to heat. The

activity that cannot processlarge amounts of sensoryinput

remaining energyis stored for all the other vital functions of

becausewe are asleepor becauqe.,,as

in infancy, the brain is

the brain and body. If people are allowed only three hours of

underdeveloped,or because,asin schizophrenia,prevailing

sleepa night, many of thesefunctions start to decline within

chemicals have limited the efficacy of large-scaledialogues

a week. If we are deprived of sleep, energy is not stored

over large banks of brain cells. Again, however, dreaming is

efficiently;more is immediately squandered,dissipatedas

a further facet of consciousness,and thus its causeand its

heat. Thus, persons who are continuously and completely

function can still only be extreme conjecture at the most.

sleep deprived would eventually, literally, be burning

temperature.

But what are the functions of ordinary sleep,when we are

themselvesout. If rats do not sleep for long periodsof time,

completely unconscious? This--is an important question

they gradually need more and more food to restore their

becausesleep is a pretty risky bu-siness

- in the Cro-Magnon

energy. Eventually such rats die, underweight and

75

16

THE HUMAN BRAI N

exhausted, despite their \uge food intake. Sleep then is

vitally important.
Another intriguing feature of the brain and its arousal
systemis that usually the brain'knows'when to fall asleep.
At least in a number of non-human animals there is one area

SYSTEH S OF SYSTEM S

sequenceof events might seem at first glance to be of little

relevanceto us sophisticatedhumans, it is worth noting that
in the United Statesmelatonin is proving to be a popular
treatment for iet lag. A tablet of melatonin taken just before
sleep in the new time zone might ensure that sleep occurs

of the brain that plays a very important part in sleep and

waking: the pineal gland. The pineal gland lies deep at the

rapidly and for a decent periorl of time.

centre of the brain: unlike most other brain structures that

a variety of factors. We can soe just how important these

outside clues are from experimr:ntswhere people have been

are duplicated on eachside of a midline, the pineal gland is

In humans, the sleep/wakecycle is normally controlledby

right over this midline, straddling t\e middle of the brain.

For this reasonthe philosopher$dn6 Descartes,over three

put in cavesand left entirely to their own devices,free from

hundred years ago, thought that the pineal gland was

actually the seat of the soul. Descartesargued that since

David Lafferty, a former Royal Air Force officer, answered

an appeal in the Daily Telegraph in 1966 for a volunteer to

there are not two pineal gla,ndsand since we have only one
t

live in isolation in a cave 3S0 l'cet underground for at least

all the demands of the outsido world.

soul, the nileal gland must be wherethe soul is located.

Today we know that the pineal gland is important in
regulatingsleep and wakefulness.Birds are stimulated

l00 days. In return he rccoived 100 ($roz;, plus an

additional 5 ($8.35)per day for.each subsequentday spent

directly by light through their skull: we know that even

a

underground for 130 days. At the end, doctors were sur-

underground. Lafferty set a new record by remaining

when the pineal gland is isolatedfrom birds completely and

prised at Lafferty's good physical and mental health. Laff-

put in a dish, it is still sensitiveto light. The pineal gland is

not responsive when it is already light and becomes dark;

erty was surprised when he honrd how long he had been

however, when it is dark and Jrt,ddenlybecomeslight, the

cockerel awakens. The pineal gland secretesthe hormone

hour schedule,causing hinr to trnderestimateslightly the

melatonin. This substancefluctuatesin the brain according

to time of day. when levels are high in the brain, sleep

underestimateseemsto occur rr.ht:npeople are isolated,so

ensues;in fact, when melatonin is injected into sparrows

they sLrbseqrrently
fall asleep. Although such a simple

but we also need to fine-trrnt it by using cues from the

underground. His biorhythms hnd settled into a twenty-five

time he had spent undergrorrnd. Generally, this modest
clearly we do have a basic antl rirther regular internal clock,
outsideworld,

Tf

irr:i
rtl'

78

THE HUMAN

BRAI N

Sleeping and waking is not the only rhythm that is

controlled by the brain or registeredin the brain. There is a

II

SYSTEM S OF SYSTEM S

ancient Chinese art of acupuncture. The basic idea of

acupunctureis to restorethe equilibrium in the functional

very macabreexperimentshowing a less obviousbut impor-

state of the body so that the so-calledlife force, 'Chi,' is in

tant daily rhythm: sensitivity to pain. Amazingly enough,

perfect balance between the various organs. The basic

people volunteered to have electric shocks through their

teeth at several times throughout the day and night; they

procedureinvolves inserting a needle 1 to 4 millimetres into

anv one of gos special points on the body. Acupuncture is

then reported on horv much pain they felt. It might be

used for many purposes- for example,to help people quit

expectedthat thesepeople experiencedthe same amount of

smoking. One effect particularly relevant to our discussion

pain all the time. Surprisingly, the pain was perceived as

is that acupuncturecan also relievepain. An idea as to how

almost twice as bad during certain times of day, most

the analgesiceffectsof acupuniture were discoveredrelates

notably in the morning, ]ust after lunchtime, the pain

to the time of ancient warfare, when people fought with

seemed to be far more bearable.

bolvs and arrows. As arrows were removed from iniured

This study gives us yet another clue as to the functioning

soldiers,they were frequently twisted in the lvound, rather

of the brain. We can sqe that experience of pain is a

as acupuncture needles are today. Paradoxically,the sol-

subjective phenomenon, that it can change according to

diers sometimesexperienceda relief from their pain.

something that is happeningwithin our brain. That event in

The relief of pain fror+acupuncture can sometimesbe so

the brain must, in turn, be changeable.Pain is usually

effectivethat surgerycan actually be carried out. To attain a

caused by something damaging,or perceived as damaging,

perfect level of analgesia,the needles will have been in

that is directly in contactwith some part of the body. We

acupuncture is
place for about twenty minutes. ,.Somehow,

saw earlier that signalsrelatedto pain were conductedvia a

.rlq1-Ql[9lfngwith

specific route up the spinal cord and into the brain. The

wherebv nerves in the skin from the affecled

signals were carried by nerves that did not change their

signalsto the brain. The analgesiceffectof acupunctureis

physical properties and hence their efficiency for conduc-

abolishedif the nervesin the skin where the needle enters

tion of electricity did not vary accordingto the time of day.

are actually treatedwith a local anaesthetic.

^lt seemsthat
when a nerve is stimulated rnechanicallyhy inserting the

Some other factor must be at play to account for the diurnal

variation.
One of several ways to study pain is to consider the

the- nor

there is a chanqe*in how rrain is nerceived in the

^geedLg,
brain. Because the analgesic effect takes some twenty

7'

80

THE HUMAN BRAI N

SYSTEM S OF SYSTEM S

8I

that in
harnessed within the brain to transmit the signals
time to
turn underscorethe functions of our daily lives. It is
send
explore what brain cells actually are and how they
signalsto each other'

minutes to come on once,theneedlesare in place, and lasts

for an hour or so after withdrawing the needleS, .S
possibility is that the needleson their own are not direetly
responsible, but rather that they must be releasing some

in turn is able
combat pain. Perhapsit is this chemical that is susceptible
to diurnal fluctuations and is even modifiable by drugs.
It was one of the greatestrecent discoveries in neuroscience,in the early 1970s,that the brain possessesits own
morphine-like substance,jlrkephalin. When this substance
is blocked by drugs, their pain perception increases and
acupuncture is less effective.By the same token, morphine
mimics this natural chemical, fooling the brain into believing that very high levels.of enkephalin have been released.
There is 4o pain centre in the brain; rather, enkephalin can
be found in a variety of locationswithin both the brain and
spinal cord.
Having failed in Chapter 1 to allocatea function to each
particular brain region, we started in this chapter with the
reverse strategy:to see how particular functions could be
accommodatedby the brain. We have seen that in all cases
several brain areasare active in parallel to enable us to
interact effectivelywith the outside world. In each description it became obvious that brain electricity and brain
chemistry somehoware vital componentsin the successful
functiclning,in general,of our sensory,motor, and arousal
systcnrs.We have not yet exploredjust how theseforcesare

t.

rt.

y'1b

i,r &./

<
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o'zt"ty'7er1'e

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lfrt4

C H APT E R 3

P U LS E , IMP U LS E

A great advance in neurosciencetook place in an Italian

kitchen in't872. Cemillo Golgi (1843-1926),a young medical graduate of Pavia University, was so fascinated by the
brain that he had set up a makeshift laboratory. The problem
plaguing Golgi concerned the very essenceof the physical
brain: the matter of which it was composed. At that time,
although the brain could be sliced into small slivers and
placed under the microscope, only a homogeneous pale
mass could be detected.Uatil its basicbuilding blocks could
be identified, it would prove impossible to discover how the
brain worked. Then one day, so the story goes, Golgi
accidently knocked a block of brain !.to a dish containing a
solution of silver nitrate, where ll rcmainsfl lost for seVeral
weeks. It turned out that Golgi had discovered a critical
reaction. When he retrieved the brain block. a bansformation had taken place. Under the microscope there appeared
to be a complex pattern of dark blobs suspendedwithin netlike tangles. We now know that once brain tissue has been
placed in silver nitrate for three hours or more it is possible
to visualize the most basic component of brain tissue: the
special type of cell refened to as the neuron.

s w 2ef9% fuo
84

T HE HUM AN BRAI N

b";Ll

of neuaos PULSE'tt{PUtsE 8s

'
i
j

i:
'
:'

,.^
^
^7 "St
t",
".Qo

.t
*)
r.V
t+

Even mcjremiraculous abgg!Golgil_s

discovery_was_thatby
-orie-is"'still
a. capricious - procesr of which-no
compietely

these parts are named'lendrites, after the Greek for 'tree.'

sure, the stain randomly marked out only one cell in every
ten-to a hundred,"so that it eppear-edblaCk againsi a pJe,

in denlity, and emanatefrom all around the neuron, giving

amber background. If every neuron had been stained, then

one or both ends of the cell body. According to the extent of

the delicate and complex silhouette of the cell would have

been buried by overlapping parts of other cells - the whole

dendritic ramification, neurons wiil vary enormously in

view of the brain tissue under the microscope would have

been transformed into almost unifonn blackness. Because

for neurons in the brain.

only one to tgn per cent of cells react to the Golgi stain,
however, these neurons stand out in stark contrast.

have a single, long, thin fibre stretching from the cell body.
This fibre is called an axon and it is many times longer than

What do neurons actually look like? tn dl casesthere is a

sgua! blob-shaped region known as the cell body or Eg,

the rest of the neruon. The normal range for diamet*

(from the Greek for 'body'), measuring some forty thousf s pillimehe in diemg[s1.Actually, the shape of
2andths
the soma is usually not as ambiguous and amorphous as a
blob, but cen come in any one of several characteristic
shapes - for exiimplS, rormd, oval, triangular, or even
fusiform (shaped like an old-fashioned spindle). It is the cell

rn" a;;*,;, oi. rr"*ffil,

oooJoo; ti";;t, ;

it a starlike appeamnce;alternatively, they can sprout from

their overall appearance:there are at least fifty basic shapes

Not only do neurons have these minibranches, but most

PY.JI
I
cell body is some twenty to one hundred thousandths'of a
^
millimetre; however, in an extreme case,a fibre such as one Y6'J
t0
running down the human spinal cord can be up to a metre
long!
fust by looking at a neuron, it is quite easy to tell the
difference between these two special features.Axons are far
harder to see,even under a microscope,since they are much

body that contains all the life.support organs for the neuron,

thinnsl than the relatively stubby, branching dendrites.

"that they tapef
-- --- ---Elendrites'are'like true'branches on a-tree in

cell in the body. However, when neurons are compared to

at the ends, whereasa;consdo not. The overall appearanceof

other cells, a very big difference is revealed as soon as one

a netuon then is of a squat central region with relatively

looks beyond the soma.Unlike other cells, there is more to a

stubby microbranchesprotruding and one long, thin sband

neuron than just the cell body.

snaking away. How might such a strange object be the

Almost as though it were some kind of microscopic tree,

tiny branches grow out of the neuronal cell body. In fact,

io,ni

building block of our personalities,hopes, and fears?

Since the cell bodv contains similar sets of internal

T
T
T
T

86

THE HUl. I AN BRAI N

PU L SE, IM PU L SE

apparatus to that of all cells, it is easyto

th.gtat leaqt
pl-o3_g_qe
sbme of its functions are i" ;;;
th"t the cell stays alive
and manufactures the appropriate ghemiclls, However,
the

,'li
!;i

i,i
iil

'i-

ggmmunication, and -how much communication between

one neuron and another cari be distorted by drugs.
ln neuroscienceresearch,this type of approach that starts

role of r,*ons and dendrites is not so obvious,

since their
very existence is excrusively linked wittr the
specific functioning of neurons alone. Moreover, such clear
physical

Jvith a, single neuron is referred to as bottom-rrp. The

strategy is to start at the bottom, with the most basic

differences between axons and dendrites suggest

that they
have very different roles to play.

betweenone single neuron and anothercan finally be built

When we saw the power of the sensitivity of the

electroencephalogram (EEG)in mirroring changing brain
statesin
Chapter 2, we ffrst encountered the idea that neurons
cen

up studv is the top-down approach. Here the underlying

theme is to start at the top with a macro system,be it a brain

generate electricity. The dendrites act as the

receiving area
for these signals, like some vast dockland zone
receiving
goods brought io by a variety o1 ships.
Just as the goods
might be unloaded from the docks and dispatched.
along
routes converging on somecenbal factory, so these
disparate
-srgnalBalti conducted along'the"a""arit"s
nl
"o"o"qgi"fo"
cell body where, if the signals are sufficiently sbong,
a new
electrical signal - o*to continue the anarogy,a new
product

gelglgle* -Iben*the -axors-aoee-inte-play:-rhey-**

;-Y$l --U-9.
conduct this new electrical signal away from the
celr body
and on toward the next target neuron in the circuit,
iust as
the product of the factory is exported to some
faraway
destination. In this chapter we will took g{-ho*-gcll1ggq
send and receive electrical signals. We will ,lro
g.

ho*

87

component,the neuron, and then seehow communication

into a complex,working whole. Ibe oppositeof the bottom"-

Eglg (Chapter 1) or q_function (Chapter 2), and see how it

mrght fft into the brain operations by travelling down,
analysing it into smaller subsystems.Often, neuroscientists
are divided on the repetitive merits of these forms of study.
In a sense, we have already been using the top-down
strategy in the first two chapters, so we are faniliar with its
advantagesand limitations. bn this chapter we turn to the
reductionist approach based on the single neuron.
tuigi Galvani (7737-1798) was &e first to demonstrate

-t[e
6at'nbiVrilAmanttingfrdm
g"*;i"
tpinJ cord
"o"fa

electricity. During a thunderstorrl, Galvani laid out frog's

legs on a metal plate. Surprisingly, the frog's legs twitched
as the thunder and lighhing resounded. Hence Galvani
concluded, wrongly as it turned out, that all electricity lay
in living tissue. He envisaged that muscles were containers
and nerves were the conductors of electricitv. lnstead. it was

"

88

'l

THE HUM AN BRAI N

the nineteenth-century pioneering physicist

Michael Faraday
who realized. hoy
fu}dhmenJal a phenomenon electricitv
really was. He concluded from his
experitrrentswith non_

m-alerial
its-source;.is:":::-,-::,;-:,--,,,,,,F^lgJ9gigal that 'electricity,--whatever
identical in nature.' Nerves were indeed
a sourceof electric_
ity, but with no particular monoDolv.

'::1

i:,lllt,1;

.;ir,$

An electric cu'ent is, literaily, the flow

of charge.In the
brain, such flow will occur due to the
movement of any one
of four everyday ions (atoms either racking
an erectron or
possessingan extra one): sodium, potassium,
chloride, or
calcium. These ions are distributed either
insrde [potassium)
or outside (sodium, calcium, chloride)
of the neuron, but
they cat'''ot readily enter and reave
randomly. Rather, a'
four ions are kept in one place by a barrier,
the membrane of
the cell' This membrane is not just
a simple wall, but
consists of two layers wi& a fatty
middle, like an oily
sandwich. Becauseions cannot penetrate
the o'iness of the
-""- inhpspitable middle-sf the"membmne_ofthe ueuroa;lio
ions
will be able to move freely into or out
of the neuron.
Consequently, ions amass inside and
outside of the
neuron. There are also other negatively
charged proteins
proteins are taken into
consideration, the net d.istribution of
chargeon either side of
the neuronal membrane turns out to
be uneven; there is not
an equal number of pluses and minuses.
The inside of the
neuron is negative with respectto the
outside. There is thus
a potential difference _ a voltage
generated _ that is

II
i

?
i

!
d

I
*
*

PU L SE, IH PU L SE

expreg-sedas a negative value q+d is usuaJfy-abou!

7/0 or
-80 thousandths of a volt (mV).
However, there is little point in having this potential
differgiice if ttib ions rein.ain trappbil in one place and are
unable to-flow, to give rise to a real electric current. Imagine,
for example, a dam that has enormous reserves of water
stackedup on one side,which you are nonethelessunableto
use. For a cell to generatean electrical signal, a current has
to flow, just like water has to be unleashedfrom the d.am.In
order for the current to flow, ihe ions need to move
temporarily into or out of the neuron. But how is it possible
for the ions to travel through the impenetrable oily middle
of the membrane?
The membranebarrier cAn, after all, be breached.Various
special sEuctures made from large molecules - proteins
span the two layers of the membrane:they serve as a bridge
-. .. . for-.a particular isn from one watery, non,oily zone (the
outside of the neuron) to another {the interior of the cell),
However, since these protein passagewaysare inserted
snugly through the middle

of.the membrane, a more

parlance it is actually referred to as a channel.

