AIDS

TO

DENTAL

ANATOMY

AND

PHYSIOLOGY

by

AIDS
TO

ANATOMY

DENTAL

AND

PHYSIOLOGY
BY

S! UNDERWOOD,

ARTHUR
LATE
MEMBER

OF"~DENTAL

PROFESSOR
OF
COLLEGE
IN

BOARD

THE

OF
DENTAL

SURGERY

OF

AT
IN

EXAMINERS

SURGEONS

OF

ANATOMY

NEW

DENTAL

OF

LONDON

COLLEGE,
SURGERY,

AND

PHYSIOLOGY

HOSPITAL

THIRD

KING'S

ENGLAND',
AND

DENTAL

WILLIAM

M.R.C.S., L.D.S.

LECTURER
AT

THE

LONDON

EDITION

YORK

WOOD

"

COMPANY

MDCCCCXIV

by

ROYAL

TO

PREFACE

which

changes

The

consist

in

the

science

our

innervation

will

last

has

been

owing

relegated

to

the

The

simple

that

with

no

less

work

the

an

modern
is

on

former

this work

going through

point

was

has
and
and

of

than

ARTHUR

May,

subject

unavoidable

was

been

accessible

giving

appearing
Mr.

A.

Street,
191 4.

270967

S.

only
the

omitted, for the

handbook
all the
from

latest

the

W."Wellings,

Birmingham.

38, Harley

The

years.

Transactions

knowledge

authority

an

this

of

scene

Philosophical

excellent

the

the

and

the

two

the

on

appendix

Microscopical chapter

developments
of

an

while

paper

reason

dealing

to

fact that

the

published
press.

latest

Mummery's

last

in

advances

been

course,

edition

present

Calcification

the

during

the

important

issue.

have, of

revolutionary changes
of Mr.

in

found

of many

the

of dentine

text

be

inclusion

since

EDITION

THIRD

UNDERWOOD.

pen
of

PREFACE

following

The

the

to

hope

study

"

vexed
the

it may

days

of

not

as

are

Dental

that, containing

many

from

will,

pages

friends,

student

FIRST

TO

questions
work

constant

prove

of

long

past.

some

"

found

be

I trust,

cram-book,

but

Histology.

it does

as

EDITION

involved
interest

ARTHUR

as

in
to

which

eight
friends

S.

by

my

introduction

an

indulge

also

opinions

opinions

useful

of

own

my

have
years

in

the

upon
resulted

of

lecturing

whose

student

UNDERWOOD.

by

AIDS
I

ANATOMY

DENTAL

TO
AND

PHYSIOLOGY

CHAPTER

adapted for
triturating

alimentary canal,
lacerating, dividing, and
the

of

mechanical
This

Teeth

that

teeth

possessor
the teeth

erupt

never

animal),

and

fall back

should

whatever

forced

be

from

instances.

the

shape

upon
to

mandibular

the

of

and

include

are

the

of

the

found

placoid

for

scales

as

latter

that
in

mastication

structure

their

female

the

teeth

Then

teeth

scene

of

teeth

to

instance,

for

in

many

are

useful

incisors

the

whales,

There

true

used

always

nor

such,

"

it is true

certainly not

never

are

classifyas

to

it is

situated

all,and

at

other

many

cysts, far
we

(and

hesitate

would

chief

The

definition

definition.

the

for any
purpose
of the baleen

dugong

narwhal,
one

in

teeth, but

always

not

described

way

is scarcely

of most,
are

food.

digestion.'

of teeth

indeed

of many,
of all.

the

agents

account

being
beginning
seizing, tearing,
as

parietes of the

the

to

Owen

Professor

by

attached

substances

firm

defined

been

have

Teeth

TEETH

OF

NATURE

THE

no

logical
patho-

; and

if

definition,we
of the

shark

skin
the
the outer
on
although situated
its
with
teeth.
identical
animal, are structurally practically
animal
starts
cell, which
as
a
presently by
Every
of

tribe, which,

repeated
cells and

subdivision
the

subsequent

matrix

activity

becomes

number

These

of cells.

by their
they are embedded,
prolongation or
growth, the

in which
and

by

AIDS

TO

DENTAL

ANATOMY

of their branches, the distortion of their shapes,

shortening
and the complexity
form
of their arrangement, eventually
the very varied tissues which, in their adult condition,
have
to
scarcely appear
qualityor feature in
any
In the

common.

similar

same

way groups

of

which
cells,

are

so

almost identical,
may, after a sufficient
and development,result in creatures
as
activity
dissimilar as the elephant,the sparrow, the dog, or man.
The
great laws of inheritance,which we group together
under the name
of evolution,
these changes,and
govern
will be alluded to again later on ; at present the point
as

to appear

periodof

which I wish to lay emphasis is that every organ,

however complex its structure, is the result of the activity
of cells in a matrix.
Now, it is very apparent that in
such a collection of tissues as go to make
up an animal
for
the
of
the
convenience
some
animal,hard, and
are,
others soft ; some
others non-elastic. This quality
elastic,
of hardness
is found
infinite degree in the
in almost
tissues of any animal.
Where
ordinaryfriction is to be
resisted the tough epithelium
of the skin suffices ; where
the friction is excessive a cornified or horny thickening
is found.
Where
great weighthas to be sustained,as in
the bones,or great shocks to be resisted,
in the teeth,
as
is
and
much
harder
this special
somethingvery
required;
in some
hardness is attained by the deposition
part of
the formed
tissue of various salts of lime, varying in
amount
according to the exact
degree of hardness
which will be
known
as calcification,
required a process
dealt with in greater detail in a subsequent chapter.
This calcification is,however, so peculiar
and essential a
characteristic of the organs we call teeth that it will be as
well to spend a little time in a general consideration of
its nature
in this preliminary
chapter. As I have said,
the varied or complex structures
which go to form an
adult animal are
formed
from
neither
cells which are
and throughout the
varied nor
very
very complicated,
and
of
changes
many
extraordinary
development the
kindred originof the different organs may stillbe traced
by the observer. The very processes of cell-development
have much
in common
than a casual glance at the
more
result would suggest, and though it has been found convenient
for teachingpurposes
to mark
to
distinctions,
upon

"

by

THE

OF

NATURE

TEETH

laydown rules and

for Nature
to follow, it is well to remember, and
trouble and confusion of mind will be saved thereby,

and differentiate forms,and

classify

to

laws
much
that the various

classes and subdivisions of structures

another by imperceptible
shade off into one
gradations,
after all only descriptions,
and
and that the rules are
Nature does not considerherself bound by them.
Thus the calcified structures
"bone, cementum, dentine,
have such
and enamel
look
so different and
though they
different patterns, and are found in separate chaptersin
text-books,are after all only the result of a depositof
lime-salts among
altered cells lying in a matrix;
some
dentine is also found in the skeletons of some
fish,
in cysts in the
and excellent teeth are found sometimes
"

pelvis.
rendered
hard in varying
*
degree by a complicatedprocess called calcification
that is,an impregnationwith salts of lime. The process
will be discussed in detail later. At present all that is
is to pointout that the cells and matrix substance
necessary
in the soft,
form the various structures
and that
the tissue thus evolved is hardened
by a
subsequently
depositof salts of lime between and around the cell
elements, or in their immediate
neighbourhood. The
and macroin microscopical
result ishard tissues differing
These

dental

tissues

are

'

"

These

structural
form of cell

are
peculiarities
due
and
to variations in the
its
activity
of the soft parts, and not to varyingmodes
arrangement
of calcification. The
lime is always brought by the
blood- stream
and
deposited as small round bodies
which coalesce and render permanent
(calcospherites),

scopicalappearance.

the newly-formed soft structures.

the fused calcospherites
Sometimes
surround,but do
the cells,vessels,nerves, and connective
not
obliterate,
tissues of the part, as in bone or osteodentine.
times
Somethe cells and their branches
alone remain in the
normal cementum
calcified mass, as in human
; sometimes
in
branches
of
cells
the
human
the
dentine ;
as
only
persist,
and sometimes
only the bloodvessels, as in pure vasodentine ; while in the hardest tissue of all,
enamel, whose
be
said
be
to
development can scarcely
quite as well
understood
the rest,nothingat all appears to remain
as

by

AIDS

TO

DENTAL

ANATOMY

In all probability
is
the process
little water.
and
the varieties are
similar in all cases,
simply the
varied
result of the
arrangement of the tissues previous
o
f
lime.
the
the
to
deposition
Just as in their firstoriginall the various creatures
and all the different parts of them
similar,so the
appear
of the special
earlier stages of development,whether
show much
animal or the specialorgan,
blance
resemmore
than their perfectstate.
The
of
infancy a hair
and a tooth present great pointsof resemblance, and the
are
nearlyindistinguishable
earlystages of the ape and man
differences
arise
in
each
when the organ
case
; the
functional life,
when the habits
or the individual reaches
by which the creature maintains itsexistence or the organ
dischargesits functions become necessities.
Teeth, then, vary in their situation far beyond the
and in their developin the definition,
limits laid down
mental
affinities
or
they display
relationships
process
other parts of the organism. Let us now
with many
how far they are, as the definition
consider for a moment
to infer,
seems
employed as mechanical aids to digestion.
doubt this is the main purpose
which they
Without
a
shall find that they also serve
other
serve, but we
many
but

such

purposes,

Speech.
"

in

the

as

speech,combat, etc.

teeth
function of

In

man

ance
great mechanical assistin
fact,
speech"
speech could
are

without them.
I say
properlybecause edentulous peoplecan make themselves
the speech of man
be to a
understood; moreover
can
birds
mimicked
certain
certain extent
correct
by
; still,
without teeth.
and exact speech is impossible
Combat."
The hugelydevelopedincisor of the narwhal
has no
reference at all to the subdivision of food, nor
the tusks of the
have the rostral teeth of the sawfish,
nor
wild boar.
Many similar developments of teeth exist
their
throughout the animal world which only serve
of offence and defence in the great
possessors as weapons
existence.
for
Moreover, we recognize
generalstruggle
dental developmentstwo great varieties
in these fighting
that are
used for killing
the prey : these are
weapons
found in both sexes
(such as the huge canines of the
that are used in combat
between
felidae)
; and weapons

scarcelybe

properlyperformed

"

by

NATURE

THE

TEETH

OF

for the possession

of the female (as the tusk of the
narwhal),which are not developed until puberty,and in

males

the female animal

are

never

eruptedat

all.

Sexual

Weapons." The teeth that are highlydeveloped

in
puberty only in the male animals serve their owners

at

breedin the ing

life.
feature of animal
are
a common
for
Sometimes
the male animal developsother weapons
cumstances
this purpose, such as the horns of deer,under which cirthe
sexual
of the teeth is
specialization
If the animal is castrated in infancy,
absent.
generally
these sexual weapons
arrested in their development.
are
The narwhal's and the dugong'sincisors and the canines
of the wild boar are instances of these sexual weapons.
The
Locomotion.
huge incisors of the walrus are
for
both
self-defence and
for assistingits
employed
the ice,while the lower incisors of an
movements
over
extinct monster, the deinotherium,which, though springing
for
those combats
which
season

of
possession

the females

"

from

the lower jaw, turned down

direction as the walrus's tusks, were

employed, and possiblyserved

animal in its watery abode
to

much

in the

same

probablysimilarly
the unwieldy
fixed point during

to anchor
some

of rest.

moments

Other Purposes. The

elephantuses his huge incisors

for
carrying things; the beaver employs his
largely
teeth ,in his buildingoperations
other creatures
; many
dental developments in diggingfor roots and
use
special
climbingtrees" in fact,the teeth,like every other part ot
the animal, serve
handy.
any purpose for which they seem
The
rule
that
the
rule
of
is
the
only
governs
process
If an
usefulness.
animal's teeth or horns or claws help
it to do anything more
or
more
effectually
quicklythan
its neighbours,
it is likely
to live and prosper and rear
up
"

and
offspring,

housed,
form

in

or

then
those peculiarities
that protected,
fed the parent will be found in an exaggerated

subsequentgenerations.

Teeth, then, are

difficultto define,
because they shade
insensible
which
gradationsinto allied structures
by
teeth. The
teeth of the shark are
not
are
like
in
so
structure
to its dermal
would
useless
that
it
seem
spines
and foolish to attempt to draw
the line of a definition
between
the
But
them.
when
to consider
come
we
off

by

AIDS

TO

DENTAL

ANATOMY

structures, and study their history

find the resemblance
than
is more
of
likeness
form
and structure, and includes a
a
mere
of
and
of
formation, so that there is
similarity origin
time
and at the same
hardly any characteristic essential,
to, teeth.
peculiar
Let the alimentary
canal be considered as a passage
the
the
animal,
through
liningof which is continuous at
each opening with the coveringof the outside,so that
there is a complete and
uninterruptedinvestment of

homologies of these
and development,we

servingas
epithelium,
outside of the

body, and

and coveringfor the

protection
continuingthroughthe passage
mouth
and ending at the anus,

which, beginningat the

the body, and is called the alimentarycanal.
perforates
This complete and continuous coating consists of epi
thelial cells in

and
varying in disposition
juxtaposition,

according to the requirementsof their

situation. In the alimentary
canal it is called
mucous
membrane
Under
'; on the skin it is called 'epidermis.'
neath this epithelial
coatinglies a co-extensive layer of
of connective-tissue vessels
subepithelial
tissue,
consisting

arrangement

and nerves,
and
this is called in the alimentary canal
'submucous
tissue,'and in the skin 'dermis* or true
skin. Between
the two there is supposed to be a thin,
this
transparent and homogeneous basement membrane;
is not everywhere capable of demonstration, but from its
in certain situations,
and notably in the foetus,
presence
it has been assumed
to co-exist with the epithelial
layer.
At one
of formative cells
stage in development the mass
into three layers,
each of which has its
range themselves
specialtract to form, and it is an importantfact that
whereas
the epiblast
or
outer
layer forms the epidermis,
the mesoblast
middle layer forms the dermis ; these
or
therefore very distinct in their origin.
two
structures
are
The
is liable to variation in the shape
coat
epithelial
and arrangement
of its component
cells ; thus, where
there is much
friction there are
layers of cells,
many
Where
in the sole of the foot or palm of the hand.
as
an
interchangeof gases takes placebetween the air on
the one
side and the bloodvessels on the other side of

epithelial
layer,the cells are singleand as flat and
thin as possible,
vide the air-cells of the lungs. Some-

the

by

THE
times the

NATURE

OF

special
arrangements

TEETH

cells are
epithelial
underlying subepithelial

of the

the
considerable,and
very
tissue may take
in
the
changes. This is the case
part
where the necessities of warmth
or
requirethe
protection
motion
formation of hairs,
where
or locoor
purposes of combat
render nails,claws or horns an
advantage, or,
where prehensionor subdivision of food requires
lastly,
of teeth.
organs of the nature
Whether
it be the scaly armour
of the armadillo,the
horn of the rhinoceros,
the tusk of the elephantor the
narwhal, or the hair and teeth of the human
being,one
and all are due to a special
arrangement or specialization
of the epithelial
and, in many cases, of the subepithelial
quired
coatings.When any such specialdevelopments are reto be very hard,the result is attained (as hardness
usuallyis in livingbeings)by the depositof a certain
proportionof salts of lime in the structure formed by the
is slight
in
of lime deposited
specialcells. The amount
cases, and the result is a horny substance, which
many
of
has an albuminous
the amount
basis ; in other cases
lime is considerable (from 60 to 97 per cent.),
the result
basis.
being a calcified tissue which has a gelatigenous
*
or
Among the firstclass rank false,'
horny teeth ; and
the
second, 'true/or calcified teeth. Thus it is
among
that teeth,in common
with hairs,nails,and other dermal
appendages, are specialdevelopmentsof the external
coveringlayerof the animal.
the comfort or safetyof an animal requires
Whenever
natural implement or weapon, the complicated
a special
evolution
influences that are included in the term
slowly
but surelytend to produce somethingof the kind required.
less of the
or
Among a generationof animals, more
'

same

kind, some

few

are

sure

to

better providedin
rest; these few, by
their rivals in the

be

slighta degree than the

reason
advantage,overcome
two
great departmentsof the strugglefor existence
pursuitof food and the pursuitof the female. As a
the next
generationinherit only from
These
fittest*of the generationpreceding them.
ever

so

of their

"

the
sequence,
con-

the
in

'
their turn undergo the same
process of natural selection,'
that each succeedinggenerationpossesses in a more
so
marked degreethan any precedingit those characteristic^

by

which

AIDS

TO

DENTAL

ANATOMY

the most suitable

favourable for procuring
the female animals,and for destroying
captivating
rivals in combat.
Thus
countless generationshave
evolved
in the descendants
of the anoplotherium,
with
his regularand even
of teeth, the tusks of the wild
row
most

are

food, for

boar, and

the peculiar
of sus babyrussa; thus
armature
the song of birds and the beautiful variety
of colour and
which
in all the animal kingdom is the
ornament
we
see
result of this process of selection and survival of the fittest.
At the same
thus becoming
time that usefulpeculiarities
are
enhanced, useless ones are undergoing
inevitably
a
retrogradechange, so that such organs in successive
generationsare found to dwindle until sometimes all that
remains is an abortive relic (this
supposed
process is now
of the human
wisdom
to be causingthe disappearance
Some
such
relic
lateral incisor).
of
tooth and upper
found
form
be
in
a lost organ
some
or
generally
may
will cause
freak of nature
a
other,and not infrequently
the growth and development of an organ that can
only
*
*
be regarded as a reminiscence
hark
back
to some
or
in
the
the
anatomical
individual
o
f
previousstage
history
period; and as on the one hand we can generallyfind
in the progress
trace of that which has disappeared
some
of evolution,so on
the other hand
we
generally
may
the germ
discover in earlier generations
or
originof that
which
has been
exaggerated. Moreover, during the
life
of
each
individual there are
numerous
embryonic
hints or suggestionsof previousstages in the existence
of the race; it would
almost seem, in fact,as if each
time that Nature producedan individual she prefacedits
of the importantpoints
birth and life by a brief summary
in the history
this particular
of the evolution by which
animal
be examined
type had been produced. If an
will
be
found
retain many
after
it
to
birth,
immediately
from
to creatures
a stage removed
pointsof resemblance
which resemblances
it in the historyof evolution,
will
into
full
decrease and fade as maturitybrings
ment
developcharacteristics of the animal,and at this
the special
there are many
pointsof difference between
infants of allied genera that have not become
prominent.
It is in infancy that the greater kinships if I may
so
of living
most
call them
creatures
easilyperceived,
are

early age

"

"

by

AIDS

io

TO

DENTAL

ANATOMY

the
easily. In this way there will be among
that are better suppliedwith
hundred, say, twenty-five
and
best supplied
from the trees.
ground food,
twenty-five

more

There

will also be

of both kinds that will go

twenty-five
to the wall in the strugglefor existence.
The
result is
the survival of those who are best fittedfor that struggle.

This

combat
results in those of each generation
who are
fittestfor their environment surviving
and therefore transmitting
the
this peculiarity
in an ento
next generation
larged
If
have
number
of
less
or
a
more
degree.
you
arboreal animals who can climb more
and a large
easily,
number
adapted by thickeningof integument to get at
in
the next generation
the same
roots,
dispute,the same
mit
struggleis takingplace,and those only will live to transwho are best providedfor the struggle.I
offspring
do not mean
that no imperfect
but the
will survive,
ones
general tendencywill be for the best to improve generation
after generation. Now, the cells that thickened the
upon another ;
examined
cells,
tions
generamany
will assume
to the casual
a definite structure
after,
observer special
of these animals
and peculiar. Some
have
had
o
f
lime
in this part of his economy
a deposit
may
hard. This would give the animal
renderingit specially
immense
an
advantage,and we should find that the
presence of the lime would in time become the rule instead

nose

were

simplyepidermiscells lyingone

the descendants

of the
The

of these

exception.

animals have
horns of the various horn-bearing
acquired this specialdevelopmentin obedience to these
will always arise to meet
laws, the peculiarstructure
peculiar
surroundings.The structures themselves come
that you will
after countless generations
to differ so much
had
believe they may have
ancestors.
not easily
common
tion
variaThe
which
known
concomitant
is
as
phenomenon
has resulted in the fact that it is unusual to find one
animal doubly armed
i.e.,possessingboth horns and
The musk
deer and an extinct
canine
teeth.
specialized
animal called Deinoceros are exceptions
to the rule, the
latter havingpossessedenormous
sword-like canines and
"

three

pairsof horn cores

Odontology,'
p. 105).
This economy

on

(seeUnderwood,

'

Comparative

the part of Nature isfurther illustrated

by

THE
in the

NATURE

trophicfunctions

OF
of vessels

TEETH
or

fibrilsin dentine

(seep. 66).
It is

this constant
inheritance and this incessant
strugglefor existence that peculiarforms arise,and as
an
they arise we shall note that whatever peculiarity
animal possesses, however extraordinary
it may be, it is
but
not
that
t
o
thing
animal,
somethingpeculiar
simplysomewhich every other animal has,more
or less developed
in a special
direction ; not a new
thing,but something

by

to all creatures

find

incisor
incisor of the narwhal.

the

common
our

own

specially
developed. So we
is homologically
the same
thing as

Let us now
conditions in our
suppose the geological
island to undergo a change. Of course, such changes
would take countless generations
and countless ages, but
and affect the island in such
suppose theydo take place,
that it is no longer an advantage to the animals
a way
it to retain the peculiarity
of teeth,or hair,or
inhabiting
limbs that was
essential to existence ; then this
once
no
peculiarity,
acquiredby a long process of selection,
longer serving the animal in helping it to obtain
food more
quicklythan its rivals,
possiblybecoming a
under
will dwindle,
hindrance,
changed circumstances,
leavingnothingat last but a feeble and abortive reminder
of a long-forgotten
Thus
time.
geologicalchanges
then
and
further changes efface
produce peculiarities,
But as they are
them.
producedby the development of
so
somethingwhich alreadyexists,
quite
they are never
effaced. We
which has not its
rarelyfind a peculiarity
in
analogue other creatures ; on the other hand, a special
but leaves an
development never
entirely
disappears,
abortive trace, though it no
its old
longer answers

In this way there are teeth which never

come
purpose.
baleen
The
to the surface and
used.
never
are
great

whales obtain their nourishment

of
by an arrangement
whalebone
plates; underneath this is a set of teeth which
are
never
eruptedand have disappearedunder changed
circumstances and environment.
So it is that in this struggle
we
constantlyfind races of
animals acquiringgreat peculiarities
and then losing
It is also true that very strange surroundingsmay
them.
and, put under a new
requirevery strange developments,

by

ANATOMY

DENTAL

TO

AIDS

12

order of

animal passes away

the very specialized
things,
and becomes
extinct ; he is outrun
by other
altogether
animals,and has no time to adapt himself. The extinct
able
of the older geological
creatures
periodsafford innumerinstances of

specialized
types to
races
are
persistent
an

of the most
the
whereas
disappearaltogether,

general tendency

the

those which

do

not

vary to

so

great

extent.

be always borne in mind that all these changes

It must
of incalculable slowness ; that they affect every part
are
of the creature
and every attribute it possesses. The
of their plumage is as
song of birds and the brilliancy
much
the result of evolution as the tusk of the elephant
the colour of
of the rhinoceros.
the horn
Thus
or
creatures

becomes
frequently
to serve

assimilated

to

their

roundings
sur-

of the
: in the case
a twofold purpose
it enables them to conceal themselves

feebler creatures
from their pursuers, while the predatoryraces
are
by a
enabled
similar stratagem, if we
the
unuse
word,
may
ithas
their
Hence
t
o
happened
perceived approach
prey.
that gradually
have come
to resemble
the living
creatures
perceptible
their inanimate surroundingsso as to be almost imThus
the
objectsin their native haunts.
birds and beasts in polarregionsare white,the colour of
countries are
the snow, and some
that migrate to snow
The
white in winter and other colours in summer.
lion,
colour
and
sand
of
the
are
;
antelopefamily
puma,
many
in which they live ; the
birds resemble the foliage
tropical
whose jungleis mostlyreedy,is striped
; while the
tiger,
the leopard,
the great cats
tree- climbers among
panther,
shadows
the
cast
etc.
simulate
to
are
by
spotted
ounce,
The
their home.
the leaves of the trees which
form
forests where littlelight
fauna found in the impenetrable
curious exOne
ception
penetrates are mostly neutral-tinted.
to emphasize the rule : the carrion crow
seems
but he never
travels annuallynorth into snowy
climates,
for
changes his coat because there is never
any reason
"

"

such

change

attack,and

; his

his

natural

food

secures
ferocity

being always

dead

him
from
need not be

stealth.

approached by
Among all the
of

natural

giftswhich

few individualsover

affect the preeminence

most

their contemporaries.

by

THE
the

NATURE

OF

TEETH

13

first. They are essentials in all

obtainingand consuming of food, and
they are the most formidable weapons in the armoury of
Nature.
It is not, therefore,surprising
that, having
the
that
law
accepted
surroundingcircumstances in time
in the form of animals,
producedeviations and specialities
in the teeth.
So
find these variations most
we
marked
great are the varieties that time has produced,and so
teeth

to
relating

must

rank

the

thoroughlyhas

evolution providedappropriate
apparatus
for the various families of the animal kingdom, that it is
to
comparatively
easy, from a glance at an animal's teeth,
decide what were
its habits,its generalcharacteristics,
and its surroundingcircumstances.
The
teeth of the
animal are a kind of imperishable
diaryor autobiography
;
harder and less subjectto change than any other
of
the animal, and
often represent all that remains
part
sufficient
of extinct races, but they are
to us
evidence of what their possessors were
like to enable
anatomists to restore with tolerable accuracy the rest of
the animal's anatomy and to form a close guess at its
habits and mode of life. Not only are teeth imperishable,
but in many
the animal produces and sheds an
cases

they are

infinitenumber

these relics are

also very numerous.
the teeth afford evidences of
Thus
the characters and habits of the animals that possessed
them that are at once reliable,
under ordinary
imperishable
be
and
to
circumstances, accessible,
numerous,
easy
examined.
combined
All these circumstances
have
to
with
render odontologya very useful branch of zoology,
the natural result that the Kterature of the subjectis
so that
duringits lifetime,

and that comparative

as well as human
considerable,
odontologyforms an essential element in the education of
the dental student. Seeing, however, that it is with
very

and in
human
teeth that the student has to do both now
the future,it is chiefly
that this
with human
anatomy
of the human
book will concern
itself. The description
the
involve
will
continual
to
reference
comparative
organ
for purposes
but the human
of analogy and illustration,
I shall
tooth will be the principal
business of the book.
of
t
he
endeavour
to follow as nearlyas possible
course
firstof the development of teeth,and
Nature, treating
when
afterwards of their form and structure
mature,

by

AIDS

14
and

TO

ANATOMY

DENTAL

finallydiscussingthe

attendant

changes

upon

senility.
A leisure hour

or

two

spentin

the

study of Wallace's

'

will well repay

delightfulDarwinism
odontologyat this stage of his studies.

In the last

student

of

II

CHAPTER

DEVELOPMENT

the

OF

THE

TEETH

of the
glance at some
slight
moulding and modifyingthe
Producing continents and
then effacing
them
with very strange and
them, peopling
them
specialized
livingcreatures and then obliterating
with relentless completeness,the great laws proceed on
their way, and we, who can only observe duringthe brief
of a human
moment
life,have to build up our history
from geological
remains, or piecetogetherfrom a few

took a
to be
great laws which seem
world
and
its inhabitants.

chapterwe

scattered remnants
the almost endless succession of slow
have
world as we see
that
changes
produced the existing
it,and are on their leisurely
way to produce something
future. To have the faintest
very different in the remote
the
of
of
this vast system of evolution
nature
conception
it is essential to appreciate,
far as we can, its extreme
as

realize that it is stillproceeding

ing
and mouldeverythinganimate and inanimate justas much as in
plioceneor miocene ages. Geologicalrecords enable
in imaginationmany
of the weird and
to restore
us
that have played their part in this
monstrous
creatures
world's
ceaseless
long
pageant and passedaway, but we
have another curious source
of knowledge in this connection,
which is of specialvalue as an introduction to
this chapteron development.
exhaustive
scientific treatises it is
Just as in many
customary at the end of each chapterto summarize the
facts therein worked out, and present to the reader's mind
of what has gone before,
or brief recapitulation
a synopsis
slowness

; to

by

OF

DEVELOPMENT
so

does

Nature, whenever

an

THE

TEETH

individual is

15

marize,
formed, sum-

and run over the heads of the stages

recapitulate,
of the evolution of that individual duringits intra-uterine
life. A human
being, at first a cell,graduallyevolves
to gillitself,
passing through stages of resemblance
(the branchial arches) and showing
breathingcreatures
and lower mammalian
affinitiesfor reptilian
forms, and
finally
justbefore birth displaying
pointsof strong kinship
with the simian race.
This re'sume* that accompanies
existence of an individual is equallynotable
the praenatal

in the formative stages of each organ.

The transitions of
the evolution of a tooth from a littlethickened epithelium
in brief in the developupwards are shown most perfectly
menal
stages of each tooth that is formed, and, therefore,
consider
and watch
the changes that result in the
we
as
human
tooth we must
that we are watchingan
remember
epitome of the evolution through countless ages of the
tooth itself. We
shall find it presentingpointsof great
lower forms at various stages of its
much
to
similarity
growth,and while this reflection will enable us to take a
broader view of,and a more
sensible interest in,dental
apparently
anatomy, so will itassist us to remember
many
trivial details,
historical
view
value
because in
of their
they become of large importancein proportionas they
of wide application.
are
This
general kinship between all livingcreatures,
dependent upon a common
ancestry in the remote
past,
will regard as beinggenerallyaccepted,
shall
and we
we
in the
expect to find many hints and traces of this affinity
earlier stages of the development of each individual.
It is therefore possible
to consider,
to a
a preliminary
as
of tooth development in man,
detailed description
more
that may be regardedas applicable
to
some
generalities
tooth developmentin general.
of teeth are formed
The germs
little
firstof all some
way below the surface, and afterwards,as their formation
proceeds,graduallyapproach the situation where in their
useful. This move
completed state they will become
of action is called
ment
or act of travelling
to the scene
about, and what the mechanism
eruption. How it comes
is which underlies the movement,
understood
is scarcely
;
but itvaries very much in amount, beingalmost insignifi-

by

AIDS

16

TO

while in

DENTAL

ANATOMY

sharks the distance traversed

is many
times the length of the tooth that performs
the changes that result in
the journey,but in all cases
of the epithelial
tooth begin by a dipping downwards
a
into the subepithelial
membrane
coveringof the mucous
cant

in man,

some

tissues.
The

lower

description
following
may
jaw, so

that there may

the
meaning^of

words

be

be taken to
no

applyto

confusion

about

the
the

upwards,' downwards,' etc.