For a neuron to send an electrical signal, positively
chargedsodium ions are briefly able to enter the cell, thus
making the potential difference temporarily more positive
inside than outside (depolarization). However, as soon as

8'

THE HUI ' I AN BRAI N

'O

PU L SE. II.IPU L SE
'I

this voltage becomespositive, say to +20 millivolts,

posil

liyslv chargedpotassirrmions.-thenleave"the cell,-actually

-

making the voltage temporarily more negative thaa

normal
(h"yp"rpolarization|.Hence,
when theneuron is activated in
"
this way, there is a brief and characteristic change
in the
potential difference,a positive pulse, followed by
a negative
overshoot. This transient positive_negative wave
usually
lasts for about one or two thousandths of a second
and is
known as an action potential. The action potential
is
distinguished from the vortage(the resting potential)
that is
normally generatedall the time when the cell is not
sending
a signal. (See Figure S.)
Why should the sodi
'm channel suddenly open in the
first place? To pose the question in another way,
What
triggers an action potential? After a[, it would fs gempletely
pointless and almost a paradox to have a signal that
was
generated completely at random. Imagine the
telephone
*inging at odd times throughout the night, but theie being
no
caller on the other end of the rine. Let us return
to the
dendrites, the branch-rike processessprouting out of
the cel

j;:r'*Jt;:;T

FIG URE5
Actual recordingof a responseof a neurou in a rat brain (the hippocampus)to
stimulation (shown by an asterisk). Stimulation at this intensity can rezult
either in an EPSP(excitatory postspaptic potential) or an actiou potential. In
casesof action poteutial generation, a sharp chengein voltage can be seen
arising frm the peak of EPSP.1te action potential is the electrical signal via
which neumns communicate, the membranepotential (the potential difference between the inside of the neuron and the outside) becomesbansieutly
more positive (depolarization) as positively charged sodium ions enter the
neuron Oncethe cell is depolarized,this newvoltage causesttre opening of
potassium chan:rels,allowing positively chargedpotassium ions to leave'the-cell (repolarization). (Figure courtesy of Fenella Pike, MRC Anatomical
NeuropharmacologyUnit, Oxford.)

body. Dendrites act as receiving,*ll$9glf_o:

Ugo_ging_ _
-^,

signals from other neurons. If the signals are strong

and/or
sustained, they will be conducted along the branch
of the

central cell body at any one time. This barrageof signals at

dendrite down to the cell body, like a domestic elecbic

current down a very imperfect leaky cable.

generating an electical signal, an action potential, of its

Like some massive and frenetic railway station,

tens,
hundreds, or even thousands of signals can convergeon
the

make up one grand super-value,one final changein voltage.

any one moment will be contributing to its likelihood of

own. As these incoming signals arrive in the cell body they
If this new net difference in voltage is large enough within

TH HUM AN BRAI N

PU L SE. I].IPU tSE

the target cell, it causesthe sodir+mgh,^.l4els,

_whichare
.r'anges-to
to
*o." positive vbltage;to ciliiiii.-""""-].'q-nsitiv-e
"
Onceagaina new actionpotentialis generated,
this rime in
.$e se-condneuron. _: , _.An action potential for any given neluon
is always the
same size, typically some g0 millivolts. But
such consistency posesa problem: If the incoming signals
becomemore
numerous or stronger, then how is the receiving
Deuron,
limited to generating only one type of signal,
ever going to
convey these differences?Becausean action
potentiar cannot be made any bigger, tbe neuron will
instead generate
more and more action potentiars as the signals
it receives
become more intense. When this happens,
a neuron is said
to be excited. The way a neluon signals
more or less
vigorously is reflected by a change in the frequenry
of action
potential generation. Some [eurons can
ffre up to five
hundred action potentials a second (500 Hertz),
although it

!.-.--=*------ - -.is-more..norma]
to-recoftriat6s-of-so-me
d0 totoo H;.
t:

neuron that only generates one or two action

potentials a
second is considered to be slow ffring

thg actiga pofential =_i1 trllpmitted

93

varigs acgording to-,the

diameter of the axon and whether or not it is insulated with

a !,-atty +e._qq kooyq as myefl4. ff the, myelin sheath
deteriorates,neflre fibres will conduct electrical signals less
efficiently, as is the case,for example, in multiple sclerosis.
Normal movements are so fast and automatic that it is hard
to accept a delay between a thought in the brain and a
muscle contraction. The effortless rapidity of our brain
processesand our movements canbe explained by the speed
of nerve conduction, up to approximately 220 miles per
hour!
Even though it might be clear how a signal is generatedin
a neuron and then sent down its axon, it is far from obvious
what happens next. We now need to know how one neuron
actually makes contact with another,in order to pass on the
signal. Ever since nen/e cells have been stained and visual.ized, scientists have--pondereilthis problem.'For exartple,
Golgi thought that all neurons were joined together, a bit

like a hair net. At the time he was to meet with ferocious

opposition from the great Spanish anatomist Ramon y Caial.
Mostneuronsinmgjrl*-ltllg,9rr.,e,r31tg=ap00*4-Ps:tmtia]s-,--rrr::;:lFh:jes:*truo-ptbueernamoseientstg-p-olelizdd'-mftt -a longl--*"'i.:;;:=**.i:i-'-:+1.*
- -.- -. " :-'
rn this way in order to communicate with
their respective
standing conflict because Cajal, contrary to Golgi's vision,
target cells. The next vital step is for the action
potentiar to
was convinced that there was a gap between neurons. The
reach its intended destination.
Just as the dendrites act as
issue was not to be defrnitively resolved until the 1950s,
the receiving area of the neuron, so the
thinner, solitary
when there was an awesomebreakthrough:the advent of the
txon serves as the output path for exporting
electrical
elecbon microscope.
signals onward. The speed at which the electrical
signal _
The elecgon microscope enablesthe study of cells with an

I
I
:.1 .:

94

T HE HUM AN BRAI N

PU tSE. If'IPU L SE

astonishing magnification factor. A light microscope,

using
n.rxmal light waves and powerful lensei, can only rnignify
up to fifteen hundred times; an electron microscope
can
,,Iqffitfy some ten thousand times. siices of the brain are
coated with a special,substancethat blocks electrons,
and
this substance is then'atsorbed to a different extent
by
different parts of the neuron. In electron microscopy,
a beam
of electrons is passed through brain tissue onto photo_
graphic film. The more a part of the cell is electron
dense,
the blacker it appears on film. ln electron micrographs,
neurons lose their fragile, almost flower-rike appearance
and
instead take on the abstractedbeautyofamonochrome
form of
modern art. Now sturdy brack ri nss and sircles form unambiguous patterns not obvious to the untutored eye as parts
of
rD.o's, dendrites, or soma. But at last, to neuroanatomists
at
any rate, different parts of the neuron, including its internal
machinery, can finally be distinguished. (SeeFigure 6.)
once scientists were able to peer at the brain with
this
degreeof accuracy,neurons yielded one oftheir secrets.
The
final answer was that Cajal was right:.Fre is :i:ndeed
a gap _

F I GU R E 6 o p p o s i te
An electron micmgraph of a slaapse (main picturel
To the left-haudside, a
nerve is 666ing iuto closecontactwith the second
cell to the right m tne cett
on the Ieft are small packetsof a hansmitter chemical,
seenas small circles,
that will empty their contelts iato the narow gap,
the synapse.The areas
where the transmitter wil diffrrse are seenas aait
tutre"i"gr. a s"mm" or
those two neurons is shown in iuset. (Adapted
from L B. Levitan and L. K.
Kazmarek, The Neurcn,Oxford University press,1991.)

*l;
t

95

95

T I { E HUM AN BRAI N

the brEn, neuronsmake

-asvnapse- betweenngqfo,,ns..In
sSrnapticcontact,with .eachotrer by the-apprisitionof all
manner of differentparts of the cell: dendritescan form a
f, ,_-synapsew_ithotherdendrites; aXorislttth iither axons.
; and
.rxons can make contact directly withthe cell body of the
r-rf,
\'\
target cell. The most common form of synapseoccurs when
;
the outgoing part of the cell, the axon, reachesout so that its

I
t

Ii

end point, the axon terminal, makes a slmapse with the

p.fi stubby branching section of the target cell, the dendrite.

*'Fv

l**

The concept of the synapse immediately presents a

n^{:gnroblem. }ust imagine that a signal - an electrical impulse s\P travelling at some 220 miles per hour, arrives at the end of

t{s
NrF

*" a)con,and hence at the synapse.True, the end of the

)CU
T\
axon (the a:ron terminal) is now excited; the potential is
briefly more posiuve. But where is this wave of excitation
goingto go?How canit beusedasa signalto anotherneuron

-(r"\
*

Iiin$

when it is stoppedshortby a gap,the synapse?

It is a little

{r

-like. &iving in a car and Cornin!"to-i river;

an ideal, albeit
exhavagant, strategy would be to abandou the car and find a

Nu$
{:i

more appropriate menns of bavel: a boat. We need a way of

o
-Fq
*-N
'(t -

s 3$
tlt

would dip their arrowheads into a substancecalled curale.

When the arrow embeds into the body of the prey' it does
not kill immediately, but paralyses instead' Bernard suggested that the deadly toxin works by interfering in some
way with nerves in the bodY.
It was not until the beginning of the twentieth century that
Bernard was proved right. Curare was discoveredto workby
blocking a naturally occurring chemical that is releasedfrom
nerves on to muscles.When you breathe' your diaphragm is
operated by a nerve, causi:lg it to move up and down'
Obviously, if the signals from this nerve are blocked' your
diaphragm will not work and you will be unable to breathe.
Hence the lethal effects of the toxin: eventual death tbrough
suffocation. A clear demonsdJtion of how this naturally
occurring chemical provided the vital link in nerye commu'
nication was performed in 1929 by an Austian, otto Loewi.
The story runs that Loewi acted on an inspiration from a
dream he had several nights in a row. His first step was to
repeatwhat was already known: if a nerve that enervatesthe
heart (the vagus)is stimulated, then the heart slows down. It

synapse.

is. irnportans to realize &at the hear't and- aceornpanying

nerve on which Loewi was experimenting was no longer in

Since the nineteenth century, chemicals have pricked the

imagination as being involved, in some way, in neuronal

the body. Rather, the organ was kept alive but isolated in a
special oxygenatedchamber,bathed in a fluid similar to that

communication. A Frenchman, Claude Bernard, was fasci_

nated by the effects of a poison that is used in South

normally found in the bodY.

'.The master stroke of this experiment was that Loewi took

America by Indians when they hunt their prey. Hunters

the fluid that bathed the original heart and hansferred it to a

translatinge,:""Ig:H_"d$pg!_r_nf-o_.98e_tbat-ca,,crossrle.
\

PU L SE, II'IPU L SE 9 7

T
I
I

9E

THE HUl. I AN BRAI N

PU L SE. IT.tPU L SE 9 '

second, unstimulated heart. He found

that even though this

second heart had not been stimulalgd,it
tpo slowed down.
_
'-_
-rr*f
r ne only explanation,it seemed,to
accountfor this discov_
ery was that there must be somechemical-cejeose
d.intsthe _

::::::::" '
rtuid asa resultoi,fr" ,J*,rt"uo' of th; n*t i.*t. Hence,
when the fluid was applied to the
secondheart, the effects
were the same as on the first heart,
because the same
chemical was present. we now know
that this chemical is
the one that is blocked by curare:
@
acetvlcholine. Acetylcholine is the
prototype of many
diverse chemical substances that
can be released from
diverse nerves and neurons in the brain
as the all-important
link in the signaling process. They
are referred to bv the
generic term
JFrnsnjffers.
This discovery of the action of acetylcholine
on the heart
was to have profound implications
for understanding how
brain cells comm,nicate at the synapse.
Once we realize

stipulaUon_releasesa naturally occurihg

]. gt- g.fe_gtrical

chemical from a nerye, we can see

more readily what might
happen at a brain synapse, when
the end of an axon is
effectively stimurated by the electricar
signal. As soon as the
axon' it createsthe right cond.itions
by which acetylcholine
is releasedinto the synapse.
At the end of the axon of the signalling
cell, within the
terminal, acetylcholine is stored
leq in
m many small
small
packets.

the axoq and invades this end region, this transieut change
in voltage acts as a-tligger for some of the packetsto emBtv
out their contents - the dll-important acetylcholine - into
the synapse.The more electricaisignalsthat arrive,the more
packets will empty out, and the more acetylcholine will be
released.In this way the original electricalsignal is converted
faithfully into a chemical one: $e higher the frequency,
Once released, acetylcholine diffuses easilv through the
watery, salty iiquid outside all neurons (the extracellular
fluid), crossingthe slmapseas readily as a boat might cross a
river. The time scales are vastly different, however: since
such chemicals are relatively small molecules, the synaptic
gap is crossedwithin thousandthsof a second.But how does
a chemical, a mere molecule such as acetylcioline, actually
transmit a message?
Let us return to the car and boat analogy:once the river is
negotiated and we are to continue our journey on dry land,
the boat will ideally be relinquished in favour of a car once
more. The original electrical signal, which was converted
into-a chemical p:re, now needs to be-convertedback.to-anelecbical impulse. We need to find out how acetylcholine,
or any bansmitter, might cause a transient change in the
elecbical stafus - the voltage - in the target neuron.
Once it reaches the other side of the synapse, each
molecule of the bansmitter has to make some kind of
contact with the target neuron. The transmitter, like a boat,

PU L SE. IM PU L SE IOI

will need to dock. on the outside of the target neuronlhere

.are special large molegu_Igp,
prgtetrs called rebeptors.that

di5 tailiii-iiiaaei; ;;#;id;.rr"*i."r

";i;r;ru"g"h

asprecisetyasa key

Within,our brains, there are some hundred billion neurons. To get an idea of iust how big q hun&ed billion is, the

is rnade for a lock or as a hand fits a glgve,.

- * A rdibptor-will not jusi let any
old chemical fit into it; it
has to be a specific match; where the molecular configura-

Amazon rain forest offers an appropriate analogy. The

Amazon rain foreststretchesfor 2,700,000squaremiles and

tion correspondsperfectly. gnse.thq transmitter i.s,rocked

i+to the receptor qnd bound to it. the qreation of a new

it contains about a hundred billion trees.There are about as

many trees as neurons in the brain. But the metaphor need
'

trigger for the next series of events to unfold.

not stop there: if we now consider the huge number of

connections between neurons, then we could say that there

Ieceptg,Tprotein on the target cell acts as a mqleculat

are about as many as leaves on the trees in the Amazon

jungle. It is virtually impossible to imesne on a global scale

,or
alternativelv one of the other ioas. The entry or exit of any of
these charged atoms
,in potential difference in the target ce[. In tum, this chenge
in potential difference becomes just one of the many

glg-ctLgafgign-a_ls
.gogductqddowa the.dendrites.toward.the
cell body.
bn a sense,we have comefull cycle.
@

the fervour of chemical and electrical activity, even if only

10 per cent of our hundred billion neurons were signalling
at any one moment.
hr any case,it is not immediately obvious why the brain
should work in this way. After all, it requires a lot of energy
"-' ' toassernble'hansmitter substances;"whichrequires a com'
plex series of chemical reactions. Furthermore, the chain of
electrical-chemical-electrical signalling only works unarnbiguously if the transmitter is cleared rapidly from the
;

. o_ssqjlgaiLif
the net

marked.

sodium channels will open near the cell body, instigatine an

ac9on potential in this new tareet cell. Hence the new target

removal again requires energy either becausethe transmitter

is reabsorbedinto the cell that consumesenergy, or because
the trnnsmitter will be broken down outside of the neuron
by energy-consumingenzJrrnes.
Another problem with this system of chemical signalling

t
I

I02 THE HUM AN BRAI N

PU L SE, IM PU L SE IO3

is time. We have seen that

small molecules will diffuse
rapidly across the synapse,
but the wh-ole .process ofs-ynaptictransmissiontakes a
few milliseconds.If neurons

as it happensl
the neuchemical svnapsp.Ironically,
Goigi was, at reast in these
cases' correct after aI. In such
scenariosno transmitter is
used, but the electrical signal
is conducted easily and
rapidly acrossthese low-resistance
contacts(gap junctions).
Not only is electrical hansmission
of this sort much more
rapid, but it does not entail the
use of energy_consuming
chemicals. And yet the majority
of synapsesin the brain are
chemical.

limited by the passive conduction properties of each neuronal"junction; ggnipared to chemibal transmission, electrici]
transmission, though fast arid economical, wiil be far less
fari,able and diveiCe. Ch.qlniial uansmiision, on the other,
hand, endows the brain with enormousversatilitv: different
q.hemicalshave different actions to different extents at
different times.
Sometimesa transmitter can be even more subtle in the
role it plays in neuronal communication. It can bias how the
target cell eventually responds to an incoming message,
even though it does not pass on a message.This biasing of
geulonal sisnalling is known as leuromodulation. The
concept of neuromodulation, which is relatively recent
compared to the now familiar events of slmaptic transmission, adds still more power to chemical signalling. Unlike

gf time and enersv.

,
Consider once more how many
inputs can forrrqmapses

ilh a targetceu,asmany ;;;;;

", "
case, as we have seen,

iil,rr*a. rn each

different amounts of transmitter

will

be rereased according to the

number of action potentials
,sLj;.e:i::..r-r5-,. -invadingthe end of each of the-inputj. TIie acti";ti*
of th;
neuron is not fixed but can be
varied in rp to a hundred
thousand individual cases.
Moreover, by having many
different chemicals, each working
on its own custom-made
target, different transmitters will
have different effects on
the finar voltage. By contrast,
elechical bansmission w'l be

classic slmaptic transmission, where we have looked at a

single event at some disembodied moment, _thg_jdg__cl
biasine the iesponse of a target cell embracesthe additio''al
.dimension of a particular time frame: first one thing
happens, the biasing, then another. the actual signal that
Mn*ea.Ngrrromodulationcompl,e._'.-:-.*-'^.''::].'.'.''''
ments classic synaptic bansmission as a video film might a
snapshot.This introduction of a bias for a particular period
of time could hardly have been accomplishedby the simple
passive diffusion of electric current from one cell to the
ne)ft.
It is this chemrcal-specificfeature

PU L SE. IH PU L SE I0 5

direc.t electrical counterpart or any easy analogy with a

comPuter.
Second, the chemical composition of the neurons themselves is changing, and hence there is no separate and

at a network of neurons using a high-powered electron

unchanging hardware, incontrast to-a programmable range

of software. Moreover, the ability for incessant change
within the brain leads to a third distinction from systemsin

celli-like tangles than an integrated circuit.board. And yet,

the brain has a precision of connectivity of which the circuit

silicon: of course, computers can 'learn,' but few are

shnnging alt &e time to give novel rcsponsesto the sane

board is only a pale echo, as well as the diversity of a

chemical factory. Dffirent chemicals will be released from

commands.

different inputs converging sn x single cell and active at any

True, advanced robotic devices can seemingly organize

and reorganizetheir own circuits to adapt to certain inputs,

moment. In addition to the degreeof activity of these inputs,

different amounts of transmitter will be released. Finally,

but they are still following a set of rules - algorithms - that

have been progremmed. The brain does not necessarily
operate according to algorithms: What would be the rule for

iach transmitter will dock into its own reieptor that has its
own characteristic way of influencing the voltage of the
target cell. Thus, at every stage there is room for an
enormous flexibility

courmon sense,for example?The physicist Niels Bohr once

and versati[ty in the brain, using

different combinations of transmitter gbgmi"sals.

-*firi;

;"r""J*

il*t*t

"*

beine logical.' ln f"ct, no external intelligence programsthe

brain at all: it is proactive, operating spontaneouslywhen it

o"r*, be regarded
as

comparable to the scenario inside a computer. First, and

most obviously, the brair.r is fundamentally a chemical
system-:. +rre*-the--electtricity it- generdt.C'cijmiis- hdn

--

it.'Com

chemicals. More signiffcantly, beyond the fluxes of ions into

A computer that does

and out of the neuron, a wealth of chemical reactions are

defies its prime function; a Person who does

may well be experiencinga revelation.

occurring incessantly in a bustling but closed world inside

the cell. These events, some of which determine how the
cell will respond to signals in the future, do not have a

I
I
T

decides to take its body for a walk iust becauseit 'feels like

Another insight that is generated by looking at the

ical communication between neurons is to be ab!g-[g

ckal/enyerl
g

,rtcs
"/r:
tgortsJAh8l'

t
^^ifho.,

I
I

I 06 THE HUM AN BRAI N

**"'--**buttom-up

approabhes.-toeJ<traporate
from an event
tt"
"t
single synapse,to a function of the brain. The
brain isTfiilt
::::':r'r--'
ii:p from single rieurons in inireasingiy'compiex
circuits.
These connectionsare not like a row of peopiejust
holding
hands in a line or like the children's gamewhere
a message

PU L SE. IM PU L SE IO7

the output of a single trumpet. One Wav of looking at the

from. the top-down and bottom-up
PgiSjimult?neouslv
SPectiYes-ir Ig.consider the action of drugg.We can sge
how drugs influence behaviour while thev also change
chemical communicationat the single synapse.After all, a
seeminglyindividual and unchangingmind is completeiy at

is passedfrom one end of a row to the other

and ends up
completely distorted. bnstead,remember that
between ten

the mercy of our physical brain, our neurons.

thousand and one hundred thousand neurons

make contact
with any particular neuron. In turn, any particurar
neuron

feel, perhapsthe most courmonis nicotine.Nicotine reaches

will become one of many thousands of inputs

for the next
cell in the network. If we took a piece of brain
the size of a
match head alone, there could be up to a billion
connss6sn,
on that surface.
Consider just the outer layer of the brain, the cortex.

If you
counted the connections between neurons in this
outer laver
at a rate of one connection a second,it would take
thirtv-two
seven million years ago, so you would have to
count over
four times longer than it has taken human beings
to evolve.

Of the many drugs that people take to change how they

the brain within ten secondsof the first puff on a cigarette
and it is possibleto seeimmediate g[nngesin the smoker's
EEG,which becomesdesynchronized(seeChapterz), indicating a less relaxed mode.
Nicotine actually works at one type of receptor - one of
the docking areas - that is normally reserved for the
transmitter acetylcholine. This action servesas an example
of gqe way in which a drug can work to mimic-theeffects of.
a natural hansmitter. However, this mimicry is more of a'i
caricature than an emulation of the nonnal actions of
acetylcholine, for two reasons.First, the nmount of stimulation of the receptoris far greaterthan wo'uld normally be tlie-

in the cortex alone, it would exceed the aumber

of positively charged particles in the whole universe!
Holistic brain function does not have a simple one-to-one
correspondence with any one synapse or class
of transmit_
ters. A crude and highly simplistic analogy would
be to say
that a symphony does not have a d.irectcorrespondence
with

case for acetylcholine itself. Within the brain the repeated

and exaggeratedstimulation of the receptorfor acetylcholine
will have long-terrr effects on brain functioni'g: the receptors will become less and less sensitive as they are stimulated far more than they would normally be by acetylcholine. Because the target neurons become used to these

PULSE,I M PUTSEI O '

artificially high levels of chemicar, the ne'rons gradually

last two actions it used to be sold as an effective ingredient

: " "
'
inanti-;ougL"oa
"nti'dianhse.;"di"r$";;A pernicious action of morphine and heroin is to slow

aur.o.rmaramounrc or srimula[gg.prgvide-dly-thq drug.

This

""="'-liiirro'.[o*il"i'u"ri,;i
;aai"iioo.'
second, *u"r".,

on leveral different
"*ayl"h"nnr ".ts
types of receptor, to have more balancedactions, nicotine
works on only one receptor file, again resul.,.g in rather
one-sided effects. These unbalanced effects will also occur
in a more global way outside of the brain: nicotine puts the
body into a war mode, ready for fighting or flying. Heart rate
and blood pressrue increase. perhaps the feedback to the
brain that the smoker is in a flight or fight situation is in
itself exciting or pleasurable.Most usually, however, smokers will light up simply becausetheir brain is signalling that
its respective receptors need more stimulation.

within the basic part of the brain, the respiratory centre in

the brain stem, just above th" spio"l cord, which controls
breathing. Sometimes this action can be so severe that
breathing stops altogether and the person dies. In fact,
inhibition of breathing is the most common acute causeof
death in heroin abuse.
Despiteits obvious dangers,the clinically most valuable
action of morphine is relief from pain. It is the most effective
analgesicknown, but becauseof its addictive properties, it is
the treatment of choice only to those in very severe or
chronic pain, or the terminally ill. A fascinating insight into
the nature of pain comes from patients who take morphine:

Another drug that works as a caricafirre of a natural

they often say they can feel the pain but it no longer bothers

!13rymitt91 alleit a*different -ona fton. acetylcholine,is

them. Could this action explain why healthy iieople in ho

pain still take heroin for 'pleasure'? Perhaps, in a similar

momhine. Morphine is made from a special type of poppy:

it is named after the god for sleep, Morphos, because it
makes people feel drowsy and relaxed..gglglg is a deriva_
;*.3:;;'i..,:::-:-;:;-;-:titrr8-of.morphiae;ehemficaltfn'diffb&SO that it is Easier tb

way, the heroin abuser is no longer bothered by the

everydayworries and anxieties of life. This reasonfor taking
heroin soon gives way to another,which is simply to satisfy

pass into the brain - hence heroin addicts prefer this

substancebecauseit enters the brain more readily and thus

a craving.

gives them a quicker effect, a 'rush.' The side-effects of

heroin include constriction of the pupirs, constipation, and

mimic a certain classof naturallv occurring transmitters that

suppression of the coughing reflex. In fact, becauseof these

Let us revisit the synapse.

Morphine is of a sufficiently similar
@
molecular gsnffguration to fit readily into the receptor

T HE HUM AN

BRAIN

PU L SE, II'l PU L SE III

protein that is custom-madefor theseE@lartlansmitters

(named enkephalin,glrdorphin,and dJrnorphin;
seeChapter
The
drug-can
thus fool the targetneuron that it is being
-'4.
activated by its natural chemicai messenger,It was a great
discovery in the 1970sthat we have thesenatural agents
in
our body that work, just as we saw for

ffg[yjfulige.

to

signal betweencertain neurons.Moreover, at least some

of
this normal signalling appearsin normal situations
to be
important for relieving pain; for example,
.if_*re action of
perception of pain is worse.Similarly, naloxone

preventsat
least some of the analgesic effects of acupuncture (see
Chapter 2). Mediation of acupuncture analgesiaby endor_
phins would explain the slow onset of the effect, as well
as
its persistence, even when the needles are withdrawn.

FIGU RE7
The fitting of a chemicaltransmitterinto its owa specializedreceptor.On the
left-hand side a naturally occu:ring zubstancein our braia fits perfectly into a
special receptor.This interaction will trigger electrical signais within the
neruon. Drugscan be made to copy the action of a bansmitter by having a
similar shape.In this casethe drug that hasa similar shapeto the transmitter
enkephalin is morphine. (Adaptedfrom C. Blakemore,The Mind Machine,
BBCBooks,1988.)

enkeFhelin ^n.{ relnted eomlorrnrls (See Figure

7.)

-'what would be the differencebetweenheroin

use and the

all possiblebrain areasat once, at each and every respective

natural action of enkephalin in the brain? we are not

all at
risk of becoming drug addicts just because we
have a
nlturally occurring morphiae/heroin analogue.in our
bfain.
Again, the difference between opioid peptides and
morphine is comparable.tothat with acetyrchorineand nicotine.
Opioid peptides will be releasedin different placesin
the

synapse: it will swamp the normal receptor sites. The

brain at different times, and in small amounts. When

a drug
such as morphine or heroin is taken, however, it will act
on

addiction is an inevitable consequence.

consequenceof such
a_ctiv4tionof the receptor site
_exce!-giyg
is that it becomesaccustomedto these higher amounts of
chemical and thus far less sensitiveto normal amounts.This
process of down regulation continues until more and more
drug is required to have the same initial effects. Again,
Another dangerous and addictive drug is gocaine. It is

PU L SE, l l 'tPU L SETl l 3

T' I AN BRAI N

similar to @
except that crack is
chemically modiffed so tha!.it.can be smoked
to
"r-oppnr"a
snorted. Cocainecomesfrom the cocashrub, which grows in
'- ---'
'

'

the Andes Mountaias

1"000to 3,0O0.me$esabovssea level.
- : - ' :- :7
.- .-

.- .l

chemical, E9 transgritter dopamine. In aildition, amphetamine increasesthe availability of these powerful transmit'
ters even more, by preventing their absorption into neurons'
Dopamine and noradrenalinb are thus active in the synaPse

wearly 9 million kilograms of these leaves are consumed

for far longer.than they should be.

annually by the 2 million inhabitants of the highlands of

Dopamine, noradrenaline, and even acetylcholine are

releasedfrom clumps of neurons in the primitive part of the

Peru, who chew or suck the leaves for the feeling of wellbeing they produce.@-works

on a chemical transmit-

ter in the brain called noradrenaline, Unlike nicotine or

morphine, which directly mimic the actions of the transmitters acetylcholine and endorphin respectively, cocaine
works in a different way. The drug increasesthe availability
of the naturally occurring chemical itself. Cocaineblocks the
process by which noradrenaline is normally removed from
the site of action once its role is ended: the drug blocks the

brain (the brain stem) in a fountain-like diffirse arrangement

onto the more sophisticated regions of the cortex and
immediate subcortical sbuctures (see Chapter 1). These
wide-ranging systemsof chemical transmitters are related to
arousal levels, including sleep and wakefulness.In addition,
they can bias, modulate, neuronal activity all over the brain'
It is not surprising that drugs that modify these systemswill
also modify arousallevels. For example, amphetamineusers

transmitter being absorbedinto the interior of the neuron, so

cannot remain still. They cannot concentrate and are

incessantly distracted by segmingly neutal objects and

that it has unnaturally sustained effeclg.

events in the

availability of noradrenaline in the brain but also elevates

environment. In many regards

"*tt*J
amphetamine users resemble schizophrenics in that they
are constantly at the mercy of the outside world, with no

the levels of this transmitter where it works more globally

is
inner resources of grtnd to .assess,-appropriately'-what'

Cocaine is dangerous because it not only increases the

;**;];rai::i':::;'-fgi{hilL.lha,,bOd5,,41'the

pOiUt 'wtibfe*thcl"varioUS

netves

happening.

control vital organs. The effect of noradrenaline is to

A third drug of abuse,g$Ieg4 (a,e-methylenedioxymeth-

introduce the body into a false stress situation. Heart rate

amphetamine, MDMA), targets yet a fourth bansmitter

that also pushes
.svstem ts+n, Aso nown as serot

and blood pressrue can increase, thus introducing a risk of

stroke. @

has a similar effect in that it

causes excessiverelease of noradrenaline and its precursor

upward and outward from the brain stem. Ecstasy is

frequently referred to as a hallucinogen because it gives

I
I

I t.

TH E H U M AN BRAI N

PU TSE. II,tPU L SE ! I5

feelings of disembodiment as well asan overriding feering of

elatedness.This drug causesan excessivJrefease"i***t"-

neurons are associatedwith the regulation of a mnge of very

I

'-*'*-'bnsicfunctions; including sleep.'

nin. The flooding of the brain with serotonin results in

dg-amatic effects on metabolisrnr .on:how temperature is

.:'--'thetarget for much anti:depressantmedication.Many anti-

regulated. In addition to feelings of euphoria, the drug

causes hyperactivity; it is this incessant and repetitive

depressantsact by enhancing the availability of serotonin,

albeit by a different mechanism and in a way that does not

movement that often characterizesthe dancers at raves,

where ecstasy is frequently available. A similar effect can

result in the neurons being killed. The most popular antidepressantto date,Prozac,works in this way' However,if a

actually be seen in rats. A normal rat placed in a box will

steadily explore the territory, displaying a variety of differ-

net increased availability of serotonin is associatedwith

'happiness,'it follows that taking a drug such as ecstasythat

eD.t movements, such as rearing, sniffing, walking, and

washing. However, when a rat has been given ecstasy, it
makes the same movement over and over again, out of the
context of its normal behavioural repertoirc. This repeated
movement is chillingly reminiscent of the type of repeated
dancing movements that drug-influenced humans demonstrate.
It is not yet known whether the effectsof ecstasyare due
to the fact that there has been an explosive release of
serotonin, or whether it is the subsequentdepletion of the
:l:l:-:iii:::'.&er*smitter-thatshas'caused-the-probleni: In dri' event, orie

These fountain-like serotonin-reieasingneurons are also

enhancesreleaseof serotonin in the short term wiii have a

similar effect. If, unlike anti-depressants,repeated use of
ecstasy leads to death of the nerve endings, and thus
pennanent depletion of serotonin, then sustained use of
ecstasywould be predicted to have a depressantside-effect.
In fact, there is some data suggestingthat long-term-use of
ecstasycan be followed by depression and suicide.
We can ie-ethat drugs affect the brain in many ways due to
the variety of brain chemicals upon which they act and the
different .stages in synaptic transmission at which they

tslg"ryq+9.Niqglpp__qg{gtglp,hingmimiss-pa{ictlar-natu-rally occurring compound by acting at its receptor, whereas

serious consideration is that there is now evidence in rats

that repeated use of ecstasyover the long term ca'ses death

cocaine increases the availability of another type of sub-

of the clump of neurons (the raphe nuclei) that send a

fountain-like spray of axons from the brain stem up and out

different in that it actually depletes the brain of yet a third

type of transmitter chemical. Because we have so many

in a diffuse array to the higher regions of the brain. These

transmitter chemicals in our brain, there are highiy specific

stance, before it diffuses across the synapse. Ecstasy is

II6

PU L SE. IM PU L SE I 1 7

THE HUFTAN BRAI N

targets as well as these higNy diverse ways in which drugs

-em-act.
We know to a certainententnAatdrugseardo;'but
we do not really know the full effects in the long term or the
side-effects on"the rest:of-the'bodv; -,::.- -'
Perhaps the most tantalizine issue is the link between the
known molecular/cellular changeand the changein the wa]r
we actually feel. Why should over-stimulation of enkephaiin
ve rise to the subiective
ggnsation of euphoria and a disregardfor pain? How might
.anti-depressantswork at the level of the synapsesuch that
enhanced availabilitv of serotonin leads to an alleviation of
bear in mind that anti-depressantsare biochemically effectake some ten days to
$ive within hours but therapeutically

work. There is obviously no simple one'to-one relation

between g single class of molecule and a particular mood.
,- lr.ftig way, consideration of the acti,onof'drugs highlights
the still elusive nature of the link behdreenspecific and welldefined events at the synapseand how thole synqptic events
actually constitute an emotion. 9ne of the greatest chal-

gbemical modulatioq. It is truly tantalizing to inagine how

our unique consciousnessis really at the mercy of a cocktail
of brain soup and spark. The issues of how a unique,

individual brain is assembled,andhow it is expressed,are

-'
*
tackled'reSiibctiVelyin the-next-two cliapfei;

C H APT ER 4
C EL L S U PON C EL L S

Someoneonce askedthe physicist Michael Faraday,'What

useis electricitv?'He replied, 'What use is a newbornbabe?'
It is easy to take Faraday's point: human babies seem
parthulariy helpless. It takes us some sixteen years to
realize our potential as adults, whereas a rat bom within
twenty-six days of conception takes oniy two months or so
to become fully mature. An eiephant spends over twice as
long in the womb as we do, some twenty'.to twenty-two
months, but is then mafure wi'hin eleven yea.rs.So why do
we serve, by comparison with other animals, such a long
apprenticeship for life? In 1883,the philosopher and historian John Fiske, who popularized evolutionary theory in the
United States,asked,'What is the meaning of infancy? What
t..ol:r

is the meaning of the fact that man is bom into the world
more helpless than any other creature,and needsfor a much
Ionger seasonthan any other living thing, the tender care
and wise counsel of his elders?' We shall erplore this
question as we look at the development of the brain and
identify the factors that make an individual.
Life starts with the fertilization of the mottrer's egg,when
a single sperm from the father burrows into it. This act

I20 THE HUM AN BRAI N

C EL L S U PON C EL L S 1 2 I

induces chemical changes that preclude the entry of any

embryonic disc, an oval-sheetthat is two-cell layers thick; It

is incredible to contemplate that this thin little d.iscis the

amassing around the egg. But it is a long way from an egg

ultimate source of all the different cells that will make up

with a-diameter of aboul 0.005 inche,s-te.a,brairrThe-first

the human body, but already these early pioneers have

step toward building a brain, and indeed the rest of a body,

started to diversify, albeit rather crudely.

is to form a single cell from the eggand the sperm: a zygote.