*

in the formation
Thejrrst changes that result eventually
which covers
the
placein the epithelium,
alveolar margin of the jaw. It is importantto note
that
affect
tfpto a certain pointthese changes do not specially
'certain pointsin the jaw (where teeth are to be formed),
but are continuous along the whole margin ; they are not
series of changed dots, but an uninterrupted
line of
a
of autooth take

which forms the mucous

brane
memchange. The epithelium
clothingthe jaw is,like all other thick epithelia,
capableof a rough division into two layers; that is,the
deeper layer consists of elongatedor columnar cells,
In the
the surface the cells are flattened out.
tained,
mainare
changes about to be noticed these peculiarities
the deeper cells *."?., those next
the basement
their
columnar
nature
membrane
preserving
throughout
functions in the formation of the
and performingspecial
confusion has been engendered in the minds
Some
germ.
of students concerningthese earlychanges,owing to the
in early textof loose and inadequateexpressions
books.
use
'
dental groove
For instance,the expression
has
because the thingso called
givenrise to much obscurity,
when
is full of tissue,and only becomes
the
a
groove
contained tissue is destroyedor removed
by reagents.
The greatest care will therefore be given to employ words
and similes in a very exact sense.
About the end of the sixth week of intra-uterine lifean
cells all along the
of the epithelial
active proliferation
margin of the embryonic jaw takes place, and not
raised by heaping
only does the crest of the ridgebecome
but
sinks
into
the
the
latter
of
epithelial
subepithelium,
up
tissue in such a manner
that,if it were liftedoff
margin would be found to be
bodily,the subepithelial
impressedand to present a shallow groove all along its

whereas

on

"

"

'

by

AIDS

18

TO

DENTAL

ANATOMY

observers have described,in

buds given
that only developtwo sets of teeth,
creatures
off from the lamina before and in front of those that form
for which the name
of pre-milk'
has been
the milk teeth,
suggested,and others that succeed the permanent germs,
Such
and
have been
called * post-permanent.'
germs

that have

two

Other

sets.

very

seldom

arrive

supplementalor

at

maturity,unless, indeed,

supernumerary

teeth of

the

pathology are

of their successful
occasional instances
regarded
of the
growth. Dr. Marrett Timms in 1896(Transactions
and
OdontologicalSociety)discussed the probabilities
and
evidence
concerning a pre-milk
post-permanent
dentition in man, and Dr. Gibbs of Edinburgh, in the
International Medical Congress, 19 13, described a case
without
to my
a
knowledge, in which four
parallel,
successive
first permanent
temporary dentitions and
molars coexisted in the rightmandible
of a child.
The zahnleiste is a cellular structure, and it is not surprising
of cells attached to it,neither are
to find bundles
as

in calling
unless we
a thinga tooth germ
quitejustified
that
it
show
teristics
dental characcan
developsmore
definitely
than have yet been shown to be possessedby
the 'pre-milk'
or
buds; but if these
post-permanent
shown
be
in
could be
abortive tooth
structures
to
reality
from
the
band there
common
springing
epithelial
germs
in it in fact,it would only
would be nothingsurprising
tend to show that the difference in the mode
of development
in mammalia
and those creatures
possessingan
we

'

'

"

endless succession of teeth was

more
apparent than real,
like
differences in nature, a difference in degree
most
and,
rather than in kind.

According to
the time

of

authority(Dursy),there occurs,
the formation of this lamina,a change in
one

at

the

tissue. He describes a line of opacity(not

subepithelial
series of dots correspondingto the future teeth in
a
number) due to an increase in the cell elements,at the
epithelial
expense of the fibrillarelements, in that part of the subtissue which is directly
underneath the edge of
So far, then, we
the lamina.
have a thick band
of
from
epithelium,
and
and

the

deepestpart

of which

band

springsa deep

continuous all round the jaw,

(lamina),
dippinginto the subepithelial
tissue,
a
and,'possibly,
narrow

by

DEVELOPMENT

OF

THE

TEETH

19

beneath the edge of the lamina.

All this
line of opacity
of teeth ; but as yet there
is to end in a certain number
how
where
of
the teeth are to be.
or
is no sign
many,
The next change givesthis indication. From
the deepest
a series
part of the lamina (whichI have called its edge),
of
invasions
the
tissue takes
of finger
subepithelial
shaped
those
in
of
the
while
towards
subepithelial
tissue,
spots
place,
which these fingerspoint,ceil development takes place
cellular masses
and opaque
are
formed,as ifto
quickly,
encroachment
further
of
the
the
obstruct
epithelium.The
to
however,
continues,
encroach,and, instead
epithelium
the littlecellular islands,
of displacing
envelopsthem, the
in
much
each
the
result being,
same
indiaas if an
case,
forced down on to the finger-point.
rubber ball were
The
structure
at this stage has been aptlycompared
epithelial
of the under-part
of the
flask,the concavity
to a Florence
flask beingoccupiedby the subepithelial
cellular mass.
be remembered
that the flask* is full of
it must
But
which lines itis of a
and that the epithelium
epithelium,
cellular
The
little
kind.
which is thus
columnar
mass,
tissue everyisolated from its surroundingsubepithelial
where
but at its base, is called the dentine germ^
and
to'
the
of
it by its
sometimes, because
shape given
*
and
dentine
its
t
he
substance
is
papilla/
isolation,
series
of
transformed
into
a
dentine
by
changes
gradually
invasion that has descended
and pulptissue. The epithelial
and envelopedit on all sides is called the enamel organ,
and in a portionof it namely, that which
immediately
the enamel
overlies the dental germ
coatingof the tooth
is formed ; the two are supposed by some
to be separated
another by a basement
or
from one
membrane.
limiting
So far,then,we have a series of cellular spots, which have
tissue,about to form dentine
arisen in the subepithelial
of
series
and
and pulp,
a
envelopingcaps of epithelium
them from their surroundings
coveringthem and isolating
mechanical
exceptat their bases. The one has the same
has
ball
the
to
seat of a flask
relation to the other that a
which
it
is
Both
under
bottle
imprisoned.
structures
or
of
cells
but
the
composed
are
;
chiefly
marginal cells
of each, those which are next the basement
membrane,
in
form.
columnar
The
or
dentine
is
are
elongated
germ
cells
the
the
but
on
outside
of
of
which
it,
cells,
a mass
*

"

"

by

AIDS

20

TO

DENTAL

ANATOMY

the basement
lie justunderneath
membrane, are rather
differas to
specialin form ; in or around them (opinions
is
substance
dentine
known
formed
and
as
which) the
afterwards calcified. The cells immediatelyunderlying
them and the
them are intermediate in character between
simplercentral cells,and perhaps reinforce them when
The enamel organ isalso a mass
their energies
are spent.
in fact,
but for the basement
nearest
to
of cells. Those
membrane,
contiguous with the specialcells of the
also columnar, large,
and specialized
dentine germ
are
;
is formed, and these,too,
in or around these enamel
out
are
by the cells
probably reinforced when worn
The
immediately behind them.
secondary ceils may
"

"

reinforce the

primary ones,

but

it is

not

certain that

they do.
of the
enamel organ is therefore a development
which
originally
dipped
pointof the fingerof epithelium
in from the lamina.
The
next
change is that the opacity,due to cell
the dentine germ
which has so far distinguished
activity,
in
which
it
tissue
from the subepithelial
arose, beginsto
spreadfrom the edges of the base of the germ, and, passing
the deepestedge
outward in ail directions underneath
of the enamel organ, curls up outside it and envelopsit,
surface of the
lost upon the outer
until its cells become
confused
with the outer
enamel organ, and inextricably
the cementum
is
In this structure
cells of that structure.
and
in
whose
teeth are
animals,
some
eventuallyformed,
with
investment
of
the
a
cement,
complete
provided
formation of this tissue is preceded by a formation of
of cement
and the name
cartilage,
organ has been applied
The
this
dentine
it
at
to
by Magitot.
stage is represented
the pulp
outer
cells,
by a dentine germ with specialized
by
by the middle part or core of the germ, the cementum
the
dentine
from
the cement
germ,
organ or expansion
of
the
cells
the
internal
enamel
the enamel
by
organ
the
overlie
dentine
which
those
papilla.
immediately
*".,
used
be
to
regarded as thin
Nasmyth's membrane
thin
of the cement
the
formed
prolongation
by
cementum,
the external
which loses itself (in man) upon
organ,
epitheliumof the enamel organ. The view of Waldeyer
of the external cells of the
that it is due to the persistence
The

"

by

DEVELOPMENT

TEETH

THE

OF

21

enamel
is now
more
generallyaccepted; perhaps
organ
it is formed in an union of the two.
The original
from the end of which
fingerof epithelium,
the enamel
become
has
now
long
developed,
organ was
and thin ; it is no longer straight,
and it is dotted over
at

intervals with minute

neck

buds.
epithelial

It is called

the

of the enamel

One of the buds is eventually

organ.
enamel
a new
sponding
organ, that of the correthe
tooth.
Whether
permanent
permanent
is
derived
from
neck
of
the
enamel
the
germ
organ
be regarded as certain. I do not think Nature
cannot
binds herself by rules; the permanent germ is an offshoot
of the epithelial
ingrowth,and is probablygivenoff from
the most
convenient point. It is possible
that sometimes

developed into

than

more

rise

one

bud

the

give

develop to maturity,and

may
supernumerary

and supplementalteeth which

such
feature in dental pathology.
an
interesting
M. Malassez
has described these buds with great care
and
minuteness.
It will now
be necessary
to consider
these parts of the tooth germ
in detail.
more
to

form

The

Enamel

peculiarshape

Organ,
which

as

soon

I have

indiarubber

resembling an
pressed downwards
such

as

on

to

it has
endeavoured

ball

describe,

to

its base

syringe with

body,
resisting
in
development
special
that is,
differentiation
smaller

some

finger-tip,
undergoes

acquired the

as

its various parts, which is called

the cells,which were
of a muchness, except
at firstmuch
that the peripheral
ones
were
more
columnar, develop
distinct peculiarities,
increase and render them
which
"

of

Before
being easilyseparated int,ogroups.
word
more
discussingthe several parts in detail,one
be said about the shape of the organ itself" namely,
must

capable

that,

seen

in transverse

section,it presents

oval

an

or

rather ovoid

organ

while the section of the dentine

appearance,
stretches across
the widest diameter,the internal

epitheliumapproachingmuch
consequentlyleaving much

and
the external,
for
room
intervening
tissue at the ends of the long axis of the oval.
This point
has not, I believe,
been emphasizedbefore. The limiting
cells those,that is to say, which
form that part of the
nearer

less

"

the
representing
columnar.
and
elongated
organ

indiarubber
The

outer

in the
ones

are

syringe
"

are

continuous

by

AIDS

TO

with the inner

ones,

22

they

DENTAL
which

ANATOMY
the

cap

dentine

distinguished by the specialname

are

but
germ,
'
of external

'

epithelium; while the inner ones, which are called the

much
internal epithelium,'
become
longer,with
very
largernuclei,in those situations where enamel is to be
*

formed,

that it is easy, from

so

glance

at

the enamel
of enamel

of any animal, to predictthe extent

covering that the tooth will eventually possess. The
is enclosed
which
by this continuous layer of
space
internal merging into external epitheliumis filled by

organ

cells not

contiguousto

one

another,but

more

or

less

in

albuminous
an
jellyof a similar
in
humour
vitreous
of the eye
the
nature
nected
(gelatinousconnective tissue). All these cells are con-

sparselyscattered

to that found

with
of

name

their

stellate

neighbours by long branches.

reticulum' was
given to this

The
tissue

the action of the chemicals used in the preparation

of sections caused a shrinkageof the cell envelopes
their connecting branches, and
until recently all

because
and

in this tissue. In
sections showed a stellate appearance
the International Medical Congress,'
191 3, Stomatological
Section,Mr. Wellingsand myself showed that in sections
treated by heat, alcohol,or acids, the processes are
not
and the cells have
stellate outline whatever.
no
wavy,
*
Between
the widelyseparated stellate cells and the long
internal epithelial
cells there lies a narrow
layer where
cells
and
roundish
the
huddled togetherso
are
or
oval,
but slightinterspaces.
The
to show
as
are
interspaces
lie close to the internal
less between
the cells which
enamel
cells. The
area
occupied by these latter cells
has therefore been called the stratum
intermedium^while
the apparentlystellate tissue beyond has been called the
stellate reticulum.
The
real difference between
these
and
cells is very slight,
their kinship is shown
by the
fact that each layer appears
to reinforce the neighbouring
of action ; thus the internal
to the scene
layer nearer
enamel
the stratum
from
recruit
to
intermedium,
appears
and this,in turn, from
In the
the stellate reticulum.
enamel
first formed, the cells are all in conas
tiguity
organ
; possibly
they are all interconnected by branches,
but these have not been
shown because there are hardly
In
the
regionof the stellate reticulum,
any interspaces.
'

by

DEVELOPMENT

OF

TEETH

THE

23

considerable
matrix arises
formation
of jelly-like
between the cells. The function of the stellatereticulum
and stratum
intermedium
has been regardedas obscure
because
both
where
animals
absent
in some
(1)
are
is formed ; (2)in mammalia
enamel
they disappearbefore
that
the enamel
formation is complete. The suggestion
the reticulum is merely an area
of reserve
to reinforce the
internal epithelium
both
to me
to meet
seems
objections,
in
the
first
be
it
small
amount
taken
that
a
as,
case,
may
and in
of enamel can
be formed without reinforcements,
that
the second,
towards the end of enamel formation the
reinforcements have been drawn upon and exhausted.
cells of the internal epithelium (enamel cells,
The
adamantoblasts, ameloblasts)are about twelve times as

long

as

they

are

broad, and
they are

againsteach other ;
shortest at the neck

closely packed

very

longestat

of the

crown

of

the apex, ^nd

In
the tooth.

are
granular,like the prisms which they
intermedium
also the nuclei of the stratum

structure

they

form,

are

as

are

stellatereticulum.
Waldeyer thinks the ends of these
enamel
are
open, and that the enamel

and

cells facing the

is shed out from

them.

Magitot thinks there is a plateau'or thin layer of

something (he does not say exactlywhat) through which
akin to
the future prism exudes by a process apparently
*

exosmosis.
The nuclei of these cells are largeand well defined,
and
occupy the end of the cell furthest from the seat of enamel
formation.
been
external epithelium,which, as
has
The
said,
consists of the same
of cells,only shorter and less
sort
definite in shape,is chieflyinteresting
of the
account
on
theories regardingits destinyor function.
as
Waldeyer thinks it persists
Nasmyth's membrane,
of a number
which tissue he regards as consisting
of
which
flattened cells,the cementing substance between
of
with
silver.
nitrate
be demonstrated
by staining
may
in the right
is morphologically
The
external epithelium
place to fulfilsuch a function.
researches
Mr.
recent
Wellings and myself,in our
Medical
International
Congress, iqj3)"
(Transactions,

by

AIDS

24

TO

DENTAL

ANATOMY

which
of the functions of these cells,
explanations
will be presentedin the chapteron Calcification.
for
made
The
elder Tomes
out
a very
strong case
thin cementum,
was
supposingthat Nasmyth's membrane
appeared.
disthat these cells entirely
and
therefore concluded
his
he
this
followed
In
was
by
opinion
son,
Charles Tomes, and a majorityof writers. Since the

offered

Paul's observations on Nasmyth's membrane,

has
which will be described later,Charles Tomes
in
his
and
has
the
that
fact
altered
views,
acquiesced
nucleated cells,
the
remains of this external epithelium,
probably
do enter largelyinto the formation of Nasmyth's

of
publication

and
methods.

membrane,
modern

may
The

by
readily demonstrated
will
touched
be
question
again

be

consider Nasmyth's membrane, but in

when
we
be well to point out that long
it may
this connection
before the completionof the enamel
formation the cells
fused with the
of the external epitheliumare
practically
continuation of the sac in which the cementum
isformed,
and though Paul's observations are pretty conclusive as
of the nucleated
to the presence
cells,so is Tomes'
and it
the
of
the
cementum,
reasoningas to
morphology
does not seem
that both views may
to me
at all unlikely
upon

be

true.
substantially

is vascular.
organ
far 1 have
and
Wedl,
so
Legros deny this,
with no confirmation of Dr. Beale's statement.*
met
These
observations were
made
before the accurate
modern
available. I have
instruments were
doubt
no
the
tissue
outside
that the vessels observed
in
the
were
believes

Beale

that

the

enamel

Magitot,and

stellate reticulum disappears

vessels which are really
early,and after its disappearance
in the surroundingtissue look very much as if they were
external

epithelium.The

in the enamel organ.

The
Dentine
Organ, or Dentine Papilla,like the
enamel
and,indeed,their arrangement
organ, consists of cells,
is
and differentiation
very similar in both organs.
find a mass
In the dentine organ
too
of oval cells,
we
as definitely
a stellateform as the
closely
packed to assume
stellate reticulum,yet leavingplenty of room
for other
*

Howes
the enamel

and

Poulton professto have

organ of the rat.

this

observed

by

in
vascularity

26

AIDS

TO

DENTAL

ANATOMY

form some
secondary dentine to protect and strengthen
the threatened
districts. Perhaps they are
not
quick
and
breach
and
is
the enemy
a
enough,
effected,
organisms
(micro-

this

time)pours

in

; but

in such

case

and our metaphor, such

trenchingon pathology,
has, 1 hope,served the present purpose.

we
as

are

it is,

The

cells of the dentine organ, like those of the enamel

geneous
it were, in a homoas
organ, are all embedded, floating,
substance
which does not easilystain.
jelly-like
The
have been called odontoblasts
outer
or
ones, which
*
dentine cells,
and
membrana
the
ebons/
collectively
fn
somewhat
The
ends
towards
which
are
shape.
vary
dentine are mostly wider than the inner ends ;
the new
the shapes vary in different specimens and in different
specimen. To a certain extent this
parts of the same
of preparation.
cells
be due to the method
The
may
the
may be distorted and stretched in the act of cutting
and reagents used duringthe process of staining
section,
and

mounting

there
still,

are

may
modes

cause

them

swell

to

of observation
avoided, and it may

or

in which

to

shrivel

these

plications
com-

safelysaid

that
of a shape varyingfrom that of a
active odontoblasts
are
of
that old cells are shorter
a banana, and
pear to that
The
the work
cells which commence
than active ones.
smaller
of dentine formation are
supposed to be much
than those which complete it,
and this has been suggested
Mr.
of the greater
by
Hopewell Smith as an explanation
calibre of the fibrilsat the pulp end
a view which
seems
the
received
than
to
more
me
more
acceptable
widely
that as the dentine grows
older the fibrilbecomes
one
are

be

"

converted
matter

into

sheath, the

will be touched

upon

into matrix. This

under the head of
fully

sheath
more

Calcification.
that the process of dentine formation
All observers
agree
is due in some
of these cells,
degree to the activity
but the precisepart played by the cells is a question
be
of opinion. It must
which involves much
diversity
that the tissue is formed before it is
in mind
borne
that it is easy

to show
a
layerof formed,
d
entine
as
separatingthe cells from
yet uncalcified,
completeddentine.
Until recentlythe views most
generallyreceived in

and
calcified^
but
the

by

DEVELOPMENT

OF

THE

TEETH

27

this country were,

cell became
that each odontoblast
converted into dentine by the calcification of its outer
while its core
soft as the fibril,
axis remained
or
portion,
and
between
semi-calcified
the two
a
was
layer or
sheath of Neumann.
The subsequentfusion of the outer
portionsof contiguouscells,leavingno trace or line of
demarcation
cell ended
and the next
to hint where
one
less homogeneous matrix,
or
began, resulted in a more
while the fibrils remained
in connection with the cells
that formed
them.
The
odontoblasts
only were
posed
supin the process.
to be concerned
Klein considers that the odontoblasts form the matrix,
while the cells below them form the fibrilswhich
pass
between
the odontoblasts ; this view also requiresthat
between
the fibrils the adjacent odontoblasts
must
sections would show lines of
fuse,or transverse
absolutely
demarcation.
Magitot points out that there is another element to
stance
be taken into account
namely, the homogeneous subin which the cells are embedded.
This he regards
is
the seat
of the formation
of the matrix,which
as
calcified around the soft parts, the fibrilremainingconnected
lacunae
of
the
with the cell,
contents
the
justas
in calcified
and canaliculi of bone
embedded
remain
matrix.
Accordingto this view, which is rapidlygaining
general acceptance, no part of the cells is converted into
matrix.
This questionwill requirefurther discussion under the
head of Calcification.
The Cement
Organ is simply the continuation of the
under
the
surface of the dentine organ,
which
passes
outside
of
the
enamel organ, and, rising
deepestparts
up
mixed
it
and
becomes
it,envelops
finally
inextricably
of the
with the upper portionof the external epithelium
it is formed
enamel
The
cells of which
are
organ.
but
derived from the same
the dentine cells,
are
source
as
from
in
much more
irregular shape. They are separated
each other by matrix, in which
calcification takes place,
result that one
would expect if the
producing the same
matrix of the stellaterecticulum were
to calcify namely,
less
cells
stellate nucleated
more
or
anastomozing with
in a calcified matrix,the cells
each other and embedded
"

"

by

28

AIDS

as
persisting

TO

ANATOMY

DENTAL

of lacunae and the anastomozing

is cementum.
canaliculi. This substance

the contents

as
processes
Be it observed

that these

cells,
though

closelyrelated

so

curious
fusion of substance which
attributed to the
has been
latter; in fact,this fusion,
does
take place,is,as far
if it
I know, a unique phenomenon
in as far as the total
as
obliteration of all trace of individuality
of the original
cells is concerned.
Those cells which stretch out from the deeper portion
of the dentine germ, and envelop the sides of the enamel
sac/ or 'dental
organ, constitute what is called the
follicle.' They completelyfillthe space which intervenes
between
the germ
and the bony crypt which contains it,
and if at an
earlystage of developmentthe soft tissues
torn
are
away from the hard, this tissue goes with the
and
other soft tissues,
and leaves the bony crypt
germ
bare.
is divisible
The sac tissue,
when
fullydeveloped,
into two layers an outer, more
an
or less compact, and
inner,more
less loose and soft. The sac is,like all
or
the dermal structures
(as compared with the epidermal),
and on its surface are often
very vascular and nervous,
found indentations,which have been described as ' villous*
and which the exuberant
or
fancy of some
papilliform,
observers has endowed
with the property of determining
the direction of the enamel
prisms. The processes, when
to

the

make
odontoblasts,

no

at

attempt

the

'

"

are
they exist,

separatedfrom

the enamel

organ,

logically
physio-

and the idea that theyinfluence

anatomically,
fetched.
the direction of the prisms is,to say the least of it,farand

While

brains about these layersand

our
puzzling
as well as refreshing,
processes, itwill perhapsbe salutary,
to recall the fact that the morphology as
well as the
anatomy of this follicleis extremelysimple. The enamel
from the epidermal
layeris thrust into the
organ arising
in section)
tissue in the familiar form (asseen
subepithelial
of a Florence flask ; the subepithelial
tissue adjacentto
and touchingthe epithelial
the seat
invasion becomes
of a
the

we

are

cell excitement" that is, the cells nearest

in form, and
invasion become
epithelial
specialized
arranged in rows ; those oppositethe centre of the flaskshaped invasion become odontoblasts,and form dentine ;
sort

of

by

DEVELOPMENT
those

OF

enwrappingits outer

THE

TEETH

walls become

'

29

sac,'and form

cementum.

When
is requiredto be very thick,
the cementum
as in
the herbivorous animals,where it is destined to cover
the
whole tooth,according to Magitot,it first becomes
the
afterwards
of
formation
which
seat of a
hyalinecartilage,
ossifies like any other bone developedafter the intraWhere
it is to be thin,and only
method.
cartilaginous
the crown,
the relic known
cover
as
as
Nasmyth's
it
is
intramembranous
the
of
seat
membrane,
ordinary

ossification.
The remains
endowed
with
times was

of the neck of the enamel organ has been

and in bythe ' gubernaculum,'
a name,
gone
of
assist
in
the
to
eruption the
supposed

tooth.

The

Dental

Follicle

extension of the cell

dentine
papilla
; it

or

at

first appears

which commences
activity,
and
spreads outwards

graduallyenvelopingthe
enamel
As its

Sac

rest

organ) until it meets,

to

be

an

in the

upwards,
the
(including
speak,over its head.

of the germ
so

to

development progresses its outer layer becomes

fibrous,while its inner layeris vascular and cellular.The
forms the alveolar dental periosteum,
outer layerprobably
while in the inner the cementum
The outer
is formed.
cells of the enamel
somewhat
mixed up
become
organ
*
with the tissues of the sac,'
and some
cells,apparently
been traced down
a continuation of the former,have
as
far as to cover
in which situation
the roots of the teeth,
sheath of Hertwig^
they have been called the epithelial
of which persist
cells
the nests of epithelial
remnants
as
called the * glands of Serres.'
facts may be summarized
The foregoing
as follows :
{From C. S. Tomes' adaptation
of Rose's tables.)
Age before
Birth.

5-6 weeks

"

of
First appearance
of Meckel's
trace

inflection;
epithelial
trace
cartilage
; no

first
of

bone.
7

"

lamina

defined ; deep layer

of
of epitheliumcolumnar
; first traces
bone.

Zahnleiste, or

by

3o

AIDS

TO

DENTAL

ANATOMY

by

CALCIFICATION
Age

31

before

Birth.

9 weeks"
10

"

Buds
Buds

lamina for milk-teeth.

club-shaped; eight dentine

from

papillae

present.
14

"

17

"

of milk-teeth begin to separate from

lamina ; enamel organs differentiate.
Dentine
papillaeof six-year-old molars

Germs

formed.
20

24

"
"

of calcificationin milk-teeth.
Calcification in milk canines and
molars ;
of papillae
in permanent
first trace
ines;
canenamel
molars
organ of six-year-old

Trace

fullydeveloped.
29

"

33
"
9 months"

Calcification of cusps of milk molars.

Cusps of milk molars united.
thirds of centrals,
half of laterals,
tipsof
s
urfaces
of
molars
calcified
in
milk
canines,

Two

set ; cusps

of

molars calcified.
six-year-old

CHAPTER

III

CALCIFICATION
the tissues which are
Almost
destined to
as
as
soon
become
calcifiedhave begun to be formed or plannedout
4
in the soft/the work of deposition
of lime-salts,
which
givesthem theircharacteristichardness and which iscalled
'calcification,'
begins. The process is reallythe same
it happens ; but its results are
wherever
profoundly
of the soft tissues which has
modified by the disposition
taken place,and much
confusion would be
previously
understood
spared to students if it could be once fairly
that the essential characters of the various tissues are
and that then
mapped out by the cells in soft tissue first,
the depositof salts of lime renders the structure
which
they have formed permanent and hard.
of calcificationwere
The
first
great general principles
about half a century ago by Rainey. Long
laid down

by

AIDS

32

TO

DENTAL

ANATOMY

afterwards,Harting,a German
writer, recapitulated
and elaborated the subject
siderably.
Rainey'sobservations,
very conwrote
some
Ord, a pupilof Rainey's,
Lastly,
valuable papers revising
his old master's observations,
and adding some
importantnew matter. These observers
between them may be said to have placedthe physiology

of calcificationupon a sound basis,

and to have enabled
of the process as it occurs
to givea reasonable account
us
in the various hard tissues in fact,
to have opened up the
of the formation of enamel,
way for a proper understanding
dentine,and bone.
Mummery
Recently Mr. J Howard
has communicated
result
of his
the
the
to
Royal Society
own
researches,but the paper has not yet been published.
The theoryof calcificationadvanced
by these observers
is in itselfextremelysimple; it is based upon one great
generallaw namely, that lime-salts which are deposited
in a substance containingalbumin
in a very
behave
ing
itselfexhibitspecialmanner, the deposit(or precipitate)
"

"

specialform

and

solution of salts of lime be

specialarrangement. If a
acted upon chemically
so
as

of the
the result is,that at the bottom
precipitate
it,
lime
is
vessel in which the process is going on
the
in the form of a powder ; an amorphous powder
deposited
it is called,because
its particles
without definite
are
If
albuminous substance be firstintroduced
some
shape.
into the vessel (such as a tooth, for instance,or a piece
of gelatine,
or
anythingcontainingalbumin),and then
the lime-salts be precipitated,
the resulting
is
precipitate
to

no

longer amorphous

is round, and
on
particle
consist of several layers,
rather

each

division will be found to

like an onion.
These round bodies are called * calcospherites.'
If this
be redissolved so that the lime is once
more
precipitate
in the firstcase, where no albumin
taken up into solution,
had been present, the amorphous powder would
melt
would
remains
indicate
and
the
naked
to
no
away,
eye
than if it were
that it had been there,any more
so much
dissolved in hot water ; but in the second
sugar
case,
albumin
been
where
had
would
present, something
The
remain.
consist of lime mixed
timately
incalcospherites
with albumin,and, when
the lime is dissolved
be
out of them, itcannot
removed
apparently completely
;

by

AIDS

34

TO

DENTAL

ANATOMY

dentine.

takes place
The formation of the calcospherites
in the tissue that has already been formed in the soft,
and involves the soft structures
without disturbingthem.
contrived
the cells have
Thus, whatever arrangement
receives permanence
and
the
hardness
by
process of
calcification.
Calcoglobulinis the altered albumin, as explained
above, and shares the property of an exceedinglygreat
of resistance to acids with all albuminous
tissues
power
that lie on the borderland of calcification. The fact that
this tissue is always the last to be destroyedaccounts
for
its presence
wherever a tract of calcifyingsubstance
is
destroyedby the action of acids,since it possesses a
of resistance to these agents than either
greater power
the wholly uncalcified or the wholly calcified material.
This fact has caused a great deal of confusion in the
of students of anatomy, because
minds
observers
many
have described
membranes,' which may be isolated by
the action of acids,in different situations,
of which
some
which
are
probablyonlythe half-calcified calcoglobulin
t
he
outskirts
of
calcification.
necessarily
occupies
Huxley,
when
found
the
between
enamel
describing
calcoglobulin
'
and the enamel
organ, has called it the membrana
preformativa.,
While, however, its isolation is due to the
be said not to exist ; but the mistake
reagent, it cannot
is to call it a ' membrane,' instead of recognizing
it as a
brief stage in calcification. We
shall repeatedly meet
with this tissue wherever
calcification is going on, and
shall often find that it has received a specialname
and
'

been

regarded as

Sims

specialstructure.

Woodhead

suggests that calcificationis a form of

He
degeneration.
regardsthe active cells as comparable
cells which are always found near
to the proliferating
a
cells lay down
dead tissue. These
of formed
a matrix

material,which

he regards as dead organicmatter.

The
of
combinations
molecular
phosphoricacid and lime take
the
about
active
cells,which, during their activity,
place
give off carbonic acid to render the lime for the time
being insoluble. * There is a continual process by dialysis
of separationof these salts.' He regards the
dead material,as a dialyzing
which
membrane,
the lime-salts

prepared by

the carbonic-acid

matrix,or
separates

formingcells.