After just over a day, some thtty hours, this zygote divides
into two cells and repeatsthe processagain and againso that
within

three days it has formed into a ball of cells

resembling a mulberry, hence its name

#9*I".(Latin

for

'mulberr5t').
Five days following fertilization, the ceils within the
morula split into two $oups. One group forms an outer
i ,

congregate into a tight inner mass within the sphere at one

':. ,

outer wall will provide nourishment for the developing

t.,,.,.
-- - -", . embryo, ,which-will- be ereate& from the inner-celf mass.
However, it is still only six days after conception. The next

At around twelve days,certain cells in the upper layer of

the disc start to move toward the middle, as though in some
rehearsed and choreographeddance. tn the middle of the
embryonic disc these mobile cells then insert themselves
between the original upper and lower layers, threading
along the disc and thus creating a third layer of cells. The
embryonic disc is now tlrree layers thick. It is at this stage
that we can first focus on the future brain. The middle layer
of cells appearsto send chemical signals to the upper layer
of cells so that they will diversify again,to becomeneurons.
Embryologistsrefer to this upper layer of precursor neurons
as the neural plate.
By about eighteento twenty days the neural plate startsto

vital stage is for the blastocyst to become implanted in the

change in the middle, in that the centre sinks inward and

llning of the womb, whereby the new life will have accessto

.&1dS9_r_*ov_9u^pyig$an.5l*o-.,1j,-A&qrthreerueelc,lhese
edges will start to rise up, creating a neural groove. The

*"i'rii;:!*';",*'dll-tte'dubibdts-ifi^rill iffia'ffitd;il'>ctthil;nine

weeksor

so.
Within about a day of implantation, the inner ball of cells

edgesof the groove then fold inward and fuse so that they
form a kind of cylinder, a neural tube. By the end of the ffrst

that had congregated within the blastocyst has separated

month in the womb, a primitive brain has already been

from the outer wall. where it fuses with the womb. This
jumbled mass of cells has started to flatten into the

formed. In fact, well before the neural tube took shape,the

]ottng brain was starting to manifest itself. Even at the stage

,ii

I2 2 THE HUM AN BRAI N

il
It
ll

C EL L S U PON C EL L S I2 3

of the neural plate, certain segmentswere already d.estined

interconnecting labyrinth that finally opens onto the spinal

to form specific brain regions.

cord, and tlrough which'pores allow for the-circulation of

By the fifth week in the womb it is possibleto identify two

bulges at the front that are the foundation of our hugely

the colourlessfluid that is goirig to bathe the entire brain and

deveioped cerebraihemispheres,as well as certain regions

beiow the cortex such as the basalganglia,which we saw in

philosopher Galen (see Chapter L) had thought many

chapter 2 were important for movement.Encasingall this

turbulent, burgeoning activity is the skull. The developing
skull has membranousregions to enabieit to expand and
thus allow for this frantic growth; it is only much later in
life, when the brain has reached fuil size, that the bone can
finally become fused together.
The neurons-to-bewill divide severaltimes each, so that
there is a massive proliferation in cell number: at maximum
rates, cells will be dividing to give 250,000new neurons per
minute! The primitive brain continues its development as

spinal cord for life. It is this cerebrospinal fluid that the

centuriesagowas the 'psychic pneuma,'the seatof the soul,
and which is sampled nowadaysin the routine diagnostic
procedureof lumbar puncture.
ht S. nineteentl century it was a popular idea that the
developmentof the human brain reflecteddevelopmentin
evolution: according to this idea, our brain in the womb
would first resemblethat of a reptile, then that of a fish, then
a bird, and finaily lower mammals such as a rat, via cats and
the like, to higher slammx|s. Toward the end of gestation,
the brain would be similar to that of the foremost group of

the top of the neural tube thickens into three swellings. By

the qlart of the second month, tlere are recognizablebrain

animals, literally the primates, from which destiny led

regions in place. The front stump of the neural tube initially

bends in two places set almost at right angles with the

twentieth century, such thinking continued: in one of his

developing spinal cord, as parts of the brain grow faster than

bishop's robes and proffering his bishop's ring.

-------'-others: The'very front phrt iwiillc into two tremisprreres;at

around eleven weeks, the back part sprouts an outgrowth

ultimately to humans. Even into the first half of the

novels, Aldous Huxley refers to'the 'ex-fish' standing in his
l-

__

__'- ' -:
---'-'

"

-*

_-l

*_" _'_-

However interesting and attractive this idea might be that

ontogeny (individual development)reflects phylogeny (spe-

that becomesthe easily recognizedlittle brain, the cerebellum.

cies development), it is not really valid as a blanket

It is due to the closed neural tube that cavities in the brain

- ventricles - are formed. These venhicles constitute an

that of some 'lower' species.Evolution is more like a bush

generalization.One brain is not simply more developedthan

than a ladder, with speciesdevelopingalong different lines

I2 4 T HE HUM AN BRAI N

C EL L S U PON C EL TS l zt

according to the dictates and needs of particul4r lifestyles.

these convolutions appear only relatively }ate in the devel-

.*-- * -"--*-r{t-no*time"
does-a human fetal brain-rEsembki,for'biample,

opment of the human brain, at about seven months' gesta-

that of a snake, where the regions associatedwith smell

tolfactory tubercle) are particularly well developed. Rather,

tion. The advanlag! of 1 fofded c.ortexis that more surfaco

area can be accommodated in a restricted space. Imagine

each brain has evolved for the individual lifeslvle of a

scrunching up paper into a bin: the more crumpled the

particular species. The human cerebeilum at no time in

paper, the more can be accommodated.

.,.

development occupies either a half or 90 per cent of total

In this case it would seem that the development of a

brain mass, as is the case for the cockerel and certain fish,

wrinkled cortex was a case where glggenf, did reflect

respectively. The cerebellum is a sfucture that is the least

phvlogenv. But perhaps the function of the cortex can be

changed acrossspecies;however, it is the high proportion of

directly related to the general sophistication of the brain,

brain mass devoted to cerebellum that is a departure from

irrespective of species-specificpeculiarities in lifestyle. If

the basic ground plan, the variation on the theme that fits a

the cortex is the most important areafor cognitive processes,

particular species.Presumably,the lifestyles of the fish and

as suggestedin Chapter 1, then clearly the more cortex ons

the cockerel, compared to the human, are dominated much

has, the better for being as flexible and adaptable to one's

more by the need to generatemovementsthat are exquisitely

particular environment as possible.

coordinated with the incoming senses.A disproportionately

On the other hand, dolphins have more convolutions on

large cerebellum is not a 'statutory rite of passage for

theii cortex than we do, flt r"r estimated to be only as

intelligent as dogs.
-Sizeroj cortex, and hence the number of

developing breins of all species.

On the other hand, a stiking feature of the developing

convolutions, is not the only determining feature. Dolphins

human brnin is how the immature cortex at different stases

constrained,
l*"rgpbratg.sJ-unp"Iy-hecause-thasizp-is'not
h3vS-

aoesinaeliaiJr"-ui. trr"i ot otn"i ,p*.irr. inr" *"".

as it is with humans, by the pelvic bones of the mother.

"r

rats, rabbits, and guinea pigs, for .ercample,is smooth in

Although dolphins might have a large surfacearea of cortex,

texture, whereas that of the cat has some convolutions. By

the time we arrive at the primate brain, these convolutions

it is thinner than that of h

"nans and the neurons within are
organized in a less complex pattem. Thus, convolutions of

have increased markedly, while the surface of the maturs

the cortex are clearly a factor in determining the eventual

human cortex resembles a walnut. Interestingly enough,

power of the brain, and increase as we grow in the womb

t
I
Ir
II
t
;

t
I
;

a
I
I

I2 6 TH E HUM AN ERAI N

C EL L S U PON C EL L S 1 2 7

and during evolution, but other factors are important as

-*ell*.For all species,the sequenceof eventsin growing a brain
at.the.level of the basic building blocks, the neurons,is the
same. If the brain is to grow and if the brain is made of
neurons' then those neurons must continue to increasein
number. The cells destinedto be neuronsmeet the demands
of the developing brain by splitting themselvesin two. in
order to divide, a neuron precursorwill embark on a short

different directions according to their different destinies.

"

Foi bxample,some C"tG that move just beiow this middie

region are going to become a speciai type of neuron interneuron - that connects other neurons in small. iocal

a-

circuits. Alternatively, some of the cells that move to this

region will becomeglial ceils.
9lial. cglls are not neurons at all but they abound in the
brain, where they actually outnumber neurons ten to one.
The term g/rloderivesfrom the Greekfor'giue.' When these

journey that can be recycled severaltimes. By putting out a

tentacle-like extension, the central mass of a neuron slithers

to neurons. There are various types of glial cells with

I
I
I

from the outer region of the neural tube toward the centre.
Once at the centre, the nucleus will divide so that the two

different functions. One glial type lqraqpLrygj

plays a
part in removing the debris of dead cells within the brain

new cells then push back to the outer edgeof the neural tube

following damage; another class of glial cells generatesa

to start the cycle again.

fatty sheath around many neurons to act as electrical

It is important to remember that the brain is not just a

homogeneousmass,but, as we saw in Chapters1 and 2, it is

insulation.

composed of highly specializedregions that can be distinguished by shape and by the operations they perform in

pstrocfie, and that is the most ubiquitous doesnot appearto

F
F
F
F
T

overall brain function. It is not only vital for a growing brain

to have more cells bu! to have them in the qight regions.A

cells were flrst observedthey had the appearanceof stickiag

The type of glial cell that is ne."ed after its starlike shape,
have any single role. Originally, astrocyteswere thought to
have a rather passive function, preventing the neuron from
,.. slithering around by-providing a type of biological uetting;

nouron, then, once it has undergone several cycles of

dlvlslon, must migrate to its correct location in the new

known more formally as the exhacellular mabix. It is now

braln.

and more dynamic roles. In the healthy adult, these non-

clear, however, that asbocytesplay a wide range of diverse

Initially, neuronswill simply migratefrom the outerto the

neuronal cells protect neurons by ensuring that the micro-

lnner region of the neural tube, but as this zone becomes

thtck with cells and well established,cells will move in

environment around the neuron remains benign in its

chemical composition.They act as a type of spongeor buffer

I28 TH E HUM AN BRAI N

C EL L S U PON C EL L S II'

for mopping up potentially excessiveor toxic amounts of

In theeve"t,J- *i"ur";r"r;"J;;;;"*o=
-cneiniiars.

"

cytes will redouble their efforts and increase in size and

"," n-umbet so that they;can relsxss.fuighlevels af:substancettii-

:-

further neuronsend.up misalignedand the whole cerebel- Iiiin*idinaihs-abnondiallysiiiatl. As wb iiw in Chapterz,

sincethe cerebeiiumis importantin coordinatingmovement
arid seriiei,'it is haidly surprisingthat animalswith sucha

enable subseq-uentne_urongroyq an{ rgpair,of_the injury.

compromised cerebellum extribit disorders of movement. At

How might glial ceils be important in migration of

the moment it is a mystery how eachneuron knows when to

neurons to the far cornersof the developingbrain? Although

get off the monorail, to alight in its particular piace in the

neuronal migration is a long way from being well under-

future brain.

stood, it is known that one particularly important job of glial

As more and more neurons proliferate, migrating along

cells in the development of the brain is to act as a kind of

the giial monorail and then alighting, graduaily the brain is

temporary scaffolding. Glial cells set off fron their point of

growing, accumulating cells layer upon iayer a little like an

origin ahead of neurons, as though layrng a track. In their

onion. Eventually, the outermost layer, the cortex, starts to

wake, the neurons then slither along the glial cells as though

assemblefrom an initial thin layer of cells, the cortical plate,

on a kind of monorail. If the glia are abseat, then certein

As more cells arrive, th"y have to journey through this ffrst

neluons will not be able to migrate, with dire consequences.

layer of new cortex to form a secondlayer, and so on. In the

One of the best-known illustrations of the type of prob-

mature cortex, there are six layers in all. The neurons that

lems that ensue when neurons in the brain,are unable to

were the first to arrive during development constitute what

-- migrate along their glial monoiail ciccurs in i certain abain

ends up as the deepestlayer of cortex, that furthest from the

of mutant mouse, the weaver, so named because of the

outer brain surface; the neurons in the outermost layer of

severe disorder of movement displayed- Instead of walking

cortex, forming the very surfaceof the brain, were the Iast to

in straight lines, these mice will suddenly tu-rn in random

-:----^,.,.

-. _

_*

;i ::l

_ _ - :'

-__{

directions, and are generally weak and subiect to incessant

What makes a cell become a cortical neuron? Again, as

trembling. The problem for the weaver mouse lies in the

.littte brain at the back of its head, the cerebellum. Due to a

with the actual point of descent from the glial monorail,

mutation in the genes,tne ltiat cells in this region do not

do know that in the brain there are molecules known as cell

develop as they should, resulting in a class of cerebellar

adhesion molecules that may act a little like sticky badges,

neurons that do not migrate to their rightfui place. In turn,

so that it is easyfor neurons literally to stick together to form

little is known of this means of neuronal specification. We

I30 THE HU} 4AN BRAI N

C EL L S U PON C EL L S

a group. For example,there are certain experiments

where
individual cells from different parts of,the developing
retina
have been removed and mixed togetherwith other
neurons
ftg- other populations of potential brain regions;
the
original cells taken from the retina tend to retain
their
specification in that they can find each other and
reaggre_
gate. The commitment of any neuron to a particular
size,
shape, location, and connectivity will be
occurring at
different times. In addition, it is likeiy that the chemicar
that
a cell will use as its transmitter is determinedonce
and for
all as soon as the neurons have stopped proliferating.
By
nine months' gestation,we have most of the neurons
in our
brains that we are ever likely to have. (SeeFigure
8).

25 days

@
five monrhs

40 days

six monrhs

@
100days

seven months

So oow we are born! Birth aliows the brain to go

on
growing, since otherwise the head would soon
become too
big for the birth canar of the mother. At birth, the
human
head is roughly the same size as a chimpanzee,
some 350
-

eubic"centimetres, By six months, it will be half

its eventuar
size, and by two years it wilr be three quartersthe
size of the
adult head. At four years old the human brain is
four times
the size it was at birth, some 1.,400cubic centimetres.

Evan withiii the'fiiii"month o] tiie, a babv

n",

"tr""ay
some reflexes. A reflex, as we saw in Chapter
2, is an
unchanging responseto a given, fixed,situation.
One such
reflex is a scooping movement of the arm that will
underrie
the eventual sophisticated abiiity to grasp objects.
This
reflex is seen if one tries to pull the baby,supper
arm away

FIGURE8
The developinghuman brain. (simplified from M
Cowan,ScientificAmeri-

,- canlibranr

Series, 1g7g;)^

from his body: the baby,s responseis

to retract his arm.
bringing it back into his trunk. This grasping
reflex becomes
more and more refinedas the months pass.
From curling ali

I3 2 THE HUT. I AN BRAI N

C EL L S U PON C EIL S

movement (the motor cortex, seeChapter 2) right on to the

his fingers around an object placed directly in the palm, the

obfect
.-!$j-ps-co-m{p--ahle"io- turn-.his-hana to gmsp=an-

' ' immediate nerves in the spiniil"coid that ale iesponsible.for

brought into contact with the back of his hand. Finally, he is

- .b!.t !S-perform vohrntary grasgiqg,.where he is-first able to
reach of his own accord for whatever is in view.

Thesenagesof the aru"iop*-""i

contraction of muscles in each digit.

" -_

It is this fine finger movement that is the most refined, the

most advanced.Within the motor cortex it is the digits that

p*"nJ

have some of the largest allocations of neurons. Moroovor,

changes occurring in the baby's cortex. During the first

when the motor cortex is damaged, it is the fine flnger

months of life there is a massiveincreasein the cortex in the

movement that frequently does not reappear, even though

insulati:rg substancemyelin (seeChapter 3). Once the a:<on

most other movements miraculously return.

"f ;;d;

is insulated with myelin, it conducts the electrical signal far

Once able to move each ffnger independently of the

more effrciently. Clearly, a movement as delicate 3s v6lrrn-

others, manual skills increase enormously. This improved

tary reaching can only occur when the neurons in the cortex

manual dexterity meensthat tool making is far easier,whlch

are working as efficiently as possible.Myelination continues

itr turn will help enhance the progressand survival of the

apace right up to fffteen years of age, and even beyond.

species.On the other hand, independent digit movement on

Furthermore, it is a pleasing thought that myelin can even

its own is not the secretto the sophistication and versatility

continue to increase in density to as late in life as sixty years

of the prinate lifestyle. The hamster and racoon also have

of age.

good conhol of their digits.

.".. .The*-ultimate.skill in picking up objects comes,' often

unnoticed, toward the end of the first year of'Iife. Whereas

voluntary patterns of movement. The Babinski sign, named

young babies can only move all digrts simultaneously, the

after the man who first reported it, disappears in infancy. If

older child becomesable to move his fingers independently.

the.,fqg! _Ln?4. adult .ars-stroked- then- the toeg

_o-f,
_,, F.9, !+--I-s-..
tend to curl inward, after a one-seconddelay. However, in

small object is picked up with the thumb and forefinger.The

babiesthis stroking of the balls of the feet results in the toes

6ain group of animals able to make these types of move-

being flexed upward in a fanning movement. As with the

ments is the primates. Unlike many other species, such as

the cat or dog, there is a direct connection in primates from

grasping reflex, sfunnges in the Babinski reflex during

developmentreflect x gfuengingnervous system.The Babin-

the area of cortex directlv associated with control of

ski sigu qhangesas soon as netuons in the motor cortex (see

l tt

fli

I
li

t
t
I

C E LLS

-J::;.**=*.-

Chapter 2) becomeproperly connectedwith the neurons in

s-pinal cord that control muscle contraction. In normal,
"......-t"h9-

U P ON

C E LLS

' that the brain is processing information from the outside

world, enabling the body to move about in its particuiar

mature nervous systems,the motor cortex sends impulses

lif911VfeThe brain receivessignalsnot just from the outside

do.v'm-the spinal cord to cause contraction of the muscre,

seen as a curling of the toes - hence the short delay of a

bombardingthe brain incessantly,although we are for the

second in which the input of the stroking is transmittedup

most part oblivious to what is happening. We do not, for

the spinal cord to the brain, processed,and then sent back

down the spinal cord. Since the curling of the toes in

example,continuously and deliberatelyhave to control our

breathing,heart rate, or blood pressure;such repetiiive and

response to stroking the ball of the foot requires the integrity

tedioustaskswould leave us no time for any other activitv,

of certain parts of the brain and the ascendingand descend-

including sleep.

world but also from the body. These intemal signals are

ing spinal tracts, this test is also used for diagnosisin adults
with suspected damageto the spinal cord or brain. If certnin

Becausecommunicationto and from the brain to our vital

orgnnsseemsto work mostly of its own accord,it is referred

parts of the brain or the descending spinal tracts are

damaged,then the positive Babinski sign will re-emerge,the

to as the autonomic nervous system.The autonomic nervous

system is under direction from the brain, but physically
extends beyond it, in that it comprises sets of neryes

local circuitry in the foot will predominate, and the adult

will revert to the reflex of the baby and fan out his or her

emanatingfrom the spinai cord that are in contact with all

"_99 vital organs. These nerves have developed separately

toes.
Some reflexes disappear altogether; for example, if supported and held with their feet touching a surface, very
young babies will make walking movements.No one really
knows the point of this walking reflex. There ,i{g-s.,^_o_nj9_

I35

from the centrai neryous system,the brain and spinal cord,

at a very early stage in gestation. As the neural plate is
-

closing to fofm the neural tube at about three weeks after

-o{reptioll;

fr small $olrp=of Woul-d:-beneurons on either

idea that the more a child was able to practise the walking
reflex, the more efficient and more quickly it would learn to
walk. It is now known that this is not true.

side is not included. Thesecell groups are referredto as the

neural crest and they will give rise to the neryes that make

up the autonomic nervous system.

It is not just our voluntary functions,such as graspingand

The autonomic nervous system is not just some sort of

steady-statemechanism but actually has two modes in
which it works: a war mode and a peacemode. When a loud

walking, that gradually develop after birth. Another system

trlr involuntary functions has also come into play. We know

*
*
*
!:

.<

I36 TH E HU] . I AN BRAI N

C EL L S U PON C EL L S 'r 3 7

noise surprisesyou, your heart rate increasesautornatically.

Ihe-survival vaiue {or--at irrcreasedheafi iate iS thet your body is preparing for an emergencyaction where you might
,have-to exert yourself by running or fighdng; and theiefore
your biood is pumping much more quickly to give you more
oxygen.

.thetic division of the autonomic nervous system. The

the ability of the skin to conduct-electricity and thus be

aeiecleaas i changing signal
fi" a.t""tiJo *ifrto"r.
-itf,

the defauit condition and operateswhen we do not need to

- place immediatesurvival as the top prioritv. We have time
to reiax and to digest food without being on red alert to
respond to a rapidly changing situation. The heart rate is

activities, which are not necessaryfor immediate survival,

slow and steadyand food is being steadiiv digested;we do

not need sweatto cool down or dilated airwaysto maximize

are suspended.In this condition, you need sweat to cool

breathing.

down, there is no time for digestion, and the blood carrying

f,lft16rrgh it is normally the brain that controis the vital

organs,the stateof the body can also feed back to influence

sympathetic division is activated when the usual daiiy

oxygen is needed in vital organs,so that your heart rate and

blood pressure go up; airways are dilated to that you can
breathe more easily and an organ known as the adrenal
medulla releasesadrenaline to circulate around the body to
help keep the organsresponding appropriately.
.. Nowadays we are not necessarilyhunting or being hunted
in a literal sense; however, when you feel nervous or
excited, your body has all the atavistic reactions needed on
the early cro_Magnon savannah.
IgyW
iJthdrigh
y";ilgoing
to fight or flee,even
,5i3lir'-iiiopArates
if you do not actually do anything. As you pick up the
phone to hear the result of an exam, job interview, or
medical test, you might senseyour heart thud and you might
feel hot and a little sweaty, especially in the palms of your
hands. It is this response of sweaty palms that can change

the state of the brain. For example, a certain drug such as

plqpranolol, (a 'beta-blocker')that slows down the heart rate
does not itself gain accessto the brain. Nonetheless,this
drug can be taken as a calming agent simply because the
..slowed heart rate, rather like deep breaths, registers in the
brain that a non-stress situation prevails - that is,. the
: Parasympatheticneryous system is at work.

.to situationsthat are*rfrbfdStteeifie

include-adaptations
to certain organs,rather than the global conditions of being
on'red alert or being relaxed. One reflex that everyonecan
observeis the response of the iris to light. We know that
when a light is bright, the pupil automatically constricts,
and dilates again when the light is dim, thus ensuring the
correct amount of light is let into the eye. The iris also

t\

I3 8 THE HUHAN BRAI N

responds automatically to emotion. If we are admiring

something or feeling affeetionateand tender, then our pupils

ll
ll

C EL L S U PON C EL L S I3 9

--*,

automatically dilate. Apparently some wily saiespeople

have trained themselvesto observea customer,seyes to
assessthe chancesof a potential sale.Moreover,dilation of
the pupils is considered a featurein enhancing attractiveness.

4. i

wouid consciousnesssuddenly intervene? And of what

iould a fetus be conscious?Another idea might be that the

It seems
the act of birth itself that evokesconsciousness?
hard to accept this line of thought as the brain itself is

asked to sort photographsinto one of two piles accordingto

whether the females in the photographswere attractive or

completelyunaffectedby the birth process.

not. There were many photographs,so the subjectswould

have been unable to rememberindividual faces.The result

scious sometimeafter birth. Not only would this scenario

was that they frequently put duplicate photographs of the

same woman onto different piles. The only difference was

problem is again posedof identifying the critical stagewhen

that in one casethe pupils of the women had been touched.

up to look as though they were dilated. The men were

both the fetus and the neonate is a gradual process.It is

subconsciously registering dilated pupils as the decisive

factor in assessingattractiveness.in the nineteenth cenfury,
women contrived this reflex by taking a drug that blocked a
certain type of receptorto the transmitteracetylcholine(see
"

identifying exactly when this momentous event occurs.

Ctearty,the single fertilized egg is not conscious,so when

babybecomesconsciouspreciselyas it is born. Again, this is

a strangeidea as many babiesare born prematurely.So is it

In a well-known, classic experiment,male subjectswere

IJ

'

Chapter 3);hencethe colloquial nameof the drug(

beiia donna,*""nirrg 'beautiful lady.'
And so we are born equipped to adaptto stressand.with a
certain number of reflexes. lut are we conscious? This
perplexing questionhas never beensatisfactorilyanswered.
All the possibilities seem bizarre. one scenariois that we
,re' consciousin the womb, but then the problem arises of

An alternative idea is that the child may become coni*ply that the neonatewas mereiy an automatonbut the
consciousnessoccuued. The development of the brain in

'

thereforeimpossibie to identify some distinct and conspicuous event that could be correlated with the onset of
consclousness.
There is another possibility. St"ce_th", br"t" d
the casetha
,
Phenomennnhrrtthar it grnwsas hreins do. If we acceptei
I
that consciousnesswere a continuum in this way, then it

would follow that the fetus is conscious.but consciousto a

far lesser degreethan the human adult, or even the human
newborn.This way of looking at consciousnesswould also
help with regard to the riddle of whether non-human

^-tl t--

I
.1
.,-"!.-".

I
I
It

' '.i:

| aaE FIUFIAN

C EL L S U PON C EL L S I'I

ERAIN

animals were conscious. The less sophisticated the brain,

_theenviroment in which we. are plaeed-Judging from the

*o.rla-l
$-9-!e,9sthg deglee of consciousness-ffr*"rr*r.i*"i,
be conscious, but a chimpanzee would be conscious to a

and the parallel deveiopment of behaviour, ii is clearly

lgr,setdegreethan itshuman,counterpath:asthstfiains of the

two species, so similar at birth, then follow different fates. -

-- By nins months after conception; most of the neurons that

that ;i the chimpanzeeare of

"rra
comparable weights at birth. A vital difference is that the

will constitute our brain have proliferated to the appropriate

primate brain, including that of the chimpanzee,undergoes

tively sets down roots and initiates communications with

most of its develop*sn1 wi'hin the womb. For the human,

neighbouring neurons by establishing a synaptic circuitry

much of its development - some might say most - occrus

(seeChapter 3). All the time in the aew brain, the axons are

outside the womb. Might this postaatal growth of the brain

now growing out from the neurons to connect them to other

give us any advantagesover our chimpenzee cousins, who

neurons.Much of the spectacularincreasein brain size after

differ from us by only 1 per cent of their DNA (deoryribo-

birth is actually attributable to the development of these

nucleic acid)?

processesacting as lines of communication between neu-

The biain of the U,r*"r,

Our brain, and its process of development in the womb,

has been the sr-e for thirty thousand years.With the brains

brain region. Once at their destination, each neuron effec-

mns, rather than simply to the addition of more neutons.

[See Figure 9.)

we have as newborn babieswe could have just as easily been

Even when grown in a dish - in tissue culture : brain cells

ushered into the world of early Cro-Magnon man.'stated

will send out their iu<ons.Therefore, using a time-lapse

another way, the brain of an early Cro-Magnon baby could

video, it is possible to observe directly brain cells pssshing

agile and comfortable with

out to their neighbours to establish contact. Viewed in this

com,putersas are many young people in developedcounfies

YlY,*iLiS "Lrard-noltoanfhropenorphize- d:r-devetoptng

have been as intellectually

'today*Tfie $iticil;Liil6!;

--

- .- - - i

for the adaptableand impres-

sionable young humanbrain is to develop and mature under

the particular stimulation and constraints of the environnent in which it is going to have to survive, whether it is
one of jungles or computers. Unlike 6u1 shimpnnzee relatives, our brains have an extraordinary capacity to adapt to

cells. As they move on film, they appear highly purposeful,

yet with the fragility of spun sugar,moving acrossthe screen
with alarming speed while literally feeling their way by
Itreansof fluted, weblike endings that undulate and flutter as
they make their inexorable.progress.Almost a century ago
Ramony Cajal (see Chapter 3) referred to these endings of

t
t

I
ii
b

F,

I4 2 THE HUM AN BRAI N

lr
li

C EL L S U PON C EL L S I4 3

chemical. The concentrationof this chemicalwill be strongest near the target and weaker as it diffuses awav. Bv

l{,:/!trjll

oYt
ilr[i

:::--::flroving in the direction where the concentration of the

iiltFl/

I
I

arrive at the target.

The prototypechemicalidentified as important in control-

r \' /

l,r\{"f
il

Ll

ffi#i
{i

jl' '

:;;ii

"_

inside a cell once contact between neurons has been

established.Once transportedback in this wav, NGF possi-

tr

udf. \

3 months
F IG U R E 9
The deveiopment of the human cortexis characterizedby an increasein the
number of connections.Although individual cells can be readily seenin the
newborn cortex, by two years the ner.rronsare hard to discern in a dense
network of connections.(Fromi. L. Conel,The Post-NatalDevelopmentof the
Human CerebralCortex,Vol. 1, HarvardUniversity Press,1939.)

It is
ling the gro"r'thof nervesis@),
thought that NGF works primarily by being carried back

.r.,:, bly invades the nucieus to interfere with the expressionof

;", genes,in that it switchesoff a geneticallyprogrammedselfi-1::1,.. -

I:{f,,destruct mechanism. Conversely, if antibodies to NGF are

given, then the neurons in which it normally operateswill
t

,',;ifi at". But brain developmentdoes not reiy exclusiveiyon

NGF. While NGF is used in neurons outside the brain and
str certain neurons within the brain, it is only one example
of many such probable guiding chemicals.

the nerves by the highly apposite term boffering rams;

It is hard to imagine how such processeswould work for

however, their scientific name is growth cones. When

i"T.i-: some of the very long distances that axo,-n!ate known.to

:.:;.: bavel. A further possibiliqr, during early developmentwhen

looking at such fiIms; it is hard to undersfand how anvone

could view the brain as a computeror even compareit witb

.::..

"lii'.

one.

toffee, but along which other axons will be able to follow.

How do these vounq neurons know where to eo? It is

thought that their primary orientation is probably genetic,
but that any final routing will be fine-tuned later by local
lirctors.Another idea is that the targetcells releasea guiding

the brain structures are still close, is that there are a few
:.pioneer fibres that then become stretched out like melting

;ti,r,,,, Io *y event, it is certainthat thesegrowing axonsare not

.,,; inefutably fixed in their course. Rather, growing neurons
-.;,

can adapt sensitivelyto changingcircumstancesto makethe

THE HUT. I AN BRAI N

C EL L S U PON C EL L S I.I5

l{dI
,.{il

best of a situation; for example, if the outgrowing-axo+sare

or thg.!.ntendedtarget is partially destroyed: Such
1e-_dqcedl
scenarios have been demonstrated in relatively simple

..i-..--

t
I
1

t
iI

't

J:
I
I

space, but stiil in a top-ographicalmanner: thel' will thus

each have halt"as much target tenitory as normal.
Another way we can appreciate the adaptabilitir of neu-

systemsthat are tractable to expelinental manipulation and -

rons is by looking at a part of the body that is very important

*rhere it is already known that the outgrowing neurons form

to animals: whiskers. Whiskers allow animals to pass

an orderly one-to-one connection with their target.

through narrow openings.As soon as they are touched,the

In the frog's eye, for instance,each axon leaving the eye

has a particular target territory that it will invade within the
relevant target structure (the tectum) in the frog's brain. If a
neuron is coming from the extreme right of the eye, it will
invade territory in the extreme right of the tectum. The next
neuron, just to the left of the first cell, will occupy target
territory just to the left of that of the first cell, and so on. This
type of organizadon is referred to as topographical. The
topographical connectionsofneurons can be so precisethat
even if the eye is rotated 180 degrees,the axons will still
grow into their intended patch, with devastating results..

whiskers,which are the samewidth as the widest part of an

animal's body, send signais via nerves to the brain. If the
whiskers are touched simultaneously on both sides, it is
ciearthat the animal's head is in contactwith a passagethat
will be too narrow for the rest of its body to enter.
Whiskers are served by nerves that project into the brain,
where they are allocated neurons that are grouped together
in arrangementsthat resemble the silhouette of a barrel.
These neuronal barrels can be seen easily, even under the
light microscope. It is relatively sbaightforward to. tamper

Anv, frog so nodified will now see the world upside-down

experimentally with &is particular neuron-to-target rela-

and thus will have enormous trouble sxlching flies.

tionship by extirpating some of the whiskers. In the mouse

However, a topographical arrangement of neurons can

adaptto extractthe bestfrom a changings1_qlaliog_lf

b{!,the.
't-'i;i.:'-;ar';-:r=-"ciiitgfOrniiiigneurons are deshoyed, then the remaining 50

brain we can seean adapting situation comparableto that in

ftg. tgg's brain- If the"system .wasrigidly-prc$amlriedaflif
inaccessible to change, then one would expect to see

I
i
I

per cent of neurons - which would in normal circumstances

conspicuousgaps in the barrel-like organization of neurons

have occupied respectively only 50 per cent of the tectum -

in the brain. Thesegaps would presumably occur becauseit

will now invade the entire territory. Conversely, if half of

wils no longer necessary to have neurons allocated to

the tectum is destroyed, then the full complement of

whiskers that had been puiled out. In reality, just as we saw

outgrowing neruons will crowd into the smaller target

for the frog, all target territory still ends up being used: there

'

t
t
t
J

14 6 THE HUM AN BRAI N

C E LLS U P ON

are much larger groups of neurons allocated to the remain,

-aeeordingto the working cells, which reflecte-din. tufn the

'ing;neurons- The barrels'of neurons become enlarged to fiII

the available space.

environmental requirements in which the person had to

il9, F th" expandinghumanbrain,suchsensitivityto local

factorswithin brain circuitry is rampant.As our develop-

.Alrlrough this experinent might have little rdleVanCefor

humans at first glance, a similar chain of events caused a

I
I

--

the brain are very reactive as they form circuits to reflect

in one eye. The cause of his blindness was a medical

whatever is happening in the individual outside world.

mystery. As far as the ophthalmologists could tell, his eye

Inside the brain, right up to sixteen years of age, a bloody

finally emergedthat when he was a baby, the boy's eye was

bandaged for two weels as part of the treatment for a minor
infection. Such treatment would have made no difference to
our older brains, with their more established
But so soon after birth the connections of the eye were at 6
critical period for the establishment of eye-to-brain circuits.'
Since neurons sewing the bandaged eye were not *o\
ing, their normal target became taken over by nerves from
the'normal; working eye, as we have seen already with the

-:;;j

battle is being waged between our neurons. It is a battle for

i

establishing connections. If a new neuron does not make

--contactwith a target neuron, or is insufficiently stimulated,

;-!1"" it dies.
9 ". i :i _

Flirr.Aswe interact in this way with the enviroument, we

becomemore adept at surviving in it, as more and more of
the appropriate (that is, the most hardworking) neurons ire
connected to enable the most effective signalling. Even
a specific brain lsgisn, some brain circuits grow

frog's eye and mouse whiskers. In this case,tl.e neurons that

than others.Such circuits are also the most electricallv

were not working were treated by the brain as though they

,*tive (see Chapter 3), as*well as being the most metabol-

non-existent neurons was therefore readilv invaded bv the

H["3,"ti1t-r*ig-lhp.t"they*.are-rieher-in.-ehenicalsfordreoanufacture of the energy-storingchemical adenosine tri-

active brain cells. Sadly, the bandaging of the eye was

In the b1ain,then, activity

werenot thereat all: th9 tqggl_9f thgse_

inactive,fui

;;-^":;;;:.

ment continues after birth, the jostling, restlessneurons in

tragedy for a young Italian boy. This six-year-oldwas blind

was totally normal. Eventually, the enigma was solved. It

C E LLS

:.:::;::--

:'-

''-'

' *-

misinterpreted by the brain as a clear indication that the

would not be using that eye for the rest of his life.
Normally, this 'use it or lose it'rule would be beneficialas
it would mean that neuronal circuits were being established

growth go hand in hand: it is not only a question of 'use

t[ or lose it' but 'use it as much as you can.,(SeeFigure 10.)
itt.Aoy small change in the normal lifestyle or environment
rill be reflected in a change in neuronai circuitry. For

117

C EL L S U PON C EL L S I''

determineyour individual memoriesand.-so;-as

we shallsee
ln the-bea-chapter,makeyon intii-tne p;n;; t;u are.
It is a popular idea that this selectionof connectionsis
takeri from a wider pool of exibting connectionsthat are then
'lost,' rather like making a sculptue byremoving or losing
the extraneousmarble or granite. Although many neuronal
connectioruido undoubtedly die off in development, such a
loss is more than offset by the runaway growth of the brain,
as it forgesthe appropriate connectionsbetween neuroruias
a result of how much thev are being used and consequently
F I G U R E IO
The incessant adaptation of the brain to experience.Each of a monley's five
digits (D1-D5) are rcpreseuted in two adfacent areasin the part of the cortex
devoted to processing touch: the 'somatosensorycortex' (a).Diagrams (b) and
(d) outline the neuronal allocations for each digit of an adult moukey
corresponding to digits in (c), betre (b) and after (d) naining. During bainlug
the monkey rotated a disc for one hour a day, with digits 2 and 3 and
occasionally 4. Three montbs later (d), the arearepresentiag the fiagers that
bad received the exba stinulation hom the mtatilg task in the brain, has
increased markedly. (Simplified from E Kandel and R. D. Hawlin, Mind and
Bmr'n,W. H. Freeman& Co., 1993.)

example, kittens were siTply hained to lift up one paw to

show they could discriminate between patterns of horizon:"r::.,:";:;i:;!:rJ,tatf6rlusVgrdChflfie3:Ea-riiinatioii cif their biain;"eal"d
an increase of about 30 per cent more corulectons in the
specific part of the cortex relating to sensationsin that paw.
Thus, it is connections that are important, and the degreeof
the stimulation from the environment will determine how
the connections between neurons are formed, and thus

how active they are. Hence there is no genericbrain that gets

shaped into an individual one. Rather, the individual grows
'.!rii!".
:{r':,

as the brain does, over some sixteen years in all.