CALCIFICATION

Calcification of Bone.
which

"

calcification of bone,

called, may

take

35

There

methods
by
it is usually
as
ossification

or

place,viz.:

"

are

two

intramembranous

and

ossification
intracartilaginous
that in both cases
It is important to remember
a
that
of connective tissue is first formed
membrane
so
speakingall ossification takes place in membrane
strictly
;
second
but in the
or
intracartilaginous
variety,
cartilage
is formed and reabsorbed to make way for the bone.
of
All ossification is the direct result of the activity
cells called osteoblasts. These cells do
certain specialized
but wherever
bone
is formed
themselves
not
calcify,
the calcification
osteoblasts are present. In discussing
that in those cases
of enamel and dentine, it will be seen
cells are always present, which do' net
also specialized
but are, in some
themselves calcify,
not
yet fully
way
the
calcification
f
or
understood,responsible
takingplace
in their neighbourhood.
bones
Membrane
Ossification in Membrane."
are
formed as the result of calcificationin fibrous connective
All the bones
tissue,without any formation of cartilage.
of those of the face,
of the vault of the cranium, and most
bones.
includingthe superiormaxillae,are membrane
The first stage of the process is the formation of a centre
In this region,specialized
bone
cells
of ossification.
make
their
the
tissue
osteoblasts
in
the
or
appearance,
tion
Calcificaneighbourhoodbecoming somewhat
opaque.
then takes place in the tissue around
the osteoblasts.
the bone is fullyformed, the
of time,when
In course
bone corpuscles,
remain
osteoblasts
as living
imprisonedby
called
the bone they have manufactured, in the spaces
L
bone.
lacuna
so characteristic of normal
is in a sense
Ossification in Cartilage."This term
that
it
does
the
misleading,suggestingas
cartilageis
Such a thingnever
into bone.
takes place.
transformed
In the fibrous connective tissue membrane,
cartilage
is firstformed.
Cartilageconsists of white fibrous tissue
*
as
together with a compound known
chondrosulphuric
with
lime-salts to a
whole
the
being impregnated
acid,'
Previous to the formation of bone,
less degree than bone.
the cartilagecells become
arranged in regular rows,
in
The tissue is then invaded
somewhat
size.
and increase
.

"

by

AIDS

36

by bloodvessels

ANATOMY

TO

DENTAL

and

by osteoblast cells. With the

cells called osteoclasts.
specialized
of
latter is to produce absorption

other
osteoblasts come
function of these

The
the

and
also, to a certain extent, of the
cartilage,
newly-formedbone ; in this way the medullaryor marrow
The
actual ossification takes place
cavityis formed.
around
the osteoblast cells,
justas was described above
intramembranous
ossification.
in discussing
What the precise
Calcification of Enamel.
process is
of the calcified tissue known
that results in deposition
as
much
but
be
be
said
enamel cannot
to
settled,
entirely
"

work
valuable
firstappeared.
of the problems
Charles
has worked
Tomes
out
many
still
connected
with the subject,
but
at present it will be

and most
evidence has been
collected,
done in this direction,
since this handbook

wise to regard
transition stage.

our

knowledge

as

passing through

The facts about which no one

disputesare, roughly,as
follows : Where
enamel is to be formed, we find specialized
elongated cells,called ameloblasts,of a granularappearance
and with largenuclei,
at the end farthest from where

is to be formed.
Beyond these cells,on the
side away from the enamel formation,are the cells of the
interval occupiedby the
intermedium
stratum
; then an
the external cells
cells of the stellate reticulum ; and lastly,

the enamel

enamel
Gradually the big ameloblasts
organ.
blasts,
recede from the line which divided them from the odontoand in their track is found enamel.
The cells of
and so do the cells
intermedium
the stratum
disappear,
while the external cells of the
of the stellate reticulum,
of

the

possiblypersistas Nasmyth's membrane.

organ
smaller as the process
ameloblasts
do not become
If they are
rate.
at any
at first,
not
pulled
goes on"
end
in
from
the
to
enamel,
they
forming
appear
away
short processes, first described
that direction in tapering,
called
after him Tomes'
and
by John Tomes,
processes ;
less soft. The surface of the
these processes are more
or
have been
formed
enamel from which the ameloblasts
to resemble
a
honeycomb.
pulled appears somewhat
formed
tissue
when
The
(in man), is so
fully
enamel,

enamel
The

completelycalcified that,though it has definite structure,

in its
find any trace of organicmatter
C. S. Tomes
cannot

by

CALCIFICATION
chemical

composition,but only 3 per

Lovatt
Evans'
analysis does not
Mr. Tomes, and it will be givenat a

37

Dr.
of water.
with that of
agree
later page in detail.
No
in
doubts that the ameloblasts are
one
concerned
the process of enamel
formation"in
fact,that they are
think entirely,
for it; the
some
principally,
responsible
doubtful pointis how they do it,
and about this authorities
cannot
yet be said to agree.
Tomes, followed by Schafer and others,used to teach
that the ameloblasts,or enamel
cells (cells
of the internal
become
into
converted
enamel
epithelium),
prismsby a
of
salts
of
in
lime
their
substance.
deposit
According to
this view,the cells are calcified from without inwards,the
axial portionbeing the last to calcify,
and so far the
resembles
the
of
older
description the calcification
process
of dentine.
There were, however, three great differences
between the two processes: (r) That in enamel the part
formed by each cell always remained
distinct from that
formed by its neighbours,
and the result was
separate
of
each
had
which
a'
been
formed
cell
or
prisms,
by a
succession of cells ; whereas
in dentine the tract of
calcification taking place in each cell fused with its
immediate
neighbours,so that no line of demarcation
leftto suggest where the division between neighbourwas
ing
cells existed. (2) The axial portionof the enamel
did eventually
so
cells,
though the last to calcify,
calcify,
that attached to each cell there is a short,tapering
spike
of semi-calcified material,
which dippedinto the new
prism
cent,

length not exceedingthe length of the cell (Tomes'

process),whereas in dentine the axial portionremained
uncalcified as the dentinal fibril. (3) In the resulting
tissues the fibrilsbranch freelyand inosculate with their
to

whereas
neighbours,
The

enamel

enamel
cells,when

prismsdo
torn

enamel, generallybring these

branch.
the young
away
taperingprocesses with
not

from

them, and

when
subsequentlythe cell shrinks, the
portionof it that lies nearest to the young enamel does
not shrink,
owing to its beingpartially
impregnatedwith
lime-salts ; the result is that the cell,
with its process and
looks rather like the blade of,a
the unshrunken
portion,
detached from the handle,and if the
dinner-knife when
handle be regarded as a young
prism,the relation of the

by

AIDS

38

TO

ANATOMY

DENTAL

parts is not unlike that of a knife handle and blade. If

the parts be separated
by the action of an acid,a layerof
"

discovered between the cells and the

enamel, which will be piercedwith small holes where
the processes
dipped into the prisms; this is called
the foraminated sheet,or fenestrated membrane.
less,
Doubtif the separation
carried
without
out
were
tearing,
this sheet would be found not pierced,
but invaginated
willbe
calcoglobulin

these

at

the
and

must
we
points,as, to be consistent,
between
intervene
the
to
calcoglobulin
of
the prism throughout. This
view

calcification received for many

'

suppose
process

'

enamel

the widest acceptance.

years

Kolliker regards the process as one of pure secretion.

He does not believe that the cells become
calcified or
converted
the
into enamel, but thinks that they secrete
calcifiedsubstance.
share
this
to
Waldeyer seems
view,
and describes the cells as having open mouths.
Magitotthinks there is a plateaubetween the cell and
the prism,through which a substance
passes by some
becomes
sort of osmosis, which
the seat of calcification.
prePerhapsthe * plateau'is only Huxley'smembrana
formativa and Tomes' foraminated sheet under another
thrown
observations have
More
new
name.
recent
a
this
obtain
Tomes
has
been
able
to
lightupon
subject.
series of sections of developingmarsupial enamel,
a
which offers in many
respects exceptionalopportunities
for examining the nature
of the process, for the simple
that such enamels
tubular when fully
reason
are
formed,*
and, if Tomes' conclusions are correct, represent enamel
similar in nature
to our
prismaticenamels, only not
calcified to so high a degree. He has shown, as others
in old-world enamels, that the ameloblast
have shown
cells contain
small rounded
bodies, probably calcothe cells are drawn away from the
; that when
spherites
thread-like prolongations
minute
connect
formingenamel,
of the formed
them with the axes
prisms; that these
and
threads are soft and elastic,
probably in tubular
enamels
He
of the tubes.
persistas the contents
*

has carried this research

Mummery
demonstrates
that the soft fibre is between
their axes, in marsupials.
"

stage further.

the

prisms,and

not

He
in
/

by

CALCIFICATION

39

summarizes

his general conclusions (in his edition of

follows:
1898)as
That the ameloblast itselfdoes not calcify.
That each ameloblast furnishes a fibrillarprocess, continuous
with its own
for the entire
plasm, which serves
of
enamel
length an
prism.
That
calcification and the changes preparatory to
calcification take place in or around
each fibril,
quickly
its
central
solid
axis
in
reaching
prism enamels, but
never
reachingit in tubular enamels.
It will be seen
consistent
that the appearances
are
either with the calcification of the fibrilor with calcification
going on round it and, so to speak, squeezingit
down
to smaller dimensions
it.
or obliterating
The

exceedinglysmall

present in solid enamel

amount

mi^ht seem

of

organic matter
to point to the

latter

this so, one

view, but were
being the preferable
of the prisms to
might have expected the individuality
terstitial
be lost,whereas
of inamount
there is an appreciable
substance demonstrable between the prisms.

and
The
enamel
forms
both
same
prismatic
organ
tubular enamel in certain marsupials,
which shows that
it is quitejustifiable
the development of the
to deduce
from that of the other.
one
Observers have endeavoured to find an analogybetween
the formation of the shells of certain molluscs, such as
Both processes result in the
pinna,and that of enamel.
formation of a hard prismaticsubstance,in both the
less hexagonal,and in both the
or
prisms are more
but separatedby
contiguous,
absolutely
of the
In the case
substance.
a layer of interprismatic
is
of organic matter
shell,however, a large amount
the process
Whatever
mass.
present in the fully-formed
to be, its result is so
peculiaras to be
may
prove
of
A
hard prisms,separated
mass
practically
unique.
from each other by interprismatic
substance,which must
if there is no
also be almost entirely
organic
calcified,

prisms are

not

under
the
traceable in the whole mass, formed
auspicesof cells which, though very large and active,
or remain
disappearwhen the process is finished,
totally
in a flattened condition as Nasmyth's membrane.
If,as
its
each
cell
forms
tract
or
own
seems
prism of
likely,
matter

by

AIDS

40

TO

DENTAL

ANATOMY

enamel
corresponding
exactlyto it in
of demarcation dividingthe work of one

diameter, the line

cell from

that of

itsneighbour persists
throughoutlife.
of sections of enamel of individuals suffering
In a number
Dental
tion
Associahave
shown
from erosion,
I
{British

Journal, 1 892) photographsof interglobular

spaces,
that there is no doubt that the process, in whatever
so
tissue it occurs, proceedson the same
lines as the other
calcifications.
That the key to this puzzling
questionshould liein the
of
enamels seems
study marsupial
very probable; whether
the
have
or
some
supposed,marsupialswere
no, as
of the mammalian
aucestors
races, they certainly
resent
repand
a
in
one
very ancient order of creatures
which we may
o
find
suggestionsf
reasonablyexpect to
the earlier stages of processes
that are, in their more

advanced

condition,
very difficultto study or explain.

that in
believe for a moment
selves,
enamel, or any other calcified structure, the cells themor any
part of them, are converted into calcified
tissue.
The calcificationof enamel, commencing at the edge of
the dentine,proceedsby additions on the outside. After
the enamel
is complete no trace of the cells which formed
it remains.
They entirelydisappear(unless we accept
the suggestion
of Nasmy th
of Kolliker,
that the membrane
is the remains of the last cells of the enamel
organ)
far
know.
which
I
is
a phenomenon
as
quiteunique,as
Calcification of Dentine." The calcificationof dentine
beginsa triflebefore that of enamel, and, as in the latter
means
complete agreement
process, there is by no
between authorities as to the details of the process.
It used to be taughtthat each odontoblast formed three
semi-calcified
different tissues namely, uncalcified fibril,
sheath, and completelycalcified matrix. The way in
which this was
done was
simplythat the different parts
of the cell were
impregnated with lime to a different
axis
and
the
the surface completely,
not at all,
degree
the area
the only difference
between the two partially,
between
the three tissues being a difference of degree of
calcification. There were
certain difficultiesin the way
of acceptingthis theory,
which I have for many
years
I do
Personally,

not

"

"

"

by

AIDS

42

TO

DENTAL

advancing through the

of

mass

ANATOMY

quiteirrespective

of matrix

cells,always separated from them, in fact, in

developingdentine,by a layerof formed but uncalcified
tissue,which readilystains with gold chloride or any of
the

the aniline dyes. I have

seen
enteringthe
processes
dentine as fibrilsfrom both the odontoblast
layerof cells
and the layerunderlying
it
of the process seems
to be
to me
Magitot's
explanation
the most
reasonable that has yet been offered namely,
that while the cells are occupiedwholly with the formation
of fibrils,
the transparent homogeneous substance
which embeds
them
and
all the other tissues of the
formative pulp is the seat of the calcification,
which,
from
surface
before
it all
drives
the
inwards,
advancing
of
the pulp tissue,
of the prolongation
with the exception
the cells,
which remain soft and unaltered as fibrils. If
of the process be accepted,
this account
it will be seen
in
that there is a strict analogy between
the manner
which dentine,cementum,
stiffenedand
and bone,become
with lime-salts. In each case
hardened by impregnation
epithelial
of
branches
the
cells derived from the subanastomozing
and in one
the cells themselves,become
tissue,
imprisonedby a hardening of the matrix in which they
the result is a calcified,
embedded.
In each case
are
communicating
intercontinuous matrix, traversed by soft,uncalcified,
"

the other
In enamel, on
hand, the cells are derived from the epidermallayer; the
calcifiedmatrix is not continuous,
but sharplydefined in
substance the exact
prisms between which exists some
and degree of calcification of which is not yet
nature
cell-branches.

ascertained.
The
which play a principal
cells called odontoblasts,
long,finely
part in the process, are, like the ameloblasts,
granular,with largeoval nuclei at the end farthest from

the

the
as
They are not so columnar
and
close
not nearly so
nected
conare
cells,
together,
with their neighbours,with the underlyingcells,
with the dentine itselfby branches.
new

dentine.

enamel
and

modern
views, for which we are greatly
Mr. Mummery, and which are acceptedby
Mr. Tomes, the calcificationgreatlyresembles the process
in the ossification of membrane
cells are
The
bones.

According to

indebted

to

by

CALCIFICATION

43

finelyfibrillarmatrix, which is rendered

more
apparent by the Weil process of preparingsections.
coalesce in this fibrillar
The calcospherites
form and
matrix,while the cells retreat towards the future pulp,
branches,or fibrils
leavingtheir dentinal prolongations,
of
embedded
in the calcifying
The main essence
mass.
in

embedded

the view
namely, that it is the matrix and not any part
of the cell itselfthat calcifies accords with the old view
fetched
of Magitot,and renders unnecessary
the fanciful and far"

"

cells completely fusing

imaginingsof calcifying
soft-outlined cores
or
together,but preservingdistinct,
axes.
Here, again,comparativeanatomy supports the
reasonable
is shown
as
by Mr. Tomes
explanation,
in his observations on
vaso-dentines in developmental

stages.

This

represents in brief the

teachinga

current

whether the lime

few years ago.
It has been questioned
in teeth can
be depositedin globularform,because this
is not supposedto take place in tissues where phosphates
in excess
of carbonates,as they are in enamel and
are
dentine.
It is sufficientfor me
that the salts always are
of
has ever been
other
mode
so deposited
; no
deposition
has gone over
demonstrated.
Mummery, quiterecently,
the ground with the aid of modern methods
of investigation,
and
has
his observations in a paper
embodied
communicated
the
to
Royal Society,December, 19 13.
I am
not at liberty
to describe his results until the Royal
Societyhas made them public.
Calcification in other Dentines.
Plicidentine is simply
hard dentine folded,and its calcificationis identical with
Vaso-dentine
that of hard dentine.
differs only in this,
that as the matrix is hardened
by impregnationwith
obliterated or
lime-salts the capillary
vessels are
not
pushed out of the way, but the process goes on around
them, and they are left permeatingthe calcified tissue
justas theyonce
permeated the pulp tissue in which it
formed.
Osteodentine
after the
is calcifiedsomewhat
was
of membranous
manner
bone, trabecular passingthrough
its substance,and calcospherites
formingand coalescing
round all the pulp tissues,
that the spaces contain
so
connective
and
tissues. The
same
bloodvessels,
nerves,
dentine organ may, and sometimes
does, form all three
varieties,
and, as was foreshadowed in an earlier chapter,
"

by

AIDS

44

DENTAL

TO

ANATOMY

calcification process is reallythe same,

the only
difference being that more
less other tissues are
or
in
the
so that the tissues when
entangled
calcifying
mass,
fullyformed
present very different anatomical appearances.
the

This process takes place

Calcification of Cementum."
in the folliclewall. Where
the cementum
is thick,as in
of those teeth which
the case
covered with coronal
are
cement, it is supposedto be calcifiedin a hyalinecartilage,
after the manner
of intracartilaginous
In these
bone.
*
also
been
described
has
definite
a
cement
cases
organ
the tissue is thin and conWhere
observers.
fined
by some
in man,
the ossification is intrato the roots, as
is formed the tissue
Where
membranous.
the cement
in which it is calcified remains
the alveolo-dental
as
blasts
periosteumor ligament. Large formative cells or osteohave been described upon the inner surface of this
tissue. Processes
of the connective tissue of the periosteum
into
found
the
formed
cementum
are
as
passing
fibres.
Sharpey's
'

CHAPTER

THE

IV

TISSUES

of developmentof the human

traced the course
endeavour
the minute
to describe
tooth,I shall now
it in the
anatomy of the several tissues that compose
condition. These tissues are from without
fully-developed
inwards
Cementum, which envelops the root; a thin
tissue called Nasmyth's membrane, which covers
the
of
crown
; Enamel, which, underlyingthe membrane
the crown
Nasmyth, covers
; Dentine, which, covered in
Having

"

in the root by the cementum,

forms the bulk
tooth,and decides its shape and
size ; and the Pulp, which
occupiesthe central hollow
of
the
The whole tooth is supported
dentinal mass.
core
in a bony environment,
which springsfrom the maxilla
and embraces
the root or roots of the tooth,and is conthe

crown

by the

enamel
of the

and

by

THE

TISSUES

nected with the surface of the cement

tissue continuous with the gums,
and

45

by a layerof soft
denominated
variously
The
of
consist,
therefore,

periosteum,periodontalmembrane,

etc.

have to deal
tissues with which we
three that are
calcified,
namely, Enamel, Dentine, and
the Pulp and the
Cementum
that are uncalcified,
; two
and
of
Periodontal membrane
doubtful character,
one
;

namely Nasmyth's membrane.

Chemical Composition of the Calcified Tissues."

The

calcified tissues are

variouslysuppliedwith inorganic
and
their chemical composition
salts,
may be conveniently
treated altogether.Of course, no two teeth giveexactly
the same
results after analysis,and it is both very
difficultand very unnecessary
to recollect all the decimals
and fractions discovered by various analyzers. What
the
student needs to know is an approximatestatement, and
therefore I have drawn up a table which, while not being
or any other
exactlyin accordance with Bibra or Berzelius,
and
is
remember
experimenter, near enough
easy to
"

It will be

seen

from

this table that the

amount

of

between
varies considerably
infancyand
which would seem
to suggest that that tissue
adult life,
outside the pale of nutrition. Many
is by no means
interestingdetails concerning chemical compositionof
be found in an exhaustive paper upon the
teeth may

organicmatter

by

AIDS

46

TO

DENTAL

ANATOMY

subjectby Galippein the Journal of the British Dental

1886, p. 361. Hoppe Seyler has found in
Association^
the enamel

child as much
as
22*29
and
in
the
organicmatter,
971
young pig,
in the fossilrhinoceros 3*16. Bibra gives2*00 of organic
in the adult male, and 5*00 in the adult female.
matter
A glance at the foregoing
table will show that the chief
per cent,

of

the

new-born

of

of phosphate and
variation is in the relative amounts
carbonate of lime found in the various tissues. It must
akin
not be supposedthat the said table is in any sense
It is placedhere because I feel
memoria
technical
to a
to
that whereas it is as useless as it is difficultto commit
all or any of the fractional variations in chemical
memory
analysesthat have been recorded,it is both useful and
*

the general principles

to all,
common
teeth
been
the
have
that
ever
analyzed
seeing that all
that
those
would only compare
in
the
to
ocean
as a drop
value
have not.
It would be absurd to attach too much
to the minutiae of the results,
yet it would be a pitynot to
how
know about
much
element preponderatesover
one

easy

to remember

of us whether
I do not think it matters
another.
to most
of any givenelement,
Berzelius found 1*033, or Bibra 1*038,
that everyone
that there is
but it does matter
agrees
nearly twice as much carbonate of lime in bone as in
cementum.

In

estimatingthe

exact

proportionsof

the

various

chemical constituents that go to make

up a dental tissue,
is
it
examined
should
obviouslynecessary that the mass
consist only of that tissue. If,for instance,we*have
a
littledentine or a littlecementum
to our
or both sticking
pieceof enamel, the niceties of our analysiswill suffer,
and in this way, perhaps,some
of the discrepancies
that
in
have
arisen.
In
fact,
publishedanalyses may
appear
enamel
is such a thin and brittle structure, as it exists in
that it is extremelydifficultto
human
teeth,for instance,
of
To avoid this source
isolate for analytical
purposes.
examined
has
Tomes
in
enamel,
error,
mainlyelephant's
order to be sure of gettinga fair-sizedpieceof what was
and
He
noticed that the
only enamel.
reallyenamel
of
amount
organicmatter with which these tissues were
credited was
ascertained by ignition that is,the weight
and the loss in
before and after burningwas
ascertained,
"

by

THE

TISSUES

47

the process set down

of organicmatter.
the amount
as
It occurred
that other thingscapableof being
to him
and a
weighed might be lost in the process of ignition,
careful series of experimentsled him to conclude that
which
the 3 per cent,
disappearedin the burning of
enamel
at all,but simply water
not organicmatter
was
;
that enamel
if any, organicmatter
contained so little,
worth
that this tissue
that it was
not
so
considering,
he regarded as
consistingwholly of lime -salts and
water.

International Medical
(Transactions,
of
Section
Congress, 19 13,
Stomatology) reported a
fresh analysis
of enamel
methods
in which newer
enabled
him to obtain results that greatlymodified earlier views.
His observations will be most useful if quoted rn his own
Dr. Lovatt

words
*
The

Evans

in the
the presence of organicmatter
has often been raised,and experimental
dental enamel
work
directed to the solution of the questionhas been

questionof

workers.
attemptedby numerous
Many of these workers are of the opinionthat there is
no
organicmatter in the enamel, and in support* of this
statement
theyurge the fact that,if enamel be decalcified,
coherent
no
This,
organic residue is left behind.
*

however, may
amount

of

be taken

merely as

organicmatter

is

an

small,or

indication that the

that it is soluble in

dilute acids.
of the enamel, and its durability
hardness
when
dried, are further indications of the smallness of the
*

The

amount

case,

of organicmatter.
evident
it becomes

If we
that

that this is the

assume
the usual tests for the
failfrom want of delicacy,

may
presence of organicmatter
and in order to decide the questionwe
test

capableof detectingvery

small

employ

must

amounts

of carbon.

of determiningthe loss of weight of the

The
method
for various reasons.
enamel on heatingcannot
be accurate
of the tissues would
In the firstplace,the carbonates
be
with loss of weight,which would
liable to decomposition,
due
of organic matter.
be counted
the
to
as
presence
is also undoubtedly a certain amount
There
of water
in
the tissue ; and as it is difficultto estimate the quantity
of organicmatter
of this with great accuracy, the amount

by

AIDS

48

in the enamel

TO

could

heatingthe weighed
'

ANATOMY

DENTAL

be deduced
substance.

not

Organic compounds

from

the results of

defined as the compounds

strictly
this
adhere
to
we
we
definition,
that enamel
does contain organicmatter
in
must
agree
the form of carbonates,which
would
stances.
be organic subunderstand
carbonates
Usually, however, we
to be excluded,and
thus the questionbecomes, " Does
the enamel
contain compounds of carbon other than
carbonates ?"
The question
undertaken
of one
at the suggestion
was
of us (A. S. U.),who is also responsible
for the preparation
of the samples of finely
divided enamel, a task of no
small difficulty,
in view of the fact that about 3 grms. of
the substance were
requiredfor each analysis.Normal
teeth were
first cleaned in dilute hydrochloric
acid,then
washed
and dried.
'Mr.
Underwood's
of preparationof the
method
specimens of enamel to be used for analysiswas as
of carbon

are

if

'

follows
'The

teeth chosen were

almost exclusively
canines,
because we feltthat the presence of irregularities
of form
might allow of the unintentional inclusion of fragments
of dentine.
and
divided longitudinally,
The teeth were
examined
with a lens to be quitesure
of the position
of
the dividingline of the two tissues. The dentine was
then removed
with a burr, and as soon
the enamel
as
reached
the
of
sound
the
burr
different
announced
was
the fact. Then, to make
quitecertain,a fair layer of
enamel

was

burred

away

so

that

safe distance

from

the

reached.
dentine was
examined
Every fragment was
with a lens to make
that
quitesure
by no possibility
could any dentine have remained in the mass.
To obtain
of teeth were
sufficientmaterial a very largenumber
cut
submitted
enamel
but
and
much
no
wasted,
fragment
up
have included any
could by any possibility
to Dr. Evans
dentine.
of his results after analyzing
The uniformity
so many
would
of
show
that
itself
other
material
no
samples^of
tissue of a different compositionwas
submitted to him.
in our details of this stage of
have been particular
We
the proceedings
much
as so
hangs upon it.
'

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AIDS

50

DENTAL

TO

ANATOMY

short account

be out of place
of the apparatus may
not
of
The apparatus is the Toepler pump, by means
of
the vacuum
is made.
There
is a tube
Jena
in length and
glass combustion
tubing about 20 cm.
about 1 5 mm.
end ;
which is closed at one
in diameter,
the
the other end is drawn
and
attached
to
out
pump,
and
At one
as
as
possiblemade.
complete a vacuum
end is the dried residue mixed with about five times its
weight of recentlyignited
copper-oxide
powder. Next to
this there is a layer of granular copper
oxide "from
wire,"also recentlyignited; next is a spiralof recently
heated copper gauze.
When
is satisfactory,
the vacuum
the grandularoxide is heated to brightredness ; then the
dried residue is slowly heated up to redness. Any gas
which is formed is pumped off and collected in a vessel
then analyzedin the Frankland
It was
over
mercury.

here.
which

bone-gasanalysis
apparatus.
The results of the analyseswere

Enamel.

'Analysesof Human
*

of precipitation
First sample : The
and
processes
loss.
filtration must
entailed some
have
Weight of
Volume

enamel, 3*659grms.
of"
consisting
c.c.
*

carbon
o*o

obtained,39*56c.c,
c.c.

nitrogen,
9*35

c.c.

; oxygen,
A sample of the

of carbon

of gas

dioxide,30*21

in

gas

turned

lime-water

enamel, 05 5;

centage
milky. Perpercentage of nitrogen,

04.

'Second

sample : Enamel dissolved in HC1 and twice

evaporatedto dryness. Residue combusted.
Weight of
enamel,

3*1975

grms.

Volume

of

carbon

dioxide,

of oxygen,
of
7*82 c.c. ; volume
35*38 c.c. ; volume
45*62 c.c.
nitrogen,242 c.c. ; total,
Percentage of carbon in enamel, 0*602 ; percentage of
nitrogenin enamel, 0*306.
*

Analysisof Elephant EnameL

'Treatment
as
Volume
of carbon
volume

of

above.