At last, then, we come to the mature brain that, at the age
of sixteen, has frnally increasedin size by S per cent over the
preceding elevea years or so. Although the brain is particularly impressionable while developing, such adaptability
does not ceasebut merely lessenssomewhatin maturity. It
is actually possibleto manipulate the environment and still
observelong-term changesin the mature bmin. For example,
they had an abundanceof toys, wheels,ladders, and so forth
with which to play. ln contrast, other, similar rats were kept
in an ordinary cage,where they received as much food and
water as they wanted, but had no toys.
When the brains of these two groups of rats were
exemined, it was found that the number of connections in

I50

I
I

T HE HUM AN

BRAIN

the brain had increased only in the animals in the enriched

C E LLS U P ON

C E LLS

fasterreaction times. Although the middle-agedbrain is still

not in those from the-ordinary cages. It

-_,environment,
appears that sheernumbers of neurons are not as important

in terms of certain
and reacting to the t""ito"*tnt,
"rrotrring
processesit is siowing down - for exqmple,in the acquisi-

as _the connections between them in the"brain. and these

tion of new skills such as driving. Although youngerpeople

connections are hiehlv changeablenot iust in development

do not drive better, they are better at learning to drive:

but also in adulthooC

statisticsfrom the British School of Motoring suggestthat

the averagenumber of hours tuition (the number of hours a

^connectivity in highly specific neuronai circuits.

We shouid be cautious about seemingiv obvious social
implications of this type of study. An enrichedenvironment
for a human would not simply be one with accessto more
material possessionsor increasedphysical activity, such as

I5I

personpays for at the schoolto learn how to drive) roughly

matchesthe student's age.
Our brain continues to siow down in certain ways but to
adapt and changein others, Most of us nurture the hopes of

repetitive dancing to a beat. Ralher, the key factor is

stimulation of the brain. In the study with rats. it was forrnd

known that we are living longer. Indeed, the increased

that it was the activities that involved learning and memory.

emphasison brain disordersof the elderly is one of the most

living a long time, eventually reaching old age. It is well

compelling neasonsfor studying the brain. In 1900, the life

changes in the brain. You do not have to be rich to stimulate
your brain with' enriched environment.' Human stimulation

expectancy was forty-seven, and only 4 per cent of the

can be achieved informally outside of the schoolroom by

lively conversations, meaningful relationships, crossword

12 per cent of the population was over sixty-five. Twenty per

population was over sixty-five, whereas in 1990 more thar'

cent of the population will be over sixty-five by the year

puzzles, and incessant reading, irrespective of whether such

events occur in an inner city or on a beachin the Caribbean-

.-chance.ofbeing in excellent'healthdue to"a good diet,"bdtter

Thus, as we tive out our lives, *" t"rt ion the connections

medical care, and an increasing interest in physical fitness.

between neurons that endow us with an individual, unique

brain. Nonetheless,by the time we arrive at middle age we

However, it is at this final stagein life that the brain starts

2020. More &an any other generation, we have a greater

to diminish in its mass.There is a 2o per cent loss in brain

are fairly flxed as a personality, or we think we are:

certainly, inside the middle-agedbrain someprocessesare

weight by age ninety, and even by age seventy there is a 5

starting to slow down a iittle. Younger people will have

discussedin Chapter1, we know that the remainingneurons

per cent loss in brain weight. On the other hand, as

:.1
,t

tr

I52 THE HUM AN ERAI N

C EL L S U PON C EL L S I5 3

. can take over certain roles. Whv d_oesthq brain-.agg?There people perform worse on some probiem-solvingtasks and
that run out of genetic information, or the genelic-p1ogfam.
- sudilenly becomes,,ru;r"i to **J;
;;"g;
over time, or
inactive or harmfui proteins are suddenly prodriced. We still

informationa tiftte sl-oWer.:theieis no:evidence that

abiliqy decreases with age. In fact, vocabulary
_ feqrying
improves. Politicians, heads of business, heads of the

do not know the causeof the devastatingdiseasesof old age

church, and political'leaders are very often in their sixties

- Alzheimer's disease and Parkinson's disease- where

and seventieswhen presumablyat the peak of their powers.

different parts of the brain are subiect to massive neuronal

It is perhaps telling that in ancient Rome you could be a

loss. However, it is important to realize that these diseases

judge oniy after reaching age sixty.

are actually iilnesses; they are not a natural consequenceof

old age.

Even at the physical level tlere is no reason to assume

,t","that we are all destined to become debilitated. One lady,

In a recent study of Alzheimer patients, it was found that

i, Hilda Crooks, climbed Mount Fuji at the age of ninety-one.

a certain region of the flein (the medial temporal lobe) was

Old age can be the ultimate expression of you as an

less than half the size of that of a non-Alzheimer patient of

individual. In the next chapter, we now turn to seehow that

comparable age.Even more dramatic was the discovery that

individuality can be expressed in terms of the physical

the rate of thinning of this brain region is far greater in.

Alzhermer patients than in. normal ageing.persons.ThuqAlzheimer's disease is a catastrophic event for the brain,
with devastating consequences,but it is not the natural
destiny of us all.
;;;;;;i:;;:':-,,-"-".N-gfr"ejltgless-brai*ellsdo-ehange-inifOfmal'ciftfag6. nE
believed by some that there is a reduction in the dendrites,
the receiving areaof neurons, although this idea is still hotly,

t
;

debated. If this is true, it might be thought that our

processing abilities decline, but a recent study has showu
that we can still'process an incredible amount of information. We know that older rats can still form new connections

.--.t. . .
I

VYIT H M IN D IN M IN D

l
I

I!
i
I
I

Where is the root of individuality? By looking at a singie

human brain it is educated guessworkat best to determine
whether the person was male or female. It would, however,
be completely impossible to tell whether this particular man
,fi-:

or woman had beenkind or possessed

a senseof humour. As
'we saw in Chapter 1, all brains comply with the s'-e basic
Sound plan: there are nerves carrying in information about
senses,and other nerves coming out that contract the
i: uuscles and are responsible for movement. We have also
=
seen that the flnin is made up of neurons, and that the
', circtritry in which these neurons operate is in part genetic
to a very great extent - at least in relatively complex
'hains - also influenced by the envilonment. How might
...suchcircuitry be translated into an individual? We address

1!!9 isguein this shnpter.

Identical twins are clones of each other. They are two
people with identical genesbecausethe single fertilized egg
:qllit into two. But are
they identical people? Certainly,
haging with nuclear magnetic resonance(NMR) scans{see
1) of the brains of identical twins shows a greater
dmilarit-v at the gross structural level. If identicai twins are

THE HUM AN BRAI N

WITH M IN D IN }.IIN D I5 7

questioned about their preferences,attitud,e-qr

.qn"{gxpg-rien--.

gsisthere are, perhaps-not-surprisingly;-otten"eotrsideiebli;

similarities. However, a coincideuce in tastes and ideas
might not be that-remarkable'iu any siblings-real'infin'ihe'same environnent.
Identical twins will also show signs of distinct percep-

iaeessant experiences thrown in our paF: For -expqrienCes

-"-io
fti"" *y-i"tu"g significancein this way, they needto be

' remembered.The essenceof the individual thus lies in no

='ttioi.d
*hai he or she can remember. Perhapswe
p"tt
-itn
- could start with memory as a way of approaching the secret
of the physical basis of individuality.

tions and thoughts that rnake it clear they are individuals

In English at least, memory cen serve as an umbrella term

with their own priwate consciousness,even though their

for a diverse range of processesthat may weil be quite

-distinct. Comparethe memory processesof an octopus and a

genetic make-up is the same.

gble by genes,it must, at least in part. be due to some other

human. The octopus has one of the largest brains of all

factor in the brain that is not shared even bv descendantsof

invertebrates,roughly equal in size to the brain of a fish, and

the same egg.

We have already seen in the previous chapter how

i.. composd of some 170 million nerve cells. Although this

.,[rrmber seems large, it is trivial in comparison with the

experience is a key factor in shaping the micro-circuitry of

iri,humanneuronal count of some 100 billion. Nonetheless,the

the brain. Eating something that you associate with an

{-iictopus hasprovedpopular in learning and memoryexperi-

unpleasant event could well make you dislike that particu-

i: ments,becauseit has highly developed eyes and a sophisti-

lar type'of food. Even more simply, only those e4posedto

::'catedsystemof touch via its many tentacles. In experiments,

aD odopus can clearly tell the difference between certain

the music of Mozart, for exdmple, will ever have the chance
to say that they have a preference for Mozart. Experiences
we have never had can play no part in framing our

blours

and can attach different meanings to each. For

it will readily grasp at a coloured ball it has
-leamed is associated witlr delive-iy bf a prawn,

if the person was never exposed to different languages.

but will not react to a ball of a different colour that has not
been paired with anything either rewarding or aversive.

is perhaps most

ii'i This type of memory, a simple association between a

dramatic up to and induding the teenageyears, but even

then the brain is not caught in freeze frame. Our character

lcoloured ball and a prawn, may seem a far cry ftom the
&emory of a particular hot surrmer day by the sea,or how to

continues to adapt as we respond to, and. rgcoil from, the

a bicycle, or remembering the French for 'window.'

The process of evolving a unique lpin

I58

T HE HUM AN

BRAIN
W ITH

There are many rlj:;tinct types of brain processesthat fall

under the general te'rmmemo.ry.The most basid ana famitiar
distinction is bet',areenshort-term an@.

MIN D

IN

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*;!lgg-1erm memory for nonsense words also have .a poor

long-termmemory for unfamiliar namesof toys. Short-term
srrd long-termmemory appearto work not independentlyin
parallel but in series.First, short-termmemory comesinto

Short-term memgrv operates when:we try to remember a

series of numbers. Everything is fine if there are no

operation:it is a transient,highly unstableand vulnerable

distractions, becausrrthe strategyis usually to repeat the

sequencein our mjnd over and over again.This processis

processwhere attentionand rehearsalare neededin orderto

surprisingly modest: we can only remember an averageof

seven digits.

memory. Successfulrehearsalin short-term memory will

lead into the more permanent and dormant long-term

eventuallylead to that speciaiphonenumber being retained

One of the most obvious questionsto ask about short-term

memory is how it relates to long-term memory. This less
contrived type of memory processoccurs without any need
for repetition or rehearsal. Do short- and long-term memorv
tn a com

independent

? I t is

well known that patients who remembernothing about what

has happened to them beyond the immediate present, and
who thereby exhibit an almost global amnesia,nonetheless
have a short-term memory ability indistinguishable from
non-amnesiacs. ClearLy then, the two processes can be
separated, but coulrl someone have a normal long-tern

mgryory evenwhgn sho$:teng memoryabiliry i.*

Impairments in shart-term memory are difficult to study.
Long-term memory is not a one-stepprocessbut, just as we
saw for other brain functions in Chapter 2, it can be divided
into many different aspects. For each of these different
aspects there appears t-o be a respective form of short-term
memory. For examplct, yal.lng children who have a poor

r59

without constant attention to it.

,

We all know that short-term memory is improved if

numbers, for example, have significance, such as a tele-

-. phone number or a security number to a building or a safe.

, h *y case,if an item survives in your memory for more
than about thirty minutes, it is probably not going to be
forgotten, at least for a matter of days. Patients recovering
"ftom concussion or electric shock treatment (a radical
'- therapy for severe depression) characteristically cannot
:ii:'rememberwhat happened an hour or so before the event,
,,1,whereas their long-term memory remains operational. In

early rupture in the normal course of events subse,-.frt

quently obiiterates any chance of that hour of Iife beins
s'rr..tacorded
more permanently in the mind of the individual.
Sho\t-term memory operates to serve lnng-tcrm memnry
But what do we mean by long-term memory? It turns out
t

W ITH

that, once more, this other basic categoryfor the umbrella

--*'diiitinCt ph"rro-"rr..

ih"r.

MIN D

IN

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.--brarnremoved to conbol the epilegtic seizures.Despite its

-:succeCs-"in-combiting epilepsy, this operation has never

is much that we learn and

been performed since then, becauseof the terrible conseqLence:after surgery, H.M. could remember only events

remember as we g! through life: how to drive.a car, the:French for'thank you,' and what we did when Aunt Flo last
carne to visit: All these are examples of different types of

before his operation - up to about two years beforehand.

Sincethe surgeryhe hasremainedconstantlytrapped in the

memory at work. However, the odd item in these three

present.

examples would be how to drive a car. The memory for a

It is very hard to imagineH.M.'s stateof mind. He fails to

fact, such as the French for'thank you,' or an event, such as

the recent visit of Aunt Flo, requires that we should be

- - recognizefriends or neighboursthat he has got to know after

the operation.Although he can give the date of his birth, he

making an explicit conscious effort. In contrast, driving a

cannot give his conect age, always estimating that he is

car, like many skills and habits, is almost performed on

automatic pilot. This type of memory is therefore referred to
as ,W,
because we do not need to actively and
consciously rememberhow to do something: we just get in a

16!

;;,-

younger than he actually is. During the night he might ask

,,,the nurse where he is and why he is there. He cannot
fi'''reconstruct the event of the previous day. He explains,

iffL,: Tvery day is alone by itself, whatever enioyment I have had,

car and drive. When you approach a red light your foot is
'automatically' on the brake. In contrast to this process,
memory for .events and facts. can be regarded as gxplibit
memory.
One of the most famous and intensively investigated cases
of complete loss of explicit memory is that of H.M., a young

.ffi--at

"

result of this condition,H.M. has only beenableto

,- qarryout simple acts in the here and now. Therefore,he was

. -gven monotouous iobs such as mounting cigarette lighters
g4fdboard-displays,-He eould' not givd d &isdiiption' of

man;.-who-had-severe
epilepsfime -c"onditio"
il *nifr"tnt

the place in which he worked, the nature of his job, or the

patient is stricken with a seizure accompanied by loss of

route along which he was driven every day.

consciousness.In H.M.'s case, these epileptic seizures

". H.M. can still rememberstings of up to seven digits, thus

demonstating that short-term memory is a separateprocess
tom the subsequentstageof long-term memory. Moreover,

became so frequent that it was impossible for him to live a

normal life.
In 1953, at the age of twenty-seven,H.M. had part of his

fil' although H.M. appears to have lost his ability to remember

I62 THE HUM AN BRAI N

W ITH

in the long term, his brain has retained

a different t-vpeof
memory. H.M. can actuaily perform
quite well- at eertafn
moio. skills like tracing a star. However,
this task is not as
easy as it sounds since the outiine
has to be traced while
looking in a mirror: it is a demanding
exercisein sensory
motor coordinationthat improveswith practice,
like driving
or riding a bicvcie, Everv day, FI.M.
did indeed improve,
showing that this other type of memory,
implicit memory,
was not processedin the samepart of the
brain as memories
for events.Interestinglyenough,H.M.
was not consciousof
remembering the event of learning to
draw the star (an
example of explicit memory), although
his brain was quite
happilygettingbettera t d o in g s o -imp lic it me mo rY
Particularly relevant to our present
discussion is that
although H.M. cannot remember events
occurring after, and
two years prior to, his surgery, past memories
from long ago
are still intact, trapped in the brain like
a fly in amber.
memories are obviously not dependent
on the brain areathat
has been removed. It must be the case
that no one brain
region can assume the entire responsibility
for the whole
processof .factsand events.Ratlier,
'.}em-gl]4
memoriei musi
be somehow processedthrough one region
but consolidated
elsewhere.In H.M.'s case,the damagemust
have intervened
at the stagewhere a new memory is first
processed.Hence,
all the memories that had aiready been
consolidatedwere
safe. Just as we saw in Chapter2 in our consideration
of the

MIN D

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sensesand the control of movement, different brain regions

for different aspectsof a function.
Iie r-ebponsible
The areathat H.M. had removed was the middle part of
his temporallobe, which lies on either side of the brain, as
its name suggests,by the temples, just abovethe ears.This
areaalso includes a region underneaththe cortex cailed the
hippocampus.which is Greek for 'seahorse,'becausesome
peoplethink this structurelooks iike a seahorse.It is, to my
-mind, actuaily easiestto visuaiize the hippocampus as it
nestlesin the brain below the cortex as a structure shaped
more like rams' horns, curling around the inner mass of the
''brain. A considerableamount of clinical and experimental
i,widence has shown, subsequentto the caseof H.M., that
:demageto this
brain region results in an impairment in the
Iayrng down of memories.
Even for this more specific aspect of memory, its initial
consolidatiotr, there is another area that appears to be
: tlg medial thalamus. which is vital for the
l.,relayrng of incoming sensory information onto the cortex
i fseeChapter2). Iust as the processingof hearing and vision
Ig]lt gn a,different-part,of the thalamus, so there iS a
, rpecific area of the thalamus contributing to memory.
; . We know that the medial thalamus contributes to memorv
lrecauseof one or two unfortunate and bizarre accidents in
which people have ended up with either fencing
foils or
rnooker cues up one nostril, thereby destroying
the medial
ii:':thalarnus.In such
cases,the victims of these accidents have

r63

wt r H

displayed amnesia for events. Unlike the elamples o{

iffi

;i;
):..*--,
-. ' '
"
-'":"

-..'
*-. --- ---*T::--- - - i-- ; ---'
Seen only temporary,

events are rernembered,as occrrrringiat a.certain time and

ice,-bnt-*tio i" how they are associatedwith related

however. Despite the fact that the

amnesia can be temporary, _thereis apeunannt inabilitytor':

-'-......'--'..
memory of events that occurred while the amnesia lasted,.