Weight

of

enamel,

dioxide,62*0 ; volume
nitrogen,
163 ; total volume,

3*5205.

of oxygen,
8o*i c.c.

by

i*8 ;

THE

TISSUES

51

Percentage of carbon in enamel, 0*97

nitrogenin enamel, 0*57.
4

of the

percentage of

the amount
of
is greater than in human
enamel.
4
of carbon
The
relative amounts
and nitrogen
do not
higher than
agree very well, the nitrogenbeing much
would correspondwith the carbon found.
Probably this
difference is due to the absorptionof nitrogenon
the
glasstube and pump, and on the copper-oxide
powder,
from which it would
be driven off by heat.
Great
in
r
esults
is
the quantitative
not
claimed,
accuracy
however.
The
results given here prove
simply that
is from
organicmatter is present, and that its amount
In the elephantenamel
cent,
it is
at least
1 to 2 per
cent.'
probably over 2 per
In dentine analyses,
as in enamel, the results of various
which are to me
as
analystspresent slight
discrepancies,
unimportantas they are uninteresting
; but Mr. Tomes
interest
and
of
notes
a fact
importance namely,that
contains
a
ivory
comparativelylarge proportion of
derived from the calcoglobulin
which
a substance
elastin,
surrounds
the sheath of
and is known
the fibrils,
as
of
dentine
that
the
the
also
Neumann,
elephant'stusk
contains 34*0 organicmatter, 8*5 of combined
water, and
animal's
57*5 of salts ; while the dentine of the same
molar yields
only 21*0 of organicmatter, 9*0 of combined
and
70*0 of salts.
water,
If a young tooth be immersed
Nasmyth's Membrane."
for a short time in a weak acid (1 per cent, chromic acid)
is slightly
the outer surface of the enamel
as
soon
as
be
white
will
thin
membrane
found
to be
softened, a
the
but
the
free
round
attached
neck
to
ot
enamel,
lying
first described by Alexander
the tooth. This tissue was
after him.
At
Nasmyth, and was
accordinglynamed
of the tooth,but soon
first it covers
the whole crown
becomes
at the
worn
masticatingsurface. It
away
it shows a minutely-reticulated
stained with silver nitrate,
which
led
observers to suppose it
has
some
appearance,
consists of an aggregationof flattened cells,the intercellular
the stain. It is sometimes
substance
receiving
dental cuticle or dental capsule. It
called the persistent
is about sQ"ofl
of an inch thick at its thinnest part, and
In

the

case

elephantenamel

organicmatter

"

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52

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TO

is

resistant
remarkably

DENTAL

ANATOMY

the action of acids,probably

calcified.
because it is imperfectly
considered that Nasmyth's membrane
The elder Tomes
the crown
was
over
onlya thin extension of the cementum
is a summary
of the tooth. The following
of the reasons
based :
upon which this conclusion was
is
tissue
continuous
with
the cementum,
The
i.
and
remains attached to it when set free from the enamel by
the action of acids.
In one
specimen (inthe collection of
Mr. Tomes, and figuredin his manual) the edge of the
be seen
cementum
overlappingthe enamel, and
may
taperingoff into the membrane.
the cement
2. Wherever
is quite
coveringthe crown
herbivorous
is
animals, the membrane
thick,as in the
been found co-existent with coronal
absent ; it has never
to

cement.

is specially
the membrane
thick,as when it
contains lacunae
fillsa pitin the enamel, it sometimes
and canaliculi.
The
absence of lacunae from the greater part of the
is no argument againstits cemental
membrane
nature,
fact
that wherever
is
because it is an accepted
cement
structureless.
thin
itis
very
Magitot and Wedl agreed with the Tomes.
Huxley
called the tissue 'membrana
which, as we
preformativa,'
that he could isolate it with a weak
have seen, meant
acid. Waldeyer was
unconvinced
by the arguments that
satisfied most
writers,and maintained throughout that
due to the persistence
the membrane
of the external
was
and
it has always been a
cells of the enamel
organ,
that all the cells of the enamel
of wonderment
matter
work was
finished,
organ should,after their extraordinary
entirelydisappear,leaving neither trace nor vestige
Its resistance to reagents suggests that it is of
behind.
of calcoglobulin.
the nature
fully
Recently Paul has successwith eosin and haematoxylin,
stained the membrane
that it does consist of flattened
and has demonstrated
cells ; that they are so much
largerthan the ends of the
prisms as to bear no relation to them at all ; and that
they possess large nuclei. Similar cells Paul found in
of the enamel organ.
Inside this
the external epithelium
the
between
it
and
prisms,there appears to be
structure,
3. Where

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THE
a

TISSUES

53

of the ends of the

layerwhich does receive impressions

as though everyone
prisms. It seems
Waldeyer,certainly
; the Tomes
way.

had

been

rightin a

also,because

the

tissue is continuous with cementum,

and the outer portion
of the enamel organ is also continuous with the follicle
wall ; and those who described the pitting
of the under
surface by the prism ends and stained it with nitrate of
as
silver,
this,too, is confirmed by the research above
quoted. At any rate, happilyfor the student,the fact
that Nasmyth's membrane
is mainly composed of the
cells of the outer part of the enamel organ is
epithelial
and everyone
else, and for this
accepted by Tomes
definite state of affairs we are indebted to Dr. Paul.
Cementum."
At the neck of the tooth, the thin layer
of which I have justwritten,
where itoverlies the cervical
edge of the enamel,graduallybecomes thicker,and in its
for them) a
substance appear
(as soon as there is room
few lacunae
that is,
spaces in the calcified matrix which
in life contain each a nucleated cell. These
cells communicate
of branches,
with neighbouringcells by means
and when calcificationhardens all the matrix round them
communicatin
and imprisonsthe cells,
it does not obliterate the interBut
these latter persistin
branches.
which
are
great numbers, occupying minute channels
called canaliculi. So far the tissue resembles bone, in
which
also the cells are
encased
togetherwith their
In both tissues these contents
anastomozingbranches.
of the lacunae and canaliculi are easilydemonstrable, as
be
with gold chloride and can
they stain beautifully
examined
with low powers.* Cementum
differs from
ordinarybone in the following
respects :
vessels,
In
its
normal
it
is
traversed by bloodstate
1.
never
"and when a bloodvessel is present the occurrence
is pathological,
and there is no
specialarrangement of
the lacunar system round
Such
the vessel.
irregular
vessels occur
in abnormally thickened
but
cementum,
be mistaken for the regularand systematized
can
never
vascular arrangement of bone.
lacunae of cementum
2. The
are
larger and more
"

that

The
the

remained

which
in certain reliable text-books,
appears
have
of
these
is
not
nature
contents
yet decided, must
from earlier editions by inadvertence.
statement

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54

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DENTAL

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ANATOMY

the
those of bone.
They often assume
wildest shapes,
in hypertrophiedcementum.
especially
and irregular,
Their canaliculi are also very coarse
times
somerushingoff from one side of the lacuna in crowds,
instead of lookingas a bone-lacuna does" like an
so that,
insect with ten or twelve legs the object
rather resembles
that is being squeezed,and
from which
the
a
sponge
is rushing off on one
I possess a
side in streams.
water
specimen of a lion's cementum, in which the lacunae are
far apart, and the canals unitingthem
so
so
straight,
substance
that the intervening
is indistinguishable
from
dentine.
Mr. Hopewell Smith considers that normal cementum
I do not know
does not contain lacunae.
of any other
observer who shares this view.
is structureless.
The
outermost
layer of cementum
is exceedingly liable to increase with advancing
Cementum
additional
if
one
layersbeing plastered
on,
age,
such
be
This
condition
will
cussed
disuse
an
expression.
may
when treating
of the changes effected in the dental
tissues during advancingage.
Sometimes
found contained in a kind
a few lacunae are
described by Tomes
of envelope; these
as
enare
capsuled lacunae, and defined by him as 'individual
osteoblasts
with a connective-tissue envelope, which
to
some
have,
during
extent, preservedtheir individuality
calcification.'
It is importantto remember
that the lacunal cells all
is
a
nd
that
therefore the cementum
intercommunicate,
than
irregular

"

entirelypermeated by

network,the
livingprotoplasmic
have been supposed to be in

terminal fibrilsof which

direct continuity
with the terminations of the dentinal
the
side
fibrilson
and with the nervous
one
supply of the
the anatomy
of
dental periosteumon the other. When
the other calcified tissues has been considered,it will be
have a great
that this protoplasmic
network
seen
may
which
shall
I
endeavour
to clearly
lay before
significance,
my readers in its proper place.
When
is young,
cementum
globularforms may be
in
thin it is apparently
its substance,and when
traced
structureless.
The
lacunae

are

arranged more

or

less in rows,

by

and

THE

TISSUES

55

sometimes
far apart. In the before-mentioned
these rows
are
section of a lion'stooth the rows of lacunae are so far
them so straight,
apart, and the canaliculi which connect
viewed
with
(one quarter),
a moderately high power
that,
of
it is possibleto isolate a patch between
two
rows
It is
dentine.
lacunae that is indistinguishable
from
ing,
containcanaliculi
that
lacunae
obvious
and
these
fairly
as
do,
they
protoplasm during life represent the
softer elements
of the tissue,and that therefore where
they abound the whole tissue should be comparatively
easy to cut, and vice versd. This, however, is by no
*
of the outer
means
always the case.
Ivory' exostoses
the hardest tissues
of the ear, which are among
meatus
in the body, ranking next to enamel
in this respect, are
exostoses
abundantly traversed by lacunae,and many
almost
which resisted the section-grinding
as
process
such
full of
much
have proved microscopically
as enamel
the degree
This would seem
indicate
that
structures.
to
of calcification of the matrix is a very variable quantity,
the
hard exostoses
and
that in these preternaturally
of
balance
counterthan
lime-salt in the matrix may more
excess
tissue throughthe plentiful
distribution of living
out
the mass.
is the hardest tissue in the human
Enamel."
Enamel
it is (with the exception
ing
of the thin coatbody. In man
alreadyalluded to as Nasmyth's membrane) the outer
of the tooth. When, however, the
coveringof the crown
of the tooth,as is
is continued over the crown
cementum
lies between
in certain animals, the enamel
the case
it and the dentine.
the cusps, and
It is thickest over
thinnest at the necks of the teeth. Its chemical
position
comhas alreadybeen discussed.
The views as expressedin the paper communicated
by
Mr. Wellings and myself to the International Medical
the results of a prolonged
Congress, 1913, embodied
and may be appropriately
series of observations,
quoted
here :
'
of calcified
consists of rods or columns
Adult enamel
intercellular
cement
in a calcified
material embedded
in section
substance.
The intercolumnar substance seen
about one-eighth
to one-tenth the thickness
appears roughly
that nearest
of the column.
The first-formed enamel"/.*.,
"

"

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AIDS

56

TO

DENTAL

ANATOMY

later ;
than that formed
is more
irregular
and though it is difficultto be dogmaticon the subject,
it would appear
that the rods begin with tapered ends,
and do not afterwards
but soon
a uniform thickness,
assume
appreciablyvary in thickness throughout their course.
We have never
a column
seen
givingoff branches. Very
earlysections of ameloblasts at the apex of the toothfavour this view,the cells themselves beingtapered
crown
in section
towards the dentine. This is well marked
and nigrosin.
stained with malachite green, fuchsin,
4
Obviouslythe number of the rods at the outside is
rods
new
greater than at the inside surface ; when
is taperedto a fine point.
appear, their commencement
This might be due to the gradualcoming into focus of
the dentine

to

"

waving rod, but

start

of

we

rod.
(stainablewith
a

new

edges
method), as we shall
stainable margins.

have
never
The
rods

seen

square-ended

have
strongly marked
nitrate of silver,Bielskowsky's
have also
the ameloblasts
see

these striations
rods are
transverselystriated,
coincidingand traversingseveral columns, but not
the intercolumnar spaces.
traversing
*
If this complicated
were
structure
inorganic,
absolutely
in a strong acid would
the effect of prolongedimmersion
be to destroyit altogether.
We
submitted a thin section of enamel for twenty-six
hours to the action of strong acetic acid. The acid was
it
introduced under
as
the cover-glassand renewed
evaporated,the specimen being under observation the
whole time.
4
The experimentwas
started at 9.55 p.m. on April26.

|The

The edges
had become
plainer.
10.15 the structure
of the columns
very distinct. A clean layerbegan to
appear inside the edges of the columns, as ifthe part of
the edge had been clarified.
the column nearest
4
acid was
11.27 P-m. no
great change ; about 2 a.m.
4

At

renewed.
1
At 9
acid was
at

the 27th the acid had

The structures
added.
were

a.m.

evaporated. Fresh
still quitedistinct
added at intervals,

fresh acid having been

stillquiteobvious.
was
the transverse
midnightno change had taken place,

4-35 P"m"
the structure

By

on

by

58

AIDS

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DENTAL

ANATOMY

cells,
singlecells appear to contain one, two, or even
three,nuclei apparentlypassingalongthe cell body.
Sometimes
in the ameloblast
there are
areas
layer
*

where

it is

impossibleto

but collections of
define cells,
wandering nuclei appear to be shut off at intervals by
filaments of tissue like cell wall material.
*
The ameloblast layerappears to be separatedon the
outside from the stratum
intermedium, and on the inside
from
the forming enamel, by an
continuous
almost
is
in a very earlystage, before any enamel
membrane;
alone is stained. The
formed, the internal membrane
cells themselves have stainable envelopes.
*
The intervals between the cells of the ameloblast layer
where these cells appear in a definite row
to correspond
seem
in diameter with the intercolumnar spaces in the
formed material.
'
of the
It would
appear that the result of the activity
ameloblast
layeris the formation firstof a honeycomb of
intercolumnar substance into the spaces of which something,
of
of
consisting rounded bodies
varying sizes,is
poured : these bodies are no doubt calcospherites.
*
In some
sections the ameloblast layer is apparently
the
dragged away from the forming enamel, and across
interval fibrous strands interspersed
with calcospherites
These
sections have
to
cross.
plainlybeen
appear
and the appearance
well be an artifact.
distorted,
may
4
We
do not wish to form conclusions at this stage of
the investigation
would preferto submit facts and
we
let the theories come
later ; but it would
able
appear reasonin the
that the intercellular substance
to suppose
with the
ameloblast layercorresponding
in measurement
intercolumnar substance in the formed enamel, that the
intercolumnar substance is the result of a change in the
the result of
and
intercellular substance
the columns
the productof the cell activity
being depositedin the
"

spaces
It is

so^ormed.'

than allude to the

and
of Heitzmann
views associated with the names
of
Bodecker.
described a network
These
observers
Such
enamel.
human
livingstainable matter traversing

scarcelynecessary

to

do

more

would be wholly inconsistent with chemical

and is not demon st rat able by any staining
analysis,
process.

network

by

THE

TISSUES

59

is convinced that enamel

is penetratedto a
by dentinal
very considerable extent,althoughirregularly,
outside the
and
is
tissue
that the
fibrils,
not
so
entirely
pale of nutrition as has been supposed. This penetration
is a regularphenomenon in some
marsupials.
Mr. Pickerill of Otago has drawn
attention to surface
human
in
demonstrable
enamel
by rubbing
inequalities
and then polishinglightly
the surface with graphite,
so
that the valleysare black and the mountain-tops white.
These black lines he calls .'imbrication lines,'
and believes
of the tissue to invasion by caries varies
that the liability
according to their thickness and frequency.He has
evidence of the existence of these lines on the under
seen
He proposes to divide
surface of Nasmyth,s membrane.
teeth,
accordingto their enamel surface arrangement, into
malacotic (soft)
and sclerotic (hard) types. The
meability
perthe
by stainingagents such as silver nitrate,
all
density,and hardness,
solubility,
vary accordingto
the proportionate
thickness of these lines. He believes
that it is possible
to influence the difference of hardness
in the tissue after birth. This author appears to consider
of enamel as pathological
(Cosmos, October,
pigmentation
which
view
would
to
seem
implyan appalling
i9i3j P* 973)i
degree of disease in the rodent family.
The
problem, which still remains a problem, that
arises from the obvious fact that the outer surface of the
is greater than the inner surface,has
been
enamel
It has been suggestedthat
explainedin various ways.
the columns giveoff branches,it has been suggestedthat
supplementalor extra columns are formed to fillthe gaps
(Hopewell Smith), and it has been suggestedthat the
columns
increase in width towards the surface (Pickerill).
Mr. Pickerill has measured
the columns,and has satisfied
himself that he has detected a sufficient widening in
each
for the required
individual column
account
to
increase of space.
I have never
seen
a column
giving
has
far
I am
off branches,nor, as
as
aware,
any other
It is unwise to be dogmatic about
observer.
mental
supplecolumns, but this appears to me less difficultof
belief than the other hypotheses. The increasing
width
of the columns towards the surface,according
to Pickerill,
difference of 0*0025 mm.
at the ameloto a
amounts

Mummery

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ANATOMY

DENTAL

dentinal junction,
and 0*0065mmsurface.
at tne enamel
Kolliker givesthe figures
as 0*0051 and 0*0064 respectively,
and Tomes
and Hopewell Smith figurethe difference at
These
do not agree, and no
observations,plainly,
0*005.
doubt the wisest course
for the student to pursue is to
the difference of opinionand wait for more
remember
lighton the subject.
At present, therefore,
in the absence of more
definite
wisest to refrain from dogmatic
evidence, it seems
to me
statements
about the nature
of the component parts of the
tissue in question,
and to content
ourselves with describing
its appearance
under the microscope.
Enamel
consists of prismsor rods,the average diameter
of which is about -^^ of an inch ; they vary in thickness a
section. They do not touch
good deal,even in the same
each other,but are separated
about fo to J
by an interval,
the thickness of a prism,from one
another.
In transverse
section the prisms appear sometimes
hexagonal,quadrangular,triangularsometimes, and it
The
oval or round.
much more
to me
seems
frequently,
generalappearance under a low power isalwayshexagonal,
In
but so is the appearance
of circles in juxtaposition.
marked.
the rat the angular form of the prismsis strongly
Each prism is transversely
these
striated ; sometimes
striations are
but
sometimes
imperceptible,
very faint,
generallypretty plain,and in defective tissue specially
when observed under a high
pronounced. They consist,
the whole prism
power, of small spots or granules that is,
at the points
so
beingslightly
granular,it is much more
of striation and the granulesformingthe stripes
are larger
and coarser
than those in the intervening
layers. These
do not alwayscross
the prismsat right
stripes
angles,but
tions
are
obliquein direction ; neither do the striafrequently
of
its
of one
with
coincide
those
prism necessarily
though they often do so, and Leon Williams
neighbours,
thinks theyusuallydo.
it appears as though
Sometimes
tfre stripes
did not quitecross the prisms. These markings
and lie close together,somewhat
after the
are
narrow,
of the striations in muscular
fibre. As far as I
manner
of
can
discover,
they have no relation to any varicosity
the prisms,as has been suggested by some
observers.
It may be added here that some
of the acceptedfigures
"

"

"

by

of enamel

61

TISSUES

THE

prisms,which

are

text-book to
whatever
resemblance

copied from

bear no
itselfas seen
under the microscope.
to the structure
and Retzius describe the striation as due to
Tomes
of the prisms; Kolliker and Waldeyer consider
varicosity
of
is produced by mutual
that this varicosity
pressure
thinks the stripes
indications
the fibres. Hannover
are
the
the
successive
formation
of the
of
prisms;
parts of
striation
a
nd
Hertz is of the same
the
attributes
opinion,
to intermittent calcification.
Micro-photographyhas made great strides since the
above
lines were
and Mr. Leon Williams has
written,
untouched
figuresof enamel
publishedsome
exquisite
under highand low powers which seem
to leave little
doubt
that there are varicosities in the prisms,and that these
varicosities do coincide with the striations. These effects
are
prominenceby washing the tissue
broughtinto special
with dilute hydrochloric
acid,and if this be so we are
whatever the chemical composition
forced to conclude that,
of enamel
of the prismsis not
as a whole, the composition
identical with that of the interprismatic
substance,seeing
know
tends to remove
limethat the solvent which we
salts brings out the demarcation
of the structures.
If
there is no organicmatter
present at all they cannot
vary
in respect of its amount, therefore they can
only vary
of water
either in the amount
they contain,or in the
The varicosities in
varietyof the lime-salts themselves.
close
Mr. Williams' photographsdo not appear to me
so
familiar with,
as the striations I am
togetheror so narrow
but I cannot
at any rate
any longer doubt their existence,
decalcified. The striations
when the tissue has been slightly
of muscular
fibre do not reallyafford a reliable
and though these latter appearances
have been
parallel,
and thereshown
fore
to leave their impression
upon collodion,
be
due
than
rather
well
to shape
to
or as
as structure,
be
must
to
not
too much
weight
given analogiesbetween
contractile and wholly calcified tissues.
of
Course of the Fibres. In human
enamel the course
the prisms varies immensely, sometimes
being nearly
of the
outside
from
the
the
dentine
surface
to
straight
and curl
the other hand, they wave
tooth ; sometimes,
on
round each other in apparent confusion,and it is in the

text-book,seem

to me

to

"

by

62

AIDS

TO

curly parts that

most

ANATOMY

DENTAL
the

stripingis

transverse

most

observable.
clearly
of the fibres varies considerably
The course
throughout
the animal kingdom ; a few of the most
notable varieties
are

subjoined.
Manatee, the fibres are

straight.

for pardefinite and constant

are
ticular
families. In this order the enamel
is generally
divided into an inner and an outer layer(except
in hares and
in longitudinal
as
seen
section,the
rabbits). In Sciuridce^
outwards for about " of the thickness
prismsgo straight
of the tissue,and then turn abruptlyupwards at an angle
of 459. Seen in transverse
the fibres of alternate
section,
layerstake opposite
directions,
givinga sort of * criss-cross
to that portion of the tissue,
because
appearance
any
consist of more
than one
section,however
thin,must
layer of prisms; if it were
possibleto isolate one
would
be seen.
not
layer,the criss-cross appearance

Rodents,the patterns

'

In the outer

outwards.

third,they all take

the same
course, straight
In the porcupine,
the prismsare
individually

and cross
each other throughout about " of the
flexuous,
inner part of the enamel
(thatnext the dentine),while
towards the outside they run
parallel.Leporidce(hares
and rabbits)do not possess
two
layers; the fibres are
wavy.
The

enamel
surface of human
is often traversed
by striations that are visible to the naked eye; these
attributed by Tomes
are
to the outcrop of the fibres
that is,that some
layersof fibres stick out further than
others.
Since the outer surface of the enamel
is greater in extent
than the inner,and the fibres are not thicker nor further
be more
apart from each other,it follows that there must
of them.
I think the most
of this
probableexplanation
is that the fibres split
into
But
I
branches.
cannot
at
up
feel
all
I
about
have
at
sure
this,
seen
though
present
indicate
that
that it is so ; it is
to
seem
appearances
generallybelieved by the best authorities that the prisms
do not branch.
If enamel
is broken,the line of fracture is said to run
through the axis of the prisms,and not between them.
And
this is explainedon the ground that the axial portion
outer

"

by

THE

63

TISSUES

is the first to yield.

calcify,
This comparativesoftness of the axial portionof the
prisms is illustrated by Tomes, on page 56, Fig. 22
fenestrated
where
a
('Dental Anatomy,' third edition),

of each

fibre,being the last

to

in a transverse
section of enamel,
is shown
appearance
thinks the
treated with dilute hydrochloric
acid. Tomes
of
of
due
the
axial
is
to the removal
portions
appearance
the prisms. It looks to me
like the result of a total
removal of the prisms,the interstitialsubstance
being

left.
Striae of Retzius-^Very
often longitudinal
Brown
in
sections of,
enamel show dark lines,runningsomewhat
of the tooth, and therefore
the direction of the contour
the prisms. These
lines vary, both in number,
across
and thickness ; but they are of a brown colour,
frequency,
and are called,after the distinguished
anatomist who first
described them, the brown
striaeof Retzius. The appearance
is generallyascribed to the effects of intermittent
calcification that is,that enamel
calcification not being
continuous
but
a
an
interrupted
process, the pauses are
indicated by these markings.
After some
of the * lines,'
years of careful investigation
I feel convinced that they are due to an excessive granularity
of the prisms,and that,this granularity
in each
with that of its neighbours,the effect of
prism coinciding
result of my observations in this
a line is produced. The
direction was
fullylaid before the Odontological
Society
in January, 1888.
Enamel
Spindles. Sometimes
long irregular
cavities,
in
with
communication
the
of the
terminations
apparently
be observed
dentine fibrils,
in that portionof the
may
enamel
immediatelyoverlyingthe apex of the dentine
of these bodies has not been decided ;
The exact
nature
the direction of their long axes
is the same
as that of the
prisms. Klein calls them interglobular
spaces.
Pigment is sometimes present in enamel, notably on
the incisors of some
rodents.
It has been suggested that
this is due to direct staining
of the tissue by substances
which the animals feed.
It is present, however, in the
on
sockets of the teeth of many rodents (beaver,
capybara,etc.)
In sargus, the sheep's-headfish,
is traversed
the enamel
by tubes opening on the outside.
"

"

by

AIDS

64

TO

DENTAL

ANATOMY

is penetratedby
certain marsupials the enamel
fibrilscontinuous with those of the dentine.
varieties of tubed
describes and figures
Tomes
many
ture
enamels, and sees both in their development and structissue. In
the key to the proper study of the human
In

in the
fish,especially

shark

family,spherodus,
galeus,the porbeagleshark, cestracion (theonly survivor
all present systems
of a once
widely-distributed
family),
doubt is
of tubes in the enamel, but at present a slight
in
whether
all these structures
are
expressedby Tomes
many

enamels.
reality

Dentine.

is the tissue which reallymakes

up
If everything
else but the dentine
the bulk of the tooth.
taken away from a tooth, the tooth would stillretain
were
so

much

"

Dentine

of its

generalshape

and

appearance

as

to

be

recognizable.
much in its structure
that it has been
definitely
grouped into several varieties. I shall beginby
Dentine

varies

so

hard or unvascular
describing
the human
subject.

dentine,such

as

is found

in

dentine consists of a calcified matrix,permeated

channels,
throughoutby branching and intercommunicating
contains a soft protoplasmic
fibril.
each of which
Mr. Howard
Mummery's recent researches (Philosophical
Transactions, 19 12)have demonstrated
bility
beyond any possiof questionthe fact that the fibrilsare accompanied
the plexus of
filaments given off from
by fine nerve
Hard

permeating the dentine throughout.

has
this demonstration
After much
searchingcriticism,
and
been practically
universally
acceptedby histologists,
must
now
teaching. Mr. Mummery
pass into current
terminations
in staining
has succeeded
these fine nerve
Raschkow

and

which
nitrate of silver may
with many
reagents, among
be mentioned
as
having given very convincingresults.
of dentine is now
cleared
The problem of the sensibility
up, and is no longera problem. There are generallytwo
fibril. Cajal'ssilver nitrate
filaments with each
nerve
successful
method
stain, and Ranvier's
as
a
was
very
modification of Lowit's formic acid process was
employed
for gold impregnations.
In any
meet
section,therefore,
we
and
(b)tubes,(c)fibrils,
(d) nerves
;

with
much

so

by

(a) matrix,
everyone

66

means
common

AIDS

ANATOMY

DENTAL

TO

it is so, it is more
than
which underlies cementum
It consists of a row
underlies enamel.

always present,

and

when

in the dentine

in that which
of small interglobular
not, and have
spaces ; they are
with,lacunae,but under a low power
nothing in common
like
look
them.
rather
common
They are excessively
they
in certain cetaceans, where there exist sometimes
not one
It is important
but several of such rows.
not to confuse
and it will be well for every
these spaces with lacunas,
student to examine
under a high power,
them carefully
their real nature will become apparent. The space
when
occupiedby a cell must, in a fresh specimen,contain the
and from this cell,
an obvious and stainable
cell,
possessing
nucleus, branches proceedin many directions,
connecting
the cell with its neighbours. The
interglobular
space
contains no cell and no nucleus ; it is simplya spot where
failure on the part of the calcospherites
to entirely
a
in
coalesce has resulted
outlines.
a hiatus with rounded
heated
If a million round lead bullets in an iron box were
the resulting
be solid
would
mass
up to melting-point,
If the meltingprocess stoppedjustshort of comlead.
plete
there
would
be
of
fusion,
interglobular
spaces left,
but always with rounded
outlines. The
varying size,
be of all sizes to carry out the simile. This
bullets must
is the best I can
do to explain
an
interglobular
space.
This portionof the dentine is often extremelysensitive to
be removed
with an excavator
touch, so that it cannot
without causing considerable
pain ; but as soon as the
been
has
removed, the tissue underlying
layer
superficial
be found to
to the pulp,will frequently
it,though nearer
lower degree of sensibility.
exhibit a much
Another
function has been conjectured
to appertain
to
the dentinal fibrils,
and that is a trophicone
that is to
to the nourishment
manner
say, that theyminister in some
of the tissue which
is no doubt
There
they permeate.
that the division of certain nerve
trunks is followed by a
in
apparentlydue to impaired nutrition,
degeneration,
the peripheral
of
the
which
tract
portions
they supply.
Such changes have been demonstrated in the consequences
of operationson
spinalnerves, the Gasserian ganglion,
"

etc.;

moreover,

destruction,whether

the result of disease

"

or

pathological *'.*.,
"

as
accidental,

in

by

the

case

of

THE

TISSUES

67

blow, etc., or followingon the deliberate

of treatment
annihilation of the pulp in the course
of the
is
of
followed
central nervous
by changes
supply a tooth,

injuryfrom

in the nutrition of the organ.

This

undoubted

clinical
conditions

fact is further emphasized by the anatomical

observable in vaso-dentine.
Among fishes we find vasointermediate
dentine, hard or tubed dentine,and an
In the hake the dentine is ail vascular with
condition.
in the flounder tubed dentine becomes
vascular
no
fibrils,
and
varieties
the pulp chamber,
are
common
near
fairly
find
both
fishes.
But
never
we
trophicapparatus
among
fades the
tooth in full development. As one
in the same
that we are
other grows.
May we not therefore assume
of
a
nd
schemes
that
two
nourishment,
considering
Nature,
refuses
bent
on
to
economizingenergy,
employ
always
forces to do the same
two
thing? It is the old story of
the horns and the canine teeth over
again. It would be
of energy to encourage
two
concurrent
Nature
the
handiest.
of irrigation,
so
employs
the tooth of the flounder,the tip consists of
a

waste

systems

Thus* in

ordinary

dentine ; a littlelower down

the tubes
hard,thickly-tubed
wider
here
while
fewer and
and
become
there
apart,
Stillnearer
vessel appears.
to the base of the tooth the
while the vascular network
fibrils disappearaltogether,
dense
and
thick.
becomes
In the hake, the
of capillaries
tissue is freelyvascular throughout,but destitute of fibrils.
As a generalstatement, it may be said with truth that the
of fibrilsin dentine is in inverse
frequencyand regularity
of capillaries,
ratio to the frequencyand regularity
and it
deduction from these observations that
seems
a justifiable
the same
both are
ends in the
extent
servingto some
and
that
of
the
the
physiology
part,
presence of both in a
in
of
the
tract of tissue would
state
same
efficiency
high
be a redundancy and a solecism,
not in accordance
with
that
almost always to result
the laws of economy
seem
from the workingsof evolution.
fibrils taper towards
the outside of the dentine,
The
and give off throughout their course
fine inosculating
These
branches
branches.
are
more
the
numerous
as
distant from the pulp, and
fibrilbecomes
in the
more
ingly
portionsof the tissue the network is exceedperipheral
fine,and no doubt better instruments will soon
a

by

demonstrate

ANATOMY

DENTAL

TO

AIDS

68

stillmore

delicate system of ramification.

the older writers that the fibrillar

It was
supposed by
elements did not penetrate to the actual outer
margin of
anatomists are agreed that they do.
but modern
the dentine,
These
Dentinal Tubes.
are
simply the holes which
and in life are
contain the fibrils,
quitefilled by them.
if
After death, especially astringent
reagents have been
"

employed, the

soft tissues shrink

somewhat, and

times
some-

less
or
coagulatedinto a series of more
Under
these circumstances
interval
an
between
the
contained soft tissue and
may be perceived
filled.
the tube which it once

become
elongated dots.

the changes due to

for much
erroneous
reagents employed are responsible
the
safest fixative
and misleading histology.Perhaps
These

changes

post-mortem

and

available is formalin.
and the effect of the

heat,
Decalcifying
agents, alcohol,
all
of
are
sources
razor,
possible
this subject
Mr. Wellingswriting
on
error.
says :
are
Agents. These
always likelyto
'Decalcifying
of
the
and
to bring about
tissues,
produce swelling
affect
chemical changes in the tissues that must seriously
the reaction of staining
reagents. Very earlycalf embryos
"

be cut without tion.

decalcificaconsiderable
of
a quite
amount
germs
be cut at once.
When
formed dentine and enamel
it
can
have found that,in our
to employ acids, we
is necessary
solution is 5 per cent,
the most
hands at least,
satisfactory
formalin.
in
acid
A largeamount
cent,
1 per
hydrochloric
and

rats

and

mice

at birth can

with

Tooth

of the fluid was

found
necessary.

used,and

changed

this way
attained without
In

as
was
frequently
calcificati
dequite satisfactory
as

interference with
was
any
and
the
with
minimum
fixation
of swelling.
the original
'
Alcohol frequentlycauses
shrinkage and distortion
through too rapid diffusion. To control this effect we
of impregnationthat do not
employed methods
of
alcohol.
Gum
dextrine mass,
mucilage,
requirethe use
tried and
found
each
serviceable in
and gelatinwere
with the Aschoff C02 freezing
microtome.
conjunction
'
The
temperature of the paraffinbath is also the
distortion from shrinkage. In addition
of much
source

have

to

this,the effect of

decalcified bone

and

temperature above
teeth

is

to

harden

400C.
them

by

on

so

adult
that

THE

TISSUES

69

it has been necessary

to
cuttingis impossible.Where
in the vital staining
for instance,
examine such tissues (as,
have
been
without
cut
embedding or
they
experiments),
with gum mucilage,
after impregnation
or gelatin
dextrine,
;
of sections through the entire skull)
or (inthe case
they

in celloidin.
been embedded
distortion
due
'The
it is impossibleto
to the razor
of qualityand
avoid. Every precautionin the matter
sharpness of instruments,
etc., has been taken,but there
is always a liability
to the productionof artifacts where
knife
has
the
to pass through tissues of different densities
the section.' *
in cutting
have

of the tubes is,naturally,

the course
of the
and varies from beingperfectly
to curving
fibrils,
straight
Two
have been
like a bent corkscrew.
sorts of curves
which
in
the
consist
described,
one
primary curves^
sweep
like the italic letter /, stretching
somewhat
the whole
these are
more
length of the tubes
easilyobserved in
The

course

"

the

of the tooth ; and

undulations of

crown

the

secondarycurves,

which

less corkscrew
or
are
a
more
in the root tissue.
and are better demonstrated
character,
the primary curves
When
perform a sudden wrigglein
smaller

their course,
and these wrigglesof neighbouringtubes
coincide
in the same
i.e.,occur
plane the result is an
of a line running at right angles to the
appearance
of the tubes.
had been
It is as ifsomeone
generalcourse
and had had his
tryingto draw a series of parallellines,
elbow jogged midway in each line,and the coincidence
"

"

of the deviations resulted in a line more

or less at right
lines
of the fibrils. These
angles to the general course
called the contour
and are defined
lines of Schreger^
are
due
the
of
coincidence
the primarycurves.
to
as
include both these and the lines
The
lines of Owen
in certain
formed
by rows of interglobular
spaces common
The openingof each tube towards
the pulp
called
the
is
lumen.
sheaths of Neumann^
the existence of which
is
The
denied by Magitot and Kolliker, consist,accordingto
and others,of the semi-calcified layer of matrix
Tomes
the tubes.
which
Being of the
immediatelysurrounds
cetaceans.

of

nature
*

they
calcoglobulin,

Transactions, International

exhibit all the indestructiMedical

Congress, 1913.

by

AIDS

70

bilityof

TO

that

ANATOMY

DENTAL

Although they can best be

of the tissue,
they may
be observed
in transverse
sections of incipient
caries,
when
they remain of a faint yellow colour,all the rest
of the tissue being stained with an aniline dye. They
have been described
as
possessinga double contour
that is,an observable
limit outside and inside. But this
demonstrated

substance.

by decalcification

"

of the section
the upper and
lower surfaces of the slice the one
appearing to be
outside and
distinct from
Howard
the other.
Mr.
has
demonstrated
these
sheaths
that
Mummery
recently
do unquestionably
structures.
exisi-"e-separate
be due to the thickness
appearance
may
showing the inside margin twice /.*?.,on
"

"

'

Vascular patches1in human

dentine
stated.
I have
rare
as is sometimes

by no means
already collected

are

specimens, and
many
in
the arrangement
peculiarities

have
observed
some
in
of the dentine
their
immediate neighbourhood,which have not, as far as I
The
know, been described by other authors.
general
under
of a vascular patch in
low power
a
appearance
human
dentine stronglyresembles
that of a knot in a
eat

pieceof

wood.

The

the
tubes,as they approach

vascular

canal,deflect in all directions,crowding togetheras if

to

give the objecta wide berth,and after having passed it

their previousrelative positions.At first
they resume
the
of the section suggests that the
sight
appearance
small area
round
the vessel is destitute of tubes, but
minuter investigation
reveals a very fine system of tubes,
from the vascular centre to the periphery
of the
radiating
thus
additional
for
reason
an
patch/
supposing
affording
that the fibrillarsystem
has some
trophicbusiness to
in the
vascular
perform. These
patches are normal
the
is
dentine
inner third of whose
walrus,
entirely
composed of them.
Mummery holds
Touching the termination of the fibrils,
that the dentine is everywhere permeated by very fine
branches. Hopewell Smith and Tomes
both describe these
branchingsas more frequentin the root than in the crown
of the tooth.
more
Mummery
readily
says they are
*

stained
'At

the

in the

margin

loopsand

very

root, but not

of the cementum

more

numerous.

He

says

fibrilsterminate in
v
isible
delicate,
branches,many
scarcely
the

by

THE

TISSUES

71

of which enter the granular layer and are

to their extremities.
Under
the enamel

finelybeaded

of the
many
tubules are much
coarser, and some
appear truncated
if there had been
alternation of absorptionand
as
an
deposition
during the firststages of development of the
tissues. Many tubes cross the border line and terminate
in various ways in the enamel, some
like
by small round,knob.

bodies,some

by tinyflattened

while
expansions,

others
the prisms in fine whip-like
pass between
processes, but
I have not been able thus far to trace the fine beaded
fibres into the enamel.'
There are three recognized
varieties of dentine,besides
hard or unvascular
dentine
vaso*
namely, plicidentine,
"

and
dentine,

osteodentine.
Plicidentine differs from hard dentine in that, the pulp
from which it is formed
form, the
being of irregular
dentine has a doubled
in order to
folded appearance
or
fitinto the convolutions of the pulp. It is practically
the
difference between
the skin of a filbert and the skin of a
walnut
This
in the shape,of the pulp,
complication,
varies very much
in degree. Thus, in certain lizards

(varanus,tor
rather like the

example) the amount

foldingof the silk

of foldingis slight,
between
the ribs of a

extinct
an
half-openumbrella, while in labyrinthodon,
amphibian,it looks like a wild confusion of small pulpchambers
and systems of tubes surroundingthem.
Sometimes

the pulp recalls the shape of the tree

in the rays mylioas
commonly called a ' monkey puzzle,'
and zygobates,
and in the rostral teeth of
bates,aetobates,
the saw-fish.
pristis,
Vasodentine
hard dentine and plicidentine,
resembles
in so far as that it is,like them, developedfrom a row

in not
cells. It differs from plicidentine
varietiesin
being folded,and from both the above-named
less
or
by vascular
regularity
being traversed with more
and nothing
canals.
Each
canal contains one
capillary,
else ; and each capillary
fits and wholly fillsits canal.

of odontoblast

Vasodentine

varies much

in

vascularity.Sometimes,

as

hake, the vascular supplyis very rich, and then

The
two
seldom
are
any fibrils distinguishable.
designed
elements, fibrilsand capillaries,
being apparently
namely,
the same
functional end
and
to
serve
one
in the
there

"

by

AIDS

72

nutrition

"

tract

same

never

are

DENTAL

TO

ANATOMY

present, to any

of tissue.

great extent, in the

If the vessels

are

numerous

the

In the flounder the tip

fibrilsare absent, and vice versd.
tubed ; the harder varieties
is pure, hard dentine,finely
with in exposed situations,
such as the
tip and outside of the tooth. A littlelower the tubes
At the base
become
scanty, and a few vessels appear.
the tubes are absent and the capillary
system abundant.
The two forms of dentine present in this creature, as

always

are

met

intermediate
similar odontoblast

also

the

condition,are

obviously formed

cells. The
manatee, a Sirenian
enamel
animal,whose straight
prisms have already been
noticed,is possessedof vasodentine,as are the tapirand

by

extinct megatherium
Indian tapirhas vascular

the

(a giganticedentate).

The

of his
canals in the crowns
varietyin the roots.
teeth,and the American
In teeth of perpetual
growth, such as rodent incisors,
that
the dentine
occupiesthe situation of the obliterated
that
pulp is generallyvascular. It must be remembered
and hard dentine are found in the same
tooth
when
vaso
the hard is always in the exposed situations the tipand
"

the
tissue forming the central
and base.
core
hard
It will have been noticed that in human
/."?.,
vessels are
unvascular
an
dentine, when
present as
modifies
their
abnormal
phenomenon,
presence profoundly
of the tubes in their neighbourhood. In the
the course

outside

"

softer vascular

"

dentine of the

walrus,where

the presence

"

of these

canals
is not an abnormalitybut a constant
phenomenon, the
fibrillar
of
the
disturbance
same
system takes place,
of the fishes where
both fibrilsand
in the case
whereas
fluenced,
uninvessels are present the fibrils pursue their course
by the presence of the capillaries.
apparently,
rule
no
The
nerves
accompany
appears to be that when
the vessels the course of the neighbouringtubes is not
modified,but when nerves
go with the vessels (and the
the dentinal
and not vaso-dentine)
tissue is strictly
osteoand
with
direct
focussed
is
reference
to
arranged
system
the canals.
Osteodentine differs in a broad and essential way
from
the forms of dentine already mentioned.
It is not, like
hard dentine, plicidentine
and vasodentine,
the product

by

AIDS

74

TO

ANATOMY

DENTAL

observation,
though he may pass examinations,he
never
knowledge.
possess anythingworth calling
THE

UNCALCIFIED

DENTAL

can

TISSUES.

The Pulp. We
have alreadydiscussed the functions
of the pulp as a formative organ.
It remains
to consider
its anatomy
is
that
and
the
finished
now
development
remains
of the dentine germ
as the nutrient and
persist
sentient centre of the tooth.
The
pulp,which is sometimes looselyand erroneously
of vascular and nervous
spoken of as the nerve, is a mass
connective tissue,
situated in the centre, or
it
core, as
were, of the tooth, and roughly resemblingit in shape,
three
scale.
It is fed by two
or
only on a reduced
and
and
few
smaller
The
one
nerves.
a
arteries,
large
latter break up near
the surface into an
exceedinglyfine
plexuscalled the plexusof Raschkow.
Up to quiterecent
"

times various conjectureswere

for
advanced
to account
the final distribution of the nerve
tine
denfilaments.
That
sensitive
all
knew.
An
was
we
exposed surface of
dentine at some
distance from the pulp cavitycould be
shown

to be

not
quitecapable,at times, of appreciating

only heat

but the
and cold and the touch of instruments,
This
sensation
of
acids.
of
could
presence
conveyance
filaments
due
be
either
of
the
to
nerve
or
only
presence
the presence
of nerve
endings. I think most observers
inclined with Hopewell Smith to the conclusion that the
fibrils were
in themselves
nerve
endings. I certainly
than forty years ago,
thought so myself. Boll,more
blasts
thoughthe saw fine filaments passingbetween the odontoas

far

as

theidentine; and

when

the

fibrils were

filaments
the nerve
seemed
to accompany
which
he
concluded
from
fact
that
them,
they penetrated
with the fibrils,
but he did not
the dentine in company
Klein (1883)
them in the hard substance.
see
guessed that
cells and
The fact that all nerve
this might be the case.
derived from the epiblast
fibres are originally
(Schafer)
conclusive
the
odontoblasts
a
to
seems
objection

pulledout,

(mesoblastic)
being nerve
endings. Weil, Magitot,and
others,thought the objectionnot insuperable
; Tomes

thoughtit was

almost

so.

by

THE
To

Mr.

TISSUES

75

belongs the glory of demonstrating

Mummery

filaments
these fine non-medullated
accompanying the
fibrils throughout the dentine.
As Tomes
pointed out,
the easier a tissue is to examine, the finer the nerve

distribution observable.
in the

of

case

difficultiesof observation
calcified tissue like dentine no
doubt

The

lor the delay of this discovery.

basal layer of Weil
is a clear space in the pulp
the odontoblasts.
vessels
bloodimmediately beneath
Many
in this layer. Mummery
found
stained it
are

account

The

gold chloride,and

with
fine

the space

by
basal
a
really
layerat

have
not
yet been
difficult to believe
will doubtless
methods

point.

The

pulp

is it

all.

Lymphatics
pulp,but it is
Modern

filled with
and it is
fibres,
to be

connective-tissue
distortion in section-making,
nor

nerve

caused

not

found

fibrilsand

in

demonstrated
that they are

the
absent.
lightupon this

throw
which
of
largely cells,
round
oval nucleated
or

consists

for
bodies.
are

the most
part simple
Those
the surface are, however, somewhat
very near
in
specialized form,tendingrather to the columnar shape ;
with the
the cells actually
the surface and in contact
on
'

calcified dentine

elongated,and

because
when
the pulpis torn out of the tooth these cells adhere to the
mistaken
dentine, they were
by the earlier writers for a
*
membrane
and called the membrana
eboris.' '1hey are

usuallycalled

are

the

very

odontoblast

cells,and

described

are

the dentine
(fibrils),
possessingprocesses towards
the cells underlyingthem, and
towards
towards
each
I have not been successful in demonstratingall
other.
as

these

but

there seems
to be no
reason
all
the
exist.
cells
of the
not
Possibly
by branches with their neighbours.

processes,

they should
inosculate
I have

miniature
the

case.

said

that the shape of the pulp is a rough

of the shape of the tooth,and this is generally
But it sometimes
happens that the pulp gives

off offshoots,
more

with no referless fine diverticula,

ence
of the tooth.
I have met with several
fine filaments of pulp tissue have shot
or

to the contour

instances
from
out

why
pulp

in which

the confines of the pulp as far

of
the enamel, carryingwith them
surface

This

is

a most

the

as

acute

and
puzzlingabnormality,

by

under
bility.
sensiresults

TO

AIDS

76

ANATOMY

DENTAL

the
in all the symptoms
of the pulp,when
of an exposure
not been exposed.
pulp proper has obviously
The
pulp undergoes considerable changes as age
advances.
If decay threatens it from any
quarter, the
formative
their
odontoblast
cells frequently resume
some
rough-and-ready dentine to
functions,and make
point. At first the
protect the pulp at the threatened
their work, and the resulting
cells seem
to have forgotten
globular
dentine is very rough and irregular,
with large interdescribed
often incorrectly
as
are
spaces which
back
to
but presentlythe old art seems
to come
lacunae,
is
and
the
dentine
result.
them,
Secondary
very passable
is always
like
normal
but
dentine is never
dentine,
quite
of interglobularspaces
characterized by the presence
and more
less sphericalforms.
It does not always
or
described
be
to
formed, as
above, to protect the
appear
pulp from the ravages of the caries,but is not unfrequentlythe result of a capriciousand mischievous
the part of the pulp cells,causing the
on
activity
phenomena known as pulp-stones,'and often resulting
*

in

violent

neuralgicpain,the

difficult

cause

of which

is obscure

trace, until the tooth has been removed

and the pulp examined.
It is exceedinglydifficult to
destroya pulp in which these bodies exist.
As
advances, the pulp undergoes changes, in
age
of secondary dentine,
addition to the encroachments
and

which

to

take place. These

changes are
due chiefly
to interference with the vascular
supply of
the organ.
In all arteries throughoutthe body there is a
tendency to degenerativechanges which render the
vessel walls thicker and less elastic,
and so diminish the
lumen.
The vessels may also become
completelyblocked
or

may

may

not

blood.
this clotting
results
When
which
the
channel
clot
of
with
a
a
in size,but does not
increases
move
along with the
and
the
blood-stream, the process is termed thrombosis,
in
This is more
veins than
clot a thrombus.
common
in arteries. When
of
clot
becomes
a piece
loose,and is
carried along in the circulation,
called an embolus.
it is.
This
most
(embolism) occurs
frequentlyin
process

by
in

the

of the
clotting
gradualfilling
up

and
arteries,

it reaches

the

embolus
passes
point where the lumen

along
is too

the artery until

small to allow

by

THE

of its passage.
blocked.

The

TISSUES
vessel

77
this

in

The degenerative
from
changes resulting
vascular supplyare of various kinds,viz. :

this deficient

fibrous,
fatty,

cellular elements in the pulp

diminished
in number, the whole mass
are
becoming
of
e
ither
and
mummified
transmitting
eventually
incapable
sensation or subserving
the function of nutrition.
of that
is the familiar designation
Gum.
The
gum
the
mouth
of
of
and submucous
portion the mucous
lining

mucoid,or

becomes

way

calcareous.

The

"

which
dense

the alveolus.
The submucous
layeris very
and closelyattached
to the bone
by tendinous
which are somewhat
the handle
of
fasciculi,
fan-shaped,
the fan being towards
The
is
continuous
the bone.
gum
with the membrane
The
which
envelops the roots.
membrane
consists of many
mucous
layersof squamous
epithelium.The gum, as a rule,is deficient in nerves
but rich in vessels ; it contains several small epithelial
covers

like abortive tooth-germs,

look very much
and are called the glands of Serres.
Nothing is known
of their function,
but Malassez
has described them as the
of
the
of
seat
commencement
growths.
epithelial
The
is the tissue which
Periodontal
Membrane
intervenes between the tooth and the bony socket.
At
but this
time it was
one
supposedto consist of two layers,
view has been abandoned
since it has been demonstrated
that fibres can
be traced continuouslyfrom the bone to
the cementum.
The
cemental
part is less cellular and

nests, which

more

fibrous than the

bony portion.

itisvariously
later
names
on), consists of connective tissue,between the bundles of
which run groups of vessels and nerves, but it contains no
be stretched.
yellow elastic fibres, and therefore cannot
It is,however, necessary that the tooth should be capable
of some
in its socket,for every temporary
degree of movement
The
called

as
or ligament,
membrane, periosteum,
(I will discuss the proprietyof these
.

for the time being,the

inflammation,
by increasing,
of the part,necessitates the partial
extrusion
blood-supply
of the tooth from its socket.
The familiar feeling
so often
described by patients
longer than its
as of a tooth feeling
the
and meeting its opponent firstwhenever
neighbours,
jaw is closed,is due to such an extrusion. This movement

by

AIDS

78

TO

DENTAL

ANATOMY

the unyielding
of the tooth,without tearing
membrane, is
rendered
of the fibres.
possibleby a specialdisposition
of each band of fibres is nearer
the
The bony attachment
neck of the tooth than the cemental
attachment,so that
the tooth may
be regarded as suspended in the socket by
fasciculi of the periosteumrather than as restingupon
the membrane
some
; consequently,
playis allowed to the
be pushed out of its socket until the
tooth, which can
cemental
end of each bundle
of fibres is as
deeper or
much
above its bony attachment
as it normally is below
'
'
'
and
above
below ' must
it (the words
be considered
fibres
to apply to a lower tooth). Although the
tinuous,
conare
and there is no such thingas a demarcation
into
layers,there is a difference between the portionnearest
the cementum.
the bone and that nearest
At the bony
end they break
end the fibres are larger; at the cemental
in the latter situation the
up into a fine network, and
cellular. The tissue becomes
thinner
tissue is rather more
from
with age, being encroached
both sides,but
upon
it
f
rom
the
cemental
becomes
side;
never
especially
is very free,
obliterated. The blood-supply
entirely
being
derived from the vessels of the pulp,as well as those of
the cementum
there is a capillary
and bone ; near
the gum
'
midway
plexus. The bloodvessels are most numerous
the bone and cementum, or rather nearer
between
to the
'
latter (Tomes).
of this membrane
remains
The
exact
nature
to be
discussed.
It is most
commonly called the dental periosteum,
but Malassez {setJournal ofBritish Dental Association,
it
is
contends
that
not
a
1885,p. 484)
periosteum
but
1.

ligament,for
If

true

the

followingreasons

periosteumexisted in such a situation,

a very painful
process, owing to the

mastication would be
of nerves.
abundance

the tissue does

Microscopically,
rather
but
a ligament.
periosteum,
Mastication
not
producestraction,
3.
2.

not

resemble

pressure.
the
bone
and
attachment
is preThe
to
cementum
4.
cisely
like a tendinous insertion.
in the
distribution of the vessels and nerves
5. The
interstices between
bundles
of fibres suggests ligament
and not periosteum.

by

DEVELOPMENT

THE

THE

OF

MOUTH

79

similar view of the structure.

is directly
It is,however, true that the membrane
continuous
of
the
the
with the periosteum,
rest
covering

Ranvier and

Kolliker take

of mastication can be borne on a

bone
ridgesof jawas, for instance,in edentulous
periosteum,
i
s
it
that whatever
it is, certainly not quitea
; and
but a specially
typicalligamentor a typicalperiosteum,
maxilla

; that the shock

modified form.

CHAPTER

THE

DEVELOPMENT

OF

THE

MOUTH

the developmentof the human

Although
creature
must
be studied
elsewhere, it will be an advantage to the
student to present here a slight
sketch of those changes
in
which
and which
intra-uterine life,
lead to the
occur
of the scene
of his future operations the
formation
mouth.
It will be my endeavour
to introduce this subject
in as simpleand homely a fashion as I can.
At an early periodthe human
embryo is in shape not
unlike a small thin sausage, slightly
curled over
at one
jend. The curled end, which is bulkier than the rest, is
the head.
destined to become
From
each side of the
this
below
curled
end
be seen
body immediately
may
sproutingbuds, which advance and bend round as if to
each other and form arches. The front part of the end
meet
sends downwards
which is curlingover
a
process called
the nasofrontal process, which,as itstretches downwards,
givesoff two offshoots on each side,or four altogether.
the uppermost
lateral processes, called the
Meanwhile
maxillaryprocesses, coming forward and bending as if to
in the middle line,find themselves
unable to do so,
meet
the naso-front'l process has already projected
because
downwards
far as to find itself occupyingthe middle
so
should meet
if proline,where the maxillary
processes
longed.
It results,therefore,
that the forward movement
of upper
lateral processes
is arrested,and
the downward
themthe
naso-frontal
find
of
prolongations
process
"

by

80

AIDS

selves pinchedor
left upper lateral
the

ANATOMY

caught,as it were,

maxillary
processes
on

DENTAL

TO

or

between

the

rightor

maxillaryprocesses. Although the

quitemeet in front
cannot, therefore,

surface,they do

meet

in other words in
illustrate this with a
of your hand, with no

behind,or

the

deeper part of the face. To

homely simile,imagine the palm

the top part of the embryo ; bend forward

the palm to indicate the first curlingover
of the head;
then imagine the four fingerssprouting
downwards
the
as
naso-frontal process, firstalljoinedtogether,
then separate ;
time imaginea thumb
at the same
each
out on
sprouting
side,and curlinground as if to meet its oppositeneighbour
as
fingerson it,

until stopped by the tipof the forefinger

on
side and of the littlefingeron the other, these

the

one

thumbs
the
maxillary processes ; imagine the
represent
the
thumbs, though arrested by the intervening
on
fingers
behind
the fingers
outside,to continue their advance
and
actually meet;
lastly,imagine a second pair of
thumbs
time as the first pair but
the
a
t
same
starting
below
line as
the
them, and meeting in the middle
four fingers
The
mandibular
fusingtogether
processes.
in front,
middle
in
form
the
the
and
line,
pre-maxillary
which
middle
touch
bones ; the two
each other
ones
to

and littlefinger
form the endognathion; the forefinger
form the rightand left mesognathion,while the upper
form
the maxillae
thumbs
rightand left
proper, or
exognathion ; and the lower pair of thumbs, where
Meckel's
cartilagefirst appears, eventuallyform the
is
mandible, or lower jaw. The orifice of the mouth
the upper and lower pairsof thumbs
between
tween
*.*., bethe maxillaryand mandibular
"while
the
processes
in the space above
the first part of the
eye is formed
in cleft-palate
side.
The
cleft
each
on
was
upper thumb
the
be
between
to
and
the
endognathion
usuallysupposed
Professor
Arthur
Keith
has,
mesognathion
however,
while ihe separationin
that this is not the case
shown
"

"

"

of the right and

due to non-union
the hare
left
is
in
therefore
situated
and
the
middle
mesognathion,
line. To continue our metaphor, the upper incisor teeth
while the rest of the
formed
each finger-tip,
are
one
on
teeth
formed
in
the
are
upper pairof thumbs, and
upper
the whole of the lower teetb in the lower pairof thumbs.
is

izijby

82

AIDS

TO

DENTAL

ANATOMY

well illustrated in the growth

of the mandible.
In infancythe front teeth are high,the
back teeth few and low, and consequently the condyle
reaches the glenoidcavitywithout risingabove the level
When
the formation of the great
of the rest of the bone.
is called the antrum
of
air cell in the upper jaw, which
tooth bearing part of the
the molar
Highmore, causes

some

opposing surface,is

upper jaw to descend, and the growth and eruption of

the molar teeth separates the upper from the lower jaw, a
rapidupward growth in the bone supportingthe condyle
becomes
necessary to keep the articular surfaces in apposition
of
the
and
the
takes
thus
this
angle
jaw is
place,
;
formed
The

by a quickgrowth of the ascendingramus.

bony parts of the jaws may be roughly divided

two
portion,
parts : the alveolar or tooth-bearing
which
the
tooth
envelops
during development,
germs
and the roots of the teeth at maturity. This portion
exists only for the teeth,is formed
with the milk-teeth
and absorbed when
they are removed, and againformed
around
the permanent
by
teeth,and then removed
is
when
lost.
the
Then
there
are
absorption
they
body
of the
of the jaw,which exists and grows
independently
teeth,but is profoundlymodified in its anatomy by their
If we glance at an infant's lower
and specialization.
uses
jaw just before or during the eruptionof the first set,
and a senile edentulous
specimen from which all the
have
teeth
to
long disappeared,
they will at first seem
resemble
each other.
In both the angle is feeblymarked,
and the whole bone much
and thinner than in
straighter
notice the position
adult specimen; but if we
of the
an
where
the
inferior
dental
mental
bloodvessels
foramen,
from their canal underneath
and nerves
the teeth,
emerge
shall see that in the infant jaw it is near
the lower
we
margin of the bone, because the tooth-bearing
portionis
is
in the
but
the
whereas
not
body
there,
yet formed,
senile jaw it is almost on the upper surface,
edentulous

into

because
the tooth-bearing
portionhas been absorbed
after the loss of the teeth.
The
fact that the increase in size of the mandible
is
almost entirely
brought about by additions to its posterior

surface,and to the outside,as

which, as has been
extension,

the downward
as
is
evident from
seen,
well

by

THE

DEVELOPMENT

comparisonof

the

OF

THE

MOUTH

83

of
positions

the mental
in
foramen
relation to the upper and lower surfaces at various ages,
is demonstrated
The
by certain simple measurements.
second bicuspidsor pre-molarsoccupy
about the same
Let a line be
as the second
position
temporary molars.
drawn
from
the posterior
surface of the second temacross
porary
molar on the one
side to a similar point on the
other,and called AB ; bisect this line at C, then draw a line
forwards from C at rightangles to AB, tillit meets
the
the
of
and
left halves of the
right
symphysisor junction
mandible between the lower central teeth at D. Let similar
in the adult jaw, only AB
lines be drawn
unite the
must
surfaces of the lower second bicuspids.Measure
posterior
these lines,and it will be found
that, though the
has increased in size so as to be more
mandible
than
twice as long from the symphysisto the condyle,the line
AB
has not increased,
has the line CD.
nor
If,however,
these lines be prolongedto the outer surface of the bone
it will be found that the jaw has
each side and in front,
on
quitedoubled its thickness by additions to the outside.
*
'
Anatomy
p. 205 of Tomes'
of 1898)will show these facts with great clearness.
(edition
The head of an adult is much
largerthan the head of
if we
into a
divide the head
a newly-born infant ; but
shall find that the amount
cranium and a face, we
of
in
the
is
divisions
two
that,
growth
very disproportionate
while comparatively
little increase takes place in the
cranium or brain-pan,
there is a very considerable increase
the
face.
in
The increase in the diameter of the cranium
which
may be indicated by the followingmeasurements,
:
typical
may be taken as fairly

glance at

the

figureon

"

51

6;

12

16 to 18
Adult

6
7

slow increase in the

with this comparatively
Coincidently
cranium, the growth of the face goes on at a great rate.
The

mandible

elongates backwards, the great air-cells

by

AIDS

84

TO

DENTAL

ANATOMY

above the orbit and below it expand,the latter or antrum

shallow depressionto
of Highmore increasingfrom
a
with
become
an
a cavern
capacityof eightfluid
average
The roots of the upper teeth,which
drachms.
at
were
first in close proximityto the floor of the orbit, find
themselves an inch and a half below it,while the nose
elongatesin proportion.To estimate this growth of the
the relative proportions
of the face
face,and to measure
in various animals, some
and cranium
imaginary lines
called
which
indicate what
are
Camper's lines,
employed
is called the facialangle.
A line is drawn
from the external opening of the ear
(theexternal auditorymeatus) forward to the lower edge
line is drawn
of the ala of the nose.
A second
from the
frontal

eminence,

or

most

prominentpart of

the

forehead,

ridge,where the eyebrow grows, to

past the superciliary
lines form where
the first line. These
two
meet
they
which
called
facial
is
the
an
meet
angle,
angle Obviously,
if the face is greatlydevelopedand projects,
while the
is small, this angle will be acute, while,
in probrain-pan
portion
.

as

the

extends
brain-pan

forwards

and overhangs
in ideal Grecian

face,the angle will be obtuse. Thus,

the
art
angle was ioo ; in the European infant about 90 ;
in the European adult,80 ; in the negro adult,70 ; in the
67 ; in the chimpanzee and
chimpanzeeand orang infant,
will
be seen
It
that a great development
orang adult, 40.
in the anthropoof the teeth at puberty,such as is seen
less degree in the
morphoid apes, and to a much
lower or less civilized types of mankind, tends to cause
of the face and a reduced angle; this condition
a projection
is called prognathism^and the individual is spokenof as
the

Prognathous. The

prominence of

the chin

generally
largeangle and a high mental development,
and the buck teeth,recedingforehead and chin,
and consequent low angle,are associated with imperfect
mental development or idiocy.
associated with

is

for indicating
the relative promeasurement
portions
of the brain and face is that called the measurement
of the gnathic index.
A
line is drawn
from the
anterior margin of the foramen
the
to
magnum
pointof
This is pracjunctureof the frontal and nasal bones.
tically
of
the
measurement
base of the skull,and is
a
Another

by

THE

DEVELOPMENT

OF

MOUTH

THE

85

called the Basio-nasal line ; a second line is drawn

from
the anterior margin of the foramen
the
to
magnum
the
two
alveolus,between
upper incisors or the front edge
of the alveolus ; this line is a measurement
of the length
of the mouth, and is called the Basi-alveolar line. Then
a

is

sum

divided

mouth-line
the basio-nasal line :

done,

by

the

Basi-alveolar length x

Basio-nasal length

multipliedby

100,

and

100
,

-S^thic

mdex.

If the result is below

is called orthog98, the creature
nathous ; if between
98 and
103, mesognathous; if
above
is now
measurement
103, prognathous. This
used by anthropologists.
more
generally
Among the various races of mankind
types of
many
skull and face exist,some
or
longdolichocephaloid,

skulled,some

or short-skulled.
brachiocephaloid,

In the
cranium
and wide mandible
called pyramidal.

Mongolian

races

give rise to

the appearance
of absorption
over

The

excess

the

narrow

of advancing
deposition

in the fact that the skull has been estimated

two-fifths of its weight between
the ages of

age is shown
to lose about

or eighty.
thirty

It has been

that the

seen

jaws increase greatlyin area

when

of the
tooth-bearing
portions
duringthe periodof eruption.

bear in mind that at an earlystage in the

formation of each tooth its enamel cap is formed
and the
size of its crown
therefore definitely
decided
moreover,
that all these crowns
their full size before
are
practically
the bony alveolus is nearly large enough to contain
them
it is obvious that in the early stages there is no
for them
to be arranged in an
room
even
row, but that
be
less packed, some
inside others
more
or
they must

Now,

we

"

"

and

above

below
the others.
Thus
in both the
jaws, the lateral teeth are in crypts inside (nearer the
tongue) those of the centrals and canines, these latter
almost
touching in front,while the upper canines are
formed
above
the level of the incisors,
and the second
molars are above the level of the firstmolars.
The bony
these developingteeth
crypts which contain and encase
without tops, so that if the jaw at this
at first boxes
are
be
denuded
of its soft parts the developingteeth
period
some

or

by

86
will

TO

AIDS

drop

out

of

DENTAL

ANATOMY

their crypts.