'

] ilND t N r {t NO lat

presumably whiie the medial thalamus *",

-"mroctioning.
Hence, as for the hippocampus, the medial thalamus can be
viewed as important in the consolidation of memories.

eveotsat presumably a similar time or place.

'*flc6; as in semantic memory, need differ only from the
- events of episodic memory in that they are removed from a
specific'moment and place. Once the pink elephant is
displaced from the jungle hideaway in which you saw him
one night in the last summer vacation, he becomesreduced

an event occurred. If there is no space or time reference,

to the generic thought that elephants can be pink. Damageto

the areawhere facts have been personalized into eventsby

events cannot be differentiated, and there is no personal

+ime and space referencing would not achrally destroy

involvement of the individual with what hao happened.

rnemoryitself but rather would uncouple facts from the

Because events are unique and personal, whereas facts are

in which they occurred. Specific events would be

generic and free of time and space frr-es of reference, it '

to mere generic facts in that they would have no

Source amnesiais a loss of memory for when and where

follows that source amnesia will primarily affect menory for

or unique features in time and space.

and events appeius to rely on the integrity of the hippocam-

!,:l If the prefrontal cortex is needed for this type of timeisPaceallocation of events, and if, as we saw in Chapter 1,

pus and the medial temporal lobe, only memory for eveniC

*,+fheprefrontalcortexhasshownan extraordinarydifferential

seems affected by damage to this third area, the prefrontal ,

lgmwth during evolution, it follows that this type of memory

ifur-events would be particularly pronounced in humans,

events rather than for facts. Whereas memory for both facts

cortex, which we encountered in Chapter 1.

Interestingly enough, damage to the medial thdamus,

Iargeprefroittaliiirtiix, anAA
frr less prominence in other animals. For other animxls,

result in special types of errors in the time-spaceallocation

of memories. Memories can come forward inappropriately, :
out of context, when they are irelevant to the speech and
ideas of the present moment. The prefrontal cortex is "
presumably havirig ssms influence not just in the way

perhaps memory of an event is more generic, less anchored

trnique rime and place coordinates. A cat may not
;ltmember a specific spring day when it caught a mouse in
tie back garden just after drinking a saucer of milk and
climbing a hee, although it may well have a vaguer

I56 THE HUM AN BRAI N

W ITH

and more generalrecall of catching mice. Interestingly,

there

is a specific,contrived-situationwhere our human mbmories

also seem to be more like this generic type of memory.
. These pioneering studies were:performedin
Canadain the
mid-1900s by Wilder penfield, a surgeon.penfield
worked
with five hundred patients who were undergoing
neruosur_
gery. It is often a ghoulish surpriseto many that there
are no
sensorsfor pain within the brain itself; henceit is possibie
for the brain to be exposed in conscious patients
without

IN

MIN D

be operative during memory of an event. We have

.g-g@-ally
seenin ChapterL, and just now, that without the prefrontai
cortex our memories still exist but iue vaguer and less
specific, perhaps resembling the dream-like memories of
patients,or even normal dreams.If a reduction in
Pen-field's
the role of the prefrontal cortex, for whatever reason,did
indeed induce a more dream-like state of mind, it would
follow that animalswith a lesspronouncedprefrontalcortex
do not have the precise memories that we have. Instead,

them feeling any pain. With the patients, consent,penfield

used the operations,which had to be performedin any
case,

their memorieswould be disembodiedfactsthat lack a time-

to investigate the storageof memory in the brain. since

the
surface of the brain was exposedand the patients were
fu[y

almost become 'semantic' memory for a fact.

conscious, he was able to stimulate different parts

of the
cortex electrically while documenting the reports
of the
patients as to what they were experiencing.
Most of the time, perhaps not surprisiogly, the patientsdi*not

report any new experience.Sometimes,however, a

rather interesting phenomenon occ'rred: the patients

: - ; . * : ; i; ^- - l; , "- ' :

MIN D

claimed they could remember very vivid scenes.Thev

often
said that memories were like a .dream; they-.were*
generalized experiences that did not have particular
time
and space points of reference. perhaps in this
highly
" '

artificial situation, the electrical stimulation was locally

kick-starting the medial temporar lobe, without recruiting
other requisite but more remote regions. of these remote
brain regions, the prefrontal cortex in particular would

spacecontext: 'episodic' memory for an event would have

As we saw fur Chapter 1, the prefrontal cortex appears to
, be important in working memory, where incoming informa',- tion and ongoing behaviour are influenced by certain
internalized and individual ideas, perceptions, or rules, the
resources accrued over a lifetime that constitute an
idiosyncratic mind. These inner resources would provide
somesort of counterweight to the bombardment of the brain
by a tonent of implosive sensoryinformation. Damageto the
coitex hail.oftenbeen compared with schizophrenia and, conversely, schizophrenia has in part been attributed to a malfunctioning of the prefrontal cortex (see
Chapter1). A common and conspicuousfeature of schizophrenia is excessiveattention to the outside world, which
often appearsoverly bright and buzzing, without the sobering perspectiveand exposition-basalinterpretation of inner

167

I5 8 T HE HUM AN BRAI N

resources. Perhaps dreamers, schizophrenics, and nonh,umansshare a similar type of consciousness;characterized

by little memory for previous events, and dominated by
generic facts and the imrnediacy of the here and now. If so,
such a profile of awarenessmight have contributed to the
character changeof PhineasGage(seeChapter1) following
the severe damageto his prefrontal cortex.
So far we have seen that for explicit memory of both

169

Patientswith this diseasenot only have the same type of

riibnory impairment as H.M. - namely amnesiafor every" thing that happenedsincehis surgery(anterogradeamnesia)
'tr'bui also loss of memory for everything that happened
,--. before being taken into the hospital, and even before the
onsetof the illness (retrogradeamnesia).
The distinction between anterograde and retrograde

events and facts, clinical cases(for ocample,H.M.) suggest

amnesiawas demonstrated in a study carried out in the

"1970s.Korsakoffpatientsproved worse than H.M. at recog-

that the hippocampus and medial thalamus play a role in

nizing the faces of celebrities who had been famous in the

laying down memories for about two years; these long-term

1930sand tg+0s. The problem in learning about memory

memories are somehow 'stored' in the temporal lobe, as

Korsakoff patients is that it is hard to isolate the

revealed by Penfield's studies. Meanwhile, the preftontal

ts from other types of thinking processes.With alco-

cortex, with which the hippocampus and medial thalamus

the brain damage is so widespread that there are

both have connections, coordinates facts with an appropri-

lna"y conditions affected other than memory. Korsakoff

ate time and space context to ensure that an event is

patients, unlike H.M., would have extensive damage to a

remembered as a unique happening.

i --.-.

WITH M IN D IN M IN D

of other brain regions, incuding large areas of the

-- - --How might a fact or an eventbe stored in the brain? We

know that even though memories of past events might

tlere a particular brain region where a memory is

suwive damageto the thalamus and hippocampus, they are

y laid down? PsychologistKarl Lashley tried to answer

far from indestructible.It is helpful to cq!npar-e*_H,M_J

the-1.940s.
Lashley bained rats'offa mdriiory-* -'

niemory loss, where there was removal of the medial

in mazes,and then removed different portions of cortex

temporal lobe, with another type of memory loss from

seeif he could identify where the rnemory engram might

another group of patients. These patients have memory

stored. To his surprise and consternation, removal of

problems due to chronic alcoholism. Among the many risks

parts of the cortex did not result in a precise

of taking alcohol.in great abundanceis a diseaseassociated

between one specific area and the retention of a

with a dietary deficiency of thiamine: Korsakoffs syndrome.

memory. Rather the more cortex that was removed,

5; ; ; ; : - : ; l- : : : ; , _. i* ' - "- - . , * : ; , . - . - * - ; - ' '. - - - . '

I7 0

T HE HUM AN

BRAIN

W ITH

irrespective of specificity of region,

the worse the rats
became at the memorytask.
ntire

t
I
I
I
I

t
t

T
T

T
I

in the stor

, memory.
In line with Lashley's evidence from
rats, the clinical
casesreported by penfield would aiso
suggestthat memory
is not stored simpry; it is not laid down
directly in the brain.
Rather, as seen in penfieid's studies,a
cache of mernories
wouid be more like a nebulous series
of dreams. One
immediate problem was that the memories
themseiveswere
not like highly specific recordings on
a video and were a frr
cry from the memories on a computer.Another
problem
that if the same areawas stimulated by penfield
on
occasions, different memories were elicited.
Conversely,
same memories could be generatedfrom
stimulating di
ent areas. No one has yet showu definitively
how these
phenomena can be explained in terms
of brain
One possibiiity, however, is that each time penfield
lated the same site, he was activating a
different circuiby
neurons, where each circuit might participate
in a particul
memory. Similarly, when stimulating
in-,another
Penfield may have sometimes been activating
a circuit he
had activated before,but simply from
a different
point - once the same circuit was activated,
from

wha
triggering point, the memory would be
the same.
dPenfield's find
ehow

MIN D

IN

II4IN D

r;ould be a member of a number of di{ferent

__*Oneneuron
circuits; it would be the specific combination in each case
one circuit from another. Each circuit
*- ttrat distingrrished
would contribrrteto the phenomenonof a memory, so that
no singiebr;rirt';ell or exclusiveiycommittedgroup of ceils
instead, tlre memory would be
is rvhoily rr;:;pr.rnsible;
'l'he biochemist Stephen Rose reached this
distributed.
conclusion when he trained chicks to go against their
natural inclin;rtion and to avoid pecking at a bead.
ln brief, Ro:;efound that different parts of the chick brain
nrereprocessingand rememberingdifferent featuresof the
size as opposedto its colour.Justaswe saw
{:,;. bead,such as its
;;i for the visual processitself in Chapter 3, memory for the
tsgg
sigbt of an obiect was also laid down in parallel. &t"
si[gle region for a memorv but rather it is distributed over
According to the modality of what is remem,Egggo"r
and the associationsit triggersfor a certain context,
different levels of circuitry will be recruited over the entire
cofiex. ltis easy to seehow Lashleycould have been under
the impression that virtually all the cortex was somehow
ng togelher in the memory process.
..: How do the memories actually become consoiidatedin
',tbe cortex in thr: first place?We have seenthat all types of
''''memov first r:nter the highly transient and dissociable
Phaseof r1tol-tr:rm memory, but short-term memory only
..-lastsat most far half.an hour. In contrast is the striking case
of H.M., wherc, aithoughhe had perfectrecallof all that had

I7I

IZI

THE I . I UM AN BRAI N

wlTH

r.ilND lN r.ilND

t73

happened early in his lifg-,h9 qould not rememberanything

the hippocampus and medial thala-us to consolidate mem'--::." -::.:ories;it:is.:16g1ust.a- matfrti bf a tfew'-rntinuieJ'butof a
substantial p,eriod of ti'ne.
No one really knows exactly how the hippocampus and
medial thalamus might be working over a period of years, iu
conjunction with the cortex, to lay down memories that will
eventually no longer depeod on the integrity of these
subcortical structures. One attractive idea draws on a

memory being composedof otherwisearbibary

brought together for the first tine in the event or the fact
be remembered. The role of the hippocampus and
thalamus would be to ensure that these disparate, p

hippocampus dissectedout from a rat brain. (photo courtesv of Dr. Nick

Oxford.)

ously unassociated elements .ue now associatedand

';-'^ **

somehow bound into a cohesive memory. |ust as we saw

y, as we saw with H.M., an established memory

the simple case of the colour versus shape of the beads

*"-'
-*'"- Ko'Cli's"rixperiment," different part of the cortex would

rem_ain.
intact, fr,9g_{up from, and entirely-independent
the hippocampus. One metaphor might be that. of

involved. Some mechanism would therefore be needed

while a building is being established, the

recruit these different and remote neuronal populations

al of the ssaffs[ding would lead to the collapse of the

'1s*tonmTerdif,-the
It is possible to imagine that initially the cohesiveness
the working cortical network that represents the
depends on an ongoing dialogue with the hippocampus
medial thala-us. However, as the network becomes
lished, seemingly over a period of several years,
the subcofical sbuctures become less important, such

is redundant.
explicit memory for events and facts depends on an
dialogue between the cortex and certain subcortical
, perhaps this same saengement could also apply

the laying down of skills and habits: implicit memory.

habits, such as remembering sequencesor making a

I74 THE HUM AN BRAI N

W ITH

IN

} 4IN D

certain type of movement in an appropriate context without

of this -chapter: presentation of an erstwhile

needing to think about it, can be performed adequately in

naiiihalstimulus, siich as a ball, evoked a responseonce it

,hadbeen assocatedwith a meaningful stimulus, such as a

amnesiac patients with medial temporal lobe damage.

However, paJients guffering, from- dissrders of:::the bas
ganglia such as Parkinson'sdiseaseand Huntington,s chorea

ilratarn;Certain iypes bf cond.itioning involving an immediate rnovement of the muscles are now thought to be

(see Chapter 2) have seemingly no problem explicitly

remembering facts and events.Instead,their problem is that

controlledby the cerebellum,the little brain on the back of

the head (seeChapters1 and 2). For example,it is possible

they are no longer able to perform the habit of an appropriate sequenceof movement,or of recognizingthe next item in

in both rabbits and humans to condition the eye to blink to

a sequencethat had been shown to them over and over again

when it is associatedwith a natural trigger for the eye blink,

and normally would have been implicitly remembered.

zuch as a puff of air.

.
We can see that for habits and skills. different brain

An everyday example of a habit is the ability to generate

the right type of movement at the right time. patients with
Huntington's chorea no longer generatea movement in an
appropriate context - for example,the flinging of a limb that
can characterize that disorder might be appropriate on the
baseball pitch, but not in the shopping mall. On the other,
hand, Parkinsonian patients can no longer sequence'a
movement: the more complex the sequence- for example,.,
standing up, or turning around -the greaterthe impairment..
In both of these very different disorders of the basal

MIN D

- ---thereisa lapsein the imphCit'memory;tri;*;

; fault in

different aspects(respectively,context and sequence)of the

habit of generating movements

I75

an erstwhiie neutral stimulus, such as the sound of a bell,

-*,sEucturesare involved compared to those used in explicit

i i --,- - , r
r...
r.
^ -- of
^.c 4-:-- r events.
facts and
memory
A criticalr difference
lies not iust
iin the identity of these structuresbut also in their relation to
&e cortex. Whereasthe medial thalamus and hippocampus
have sbong reciprocal connections with the cortex, the
with the basalganglia and cerebellumare not so
or dominant. The striatum, which is the pivotal part
the basal ganglia in both Huntington's chorea and
is,-disease,reeeives an-input from' the-coiGx, but
doesnot send one directly back. Similarly, the cerebellum,
r.while indirectly linked to the cortex, does not have any
.direct connections. Hence it is tempting to imagine that

The basal ganglia are not the only brain regions to be

involved in implicit memory. Some memory tasks involve

ir-thesebrain regions, unlike those involved in explicit memare left in a sense to freewheel more autonomously.

conditioning, rather like we saw for the octopus at the

scenario might be expected of activities such as

.'',..-''.-.'" .-'?