Then

the

crypt-walls

the germs
so
as
presentlyto enclose
themselves
The
are
always
crypts
completely.
contain
the
to
just deep enough
forming teeth, and
deepen,as additions to the roots increase,the lengthof
and tops of the
the germs ; afterwards the front portions
removed
by absorption,
exposingthe
crypts are gradually
which is called
of the crowns
enamel
to view
a process

graduallyarch

over

them

"

cuttingthe teeth.
Eruption. We

of teeth
have noticed that the germs
less distant
to be formed
at a pointmore
or
commence
from
the scene
of their future activity.They move
the
further in some
animals than in others, but what
in man
is regarded
forces are which govern this movement
still a moot
as
point. Many explanationshave been
of them
offered from time to time, and none
have been
It has been suggestedthat the
acceptedas satisfactory.
teeth are forced into their final position
by the lengthening
of their roots, but many
teeth with fullyformed
roots
are
never
erupted,and many teeth with stunted roots are
formed
erupted. It is true that uneruptedteeth with fully
but
their normal
roots do not generally
position,
occupy
of fully
formed
it is also true that frequent
instances occur
in the direction of eruptionuntil
teeth that do not move
in
for some
late
and
then
life,
unexplainedreason
quite
do erupt. I would
that the key
throw out a suggestion
of this mysteriousmovement
lie
to the explanation
may
in evolutionary
and that the process may
not
inheritance,
after all be so different from that which is known
take
to
in
of
the
f
ishes.
In
the
case
shark,
place plagiostomous
from the
where the area
of tooth development is remote
the organ has to traverse
of tooth activity,
a space
area
in
arrive
the
its
order
times
to
at
own
scene
length
many
is
of its labours.
It has been proved that this movement
due to a general rotation of all the mucous
membrane,
with
teeth
it.
and
The
jaw of
along
carryingthe germs
found transfixed by the spineof a sting-ray
a shark
was
;
"

membrane
the shark had lived on, and as the mucous
had moved
forwards,so a scar with signs of arrested
development had been producedforwards from the wound
that tissue that
to the ridgeof the jaw, showing plainly
time occupied the site of the wound
had
at one
had

by

THE
moved

DEVELOPMENT

OF

MOUTH

THE

87

forwards

have
We
to the front of the mouth.
in
milk
the
and
buds
seen
post -permanent
predescribed by Kiickenthal there are hints in the human
subjectand the dog of inheritance from an ancestor with
endless.succession of teeth. A very amazing case was
an
reportedby Dr. Gibbs of Edinburgh at the Stomatological
of milkCongress of 191 3, in which several series (four)
teeth at the same
formation
in
of
appeared
stage
imposed
superin the rightmandible
of a child of five.
rows
There were
also the germs of two firstpermanent molars,
above the other.
one
directly
Nothing was known about
the other side of the mandible, and the death of the child
prevented further facts being elicited. It seems, on the
that the human
to me
to
whole,possible
germ may move
with the generalmass
of connective
its placein company
herited
tissue in which it is buried,and in obedience to an infar-off ancestor.
habit of some
notable
Jaws
in
before Birth." The
the
Changes
in the jaws before birth may
be
changes which occur
tabulated as follows :
conveniently
and mandibular processes
The
buds
of the maxillary
1.
appear about the twentieth day of intra-uterine life.
ossified in a membrane.
2. Both jaws are
cartilageappears and takes the place of
3. Meckel's
of the mandible
the lower jaw before the formation
and then wastes
proper. It beginsin two halves,unites,
of
ossification. It disappears
until the commencement
of it
about the sixth month, the only persistent
trace
that

being in the ossicles

the incus).

of the middle

ear

(themalleus

and

4. Ossification beginsso earlyand proceedsso quickly

about the exact number
that it is difficultto be precise
of ossification. Separate
and
situation of the centres
have
nuclei
been, however, seen in
Maxilla.
i.

Alveolar

process.
bone.
2.
Intermaxillary
3. Frontal process.

Mandible.
i.

Coronoid

2.

Condyle.

process.

3. Angle.
and outer part
4. Under
of ramus.
5. Inner alveolus,

by

AIDS

88

TO

In the crocodile
and
throughout life,
human
lower jaw.

DENTAL

some

ANATOMY

remain

of these centres
been

instances have

5. Ossification beginson

recorded

the thirtiethto the

distinct
in the

thirty-fifth

day.
6. The

two

halves of the lower

jaw unite by

membrane

only.
7. At

the end

of the second

month

the

parts of the

lower jaw are plainly

recognizable.
all the nuclei of the
8. At the end of the third month
coalesced.
have
upper jaw
The Jaws at Birth." The two halves of the lower jaw
not united by bone, but by fibro-cartilage.
are
The
crypts for the temporary teeth are wide open, so
removed
the teeth would drop
that if the soft parts were
The crypts are only deep enough to contain what is
out.
formed of the teeth. The septa are so arranged that the
bours,
crypt for the lateral incisor is a littlebehind its neighwhile those of the central and canine -almost meet
in front of it. The
pressions
crypts of the centrals present dethe lingualwall for the permanent germs.
on
inferior dental canal is partlyseparatedfrom the
The
sockets of the molars,underneath
which itruns diagonally
from the inferior dental to the mental foramen.
There
inferior dental foramina" (a)the largeropens to
two
are
which
is in places a canal on
the floor of the
a groove
molar sockets,which persists
the inferiordental canal ;
as
under the
(b)the smaller opens to a canal which runs
is
a
nd
the
obliterated
milk-teeth,
eventually
groove to supply
the milk-teeth are
when
lost. There
also two
are
coronoid
mental foramina
each
side. The
on
process
stands up at a right
The
condyleis level with the
angle.
alveolar ridge; it ossifies up to its articular cartilage,
it
and the sternal end of the clavicle beingthe only bones
which do so.
The movement
of the mandible
is almost a
and
the socket of the glenoid cavity
pure hinge-joint,
is deep. The
alveoli of the upper jaw descend
but little
below the general level of the palate,
sockets
the
though
fair depth, and
is representedby a
are
a
the antrum
shallow depressionon the inner wall of the maxilla. The
between
the maxillarycheek-teeth
relationship
during
formed an important
antrum
eruptionand the maxillary

by

AIDS

90

DENTAL

TO

ANATOMY

in the angle is arrested,and does

not
half
until three and a
recommence
years.
Three Months*
The crypts which contain the teeth are
closingin over them, so that the germs do not drop out
when
the soft parts are removed.
Six Months,
The
crypts begin to be opened againby
consider to be the commencewhich we
ment
absorption,
may
of eruption.The crypts for the permanent centrals
surfaces of the crypts of their
distinct on the lingual
are
wide open.
are
temporary predecessors;their mouths
The crypts of the firstpermanent molars are distinct ; the

age

the

change

"

"

from the second permanent molars

is not very marked
indeed, the upper first permanent
molar crypt has no posterior
wall.
set in ; anchylosis
Eight Months,
Teething has fairly
in the symphysisof the mandible
The
has commenced.
is
marked.
the
mental
In
jaw
prominence well
upper
the antrum
is represented
extending
by a deep depression
under two-thirds of the floor of the orbit.
Nine Months.
In the upper jaw the alveolus is partly
them
septum dividing
"

"

"

absorbed

in

exposed to

front,and

view

the

enamel

is
forwards,but
being stillnot

of the

teeth have moved

their cuttingedges

; these

centrals

much
downwards,
much
below the general level of the palate. The molar
sockets are separatedfrom the floor of the orbit by the
of Highmore.
The halves of the lower jaw are
antrum
not

stilluniting
from

below upwards. It has been suggested

before
that the reason
that the lower front teeth appear
the upper is to prevent the maternal
nipplefrom being
the tongue protectnipped between two rows of incisors,
ing
it from the lower teeth.
into a
As the teeth move
increase
afforded
of space
by the
perfectrow with the
and expansionof the tooth -bearingportionsof
the jaws, their movements
are
certainlyinfluenced to a
by the pressure of the tongue from within,
great extent
antagonizedby that of the lipsand cheek from without,
and- the
buccinator
the orbicularis oris and
muscles
them
tendingto press the teeth into a
tongue between

growth

singleregularrow.
that is
teeth
Months,
The
cut
now
are
visible to about the same
extent
as
they were calcified
that is,two-thirds of the centrals,
one-third of
at birth
Twelve

"

"

"

"

by

THE

DEVELOPMENT

OF

MOUTH

THE

91

the laterals,
of the
the tipsof the canines,
half the crowns
first molars,and the cusps of the second molars.
The
of the centrals is complete,
enamel
and looks polished.
Three and a Half Years.
All the temporary teeth are in
their places; the roots of the incisors are complete ; twothirds of the canine roots, four-fifths of the first molar
"

and

one-half of the second molar roots, are formed.

The
changes in the anglethat were arrested at two months are
resumed.
now
Five
All the temporary dentition is perfect.
Years.
The order of the appearance
of the milk-teeth is generally
follows: the lower centrals,the four upper incisors,
as
the lower laterals and first molars, the canines and the
"

second molars.
That of the permanent series as follows :
the first or six-year-old
molars, the lower centrals,the
rest of the incisors,
the first bicuspids,
the canines,the
second bicuspids,
the second molars, and the third molars

wisdom

or

The

teeth.
process is not

continuous,but
eruptionsdivided by intervals of

of

molars
the

generallyappear

wisdom

at

rest ;

about twelve
fifteen and

teeth between
these may
make
their appearance
The
actual dates,and even
to some
are
eruption,

on

goes

Temporary

eruptionof

the

years

series

second
old,and

but
twenty-five,

almost
age.
any
the order of
periodbetween nine

at

extent

very variable,but some

and
twelve
months
generally intervenes
of
each
appearance
group.

Absorption of the

as

Teeth

between

the

precedes the
This absorption
hardlybe due to

their permanent
successors.
has been attributed to pressure, but itcan
this cause, seeingthat there is always soft tissue between
the two, and that it sometimes
affects teeth which have no

described
absorbent
as
organ has been
the way for the advance
of the second
dentition.
clearing
is to be
I think that here,as in eruption,
the explanation
found in an inherited tendency,and no doubt the active
successors.

An

is the same
that first opens
and
the
removes
agency
anterior portion
of the bony crypt which at one
time enclosed
the permanent
germs.
Milk- Teeth
The
their permanent
roughly resemble
successors.
They differ from them, however, in that

theyare smaller,that

their enamel

edges are

thicker and

by

AIDS

92

TO

ANATOMY

DENTAL

that the roots of the molars

wide apart to admit of the presence of the bicuspid
are
them.
The
between
cuttingedges of the front*
germs
wider than the necks, and before the
teeth are not much
end

somewhat

eruptedtheir temporary predecessors

widelyseparated.

teeth

permanent
are

abruptly*and

somewhat

are

CHAPTER

TEETH

OF

MAN,

VI

THE
AND
TEETH

HOMOLOGIES

OF

considered the microscopical

Having
and
appearances
minute changes that result in the organs we
call teeth,it
little
will be as well to devote a
space to the consideration
of the teeth themselves
and arrangement
of the form
when in use, and their relationship
to each
other, and to
the tissues in their immediate
neighbourhood.
that
notice
be
First of ali,we
the teeth of man
may
divided into distinct varieties incisors,canines, premolars
and molars.
A convenient and quick
or bicuspids,
method
of recordingsuch varieties is by means
of a
dental formula in which, as the rightand left side are
the formula
alike,
only one side is noted. Thus in man
is In. f Can. \ Prm. " M. | (forthe two sides)
32. The
sudden
transitions between
these groups
somewhat
is
accounted
for on
the supposition
that the creatures'
in a remote
ancestors
periodpossesseda fuller series,
have been lost. Organs that are ceasingto
and that some
tend to become
be serviceable to their possessor
mentary,
rudiand finally
to disappear.
teeth are arrangedin what is termed a parabolic
These
of the upper and lower jaws are not
but the curves
curve,
the same.
Except quiteat the back, the upper teeth
outside the lower.
The
six
overlap and are slightly
upper front teeth are quiteoutside their lower antagonists
in modern
so, about oneman, the two pre-molarsslightly
third of their surface projecting
beyond the outer surface
of the lower pre-molars. This projection
beyond the
"

by

G00gl(?

OF

TEETH

MAN

93

lower
teeth is less in the first molars, less stillin the
second
in the third
molars, and scarcely noticeable
molars or wisdom
This arrangement
necessitates
teeth.
that the curves
of the tooth-bearing
region in the upper
and
lower jaws should differ. The
of the upper
curve
is
jaw
part of a greater circle than that of the lower, and

is,moreover,

gradual and

continuous,never

alteringits

generaldirection from rightwisdom tooth to left wisdom

is in
tooth,whereas that of the lower jaw or mandible
front part of a lesser circle falling
within the arc of the
in
while
the
molar
teeth,
region the right and
upper
left sides separate rather more
abruptly,so that at last
the wisdom
teeth almost
size of the canine teeth

coincide

in

position.Increased

into
changes this gradual curve
of
sides
a
gram,
parallelosomething approximatingto three
and this may
be well illustrated by comparing a
well -developed
human
jaw with that of one of the larger
apes.
Looked
be found

at in any aspect, the upper

in curves
and
to antagonize

Thus, if we

look

the closed

at

jaws

and
not

in

lower jaws will

in straight
lines.

we
profile,

notice

that the upper surfaces of the lower teeth rise gently as

the eye travels towards the molar region,
that a piece
so
of paper held between
the upper and lower teeth would
bend
but would
not lie flat,
upwards towards the back.

Lastly,if we

look at the molar

teeth from

behind

"

having

the parts behind

the teeth that would
removed
have
while
shall
that
the
obscured
the view
see
we
upper
molars
face not only downwards, but slightly
outwards
towards
molars
the cheek, the lower
face not
only
inwards towards the tongue.
Thus
upwards,but slightly
of the mandible
the movements
the
jaw
upon
upper
which are produced by the alternate contraction of the
external pterygoidand
other
muscles
partake in all
directions of a sort of ball and socket action, and do
resemble
flat plainsin movement
the
not
two
one
over
"

other.
The

wider than
the lower
central incisors are
upper
ones, and overlap part of the lower laterals ; the upper
laterals antagonizethe rest of the lower laterals and part
canines and pre-molars
of the lower canines; the upper

likewise

meet

the back

half of their lower

analogues and

by

TO

AIDS

94

DENTAL

ANATOMY

the front half of the tooth behind.

In the molar series
this discrepancygraduallyadjustsitself,
the first molar
touchingthe front portionof the lower second molar, the
molar justreachingover
the lower wisdom
upper second
wisdom
tooth,being rather smaller
fourthan its lower analogue,coincides with its posterior
is greater than the lower,
fifths. Because the upper curve
outside
and the upper pre-molarsand molars are slightly
the jaw is at rest, the outer cusps of
the lower ones
when
double
teeth
than the lower ones
the upper
less worn
are
and vice versd,so that in the upper teeth the outer cusps
while the
stick up more,
and the inner ones
are
flatter,
is true of the lower teeth.
considerations
These
reverse
render the value of a tooth greater than one-thirty-second
tooth
of the whole armature, because
the loss of any one
teeth which
the loss of service of parts of the two
means

tooth, while the upper

it

antagonizedwhen
with

mouth

hinge

simply opening and

; but when

movement

shutting the
we

into
side to

take

from
the lateral roll of the mandible
side, the loss of masticatingsurface is obviouslymuch
account

increased.
should be quitefamiliar with
It is importantthat we
the macroscopical
of the human
or
naked-eye anatomy
teeth. And
the student should be prepared to describe
certain
tooth.
There
are
accuratelyany normal human

characteristics
or

to

common

every

mandibular.
or
molar, maxillary

consists of

tooth,whether

Every

human

incisor
tooth

being
crown, and a root or roots, the crown
separatedfrom the root by a constricted portionwhich
is called the neck.
is exposedto view,while the root is hidden
The crown
The neck is nothingmore
than a convenient
by the gum.
a

term
descriptive
was

at

surfaces

one

time

of

no

morphologicalimportance.

as

the

It
various

speak
internal
anterior,
posterior,

customary

of the teeth

of

to

of the jaws,
this nomenclature
is very misleading,and
has been
discarded in favour of the following
far more
convenient
The
biting surface is called occlusal.
system : The
surface which faces towards the lips
is called labial,
that
towards
the tongue lingual,while the remaining two
surfaces are termed
mesial and distal,
accordingas they

external,
respectively.
Owing

to

the

and

curve

by

TEETH

OF

MAN

95

face towards or away from the middle line. Thus it will

that the anterior surface of a central incisor would
be seen
the anterior surface of a
be the labial surface, whereas
this point
To make
be its mesial surface.
molar would
teeth
clear,let us consider the eightright mandibular
and imagine them
to consist of a locomotive
separately,
with six coaches and a guard'svan, runninground a curve.
incisor is the engine, while the third
The
central
If the lower lip be the
is the guard's van.
molar
platform,it will be seen that the labial surface of our
train is that at which
must
alight; if they
passengers

they would
upon the lingualsurface,
mesial surface
the other line. The
find themselves
on
of each coach faces along the line in the direction in which
the train is moving, while the distal surface faces the
attempted to

do

so

The

oppositeway.

corresponds
obviously

occlusal surface

to the roof.

teeth of man,

The

which

apportioninto
very
on

hard
the

groups, are
define.
The
to

and
recognize
in
else
like everything
Nature,
are

so

easy

the teeth situated

the teeth
canines are

incisors

bones, the
pre-maxillary

to

are

suture
ifnot too far
maxillo-intermaxillary
is
which
behind
vague), the
delightfully
(a qualification
The
lower.
the
behind
canine
pre-molars
biting
upper
which
series
of
molar
the
porary
teeth
those
displacetemare
next

behind the

the molars are the teeth behind

them.
Incisors and canines are generallysingle-rooted
;
which
fused
often
sometimes
have
are
two
roots,
pre-molars
The
second
in
the
pre-molar in man.
together
upper
have usuallythree roots, an inner or
upper molars in man
and two
outer
or
buccal, the roots of the third
palatine,
lower pre-molars
molar being often fused together. The
and
the molars
have
an
are
generally single-rooted,
anterior or a posterior
root, which are often longitudinally
the result of the fusion of
grooved,as though they were
These roots incline
anterior and two posterior
roots.
two
and are in the case
of
backwards
at their apices,
slightly

while
predecessors,

the

wisdom

tooth

often

fused

together into

single

mass.

of
facts are, generallyspeaking,true, but none
These
that of the
the definitions are reallygood except, perhaps,
incisors. The lower incisors are supposed to be
upper

by

AIDS

96
the

TO

DENTAL

ANATOMY

which

teeth

The
correspond to the upper ones.
to define with any accuracy
are
reallyimpossible
of fact,they are
as a matter
simply first pre-molars,and
though where a tooth is speciallydevelopedin what we

canines

must

call

situated

caniniform

fairlyclose

direction it is generallya tooth

the maxillo-intermaxillary
behind
'

that such a tooth in the upper jaw usually

bites behind its lower antagonist,
this is not invariably
the
Thus in the mole, and in an extinct animal called
case.
Oreodon, there is a largeupper tooth of a typicalcanine
form (two-rootedin the mole),which
bites in front of a
caniniform lower tooth.
If this lower tooth be a canine,
then these creatures
incisor teeth
possess four lower
and four upper and three lower pre-molars,
which
would
look as if the lower tooth we
call a canine had remained

suture, and

in development,
insignificant
had

attained

caniniform

while the

first lower
dimensions.
If we

premolar
had

all over
begin our nomenclature
again,perhaps we
should avoid the name
canine, as leadingto so much
confusion ; but it is sanctioned by long usage, and itwould
to

be

with
to dispense
impossible

it

so

now,

that

we

must

remembering that it is in reality

but not invariably,
the
onlya specialized
pre-molar,
usually,
content

ourselves

with

first.
be almost level,
In man
the cusps of the teeth should
no tooth rising
markedly above itsfellows ; the alveolus in
which they are implantedis thin on the outside and solid
the inside until we reach the lov/er wisdom
on
teeth,where
the thickness of the bone is generallygreater on the outside.
The front teeth stand up nearly straight
; the teeth
and their surfaces,
which
all touch their neighbours,
are
coated with enamel, become
broader from the pre-molars
backwards

the second
molars.
The
total number
of
eightabove and eightbelow
permanent teeth is thirty-two,
each side.
on
The typical
formula^as it is called," \ " " 44, is the
to

have been possessedby some

of the
class. When
to the mammalian
an
animal possesses fewer than forty-four
teeth,it is spoken
of as havinglost some,
in
that
the course
of ages
meaning
teeth which were
of supreme
not
to the welfare
necessity

formula

supposedto
common
progenitors

of the animal

have

graduallybecome

more

by

in
irregular

ANATOMY

DENTAL

TO

AIDS

98

here to endeavour
by
them to students in a

of

means

homely simile

to

present

less

glaringmanner.
Upper Central Incisors." Suppose we take two wedgemeasuring6 inches in height,
shaped piecesof plasticene
Let
4 inches in breadth,i" inches in depth at the base.
of a rightand
into models
them
us
proceed to make
more

or

upper central incisor. First we cut a

inch
the cuttingedge of each, beginning
from the cuttingedge on what is to be the distal surface
in an upward direction
of each tooth,and slopingacross

tapering
|

leftnormal
slice from

the mesial side. Then

tillwe
reach the surface on
we
inch
side
of
each
the distal
slice off from
a
shaving J
the
at
the
at
cuttingedge,
neck, taperingto nothing
tooth
the
curve
on
convex
; then we do
leavinga slight
the

same

surface.

mesial
make
we
inches
3

the

on

Then

lingualsurface
edge where the

side,leaving a slightlyconvex

V-shaped depressionon the

long, reachingfrom the cutting
a

inches broad, to within j inch

it tapers to nothing; this
of the cervical edge, where
outside ridge,the cingulum. The
will leave an
act of
will
up-and-down convexityto the
givea slight
doing this
front or labial surface.
Finallywe round the edges.
resemble
the
The
Upper Lateral Incisors." These
centrals,except that they are narrower, the cingulum not
V

is

rounded.
The
cuttingedges more
and
mesial surfaces are
longer than the distal,
slightly
this givesthe indication of rightand left teeth.
The Upper Canines are pointed; their roots are longer

so

marked, and

the

The
those
of the other teeth.
than
slope from the
shoulder to the point is longer on the distal side,and this
gives the diagnosisof rightfrom left. The cingulum is
very stronglymarked.
These
have two
The
principal
Upper Premolars.
labial
t
he
and
more
labial,
beingslightly
cusps, a lingual
"

pronounced. They

are,

however, nearlyof equal height

had a plasticene
If we
model
indented
its
of an upper
bitingsurface with
H divided the
the
of
bar
the
H
that
iron letter
so
an
labial from the lingualcusp, and the whole indentation
only occupiedthe middle of the surface,with a complete
ring of enamel enclosing it,we should get a fair reproduction
of a bicuspidcrown.
The
lingualsurface of the

(unlikethe

ones).
pre-molar,and

lower

by

TEETH
teeth from

the second

less than the

have

labial,so that each

the

near
or

99

the second bicuspidis

tooth is narrower
its
on

the intervals between

narrower

are

singleroot

MAN

bicuspidto

surface,
lingual
justas
wheel

OF

two

roots

hub.
fused

the

spokes of

Mostly bicuspids
times
together,but some-

the two roots are separate. The upper firstbicuspid

is more
often two-rooted
than the second.
become
smaller in area from before
The Upper Molars
backwards, the third being the smallest. They have
labial (orbuccal) and
three roots, an anterior and posterior
-The
surface of the upper
a palatine.
cusps on the outer
than the inner ones, and the
cheek teeth are less worn
anterior internal cusp is largestin area ; the recognition
of the anterior internal cusp decides the questionof right
anterior internal cusp is joined by a ridge
left. The
or
to the

external.
posterior

Incisors.

The Lower
than

The

lower centrals

are

littlemore

of their upper
and the
antagonists,
wider than the centrals.
The
lower
slightly
generallytouch their neighbours from neck

half the

laterals are
front teeth
to

"

width

cuttingedge, leaving no

little V-shaped

interspaces

the upper incisors.

Their roots
between
as
we
see
are
generally flattened from side to side,so that the
is greater than
that from
labial to lingual diameter
of the upper
mesial to distal surface.
The
central
root
such

is often round.

The

Lower

Canine

is a narrower
and
otherwise
resembles

slimmer

tooth

it. The lower

than the upper, but
canine
the
to
molars
displays its relationship
pre
by its very marked
cingulum,and it may be noted that
caniniform
than the upper.
the lower pre-molarsare more
-

The
than

Lower
the upper

Pre-molars
ones.

and
insignificant,
indented

to the

Lower

much

more

like canines

The inner cusps are comparatively

mesial and
the
distal surfaces are

edges.

Molars are
anterior and posterior
root.
and in the case
at the apices,
The

are

two-rooted

teeth

having

an

These roots curve

backwards
of the third molars are often
fused.
molars
the outer
In the lower
are
most
cusps
is
anterior
the
internal
the
but
elevated.
most
worn,
cusp
There
three
and
inner in
outer
two
are
generally
cusps
in the second.
the first molar, and sometimes
The third

by

AIDS

ioo

TO

molar in European man

five
in primitive
races

Teeth

Missing
it is I3,or the

DENTAL

are

ANATOMY

has usuallyonly four cusps, but

often present.

in Man."

We are generallytaught
of the series,
that is missing;
outer
one
that
thinks it is Ilfand Albrecht that it is I2. The
but Baume
obviouslysharp transition between the lateral and canine,
and the noteworthy fact that the upper lateral is itself
of size and eruption
a tendency
showing by its irregularity

The

to follow

suit,pointto the

outer

member

of the series

as

based upon
the lost tooth. The other opinions
are
chiefly
of
the
o
f
the
third
observation
incisor
the
when it
position
Albrecht
for
is present in man.
somewhat
as
I2
argues
bone on both sides is divided
follows : The pre-maxillary
inner portiontouching the middle line and
into an
an
maxilla
the
The inner
outer
portiontouching
proper.

portionis called the endognathion,the outer portionthe

mesognathion,while the maxilla proper is called the
the
exognathion. In cleft palate the cleft is between
in
these
third
and
endodwarfed
cases
a
meso-gnathion;
incisor is often present, and

is situated

the inner side

the obvious suggestion
of the cleft,
beingthat itrepresents
of the lost I2. Professor Arthur
Keith
a
reappearance
the
situation
of
these
that
incisor
teeth is
has shown
be relied upon
decisive
as
variable,and cannot
very
evidence,since he has found them in the cleft itself. It is,
that in these
to
assume
irregular
perhaps, unwise
dwarfed
teeth
the
always appear in the
developments
when
accessory teeth do appear we
proper place; in fact,
much
from their situation,
too
not assume
must
as
we
the
shall see in discussing
pre-molars. At present most
incline
the
to
authorities
opinion that it is I3 that is
on

missing in man.
Missing Pre-molars."

These
are
probably Px and P2.
model
that
has been frequently
a
I have in my possession
figured,in which, the full complement of teeth being
present in all other parts of the mouth, there are no less

than six perfectly

developedbicuspidteeth on one side
taken to
of the upper jaw. If this phenomenon were
infer
it
would
that
the
mean
anything
typicalformula
of the progenitorof this individual was
two
pre-molars
received
formula
allows of. To argue
greater than the
such lines might lead us, if we wished to be consistent,
on

by

TEETH

OF

MAN

101

the possessionof
attribute to some
ancestor
remote
ovarian teeth.
clines
animal inDiet and Dentition.
In proportion
as
an
to a restricted flesh diet we find certain peculiarities
in its teeth and the surrounding parts. The
incisors are
to

functional

"

row
straight
; the canines
and recurved ; the pre-molars
and molars
large,pointed,
In typical
five in number
and
specializedin form.
carnivora there is a tooth in the upper jaw (the fourth
and one
in the lower (the firstmolar) specially
pre-molar)
food, and called carnassial.' The
developedfor slicing

six in

number, small,and in

'

lower

carnassial

tooth (the firstmolar) is shaped like the

letter M, while the first upper pre-molarhas three
capital
is the largest,
great pointedcusps, the middle, which
between
the pointsof the M, and the anterior
in
fitting
in front and
and posterior
ones
coming down respectively
behind it,
inner tubercle being the only relic of
a small
the rest of the biting
surface of the tooth.
Thus, in the
have

tigerwe

the upper
molar is a tinyabortive tooth of no use
to its possessor.
In the extinct machairodus
stillfurther
the formula
was
reduced by the loss of an upper pre-molar,9 J f J"and in
a

forirula," \ " },and

of these

happilyextinct

of this ferocious family,

member
of Smilodon, it was
name
probablyeven less,f ]f ?,
only twenty-fourteeth in all. In the hyaena we find the
inner tubercles largerand the pre-molarseries increased,
another
with the

in the mustelidae
molar added, " J

"$"I;

(stoatsand

weasels)we

have

in the cat an upper molar,

"
i f 5 m tne dog another lower molar, " \ f $ ;
the horse possesses
the full typicalformula
and, lastly,
a

lower

{ ";

As we mount
of molar
the scale in number
teeth, so
find
them
altered in character and shape,becoming
we
broader
and better adapted for triturating
or
grinding,
less formed for slicing
and tearing,and coincidently
we
notice changes in the tooth-bearingbones.
Where, as
in the felidae,
the molars
few and
the canines
are
the movement
of the mandible
is a pure hingeenormous,
the condyle is a rod placed almost
joint movement;
transverselyto the axis of the jaw and moving in a
hollow, the glenoid cavity. The external
cylindrical
pterygoidplatesare dwarfed because the action of the

by

AIDS

102

TO

external
never

ANATOMY

DENTAL

pterygoidmuscles
moving from side to

is practically
nil, the
side ; in fact,this muscle

jaw
is

The
attached
to the interarticular fibro-cartilage.
of the movements
to admit
zygomaticarch is enormous
of the huge temporal muscles, and the coronoid process
is huge, the temporal muscles
being employed to shut
the jaw suddenly and having specialreference to the
the
action of the canine teeth in snapping, whereas
not

masseters

have

more

reference

to

the process

known

as

crunching.
animals the exact
oppositeis the case.
Their herbivorous diet requires
broad molar and pre-molar
teeth,the condyles are flattened,and so is the glenoid
cavity. The large external pterygoidmuscles arise from
and are attached to the
largeexternal pterygoidplates,
lateral
interarticularfibro-cartilages,
that the extreme
so
in
dislocation
of the mandible do not result
movements
a
In ruminant

of these parts ; the coronoid

processess are insignificant,
but the attachments
of the masseters
well developed.
are
the upper
Where
there is a normal
interval between
canine tooth and the outermost
incisor the space is called
the diastema,and the lower canine tooth fitsinto it when
the mouth
is closed.
ruminants
In many
the upper front
teeth are
missing,as the animal tears the vegetable
that form its herbaceous
diet by pressingits
structures
that
lower incisors againstthe pad of mucous
membrane
occupiesthe space usually allotted to incisors in the
ruminant
formulae as
find such
we
upper jaw. Thus
molars
and
pre-molarsbeing large and flat
" 1 I h tne
the canines insignificant.
and rough on the surface,
With
regard to these latter teeth in the ruminant familya
with the
curious fact must
be noticed in connection
of
and
in many
horns
canine
teeth.
Horns
relationship
the purpose of canine teeth as weapons
of
animals serve
offence in those combats
between
the male animals which
the breedingseason
; both are weapons
in
and
therefore neither
sexual
combat,
warfare,
are
greatlydeveloped in the female animals,and both
male individual.
are
rarelyfound developed in the same
does
Nature
rule providea creature
with two
not as
a
of weapons
for doingthe same
and
sorts
thing, though
there are a few rare exceptions (such as Swinhoe's water*

alwaysaccompany
of

used

by

TEETH

OF

MAN

103

is a generalrule that if horns are

veloped
greatlydecanine teeth are not, and vice versd.
In connection
with correlation of growth, a most
mysteriouspieceof evidence with regard to the strictly
sexual character of certain canine teeth is afforded by
male when
the effects of early castration. The unsexed
it arives at maturity does
not
develop these special
tusks
of the wild boar do not
characteristics. The
acquiretheir formidable dimensions,justas the change
male
of voice which accompaniespuberty in the human
That
does not take placeunder similar circumstances.
in evolution should take place
so
strangea retrogression
in the life of a singleindividual is a matter
that is at

deer),it

present whollybeyond explanation.

in form or number
alone that teeth
It is not, however
role
that
the
are
they are called upon
special
adapted to
and arrangement of the comto play. The
ponent
disposition
is equallyremarkable
tissues of the teeth themselves
in its suitableness. If a sharp, strong, piercing
is required,
is clothed with a
the dentinal mass
weapon
sheath of enamel, as in the canine of the felidae.
If a chisel-shaped
incisor is requiredfor
(scalfiriform)
gnawing, as in the rodent family,the sharpness is
attained by a thin outer
layer of enamel, with softer
dentine behind it and stillsofter cementum
at the back ;
different
these three tissues wearing away
at
rates, the
hardest wearing most slowly,tends to form and preserve
of
while the incessant wear
a
constantlysharp surface,
the tooth necessitating
the
active
constant
a renewal,
as

pulp continues its function throughout life (teethof

persistent
growth) instead of the root closingand the
at maturity. Where
a constantly
growth ceasing
rough
the result
and grinding,
triturating
of these
is attained again by a judiciousintermingling
tissues of varyinghardness
in the surface of the tooth.
in
the
molars
of
the
Thus,
elephant,capybara,horse,
and
find
of dentine surrounded
rhinoceros
islands
we
tapir,
and
by ringsof enamel in a sort of sea of cementum,
surface is required
for

the different rates of wear

produce the rough surface in
this case justas they produced the chisel surface in the
rodent incisor. The patterns of these molars are constant
which can be infallibly
for various families,
recognized
by

by

AIDS

104

them

; but

TO

though the

DENTAL

ANATOMY

results are often so complicated

as
than anything
old-fashioned -maze
their originis extremely simple.

resemble
an
mode
of
cone
shaped,consisting
Imagine first a simple denticle,
of a body of dentine,a coatingof enamel, and a covering
show
this would
an
of cement;
after a little wear
as
line surroundingit
enamel
island of dentine with an
and cement
beyond. Imagine four such denticles with
a
common
pulp,and you get four ringedislands %% (as
in number, say
in the hog). If the denticles are
more
six or eight,
and united transversely,
we
get the molar
the cusps
of the primevalelephants(mastodon); multiply
and cuspules,
and the result is the molar of the African
givesus that of the
elephant,and further multiplication
Indian elephant. Or suppose a different series of changes.
Imagine the two anterior cusps of the hog united,and
the two
likewise,and you have a bilophodont
posterior
if
tooth (tapir)
the two outer cusps are also united you
;
have another pre-molarof tapir
; let the line of union of
the two anterior and the two posterior
be crescentcurves
shaped instead of straight,and you have the P2 of
to

more

else,the

crescent
rhinoceros ; increase the posterior

very much,
of this
the
let
ends
rhinoceros
of
have
;
P3
you
and
curl round
crescent
P4 of
meet, and you have
back
stretch
anterior
to
rhinoceros ; let the
crescent
join
internal cusp, a small island developeinside
the posterior
the anterior internal cusp, and the outer
edge become
of hipparion
molar
the
and
have
bicrescentic,
you
of
(ancestorof the horse); while a little irregularity
outline brings us to the complicatedpattern of the
and

modern

horse's molar.
Where
the various dentines are present in the same
tooth the tubed or hard varietyis always outside,the,
vasodentine next, and the osteodentine or softest inside.

THE

TEETH

OF

ANCIENT

MAN.

have altered remarkably little since

teeth of man
times.
palaeolithic
Only one instance of a mandible with
discovered so far,
any trulysimian characters has been
and that is the fragment latelyfound at Piltdown.
The
The

by

106

AIDS

TO

DENTAL

ANATOMY

the Kaffirs the third

the neighbouringteeth.
Among
lower molars are eruptedfairly
early,and are often wider
than the second molars.
In European modern
the
man
molars decrease in size backwards, only the first molar
and the third molars are often
having five cusps as a rule,
dwarfed
a
nd
and
irregularin the time of
capricious
their eruption.These
considerations pointto the fact
in
that changes in facial and dental developmentoccur
of
short
that
and
a fluctuating
vironment
entime,
quitea
space
different
and dietarymay
a
produce
type of
mandible and different teeth in what, geologically
ing,
speakwhole
brief
The
is a very
bony anatomy of the
period.
face is,I may
say, governed by the necessities of the
teeth. The
snapping closure of the huge canines of
the felidsedemands
a
very tall coronoid process for the
the cheek
insertion of the temporals,and in such cases
in form.
The condyle
teeth are few in number
and slicing
in such cases
is a wide rod fitting
in the
a deep groove
and
the
external
of
temporal bone,
pterygoidplate the
is
sphenoid is abortive,as no side-to-side movement
flat-surfaced cheek teeth of the
possible.The numerous
ruminants are associated with great lateral movement,
and big pterygoidplates. On
big external pterygoids,
the other hand, the cranial capacitygoes on
slowly
and taking,
notice of
to speakcolloquially,
no
increasing,
the face and its variations.
that
Thus, it is not surprising
find in Neanderthal
and the Piltdown
we
man
man
not
much
but a totally
difference of cranial capacity,
different
different
dentition.
of
mandible
and
a very
type

by

APPENDIX

introduced
Mr. Howard
Mummery's paper on
Calcification (Phil.
Trans., 19 14)in this Appendix because
in time to be incorporated
in the text.
it did not appear
I have
not
or
attempted to summarize
epitomizethe
I HAVE

because

paper,

such

an

importantpronouncement,

up as it does so much
valuable in the authors

more

ing
clear-

confusion,is much
The platesare
words.

ancient
own

Transactions
omitted, but the Philosophical
necessarily
is not illustrated.
contains them, and the present work
paper contains not only Mr.
but also a lucid account

The

I have
will,

of earlier

where

doubt,prove

very

IN

In

and

J.

Mummery,

Howard

enamel, I have

as
investigations

to

dentine commenced

having been

to

show

no

case

the

sense

CALCIFICATION
DENTINE

M.R.C.S., L.D.S.

of calcification in dentine
endeavoured
to
carry further my
the

structure

development of
object of that paper
and

in 1892, the
the existence of

connective
pulp,in which

from the tooth

place much as it does

actual mode

has

process

derived

takes
The

AND

ENAMEL

consideringthe

foundation

OF

PROCESS

THE

By

work, which

instructive.

otherwise unintelligible,
but in
been allowed to suffer.

ON

coveries,
dis-

recent

It has been
of
alter the sequence
to slightly
occasionally
allusions to absent plateswould
have been

no

unavoidable
words

Mummery's

in membrane

tissue

fication
calci-

bone.
of the lime-salts I did not
of deposition
107

by

108

AIDS

TO

DENTAL

ANATOMY

consider,except to express the opinion that the

calcification took place by a process of secretion from
the
the cells of the pulp. Since that date,and following

then

Ebner
and
others, the
conversion
theory of the formation of dentine has been
to a great extent
abandoned, this conversion theorybeing
that the odontoblast
cell became
actuallyconverted into
dentine matrix,its centre
remaining uncalcified as the
grees
soft fibril and the rest of the cell formingin different desheath and the matrix.
of calcification the Neumann's
The view held in this paper and by numerous
histologists
at the present day is that the cells of the pulp secrete
a
which
material
not
calcifies,
entering
they themselves
into the calcified substance, but receding farther and
farther into the pulp as calcification advances, and the
of
previousinvestigations

von

fibrilbecomes
and more
more
elongated.
had
functions assigned
odontoblast
cell
has
The
many
of them
to it,
one
being that it is a nerve end-organ,and
that its prolongation,
the dentinal fibril,
conducts sensation
Charles
from
the dentine.
As Mr.
Tomes
says,
'*
in
of
the
the
difference
to
opinion as
writing
1904,
cells never
be finally
function of the odonoblast
can
settled until the nerve
endings of the pulp are finally
demonstrated.'
As I hope it is now
proved
satisfactorily
distributed to the dentine,the
fibres are
that true nerve
of the real
ground is cleared for a further investigation
and functions of the cells and dentinal fibrils.
nature
We
no
can
longer look upon the odontoblasts as being
in any
sense
nerve
end-organs,or consider their processes,
transmitters of sensation
the dentinal fibrils,
as
substance
of the dentine.
From
their
from
the hard
c
onsider
a
nd
them
structure
we
position
may confidently
in
the
formation of dentine,and I
be
to
engaged
actively
of this and of the
shall endeavour
to produce evidence
carried out, drawing
in which
their functions are
way
certain
a
ttention
to
special
processes of a purelyphysical
in calcification and their
concerned
which are
nature
cells
co-ordination with the action of the living
of lime salts
In order to study the process of dialysis
under
I
made
some
ments
experipurely physicalconditions,
in which
of
o
f
were
particles sulphate
copper
dropped into a solution of silicate of potash in water

by

APPENDIX

containinglime salts,and
In the tubes
transverse

109

examined

under

scope.
micro-

the

of the osmotic growth so produced,delicate

membranes
After
appeared.
occasionally

interval of several days, large,spherical,

colourless
bodies
formed
the distal side of the membranes
on
;
in
these were
similar
of the
to many
exactly
appearance
in
albumin.
calcospherites
produced artificially
They
would appear in this instance to be spheresof silicateof
in a colloidal medium
lime formed
of
by the dialysis
an

their constituent
that the passage
does take

salts through the membrane, showing

of salts through these osmotic
branes
memThese
bodies
behave
to
polarized
place.
do the similar spherites
produced in

light exactlyas

albumin.
The
by calcium
penetrationof osmotic membranes
salts has necessarily
an
importantbearing upon the
and dentine,as the proof calcification in enamel
nature
cess
would appear
to depend to a great extent
upon the
lime
The
matrix
of
both
the
salts.
tissues
o
f
dialysis
being a substance to a considerable extent removed from
should expect physical
the pale of nutrition,
we
processes
to play a largepart. While, however, the role of purely
it
physicalprocesses has to be taken into consideration,
be remembered
that,as Professor Philipsays, * the
of the livingcell membrane
towards
behaviour
the
is
in
with which
it comes
in many
substances
contact

must

cases

on
incapableof interpretation

purely physical

basis.' This is instanced by the altered conditions in

dead cells which
permit the penetrationof stains to

imperviouswhen

and
living,

also

by the
that
in
the
salts
contained
instances,
differ from those in the plasma, the
the blood-corpuscle
of potassium and
corpusclescontainingan abundance
phosphate,while the plasma shows a large proportionof
and
sodium
chloride,and little of the potassium and
phosphates. These facts show that *a purelyphysical
a living
theory of the exchanges which take place across
is
is inaiequate
a
membrane
; there
physiological
meability
perwhich

they are

fact, which

as

he

well

as

physicalpermeability.'

by

AIDS

no

The

Artificial

TO

DENTAL

Production

ANATOMY

of

Calcospherites.

ular
1858Rainey publishedhis work on moleclime
that
salts
were
coalescence,showing
deposited
in
in colloidal,
but in a
not
a
substances,
crystalline,
ture
globularform, the globuleshaving a very definite strucIn the year

also showed
that in many
He
and
arrangement.
and
mollusca
similar
animal
Crustacea,
organisms,as
forms appearedduringthe process of calcification of the
shell.
Both

Rainey and Halting found that when calcium

of the
phosphatewas present in the solutions in excess
posit
carbonate,globularbodies were not formed, but the deif there was
was
only a small
; but
crystalline
proportionof phosphate to carbonate, largerand more
sphereswere produced than with carbonate alone.
perfect
This observation has a very importantbearingon the
subjectof the present paper, as in completed dentine
and enamel
the phosphates are largelyin excess.
On
repeatingthe experiment of Professor Harting
main forms of calcospherites
with albumii, 1 found that two
those in
convenientlybe distinguished,
may
which radial striaeare most
evident,and those especially
concentric
lamellae
marked
or
rings,althoughthere
by
which show both structures.
are
many
beside the
of the larger radial spherites,
In many
wider
delicate radial striae,
seen
extending
spaces are
of
the
from
the centre
to
an
circumference,
irregular
of
or dividing
shape,and evidentlyindicatinga splitting
in the
into sections. This is frequently
seen
spherite
r
eferred
to.
globulesin enamel, presently
disintegrating
in
the
of
calcification
the
In studying
Crustacea,
process
the forming shell of the crab,of the prawn,
I examined
and
the small Norway lobster (Nephrops
the crayfish,
norvegicns)a specieswhich I do not think has been
examined for this purpose.
previously
in the carapace
of the prawn
of
The calcospherites
are
the radial type, and of particularly
elegantforms ; in the
of compressedspirals
the
form
in
crayfishthey are chiefly
the

"

while in Nephrops they are radiate and of very

helices,
definite floral form,and show very clearly
the breakingup

or

of these flower-like elements

in the

of
production

by

the shell.

APPENDIX
In

can

in

ground section of the forming shell of the crab,it

be seen
by dark-groundillumination that the lines in
striae
the transverse
form
detached
calcospherites

the
visible in the formed shell.
I shall describe in dentine and
It is noticeable,
as
spherites
enamel, that there appears to be a special
type of calcoin each of these different structures.
I think it is evident in all these examples that the shell
spherites,
of the calcois ultimately
formed
by the disintegration
which
may take placeeither previousto their
after that incorporawith its substance
or
tion
incorporation
has taken place.
in
examined
I have
In all these organisms which
there is a distinct
Crustacea, mollusca, and brachiopoda,
finelyfibrillarbasis substance in which calcification takes

place,which appears to serve as a support to the material

This fibrillar substance
is of a
secreted by the animal.
clear,apparentlystructureless,homogeneous nature, and
in any
way
appear
of the calcareous
arrangement
does

to

not

serve

to

determine

Calcification

in

actual

but simplyseems
particles,

binding material. It is not

with
and shows no connection
tissue,
as

the

the

true

nective
con-

cells.

Dentine.

In young
growing teeth,as is well known, there is a
matrix forming a marginalband between
of
the
portion
the calcified dentine and the odontoblast cells.
This layer,
the odontogeneticzone, appears
to consist
of the collagenousbasis substance of the dentinal matrix
of lime salts takes place. As shown
in which the deposit
in a former paper, fine connective tissue fibres from the
in this material,
pulp can be seen to become incorporated
althoughfrom their transparent nature and their index
that of the material
of refraction so nearlyapproaching
in this zone
they can seldom be seen in its substance ;
revealed in the formed matrix
but they are occasionally
action or acids in caries.
of dentine by the decalcifying
uncalcified portionof the matrix, unlike the calThe
cified
a
nd
takes
stain
the
the
readily,
advancing
part,
calcification is seen
encroaching upon it in the form of

by

AIDS

H2

rounded

masses

material.
are

seen

TO

DENTAL

of the lime
free in the

ANATOMY
of the

salts,some

spherites
calco-

lying
surrounding uncalcified
The
in
calcospherites this advancing layer
radial or
to be quiteclear,and
to exhibit no
markings.

concentric
When
tooth is decalcified,
the rounded
a
tours
conyoung
of the calcifying
border
have
the
same
exactly
in the calcified tooth ; they appear strucas
tureless,
appearance
and consist of the calcoglobulinbasis of the
which
takes stains readily. The
well-known
spherites,
in imperfectly
interglobular
developed dentine
spaces
show very clearly
the advance
and coalescence of these
bodies
basis
the
substance.
on
globular
The
the oppositesides of the pulp
between
space
with calcospherites
in all stages of
cavityis crowded

coalescence.

Comparison with calcification in the crabshell shows

that the same
process of coalescence is in
do not show
progress, but in the tooth the calcospherites
the striaewhich
evident in the similar bodies in
are
so
the crab-shell.
In some
I prepared with Ramon
sections which
y
silver nitrate method, and
Cajal's

which were
taken from
an
unerupted human
premolar, a further stage in the
consolidation
matrix
is revealed, which
of the
appears
mode
of
the
to throw
calcification
of
a strong light
on
clear the meaning of certain
dentine, and to make
in the adult tissue of which
I had
not
appearances
hitherto met
with any explanation.
As

the calcospherites
seen
stated,
previously

dentine

appear

clear and

but it was
structureless,

in human

pointed

by Rainey, referringto calcification of the clear

in shell,
that 'as the development procalcospherites
gresses
the globuleslose their brightand structureless
out

laminae and radiating

artificialcalculi
lines just as the
do, the lines beingmore
distinct when
the globulesare suffering
disintegration?
in the dentine.
This is the case
The decalcified calcospherites
found to have
first coalesced
into larger
are
bodies
and
then to have
become
laminated, showing
marked
concentric
striae.
The
dentine isstained
strongly
is everywhere traversed by these cona deep brown, and
tours.
The circular bodies are of very various sizes,
and

character,and

begin to present

by

114

AIDS

TO

DENTAL

ANATOMY

concentric elements
beingspreadout into the laminae of
the dentine,and their lime-salts becoming equally
diffused
in the calcified matrix.
I have drawn
formed
of the
in one
a
calcospherite
albumin
with those
in the
experiments,to compare
dentine on
the same
This
shows
plate.
example
cisely
presimilar concentric markings in the calcified spherite
to those in the calcoglobulin
substance in the dentine ; it
also shows
faint
of radial lines which
indications
very
be detected in the dentine.
cannot

Calcification

in

the

Enamel.

Many observers have described globularbodies in the

and have especially
drawn
ameloblast
attention to
cells,
indeed
these bodies, which
are
very evident in many
These
authors are of
specimensof developingenamel.
the
that
elements
opinion
calcifying
theyare
separatedby
while quiteconvinced of the appearthe cells ; but others,
ance
of these bodies,do not feel in a position
to offer any
Von
Ebner
shows
positive
opinionas to their nature.
of
in
the
bodies
the
formed
a
drawing
spherical
newly
and
considers
enamelled
from
are
they
prisms,
deposited
salts dialyzedthrough the membrane
present on the
have
I
myself preparations
surface^offormingenamel.
minute
these
o
ver
a
largesurface of the
showing
globules
young enamel.
describes an outer and an inner ameloblastic
Leon Williams
and
consider
that
to
membrane,
a dialysis
appears
of lime salts takes place through these membranes.
I
this
for conhave given elsewhere in
sidering
paper my reasons
e
xist
that these membranes
after
fication
calciactually
has commenced.
the actual conditions
In the endeavour to investigate
takes place,I took some
under which the deposit
ing
developteeth from the crypts of a dried skull of a
molar
the cap
young kangaroo (Macropus rufus\in which over
of
dentine
enamel
laid
of
down.
a rough deposit
onlywas
removed
with a knife
Small piecesof this substance were
and
placed in a drop of glycerineon a slide under
the dissecting
microscope. The enamel at this stage is
and a gentletouch with the needles separates
very friable,

by

APPENDIX

115

each
small fragment into numerous
parallellaminae.
These
laminae are at an acute angle to the surface of the
and are perfectly
enamel
parallel. They can be divided
againand again into very thin layers,and all lie in the

plane.
that crossing
a highermagnifying
power it is seen
of prismsat an angleis a delicate fibrillarsubthe rows
stance,
which can in several placesbe seen
on
projecting
Ebner
shows
either side of the prisms. Von
similar
a
same

With

in a figurein his paper

enamel.
This
on
described by
to be the delicate fibrillar substance
seems
the ameloblasts,
and covers
Andrews
the
as derived from

appearance

laminae. These
or
calcified,
already calcified,
partially
like
the
of
leaves
a
book, apparently
layers are arranged
interleaved by the delicate membrane
described above.
Examination

with

high power showed

layers are
interprismatic
sizes.
of
of
various
Some
calcospherites
these

fibrillar

that many
of
covered
with

these bodies
and show radial markings,
minute granules,
are very large,
of the calcosphenteinto three or four
and a radial splitting
small globularbodies,
which are also seen
extended along the laminae in their
and apparently
result from the disintegraneighbourhood,
tion
of the largerbodies. In many
placesthe separated
with these small
laminae are seen
to be quitecovered

portions.They

are

surrounded

by

globulesat their edges.

somewhat
to find calcospherites
of
surprising
in
this
dimensions
and
to notice
situation,
large
bodies produced
their strong resemblance
to the artificial
These
in albumin.
going
to be underlarge spherites
appear
It
such

was

and
disintegration

are

breakingdown

into smaller

globules.
in calcification from the
further advanced
later
same
animal, but which was not quiteerupted,
a
of the calcific material could be
stage in the deposition
the enamel
studied. In this tooth in which
was
quite
the
and
t
he
smooth
consolidation
on
surface,
polished
stillincomplete
of the substance was
round
; very small
within
the prismsand in the interprismatic
globules are seen
substance a long way into the enamel.
that these
in
It is noticeable
large calcospherites
show
but the very faintest traces
of concentric
enamel
In

tooth

by

AIDS

u6

TO

DENTAL

ANATOMY

have
distinct j radial
very
markings. In dentine, on the contrary, one can see no
of radial markings,although the concentric rings
trace
in enamel, at all
are
spherites
very conspicuous.The
here examined,
events
at the stage in which
they were
and
have
the
clear glass-like
not
are
markedly granular,
of the artificialspherites,
nor
are
they so
appearance
in the earlystages of dentine calcification
clear as those seen
described by Rainey in
as
; and this granularity,
is an
indication of
artificially
produced calcospherites,

lamination,although they

of the globules. The largespaces

disintegration

visible

and in
of them
the same
also show
condition,
into
found
broken
four
I
a
case
one
large spherite
ments
segclose
lying
together.
of the delicate fibrillar matrix in which
The meshes
these large spherites
are
depositedare probably filled
which
Williams conLeon
with the albuminoid
matter
siders
to be poured out by the ameloblast
cells,and we
into this collagenous
that the salts are dialyzed
must
suppose
form
and
there
the
spherites,
calcoorganic substance
in the experimentwith albumin
devised
as
in

some

Professor Harting.
slides of the firstformed enamel, fibres can be
In my
in many
seen
places,crossingthe prisms
very distinctly
at an
angle ; and the whole of the enamel, prisms,and
in this
would seem
substance
to be laid down
prismatic

by

of fibres.
network
It is not always the fact that all traces of this substance
this fibrillarbasis
for in some
lost in calcification,
cases
are
in the completed enamel
of marsumaterial can be seen
pials.
in
which
in
instances
succeeded
I
In two
staining
with fuchsin,
in the one case
the enamel very completely,
be seen
it can
that
and in the other with silver nitrate,
fibrillar
matrix
have escaped calcification
small portionsof the
and have taken the stain deeply.
The fibres are
seen
crossingthe stained tubes in the
tubular enamel
exactlyas in the unstained teased preparations
of the kangaroo.
of the developingtooth
Although it has been often stated that the enamel, once
formed, undergoes no further change,I think there can
be littledoubt, as shown
by this observation and many
the
enamel
when
that
formed and functional is not
others,

by

APPENDIX

117

Pickerill,
as the result of his experiments,

fullyconsolidated.
*

the degree of penetrability

that
of enamel
surface
bears
inverse
from
the
ratio to the
to stains
an
and suggests that an
time the tooth has been erupted,'
take
after-hardeningof the enamel
place by the
may
of salts from the
dialysis
throughNasmyth's membrane
Professor
saliva. As
Harting says : * La solidification
states

secrete
est
tou jours un
phenomene
de
certain
un
secondaire,
quiexige
temps, pendant
espace
lequella secretion continue.'
which lie in this fibrillar
The
larger calcospherites
bodies
with
rounded
somewhat
a
layer are
irregular
de

la

substance

and many
are
margin ; they show radiatinglines,
finely
of more
or
granular. In many parts the minute spherites
less uniform
ing
size,which appear to result from the breakof
these
be
extended in
largerbodies, are seen to
up
and often arranged as a layer
lines,
diagonalor straight,
of minute globuleson the surface of the leaf-like expansions.
The
torn
margins of the fragments of enamel
often seen
with globulesof various
to be crowded
are
of coalesced spherites
of
sizes,and often largemasses
from
their edges.
irregularform are seen
projecting
Simple and compound sphericalbodies are also seen
lying free in the glycerinein which the fragments of

examined.
enamel
are
several examinations
of human
I made
developing
if the same
teeth to see
conditions existed as in the
teeth of marsupials,
but was
in finding
at firstunsuccessful
the large calcospherites,
in
these
teased
although

preparationsmany
prisms
depositwithin them.
globular

showed

Having

the
at

lines of

tiny
lengthobtained

suitable material,I found

that similar bodies were

present to those found in marsupial enamel, and I think
there is strong evidence that here also the calcification
is due to this deposit.The
of the cement
substance
taken
from
all
drawings were
preparationsin glycerine
in
the
those from the marsupial
same
manner
as
procured
teeth.
Most
of them
from
the developing teeth
are
from the tooth sac in the jaw of a ten-months-old
removed
subjectand preserved for a few weeks in spirit. By
I was
dividingone of these developingteeth vertically
able to take "minute scrapingswith a sharp knife persome

by

u8

AIDS

TO

ANATOMY

DENTAL

its outer
surface of the enamel
at
that the largerbodies lay for the most
both
the outer
the surface and justbeneath
part
upon
clear layerof the enamel
within its substance.
These largebodies in human
enamel
probablybear the
substance
relation to calcification of the cement
same
as
do those of the kangaroo. The difference between
the

pendicularto the
margin,and found

enamels

two

to be

appears

the

organic matrix is more

the spherites
calcified,
can

that in the

marsupial,where
developedand probablyless
be seen
deep in the enamel

enamel, where this material is

; but in human
less marked
the largercalcospherites
and sooner
calcified,
are
only found in the forming layersof enamel near the
The
surface.
of globules of various sizes
large mass
substance

the developof the enamel

between
ing
and
dentine
in
which
the
enamel
of
tooth
a
cusp
formed
small open
the pulp. These
a
large
cap over
bodies,which were found to react to polarizedlightas
do the artificialspherites,
would
to be deposited
appear
of the
membrane
inner
ameloblastic
under
the
directly
the
enamel
as
organ, where they have probablyformed
were

on

the surface

result of the dialysis

the ameloblasts
of salts from
or
the fluid by which
It is evident
they are surrounded.
that these bodies cannot
have been
present, as such,
in
the ameloblast
of
them
cells, many
having a
diameter coveringfour or five enamel
prisms; but they
clear globules.
of all sizes,
from this to very minute
In this situation it is noticeable that,as at the margin of
show
the forming dentine,the spherites
clear and
are
tive,
; they are highlyrefracvery littleindication of structure
are

careful focussingare seen

to be globular.
Many of the isolated bodies show denser portionsin
and here and
few a faint indication of striae,
some
places,
there a sphere is found breakingup into four segments, as
in the kangaroo. Many of the small globuleshave
a
central prominence like the boss of a shield. Many of
the largerones
oval, a form I have not met with in
are
and

the

by

and
marsupials,

last appear

to

be

stronglygranular; these
do not
and
to
react
disintegrating,
a

few

are

polarizedlight.
In
very

lookingfor these
importantto make

developingenamel it is
as
by this
polarizedlight,

bodies in
use

of

by

APPENDIX

with

119

from
distinguished
which
have found
substances,such as leucocytes,
their way in from the tooth sac.
is
it
think
I
able
very prob-

they

means

can

certaintybe

other

that the stage of calcification in which these large

found is a very passingone, the larger
are
calcospherites
bodies very quicklybreakingup, for in the enamel
of the
I
have
found
in
from
the
kangaroo
that, slides prepared
same
part of the tooth,three or four of the laminae will
be crowded
with the round bodies* and twenty or thirty
other

fragmentsin

the same
preparation
may show none.
in developactual presence of largecalcospherites
ing
enamel
other than the regularglobulesin the prisms
Professor
been hitherto described.
not
has, I believe,
Underwood, however, described interglobular
spaces in
human
in
of
in
of
erosion
the
cases
enamel,
1896, some
the enamel.
The
globules surroundingthe spaces are
the same
as in the
very small,but have much
appearance
that in
It would appear
interglobular
spaces in dentine.
The

these

teeth the action of the agents (whether

pathological
acids or not is disputed)
had justreached a stage in which
the process of development of the enamel
was
revealed,

acids of caries reveal the

structure
calcospheritesin dentine. They were
in these cases
of erosion.
We
can
only seen
hardly,
globular
to
however, look upon these as
corresponding interdue to an
ruption
interspaces in dentine,which are
of calcificationduringthe formation of the tooth ;

in the

same

manner

as

the

of the

and

if there

were

similar

in
interruption

the enamel

we
deposit,

should expect to find these spaces in healthy

teeth,as is the case in dentine.
One
would
feel more
disposedto consider that the
of the
erosion process had partially
dissolved away some
lime salts of the matrix in an irregular
leaving
manner,
the

presumably denser
globulardeposit.

portions representedby

the

first large calcospherites

which I have described
found
in marsupial enamel, marsupial enamel,
were
from its somewhat
a good
condition,affording
primitive
of
of
the
the
earlystages
process,
opportunity studying
calcified
obscured
in
the
which are
much
more
very
The

of human
teeth,although I have
similar deposittakes place here also.

enamel

shown

by

that

i2o

AIDS

TO

ANATOMY

DENTAL

all events, the greater part of their

be
the firstcalcifiedmaterial formed,
substance,appear to
probablyby the depositof lime-salts in the Tomes' processes
of the ameloblasts.
The lime-salts are deposited
in the interprismatic
stance
subof
formation
the
as
probably by
calcospherites,
stance
carbonate and phosphate of lime within this subnascent
of
by the chemical combination with its contents
The

prisms,or,

at

salts dialyzed
through the inner ameloblastic membrane,
the outer
surface of the colloidal matrix, which such
or
membrane
would represent.
In marsupialenamel, as I have shown
tion
above, calcificatakes placein the laminae between
the prisms,and I
think it is very evident that these largecalcospherites
are
of the prisms upon which
produced quiteindependently
in rows
The
lie.
calcified
of minute
prisms are
they
be formed within the
and are the firstpart
Tomes' processes of the ameloblasts,
of the enamel
calcific
receive
the
to
deposit; while the
is formed
substance
interprismatic
subsequentlyin the
organic fibrillarmatrix,which is seen
throughout the
in
enamel
the kangaroo. The calcification in marsupial
does not, however,proceed to complete obliteration
enamel
of the cement
substance,much of which remains
uncalcified,
as the alcoholic fuchsin method
clearlyshows.
formed
It would appear that,in less perfectly
enamels,
is
that
substance
and
the
denser
evident,
cement
a
very
and more
completelyformed the enamel, the less is this
fication,
substance in evidence,its existence beingveiled by calciin
Primates
the
that
of
enamel
the
evidence
the
so
in the perfectly
formed
of its presence
portionsof the
tissue is very slight
absent.
or altogether
and of dentine
of
enamel
t
he
calcification
In explaining
by the depositof lime-salts in the form of calcospherites
with one
The
met
we
are
analyses of
great difficulty.
enamel
of
show
dentine and
a very large preponderance
carbonates.
In dentine, accordingto
over
phosphates
of Berzelius,
the analysis
phosphatesform 62 per cent, of
the mineral
In
matter, and carbonates
5*50 per cent.
Bibra's
for
enamel
adult
enamel
von
analysisgives
8982
calcium phosphate and fluoride,
and only4*37 of calcium
carbonate.