1 76 THE HUHAN BRAI N

W ITH

implicit memory that are performed withoul attention or

- conscious-effort: such activities would nof n6ed-lonitant
referral to the cortex, which is known to play a key role in
-eonscious attentiori. As we saw inr Chapd; z, orr.. .-

I' !IN D

IN

MIN D

Again, purely for simplicity, let us think of each of these

elementsas representedby two single cells.
- During a memory, the two previously unassociatedneu_
rons would be simultaneouslyactive, and that coincidental

movement can become automated,either by internaiized-'T

activitv would eventually have some long-lastingresult far

triggers in the basal ganglia or from sensory inputs fed

outliving the periods for which each cell was active in the
first place.The easiestscenarioto imagine is that explained

through the cerebellum, the cortex is free for other functions, such as explicit memory, the rememberingof facts {
and events.
We have seen that memory can be subdivided into
different processesand that each process will be served by
different combinations of brain regions. But common to all
these memory processesis perhaps the most my
issue of all: We know that some people can rememberwhat
happened to them ninety yeius ago, but by then

in the 1940sby the visionary psychologistDonald Hebb.He

proposedthat when an incoming cell, X, was particularly
j active,and exciteda targetcell, y, then the synapsebetween
1X and I would become strengthened.By strengthening,
Hebb meant that this synapse would be more effective in
glemical signalling than the other, more dormant inputs
contact with Y. This idea is a version of what we

molecule in their body will have been turned over

already seen in &e previous chapter during development, when the hardest working neuron (in this case, X)

times. If long-term changesmediating memories are

ends up with the most effective connections.

ring continuously in the brain, how are they

i, The second, more recent proposal for an alternative wav

:ln which to strengthen a connection is for the strengthened
not to involve the target cell y directly, but to

ve of brain

how do neurons registermore

less pennanent change as a result of experiengqL_

- We have been conridering memory using top-down
egies. In order to answer this latest question, we have
travel bottom-up. Imagine a synapsethat was participa
in a memory process of any kind. Let us think,
simplicity's sake, of a memory in its simplest form as
association between two previously unassociatedel

luvolve the use of a third cell, Z. This third cell

would
onXbefore it signalled to y. Hence this strengthenwould be presynaptic, not postsynapticas in Hebb,s
e.If. Z and X were concidentally active such that Z
'ulated (seeChapter 3) the activity of & more transmitwould be releasedonto the final target y. Only when
X

177

I7E THE HUT. I AN BRAI N

W ITH

MIN D

IN

MIN D

Z (cornparable to Z above) that responds to a

naturally aversivestimulus in the tail impinges on a sensory

-a-ne*e.

--rlerv {XJ that responds to a benign stimulus. This sensory

nerve then linla up directly with the motor nerve (Irl witrr
which Aplysia retractsits gill.
Aplysia can be conditioned to retract its gill to an
otherwise neutral stimulus on the sensory nerve, rather like
can be conditioned to blink o'r eyes to the neuhal
stimulus of a bell ringing. when z and.x are concidentally
I

F IG U R E I2
Possiblebasesofadaptation ofneurons to experience,reflectedin the activity.
of incomi-ugcells.In Aplysia (a),the coincidentactivity of two cellswhere one
intercepts the other results in more bansmitter releaseonto the target cell. h
us 6am..xlian brain (b) a cell that has alreadybeenactivatedwill be ableto
respond more readily to furtler stimulation, as shown in the shaded.areas..
fDrawn by Dr. O. Paulsen, MRC Anatomical Neumphannacology Unit;
Odord.)

and Z were active at the same time would X consequently

release more trarumitter onto Y. (See Figure 12.)
- .1,=

". , .- Ttris scenario-hasbeen.most-effectively

the sea slug, Aplysia, which has the advantage of a
simpler nervous system, such that single neurons are
identified bv name. In the sim
Aplysia, there is no
bottom-up: the activi
-directlv

active (that is, when the benign and aversive stimuli occur
simultaneously),Z induces in nerve x a cascadeof chemical
t?actions that lead to the closure of potassium channels (see
Chapter 3). When the efflux of this positively chargedion is
prevented, &e voltage across the cell membrane becomes
more positive: this is precisely the voltage requirement
needed for the opening of special channels for calcium
to
the cell. When more calcium consequently enters the
tell, more trnnsmitter is released (see Chapter g).
More
"bansmitter releasedby the
sensory nerve X onto the motor
larre Y means that the motor nerve will work harder,
and
resultant behaviour, withdrawal of the gill, will be
Songer. Nerve X can remain in this potentiated
state even
Z ceasesto be active. The behaviour will have been
ned.

uni
neuronal

into demonstrablebehaviour. Here is one examp

Similarly, in the mammalian brain, it is plausible

that at
of the many synapses in the many brain regions
d in memory, there will be a strengtheningof certain

179

w tTH

hardworking synapses.A pivotal mechanismby which this

scenario is thought to be achieved is called Jons-ter-m.
potentiation &Tp'|. LTP worksby exploiting the fo*y *,*"
of a certain type of targeJreceptor (NMDA;.thatis, N-methylo.aspartate) for a certain type of bansmitter (elutamate).
Unlike the more normal sdenarioin Chapter 3, this receptor

ascoincidentactivity of incoming cells, could in either case

triggera changein the long-term responseof a target neuron.
This type of sustainedor coincident activity of incoming
neurons might occur in a memory situation. The ensuing

must also be active so that the transmitter in question - in

this case, glutamate - can be released to dock into its
receptor. Second is the unusual requirement that the cell
when these two requirements are met does the fussy
receptor allow a large amount of calcium to flood into the
target cell.
These two requirements could be achieved only if there is
coincident activity in either of two ways. One way would be
for two incoming cells to be active simultaneously, so that
each would meet one of the two requirements - one
incomi.g cell would releasethe glutamate while the other
caused a decreasein voltage by releasing another transmit- ,

, - ter-.T!.e.secondway-to fulEl both requirementiwouta fe

with just the cell that releasedglutamate. Initially the fussy :
channel would not open since, although glutemate was
released, the voltage of the cell would be normal. The
glutamate would just work, in the normal way, on a lesg
fussy tlpe of glutamate receptor. If the release of this

tN l ,i l N D

glutamate was susfarned, then the effecis of activating the

--"Iess fussy receptor would be to decrease
the voltage of the
target cell, thereby fulfflling the second.requirement. The
=-" r' fuisy::glutamatereceptor could
then open the charurel for
'
calcium ions to flood in. Hence, sustained activity, as well

will only trigger ion channels to open when two conditions

are fulfilled. First, as is normally the case,the incoming cell

must already have a more positive voltage rhnn usual. Only

H |N D

largeinflux of calcium again wiil trigger a chemical cascade

within the target cell such that another chemical is released
'

that ricochets acrossthe synapse,enters the incoming cell,

and acts to make it release even more transmitters. The
targetcell is in furn even more active and the synapseis said

F"r'' to be strengthened.when the incoming cell of the strengthl,' ened synapse is stimulated flgain, only modestly, the
. responsethat now ensueswill be greater,a little akin to the

:: enhancedgill with&awal of Aplysia.

T-je_is, tnown-E,
@on.This type of phenomenonmight accountfor short-terur
.J{owever, wd know-that.short-terui memory is just
that - lasting less than an hour. In order to account
for our
seemingly pemanent memories, more permanent qhanges
uust be taking place at the cellular level. LTp
in the
&ammalisl blain, as for the potentiation in Aplysia,
would
be a necessarybut not suffrcient factor.
If the enhanced
releaseof transmitter were sustained,
as it might well !e in

lll

ffiT

n u m AN

BRAIN

the laying down of a memory, then the hansmitter would

not only have to be passingon the messageto the targetcell

on the other side of the synapsefor a more vigorous or
potentiatedresponsein the short term. in addition, thelong- ,-,
lasting result of this enhanced activity would actually have
to be a changein what was happeninginside the targetcell.
Clearly, permanentchangescannotreiy on existing chemicais simply being releasedin greater quantities. Even if
certain enzymesbecomespontaneousiyactive, which they
do, thereby increasing the efficiency of the synapse, such
molecules only have a life span ranging from minutes to
weeks. Although much of what happens within the cell
during memory is still a mvstery, certain facts are emerging.
For both Aplysia and the LTP in marnmalian flains, ss
have seen that &e underlying corrmon event is the entry of
calcium into the neuron.
This entry can act as a trigger, in as little asthirty mfututes",
for activating certain genes by using proteins that in
themselves eire short-lived. Nonetheless, the products of
sudi genes are then able to activate other genes,which, by
the diverse ways in which they are expressed,can modify a
nsuron fol a very-long period. The effectsof activating genes
withiril the neuron might be to increasethe efficiency of the
transmitter, increase the number of receptors, or
increase the effi.ciencywith which a receptor opens an iou
channel. However, the alternative way in which a
might be changedby gene expressionis even more

WITH M IN D IN T.IIN D I8 3

We learned in the previous chapter ft"t

llg_efircS:_ql
are to r-rlsdifynot so .much the numbers of
"exqgrieng:
4guons themselvesbut rather the connections between

fr"rl, As a broad generaiization,we saw that the more

experience.the more connections.It is now known that
within an hour of training on a particular task, certain
important proteins are pressed into service. Two good
examplesof these proteins are the sticky badges,the cell
adhesion molecules that we looked at in the previous
chapter, and another aptly named growth-associatedprotein, GAP-43. Celi adhesionmoleculesappearto be important for neuron recognition and for the stabilizatioa of
neuronal contacts.As cell adhesion moiecules are made in
the brain, certain sugarsare incorporated..we know that cell
adhesion molecules are important in memory because if
sugarincorporation is preventedby administering an appro_
priate drug, then amnesiaresults.
GAP-43 is an example of another protein that may play a
role in memory - as its namesuggests,it is implicated in the
a ,growth of neurons. Growth cones (see Chapter 4) contain
GAP-43, and its synthesis is known to occur at a high rate
when neurons are extending their itxons. GAp_43 is appa_
rently activated during LTp. Hence, it is tempting
to
speculatethat the entry of calcium during the strengthening
+:of a contact during a memory task
will lead to enhanced
growth of neuronal contacts, perhaps
via GAp-43, and

:::::;=if

WITH M IN D IN M IN O

oKr{tN

saw in Chapter 4 were. during deveiopment, the way the

--- triggermany associations.For example,associatinga number ('3') with somethingthat canbe readily visualized('throo
blind mice'), and which is very familiar (an oid nursery
::r:-. ihyme), wiil improve the subsequentrecall of the number.

brain most conspicuously reflected changesin the environ-

--, An aitemative strategy is to imagine items of, say, il

ment. It would not be surprising, as we go through life, for

shoppinglist, distributed in different parts of a room, such

stabilization of those contacts, perhaps by means of ceil

.adhesionmolecules.
In this way there will be new synaptic contacts,which we

that a bar of chocolatemight be nailed to the door, the buttor

the processof adaptingto experience,namelv memory,to be

an echo in our brain of the developmentalprocess.

*"y be placed under the tabie, with the milk on the table
'
**'and the tea in the sink. An.alternative way of improving

underpin the memory process?This is a difficult question to

'

memory is to either piace yourself, or imagine yourself, in

the samecontext in which the rememberedevent originally

answer as it means bridging the gap, in the mammalian

brain, between the cellular bottom-up level we have been
discussing and the functional top-down approach addressed
earlier in this chapter. We need to know how to relate the
world of microscopic eventsin a multitude of neurons to the
macrophenomenological world of memory. Although in

occured. You might imagine you were on the beach during

,. surrmer vacation'in order to recall the name of the lifeguard
a.

.'with whom you struck up a conversation.A more sophisticatedversion of this idea is to imagine other items belonging

to the context in which the respective remembering took

' place (suntan
lotion, towel, sunglasses),In all thesecaseswe
.t:.

Aplysia it is relatiyely . straightforward to translate the'

,'would be either creating a maximal number of associations

activity of a certain neuronal circuit into a mechanical '

during the act of consolidating a memory or exploiting such

,essociationsduring recall.

behaviour such as retracting the gill, it is impossible i+ the

human brain to pin down the behaviour of having a certaiq
+neerv to a specific neuronal ciicuit. Nonetheless, some
features of the memory process do provide indications that
connectivity between neurons, albeit sophisticated and
currently unidentified, is important.

!.=ILis,-wellknown that most people cannot rememberevents

that occurred before thev were about three years old. This
iphenomenoncannot be accounted for by simple length of
, silce we are able subsequentlyto remember eventsfor
Someninsq/ years. Moreover, young children are able to
habits and skills from an early age - it is only

It is a well-known trick for improving memory to associate

the item to be remdmberedwith something that would itseU

i*plicit

memory that is the problem. On the other hand.

It'

I86 THE HUhAN gRAI N

WITH

babies as young as five months are able, arguably, to show

associations.

explicit memory by looking at a new item more than one

between the

M IN D IN I'IIN D

187

sical and the

they have previously seen, when the two items are presented together.Children under a year old may .opy g"*ri

aware of the

they saw someone else playing, even onlv once, on a

operation.lvlemory is muiti-faceted and muiti-staged.It is

more than a mere function of the brain, as it encapsulates

previous day.
It seemsthen that somesimple form of explicit memory is
available to young children, which in turn wouid mean that

individuais' :-nnerresourcesfor interpreting, in an exquisitely unique fashion, the world around them. As such,

their hippocampus and medial thalamus must be opera-

memoryis a good place to end our brief glimpseof the brain,

tional. With regard to maturity, more in doubt is the cortex.

for it is a cornerstoneof the mind.

If neurons in the cortex were unable to form man

tions, then the explicit memory of children would not be. as
jl[!eeI!.i! is not, verv robust. After the ageof three the ability
to associate items with a richer repertoire garnered froc
e:qlerience, aided in turn by an increased number
neuronal connectionsin the cortex, would make memorv,
we know it, possible.
Although these strategiesand examples might vary, the
basic theme is the same: to capitalize on associationswith
the remembereditem. At the neuronal level, these associations are certainly not a crude one-to-onea131shingof si
cells. However, within the vast interplay of neuronal cir-t
cuits of varying complexity, the basic unit of change
boil down to the modifications to connectivitv we have
discussing. We know that long-term memory is accompanied by an increasein the number of presynaptic
and we know that'memory involves establishing

[,

p
$

ff

Jff

C O N C L U SI ON
L OOKIN G F OR WAR D

In the preceding chapters we have had the chance to

appreciatesomeof the awesomequestionsconfronting brain
researchers.bn Chapters 1 and 2, we saw that the brain did
not work as a collection of mini-brains but that somehow
multiple brain regions contributed to different functions
(parallel processing).
Ng.one has aav idea, however. how
regiousin the brain manageto give rise to a whole,
_disparate
a function such as movement or vision, that is more than the
sum of the parts.
In Chapter 3, we sunreyed how a brain is built. Although
the neuronal nuts and bolts are now very familiar to most
ie-uroscienUsts,their mode of operationsis still continuing

to surprise. In the 1970s a rather dogmatic certainty had

crept into brain research that all functions in the brain were

derived from

l . the number of action potentials in any one neuron) and

I -inhibition (decreasingthe number of action potentials), so

the
ffi"Eggq_Igqqg4el!.Onlynow arewe truly appreciating
complexity of actions of these chemicals. The concept of
[euromodulation, of biasing the responsesof neurons, is

ll

tli
I!l

T HE HUM AN

BRAIN

L OOKIN G FOR WAR D I9 I

still being exploredin relation to a diverserengeofbioactive

suFstances. As we saw in Chapter 3, the fountain-like

l,

line are well placed to fulfil such functions. The challengeis

t
T

arrangementin the brain stem of aminesand of acetylchoto discover how the modulatory actions of these fountains,
so often the targetfor mood-modifyingdrugs,can be related
to the giobal functioning of the brain.
In Chapter 4 we saw how the synapsesfeatured in the
previous chapter were built into ever more complex circuits
that evolve in childhood to give rise to unique yet impressionable individuals. On the one hand, there are very
specific questions to be resolved, such as how a neuronl
knows when to alighihom its glial monorail to the correct,;
region of brain, and how it recognizgssimilar neurons with

does the

Although I attempted to suggestone possible scenario,

idea of fetal consciousnessbeing at the lower end of
The riddle of a physical basis for consciousnessand
mind was even more pressing when, in Chapter 5,
explored memory. Any study of memory will throw up
very large issuesin neuroscience.The first issue is how

number of cellular mechanismsare not just necessarybut

sufficientfor memory."trhisis becausememory apoearsto be,

ip paralJgl.
bottom-up mechanisms.Until the two can be woven into a
cohesive description, attempts to understand that rich
;- tapestrvof human memory will inevitablv flounder.
The secondbig issueposedby a considerationof memory

to equate it with the inner resources discussed in

1 and 5, We saw that this stockpile of memories,

a fetus be

continuum of consciousnessis far from proven.

mammaiianbrains, however,we cannot show that a certain

or motor functions, it would be readily regardedas

aspectof the mind. One way of regardingthe mind might

do ne

ve rise not iust to an individual brain but to an

consciousness?Of

behaviour,such as withdrawing a gill. In more sophisticated

iryi"d. Memory is clearly a product of the physical brain

fwihess the story of H.M.), but compared to more explicit

other hand, there

creep into the developine brain?

the operationsof biochemicalmachineryinto a remembered

perhapsthe most challengingof all: the relation of brain to

which it will team up into a particular circuitry.

At what

bottom-upapproach.In seaslugs,it is possibleto trarulate

:'

and experienceswould serve as a counterweight

the flood of everyday sensory experience. We also saw
in simpler brains, in schizophrenia,or in dreams,such
might be diminished. Viewed in this way, mind
be the personalization of the physical brain as it
and adaptsthroughoutlife. More complex brains,
l

)t
ifi

I'2

THE HUM AN BRAI N

as we saw in Chapter 4, would have more chancefor a more

F U R T H ER R EAD IN G

in$ividual, less stereotyped mind.

We seem to have returned to the same idea, that of a
-continuum, already consideredfor consciousness.If mind is
seen as the evolving personal aspect of the physical brain,
My particular
then how might it relate to consciousness?
view is that mind can only be realized when we are
conscious.After all,.arelose consciousnesswhen we sleep,
but we do not lose our mind. However,mind is meaningiess
if we are unconscious.Hence, consciousnesscould be seen
as the actual firsthand, first-person experience of a certain
mind, a personalized brain. Consciousnessbrings the mind
alive; it is the ultimate puzzle to the neuroscientist.It is your
most private place.
This ultimate puzzle, the subiective experience of consciousness,is perhaps a good place for any purely scientific
survey, namely one of objective facts, to cease.Although all
these issues may currently appear daunting, neuroscientists
have made exciting and fundamental discoveries, some of
which I have tried to review in this book. Slowly, we are
seeing the type of questionswe must ask and,having aa
of the" type of answers we should expect. We have seen

astoundingprogress,even since the 1970s,lgtJbaadJga-

Blakemore,C. 8., and S. A. Greenfield.Mindwaves:Thoughtson

Intelligence, Identity and Consciousness(Oxford: Basil Black,,vell,rggz).
Bloom, F. E., and A. Lazerson.Brain, Mind, and Beftoraor(New
York W. H. Freemanand Co., 1988).
Churchland,P. S., and T. |. Sejnowski. Tfie Computationol Brain
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