bodies, which
spherical

appear

to

by

AIDS

122

TO

ANATOMY

DENTAL

acid lime-salts.' The same

conditions probablyapplyto
the other calcified organictissues of the body, as enamel
and dentine.
The conditions in young growing animals,
nently
emiwhere
the metabolic changes are very active,are
and
of carbonic acid,
favourable to the production
it is quiteconceivable,
that in the
and in fact probable,
bonate
earlystages of enamel and dentine formation the carand
of the phosphates,
may be greatlyin excess
be easily
thus the depositof lime-salts as calcospherites
understood.
The after alteration of the chemical
sition
compoof the material in which calcificationoccurs, and the
addition of salts of phosphorus from the blood, would
spherites
for the breakingup of the calcoto account
serve
perhaps
the ultimate hard
as calcificationadvances,and

condition of the dentine and

the

semi

dense

-crystalline

completedstate.
aspect of the enamel in its fully
It is evident from
numerous
experiments,especially
found
those of Haltingand Ord, that in the proportions
in finished bone and

teeth the lime-salts are not deposited

in albumin
do not know
in the globularform ; but we
material
the exact
chemical constitution of the exuded
in which this deposittakes placein the livingbody, and
referred to by Mr. Tomes.
that the salt in teeth and bone is
of three equivalents
of calcium

as

Hoppe-Seylerconsiders
salt,consisting
a double
phosphate,with one of

the
in
be effective
causinga
in
whatever
but
be
globular depositmight
considered,
allow of
conditions which
way we explainthe chemical
the evidence that in
their formation,we
cannot
escape
in
both dentine and enamel
the lime-salts are deposited

in this combination

calcium carbonate.
Whether
influence of the carbonate
may

the

globularform.
With regardto the

other mineral constituents of enamel,

the fluoride of calcium is present in very small quantity,
of from
but magnesium salts are present in the proportion
Mr. Bennett mentions an experiment
1 34 to " 2*55 P^
cent"
added to
of Rainey'sin which magnesium phosphatewas
and in
in similar proportions
solutions of carbonate
gum
which no globular bodies were
produced ; but Professor
Harting describes an experimentin albumin to which a

magnesium
the

salt

was

was
precipitate

added, and

composed

he

in which

of the

that

bodies,as

same

found

by

to

APPENDIX

123

form

and structure,as those which had formed

without
the addition of magnesia; the only effect he remarked
that the formation of the precipitate
was
was
delayed.
We
thus see
that, in both dentine and enamel, the
in the spherical
form throughout,
lime-salts are deposited
forms
the larger
eventuallybreaking and being lost in
the calcified

We

mass.

up
have, as in Rainey'sexperiments,

coalescence
of the small globularparticles
leading to
integration
the production
followed by disof the largecalcospherite,
of these forms and a deposition
in the formed
tissue in a more
homogeneous condition. As pointedout
integration.
by Mr. Rainey,conditions accelerate the process of dis-

found
He
sisting
conthat, when calcospherites
and
of carbonate
placed in a
phosphate were
denser medium
than that in which
produced,
they were
did
he
but
not
they became
rapidly disintegrated,
of
find this occur
with those
formed
of
carbonate
lime

A
alone.
already formed

those
fresh depositof spheritesupon
of those first
also caused
disintegration

deposited.
In some
experimentswhich
I found
albumin

the

I undertook

for the purpose

crystalsof phosphate to
of
formed
containingalready
calcospherites
carbonate
preventedtheir further formation,and caused
of those alreadyformed.
disintegration
is
therefore noticeable that the salts of lime in the
It
condition of calcospherites
not in a very stable state
are
of equilibrium,
coalesce and disintegrate
they alternately
in
under
the effect of various conditions in the medium
which

that

addition

of

This fact has an

important
in the
conditions
of
aggregation
bearingupon
varying
of livingorganisms.
structures
calcifying

they are

produced.

their

The

existence

Sheath

of

Neumann.
of this sheath

has long
the first
of controversy. John Tomes
was
a matter
fibril
dentinal
of
in
the
soft
to describe the presence
a
tubes.
He showed
that these fibrils ran in tubules in the

The
been

dentinal

matrix,and

membrane,
and

or

named

non-existence

were

providedwith

definite

lining

the sheath firstfully

investigated
by Neumann
him.
shows
This
after
liningmembrane

by

AIDS

124

TO

DENTAL

greatresistance to the action

of dentine by
in preparations
matrix

substance

by

ANATOMY
of acids,and can be isolated
vening
the destruction of the interof
the prolongedoperation

strong acids.
considered that the
membrane
of
to the dentinal canals
a lining
appearance
Sudduth
is due to effects of refraction,
and Magit6t,
and,
Romer
more
recently,
deny its existence. Mr. Charles
*
Tomes
that the dentinal
be remembered
says : It must
sheaths
can
by processes
only be fullydemonstrated
which amount
destruction of the dentine,and
to a partial
that they are
therefore in some
degree, at all events,
artificial
it
be
existence until they
have
real
no
they
;
may
called into existence by the action of these agents.'
are
He
in stainingthem
with
has, however, succeeded
and
with
silver
have
Rose
and I
ceeded
sucnigrosin,
nitrate,
in doing the same
in many
preparations.
Professor Romer
considers the tubes to be simply
true walls ; the
spaces in the dentinal matrix with no
so-called Neumann's
sheath he affirms to be a part of
and considers that in the preparations in
the soft fibril,
which it is claimed that the sheath is stained,
what is
is
fibril.
the outer surface of the soft
He
stained
really
shows
in
the
illustrations
his
the
to
a
figure
paper on
in
which
in
fibrilhas
fallen
the
section
transverse
subject,
in the dentine with no wall
out, and there is a perforation
to it. Judging
the
it would seem
by
illustration,
probable
that,in this instance, the fibrilonly is stained,and the
sheath not ; as I have several preparations
which show
outer
an
ringenclosinganother dark-stained ring,which
is the border of the dentinal fibril.
It is somewhat
of the
difficultto obtain good staining
Neumann
having
sheath,my most successful preparations
been from young
teeth stained by a modification of the
method
of Ramon
the
y Cajal.In these preparations
sheath of Neumann
is stained a deep black,and the darkstained fibrilis seen
lyingwithin the ring,an appearance
which
be explainedon the assumptionthat the
cannot
supposed sheath is the outer border of the stained fibril.*
Some

observers, however, have

Mr.

this very
Mummery exhibited a photograph which showed
distinctly,
and, moreover,
definite proof of the
still more
gave
existence of the sheath, for at the thin margin of the section the

by

APPENDIX
The

125

section shows
line,there
a sharplydefined
diffused stainingof the surrounding matrix,
that this wall consists of some
material which

tube

being no
showing

takes the stain quitedistinctly

from the rest of the matrix.
In the specimen in which the segments of the tubes are
and the fibril has entirely
fallen away, there is no
seen
ness
of any disturbance of the image from the thickpossibility
The
sheath
of the section or from optical
refraction.
is also very evident in the projecting
ends of tubes which
are

cut

obliquely.

Erwin
Hohl
section,
pointedout that, in longitudinal
with suitable
while the sheath of Neumann
is very evident,
in the calcifiedportion
it is not to
of the dentine,
staining,
in the odontogenetic
be seen
zone, where the tubes appear
definite walls. A section of a young
to have no
growing
tooth, where the uncalcified zone is of great width, shows
this very distinctly,
the section being a portionof the
Hohl
that alluded to above.
same
siders
conas
preparation
*
the dependence of the
that this fact pointsto
sheath of Neumann
We
calcifieddentine substance.'
on
further,and suggest
might perhapscarry this statement
that this sheath of the tubes is concerned
in the calcifying
as a dialyzing
membrane, through
process, and may serve
which dialysis
takes place.
In developing
has been described
enamel
a membrane
between the ameloblast
cells and the forming enamel,
and also between
medium
interthe enamel
cells and the stratum
existence
of the enamel
of memThe
branes
organ.
in these situations has been the subject
of much

controversy.
is inclined to look upon
these
Mr. Charles Tomes
in
membranes
artificial
sections
b
ut
as
productions,
which have not been treated with either alcohol or acid
At the base of the tooth-germ,
to be seen.
theyare easily
has not yet commenced,
where the formation of enamel
of circles were
surrounded
seen
by a black-stained line,
the stained fibril had fallen away
altogether. It does not
this
at the margin of the
to
for
seem
possible account
appearance
definite
in
than
section
a
as
proof of the actual
any other way
sheath
of Neumann,
called the
existence of the structure
and,
of its existence as an integralpart of the dentine lining
moreover,

segments

and

the tube.

by

AIDS

126

TO

DENTAL

ANATOMY

the inner membrane

be seen, and the outer ameloblastic
cannot
membrane
is evidently
the
cells
quiteabsent,as
of the stratum
intermedium
are
mingled with the ameloof calcification
blasts,althoughafter the commencement
for
they are distinctly
separatedas a layerof cells,
lying,
the most
the
less
ameloat
to
or
rightangles
part, more
blasts. They certainly
to be separatedby what
appear
like a continuous membrane.
seems
ameloblastic memsection shows
the outer
Another
brane
which
of
from
of the enamel
the
Macropus
organ
ameloblasts had fallen away, and the cells of the stratum
intermedium
to be separated
brane
are
seen
by a definite memfrom the regionof the ameloblasts.
In two other
of the cells from
specimensI found a similar separation
the inner ameloblastic membrane.
A
of
small portionof the dried-up enamel
organ
Macropus which was found with the fragmentsof enamel
in the teased
preparationsin glycerinepreviously
described appears to represent the inner ameloblastic
of the honeycomb conmembrane
and
the membrane
nected
by the shrivelled Tomes' processes, and would

appear

to

be strong evidence

of the existence of these

structures.

in the dentine
regardto the sheath of Neumann
fine branches
of the dentinal tubes everywhere
the dentine,and must eitherpenetrate the sheath,
traverse
or be themselves
provided with a similar sheath. It is
difficult
to determine this pointin such minute structures,
very
With

the

althoughtheir sharpoutlines might suggest

latter is the

that the

case.

which
the arguments
view
above
the
stated
: the
to support
are
appear
evidence of the real existence of the sheath of Neumann,
its absence in the layer of collagenous
material in which
calcificationis about to take place,but has not yet commenced,
and its very conspicuouspresence
in young
growing dentine. It also suggests a function for a
structure, the uses of which have not been understood.
In growing dentine, as is well known, the calcified
dentine is separated
from the odontoblast
cells by the
of a substance considered
odontogeneticzone, consisting
of
to be elaborated
the
cells
the pulp and upon which
by
With

respect

to

the

dentine

would

by

APPENDIX

calcific

the

127

encroaches

matter

from

above

in rounded

contours.

calcific depositnever
proceeds direct from the
stand
odontoblasts,and it has been a little difficultto underin
this
direction ;
why calcification takes placeonly
but on the view that the lime-salts conveyed by the fibril
into this
are
dialyzedthrough the Nasmyth's membrane
understood.
itcan, I think,be more
material,
easily
collagenous
This

Conclusions.
I have
dentine

endeavoured

that calcificationin both

enamel
great part a physical
that
the
actual
phenomenon,
deposit in both tissues
in the form of calcospherites,
and that the process
occurs
in mammalian
tissues is identical in many
pointswith the
same
processes occurringin lower organisms.
While no doubt it is true that no artificially
conducted
in the
can
accuratelyrepresent what occurs
experiment
between
the substances
livingbody, the resemblances
in
and
those found in the
these experiments
produced
dentine
and enamel
are
very strong, and the objections
based
to this comparison have
on
been, I think,chiefly
the chemical difficulty,
the compositionof the dentine
and enamel
such
not being
as to allow the assumption of
the spherical
form in the deposited
lime-salts. As I have,
form is not
to show, the spherical
however, endeavoured
retained
in completed well-formed
dentine or enamel,
it
and for the reasons
is
given
highlyprobable that when
in the colloid material of the
the lime-salts are deposited
matrix substance they are true calcospherites,
ing
correspondto those produced in the experiments,
the carbonate
of calcium being combined
with a small proportion
only
of phosphate.
That some
altered chemical conditions of this nature
must

occur

and

show
is in

to

is, I think, evidenced

by the fact,which I
parable
comlarge calcospherites

have

described above, that

in every respect to those found in the organic
solutions employed in the experiments
in
to be seen
are
These largebodies only persist
in this state
young enamel.

of

aggregationfor

short

time,and

as

phosphates are

by

128
added

AIDS

TO

DENTAL

ANATOMY

and the resulting

form the finally
consolidated enamel.
disintegrated
particles
In dentine also,except where the process of tion
calcificahas been arrested at the interglobular
spaces, a similar
takes
breaking down of the first formed calcospherites
t
he
their
evidence
of
former
place, only
presence beingthe
lamination of the dentine,
which is also finally
obscured by
full calcificationin the completedtissue.
The evidences that the odontoblast
cells are the principal
active agents in calcification of the dentine are, I
venture
conclusive
the similar
to think, quite as
as
evidences of the function of the ameloblasts
a function
which has not yet been doubted
in their case.
As nerve
fibres traverse
the dentinal canals, the fibril
be looked
cannot
transmitter of nervous
as
a
upon
the
cells
have
impulses
granular contents, they lie in a
rich plexus of bloodvessels,
and we
know
that active
secretion is associated with an
increased blood-supply.
Like other secretingcells,
entiated
they are large,and well differfrom the surrounding
tissue elements,and they
retain their full size and characteristics duringthe actual
of the dentine in healthyteeth.
deposition
The
of the cell in the form
protoplasmicprolongation
of the dentinal
fibrilwould be considered to share in the
functions of the cell of which it forms a part, and there
is transmitted
are
strong evidences that calcific matter
by the fibril. The translucent zone in caries,which a
greatweight of evidence suggests is due to calcification
the tubes, and the peripheral
occlusion of the tubes
in
extension
of the cell
this
to
on
exposed surfaces,point
protoplasm being the channel by which lime-salts are
conveyed to the dentine.
If I am
rightin supposingthat the sheath of Neumann
a
serves
as
membrane, the comprehensionof
dialyzing
the process of calcification in the dentine is somewhat
simplified.The odontoblast cell,either alone or in
with other cells in the pulp which send processes
common
basis substance
to the dentine, would
depositthe gelatinous
which
calcification takes
in
stance
place- the subwhich
forms
the odontogenetic
limethe
zone
salts taken up from the circulating
blood by the secreting
cells would be transmitted by the fibril,
and passingby
from

the blood

they break down,

"

"

"

by

INDEX

Abortive

Boar, the wild, 4, 5, 8, 103

teeth, i
of

Absorption

the

Bodecker's

temporary

teeth, 91
Adamantoblasts,
iEiobates, 71
of

Boll
tell

to

man,

the

on

missing teeth

enamel,

pulp, 74

cells,35

in

in,

110

97

Ameloblasts, 23, 36, 57, 116,

of

Analyses
enamel,
Ancient
Andrews

servations
ob-

corpuscles,35
development, 3
Brachiopoda, calcification

the, 97
Albrecht

the

on

Bone

how
individual,

an

human

58

23

of, 71
plicidentine

Age

microscopical
of

dentine

50,

Brown

striae of Retzius, 63
Buds, post-permanent, 18
pre-milk,18

120

of

and

120

the teeth of, 104

enamel,
on
115
Angle, the facial,84

Cajal,Ramon

Anoplotherium,8

Calcification,2, 31

man,

of

Arrangement

teeth

in

64,
in
of
of
of
of

man,

the, 92
Assimilation

to

surroundings,11,

12

y, method
112, 124

the Crustacea,

ing,
of stain-

no

bone, 35
cementum,
43
dental tissues,3

dentine, 40, xii, 113

of enamel, 36, 107, 114
of the enamel
organ, 57
Baume
teeth
in
the chemistry of, 45
the missing
on
I Calcified teeth, 7
100
man,
Bealb on the enamel
tissues, A. S. Underwood's
organ, 24
I
I
table of the, 45
Bennett, F. J.,on calcification,

Baleen

whales, 1, 9, ix
Basio-nasal
line,the, 85

I2T,

Berzelius

45,

Bibra

I Calcoglobulin,
33, 34

122

the chemical
of the calcified

on

46, 120

position
com-

ICalcospherites,
3,

tissues,

32,

33,

110,

"5

I Camper

lines,84

Canal iculi,53
the chemical composion
tion
of the calcified tissues,
45, ;Canines, definition of the, 95
of the, 98
46, 120
description
the
sexual
character
Bicuspids,the eruption of the,
"

certain, 103

9i

130

by

of

INDEX

Capybara,the

molars

131
germ, the, 19
hard, 64
organ, the, 24
papilla,the, 19, 24
sensitive,64, 74

Dentine

of, 103

Carnassial teeth, 101

Carrion crow, the, 12

hyaline,29
Cartilage,
ossificationin, 35
the formation

Dermal

of, 35

Castration,effects

early, 5,

of

Development of cells,the, 1

103

Cat, the dental

formula

of

the,

of teeth,the, 3, 14
of the, 29
summary
of the human
embryo, the,

101

Cell

appendages,

Dermis, 6

development,1

Cement

organ, the, 11, 27

3, 29, 43, 44, 48, 53

Cementum,

Cestracion,64
in the jaws before
after birth,87, 88, 89
Chemical
composition of

Changes

and

mouth, 79
of,29
summary
Diet and dentition,101
21
Differentiation,

the

calcified tissues,45
of dentine, 51
of enamel, 46
acid
Chondrosulphuric

79
of the

teeth,the, 8
of the,

Disappearinghuman
the dental

Dog,

formula

101

Dugong, the, 1, 5
in ossification,

Dursy

development,

on

18

35
the

Cleft in cleft pilate,

the, 80
Concomitant
variation,10

Elephant,

Crustacea, the,no

Embryo,

the

human,

79

Curves
and

of dentinal fibrils,
primary

secondary,69

of the
formula
in the
four impulses at work
animal world, 9
Definition of teeth,the, 1
of human
teeth,86

Darwin's

brief

Deinoceros, 10
Deinotherium, 5
Dental follicle,
the, 28, 29
formulae of typicalcarnivora,
101

groove, the, 16
lamina, the, 17, 18
periosteum,the, 77
tissues,calcificationof the, 3
the uncalcified,74
Dentinal fibrils,
65, 70, 108
tubes, 64, 68
Dentine, 3, 44, 64, 73

of

molars

the,

103

developmentof the

Embryonic jaw, the, 16

Enamel, 3, 19, 20,
analysisof, 48
cells,23
fibres,60, 61

44, 55

marsupials,59, 64,

in

114,

118

interglobular
spaces in, 119
the, 20, 21,
pigment in, 59, 63
prisms,60, 61
organ,

22, 57

spindles,63
the

in

membranes

ing
develop-

125
the minute
structure
of, 60
the process of calcification
in, 107, 114

tubed, 64

Endognathion,the, 80
Environment, adaptationto,

calcificationof,40, 107, 113

Epiblast,the,6

cells,26

Epidermis,the, 6

Charles S. Tomes' s table of

the varieties of, 73

Epithelial sheath
the,29

of

by

12

Hertwig,

AIDS

*32

TO. DENTAL

Epithelium,6, 7, 16, 17,

19,

Highmore,

22

columnar, 29
of

the enamel
external and

the

organ,

of, 82
sheath

of

of the,

101

the molars of the, 103

Howes
the enamel
on
organ,

47

7, 15

Exognathion,80

of

102

Horny teeth,7
Horse, the.dental formula

22, 23

Evolution,2,

the antrum
the

Hohl, Erwin, on
Newmann,
125

internal,6, Horns,

Eruption,15, 86, 90
Evanst Lovatt, analysis
enamel, 37,

ANATOMY

Huxley's

24

fpreforma-

membrana

tiva,34, 38, 52
Facial angle,the, 84
' "
False
or horny teeth,7
"

Fenestrated membrane,

Imbrication

the, 38

Fibres, Sharpey's,
44
the

of the enamel, 61

course

Fibrils,dentinal,64, 65, 108

the

functions
trophic

the dental, 28,

Follicle,

Foraminated
Formalin

and

growth

Interglobular
spaces,

66
,
at birth,88
after birth, changes in the,

Jaws

29

of teeth,

89
before birth,changes in

the, i, 85
Formula

40, 66

ossification,
35

Intermembranous
1 1

sheet,the, 38
68
a fixative,

as

Formation

of ,

lines,59
Incisors,93, 95, 98, 99
Index, the gnathic,84

the,

87

of teeth in man,
the typical,
96

the, 92

Kangaroo,

enamel

of the, 114,

118

Keith, Professor Arthur, on the

chemical
the
cleft in cleft palate,
80
position
comGalippe
of the teeth,46
Klein on calcification,
41
of teeth,the formation of
Germs
enamel spindles,
on
63
the dentine organ, 27
on
the, 15
the pulp,74
on
post-permanent and preKollicker on
the calcification of
milk, 18
Gibbs on eruption,18, 87
enamel, 38, 60
Glands of Serres,the, 29, 77
the
on
brane,
periodontalmemGnathic index, the, 84
79
the sheaths of Neumann,
Granular layer,
on
the, 65
Groove, the dental, 16
69
the varicosity
of enamel
Growth
of teeth,the, 1
on
Gubernaculum, the, 291
prisms,61
Galeus, 64

on

Gum,

the, 77
La bio-dental band, the, 17

Hake, the, 73
Harting on calcification,
32, 33,
116, 117, 121, 122, 123

in,
Labyrinthodon, plicidentine

on
no
calcospherites,
Heidelbergman, 105

encapsuled,54
of cementum,
53
Lamina, the dental, 17, 29
of the
Legros on the vascularity

Heitzmann

on

enamel, 58

sheath of,
Hertwig,the epithelial
29

71.73

Lacunae, 35

enamel

organ,

by

24

INDEX_
Leporidse,the

Mesoblast, the, 6

the

of

course

Microscopical observations

fibres in, 62
enamel
Lime
in teeth,the amount
of, 7
Lime-salts
in the ameloblastic

membranes,
the

Wellings, 57
Milk-teeth, the, 30, 87, 91

of, 3
deposition

the, 97,
Missingpre-molars,

acid

process

64
gold-staining,

of

the

on

on
on
on

cement

on

organ, 20, 24
the cementum,
29
the internal epithelium,

on

23
the odontoblasts, 27
the pulp, 74
the sheaths of Neumann,

on
on

78
of enamel

course

on

'

on

the

brain

and

vasodentine

on

in,72

eboris,the, 26, 75

of
and

formation

dentine,
bone,
enamel, 32, 107
the mollusca, in
the sheath

of

mann,
Neu-

of the
the staining]
Neumann
sheath, 124

Musk

deer, the, 10
Mustelidae, the dental
of the, 101

formula

in, 71, 73
plicidentine
I Myliobates,

bones, 35

I
23,

24,

29,

44"Si
the periodontal,
77
125

the

123

Nasmyth's, 20,
Membranes

the Crustacea, no
of limethe dialysis

salts,108
on

on

the,34
preformativa,
Membrane

veloping
de-

enamel, 125
in dentine, 75

the brachiopoda,in
the calcification of

Meckel's cartilage,
29
Membrana

in

membranes

on

on

face,the, 83, 84

Megatherium,

dentine, 64
marsupialenamel, 38

dentine, 42, 107

the vascularity
of the
enamel
organ, 24
Malassez
enamel
on
buds, 21
the glands of Serres,77
on
the periodontal membrane,
on

fibres in, 62
Matrix, the, 108
Measurement
of

59
hard

nerves

on

on

the

development of the,

on

65, 69, 124

Manatee,

the,

in

membrane, the, 6, 16
tinal
Mummery, J. Howard, on denfibres in enamel,

38
the

103

79
Mucous

tine,
calcificationof den-

the calcification of enamel

on

rhino-

no

Mouth,

brane,
mem-

42
on

ceros,

Mollusca, calcification

52
the

on

the elephant, capybara,

horse, tapir, and

Machairodus, the dental formula

of,101
Macropus rufus, 114, 126
Magitdt on
Nasmyth's

100

teeth in man,
the, 97, 100
definition of the, 95
Molars,
I
for
descriptionof the, 99

Lippen furche, 17
formic

on

the calcification of the enamel

and
by Underwood
organ

114

Lines of Owen, 69
of Shreger,69
Lowit's

'33

in developing
enamel,

Narwhal, the non-erupting teeth

of the female, 1
Nasmyth, Alexander, 51

Nasmyth's membrane"
24. 29, 44, 51, 117

by

20,

23,

TO

DENTAL

and

growth

AIDS

134

Natural selection,7

Rainey

Nature, formation
of teeth,the, 1
Neanderthal

man,

ANATOMY
calcification,
31, 32,

on

116,

122

121,

coalescence,

molecular

on

105

112

no,

gold staining,64

Nephrops, calcification in, no

Ranvier's

Neumann
on
dentine, 65, 123
the sheath of, 27, 51, 65, 69,

Raschkow, the plexus of, 74

Retzius, brown striae of, 63
the striations of enamel
prisms,61
Rhinoceros, the molars of the,

108, 123

on

Odontoblasts,26, 27,
Ord

42, 108

calcification,
32,
Oreodon, 96
on

122

103
enamel in, 62
the sheath
on

Rodents,

Orthognathous,85

Romer

Ossification,
35

Osteoblasts,34
Osteodentine,43,
Owen

Rose

mann,
Neu-

124
dental lamina, 17

on

development. 29
the
staining

on

72, 104
the definition of teeth,1

on

of

on

of

the

dentinal sheaths, 124

Paul

Nasmyth's membrane,

on

Sac, the dental, 28, 29

24. 52. S3
Periodontal
membrane,

the, 45,

Sargus,
enamel

77

osmotic

4. 7i.

the
penetration of
membranes
by calcium

incisors,103
Scalpriform
Schafer

salts,109
Pickerill

on

enamel, 59, 117

Pigment in enamel, 59, 63
Piltdown
mandible, the, 104
Plexus of Raschkow, the, 74
Plicidentine,43, 71

prisms

Post-permanent buds,
Poulton
enamel

on

the
organ,

18
the

on

the

Sheath

24

Primeval

missing,97,
curves,
69
man,

the

of enamel, 46
the salt in teeth and

of

the

Henwig,

epithelial,

29
of Neumann,

the, 27, 51, 64,

69, 123
of Neumann,

the

of
staining

the, 124

100

Shreger, the
teeth

Prisms, enamel, 60
the sawfish,4, 71
Pristis,

contour

of, Smilodon, the dental

lines

of,69

formula of,

101

104

Prognathism, 84
Pulp, the, 44, 74
Pulp-stones,76

dentine, 74

bone, 122
Shark, the, 5, 16, 64
Sharpey'sfibres,44

Prawn, the carapace of the, no

Pre-milk buds, 18
Pre-molars, definition of the, 95
of the, 98, 99
description

Primary

in

nerves

Sensitive dentine, 64, 74, 108

Serres, the glandsof, 29, 77
Seyler,Hoppe, on the analysis

in the,

of
vascularity

calcification,37

on

Sciuridae,62

on

Porcupine,enamel
62

in, 63

Sawfish, the rostral teeth of the,

Periosteum,the dental, 45, 77

Persistent growth, teeth of, 103

Philip on

fish,

sheep'shead

the

Smith, Hopewell,on

cementum,

54.70,74
dentine

on

ti

cells,20

on
enamel, 59, 60 J
! Spaces, interglobular,
63

by

INDEX

135
chemical
tion
composiof the calcified,
45
the chemical
composition of
the uncalcified,
74

Spherodus,enamel in, 64*

enamel, 63
Spindles,

Tissues, the

Stellate cells,22

reticulum, the, 22, 23, 57

dental formula of the, Tomes'

Stoats,the

foraminated

sheet, 38

126
of
dentine,
S., analysis

processes, 36, 37,

101-

intermedium, the,

Stratum

Tomes,

22,

23. 57
Striae of Retzius, brown, 63
Striations of enamel
prisms,60
of enamel
rods, 56
Sudduth
the sheath of Neumann,
on

C.

5
of enamel,

albumen,

on

ameloblasts, 37
calcification,
36, 38,

teeth,18
Sus Babyrussa, 8
Swinhoe's
water-deer, the horns
and canines of, 102

on

dentine, 51

on

development, 29

on

enamel, 24, 29, 46, 60

Tapir, the

molars of the, 103

in the, 72
vasodentine

Teeth, abortive, 1,
or

on

on

70, 74
the

on

horny, 7

formula of the, 92
in man,
!
the missing,
in man,
100
97,
in primevalman,
the,'
104
of gramnivorous and
nivorous
car-

peoples,104
of the development
summary
of, 29
18
supernumerary,
the chemical composition of,
the definition of, 1, 95
the disappearinghuman, 8
the eruptionof,86
the form
and
arrangement

of,92
the

homologiesof, 92
nature,

the
the
11

uses
"

true

of, 4, 5
calcified,
7

foraminated

the functionof the

odontoblast
cells,108
the membranes
in developing

enamel, 125
on

on

the sheaths

69, 124
the
striations
enamel
prisms, 61
the tissues,52

of

vasodentine, 43
table of the varieties of

dentine, 73
Tomes, Sir John, on ameloblasts,

36
external

epithelium,

24

Nasmyth's

on

or

brane,
mem-

52
on

the

dentinal

fibril,

65, 70,

70

Tiger,dental formula
Marrett,

on

post-permanent

Tissues,the, 42,

mann,
of Neu-

on

on

Thrombosis, 76

and

on

formula, 96
typical

Thrombus,
Timms,

on

formation, and

growth of,

the

sheet, 38

on

45

brane,
mem-

52
chemical
como
f
position dentine,51
the chemical
compositionof enamel, 46
the dentinal fibril,
65,

|
I
,
!

Nasmyth's

on

on

the

33

42

124

false"

46

on

on

Supernumerary or supplemental

"

120,

44

of the, 101

pre-milk
teeth, 18

on

123
of
the nature

dentine,

64
Trophic functions
fibrils,
11, 66

of the dentinal

by

A L

DENT

TO

AIDS

136

A NA

TOM

Y
internal and

the

Tubed

Waldeyer

Underwood,

external epithelium,23
of enamel
the varicosity
on
prisms,61
Walrus, the use of the incisors of

dentine, 64, 104

enamel, 64
Typicalformula,the, 96
A.

of

S., method

the, 5

preparationof enamel
for

analysis,
48

Wedl

52

Weil

on

enamel, 55. 119

interglobularspaces

on

119
the brown

43

the

on

on

errors
histological

due to decalcifying
agents,
68
on
and Underwood
enamel,

striae of

calcification

enamel,

odontoblasts,
74

Wellings ou

Retzius,63
on

of

55
enamel
organ, 22
external epithelium
,

40

the enamel
organ, 22
the external epithelium,
23
of the

table

composition

23

micioscopical
chemical
the
of

on

of teeth,the, 4, 5

Variation,concomitant,

57

Whale, the, 1, 9,
Williams, Leon,

10

patches,70

Vascular

Vasodentine, 43. 71, 73, 104

in fishes,67
Von

Ebner
on

the formation

on

on

of I
34,

11

calcification

on

in the
116
the
enamel

enamel,

of
61
prisms,60,
on

tion,
calcifica-

121

'

brane,

on

Nasmyth's

20, 23, 52

Bailliire,Tindall

UNIV.

and

OF

NOV

mem-

the, 17, 18, 29

Zahnleiste,
IZygobates, the pulp in, 71

Cox, 8, Henrietta

Street,Covent

MICHIOAH,

6 11111

by

114,

striations

Sims,

Woodhead,

dentine, 108
enamel, 114, 115

Waldeyer

observations
the

calcification
of
enamel
organ,

calcified tissues,
45
Uses

the

organ, 24
of preparing sections,

process
on

enamel,

human

of
vascularity

the
enamel

omy,
anat-

10

on

of

Nasmyth's membrane,

on

on

comparative

formula

dental

the, 101

57

in

the

Weasel,

observamicroscopicaltions
tion
of the calcificaof enamel
organ,
on

on

Garden.